Plant Diversity 42 (2020) 302e311

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Plant Diversity
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Research paper

Ethiopian vegetation types, climate and topography

Mengesha Asefa a, b, Min Cao a, Yunyun He a, Ewuketu Mekonnen a, Xiaoyang Song a,
Jie Yang a, *
a
CAS, Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, 666303, Yunnan, China
b
Department of Biology, College of Natural and Computational Sciences, University of Gondar, 196, Gondar, Ethiopia

a r t i c l e i n f o a b s t r a c t

Article history: Ethiopia is land of geographical contrasts with elevations that range from 125 m below sea level in the
Received 14 November 2019 Danakil Depression to 4533 m above sea level in the Semien Mountains, a world heritage site. The diverse
Received in revised form climate of various ecological regions of the country has driven the establishment of diverse vegetation,
14 April 2020
which range from Afroalpine vegetation in the mountains to the arid and semi-arid vegetation type in
Accepted 14 April 2020
the lowlands. The formation of Ethiopian vegetation is highly connected to the climate and geological
Available online 25 May 2020
history of the country. Highland uplift and rift formation due to volcanic forces formed novel habitats
with different topography and climatic conditions that have ultimately become drivers for vegetation
Keywords:
Elevation
diversification. Due to Ethiopia's connection with the temperate biome in the north and the Arabian
Biodiversity Peninsula during the dry glacial period, the biotic assemblage of Ethiopian highlands consists of both
Climate Afrotropical and palearctic biota. In general, eight distinct vegetation types have been identified in
Ethiopia Ethiopia, based mainly on elevation and climate gradients. These vegetation types host their own unique
Vegetation types species, but also share several common species. Some of the vegetation types are identified as centers of
Topography endemism and have subsequently been identified globally as the East African Afromontane hotspot.
Ethiopia is biologically rich, with more than 6500 vascular plant species. Of these species, 12% are
endemic mainly due to geographical isolation and unique climatic conditions. However, researchers have
yet to extensively investigate the ecology, phenology, as well as the evolutionary, genetics, and con-
servation status of Ethiopian vegetations at community and species level over space and time. This lack
of research is a barrier to achieving the goal of zero global plant extinctions. Taxa extinction risk
assessment has not been extensively carried out for majority of Ethiopian species. Detailed research is
needed to explore how vegetation and species respond to rapidly growing environmental change.
Currently, human-induced climate change and habitat fragmentation are severely threatening the
country's biodiversity, and the consequences of these effects have not been studied at large. Furthermore,
we still lack scientific evidence on how micro- and macro-ecological and evolutionary processes have
been shaping vegetation structures in this climatically, topographically, and geologically diverse country.
These gaps in our knowledge represent an opportunity for ecologists, geneticists, evolutionary biologists,
conservation biologists, and other experts to investigate the biodiversity status and the complex
ecological processes involved in structuring vegetation dynamics so as to help take effective conservation
actions.
Copyright © 2020 Kunming Institute of Botany, Chinese Academy of Sciences. Publishing services by
Elsevier B.V. on behalf of KeAi Communications Co., Ltd. This is an open access article under the CC BY-
NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

1. Introduction

Ethiopia hosts the Eastern Afromontane and Horn of Africa

* Corresponding author. Key Laboratory of Tropical Forest Ecology, Xishuang-
biodiversity hotspots (Mittermeier et al., 2004). Located within the
banna Tropical Botanical Garden, Chinese Academy of Sciences, 666303, Mengla,
Yunnan, China. tropics (3 and 15ºN latitude and 33 and 48ºE longitude) (Fig. 1),
E-mail addresses: [email protected] (M. Asefa), [email protected] (M. Cao), Ethiopia is the center of the East African region that has eleven
[email protected] (Y. He), [email protected] (E. Mekonnen), Afrotropical ecoregions and has been designated a Global 200, an
[email protected] (X. Song), [email protected] (J. Yang).
ecoregion of global importance for biodiversity conservation (Olson
Peer review under responsibility of Editorial Office of Plant Diversity.

https://doi.org/10.1016/j.pld.2020.04.004
2468-2659/Copyright © 2020 Kunming Institute of Botany, Chinese Academy of Sciences. Publishing services by Elsevier B.V. on behalf of KeAi Communications Co., Ltd. This
is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
 M. Asefa et al. / Plant Diversity 42 (2020) 302e311 303

The elevational and climatic heterogeneity of Ethiopia provide

natural conditions to test the response of biodiversity to global
environmental change over space and time. However, the majority
of studies have simply reported the species richness and floristic
composition of different forests (Admassu et al., 2016; Feyera et al.,
2007; Kebede et al., 2013; Kuma and Shibru, 2015; Masresha et al.,
2015; Soromessa et al., 2004). Few studies have examined how the
natural conditions separately and in concert affect species diversity,
genetic diversity, vegetation formation, species growth, and carbon
storage (Breugel et al., 2016; Chala et al., 2016; Derero et al., 2011;
Friis et al., 2005; Kidane et al., 2019; Mokria et al., 2015; Siyum et al.,
2019). However, to better understand the ecological dynamics of
plants and animals of Ethiopia and develop effective conservation
priorities, researchers should widely investigate the ecology and
biology of plant populations, species, communities, and ecosystems
across spatial and temporal scales. This is a great opportunity for
ecologists, biologists, and conservationists to examine the re-
sponses of species, communities, and forests to the unprecedented
disturbances, including environmental and climate changes.
Furthermore, understanding the ecological dynamics of Ethiopia
may help save some rare species that are critical for ecosystem to
function.
Ethiopia provides an opportunity for a wide range of experts to
Fig. 1. Location map of Ethiopia (shown in black).
investigate the ecological and evolutionary processes driving
ecological communities and how the geological history of the re-
gion contributes for the biogeographical distribution of plants.
and Dinerstein, 2002). However, the endangered and critically Although the east African geological history, Great Rift Valley,
threatened ecoregions of Ethiopia have been poorly studied. diverse topography and climate are fascinating, few studies have
Ethiopia has the fifth highest biodiversity in Africa (Anonymous, examined how these factors act, separately and in concert, to affect
1997). This biodiversity in flora and fauna is strongly associated species colonization, speciation, and extinction (Evans et al., 2011;
with the geomorphological history of the region. The country is Freilich et al., 2016; Friis et al., 2005; Smith et al., 2017). Further-
characterized by dramatic geological history and a broad range of more, research is needed to describe the phenology, as well as the
elevations, from the Afar Depression (~125 m below sea level) in functional and evolutionary structure of plant taxa, assess the risk
the east to the spectacular world heritage mountains of Ras Dashen of extinction for taxa over time. The low percentage of known taxa
(4533 m above sea level) in the north (IBC, 2005). This large ele- for non-woody species and others have become barriers to
vational range has become the source for variation in topography designing effective conservation strategies and ultimately affects
and climate, and has resulted in a heterogenous landscape with our ability to meet the goals of the zero global plant extinction
high habitat diversity, species diversity, and centers of species project (Corlett, 2016).
endemism, particularly in the highlands. The Ethiopian highlands The aim of this review is to introduce ecologists, geneticists,
are divided by the Great East African Rift Valley into the north- conservation experts, paleobotanists, and evolutionary biologists to
western and southeastern highlands. Consequently, these high- the ecological dynamics of vegetations in Ethiopia. Specifically, we
lands have formed unique vegetation types. will provide an overview of the diversity of habitats, ecosystems
Phytogeographically, Ethiopia comprises diverse vegetation and vegetation types in Ethiopia, and discuss research on the
types, including the tropical lowland rainforest in the southwest, Ethiopian flora and its biogeographical implications. By integrating
arid and semi-arid dry woodlands in the East, and the Afroalpine our understanding of these regional issues with global data, we
forests in the north and southeast. Researchers have estimated the anticipate that this review will contribute to global conservation
diverse Ethiopian topography harbors between 6500 and 7000 efforts.
vascular plant species, of which, 12% are endemic (Tewolde, 1991).
Despite this biodiversity, the biological resources of Ethiopia are 2. Geological history, climate and landscape features
currently under critical threat mainly due to rapid population
growth. The livelihood of the population mainly depends on nat- The geological history of Ethiopia is characterized by periods of
ural products and lands, and the demand for these is consistently highland uplift and rift formation. The highlands are divided into
growing. This substantially drives the rapid decline of natural northwestern and southeastern regions by the Great Rift Valley,
vegetation in Ethiopia (Tolessa et al., 2017). The south, southwest, which started to uplift because of volcanic forces 75 million year
southeastern and northwestern parts of Ethiopia are the main lo- ago (Olson and Dinerstein, 1998). Generally, highland uplift forms
cations for some intact Afromontane rainforests for which habitat new habitats with hydrological, elevational, and topographical
fragmentation and loss has been becoming the dominant threat, as heterogeneity, and has been shown to drive diversification of taxa
it is also a threat for plant diversity at global level (Corlett, 2016). (Smith et al., 2017).
Ethiopia has been identified as one of the African regions con- The biotic assemblage of Ethiopian highlands consists of Afro-
taining large proportion of potentially threatened species with tropical and palearctic biota. During dry glacial periods, a range of
extinction (Ste
vart et al., 2019). A recent study on the global dis- escarpments and jebels along the Red Sea permits connectivity with
tribution of rare land plant species reported that Ethiopian high- the temperate biomes in the north and Arabian Peninsula, which has
lands are hotspots for large number of rare species (Enquist et al., provided a route for various palearctic species to colonize the
2019). However, the ecological and evolutionary history of these highlands. The Kenyan desert in the south and the great Nile River
vegetations and species have not been widely explored. floodplains in the west restrict immigration of new plant and animal
304 M. Asefa et al. / Plant Diversity 42 (2020) 302e311

species. Therefore, over the last one million years, the highlands have Several factors are known to regulate Ethiopia's climate.
recruited species from the surrounding dry lowlands. Generally, the climate is controlled by the Intertropical Conver-
The post-volcanic and post-glacial history of Ethiopia are re- gence Zone (ITCZ) and has a clear bimodal rainfall pattern: the rainy
flected in the distributions of plant diversity. Specifically, plant di- season is from June to September, and the dry season is from
versity is lower in the Ethiopian highlands than in the lowlands. October to April. Rainfall generally increases from north to south
However, the highlands are centers of endemism, largely due to their and east to west, with an average annual rainfall of 600 mm in the
geographical isolation and unique climatic conditions (Berit and northeast and 2000 mm in the southwest (Aerts et al., 2016). This
Linder, 2014; Friis et al., 2005; Noroozi et al., 2018). The Great Rift huge climate variability is responsible for the wide range of vege-
Valley also acts as a geological barrier, extending more than 900 km tation types across the country, which includes arid and semi-arid
between the border with Djibouti in the northeast to the border with Acacia woodland and Afroalpine vegetations.
Kenya in the southwest (Billi, 2015) and with a width between 50 Ethiopia's proximity to the equator and the complexity of the
and 60 km. The Rift Valley restricts immigration of many taxa, country's topography also play a role in regulating Ethiopia's
including plants, birds, amphibians, reptiles, and insects (Freilich climate, especially the temperature. Slight seasonal variation on the
et al., 2014; Gottelli et al., 2004) and has novel habitats (e.g., rift average monthly temperature has been reported at a given eleva-
valley lakes) that provide homes for large numbers of taxa. tion with about 2  C in the southern and 6  C in the northern part of
The Ethiopian highlands are extremely heterogenous, with the country. Importantly, Ethiopia's climate varies with elevation
steep escarpments and deep valleys (Fig. 2). The highlands are (Fig. 3). Elevation is the most important determinant of the average
known as “the roof of Africa” (in Africa the majority of land over annual temperature of the country with a reduction of 0.5e0.7  C
3000 m is found in Ethiopia) and reach 4533 m at the summit of Ras per 100 m increase in altitude (Liljequist, 1986).
Dashen in the scenic world heritage Simien Mountains (Roberts
et al., 2012). Most of the sub-Saharan Africa's Afroalpine 3. Ethiopian vegetation types and distribution
ecosystem above 3200 m is found in Ethiopia (Williams et al.,
2004). Ethiopian highlands have been designated hotspots for Four studies have classified Ethiopian vegetation into different
large number of rare land plant species (Enquist et al., 2019). The types (Table 1). Pichi Sermolli (1957) and White (1983) tried to
highlands are also rich in endemics species of birds (Redman et al., classify Ethiopian vegetation based on topography. However, these
2011), mammals (Melaku, 2011; Yalden and Largen, 2008) and frogs studies produced vegetation maps that differed in terms of the
(Largen and Spawls, 2010). geographical extent of vegetation types. Specifically, the
The highlands can be categorized into the lower elevations geographical extent of similar vegetation types on these two maps
(800e1500 m), higher elevations (1500e3000 m) and peaks (above is inconsistent. In addition, the vegetation types on these maps,
3000 m). The lower elevations of Ethiopian highlands support which were mainly associated with elevation, do not match the
woodland vegetations such as Acacia and Terminalia species. The complex topography of the country (Friis et al., 2010). Inconsistent
higher elevations support conifers such as podocarpus and Juniperus vegetation maps pose a clear challenge for stakeholders trying to
species. The peaks (Afroalpine ecosystem) support Erica species, plan and implement various development and conservation ob-
grasslands, and herb layer and are mainly characterized by giant jectives. In response, the Ethiopian Flora Project was established in
Lobelia species. Details on these vegetation types are given in the 1980 and ended in 2009 with the publication of eight volumes of
following sections. the Flora of Ethiopia and Eritrea. These books extensively describe

Fig. 2. Distribution of altitudes in Ethiopia. The small area below sea level at the Afar Depression is shown as the 0e250 m interval. With permission, this map is reproduced from
Friis et al. (2010) (Used under Creative Commons licenses).
 M. Asefa et al. / Plant Diversity 42 (2020) 302e311 305

Fig. 3. Distribution of mean annual rainfall (mm) in Ethiopia. With permission, this map is reproduced from Friis et al. (2010) (Used under Creative Commons licenses).

Table 1
The four classification systems for Ethiopian vegetation types.

Classification system # of vegetation types Criteria Note

Pichi 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

the geographical distribution of each woody vascular plant species. range and number of recorded species in each vegetation type in
The project has also produced an insightful book with the new Table 2.
potential vegetation types of Ethiopia showing the location map of
each vegetation type (Friis et al., 2010). 3.1. Afroalpine and sub-afroalpine vegetation
Here we present the 8 vegetation types of Ethiopia based on
elevation and climate (Sebsebe et al., 2004). These are the same Afroalpine and sub-afroalpine vegetation types are widely
vegetation types officially used by the Institute of Biodiversity of distributed in Ethiopia, unlike in other African habitats (IBC, 2005).
Ethiopia (IBC, 2005). These vegetation types have been re-classified These vegetation types have widely been designated as the national
into 12 major types, including sub-vegetation types based on protected areas due to the presence of several endemic plant and
elevation and rainfall pattern (Friis et al., 2010). In this new clas- animal species. The vegetation is highly restricted to the highlands
sification system, the ‘aquatic vegetation type’ has been re- with elevations ranging from 3200 to 4533 m asl (the peak of Ras
classified into three more detailed vegetation types. Moreover, Dashen). Despite diurnal temperature fluctuations in which the
one vegetation type called ‘Ericaceous belt’ has been added. days are summer-like and the nights are winter-like, seasonal
Ericaceous belt vegetation is dominated by Erica species in the temperature variation is not a characteristic of this vegetation type
highlands with a narrow range of elevations between 3000 and (Hedberg, 1995). For the past 10,000 years the Ethiopian highlands
3200 m. The upper and lower limit of this Ericaceous belt is were largely covered with Afroalpine moorland and grasslands
bordered by Afroalpine and Afromontane vegetation types (Fig. 4). The current highland Afroalpine and sub-afroalpine vege-
respectively. The other 8 vegetation types of this new classification tation represents remnants of these patches. Clearly, the remnants
system (Friis et al., 2010) are mostly similar to the officially recog- of these habitats in the complex highlands of Ethiopia demonstrate
nized 8 vegetation types. Therefore, here we present 8 vegetation the effects of unprecedented and massive anthropogenic activity in
types: Afroalpine and sub-afroalpine vegetation, Dry evergreen the last 19th and 20th centuries (Nyssen et al., 2014).
Afromontane forest and grassland complex, Moist evergreen Afroalpine and sub-afroalpine vegetation types in the Ethiopian
Afromontane forest vegetation, Acacia-Comiphora woodland highlands have not been widely studied even though they are
vegetation, Combretum-Terminalia woodland vegetation, Lowland inhabited by endemic plant and animal species (for example, the
semi-evergreen forest vegetation, Desert and semi-desert scrub- giant Lobelia rhynchopetalum). Studies have yet to investigate the
land vegetation, and Aquatic vegetation. We provide the elevational adaptive mechanisms, phenology, species interactions, genetic
306 M. Asefa et al. / Plant Diversity 42 (2020) 302e311

Table 2
Ethiopian vegetation types, number of recorded species obtained from Friis et al. (2010) and major threats.

Vegetation types Altitude # of species Threats

1 Afroalpine & sub-afroalpine 3200e4533 m 22 Agriculture, climate change

2 Dry evergreen montane & grassland complex 1800e3000 m 460 Agriculture, grazing, fuel wood
3 Moist evergreen montane forest 500e2600 m 135 Agriculture, tea & coffee plantations
4 Acacia-Commiphora woodland 900e1900 m 542 Fuel wood, charcoal, agriculture
5 Combretum-Terminalia woodland 500e1900 m 199 Fire, settlement
6 Lowland semi-evergreen forest 450e650 m 101 Fire, infrastructure
7 Desert & semi-desert scrubland <400 m 131 Invasive species, grazing
8 Aquatic vegetation Lowlands to highlands 251 Sedimentation, invasive species, change to farmlands

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

diversity, species growth dynamics, and distribution patterns of diversity and endemic bird areas primarily because of excep-
species in these vegetation types. tionally high endemism of species and habitat degradation (ICBP,
1992; WWF and IUCN, 1994). Moist evergreen montane forest is
3.2. Dry evergreen montane forest and grassland complex mainly distributed to the southwest escarpment of Ethiopia. It
consists of the Afromontane rainforest between elevations of
Dry evergreen montane forest is usually found between 1800 1500 and 2600 m and the transitional rainforest, which is
and 3000 m asl, where human settlements and activity dominate distributed from 500 to 1500 m in elevation (Sebsebe et al.,
(Friis et al., 2010). Dry evergreen montane forest is characterized by 2004). The transitional rainforest is composed of species from
a complex set of successions with wide-ranging grasslands rich in both the Afromontane rainforest and the lowland forests (Friis,
legumes, shrubs, and small to large-sized trees to closed forest with 1992; Gole et al., 2008), suggesting that it links the forest from
vertical canopy stratification (Fig. 5). These forests cover most of the highlands with lowlands. This forest is the location of the famous
mountainous topography of the Ethiopian highlands. wild population of coffee arabica (Senbeta and Denich, 2006).
The flora of Dry evergreen montane forest is, except for that of However, Moist evergreen montane forest is threatened by tea
Acacia-Commiphora woodland, the largest of Ethiopian vegetation and coffee plantations, human settlement, and agriculture (Gole
types, containing many unique species. Some of the most common et al., 2008).
species in this vegetation type are Juniperus procera, Podocarpus
falcatus, Olea europaea subsp. Cuspidata, and Eucalyptus (Friis et al., 3.4. Acacia-Commiphora woodland vegetation
2010; IBC, 2005). Eucalyptus has rarely becomes naturalized as its
presence in this vegetation is due to direct human interference Acacia-Commiphora woodland vegetation occurs in the north,
(Friis, 1995). east, central and south parts of Ethiopia between 900 and 1900 m,
One of the dominant ecological drivers of the vegetation including the rift valley, and consists of drought-resistant trees and
structure in this ecosystem is grazing. The cattle population in shrubs of evergreen or deciduous leaves mainly in the sandy
Ethiopia is the largest in Africa, and 10th largest in the world, which dominated soil type (Friis et al., 2010). Acacia-Commiphora
has considerable implications for ecological and biological conser- woodland is a complex vegetation type and the most diverse for-
vation (IBC, 2014). Thus, future studies should examine how grazing est type. It forms a complete set of stratum and has unique species
patterns shape the phenology, functional strategy, evolutionary with the ability to tolerate limited soil moisture (IBC, 2005). The
history, and planteanimal interaction of the community in com- most characteristic species of Acacia-Commiphora woodland
bination with other potential drivers. vegetation are Acacia and Commiphora genera. The most common
of these species include Acacia prasinata (endemic), Acacia bussei,
3.3. Moist evergreen montane forest vegetation Commiphora alaticaulis, Commiphora boranensis, and Commiphora
obovata. Additional common species in this ecosystem include
The Ethiopian montane forests are part of the global biodi- Fabaceae, Burseraceae, Balanitaceae, Combretaceae, and other
versity conservation priority areas as well as centers for plant families. Acacia-Commiphora woodland is also known for endemic
 M. Asefa et al. / Plant Diversity 42 (2020) 302e311 307

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

succulent species such as Euphorbia awashensis, Euphorbia omari- Trees are semi-deciduous with a height ranging from 15 to 20 m
ana, and Euphorbia burger (Friis et al., 2010). tall with continuous canopy of Baphia abyssinica. The character-
istic species of lowland semi-evergreen forest are Alstonia boonei,
Morus mesozygia, Pouteria altissima, and Trilepisium madagascar-
3.5. Combretum-Terminalia woodland
iense (Friis et al., 2010). Other species that build this forest type
include Celtis toka, Diospyros abyssinica, and Malacantha alnifolia.
Combretum-Terminalia woodland vegetation is found in the
This forest is prone to fire and human activities for agricultural
western escarpment of Ethiopia where the Gambela and Alatish
purpose.
national parks are located. It consists of widely distributed small to
moderate size trees that are adapted to fire (IBC, 2005). Commonly
observed species include Boswellia papyrifera, Anogeissus leiocarpa,
3.7. Desert and semi-desert scrubland
and other species in the genera Terminalia and Combretum. The
grass stratum of Combretum-Terminalia woodland is well devel-
This scrubland is mainly restricted to the eastern part of Ethiopia
oped and frequently burned during the dry season. Thus, this
below elevations of 400 m. It consists of drought-tolerant small
vegetation type is especially vulnerable to fire during the dry sea-
trees, shrubs, and herbs. Characteristic species of Desert and semi-
son (Friis et al., 2010) (Fig. 6).
desert scrubland are Acacia ehrenbergiana (Fabaceae), Boswellia
ogadensis (Burseraceae), Kissenia arabica (Loasaceae), and Ziziphus
3.6. Lowland, semi-evergreen forest hamur (Rhamnaceae) (Friis et al., 2010). This vegetation has the
highest endemism richness in Ethiopia, and includes species such
Lowland, semi-evergreen forest is mainly found in the lowland as the succulent Euphorbia doloensis. However, scrubland is
eastern Gambela region at elevations between 450 and 650 m. threatened by overgrazing and habitat degradation. In addition,

Fig. 6. Partial view of Combretum-Terminalia woodland obtained with permission from Friis and Sebsebe (2008) (Used under Creative Commons licenses).
308 M. Asefa et al. / Plant Diversity 42 (2020) 302e311

invasive alien species such as Prosopis juliflora also threaten native Ethiopian aquatic ecosystems have been estimated to support
and endemic species (Kelbessa et al., 1992). more than 200 species of phytoplankton (IBC, 2005), and are cen-
ters of breeding, feeding, and roosting for several resident and
migratory bird species. These aquatic ecosystems are also habitats
3.8. Aquatic vegetation for several native and endemic species of fish. Currently, however,
these ecosystems are seriously threatened by climate change and
Globally, aquatic ecosystems provide major support to flora and human activities. Furthermore, the aquatic biodiversity of Ethiopia,
fauna. Ethiopia, where the headwaters of the Blue Nile originate, is particularly in Lake Tana, is threatened by invasive water hyacinth.
one of the world centers of aquatic resources. Aquatic vegetation Although Ethiopia is rich in aquatic resources, the biodiversity of
includes plants found in rivers, reservoirs, lakes, floodplains, and this ecosystem has been poorly investigated. Moreover, many
wetlands (Fig. 7). ecological questions remain to be explored, including those
Freshwater lakes, shores and floodplains represent a transition regarding diversity, distribution patterns, and the impacts of
zone between aquatic and terrestrial habitats and are primarily climate change and invasive alien species.
characterized by highly diverse biological communities and centers Table 2 shows the number of woody species recorded in each
of biogeochemical activity (Strayer and Findlay, 2010). This vege- vegetation type. The composition of species in each vegetation type
tation type is geographically restricted to lakes, lake shores, raises several broad and specific ecological, evolutionary, and
marshes and floodplains. Several freshwater lakes are distributed conservation questions. What factors lead to the presence of unique
throughout Ethiopia, including Lake Tana in the northern high- species in these vegetation types? Why are some species confined
lands, and lakes Abaya and Chamo in the south. to narrow ecological ranges? What are the adaptations that allow
The species composition, structure and density in aquatic eco- unique species to tolerate environmental stress? How do these
systems vary greatly and depend on the elevation and geographical unique species interact with other neighboring non-unique spe-
location. Aquatic vegetation is characterized by poor floristic cies? What do unique species contribute to ecosystem functioning?
composition in relation to woody species. Species in aquatic eco- How does evolutionary history of species shape their current
systems are not unique, but rather shared by other adjacent vege- spatial distribution? What are the values of these species for
tation types. According to Friis et al. (2010), eight species have been biodiversity conservation?
recorded in this vegetation type. Lemna aequinoctalis, Wolfia The vegetation map of Ethiopia is derived from work by Friis
arrhizal and Pistia stratiotes are the characteristic species particu- et al. (2010) and is based on topography (elevation) and climatic
larly for freshwater lakes. On other hand, Phoenix reclinata, Lannea conditions (Fig. 8).
edulis, species in the genera Aeschynomene and Sesbania from
woody species; Cyperus digitatus, Cyperus denudatus, Cyperus
dichroostachys, Cyperus elegantulus, and Cyperus latifolius from the 4. Plant diversity and endemism in Ethiopia
sedges are found to be the characteristic species for freshwater
swamp, lake shores and floodplains. 36 plant species were recorded Ethiopia is located at the center of two of the world's 36
just from two wetland sites in the southwesten part of Ethiopia biodiversity hotspots, the Eastern Afro-Montane and Horn of Africa
(Woldemariam et al., 2018), suggesting that wetlands are rich in biodiversity hotspots (CEPF, 2016; Mittermeier et al., 2004). For
species diversity. both of these biodiversity hotspots researchers have identified
Riverine vegetation commonly includes Celtis africana (Ulma- significant threats and levels of endemism. The number of endemic
ceae), Ficus sycomorus (Moraceae), Mimusops kummel (Sapotaceae), plant species and near-endemic species in Ethiopia is high (Friis
and Tamarindus indica. Below elevations of 1800 m, species et al., 2005). Specifically, 12% of the total woody species in
composition of riverine vegetation differs greatly from the sur- Ethiopia has been found to be endemic to different vegetation types
rounding forest in lowland dry environments. Interestingly, how- (Tewolde, 1991). High species diversity and endemism in Ethiopia is
ever, this dissimilarity in species composition decreases as most likely associated with its diverse topography and climatic
elevation increases (Friis et al., 2010). Well-framed ecological conditions. Levels of endemism can be predicted using various
questions should be examined to understand the mechanisms environmental gradients such as climate, temperature, and
responsible for the spatial variation in species composition, rich- microhabitat. These predictions help identify centers of endemism
ness and diversity of different vegetations at a wide range of spatial and improve efforts to conserve biodiversity. In tropical and
and temporal scales. temperate regions, endemism increases along an elevational

Fig. 7. Vegetation of freshwater lakes and marsh.

 M. Asefa et al. / Plant Diversity 42 (2020) 302e311 309

Fig. 8. Map of the vegetation types of Ethiopia obtained with permission from Friis et al. (2010) (Used under.Creative Commons licenses). This map contains additional potential
vegetation types which are not explained in the article.

gradient, suggesting that mountains are rich in endemic plant uploads/downloads/Endemic_plants_of_Ethiopia-Reported.pdf).

species (Matthews et al., 1993). To prioritize conservation efforts, the distribution of these endemic
In Ethiopia, centers of endemism include the highlands and species should be matched to the eight identified vegetation types.
southern part of the country (Ogaden). In southern Ethiopia, near- Furthermore, the population demography of most of the endemic
endemic species include taxa that have narrow geographical species has not been well characterized.
ranges shared between the horn of Africa and north Kenya.
Endemic species are also linked to the environmental heteroge-
neity of Ethiopia's complex topography, which creates diverse 5. Conclusion
local habitats that provide micro-refugia for species during
extreme environmental changes (Steinbauer et al., 2013). Ethiopia is rich in biodiversity, topographical complexity and
Furthermore, complex topography is also expected to limit gene climate variability which jointly results in different vegetation
flow between isolated populations, thus promoting genetic dif- types. The geological history and climate of the country are
ferentiation that may lead to speciation (Gillespie and Roderick, mainly responsible for the formation of different vegetation
2014). Friis et al. (2005) found that high species diversity in types ranging from drought-tolerant vegetation in the lowlands
Ethiopia is correlated with high elevations. This is consistent with to cold-tolerant species at the peak of the mountain in the
previous studies that have reported that endemic plant and ani- highlands. Ethiopia represents the Afrotropical and palearctic
mal species increase as elevation increases (Ghimire, 2005; species due to the connection established in the dry glacial
Noroozi et al., 2018), likely because isolated high mountain re- period. Most previous studies have focused assessing the
gions promote population divergence (Heaney, 2001). However, floristic composition of forests. Studies remain needed to
research is still needed to explore the population status, and the quantify and predict the effect of disturbance, environment,
ecological and evolutionary drivers that underlie the distribution climate change, geology and topography on the ecology and
of endemic species in space and time, especially in response to biology of species. To understand the response of species and
rapidly changing environments. vegetation to climate and environmental changes at a global
In total, there are 476 endemic species in Ethiopia belonging to level, researchers are currently able to generate data on a
69 families and 224 genera (Edwards et al., 2000, 1997; 1995; global scale. This ultimately helps identify biodiversity hotspots
Hedberg et al., 2006, 2003; Hedberg and Edwards, 1989; Phillips, and ecoregions, and thus design comprehensive conservation
1995; Tadesse, 2004). Five genera but no families are wholly strategies that save threatened species. Ethiopian vegetation
endemic to Ethiopia. Based on the studies cited above, a list of types provide an opportunity for ecologists, biologists, conser-
endemic species has been compiled by Awas in the Institute of vationists and other researchers to investigate how complex
Biodiversity Conservation (http://www.ebi.gov.et/wp-content/ biotic interactions, physical environment, ecological, and
310 M. Asefa et al. / Plant Diversity 42 (2020) 302e311

evolutionary processes determine vegetation dynamics at Friis, I., 1992. Forests and forest trees of northeast tropical Africa: their natural
habitats and distribution patterns in Ethiopia, Djibouti and Somalia. Kew Bull.
regional scales and beyond.
Addit. Ser. 15, 1e396.
Friis, I., 1995. Myrtaceae. In: Edwards, S., Tadesse, M., Hedberg, I. (Eds.), Flora of
Ethiopia and Eritrea: Canellaceae to Euphorbiaceae, 2, part 2. Addis Ababa
Author contributions university, Uppsala university, pp. 71e106.
Friis, I., Sebsebe, D., Breugel, P., 2010. Atlas of the Potential Vegetation of Ethiopia
MA and YJ wrote the manuscript, and all authors provided Atlas of the Potential Vegetation of Ethiopia.
Friis, I., Thulin, M., Adsersen, H., Burger, A., 2005. Patterns of plant diversity and
comments. endemism in the Horn of Africa. Biol. Skr. 55, 289e314. ISSN 0366-3612. ISBN
87-7304-304e4.
Ghimire, S.K., 2005. The endemic flora in Dolpo, north-west Nepal: distribution
Declaration of Competing Interest patterns, life forms, habitat specificity and conservation status. Bot. Orient. 5,
30e39.
Gillespie, R., Roderick, G., 2014. Evolution: geology and climate drive diversification.
Authors declare no conflict of interests. Nature 509, 297e298.
Gole, T., Thomas, B., Manfred, D., Demel, T., 2008. Floristic composition and envi-
ronmental factors characterizing coffee forests in southwest Ethiopia. For. Ecol.
References Manag. 255, 2138e2150. https://doi.org/10.1016/j.foreco.2007.12.028.
Gottelli, D., Marino, J., Sillero-Zubiri, C., Funk, S., 2004. The effect of the last glacial
Admassu, A., Teshome, S., Ensermu, K., Abyot, D., Alemayhu, K., 2016. Floristic age on speciation and population genetic structure of the endangered Ethiopian
composition and plant community types of Agama forest, an afromontane wolf (Canis simensis). Mol. Ecol. 13, 2275e2286.
forest in southwest Ethiopia. J. Ecol. Nat. Environ. 8, 55e69. https://doi.org/ Heaney, L.R., 2001. Small mammal diversity along elevational gradients in the
10.5897/jene2015.0547. Philippines: an 333 assessment of patterns and hypotheses. Global Ecol. Bio-
Aerts, R., Overtveld, K. Van, November, E., Wassie, A., Abiyu, A., Demissew, S., geogr. 10, 15e39.
Daye, D.D., Giday, K., Haile, M., Tewoldeberhan, S., Teketay, D., Hedberg, I., Edwards, S., 1989. Flora of Ethiopia. In: Pittosporaceae to Aral-
Teklehaimanot, Z., Binggeli, P., Deckers, J., Friis, I., Gratzer, G., Hermy, M., iacae.the National Herbarium, vol. 3. Addis Ababa University, Addis Ababa &
Heyn, M., Honnay, O., Paris, M., Sterck, F.J., Muys, B., Bongers, F., Healey, J.R., Uppsala.
2016. Conservation of the Ethiopian church forests : threats , opportunities and Flora of Ethiopia and Eritrea. In: Hedberg, I., Edwards, S., Nemomissa, S. (Eds.), 2003.
implications for their management. Sci. Total Environ. 551e552, 404e414. Apiaceae to Dipsaceae. The National Herbarium, vol. 4. Addis Ababa University,
https://doi.org/10.1016/j.scitotenv.2016.02.034. Addis Ababa & Uppsala (2).
Anonymous, 1997. Ethiopia: National Conservation Strategy, Phase I Report. Ethio- Hedberg, I., Kelbessa, E., Edwards, S., Demissew, S., Persson, E., 2006. Flora of
pian Environmental Authority, Addis Abeba. Ethiopia and Eritrea. Vol 5, gentianaceae to cyclocheilaceae. The National
Awas, T., n.d. Endemic plants of Ethiopia: preliminary working list to contribute to Herbarium, Addis Ababa University, Addis Ababa and Uppsala.
National plant conservation. Retrieved on June 25, 2019 from http://www.ebi. Hedberg, O., 1995. Features of afroalpine ecology. Facsimile edition. Acta Phyto-
gov.et/wp-content/uploads/downloads/Endemic_plants_of_Ethiopia-Reported. geogr. Suec. 49, 1e144.
pdf. IBC, 2014. Ethiopia's Revised National Biodiversity Strategy and Action Plan.
Berit, G., Linder, P., 2014. Species richness, endemism and species composition in IBC, 2005. Institute of Biodiversity Conservation, National Biodiversity Strategy and
the tropical Afroalpine flora. Alpine Bot. 124 (2), 165e177. Action Plan, Government of the Fedral Democratic Republic of Ethiopia, vol. 115.
Billi, P., 2015. Geomorphological landscapes of Ethiopia. In: Billi, P. (Ed.), Landscapes https://doi.org/10.1016/S0006-3495(96)79498-5.
Landforms Ethiop. Springer, Dordrecht, The Netherlands, pp. 3e32. ICBP, 1992. Putting Biodiversity on the Map: Priority Areas for Global Conservation.
Breugel, P. Van, Friis, I., Demissew, S., Lillesø, J.B., Kindt, R., 2016. Current and future Int. Counc. Bird Preserv. Cambridge, U.K.
fire regimes and their influence on natural vegetation in Ethiopia. Ecosystems Kebede, M., Yirdaw, E., Luukkanen, O., Lemenih, M., 2013. Plant community analysis
19, 369e386. https://doi.org/10.1007/s10021-015-9938-x. and effect of environmental factors on the diversity of woody species in the
CEPF, 2016. Announcing the World's 36th Biodiversity Hotspot: the North American moist Afromontane forest of Wondo Genet, South Central Ethiopia. Biodivers.
Coastal Plain. Retrieved on June 04, 2019 from. http://www.cepf.net/news/top_ Res. Conserv. 29, 63e80. https://doi.org/10.2478/biorc-2013-0003.
stories/Pages/Announcing-the-Worlds-36thBiodiversity-Hotspot.aspx. Kelbessa, E., Demissew, S., Woldu, Z., Edwards, S., 1992. Some threatened
Chala, D., Brochmann, C., Psomas, A., Ehrich, D., Gizaw, A., Masao, C.A., endemic plants of Ethiopia. In: Edwards, Sue, Asfaw, Zemede (Eds.), The
Bakkestuen, V., Zimmermann, N.E., 2016. Bye to tropical alpine plant giants Status of Some Plant Resources in Parts of Tropical Africa. Botany 2000: East
under warmer climates ? Loss of range and genetic diversity in Lobelia rhyn- and Central Africa. NAPRECA Monograph Series No. 2. Published by NAPRECA.
chopetalum. Ecol. Evol. 2016 6 6, 8931e8941. https://doi.org/10.1002/ Addis Ababa Univ.
ece3.2603. Kidane, Y.O., Steinbauer, M.J., Beierkuhnlein, C., 2019. Dead end for endemic plant
Corlett, R.T., 2016. Plant diversity in a changing world: status, trends, and conser- species? A biodiversity hotspot under pressure. Glob. Ecol. Conserv. 19, e00670
vation needs. Plant Divers. 38, 10e16. https://doi.org/10.1016/j.pld.2016.01.001. https://doi.org/10.1016/j.gecco.2019.e00670.
Derero, A., Gailing, O., Finkeldey, R., 2011. Maintenance of genetic diversity in Cordia Kuma, M., Shibru, S., 2015. Floristic composition , vegetation structure and regen-
africana Lam., a declining forest tree species in Ethiopia. Tree Genet. Genomes 7, eration status of Kimphe Lafa natural forest. J. Bot., Le 5, 19e32.
1e9. https://doi.org/10.1007/s11295-010-0310-1. Largen, M., Spawls, S., 2010. The Amphibians and Reptiles of Ethiopia and Eritrea.
Flora of Ethiopia and Eritrea. In: Edwards, S., Demissew, S., Hedberg, I. (Eds.), 1997. Edition Chimaira, Frankfurt am Main.
Hydrocharitaceae to Arecaceae, vol. 6. The National Herbarium, Addis Ababa Liljequist, 1986. Some aspects of the climate of Ethiopia. Symb. Bot. Ups. 26 (2),
University, Addis Ababa & Uppsala. 19e30.
Edwards, S., Tadesse, M., Demissew, S., Hedberg, I., 2000. Flora of Ethiopia and Masresha, G., Soromessa, T., Kelbessa, E., 2015. Status and species diversity of
Eritrea. In: Magnoliaceae to Flacourtiaceae. The National Herbarium, vol. 2. Alemsaga forest , northwestern. Adv. Life Sci. Technol. 34, 87e100.
Addis Ababa University, Addis Ababa & Uppsala (1). Matthews, W.S., van W, A.K., Bredenkamp, G.J., 1993. Endemic flora of north-eastern
Edwards, S., Tadesse, M., Hedberg, I., 1995. Flora of Ethiopia and Eritrea. In: Can- transval escarp- ment, South Africa. Biol. Conserv. 63, 83e94.
ellaceae to Euphorbiaceae. The National Herbarium, vol. 2. Addis Ababa Uni- Melaku, T., 2011. Wildlife in Ethiopia: endemic large mammals. World J. Zool. 6 (2),
versity, Addis Ababa & Uppsala (2). 108e116.
Enquist, B.J., Feng, X., Boyle, B., Maitner, B., Newman, E.A., Jørgensen, P.M., Mittermeier, R.A., Gil, P.R., Hoffman, M., Pilgrim, J., Brooks, T., Mittermeier, C.G.,
Roehrdanz, P.R., Thiers, B.M., Burger, J.R., Corlett, R.T., Couvreur, T.L.P., Dauby, G., Lamoreux, J., da Fonseca, G.A.B., 2004. 2005. Hotspots Revisited: Earth's Bio-
Donoghue, J.C., Foden, W., Lovett, J.C., Marquet, P.A., Merow, C., Midgley, G., logically Richest and Most Endangered Terrestrial Ecoregions. Conservation
Morueta-Holme, N., Neves, D.M., Oliveira-Filho, A.T., Kraft, N.J.B., Park, D.S., International, Washington.
Peet, R.K., Pillet, M., Serra-Diaz, J.M., Sandel, B., Schildhauer, M., Símov 
, I.,
a Mokria, M., Gebrekirstos, A., Aynekulu, E., Bra €uning, A., 2015. Tree dieback affects
Violle, C., Wieringa, J.J., Wiser, S.K., Hannah, L., Svenning, J.C., McGill, B.J., 2019. climate change mitigation potential of a dry afromontane forest in northern
The commonness of rarity: global and future distribution of rarity across land Ethiopia. For. Ecol. Manag. 344, 73e83. https://doi.org/10.1016/
plants. Sci. Adv. 5, 1e14. https://doi.org/10.1126/sciadv.aaz0414. j.foreco.2015.02.008.
Evans, B., Bliss, S., Mendel, S., Tinsley, R., 2011. The Rift Valley is a major barrier to Noroozi, J., Talebi, A., Doostmohammadi, M., Rumpf, S.B., Linder, H.P.,
dispersal of African clawed frogs (Xenopus) in Ethiopia. Mol. Ecol. 20, Schneeweiss, G.M., 2018. Hotspots within a global biodiversity hotspot-areas of
4216e4230. endemism are associated with high mountain ranges. Sci. Rep. 8, 1e10. https://
Feyera, S., Tadesse, W., Sebsebe, D., Denichi, M., 2007. Florestic diversity and doi.org/10.1038/s41598-018-28504-9.
composition of sheko, southwest Ethiopia. Ethiop. J. Biol. Sci. 6, 11e42. Nyssen, J., Frankl, A., Haile, M., Hurni, H., Descheemaeker, K., Crummey, D., Ritler, A.,
Freilich, X., Anado
n, J., Bukala, J., Calderon, O., Chakraborty, R., Boissinot, S., 2016. Portner, B., Nievergelt, B., Moeyersons, J., Munro, N., Deckers, J., Billi, P.,
Comparative phylogeography of Ethiopian anurans: impact of the Great Rift Poesen, J., 2014. Environmental conditions and human drivers for changes to
Valley and Pleistocene climate change. BMC Evol. Biol. 16, 206e225. north Ethiopian mountain landscapes over 145 years. Sci. Total Environ.
Freilich, X., Tollis, M., Boissinot, S., 2014. Hiding in the highlands: evolution of a frog 485e486, 164e179. https://doi.org/10.1016/j.scitotenv.2014.03.052.
species complex of the genus Ptychadena in the Ethiopian highlands. Mol. Olson, D.M., Dinerstein, E., 2002. The global 200: priority ecoregions for global
Phylogenet. Evol. 71, 157e169. conservation. Ann. Mo. Bot. Gard. 89, 199. https://doi.org/10.2307/3298564.
 M. Asefa et al. / Plant Diversity 42 (2020) 302e311 311

Olson, D.M., Dinerstein, E., 1998. Issues in international conservation the global Couvreur, T.L.P., 2019. A third of the tropical African flora is potentially
200 : a representation approach to conserving the earth ’ s most biologically threatened with extinction. Sci. Adv. 5 https://doi.org/10.1126/sciadv.aax9444.
valuable ecoregions. Conserv. Biol. 12, 502e515. Strayer, D.L., Findlay, S.E.G., 2010. Ecology of freshwater shore zones. Aquat. Sci. 72,
Phillips, S., 1995. Poaceae (gramineae). In: Hedberg, I., Edwards, S. (Eds.), Flora of 127e163. https://doi.org/10.1007/s00027-010-0128-9.
Ethiopia and Eritrea, vol. 7. The National Herbarium, Addis Ababa University, Tadesse, M., 2004. Asteraceae (compositae). In: Hedberg, I., Friis, I., Edwards, S.
Addis Ababa & Uppsala. (Eds.), Flora of Ethiopia and Eritrea. Vol 4 (1. The National Herbarium, Addis
Pichi Sermolli, R.E.G., 1957. Una carta geobotanica dell'Africa Orientale (Eritrea, Ababa University, Addis Ababa & Uppsala.
Ethiopia, Somalia). Webbia 13, 15e132. Tewolde, E.B.G., 1991. Diversity of Ethiopian flora. In: M Engels, J.M., Hawkes, J.G.,
Redman, N., Stevenson, T., Fanshawe, J., 2011. Birds of the Horn of Africa: Ethiopia, Worede, Melaku (Eds.), Plant Genetic Resources of Ethiopia. Cambridge Uni-
Eritrea, Djibouti, Somalia and Socotra. versity Press, Cambridge, pp. 75e81.
Roberts, E., Stevens, N., O'Connor, P., Dirks, P., Gottfried, M., Clyde, W., Armstrong, R., Tolessa, T., Senbeta, F., Kidane, M., 2017. The impact of land use/land cover change
Kemp, A., Hemming, S., 2012. Initiation of the western branch of the East Af- on ecosystem services in the central highlands of Ethiopia. Ecosyst. Serv. 23,
rican Rift coeval with the eastern branch. Nat. Geosci. 5, 289e294. 47e54. https://doi.org/10.1016/j.ecoser.2016.11.010.
Sebsebe, D., Cribb, P., Rasmussen, F., 2004. Field Guide to Ethiopian Orchids. Royal White, F., 1983. The vegetation of Africa. A descriptive memoir to accompany the
Botanic Gardens, Kew. UNESCO/AETFAT/UNSO vegetation map of Africa. With 4 coloured vegetation
Senbeta, F., Denich, M., 2006. Effects of wild coffee management on species di- maps (1:5 000 000). Nat. Resour. Res. 20, 1e356.
versity in the montane rainforests of Ethiopia. For. Ecol.Manag. 232, 68e74. Williams, S.D., Vivero, J.L., Spawls, S., Anteneh, S., Ensermu, K., 2004. Ethiopian
Siyum, Z.G., Ayoade, J.O., Onilude, M.A., Feyissa, M.T., 2019. Climate forcing of tree highlands. In: Mittermeier, R.A., Robles-Gil, P., Hoffmann, M., Pilgrim, J.D.,
growth in dry afromontane forest fragments of northern Ethiopia: evidence Brooks, T.M., Mittermeier, C.G., Fonseca, G. (Eds.), Hotspots Revisited: CEMEX,
from multi-species responses. For. Ecosyst. 6 https://doi.org/10.1186/s40663- Mexico City, Mexico. Earth's Biol. Richest Most Endanger. Ecoregions,
019-0178-y. pp. 262e273.
Smith, M.L., Noonan, B.P., Colston, T.J., 2017. The role of climatic and geological Woldemariam, W., Mekonnen, T., Morrison, K., Aticho, A., 2018. Assessment of
events in generating diversity in Ethiopian grass frogs (genus Ptychadena). wetland flora and avifauna species diversity in Kafa Zone, Southwestern
R. Soc. Open Sci. 4, 170021. https://doi.org/10.1098/rsos.170021. Ethiopia. J. Asia Pac. Biodivers. 11, 494e502. https://doi.org/10.1016/
Soromessa, T., Teketay, D., Demissew, S., 2004. Ecological study of the vegetation in j.japb.2018.08.003.
gamo gofa. Trop. Ecol. 45, 209e221. WWF, IUCN, 1994. Centres of Plant Diversity: A Guide and Strategy for Their Con-
Steinbauer, M.J., Dolos, K., Field, R., Reineking, B., Beierkuhnlein, C., 2013. Re-eval- servation, vol. 3. IUCN Publications Unit, Cambridge, U.K.
uating the general dynamic theory of oceanic island biogeography. Front. Bio- Yalden, D.W., Largen, M.J., 2008. Endemic mammals of Ethiopia. Mamm Rev. 22,
geogr. 5, 185e194. 115e150.

vart, T., Dauby, G., Lowry, P., Blach-Overgaard, A., Droissart, V., Harris, D.J.,
Ste
Mackinder, A.B., Schatz, G.E., Sonke
, B., Sosef, M.S.M., Svenning, J.C., Wieringa, J.,