Location and General Description
Montane Fynbos and Renosterveld, encompassing an area of 45,802 km2, comprises the upland and relatively untransformed portion of the world-renowned Cape Floristic Region (CFR) located in southwestern South Africa. Fynbos comprises 29,475 km2 (81.5 percent) of this ecoregion and renosterveld comprises 6,684 km2 (18.5 percent). It is largely contained in South Africa’s Western Cape Province, although a small portion extends to the Eastern Cape Province. This ecoregion is located in the mountains and uplands of the CFR, which are distributed throughout the region; some mountain ranges running inland, and others lining the coast.
Most of Montane Fynbos and Renosterveld ecoregion receives annual rainfall between 300 mm and 2,000 mm, although some sites in the southwest receive as much as 3,000 mm (Deacon et al. 1992). West of Cape Agulhas, rainfall is concentrated in the winter months, associated with cold fronts budding from the circumpolar westerly system. East of this zone, rainfall distribution is less seasonal. Post-frontal events, chiefly the advection of moist air across a relatively warm Indian Ocean, produce rain throughout the year, largely in the spring and autumn. Temperature lows are more extreme than on the adjacent lowlands. Frost is widespread on upper peaks where snow may lie for several weeks in the winter months. High summer temperatures seldom exceed 25oC, except in the interior valleys.
The Montane Fynbos and Renosterveld ecoregion comprises four major physiographical regions.
•The Western Mountains extend for 375 km from the Cape Peninsula in the south to the Bokkeveld Mountains in the north. Starting in the south, principal mountain complexes and maximum elevations are: Cape Peninsula (1,086 m in elevation), Kogelberg (1,269 m), Hottentots Holland (1,590 m), Du Toits (1,995 m), Slanghoek (1,694 m), Hex River (2,249 m), Witsenberg (1,630 m), Groot Winterhoek (1,884 m), Skurweberg (2,071 m), Koue Bokkeveld (1,800 m), Olifants River (1,015 m), Cederberg (2,026 m), Gifberg (1,016 m), and Bokkeveld (982 m). Rainfall arrives exclusively in the winter months and shows great variation, ranging from as little as 200 mm along the Succulent Karoo ecoregion border in the northeast, to as much as 3,000 mm on the upper slopes facing the coast in the south.
•The South Coastal Mountains extend from the Elgin Basin in the west, to near Port Elizabeth in the east, an unbroken chain of some 800 km. Principal mountain complexes and maximum altitudes, starting in the west, are as follows: Groenland (1,201 m), Riviersonderend (1,654 m), Klein River (964 m), Langeberg (1,710 m), Outeniqua (1,579 m), Tsitsikamma (1,675 m), Kouga (1,757 m), Groot Winterhoek (1,758 m), and Elandsberg (987 m). In the extreme west, most rain falls in the winter months, while elsewhere, rain falls year-round. Annual rainfall is mostly in the vicinity of 800 mm to 1,500 mm, although less than 400 mm has been recorded along the inland foothills, fringing the Little Karoo.
•The Interior Mountains are located to the north of the Little Karoo Basin, and run from Laingsburg in the west to the vicinity of Willowmore in the east, a distance of some 400 km. Starting in the west, principal mountain complexes and maximum altitudes are: Witteberg (1,504 m), Klein Swartberg (2,325 m), Groot Swartberg (2,085 m), and Baviaanskloof (1,626 m). Annual rainfall averages about 500 mm to 750 mm, although interior foothills may receive as little as 200 mm and upper slopes as much as 1,000 mm. The extreme west forms part of the winter-rainfall zone while in the extreme east, rainfall is bimodal with spring and autumn peaks. Most of the area receives rain throughout the year.
•The Little Karoo Inselbergs comprise several isolated, fynbos-clad ranges within the Little Karoo, a basin wedged between the South Coastal and Interior Mountains. Principal mountain complexes and maximum altitudes are, from west to east, Anysberg (1,622 m), Warmwaterberg (1,350 m), Touwsberg (1,491 m), Rooiberg (1,490 m), Gamkaberg (1,100 m), Kamanassieberg (1,955 m), and Antoniesberg (1,720 m). Annual rainfall averages about 400 mm to 750 mm, although the northern foothills may receive as little as 200 mm. Rain may fall at any time of the year, except in the extreme east where the distribution is bimodal.
The Cape Floristic Region overlies the Cape Supergroup, a Devonian-Ordovician series of sedimentary strata, where alternating quartzitic sandstones such as Table Mountain and the Witteberg Groups alternate with fine-grained shales, as seen in the Bokkeveld Group (Deacon et al. 1992). These sediments accumulated in a basin depository with one billion year-old metasediments of the Malmesbury, Kango, Kaaimans, and Gamtoos Groups lying beneath. Faulting associated with the break up of Gondwanaland exposed this ancient layer, as evident on the West Coast forelands where Malmesbury shales predominate. Over a period of some 50 million years, the cape sediments were violently folded but since then the region has remained relatively stable. During this period, erosion wore down the once extraordinarily high landforms into today’s resistant quartzitic sandstone mountains while depositing another sequence of sediments in the fault basins, especially during the Cretaceous. The softer shales in the Cape Supergroup and remnant Cretaceous sediments formed the low valleys and undulating plains adjacent to the sheer-sided mountains. Due to erosion the quartzitic sandstone fold mountains that characterize this ecoregion do not reach exceptional heights. However, the topography is nonetheless impressive since the mountains rise very steeply from the adjacent lowlands. Another, albeit minor, feature of cape landscapes are scattered outcrops of Precambrian granite, especially along the West Coast forelands. Mountain building in the Cape region formed megafolds and monoclines, with the highest terrain diversity seen where two trends of folding meet (syntaxis). Erosion and the linear formation caused by folding mean that the mountains form strings of isolated, elevated land masses of quartzose substrates, similar to islands scattered throughout an archipelago (Deacon et al. 1992).
Fynbos and renosterveld are the main vegetation communities found in this ecoregion. Fynbos covers 19,227 km2 (54 percent) of the region, and is commonly identified as a hard-leaved, evergreen, and fire-prone shrubland characterized by four major plant types: restioids, ericoids, proteoids, and geophytes (Cowling and Richardson 1995). Among these four plant types, restioids, evergreen reedlike plants, are uniquely diagnostic of fynbos (Campbell 1985). Ericoids are small-leafed shrubs from 0.5 m to 2 m tall which give fynbos its heathlike appearance. The proteoids belong to another Gondwanan family, the Proteaceae, and are known for their showy blooms and their height. At two to four meters, they are the tallest fynbos shrubs. Finally, geophytes or bulblike plants, are usually most conspicuous after fires, and also have attractive blooms. Geophytes are especially prized as horticultural plants. Fynbos thrives on the low nutrient soils of the rocky sandstone mountains. Trees are rare in fynbos, although the Clanwilliam cedar (Widdringtonia nodiflora) is restricted to this ecoregion, found only in the Cedarberg Mountains.
Montane fynbos encompasses a bewildering diversity of plants, especially in the winter-rainfall west where high turnover along habitat and geographic gradients results in extremely complex vegetation patterns (Cowling et al. 1992). The most recent region-wide treatment of the vegetation of the Cape Floristic Region (Cowling and Heijnis 2001) has identified a number of mountain fynbos complexes, which are habitats characterized on the basis of homogeneous climates, topographies, and geologies, but including a wide range of vegetation types. Although these complexes, termed Broad Habitat Units (Cowling and Heijnis 2001), are surrogates for biodiversity, they also reflect the major biogeographical patterns of the montane flora. Thirty such mountain fynbos complexes have been recognized.
Three major environmental gradients determine the distribution of montane fynbos communities: a west to east gradient, a coast to interior gradient, and an altitude-aspect gradient (Cowling and Holmes 1992). Montane fynbos vegetation changes in response to increasing elevation and the aspects of the slope it grows on. Sheltered valleys harbor different plants than exposed ridges, and communities below the snowline differ appreciably from those on mountain peaks.
Proteoid fynbos (fynbos with a high cover of proteoid shrubs) is commonly found on the deep and fertile soils at the base of mountains. As elevation increases, ericaceous communities dominate, growing on moist, fine-grained, and organic carbon-rich soils. The north-facing hillsides are normally drier, and support different communities than the southern slopes. The warmer north-facing slopes have low and drought-prone soils where shallow-rooted restioids dominate. At lower elevations on north-facing slopes, soil depth and restioid root lengths increase. Moving in from the coast, rainfall decreases so that coastal mountains support mainly proteoid and ericaceous fynbos while interior mountains harbor restioid and asteraceous, or dry, fynbos. Small-leaved ericoid shrubs able to obtain soil moisture from deep underground dominate dry fynbos (Cowling et al. 1997; Cowling & Holmes 1992).
Renosterveld means rhinoceros veld in Afrikaans, as a possible reference to the historic habitat of the black rhinoceros (Diceros bicornis). It covers 16,940 km2 (46 percent) of the ecoregion. Unlike fynbos, renosterveld lacks restioids, and proteoids are very rare (Cowling and Richardson 1995). This vegetation type comprises a low shrub layer from 1 m to 2 m tall, mainly made up of ericoids, and usually dominated by the renosterbos (Elytropappus rhinocerotis, Asteraceae), with a ground layer of grasses and seasonally active geophytes. In this ecoregion, renosterveld is most widespread in the western and interior mountain regions where the annual rainfall is between 250 mm and 650 mm. It is replaced by fynbos and succulent karoo at rainfalls higher and lower than this, respectively. In the mountains, this vegetation occupies the fine-grained and relatively fertile soils associated with shale bands, mountain basins, and footslopes. Disturbance regimes are widely credited with setting the boundaries of renosterveld shrubland communities, but recent studies suggest the boundaries may be edaphically determined (Cowling and Holmes 1992).
Both the lowland and montane forms of fynbos and renosterveld share similar biodiversity features. Lowland and Montane Fynbos and Renosterveld ecoregions comprise about 80 percent of the Cape Floristic Region, the smallest of the world’s six floral kingdoms. At 90,000 km2, the Cape Floristic Region is comparable in size to Malawi or Portugal, yet holds 9,000 plant species, of which almost 6,000 are endemic (Cowling & Richardson 1995). Combined, the Lowland and Montane Fynbos and Renosterveld ecoregions harbor approximately 7,000 of the CFR’s 9,000 species (Cowling et al. 1996). Regional richness is amongst the highest in the world, and certainly the highest outside of some tropical rain forest areas (Cowling et al. 1992). The Cape Mediterranean-climate ecosystems are more diverse per set area than all other Mediterranean-climate shrublands (which are all recognized as biodiversity hotspots by Myers et al. (2000)) (Cowling et al. 1996). This high regional richness is a consequence of the extremely high turnover of moderately rich communities along habitat and geographical gradients (Cowling et al. 1992).
In total, the Cape Floral Kingdom contains 526 of the world’s 740 Erica spp. and 69 proteas out of the world’s 112 species (Cowling & Richardson 1995). Genera tend to be exceptionally speciose: Erica and Aspalathus (Fabaceae) and 11 other genera have more than 100 species each (Goldblatt and Manning 2000). As a result, the species per genus ratio is 9.1, one of the highest in the world and more typical of an island biota than a continental area. Geophyte diversity is remarkably high; the lowland and montane fynbos ecoregions support about 1,500 species, most belonging to the petaloid monocot families, notably Iridaceae, Orchidaceae, Hyacinthaceae, and Amaryllidaceae.
About 80 percent of the species found in fynbos and renosterveld are endemic. This is a spectacularly high level of endemism for a continental flora; comparable levels have been recorded only for islands such as Madagascar and New Zealand (Cowling and Hilton-Taylor 1994). A large, but as yet unknown numbers of these endemics are point endemics, restricted to areas of 100 km2 or less (Cowling and McDonald 1999). Most endemic species grow in fynbos vegetation, although locally endemic geophytes are relatively common in renosterveld vegetation in the western, winter-rainfall part of the lowland and montane fynbos ecoregions.
The physiography of the Cape Fold Mountains make them important habitat for fynbos vegetation. The linear sections of the mountains have promoted landscape-level diversity in plant communities, rather than on a local community scale. Regional richness is more important than alpha diversity (number of plant species in a homogenous community). Because these mountains were only of moderate height after erosion, extreme cold has not been a limiting factor, even during the reduced temperatures of the Pleistocene. The maximum depression in mean annual temperature under glacial conditions was approximately 5ºC lower than present temperatures. However, the increased elevation and resultant increased precipitation (also associated with the orientation of the mountains and maritime conditions), of the mountains meant that these regions were important in offsetting the effects of arid periods. Terrain and relief diversity have combined with edaphic and climatic factors to create and preserve upland centers of species richness in the Cape mountains (Deacon et al.1992). In the lithosols of the mountains, podzolization is the main soil-forming process, and a rich diversity of variable and localized soil conditions have selected for microhabitat plant specialists. This contrasts with the slightly richer soils found in the residual duplex and alluvial soils on the coastal platform of the Lowland Fynbos and Renosterveld ecoregion. Here, generalist taxa are favored over specialist species (Deacon et al. 1992).
There are many cases where a single pollinator is responsible for pollinating many species of plants in the CFR. One example unique to the Montane Fynbos and Renosterveld is the mountain pride butterfly (Meneris tulbaghia), which pollinates phylogenetically disparate autumn-flowering, high-altitude geophytes. M. tulbaghia visits only red flowers, and is the exclusive pollinator of at least 15 species (Cowling & Richardson 1995). There is also good evidence that pollinators have driven adaptive radiation in many plant groups, e.g. Gladiolus (Goldblatt and Manning 1998). Insects pollinate the vast majority of fynbos plants, but birds pollinate approximately 430 fynbos plants, so that 75 percent of all plants adapted to bird pollination in southern Africa are found in the fynbos and renosterveld ecoregions. Non-volant mammalian pollinators are also found here, as well as in the ecologically similar kwongan vegetation of southwestern Australia (Cowling & Richardson 1995).
The Montane and Lowland Fynbos and Renosterveld ecoregions are essentially coincident with the Cape Faunal Center (CFC), a discrete zoogeographic zone characterized by the phylogenetic antiquity of much of its invertebrate fauna. (Struckenberg 1962). Many of the species found here have ancient lineages, suggesting that they are Gondwanaland relicts. Examples include freshwater crustaceans (Phreatoicidea, Paramelitidae and the unique, cave-dwelling Spelaeogrypus lepidops); harvestmen (the endemic Triaenonychidae); flies (Pachybates, Trichantha, and Peringueyomina); Megaloptera, Dermaptera, bugs of the tribe Cephalelini, caddisflies (Trichoptera) and various beetles, notable stagbeetles (Lucanidae) of the genus Colophon.
The vertebrate fauna of the CFC is neither especially rich nor distinctive (e.g. Crowe 1990, Branch 1988). However, the fynbos fauna is unique, with some species specifically adapted to living on the low-nutrient vegetation available 127 of southern Africa’s 280 mammal species have been recorded in the Montane Fynbos and Renosterveld ecoregion. Among the ecoregion’s mammal species, 6 are endemic or near-endemic to this ecoregion, including five rodents and one larger mammal, the Cape mountain zebra (Equus zebra zebra). Sadly, two other large mammls once found here are now extinct: the blue antelope (Hippotragus leucophaeus) was hunted to extinction by 1800, and the famous quagga (Equus burchelii quagga) was hunted to extinction the 1850s. Endemic rodents include the Cape spiny mouse (Acomys subspinosus), Cape golden mole (Chrysochloris asiatica), Cape molerat (Georychus capensis), Verraux’s mouse (Myomyscus verrauxii), and the Cape gerbil (Tatera afra). The Cape spiny mouse and Verraux’s mouse are found only in this ecoregion and the adjacent Lowland Fynbos and Renosterveld ecoregion.
The most common antelope species in the Cape mountains are the Cape grysbok (Raphicerus melanotis), common duiker (Sylvicapra grimmia), and klipspringer (Oreotragus oreotragus). The leopard (Panthera pardus) is the only large predator left in the CFC, and is increasingly rare, with most individuals confined to the mountains. Nutrient-poor plants mean that there are few herbivores for large predators like leopards to prey on. As a result, leopards in this region have large home ranges, in some cases ten times larger than the average home range in a savanna environment. Medium-sized mammals, such as chacma baboon (Papio ursinus), honey badger (Mellivora capensis), aardvark (Orycteropus after), and rock hyraxes (Procavia capensis) are all found in this ecoregion. Small herbivores, especially rodents, are numerous in the fynbos and renosterveld ecoregions because they eat seeds, one of the few readily abundant and nutritious food sources in fynbos. Fynbos plants endow their seeds with high concentrations of nitrogen, phosphorus, and potassium to ensure that seedlings will grow in low-nutrient soils. Geophytes, found throughout fynbos and renosterveld communities, also provide a rich source of food in their underground bulbs, corns, and tubers. Baboons and molerats are dependent upon geophytes (Cowling & Richardson 1995), and molerats are capable of eating toxic geophyte bulbs, such as Boophane spp., Amaryllis spp., and Morea spp., and even the highly toxic Ornithogalum thyrsoides, which can kill humans and livestock (Johnson 1992).
Bird diversity in fynbos and renosterveld is not particularly high, because the structurally uniform vegetation offers fewer niches and little high-quality food, such as fleshy fruits and large insects (McMahon and Fraser 1988). Only 288 species (excluding seabirds) have been recorded from the region, and only one of these, the forest canary (Serinus scotops) is strictly endemic. There are six other endemics, which are also found in the Lowland Fynbos and Renosterveld ecoregion: Victorin’s warbler (Bradipterus victorini), Cape rock-jumper (Chaetops frenatus), orange-breasted sunbird (Nectarina violacea), Cape sugarbird (Promerops cafer), and Cape siskin (Serinus totta). Even these fynbos specialists often have broad or flexible feeding habits to better capitalize on available food sources. Orange-breasted sunbirds are commonly found in Lowland Fynbos and Renosterveld from autumn to spring, where they feed on nectar from proteas, but then move into Montane Fynbos and Renosterveld in the summer when the Erica spp. common at higher elevations will flower (Cowling & Richardson 1995). Finally, approximately one third of the birds recorded in the fynbos and renosterveld ecoregions are not native to fynbos and do not utilize fynbos resources in any way. These birds have arrived within the last 100 years, and many in the last 50 years, as a result of habitat transformation. They are totally reliant on transformed environments (Cowling & Richardson 1995).
This ecoregion is a major zone of endemic species richness for freshwater fish, especially the drainage systems of the Olifants, Berg, Breede, and Gouritz Rivers (Skelton et al. 1995). The CFC is home to 109 reptile species, 19 (17.4 percent) of which are endemic (Cowling and Pierce 1999). Tortoise diversity is especially impressive, and South Africa has more terrestrial chelonians than any other country in the world. Although amphibians are low in overall diversity, they exhibit high endemism. In all, half of the 38 species present in both fynbos and renosterveld ecoregions are endemic.
Combined, the Lowland and Montane Fynbos and Renosterveld ecoregions include 1,435 Red Data Book plant species and 112 Red data Book animal species (Rebelo 1992). While many of these species are endangered because they occupy extremely small ranges within natural landscapes, others are highly threatened because they occupy small remnants within urban and agricultural areas (Rebelo 1992). Habitat conversion is less problematic in the Montane Fynbos and Renosterveld ecoregion.
Due to its inaccessibility and poor agricultural potential, this ecoregion has undergone less transformation than its lowland counterpart (Cowling et al. 1999). Only 9.1 percent of the original extent of montane fynbos, and 11.7 percent of montane renosterveld have been transformed, principally by agriculture and forestry. The conservation status of this ecoregion is among the best in South Africa, a legacy of protection aimed at safeguarding mountain catchments for sustainable water production. As of 1999, approximately 7,700 km2 or 26.2 percent of the original extent of fynbos was conserved in statutory reserves. The situation is not nearly as good for renosterveld where corresponding data are 320 km2 or 4.76 percent (Cowling et al. 1999). Many of the protected areas in this ecoregion are relatively large, especially when compared to the situation in the lowlands. However, none is large enough to sustain the full suite of ecological and evolutionary processes required for the long-term conservation of the biota (Cowling et al. 1999). Protected areas wholly or partially contained in this ecoregion include the Anysberg and Baviaanskloof Nature Reserves, Cederberg Wilderness Area, the Groot Winterhoek Wilderness Area, the Grootvadersbosch State Forest complex, the Outeniqua protected areas complex, and Tsitsikamma National Park.
Of the 30 mountain fynbos Broad Habitat Units (BHUs), 20 have more than 10 percent of their original extent conserved; 16 have more than 25 percent conserved, and 15 have achieved reservation targets derived on the basis of biodiversity patterns and retention of habitat (Cowling et al. 1999). Only in the case of 9 BHUs do extant protected areas achieve less than 50 percent of the recommended reservation target. The situation for renosterveld is not nearly as encouraging. Only 2 of the 6 Inland Renosterveld BHUs have greater than 5 percent conserved, and the reservation target has been achieved for only one of these.
Types and Severity of Threats
The three major threats facing montane fynbos and renosterveld habitat are: invasive alien trees and shrubs, novel forms of agriculture (e.g. the cultivation of indigenous species for cut flowers, cultivation and collection of plants for beverages and for their medicinal properties, such as rooibos and honeybush tea) that will transform otherwise marginal agricultural land; and global climate change, which is likely to have a major negative influence on the biodiversity of fynbos, given the specialized habitat requirements of the numerous local and point plant endemics (Rutherford et al. 1999).
Justification of Ecoregion Delineation
This ecoregion was delimited by amalgamating all of the fynbos and renosterveld Broad Habitat Units (BHUs) within the Cape Floristic Region that are associated with montane habitats (Cowling and Heijnis 2001). BHUs are surrogates for plant and animal biodiversity that were identified on the basis of concordant patterns of geology, topography, climate and, in some cases, vegetation types (sensu Low and Rebelo). Montane Fynbos and Renosterveld comprises all of the Mountain Fynbos Complex and all but two of the Inland Renosterveld (see Justification for Lowland Fynbos and Renosterveld ecoregion) BHUs.
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Prepared by: Shirley Cowling
Reviewed by: In progress