Location and General Description
The Southern Rift Montane Forest-Grassland Mosaic consists of a series of mountains and plateaus centered on the eastern and northern shores of Lake Malawi/Nyasa. Altitude and climate are the main factors separating this ecoregion from the surrounding miombo ecoregions (Central Zambezian and Eastern Woodlands). These woodlands occupy the plains and lower-lying areas surrounding the Rift Mountains and are characterised by the dominance of trees in the genera Brachystegia, Isoberlinia, and Julbernardia (White 1983). The major differences between this ecoregion and the adjacent and superficially similar Eastern Arc Forest ecoregion in Tanzania and Kenya relate to the differing climate regimes. The highland areas of the Southern Rift ecoregion receive rainfall from the surface convection off Lake Malawi/Nyasa, while precipitation in the Eastern Arc Mountains is derived from the Indian Ocean (Lovett 1993). This factor may have been responsible for the Eastern Arc mountain forests remaining stable during the Pleistocene droughts, during which time, evergreen forests were drastically reduced elsewhere in East Africa, resulting in floristic impoverishment (Wasser and Lovett 1993). The Eastern Arc Mountains thus have a much higher degree of both generic and specific endemism than the Southern Rift montane areas (Lovett 1993). The Southern Rift Montane ecoregion is similarly separated from the Albertine Rift Mountains ecoregion to the northwest on the basis of endemicity and overall biodiversity, as the latter is richer in each category. Finally, this ecoregion is distinguishable from montane areas to the south because many northern plant taxa reach their southernmost distributions within the ecoregion, while Mount Mulanje in the South Malawi Montane Forest-Grassland Mosaic forms the northern limit for many species belonging to the southern flora (Williamson 1979; Chapman and White 1970, Kerfoot 1963-64b).
The Southern Rift Montane Forest-Grassland Mosaic consists of several discontinuous mountain chains and plateaus that are centered on the Southern Highlands of Tanzania, where the East and West Rift escarpments meet. This ecoregion’s westernmost section lies on the Ufipa Plateau between Lakes Tanganyika and Rukwa, while the extensive north-central and north-eastern sections (collectively referred to as the Southern Highlands) are made up of several different ranges, including the Mbeya and Kipengere Ranges, the Umalila Highlands, the Poroto and Livingstone Mountains, and the Kitulo Plateau. The northernmost portion of the ecoregion in Malawi is found in the Misuku Hills, while the southernmost point of the ecoregion both within Malawi and Mozambique and overall lies in the Kirk Range. Between these two points are the extensive Nyika and North and South Viphya Plateaux, and a number of smaller mountains. Portions of the ecoregion within Malawi, including the Nyika Plateau and Kirk Range, overlap into either Zambia or Mozambique.
The general climate pattern around the Southern Rift Mountains is dictated by Lake Nyasa, from which winds carry moisture into the highlands (Lovett 1993; Cribb and Leedal 1982; Chapman and White 1970). Average annual rainfall may be as low as 823 mm on the Ufipa Plateau in the northwest of the ecoregion, to as high as 2,850 mm in the wet Livingstone and Poroto mountains at the head of Lake Nyasa (Cribb and Leedal 1982) with a mean rainfall of approximately 1500 mm for the entire ecoregion (Cribb and Leedal 1982; Dowsett-Lemaire 1989; Chapman and White 1970). Precipitation is largely confined to the wet season months of November through April (Cribb and Leedal 1982; Dowsett-Lemaire 1989), although light rains or mist may occur at the higher altitudes during the dry season months of May through August (Kerfoot 1963-64a; Dowsett-Lemaire 1989). Mean annual temperatures in the ecoregion fall between 13?C to 19?C (Dowsett-Lemaire 1989), with an average maximum of 22?C and an average minimum of 9.8?C (McKone and Walzem 1994). At the highest altitudes, temperatures as low as –5?C have been recorded, and frosts are common, with widespread, destructive frosts recurring every 25 years in the Southern Highlands of Tanzania (Cribb and Leedal 1982).
The Southern Rift montane areas are composed primarily of Pre-Cambrian granites, gneisses, and schists formed 3,250 million years ago (Cribb and Leedal 1982; Kerfoot 1963-64a; Chapman and White 1970). Early uplifting and erosion of these original formations caused the addition of sedimentary rock, and additional granites were subsequently intruded in places (Cribb and Leedal 1982). The Rift Valley faulting began during the late Cretaceous, creating the mountains and plateaux of this ecoregion adjacent to the great Rift Valleys, which are now inundated by Lakes Nyasa, Tanganyika, and Rukwa (Cribb and Leedal 1982). During this time, localized volcanism deposited lavas and ash, chiefly in the Rungwe area north of Lake Nyasa (Cribb and Leedal 1982). The ecoregion’s soils are primarily well-drained andosols and ferrisols (McKone and Walzem 1994; Chapman and White 1970), and its topography is characterised by large plateaux surrounding high peaks and ridges, bounded on all sides by escarpments or deeply dissected hill country (Chapman and White 1970; Cribb and Leedal 1982). Plateau altitudes range from 1,400 to 2,400 m, with the two highest peaks in Tanzania (Rungwe and Mtorwi) attaining 2,960 m.
The Southern Rift mountains and plateaus are important watersheds for Lake Tanganyika via the Kalambo River, Lake Rukwa via the Kafufu, Mtembwa, and Rukwa Songwe Rivers, the Ruaha River via most north-flowing rivers, and Lake Nyasa via the Kiwira, Mbaka, Lufirio, Lumbila, Ruhuhu, Livulezi, Ketewaka and Nyasa Songwe Rivers.
The Southern Rift Montane Forest Grassland-Mosaic ecoregion is composed of several structurally and compositionally distinct vegetation communities, the most dominant of which is grassland (White 1983). The preponderance of this community is commonly attributed to the high frequency and extent of range fires, which have swept the ecoregion’s grasslands for centuries, continuously eroding the margins of the once abundant Afromontane forests (White 1983; Dowsett-Lemaire 1989). Dominant grass species in this vegetation type are Loudetia simplex, Exotheca abyssinica, Monocymbium ceresiiforme, Themeda triandra, Andropogon spp., Pennisetum spp., Setaria spp. (White 1983; Chapman and White 1970; Cribb and Leedal 1982). A number of herbs, sedges, and geophytes also occur within the grassland community (Kerfoot 1963-64a), as well as the occasional fire-resistant shrub, usually of the genus Protea (White 1983). In areas of impeded drainage, permanent and seasonal bogs known as dambos may be found. These habitats, dominated by grasses and sedges, contain a remarkable abundance of species relative to their area, and are known for having rich orchid floras (Cribb and Leedal 1982; Kerfoot 1963-64a).
Several other vegetation types are set within the grassland matrix, the most prominent of which is Afromontane forest, although this constitutes less than 5 percent of the landscape, and is confined to fire-sheltered pockets, moist escarpments, valleys and watercourses (Chapman and White 1970; Kerfoot 1963-64a, Dowsett-Lemaire 1989). Afromontane forests can vary considerably in structure and composition, depending on available moisture, altitude, and disturbance regime (White 1983). The most favourable circumstances produce tall, triple-canopied rain forest, in which the trees are festooned with epiphytes and lianes, while the patches found at higher altitudes and under poorer conditions are shorter and less developed in structure, and compositionally less diverse (White 1983; Dowsett-Lemaire 1970). Dominant tree and shrub species are Pouteria adolphi-friedericii, Apodytes dimidiata, Bersama abyssinica, Chrysophyllum gorungosanum, Cola greenwayii, Cylicomorpha parvifolia, Entandophragma excelsum, Ficalhoa laurifolia, Garcinia spp., Ilex mitis, Kiggelaria africana, Myrianthus holstii, Ocotea usambarensis, Parinari excelsa, Podocarpus latifolius, Polyscias fulva, Rapanea melanophloeos, and Syzygium guineense (Chapman and White 1970; Dowsett-Lemaire 1989; Kerfoot 1963-64a, White 1983). The most common constituents of the herbaceous layer are species of the family Acanthaceae, and of the genera Impatiens, Begonia, Streptocarpus, Plectranthus and Peperomia (White 1983; Dowsett-Lemaire 1989; Kerfoot 1963-64a). A fringe of smaller trees and shrubs, usually pioneer species, often surrounds forest patches, the width of which varies according to its exposure to fire (Chapman and White 1970; Kerfoot 1963-64a). After some form of disturbance Afromontane forests are frequently found in states in which a single tree species dominates (White 1983; Kerfoot 1963-64a). Hagenia abyssinica and Juniperus procera are commonly associated with this condition (White 1983), although the latter is now rare (Kerfoot 1963-64b). Communities of the bamboo Arundinaria alpina are also found on slopes with high rainfall, primarily in the northern portion of the ecoregion, in stands of varying height and density (White 1983; Cribb and Leedal 1982). Certain forest tree species such as Hagenia abyssinica, Juniperus procera, Podocarpus latifolius, Prunus africana, and Rapanea melanophloeos may be associated with bamboo (White 1983). Bamboo patches are sometimes found within Afromontane forest (Chapman and White 1970), perhaps as a result of post-fire invasion (White 1983).
The third distinguishing Afromontane vegetation type found within the ecoregion can be termed the "Ericaceous" zone, originally identified by Hedberg (1951). White (1983) divides this zone into two types, based on height, occasional variations in species dominance, and substrate. These communities, usually dominated by sclerophyllous shrubs of the family Ericaceae, occur on the higher mountains above the forest zone in a range of moisture and substrate conditions, although examples may be found at lower elevations in areas of shallow soil and frequent mists (White 1983). Density of the shrub component varies according to environmental factors, and height ranges from 3 to 13 m (White 1983).
Lastly, the lower reaches of the Southern Rift montane areas—the escarpments and broken hill country—are often occupied by vegetation associated with the miombo ecoregions. Most of the land between 1,200 to 1,800 m is clothed in woodlands dominated by Brachystegia, Julbernardia, and Isoberlinia (Cribb and Leedal 1982), which ascend as high as 2,050 m on the xeric western escarpment of the Nyika Plateau (Dowsett-Lemaire 1989). These woodlands may comprise a significant part of the ecoregion, as they constitute more than half of the Nyika Plateau National Park (Johnson undated).
The Southern Rift Montane Ecoregion is both poorer in species richness and endemicity than the Eastern Arc Forest and Albertine Rift Montane Forest ecoregions to the north. For instance, the Eastern Arc’s moist mountain forests alone contain 67 endemic trees, while all of Malawi’s evergreen forests contain only 10 unique species (Dowsett-Lemaire 1989). One possible factor contributing to this ecoregion’s relative poverty may be its greater distance from the Equator. An inverse relationship between distance from the equator and species diversity is noticeable in Malawi’s Afromontane woody flora, where 232 species are recorded from northern Malawian forests, while southern forests have only 149 species (Dowsett-Lemaire 1989). The forests of the Misuku Hills, the most northern portion of the ecoregion within Malawi, are the most diverse (Dowsett-Lemaire 1989).
However, the ecoregion’s biota is by no means impoverished. The Kitulo Plateau in the Southern Highlands of Tanzania is considered botanically rich, boasting 350 plant species and several endemics (Lovett and Prins 1994). The highlands of Malawi are that country’s biologically wealthiest habitat type, particularly the Nyika Plateau (Stuart et al. 1990), which has never been permanently settled (Chapman and White 1970) and is now well protected by Malawi’s 3134 km2 Nyika National Park and the 80 km2 Zambian Nyika National Park (Carter 1987; Stuart et al. 1990). The Nyika Plateau is undoubtedly one of the most significant areas of the ecoregion, as it is home to south-central Africa’s richest orchid flora, totalling 214 species (Kurzweil 2000), as well as 400 bird species and important populations of reedbuck (Redunca arundinum), roan antelope (Hippotragus equinus), zebra (Equus burchelli) and eland (Tragelaphus oryx) (Carter 1987; Johnson undated). Lion (Panthera leo), elephant (Loxodonta africana), and buffalo (Syncerus caffer) inhabit the lower woodland areas of the park (Carter 1987; Johnson undated). Significant populations of large mammals are unlikely to be found elsewhere in the ecoregion. No large-scale migrations within the Southern Rift montane areas, with the exception of the seasonal migrations of zebra, roan, and eland between the Nyika Plateau and surrounding low-lying woodlands (Johnson undated). Oribi (Ourebia oribi) are also found within the ecoregion. The top predator species include leopard (Panthera pardus), cheetah (Acinonyx jubatus), serval (Felis serval), and spotted hyena (Crocuta crocuta).
A number of restricted range species are found in this ecoregion, including 5 mammals, 18 birds, 6 amphibians, and 20 reptiles. The five near-endemic mammals include one small primate, the Mozambique galago (Galagoides granti), an insectivore, Crocidura desperata (CR), and three small rodents, Beamys major, Tanganyika mountain squirrel (Paraxerus lucifer), and Swynnerton’s squirrel (Paraxerus vexillarius, VU). Among the 18 birds considered endemic or near-endemic, only one is entirely restricted to this ecoregion, the Namuli apalis (Apalis lynesi, VU). Other near-endemic species include the churring cisticola (Cisticola njombe), the buff-shouldered widowbird (Euplectes psammocromius), Chapin’s apalis (Apalis chapini), black-lored cisticola (Cisticola nigriloris), olive-flanked robin-chat (Cossypha anomala), Fuelleborn’s boubou (Laniarius fuelleborni), uhehe fiscal (Lanius marwitzi), Iringa ground robin (Sheppardia lowei, VU), and the long-billed tailorbird (Orthotomus moreaui, CR) (Hilton-Taylor 2000).
Among the herpetofauna, there a number of strict endemics: Stewart’s river frog (Phrynobatrachus stewartae), Ngosi volcano chameleon (Chameleo fuelleborni), Poroto mountain chameleon (C. incornutus), Cordylus nyikae, Ukinga spinytail lizard (C. ukingensis), Eumecia johnstoni, Whyte’s water snake (Lycodonomorphus whytii), Braun’s mabuya (Mabuya brauni), and the Southern African stumptail chameleon (Rhampholeon nchisiensis).
Besides these species, there are numerous examples of plants unique to the ecoregion. Six protea species are limited to the ecoregion (Beard 1992). Four orchid species and 2 sub-species are unique to the Nyika Plateau, and 4 more species are near-endemics, as they are also found on several of the highland areas adjacent to the Nyika Plateau (Kurzweil 2000). Besides these, the Nyika Plateau also holds 13 other endemic plant species, and 7 sub-species (Willis et al. 2000). Tanzania’s Kitulo Plateau hosts 3 strictly endemic plants, plus 15 others restricted to the Kitulo Plateau and other Southern Highlands mountain ranges (Lovett and Prins 1994). Examples of endemic invertebrates are the dragonfly Teinobasis malawiensis, which is known only from montane streams in northern Malawi (Stuart et al. 1990), the Kitulo-endemic satyrid butterfly Neocoenyra petersi (Kielland 1990), and a further three butterflies confined to the Nyika Plateau (Carter 1987). The ecoregion also contains a number of sub-specific endemics, particularly on the Nyika Plateau, where the red-winged francolin (Francolinus levaillanti crawshayi), the greater double-collared sunbird (Nectarina afra whytei), the Baglafecht weaver (Ploceus baglafecht nyikae), the rufous-naped lark (Mirafra africana nyikae), the chequered elephant-shrew (Rynchocyon cirnei hendersoni), 3 amphibian sub-species, two sub-species of skink (Stuart et al.1990; Johnson undated), and four sub-species of butterfly (Carter 1987) are found.
A number of species of special concern occur in the ecoregion. The declining wattled crane (Bugeranus carunculatus) has an important breeding site in the Nyika National Park, and is also found in the Southern Highlands of Tanzania and on the Ufipa Plateau (Stuart et al. 1990). The Nyika National Park also hosts the locally rare Denham’s bustard (Neotis denhami)(Johnson undated), the world’s largest breeding population of blue swallows (Hirundo atrocaerulea, VU), numbering 5000 pairs, and one of Malawi’s two important populations of cheetah (Stuart et al. 1990). The southernmost populations of the globally threatened Abbott’s duiker (Cephalophus spadix, VU) occur in the Southern Highlands of Tanzania (East 1998; Stuart et al. 1990), where the threatened Iringa ground robin (Sheppardia lowei, VU) and 7 amphibian species of conservation concern are also found (Stuart et al. 1990) (Hilton-Taylor 2000).
The archipelago-like nature of the Southern Rift Highlands means that its component habitat blocks are naturally isolated from one another by topography. However, human intervention has caused further fragmentation and degradation within these habitat islands. Cultivation has been widespread throughout the ecoregion, and is rapidly increasing in places (Cribb and Leedal 1982; McKone 1995). Most of the Mbeya range below 2200 m has been subjected to shifting cultivation (Kerfoot 1963-64a), and the montane grasslands of southern Tanzania are considered to be in a state of rapid decline (Stuart et al. 1990). In the Rungwe District of the Mbeya Region, remaining natural vegetation is largely confined to government proclaimed and traditional forest reserves, but even these conserved areas are subject to anthropogenic disturbances (McKone and Walzem 1994; McKone 1995). An analysis of satellite images spanning twenty years up to 1989 has shown that at least 24 percent of the Kitulo Plateau has been transformed by cultivation and pasture improvement (Lovett and Prins 1994). The majority of grassland on Malawi’s second largest plateau, the South Viphya, has been planted over with exotic Pinus spp. (Dowsett-Lemaire 1989), and other areas of the ecoregion have been similarly afforested (Dowsett-Lemaire 1989; McKone and Walzem 1994). In addition to the destruction caused by frequent wildfires, forests and woodlands have been further reduced by charcoal production, fuelwood harvesting, and cultivation (Cribb and Leedal 1982; Dowsett-Lemaire 1989; McKone and Walzem 1994). In Malawi this deforestation is particularly pronounced, as all that remains of the once extensive mid-altitude montane forests are small relictual groves used as graveyards by local people (Dowsett-Lemaire 1989).
The protected area network throughout most of the ecoregion is woefully inadequate with the exception of the Nyika Plateau area, the majority of which is protected inside Zambia and Malawi’s contiguous Nyika National Parks. None of the other portions of the ecoregion are protected in Zambia, and Mozambique applies no known conservation measures to its share of the ecoregion (Stuart et al. 1990). Part of Chipata mountain is protected in Malawi’s Nkhotakhota Game Reserve (Carter 1987), while Chirobwe mountain in the Dedza-Chirobwe Highlands has a forest reserve, although this is under pressure from wood collectors (Dowsett-Lemaire 1989). The grasslands of the Southern Highlands of Tanzania, including the botanically important Kitulo Plateau, are almost entirely unprotected, as are those of the Ufipa Plateau (Stuart et al. 1990; Lovett and Prins 1994). The Mbeya Region of Tanzania’s Southern Highlands contains 28 forest reserves of 1,350 km2 (although several of these fall outside of the ecoregion’s boundaries), but these have low levels of management and are often subject to illegal pitsawing, fuelwood collection, grazing, hunting, and uncontrolled burning. Many of these reserves are completely surrounded by and somewhat encroached upon by cultivation (McKone and Walzem 1994). Besides these official forest reserves, there are numerous smaller traditional forest reserves in the Southern Highlands, established by local communities for a variety of cultural reasons (McKone 1995). At least 94 are known from the Rungwe district (McKone 1995). Although many of these reserves are no longer maintained, and are under increasing pressure from cultivators, evidence suggests that they could play a valuable conservation role if appropriately supported by government (McKone 1995).
Types and Severity of Threats
The most significant force of natural disturbance within this ecoregion is undoubtedly fire. Each year, large areas of the Southern Rift Highlands are swept by fires of primarily anthropogenic origin (Chapman and White 1970; Cribb and Leedal 1982; Lovett and Prins 1994; Kerfoot 1963-64a). The intensity of this burning regime, which has been practised for centuries, if not millenia, is believed to have been the main cause of the replacement of previously extensive areas of Afromontane forests with grassland and scrub-grassland (Dowsett-Lemaire 1989; Kerfoot 1963-64a; Chapman and White 1970). Although it is unknown exactly how long fire has been a driving force in the ecoregion, and to what extent it has caused grassland to replace forest (Chapman and White, 1970), fire has definitely reduced forests within the last century, and continues to do so (Dowsett-Lemaire 1989; Chapman and White 1970, Kerfoot 1963-64a; McKone and Walzem 1994). Of special concern are the few surviving pockets of the historically widespread Juniperus procera forests, particularly the patch on Nyika Plateau, as these have been reduced not only by fire but also by recent indiscriminate felling for charcoal production and other uses (Kerfoot 1963-64b; Chapman and White 1970).
Cultivation poses the other severe threat to the integrity of the ecoregion, as land is increasingly being converted to crops such as tea, coffee, banana, finger millet, potatoes, and pyrethrum (Chapman and White 1970; Lovett and Prins 1994). Although shifting cultivation is practised in places (Kerfoot 1963-64a; Chapman and White 1970), allowing secondary succession to take place on previously worked fields, fallow lands are generally recolonized by widely distributed species which may preclude the regeneration of the unique elements of Afromontane primary flora (Dowsett-Lemaire 1989; Lovett 1993). In soil-disturbed areas of the Kitulo Plateau, grassland tends to be replaced by a "shrubby sward" (Lovett and Prins 1994). Cultivation does not only change the composition of the ecoregion’s flora, but it can create serious erosion problems, as fields are often plowed on steep slopes. Overgrazing by large numbers of livestock also cause erosion problems in areas of high human population, such as parts of Malawi’s Kirk Range and Tanzania’s Mbeya and Rungwe districts, where densities reach 134 people/km2 (Dowsett-Lemaire 1989; McKone and Walzem 1994). The continuing destruction of Afromontane grassland habitat threatens the locally occurring churring cisticola (Cisticola njombe) and the blue swallow (Stuart et al. 1990).
Alien organisms pose a threat to the ecoregion, chiefly in the form of exotic timber trees of the genera Pinus and Eucalyptus, which have been used in afforesting montane grasslands (McKone and Walzem 1994). Pinus is reported as being invasive in parts of the Southern Highlands (McKone and Walzem 1994). The bramble Rubus sp. has spread extensively throughout the Nyika National Park, where exotic rainbow trout were also introduced into dams and streams on the Plateau (Johnson undated).
Justification of Ecoregion Delineation
The linework for this ecoregion follows the ‘undifferentiated montane vegetation’ unit within the Afromontane archipelago-like regional center of endemism (White 1983), with one refinement of linework for the montane area to the northwest. Although White’s Afromontane areas are not distinguished into finer units, the montane areas of the Kipengere Range and the high elevation areas to the west of Lake Malawi share biological similarities (WWF 1998). These montane areas form part of the Tanzania-Malawi Endemic Bird Area (Stattersfield et al. 1998). However, the Eastern Arc forests, Albertine Rift forests, and Mount Mulanje are distinguished as separate ecoregions due to high levels of local endemism.
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Prepared by: Lyndon D. Estes
Reviewed by: In progress