Biome: Montane Grasslands and Shrublands
Size: 28,700 square miles
Conservation Status: Vulnerable
Location and General Description
This ecoregion comprises a long narrow strip of land running from about 6° to 14° S latitude between the Atlantic Ocean, the Southwest Arid biome of Angola and the top of the scarp face of the Central African Plateau. The area is characterized by steep topographic, climatic and vegetation gradients along both the east-west and north-south axes. These gradients have resulted in high biological diversity and endemism, and are of great evolutionary significance. This is illustrated by the fact that the ecoregion lies at the convergence of all six of White’s (1983) phytochoria that are found in Angola. Three of these phytochoria are represented within the ecoregion: the Guineo-Congolian and the Zambezian centers of endemism; and the Guineo-Congolia/Zambezia regional transition zone. Three further phytochoria border on the ecoregion: the Kalahari-Highveld regional transition zone and the Karoo-Namib regional center of endemism abut it in the more arid southwest, while the Afromontane archipelago-like regional center of endemism lies along its southeastern border at the highest elevations. The vegetation is highly varied and ranges from dry woodland and wooded grassland to humid mist forest.
Though narrow, the ecoregion covers an elevational range from sea level to about 1000 meters. It includes two main geomorphologic regions: the Coastal Belt and the Transition Zone (Huntley 1974a). The Coastal Belt varies in width from 200 km in the north to less than 40 km before the ecoregion continues inland, and does not exceed 300 m elevation. The Transition Zone is a discontinuous escarpment belt formed through erosion of the ancient massif, which runs more or less parallel to the coast. It rises sharply in the south, while the increase is more gradual farther north, forming a series of steps (Huntley 1974a). Major rivers draining the ecoregion are the M’Bridge, Loge, Cuanza, and Cuvo.
The ecoregion overlies two main geological divisions: the coastal sediments and the ancient massif (Texeira 1968). The coastal sediments are made up of marine sediments (marls and limestones) and recent sands laid down in different sedimentary basins. Inland these are replaced by gneisses, gneissic granites, and metamorphosed sediments of the Precambrian Basement Complex that forms part of the ancient massif. A great variety of soil types, some very fertile, occur in the ecoregion, and their distribution is determined by geology and climatic factors. Soil types encountered include heavy black cotton soils, red or reddish sandy soils, calcareous patches on the coastal plain and lower escarpment, and the fertile paraferralitic and ferralitic soils that are the substrate of the moist escarpment forests (Texeira 1968, White and Werger 1978).
The ecoregion has a tropical climate with summer rain. Along the coast, the cold Benguela Current strongly influences the climate so that humidity is high year-round while annual rainfall is low, ranging from 400 mm to 800 mm. Offshore, the Benguela Current meets warm equatorial waters and produces mists that are precipitated by the escarpment. A narrow belt on and below the escarpment thus combines the high summer rainfall of the inland areas with the year-round humidity of the coastal plains (Hall 1960). Total precipitation in some of these areas is among the highest in Angola and is thought to exceed 1,600 mm (Huntley 1974a). Temperatures vary within the ecoregion, depending on elevation and latitude. The highest mean annual temperatures are encountered along the inner margin of the Coastal Belt north of the Cueve River, where they exceed 25° C (Huntley 1974a). The lowest mean annual temperatures in the ecoregion occur along the border with the highlands of the Angolan Montane Forest-Grassland Mosaic ecoregion and are approximately 20° C.
The ecoregion is comprised of three main vegetation zones. The first, north of the Cuanza River, is a mosaic of tall, tropical gallery forest and tall grassland, interdigitated by mangrove and swamp communities along the major rivers and their mouths. The second is a discontinuous series of semi-deciduous humid forest patches along the higher slopes of the escarpment. The third, south of the Cuanza River, includes arid to semi-arid undifferentiated woodlands and wooded grasslands on the Coastal Belt and lower slopes of the escarpment. Two azonal vegetation types, mangrove communities at the major river mouths and swamp vegetation in floodplains on the lower reaches of the larger rivers, are also found within the ecoregion.
Human population density in the ecoregion is highly variable, being densest in and around Luanda, the country’s capital city. More than 1.5 million people live in Luanda, almost one-tenth of Angola’s entire population. The city’s population is increasing at a high rate as people move to the larger cities as a result of wartime instability and lack of security (Moyo et al. 1993). The southern parts of the coastal belt are very thinly populated, while the fertile, high-rainfall parts of the escarpment have relatively high population densities.
The escarpment of Angola forms a tension zone among the surrounding biomes. Its evolutionary importance was recognized by Hall (1960) in her study of Angolan birds and supported with mammalogical evidence by Cabral (1966, in Huntley 1974a). Firstly, the escarpment allows subspecies to develop in the drier southwest arid and Brachystegia biomes by forming a barrier between them. Secondly, the escarpment zone with its great range of elevations and relatively high humidity has provided refugia for forest species in periods of climatic desiccation. This has allowed species which remained in these refugia islands to survive climate changes and has facilitated the evolution of new species when their ancestral forms became isolated in these escarpment forests (Hall 1960, Hawkins 1993).
The vegetation in the ecoregion is highly varied with many unique plant communities, which in turn support distinctive faunas. The escarpment forests are important centers of endemism and are listed as critical sites for biodiversity conservation in Angola (Stuart et al. 1990, Stattersfield et al. 1998). Despite being of great biological interest, the vegetation and fauna of these areas have been poorly studied. For example, no plant lists exist for any of the ecoregion’s many vegetation communities (B.J. Huntley, personal communication). Plant diversity, species richness and endemism are high, but no reliable data exist at present. Of the ecoregion’s fauna, only birds have been studied and documented in any detail.
Thirteen bird species are either strictly endemic or near-endemic to this ecoregion, of which seven are threatened according to IUCN (2000). These include the grey-striped francolin (Francolinus griseostriatus VU), red-crested tauraco (Tauraco erythrolophus), Angola helmetshrike (Prionops gabela EN), white-fronted wattle-eye (Platysteira albifrons), Angola slaty-flycatcher (Dioptrornis brunneus), Gabela akalat (Sheppardia gabela EN), Angola cave-chat (Xenocopsychus ansorgei), Pulitzer’s longbill (Macrosphenus pulitzeri EN), golden-backed bishop (Euplectes aureus), orange breasted bush shrike (Laniarius brauni EN), Gabela bush shrike (Laniarius amboimensis EN), and Monteiro’s bushshrike (Malaconotus monteiri DD), the last of which is also found in Cameroon (Hilton-Taylor 2000). Many of these species are included in the restricted range species listed for the Western Angola Endemic Bird Area by Stattersfield et al. (1998), although they included montane forests, which we have separated into another ecoregion.
There are also a number of endemic bird subspecies, the brown-chested alethe (Alethe poliocephala hallae), yellow-necked greenbul (Chlorocichla falkensteini falkensteini), grey-backed camaroptera (Camaroptera brevicaudata harteri) and two subspecies of Lühder’s bushshrike, Laniarius luehderi brauni and L. l. amboimensis. The latter two are considered full species by some taxonomists (Dean 2000).
There are two strictly endemic reptile species, Barbosa’s leaf-toed gecko (Hemidactylus bayonii) and (Monopeltis luandae), and four strictly endemic amphibian species, Cuanza reed frog (Hyperolius punctulatus), Congulu forest tree frog (Leptopelis jordani), Quissange forest tree frog (Leptopelis marginatus), and Congolo frog (Hylarana parkeriana). As with the birds, most of the strict endemics are found in the Angola scarp forests, although some are also restricted to the drier areas in the lowlands.
In the northernmost part of the ecoregion, between the Zaire and Cuanza Rivers, a complex vegetation mosaic is found on highly dissected topography. Large areas of tall grassland are interspersed with gallery forests, which represent the southwestern corner of the Guineo-Congolia/Zambezia regional transition zone. The forest patches are 20 m to 40 m tall and are dominated by tree species such as Piptadeniastrum africanum, Milicia excelsa, Ceiba pentandra, and Musanga cecropioides. The forest fauna is rich, though better represented in the extensive Western Congolian Forest-Savanna Mosaic ecoregion. Mammals include Beecroft’s flying squirrel (Anomulurops beecrofti), giant forest squirrel (Protoxerus stangeri), forest elephant (Loxodonta africana cyclotis), golden potto (Arctocebus aureus), Bosman’s potto (Perodicticus potto), bay duiker (Cephalophus dorsalis), and water chevrotain (Hyemoschus aquaticus). The grasslands are 2 m to 4 m high with few scattered woody plants which are limited to species that tolerate the annual intense grass fires. Examples of these include Hymenocardia acida, Erythrina abyssinica, Piligostigma thonningii, and Cussonia angolensis (Huntley and Matos 1994). Mangrove and swamp vegetation forms large tongues inland from the coast along the larger rivers.
On the upper slopes of the escarpment, where rain and mist provide year-round moisture and the fertile soils favor tree growth, mist (or cloud) forest occurs in a discontinuous band 1 km to 15 km wide (White and Werger 1978). These forest patches total between 1,300 km2 and 2,000 km2 and are most extensive in the Gabela and Amboim areas (Hawkins 1993). They are also known as coffee forests, as they have been used extensively for coffee cultivation. These semi-evergreen humid forests are of Guineo-Congolian affinity and are thought to be impoverished outliers of Congolian rain forest (Hall 1960) with tall macrophyll or mesophyll canopy trees, extensive growth of epiphytes and lianes and many fruiting species (Huntley 1974b, Hawkins 1993). Dominant tree species include Celtis prantlii, Morus mezozygia, Albizia glaberrima, A. gummifera, Ficus mucuso, and F. exasperata, and canopy height rarely exceeds 30 m (Texeira 1968, Huntley and Matos 1994). Two wild coffee species, Coffea canephora and C. welwitschii, are among the understory species.
Among the mammals naturally occurring in the escarpment forest are yellow-backed duiker (Cephalophus sylvicultor), black-fronted duiker (C. nigrifrons), blue duiker (C. monticola), tree pangolin (Phataginus tricuspis), and red buffalo (Syncerus caffer nanus) (Huntley 1974a). Grazing antelopes are largely absent. Besides the narrow endemics listed above, characteristic birds of the escarpment forests and woodlands include red-crested turaco (Tauraco erythrolophus), red-backed mousebird (Colius castanotus) (both endemic to Angola), mottled spinetail (Telacanthura ussheri), batlike spinetail (Neafrapus boehmi), naked-faced barbet (Gymnobucco calvus), red-tailed palm thrush (Cichladusa ruficauda), and yellow-bellied wattle-eye (Dyaphorophyia concreta) (Dean 2000).
South of the Cuanza River, the Zambezian component of the ecoregion comprises a mosaic of closed woodlands, grasslands and palm savanna, which is found along the lower and drier slopes of the escarpment and along the coast (Huntley and Matos 1994). The distribution of the different vegetation communities within this zone is strongly influenced by soil type (Werger and Coetzee 1978). The woodlands are floristically rich and are more easily defined by the absence of mopane and miombo dominants than by their own floristic composition. Despite their small area, they are composed of many more tree species than either miombo or mopane (White 1983) and include several endemic species (White and Werger 1978). Dense dry thicket and scrub forest are the typical woodland physiognomies, with Sterculia setigera, Euphorbia conspicua, Strychnos spp., Acacia welwitschii, and baobab (Adansonia digitata) being the dominant woody species. Between the closed woodlands, there are grasslands dominated by Andropogon gayanus, Heteropogon contortus, and Hyparrhenia spp. Extensive sand plateaus nearer the coast are occupied by Hyphaene gossweileri palm savanna, where the grass layer is dominated by Eragrostis superba, Schizachyrium semiberbe, and Digitaria milanjiana. To the north and south of Luanda, a remarkable grassland occupies large areas on smoothly rounded hills of marine sediments with heavy black and dark brown clays. This is dominated, to the virtual exclusion of all other species, by Setaria welwitschii, which forms dense swards 1 to 1.5 m high (Texeira 1968, Barbosa 1970, Huntley and Matos 1994).
The large mammal fauna in this area includes roan antelope (Hippotragus equinus), red buffalo (Syncerus caffer nanus), elephant (Loxodonta africana), reedbuck (Redunca arundinum), bushbuck (Tragelaphus scriptus) and eland (Taurotragus oryx), which are represented in the Kisama National Park (Huntley 1974a). Large predators used to occur in the coastal plains but their populations, like those of all large mammals, have been severely reduced if not entirely eliminated as a result of uncontrolled hunting. Many of the bird species that occur in the escarpment forests are found in the woodlands here. The white-fronted wattle-eye (Platysteira albifrons) is endemic to the northernmost part of the coastal belt in this ecoregion. Large populations of the woolly-necked stork (Ciconia episcopus) occur on grassy plains and gallery forest along rivers.
The Angolan Coast, the northern half of which is part of the ecoregion, includes several specialized maritime and estuarine ecosystems which do not form part of any of the biogeographic divisions outlined above, but which are of great biological interest. These communities are described in the Central African Mangroves ecoregion.
Due to its low agricultural potential, the dry coastal belt and lower escarpment are relatively sparsely settled, and large areas remain unfragmented. Huntley and Matos (1994) report large stretches of completely undisturbed habitat, for instance, in Kisama National Park. Around the larger urban centers, particularly Luanda, human settlement and activities such as woodcutting and livestock grazing have had considerable, though mostly localized, impacts on the vegetation and soils.
The escarpment forests were almost entirely utilized for coffee production that peaked between 1950 and 1970, when an estimated 95 percent of the forest area was affected (Hawkins 1993). This involved clearing the understory while leaving canopy trees for shade. Coffee-berry disease in the 1950’s, a drop in coffee prices in the mid-1970’s, and the upheaval of the civil war since 1974 have resulted in the abandonment of many of the coffee plantations, with the result that much of the forest has recovered today (Hawkins 1993, Huntley and Matos 1994). However, subsistence cultivation on the fertile forest soils is increasing and Hawkins (1993) estimated that some 30 percent of the forest areas were affected, especially around Gabela where many of the ridges and valley bottoms have been cleared.
Two areas in the ecoregion are protected, with three more areas proposed for protection but not yet established (Huntley 1974b, Huntley and Matos 1994). The large Kisama National Park is bordered by the Atlantic Coast and the banks of the Cuanza and Longa Rivers and is listed by Stuart et al. (1990) among critical sites for biodiversity conservation. Marine, estuarine, floodplain, grassland and thicket habitats are represented here (Texeira 1968). Ilheu dos Passaros Integral Nature Reserve is a small offshore island with mangrove communities and mud flats which are of great botanical interest and provide key breeding habitats for water birds.
The proposed Gabela and Chingoroi Strict Nature Reserves, both representing patches of escarpment forest, are needed to protect the area’s rare endemic bird species, because their small forest habitats are declining due to agricultural activities. Without these two protected areas, the escarpment forest vegetation and its fauna are unprotected despite their vulnerability and great biological interest. The Pungo Andongo Natural Monument comprises a series of large rocky outcrops between Gabela and the coast, which should be protected for their biological and aesthetic value (Huntley 1974b).
The almost continuous civil war in Angola since 1974 has led to great instability, poor security, economic depression, massive displacement of the rural population and a lack of infrastructure and basic services. Its effects on conservation, particularly that of large mammals, have been devastating. Most of Angola’s protected areas have been abandoned as their wardens were forced to leave for economic or security reasons and have become open areas for poachers and settlers (Huntley and Matos 1994).
Types and Severity of Threats
The most immediate and important threat to the ecoregion’s biodiversity is the encroachment of subsistence agriculture in the fertile escarpment forest areas (Stuart et al. 1990). These are, at present, not even nominally protected. This threat is expected to escalate once stability returns to Angola and displaced rural people return to farming areas (Hawkins 1993). It is also possible that coffee plantations will be re-established in the escarpment forests.
Hunting is virtually uncontrolled in most of Angola, including the protected areas. Huntley and Matos (1992) estimated that 21 species of larger mammals are close to extinction in Angola, including lion, cheetah, West African manatee and forest elephant. There are no data on the extent and impact of subsistence hunting on populations of smaller mammals and birds, but these species may be an important source of protein in the more populated rural areas (Hawkins 1993). Marine turtles and their eggs are also harvested along the coast.
It seems that commercial exploitation of timber has not taken place so far, probably due to the importance of canopy species for coffee production. However, the network of wide tracks established for coffee plantations means that it would be very easy to start timber exploitation, especially if coffee production is not resumed on the previous scale (Hawkins 1993).
Justification of Ecoregion Delineation
This ecoregion is primarily based on Birdlife’s Western Angola Endemic Bird Area (Stattersfield et al. 1998). It encompasses portions of White’s (1983) ‘North Zambezian undifferentiated woodland’ and ‘mosaic of lowland forest and secondary grassland,’ extending inland 1 km to 15 km along the Angola escarpment, which forms its eastern border. Although included in the Western Angola EBA, the Bailundu Highlands have been separated into the Angolan Montane Forest-Grassland Mosaic ecoregion since the characteristic elements of the ecoregion’s fauna and flora are more closely related to other Afromontane areas than to the surrounding Angolan biomes (Huntley 1974a, Dean 2000).
Barbosa, L.A.G. 1970. Carta Fitogeografica de Angola. IICA, Luanda.
Dean, W.R.J. 2000. The birds of Angola: An annotated checklist. BOU Checklist No. 18. British Ornithologists’ Union, Herts, U.K.
Hall, B.P. 1960. The faunistic importance of the scarp of Angola. Ibis 102: 420-442
Hawkins, F. 1993. An integrated biodiversity conservation project under development: The ICBP Angola Scarp Project. Proceedings of the VIII Pan-African Ornithological Congress: 279-284
Hilton-Taylor, C. 2000. The IUCN red list of threatened species. Gland, Switzerland and Cambridge, U.K.
Huntley, B.J. 1974a. Outlines of wildlife conservation in Angola. Journal of the southern African Wildlife Management Association 4: 157-166.
Huntley, B.J. 1974b. Ecosystem conservation priorities in Angola. Ecologist’s Report No. 28. Servicos de Veterinaria, Luanda. 21 pp.
Huntley, B.J., and E.M. Matos. 1994. Botanical diversity and its conservation in Angola. B.J.Huntley, editor. Botanical Diversity in Southern Africa. Strelitzia 1: 53-74. National Botanical Institute, Pretoria.
Huntley, B.J., and G.C. Matos. 1992. Biodiversity: Angolan environmental status quo assessment report. IUCN Regional Office for Southern Africa, Harare.
Moyo, S., P. O’Keefe, and M. Sill. 1993. The southern African environment: Profiles of the SADC countries. Earthscan, London.
Stattersfield, A.J., M.J. Crosby, A.J. Long, and D.C. Wege. 1998. Endemic bird areas of the world: priorities for conservation. BirdLife International, Cambridge, UK.
Stuart, S.N., and R.J. Adams. 1990. Biodiversity in sub-Saharan Africa and its islands. Occasional Papers of the IUCN Species Survival Commission No. 6.
Texeira, J.B. 1968. Angola. I. Hedberg, and O. Hedberg, editors. Conservation of vegetation in Africa south of the Sahara. Acta Phytogeographica Suecica 54: 193-197.
Werger, M.J.A., and B.J. Coetzee. 1978. The Sudano-Zambezian Region. M.J.A. Werger, editor. Biogeography and Ecology of Southern Africa. W. Junk, The Hague.
White, F. 1983. The vegetation of Africa. A descriptive memoir to accompany the UNESCO/AETFAT/UNSO Vegetation Map of Africa (3 Plates, Northwestern Africa, Northeastern Africa, and Southern Africa, 1:5,000,000). UNESCO, Paris.
White, F., and M.J.A. Werger. 1978. The Guineo-Congolian transition to southern Africa. M.J.A. Werger, editor. Biogeography and Ecology of Southern Africa. W. Junk, The Hague.
Prepared by: Suzanne Vetter
Reviewed by: In progress