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Ethiopian vegetation types, climate and topography

Mengesha Asefa, Min Cao, He YunYun, Ewuketu Mekonnen, Xiaoyang Song, Yang
Jie

PII: S2468-2659(20)30037-8
DOI: https://doi.org/10.1016/j.pld.2020.04.004
Reference: PLD 197

To appear in: Plant Diversity

Received Date: 14 November 2019

Revised Date: 14 April 2020
Accepted Date: 14 April 2020

Please cite this article as: Asefa, M., Cao, M., YunYun, H., Mekonnen, E., Song, X., Jie, Y., Ethiopian
vegetation types, climate and topography, Plant Diversity, https://doi.org/10.1016/j.pld.2020.04.004.

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Elsevier B.V. on behalf of KeAi Communications Co., Ltd.
Ethiopian vegetation types, climate and topography

Mengesha Asefa1,2, Min Cao1, He YunYun1, Ewuketu Mekonnen1, Xiaoyang Song1,

Yang Jie1*
1
CAS, Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden,

Chinese Academy of Sciences, 666303, Yunnan, China

2
Department of Biology, College of Natural and Computation Sciences, University of Gondar,

196, Gondar, Ethiopia

Asefa, M. ([email protected])

Cao, M. ([email protected])

YunYun, H. ([email protected])

Mekonnen, E. ([email protected])

Song, X. ([email protected])

Jie, Y. ([email protected])

*Corresponding author: Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical

Botanical Garden, Chinese Academy of Sciences, 666303, Mengla, Yunnan, China. email:

[email protected]; Tell: +86 183 8715 6270

1 Abstract

2 Ethiopia is land of geographical contrasts with elevations that range from 125 m below sea level

3 in the Danakil Depression to 4533 m above sea level in the Semien Mountains, a world heritage

4 site. The diverse climate of various ecological regions of the country has driven the

5 establishment of diverse vegetation, which range from Afroalpine vegetation in the mountains to

6 the arid and semi-arid vegetation type in the lowlands. The formation of Ethiopian vegetation is

7 highly connected to the climate and geological history of the country. Highland uplift and rift

8 formation due to volcanic forces formed novel habitats with different topography and climatic

9 conditions that have ultimately become drivers for vegetation diversification. Due to Ethiopia’s

10 connection with the temperate biome in the north and the Arabian Peninsula during the dry

11 glacial period, the biotic assemblage of Ethiopian highlands consists of both Afrotropical and

12 palearctic biota. In general, eight distinct vegetation types have been identified in Ethiopia, based

13 mainly on elevation and climate gradients. These vegetation types host their own unique species,

14 but also share several common species. Some of the vegetation types are identified as centers of

15 endemism and have subsequently been identified globally as the East African Afromontane

16 hotspot. Ethiopia is biologically rich, with more than 6500 vascular plant species. Of these

17 species, 12% are endemic mainly due to geographical isolation and unique climatic conditions.

18 However, researchers have yet to extensively investigate the ecology, phenology, as well as the

19 evolutionary, genetics, and conservation status of Ethiopian forests at community and species

20 level over space and time. This lack of research is a barrier to achieving the goal of zero global

21 plant extinctions. Taxa extinction risk assessment has not been extensively carried out for

22 majority of Ethiopian species. Detailed research is needed to explore how vegetation and species

23 respond to rapidly growing environmental change. Currently, human-induced climate change and

1
24 habitat fragmentation are severely threatening the country’s biodiversity, and the consequences

25 of these effects have not been studied at large. Furthermore, we still lack scientific evidence on

26 how micro- and macro-ecological and evolutionary processes have been shaping vegetation

27 structures in this climatically, topographically, and geologically diverse country. These gaps in

28 our knowledge represent an opportunity for ecologists, geneticists, evolutionary biologists,

29 conservation biologists, and other experts to investigate the biodiversity status and the complex

30 ecological processes involved in structuring vegetation dynamics so as to help take effective

31 conservation actions.

32 Keywords: Elevation, Biodiversity, Climate, Ethiopia, Vegetation types, Topography

33 1. Introduction

34 Ethiopia hosts the Eastern Afromontane and Horn of Africa biodiversity hotspots (Mittermeier et

35 al., 2004). Located within the tropics (3º and 15ºN latitude and 33º and 48ºE longitude) (Fig.1),

36 Ethiopia is the center of the East African region that has eleven Afrotropical ecoregions and has

37 been designated a Global 200, an ecoregion of global importance for biodiversity conservation

38 (Olson and Dinerstein, 2002). However, the endangered and critically threatened ecoregions of

39 Ethiopia have been poorly studied.

40 Ethiopia has the fifth highest biodiversity in Africa (Anonymous, 1997). This

41 biodiversity in flora and fauna is strongly associated with the geomorphological history of the

42 region. The country is characterized by dramatic geological history and a broad range of

43 elevations, from the Afar Depression (~125 m below sea level) in the east to the spectacular

44 world heritage mountains of Ras Dashen (4533 m above sea level) in the north (IBC, 2005).

45 This large elevational range has become the source for variation in topography and climate, and

46 has resulted in a heterogenous landscape with high habitat diversity, species diversity, and

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47 centers of species endemism, particularly in the highlands. The Ethiopian highlands are divided

48 by the Great East African Rift Valley into the northwestern and southeastern highlands.

49 Consequently, these highlands have formed unique vegetation types.

50

51 Fig. 1. Location map of Ethiopia (shown in black)

52 Phytogeographically, Ethiopia comprises diverse vegetation types, including the tropical

53 lowland rainforest in the southwest, arid and semi-arid dry woodlands in the East, and the

54 Afroalpine forests in the north and southeast. Researchers have estimated the diverse Ethiopian

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55 topography harbors between 6500 -7000 vascular plant species, of which, 12% are endemic

56 (Tewolde, 1991).

57 Despite this biodiversity, the biological resources of Ethiopia are currently under critical

58 threat mainly due to rapid population growth. The livelihood of the population mainly depends

59 on natural products and lands, and the demand for these is consistently growing. This

60 substantially drives the rapid decline of natural vegetation in Ethiopia (Tolessa et al., 2017). The

61 south, southwest, southeastern and northwestern parts of Ethiopia are the main locations for

62 some intact Afromontane rainforests for which habitat fragmentation and loss has been becoming

63 the dominant threat, as it is also a threat for plant diversity at global level (Corlett, 2016).

64 Ethiopia has been identified as one of the African regions containing large proportion of

65 potentially threatened species with extinction (Stévart et al., 2019). A recent study on the global

66 distribution of rare land plant species reported that Ethiopian highlands are hotspots for large

67 number of rare species (Enquist et al., 2019). However, the ecological and evolutionary history

68 of these vegetations and species have not been widely explored.

69 The elevational and climatic heterogeneity of Ethiopia provide natural conditions to test

70 the response of biodiversity to global environmental change over space and time. However, the

71 majority of studies have simply reported the species richness and floristic composition of

72 different forests (Admassu et al., 2016; Feyera et al., 2007; Kebede et al., 2013; Kuma and

73 Shibru, 2015; Masresha et al., 2015; Soromessa et al., 2004). Few studies have examined how

74 the natural conditions separately and in concert affect species diversity, genetic diversity,

75 vegetation formation, species growth, carbon storage (Breugel et al., 2016; Chala et al., 2016;

76 Derero et al., 2011;Friis et al. 2005; Kidane et al., 2019; Mokria et al., 2015; Siyum et al., 2019).

77 However, to better understand the ecological dynamics of plants and animals of Ethiopia and

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 78 develop effective conservation priorities, researchers should widely investigate the ecology and

79 biology of plant populations, species, communities, and ecosystems across spatial and temporal

80 scales. This is a great opportunity for ecologists, biologists, and conservationists to examine the

81 responses of species, communities, and forests to the unprecedented disturbances, including

82 environmental and climate changes. Furthermore, understanding the ecological dynamics of

83 Ethiopia may help save some rare species that are critical for ecosystem to function.

84 Ethiopia provides an opportunity for a wide range of experts to investigate the ecological

85 and evolutionary processes driving ecological communities and how the geological history of the

86 region contributes for the biogeographical distribution of plants. Although the east African

87 geological history, Great Rift Valley, diverse topography and climate are fascinating, few studies

88 have examined how these factors act, separately and in concert, to affect species colonization,

89 speciation, and extinction ((Evans et al., 2011; Freilich et al., 2016; Friss et al., 2005; Smith et

90 al., 2017). Furthermore, research is needed to describe the phenology, as well as the functional

91 and evolutionary structure of plant taxa, assess the risk of extinction for taxa over time. The low

92 percentage of known taxa for non-woody species and others have become barriers to designing

93 effective conservation strategies and ultimately affects our ability to meet the goals of the zero

94 global plant extinction project (Corlett, 2016).

95 The aim of this review is to introduce ecologists, geneticists, conservation experts,

96 paleobotanists, and evolutionary biologists to the ecological dynamics of forests in Ethiopia.

97 Specifically, we will provide an overview of the diversity of habitats, ecosystems and vegetation

98 types in Ethiopia, and discuss research on the Ethiopian flora and its biogeographical

99 implications. By integrating our understanding of these regional issues with global data, we

100 anticipate that this review will contribute to global conservation efforts.

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101 2. Geological history, climate and landscape features

102 The geological history of Ethiopia is characterized by periods of highland uplift and rift

103 formation. The highlands are divided into northwestern and southeastern regions by the Great

104 Rift Valley, which started to uplift because of volcanic forces 75 million year ago (Olson and

105 Dinerstein, 1998). Generally, highland uplift forms new habitats with hydrological, elevational,

106 and topographical heterogeneity, and has been shown to drive diversification of taxa (Smith et

107 al., 2017).

108 The biotic assemblage of Ethiopian highlands consists of Afrotropical and palearctic

109 biota. During dry glacial periods, a range of escarpments and jebels along the Red Sea permits

110 connectivity with the temperate biomes in the north and Arabian Peninsula, which has provided a

111 route for various palearctic species to colonize the highlands. The Kenyan desert in the south and

112 the great Nile River floodplains in the west restrict immigration of new plant and animal species.

113 Therefore, over the last one million years, the highlands have recruited species from the

114 surrounding dry lowlands.

115 The post-volcanic and post-glacial history of Ethiopia are reflected in the distributions of

116 plant diversity. Specifically, plant diversity is lower in the Ethiopian highlands than in the

117 lowlands. However, the highlands are centers of endemism, largely due to their geographical

118 isolation and unique climatic conditions (Berit and Linder, 2014; Friis et al., 2005; Noroozi et al.,

119 2018). The Great Rift Valley also acts as a geological barrier, extending more than 900 km

120 between the border with Djibouti in the northeast to the border with Kenya in the southwest

121 (Billi, 2015) and with a width between 50 to 60 km. The Rift Valley restricts immigration of

122 many taxa, including plants, birds, amphibians, reptiles, and insects (Freilich et al., 2014; Gottelli

6
123 et al., 2004) and has novel habitats (e.g., rift valley lakes) that provide homes for large numbers

124 of taxa.

125 The Ethiopian highlands are extremely heterogenous, with steep escarpments and deep

126 valleys (Fig. 2). The highlands are known as “the roof of Africa” (in Africa the majority of land

127 over 3,000 m is found in Ethiopia) and reach 4,533 m at the summit of Ras Dashen in the scenic

128 world heritage Simien Mountains (Roberts et al., 2012). Most of the sub-Saharan Africa’s

129 Afroalpine ecosystem above 3200 m is found in Ethiopia (Williams et al., 2004). Ethiopian

130 highlands have been designated hotspots for large number of rare land plant species (Enquist et

131 al., 2019). The highlands are also rich in endemics species of birds (Redman et al., 2011),

132 mammals (Melaku, 2011; Yalden and Largen, 2008) and frogs (Largen and Spawls, 2010).

133 The highlands can be categorized into the lower elevations (800–1500 m), higher

134 elevations (1,500-3,000 m) and peaks (above 3000 m). The lower elevations of Ethiopian

135 highlands support woodland vegetations such as Acacia and Terminalia species. The higher

136 elevations support conifers such as podocarpus and Juniperus species. The peaks (Afroalpine

137 ecosystem) support Erica species, grasslands, and herb layer and are mainly characterized by

138 giant Lobelia species. Details on these vegetation types are given in the following sections.

139

140

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141

142 Fig. 2. Distribution of altitudes in Ethiopia. The small area below sea level at the Afar
143 Depression is shown as the 0-250 meters interval. This map is taken from Friis et al., 2010 (Used
144 under Creative Commons licenses).
145 Several factors are known to regulate Ethiopia’s climate. Generally, the climate is

146 controlled by the Intertropical Convergence Zone (ITCZ) and has a clear bimodal rainfall

147 pattern: the rainy season is from June to September, and the dry season is from October to April.

148 Rainfall generally increases from north to south and east to west, with an average annual rainfall

149 of 600 mm in the northeast and 2000 mm in the southwest (Aerts et al., 2016). This huge climate

150 variability is responsible for the wide range of vegetation types across the country, which

151 includes arid and semi-arid Acacia woodland and Afroalpine forests.

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152 Ethiopia’s proximity to the equator and the complexity of the country’s topography also

153 play a role in regulating Ethiopia’s climate, especially the temperature. Slight seasonal variation

154 on the average monthly temperature has been reported at a given elevation with about 2 oC in the

155 southern and 6 oC in the northern part of the country. Importantly, Ethiopia’s climate varies with

156 elevation (Fig. 3). Elevation is the most important determinant of the average annual temperature

157 of the country with a reduction of 0.5-0.7 oC per 100 meters increase in altitude (Liljequist,

158 1986).

159

160 Fig. 3. Distribution of mean annual rainfall (mm) in Ethiopia. The map is taken from Friis et al.,
161 2010 (Used under Creative Commons licenses).
162

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163 3. Ethiopian vegetation types and distribution

164 Four studies have classified Ethiopian vegetation into different types (Table 1). Sermolli (1957)

165 and White (1983) tried to classify Ethiopian vegetation based on topography. However, these

166 studies produced vegetation maps that differed in terms of the geographical extent of vegetation

167 types. Specifically, the geographical extent of similar vegetation types on these two maps is

168 inconsistent. In addition, the vegetation types on these maps, which were mainly associated with

169 elevation, do not match the complex topography of the country (Friis et al., 2010). Inconsistent

170 vegetation maps pose a clear challenge for stakeholders trying to plan and implement various

171 development and conservation objectives. In response, the Ethiopian Flora Project was

172 established in 1980 and ended in 2009 with the publication of eight volumes of the Flora of

173 Ethiopia and Eritrea. These books extensively describe the geographical distribution of each

174 woody vascular plant species. The project has also produced an insightful book with the new

175 potential vegetation types of Ethiopia showing the location map of each vegetation type (Friis et

176 al., 2010).

177 Table 1. The four classification systems for Ethiopian vegetation types

Classification system # of vegetation Criteria Note

types
Sermolli (1957) 24 altitude includes
Ethiopia, Eritrea
& Somalia
White (1983) 13 altitude
Sebsibe et al. (2004) 8 altitude & climate
Friis et al. (2010) 12 altitude & climate
178

179 Here we present the 8 vegetation types of Ethiopia based on elevation and climate

180 (Sebsebe et al., 2004). These are the same vegetation types officially used by the Institute of

181 Biodiversity of Ethiopia (IBC, 2005). These vegetation types have been re-classified into 12

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182 major types, including sub-vegetation types based on elevation and rainfall pattern (Friis et al.,

183 2010). In this new classification system, the ‘aquatic vegetation type’ has been re-classified into

184 three more detailed vegetation types. Moreover, one vegetation type called ‘Ericaceous belt’ has

185 been added. Ericaceous belt vegetation is dominated by Erica species in the highlands with a

186 narrow range of elevations between 3000 and 3200 m. The upper and lower limit of this

187 Ericaceous belt is bordered by Afroalpine and Afromontane vegetation types respectively. The

188 other 8 vegetation types of this new classification system (Friis et al., 2010) are mostly similar

189 to the officially recognized 8 vegetation types. Therefore, here we present 8 vegetation types:

190 Afroalpine and sub-afroalpine vegetation, Dry evergreen Afromontane forest and grassland

191 complex, Moist evergreen Afromontane forest vegetation, Acacia-Comiphora woodland

192 vegetation, Combretum-Terminalia woodland vegetation, Lowland semi-evergreen forest

193 vegetation, Desert and semi-desert scrubland vegetation, and Aquatic vegetation. We provide the

194 elevational range and number of recorded species in each vegetation type in Table 2.

195 Table 2. Ethiopian vegetation types, number of recorded species obtained from Friis et al., 2010

196 and major threats.

Vegetation types Altitude # of Threats

species
1 Afroalpine & sub- 3200-4533 22 Agriculture, climate change
afroalpine m
2 Dry evergreen 1800-3000 460 Agriculture, grazing, fuel wood
montane & grassland m
complex
3 Moist evergreen 500-2600 135 Agriculture, tea & coffee plantations
montane forest
4 Acacia-Commiphora 900-1900 m 542 Fuel wood, charcoal, agriculture
woodland
5 Combretum- 500-1900 m 199 Fire, settlement
Terminalia woodland
6 Lowland semi- 450-650 m 101 Fire, infrastructure

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 evergreen forest
7 Desert & semi-desert < 400m 131 Invasive species, grazing
scrubland
8 Aquatic vegetation Lowlands to 251 Sedimentation, invasive species,
highlands change to farmlands
197

198 3.1 Afroalpine and sub-afroalpine vegetation

199 Afroalpine and sub-afroalpine vegetation types are widely distributed in Ethiopia, unlike in other

200 African habitats (IBC, 2005). These vegetation types have widely been designated as the national

201 protected areas due to the presence of several endemic plant and animal species. The vegetation

202 is highly restricted to the highlands with elevations ranging from 3200 to 4533 m asl (the peak of

203 Ras Dashen). Despite diurnal temperature fluctuations in which the days are summer-like and the

204 nights are winter-like, seasonal temperature variation is not a characteristic of this vegetation

205 type (Hedberg, 1995). For the past 10,000 years the Ethiopian highlands were largely covered

206 with Afroalpine moorland and grasslands (Fig. 4). The current highland Afroalpine and sub-

207 afroalpine vegetation represents remnants of these patches. Clearly, the remnants of these

208 habitats in the complex highlands of Ethiopia demonstrate the effects of unprecedented and

209 massive anthropogenic activity in the last 19th and 20th centuries (Nyssen et al., 2014).

210 Afroalpine and sub-afroalpine vegetation types in the Ethiopian highlands have not been

211 widely studied even though they are inhabited by endemic plant and animal species (for

212 example, the giant Lobelia rhynchopetalum). Studies have yet to investigate the adaptive

213 mechanisms, phenology, species interactions, genetic diversity, species growth dynamics, and

214 distribution patterns of species in these vegetation types.

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215

216 Fig. 4. Partial view of Afroalpine and sub-afroalpine vegetation

217 3.2 Dry evergreen montane forest and grassland complex

218 Dry evergreen montane forest is usually found between 1,800 and 3,000 m asl, where human

219 settlements and activity dominate (Friis et al., 2010). Dry evergreen montane forest is

220 characterized by a complex set of successions with wide-ranging grasslands rich in legumes,

221 shrubs, and small to large-sized trees to closed forest with vertical canopy stratification (Fig. 5).

222 These forests cover most of the mountainous topography of the Ethiopian highlands.

223 The flora of Dry evergreen montane forest is, except for that of Acacia-Commiphora woodland,

224 the largest of Ethiopian vegetation types, containing many unique species. Some of the most

225 common species in this vegetation type are Juniperus procera, Podocarpus falcatus, Olea

226 europaea subsp. Cuspidata, and Eucalyptus (Friis et al., 2010; IBC, 2005).

227 One of the dominant ecological drivers of the vegetation structure in this ecosystem is

228 grazing. The cattle population in Ethiopia is the largest in Africa, and 10th largest in the world,

229 which has considerable implications for ecological and biological conservation (IBC, 2014).

230 Thus, future studies should examine how grazing patterns shape the phenology, functional

231 strategy, evolutionary history, and plant-animal interaction of the community in combination

13
232 with other potential drivers.

233

234 Fig. 5. Dry Evergreen Montane Forest and Grassland complex

235 3.3 Moist evergreen montane forest vegetation

236 The Ethiopian montane forests are part of the global biodiversity conservation priority areas as

237 well as centers for plant diversity and endemic bird areas primarily because of exceptionally high

238 endemism of species and habitat degradation (ICBP, 1992; WWF and IUCN, 1994). Moist

239 evergreen montane forest is mainly distributed to the southwest escarpment of Ethiopia. It

240 consists of the Afromontane rainforest between elevations of 1500 and 2600 m and the

241 transitional rainforest, which is distributed from 500 to 1500 m in elevation (Sebsebe et al.,

242 2004). The transitional rainforest is composed of species from both the Afromontane rainforest

243 and the lowland forests (Friis, 1992; Gole et al., 2008), suggesting that it links the forest from

244 highlands with lowlands. This forest is the location of the famous wild population of coffee

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245 arabica (Senbeta and Denich, 2006). However, Moist evergreen montane forest is threatened by

246 tea and coffee plantations, human settlement, and agriculture (Gole et al., 2008).

247 3.4 Acacia-Commiphora woodland vegetation

248 Acacia-Commiphora woodland vegetation occurs in the north, east, central and south parts of

249 Ethiopia between 900 and 1900 m, including the rift valley, and consists of drought-resistant

250 trees and shrubs of evergreen or deciduous leaves mainly in the sandy dominated soil type (Friis

251 et al., 2010). Acacia-Commiphora woodland is a complex vegetation type and the most diverse

252 forest type. It forms a complete set of stratum and has unique species with the ability to tolerate

253 limited soil moisture (IBC, 2005). The most characteristic species Acacia-Commiphora

254 woodland vegetation are Acacia and Commiphora genera. The most common of these species

255 include Acacia prasinata (endemic), Acacia bussei, Commiphora alaticaulis, Commiphora

256 boranensis, and Commiphora obovata. Additional common species in this ecosystem include

257 Fabaceae, Burseraceae, Balanitaceae, Combretaceae, and other families. Acacia-Commiphora

258 woodland is also known for endemic succulent species such as Euphorbia awashensis,

259 Euphorbia omariana, and Euphorbia burger (Friis et al., 2010).

260 3.5 Combretum-Terminalia woodland

261 Combretum-Terminalia woodland vegetation is found in the western escarpment of Ethiopia

262 where the Gambela and Alatish national parks are located. It consists of widely distributed small

263 to moderate size trees that are adapted to fire (IBC, 2005). Commonly observed species include

264 Boswellia papyrifera, Anogeissus leiocarpa, and other species in the genera Terminalia and

265 Combretum. The grass stratum of Combretum-Terminalia woodland is well developed and

266 frequently burned during the dry season. Thus, this vegetation type is especially vulnerable to

267 fire during the dry season (Friis et al., 2010) (Fig. 6).

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268

269 Fig. 6. Partial view of Combretum-Terminalia woodland from Friis and Sebsebe, 2008 (Used
270 under Creative Commons licenses).
271 3.6 Lowland, Semi-evergreen forest

272 Lowland, semi-evergreen forest is mainly found in the lowland eastern Gambela region at

273 elevations between 450 and 650 m. Trees are semi-deciduous with a height ranging from 15 to

274 20 m tall with continuous canopy of Baphia abyssinica. The characteristic species of lowland

275 semi-evergreen forest are Alstonia boonei, Morus mesozygia, Pouteria altissima, and Trilepisium

276 madagascariense (Friis et al., 2010). Other species that build this forest type include Celtis toka,

277 Diospyros abyssinica, and Malacantha alnifolia. This forest is prone to fire and human activities

278 for agricultural purpose.

279 3.7 Desert and semi-desert scrubland

280 This scrubland is mainly restricted to the eastern part of Ethiopia below elevations of 400 m. It

281 consists of drought-tolerant small trees, shrubs, and herbs. Characteristic species of Desert and

282 semi-desert scrubland are Acacia ehrenbergiana (Fabaceae), Boswellia ogadensis (Burseraceae),

16
283 Kissenia arabica (Loasaceae), and Ziziphus hamur (Rhamnaceae) (Friis et al., 2010). This

284 vegetation has the highest endemism richness in Ethiopia, and includes species such as the

285 succulent Euphorbia doloensis. However, scrubland is threatened by overgrazing and habitat

286 degradation. In addition, invasive alien species such as Prosopis juliflora also threaten native

287 and endemic species (Kelbessa et al., 1992).

288 3.8 Aquatic vegetation

289 Globally, aquatic ecosystems provide major support to flora and fauna. Ethiopia, where the

290 headwaters of the Blue Nile originate, is one of the world centers of aquatic resources. Aquatic

291 vegetation includes plants found in rivers, reservoirs, lakes, floodplains, and wetlands (Fig. 7).

292 Freshwater lakes, shores and floodplains represent a transition zone between aquatic and

293 terrestrial habitats and are primarily characterized by highly diverse biological communities

294 and centers of biogeochemical activity (Strayer and Findlay, 2010). This vegetation type is

295 geographically restricted to lakes, lake shores, marshes and floodplains. Several freshwater lakes

296 are distributed throughout Ethiopia, including Lake Tana in the northern highlands, and lakes

297 Abaya and Chamo in the south.

298 The species composition, structure and density in aquatic ecosystems vary greatly and

299 depend on the elevation and geographical location. Aquatic vegetation is characterized by poor

300 floristic composition in relation to woody species. Species in aquatic ecosystems are not unique,

301 but rather shared by other adjacent vegetation types. According to Friis et al., (2010), eight

302 species have been recorded in this vegetation type. Lemna aequinoctalis, Wolfia arrhizal and

303 Pistia stratiotes are the characteristic species particularly for freshwater lakes. On other hand,

304 Phoenix reclinata, Lannea edulis, species in the genera Aeschynomene and Sesbania from woody

305 species; Cyperus digitatus, Cyperus denudatus, Cyperus dichroostachys, Cyperus elegantulus,

17
306 and Cyperus latifolius from the sedges are found to be the characteristic species for freshwater

307 swamp, lake shores and floodplains. 36 plant species were recorded just from two wetland sites

308 in the southwesten part of Ethiopia (Woldemariam et al., 2018), suggesting that wetlands are rich

309 in species diversity.

310
311

312 Fig. 7. Vegetation of freshwater lakes and marsh

313 Riverine vegetation commonly includes Celtis africana (Ulmaceae), Ficus sycomorus

314 (Moraceae), Mimusops kummel (Sapotaceae), and Tamarindus indica. Below elevations of 1800

315 m, species composition of riverine vegetation differs greatly from the surrounding forest in

316 lowland dry environments. Interestingly, however, this dissimilarity in species composition

317 decreases as elevation increases (Friis et al., 2010). Well-framed ecological questions should be

318 examined to understand the mechanisms responsible for the spatial variation in species

319 composition, richness and diversity of different vegetations at a wide range of spatial and

320 temporal scales

321 Ethiopian aquatic ecosystems have been estimated to support more than 200 species of

322 phytoplankton (IBC, 2005), and are centers of breeding, feeding, and roosting for several

18
323 resident and migratory bird species. These aquatic ecosystems are also habitats for several native

324 and endemic species of fish. Currently, however, these ecosystems are seriously threatened by

325 climate change and human activities. Furthermore, the aquatic biodiversity of Ethiopia,

326 particularly in Lake Tana, is threatened by invasive water hyacinth. Although Ethiopia is rich in

327 aquatic resources, the biodiversity of this ecosystem has been poorly investigated. Moreover,

328 many ecological questions remain to be explored, including those regarding diversity,

329 distribution patterns, and the impacts of climate change and invasive alien species.

330 Table 2 shows the number of woody species recorded in each vegetation type. The

331 composition of species in each vegetation type raises several broad and specific ecological,

332 evolutionary, and conservation questions. What factors lead to the presence of unique species in

333 these vegetation types? Why are some species confined to narrow ecological ranges? What are

334 the adaptations that allow unique species to tolerate environmental stress? How do these unique

335 species interact with other neighboring non-unique species? What do unique species contribute

336 to ecosystem functioning? How does evolutionary history of species shape their current spatial

337 distribution? What are the values of these species for biodiversity conservation?

338 The vegetation map of Ethiopia is derived from work by Friis et. al (2010) and is based

339 on topography (elevation) and climatic conditions (Fig. 8).

19
340

341 Fig. 8. Map of the vegetation types of Ethiopia obtained from Friis et al., 2010 (Used under
342 Creative Commons licenses). This map contains additional potential vegetation types which are
343 not explained in the article.
344 4 Plant diversity and endemism in Ethiopia

345 Ethiopia is located at the center of two of the world’s 36 biodiversity hotspots, the Eastern Afro-

346 Montane and Horn of Africa biodiversity hotspots (CEPF, 2016; Mittermeier et al., 2004). For

347 both of these biodiversity hotspots researchers have identified significant threats and levels of

348 endemism. The number of endemic plant species and near-endemic species in Ethiopia is high

349 (Friis et al., 2005). Specifically, 12% of the total woody species in Ethiopia has been found to be

350 endemic to different vegetation types (Tewolde, 1991). High species diversity and endemism in

20
351 Ethiopia is most likely associated with its diverse topography and climatic conditions. Levels of

352 endemism can be predicted using various environmental gradients such as climate, temperature,

353 and microhabitat. These predictions help identify centers of endemism and improve efforts to

354 conserve biodiversity. In tropical and temperate regions, endemism increases along an

355 elevational gradient, suggesting that mountains are rich in endemic plant species (Matthews et

356 al., 1993).

357 In Ethiopia, centers of endemism include the highlands and southern part of the country

358 (Ogaden). In southern Ethiopia, near-endemic species include taxa that have narrow

359 geographical ranges shared between the horn of Africa and north Kenya. Endemic species are

360 also linked to the environmental heterogeneity of Ethiopia’s complex topography, which creates

361 diverse local habitats that provide micro-refugia for species during extreme environmental

362 changes (Steinbauer et al., 2013). Furthermore, complex topography is also expected to limit

363 gene flow between isolated populations, thus promoting genetic differentiation that may lead to

364 speciation (Gillespie and Roderick, 2014). Friis et al. (2005) found that high species diversity in

365 Ethiopia is correlated with high elevations. This is consistent with previous studies that have

366 reported that endemic plant and animal species increase as elevation increases (Ghimire, 2005;

367 Noroozi et al., 2018), likely because isolated high mountain regions promote population

368 divergence (Heaney, 2001). However, research is still needed to explore the population status,

369 and the ecological and evolutionary drivers that underlie the distribution of endemic species in

370 space and time, especially in response to rapidly changing environments.

371 In total, there are 476 endemic species in Ethiopia belonging to 69 families and 224

372 genera (Edwards et al., 2000, 1997, 1995; Hedberg et al., 2006, 2003; Hedberg and Edwards,

373 1989; Phillips, 1995; Tadesse, 2004). Five genera but no families are wholly endemic to

21
374 Ethiopia. Based on the studies cited above, a list of endemic species has been compiled by the

375 Institute of Biodiversity Conservation (http://www.terrestrial-

376 biozones.net/Endemic%20Floras/Ethiopian%20Endemics.html). To prioritize conservation

377 efforts, the distribution of these endemic species should be matched to the eight identified

378 vegetation types. Furthermore, the population demography of most of the endemic species has

379 not been well characterized.

380 5 Conclusion

381 Ethiopia is rich in biodiversity, topographical complexity and climate variability which jointly

382 results in different vegetation types. The geological history and climate of the country are mainly

383 responsible for the formation of different vegetation types ranging from drought-tolerant

384 vegetation in the lowlands to cold-tolerant species at the peak of the mountain in the highlands.

385 Ethiopia represents the Afrotropical and palearctic species due to the connection established in

386 the dry glacial period. Most previous studies have focused assessing the floristic composition of

387 forests. Studies remain needed to quantify and predict the effect of disturbance, environment,

388 climate change, geology and topography on the ecology and biology of species. To understand

389 the response of species and vegetation to climate and environmental changes at a global level,

390 researchers are currently able to generate data on a global scale. This ultimately helps identify

391 biodiversity hotspots and ecoregions, and thus design comprehensive conservation strategies that

392 save threatened species. Ethiopian vegetation types provide an opportunity for ecologists,

393 biologists, conservationists and other researchers to investigate how complex biotic interactions,

394 physical environment, ecological, and evolutionary processes determine vegetation dynamics at

395 regional scales and beyond.

396

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Conflict of interest

Authors declare no conflict of interests