Location and General Description
The Lowland Fynbos and Renosterveld ecoregion is located at the southwestern tip of the African continent where it forms part of the Cape Floristic Region (CFR). It is almost entirely restricted to South Africa’s Western Cape Province, although a small portion is included within the Eastern Cape Province. As the name implies, this ecoregion is located on the coastal lowlands and interior valleys of the CFR, while the Montane Fynbos and Renosterveld ecoregion comprises the uplands and mountains of the CFR.
Both of the fynbos and renosterveld ecoregions are easily distinguishable from neighboring ecoregions by their climate, soils, and the resultant vegetation and flora. These fire-prone ecosystems, where small-leaved evergreen shrubs are the most common growth form, are associated with predominantly winter rainfall of between 250 mm and 2,000 mm annually, and mostly infertile soils (Cowling et al. 1997). The adjacent Succulent Karoo ecoregion, dominated by dwarf, succulent shrubs, occupies drier areas, whereas the Albany Thicket, a large-leaved and semi-succulent shrubland, occupies fire-protected sites in the eastern part of the CFR and into the Eastern Cape, where a greater proportion of rain falls in the summer months (Midgley et al. 1997). The Knysna-Amatole Montane Forest ecoregion occupies a relatively large tract of the southern CFR, and is associated with high, year-round rainfall (more than 800 mm annually) and topographies that afford protection from pervasive fires (Midgley et al. 1997).
The Lowland Fynbos and Renosterveld ecoregion, some 32,756 km2 in extent, encompasses most of the heavily transformed lowland portion of the world-renowned Cape Floristic Region (CFR). Fynbos covers 19,227 km2 (53.8 percent) of this ecoregion and renosterveld covers 16,490 km2 (46.2 percent). Floristically and structurally, the distinction between lowland and montane forms of fynbos and renosterveld is weak. In many cases, there is greater biotic similarity between adjacent lowlands and mountains than there is between disjunct montane or lowland areas. There are other cases, such as the communities on calcareous coastal substrata, where biotic similarity remains relatively high across substantial geographical gradients. However, their geological histories, different climates, and different levels of threats separate the lowland and montane forms of fynbos and renosterveld.
Most of this ecoregion receives annual rainfall between 300 mm and 750 mm annually (Deacon et al. 1992). West of Cape Agulhas, rainfall is concentrated in the winter months and associated with cold fronts budding from the circumpolar westerly system. East of this zone, rainfall is less seasonal because post-frontal events, especially the advection of moist air across a relatively warm Indian Ocean, produce rain throughout the year, but especially in the spring and autumn. Temperatures are generally mild: frost is seldom recorded and summer maximum temperatures seldom exceed 30oC, except in the interior valleys. Coastal areas are generally windy, especially in summer when the pervasive southeasterly trade winds blow. The forelands along the West Coast are influenced by the cold Benguela Current and are prone to fog.
The Cape Floristic Region is more-or-less coincident geologically with the Cape Supergroup, a Devonian-Ordovician series of sedimentary strata, consisting of alternating layers of quartzitic sandstones (the Table Mountain and Witteberg Groups) and fine-grained shales (Bokkeveld Group) (Deacon et al. 1992). These sediments accumulated in a basin depository underlain by one billion-year-old metasediments of the Malmesbury, Kango, Kaaimans, and Gamtoos Groups. Faulting associated with the break up of Gondwanaland has exposed this ancient floor, notably on the West Coast forelands where Malmesbury shales predominate. Over a period of some 50 million years, the cape sediments were violently folded. Since then, the region has remained relatively stable geologically. During this time, the once extraordinarily high landforms have been eroded, shaping today’s mountains of resistant quartzitic sandstone, and depositing another sequence of sediments in the fault basins, especially during the Cretaceous. The softer shales in the Cape Supergroup and remnant Cretaceous sediments now form the gentle valleys and undulating plains adjacent to the steep-sided mountains. Deposits of Tertiary limestones, Pleistocene sands and Holocene dunes mantle the coastal margin.
The Lowland Fynbos and Renosterveld ecoregion comprises nine physiographic regions.
•The West Coast Forelands comprise an undulating plain (from sea level to 300 m) which extends from the Cape Flats to the Olifants River that forms the northern boundary of the ecoregion. Near the coast, acid, aeolian sands, calcareous sands, and limestone underlie this plain, while Malmesbury shales underlie inland areas. Annual rainfall, which occurs exclusively in the winter months, varies from about 200 mm in the north to 500 mm in the south. Another feature of this region is the existence of several plutons or outcrops of Cape granite.
•The Warm Bokkeveld is a small intermontane basin (500 m to 600 m in elevation) centered on the town of Ceres. The underlying rocks are Cape shales, and the winter rainfall varies from 300 mm to 500 mm annually.
•The Elgin Basin is another small intermontane basin (250 m to 350 m in elveation) centered on the town of Elgin. The underlying rocks are Cape shales and the winter rainfall varies from 500 mm to 750 mm annually.
•The Agulhas Plain comprises the southwestern coastal lowlands from Danger Point to the Gouritz River. It includes undulating sandplains and gravelly bottomlands in the west, and extensive limestones and calcareous dunes throughout. Rainfall varies from about 600 mm annually in the west (where most rain falls in the winter months) to less than 400 mm annually in the east (where it is more evenly distributed throughout the year).
•The mid-Breede River Valley is an intermontane fault basin (250 m to 300 m in elevation) with the town of Worcester as its center. Much of the valley is mantled in alluvial deposits, principally gravels and sands, although Cape shales outcrop along the valley walls. Rainfall, which occurs mainly in the winter months, ranges from 250 mm to 350 mm annually.
•The South Coast Forelands are distributed from Caledon in the west to Mossel Bay in the east. They are underlain predominantly by Cape shales, and form an undulating plain 150 m to 400 m above sea level with occasional silcrete- and ferricrete-capped hills, remnants of a Tertiary land surface. Additional features include residual Cretaceous deposits and low hills of Cape sandstone. Rainfall varies from 300 mm to 500 mm annually and falls predominantly in winter in the west, but grades into a nonseasonal pattern of distribution in the east.
•The Little Karoo is an intermontane basin in the southern CFR. Lowland Fynbos and Renosterveld habitats are largely confined to its southern and eastern fringes where they are associated with shale-derived hills and plains (400 m to 1,000 m in elevation) and receive a non-seasonal rainfall of 250 mm to 400 mm annually.
•The Langkloof is an intermontane valley (600 m to 700 m in elevation) in the southeastern CFR. The underlying rocks are shale and sandstone, and the nonseasonal rainfall varies from 400 mm to 650 mm annually.
•The Southeastern Coast Forelands extend from the Tsitisikamma region in the west, to Port Elizabeth at the eastern boundary of this ecoregion, coincident with the eastern boundary of the CFR. This region is relatively complex topographically, comprising shale- and conglomerate-derived (Cretaceous) valleys and plains, extensive sandstone mountain footslopes, sandplains and calcareous coastal dunes. Elevation ranges from sea level to 500 m although most of the area is between 100 m and 200 m in elevation. Rain falls throughout the year and varies from 400 mm to 650 mm annually.
There are five major perennial river systems in the Cape Floristic Region, all of which traverse the Lowland Fynbos and Renosterveld ecoregion. All of these river systems are important habitats for locally endemic freshwater fish (Skelton et al. 1995). They are also important migratory routes for fauna and flora and provide opportunities for exchange between the biotas of the coastal forelands and interior basins (Cowling et al. 1999). The Olifants River forms the northern boundary of the West Coast Forelands while the Berg River drains most of this region. The Breede River is the largest river on the south coast. The Olifant-Gourits-Groot River System, which bisects the Little Karoo and drains much of the South Coast Forelands, is a particularly important migratory corridor. Finally, the Groot-Baviaanskloof-Gamtoos system is the major river system in the east.
The predominant vegetation types of this ecoregion, as implied by its name, are fynbos and renosterveld. Fynbos is a hard-leaved, evergreen and fire-prone shrubland characterized by four major plant types: restioids, ericoids, proteoids, and geophytes (Cowling and Richardson 1995). Restioids, members of the Gondwanan family Restionaceae, are evergreen rush or reedlike plants that are the uniquely diagnostic plant type of fynbos (Campbell 1985). The ericoids include many small-leafed shrubs, ranging from 0.5 m to 2 m tall, which give fynbos its heathlike appearance. The proteoids are the tallest fynbos shrubs from 2 m to 4 m in height, and comprise showy members of the Proteaceae, another Gondwanan family. Geophytes, or bulblike plants, are usually most conspicuous after fires, have blooms which are especially attractive. Many of these have been developed worldwide as valuable horticultural plants. Fynbos thrives in several locations in this ecoregion: on leached, aeolian sands of the coastal forelands and on the nutritionally imbalanced coastal dune and limestone sands of the coastal margin.
Renosterveld is Afrikaans for rhinoceros veld, as a possible reference to the historic habitation by the black rhinoceros (Diceros bicornis). 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, composed mainly of ericoids, and usually dominated by the renosterbos (Elytropappus rhinocerotis) (Asteraceae), with a ground layer of grasses and seasonally active geophytes. Renosterveld always grows on fine-grained, shale-derived soils of the coastal plain and inland valleys where the annual rainfall is between 250 mm and 650 mm. At rainfalls higher and lower than this, it is replaced by fynbos and succulent karoo, respectively.
The vegetation of the ecoregion may be further subdivided into eight major types, or primary Broad Habitat Units (Cowling and Heijnis 2001).
•Dune Pioneer is an azonal type associated with mobile coastal sands throughout the region. It comprises an open, scattered community of low, largely semi-succulent shrubs and herbs, most of which are widespread.
•Fynbos/Thicket Mosaic, as its name implies, is a mixture of fynbos elements (mainly ericoid shrubs and restioids) and large-leaved subtropical shrubs (Cassine and Maytenus, both in the family Celastraceae; Sideroxylon spp., in the family Sapotaceae; Rhus spp. in the family Anacardiaceae). This vegetation type is confined to calcareous coastal sands throughout the ecoregion.
•Sand Plain Fynbos has the typical fynbos structure, with proteoids in the overstory and an understory of ericoids and restioids. It occupies the aeolian sandplains of the West Coast Forelands, and the Agulhas Plain.
•Limestone Fynbos, also with a typical fynbos structure, is associated with the Tertiary limestone habitats of the Agulhas Plain. This type includes numerous calcicole endemics (Willis et al. 1996).
•Grassy Fynbos, characterized by the high cover of summer-growing, or C4, grasses in the field layer, is associated with the wetter, montane margins of the South and South-Eastern Coast Forelands.
•Fynbos/Renosterveld Mosaic comprises a mosaic of fynbos and renosterveld communities associated largely with shale-derived soils, especially where silcrete/ferricrete material is present in the profile or occur as remnant outcrops. Renosterveld often develops as a result of the degradation of fynbos, through overgrazing or burning at unnaturally short intervals. This community is the dominant vegetation of the Elgin Basin and the mid-Breede River Valley and occurs patchily on the West Coast Forelands, the Agulhas Plain, the Langkloof, and the Southeastern Coast Forelands.
•Coast Renosterveld, an especially grassy form of this vegetation, occurs in two main blocks, on the West and South Coast Forelands. Rainfall is concentrated in the winter in the western block, many of the component grasses are winter-growing, and bulb diversity is especially high. The eastern block is grassier than that in the west, has a higher component of summer-growing species, and has a lower diversity of bulbs.
•Inland Renosterveld occupies the shale-derived soils of the drier, interior basins, namely the Warm Bokkeveld and Little Karoo. Grass cover is lower than in Coast Renosterveld and the cover and diversity of succulents is higher, indicative of a transition to the Succulent Karoo ecoregion.
Both the lowland and montane forms of fynbos and renosterveld share similar biodiversity features. Lowland and montane fynbos and renosterveld comprise about 80 percent of the Cape Floristic Region, the smallest of the world’s six floral kingdoms. This relatively small region, a mere 90,000 km2, is also recognized as one of the world’s six floral kingdoms. The CFR is home to about 9,000 vascular plant species, 69 percent of which are endemic (Goldblatt and Manning 2000), and 1,435 (16 percent) of which have Red Data Book status in the South African plant Red Data Book (Hilton-Taylor 1996). The CFR includes 5 endemic plant families and 160 endemic genera. A spectacular feature of the flora is the massive diversification of some taxa: 13 genera have more than 100 species and a very large genus, Erica, has 658 spp. In total, Erica and Aspalathus in the family Fabaceae, as well as 11 other genera have more than 100 species (Goldblatt and Manning 2000). A consequence of this is an extremely high species per genus ratio of 9.1, which is one of the highest in the world and is more typical of an isolated island biota.
Together, 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 among the highest in the world, and certainly the highest outside of some tropical rain forest areas (Cowling et al. 1992). When compared to other Mediterranean-climate regions (all of which are recognized as biodiversity hotspots) (Myers et al. 2000), for a given area, the Cape Mediterranean-climate ecosystems support 1.7 times the diversity of southwestern Australia, 2.2 times the diversity of California and the Mediterranean Basin, and 3.3 times the diversity of Chile (Cowling et al. 1996). This high regional richness is a consequence of the extremely rapid turnover of moderately rich communities along habitat (beta turnover) and geographical (gamma turnover) gradients (Cowling et al. 1992). Other distinctive features in comparison with other Mediterranean-climate floras include the large number of geophytes or bulblike plants (1,552 spp.), especially among the petaloid monocots, and the relative paucity of annuals (6.8 percent of the flora) and trees (2.4 percent). Geophyte diversity is particularly high; the lowland and montane ecoregion support about 1,500 species, most belonging to the petaloid monoct families, notably Iridaceae, Orchidaceae, Hyacinthaceae, and Amaryllidaceae.
About 80 percent of the species complement in fynbos and renosterveld is 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 number 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 combined lowland and montane fynbos ecoregions. Over the whole area generic endemism is modest with 160 endemics, or 16 percent of the total (Goldblatt and Manning 2000). There are four endemic families (Geissolomataceae, Grubbiaceae, Roridulaceae, and Stilbaceae) and three near-endemic families (Bruniaceae, Lanariaceae, and Prionaceae). Goldblatt and Manning (2000) have recognized six centers of endemism for the Cape Floristic Region (both lowland and montane areas), all of which are dominated by fynbos and renosterveld ecosystems. These are:
•Northwestern Center: 22,000 km2; 4 062 spp.; 26.1 percent endemic.
•Southwestern Center: 23,000 km2; 4 654 spp.; 31.9 percent endemic.
•Agulhas Plain Center: 3,000 km2; 1 374 spp.; 14.9 percent endemic.
•Karoo Mountain Center: 19,000 km2; 2 148 spp.; 15.4 percent endemic.
•Langeberg Center: 7,000 km2; 2 367 spp.; 11.7 percent endemic.
•Southeastern Center: 18,000 km2; 2 832 spp.; 9.7 percent endemic.
These data reflect the well-known pattern of a decline in regional-scale species endemism from the western to the eastern parts of the ecoregion (Cowling et al. 1992). This is largely a result of lower numbers of local endemics (habitat specialists and geographical vicariants) in eastern landscapes relative to western areas (Cowling and McDonald 1999). Owing to the spectacularly high plant diversity and endemism of the fynbos and renosterveld, this ecoregion has been previously identified as a biodiversity hotspot by Conservation International (Myers et al. 2000) and as a WWF-IUCN Center of Plant Diversity (WWF and IUCN 1994).
Several studies (as cited in Cowling and McDonald 1999) have shown that local endemics are not a random assemblage phylogenetically, ecologically, and biologically. The odds are high that a local endemic will be a member of a limited number of families (e.g. Ericaceae, Proteaceae, Rutaceae); will be a habitat specialist (usually edaphic); and will be a low, fire-killed shrub with restricted gene dispersal.
The explosive speciation of many fynbos lineages has long fascinated Cape botanists. Two speciation models have been proposed. One suggests that fire-induced population turnover and fragmentation, in combination with edaphic specialization and low vagility, is the driving force behind speciation (Cowling et al. 1992). This would explain the high numbers of non-sprouting species, since lineages with this trait are more vulnerable to turnover than those that sprout (Schutte et al. 1995). However, Johnson (1996) suggests that pollinators have driven speciation in the Cape flora. Two lines of evidence support this hypothesis: widespread pollinator limitation for fruit set (Johnson and Bond 1996); and the highly specialized pollination systems for many groups (Johnson and Steiner 2000). There are many cases where a single pollinator is responsible for pollinating many species of plants, for example long-tongued flies (Tabanidae and Nemestrinidae) are the exclusive pollinators of many genera of petaloid monocots as well as Pelargonium and Erica; a butterfly is the exclusive pollinator of a phylogenetically disparate group of red-flowered and autumn-flowering species; hopliine (monkey) beetle pollination has evolved convergently in many genera etc. (Johnson and Steiner 2000). There is good evidence that pollinators have driven adaptive radiation in many plant groups, e.g. Gladiolus (Goldblatt and Manning 1998).
Other evolutionary phenomena of interest in fynbos and renosterveld ecosystems include plant adaptations to fire (Bond and Van Wilgen 1996) and the widespread occurrence of myrmecochory (Bond and Slingsby 1983). Both of these, alone and in combination, have played an important role in shaping the flora and promoting lineage turnover (Cowling 1987).
The lowland and montane fynbos and renosterveld ecoregions include 1,435 South African Red Data Book plant species and 112 South African Red Data Book animal species (Rebelo 1992). The Cape Peninsula alone, an area of 471 km2 that includes the sprawling metropolis of Cape Town, supports 141 plant and 19 animal South African Red Data Book species (Trinder-Smith et al. 1996, Picker and Samways 1996). 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). This is especially true of lowland fynbos and renosterveld.
The fynbos and renosterveld ecoregions are more-or-less coincident with the Cape Faunal Center (CFC), a distinct zoogeographic zone characterized by the phylogenetic antiquity of much of its invertebrate fauna (Struckenberg 1962). Many of these ancient lineages are Gondwanan 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. Not only is the CFR characterized by the phylogenetic antiquity of its invertebrate fauna, this center is a region of endemic species richness for reptiles, amphibians, and freshwater fish.
The vertebrate fauna of the CFC is neither especially rich nor distinctive (e.g. Crowe 1990, Branch 1990). However, the region 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). Barbine fish account for 81 percent of the fauna with 30 species: total endemicity is 50 percent with 45 percent of endemics occurring in a single drainage system.
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 the Cape Floristic Center makes a major contribution to South Africa’s position as a world leader for this group of terrestrial chelonians. Amphibians, although low in overall diversity, exhibit high endemism. In all, there are 38 species of amphibians, 19 of them endemic.
Bird diversity is not particularly high, owing to the structural uniformity of the vegetation and the shortage of food (McMahon and Fraser 1988). Only 288 (excluding seabirds) species have been recorded from the region, and just 7 of these are endemic or near-endemic. Among the endemic species, most are found in both the Lowland and Montane Fynbos and Renosterveld ecoregions: Victorin’s warbler (Bradipterus victorini), Cape rock-jumper (Chaetops frenatus), orange-breasted sunbird (Nectarina violacea), Cape sugarbird (Promerops cafer), Cape siskin (Serinus totta), and the Cape francolin (Francolinus capensis). Among the ecoregion’s approximately 100 mammal species, 5 endemic or near endemic species remain. Sadly, two others are now extinct: the blue antelope (Hippotragus leucophaeus), hunted to extinction by 1800, and the famous quagga (Equus burchelii quagga), hunted to extinction the 1850s. A charismatic flagship mammal is the strictly endemic bontebok (Damaliscus dorcas dorcas) which once grazed the extensive renosterveld plains of the South Coastal Forelands and is now mainly found in protected sanctuaries. Other strict endemics include the Cape spiny mouse (Acomys subspinosus), the Cape dune molerat (Bathyergus suillis), and Duthie’s golden mole (Chlorotelpa duthiae). Verraux’s mouse (Myomyscus verrauxii) is also considered near-endemic to this ecoregion.
Lowland fynbos and renosterveld have been severely impacted by agriculture, invasive alien plants and urbanization (Cowling et al. 1999). About 41 percent of the original extent of fynbos, and 75 percent of renosterveld have been transformed, principally by agriculture. Given its global significance as a biodiversity hotspot (Cowling and Pierce 1999) and its long-standing recognition as a regional conservation priority (Rebelo 1997), the current conservation status of this ecoregion is very poor. As of 1999, only 827.5 km2 or 4.3 percent of the original extent of fynbos was conserved in statutory reserves; corresponding data for renosterveld are 92.5 km2 or 0.6 percent (Cowling et al. 1999). Most of the protected areas in this ecoregion are small; exceptions are the West Coast National Park and De Hoop Nature Reserve, which are 260 and 400 km2, respectively.
Of the 29 Broad Habitat Units (BHUs - Cowling and Heijnis 2001) recognized in lowland fynbos, only six have more than 10 percent of their original extent conserved, and none has achieved a reservation target proposed by Cowling et al. (1999), derived from the basis of biodiversity patterns and retention of habitat. Six of these BHUs require more than 75 percent of the extant habitat to achieve a reservation target. The situation is even worse for renosterveld. Only one of the seven BHUs has greater than 1 percent conserved, and four require more than 80 percent of the extant habitat to achieve a reservation target.
Types and Severity of Threats
There are five major threats facing remnant lowland fynbos and renosterveld habitat. Invasive vegetation, such as alien trees and shrubs, threatens native vegetation. Novel forms of agriculture can utilize otherwise marginal argricultural land. Some examples of these new forms include the cultivation of indigenous species for cut flowers, beverages, and for their medicinal properties. Urbanization is a serious concern, especially in the two metropolitan centers (Cape Town in the west and Nelson Mandela in the east), and along the coastal margin. Habitat loss is compounded by fragmentation effects, which lead to biodiversity loss on small remnants of irreplaceable habitat. However, research has shown that plant species can persist in very small fragments, even in an agricultural matrix (Cowling and Bond 1991, Kemper et al. 1999). Finally, global climate change 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 those fynbos and renosterveld Broad Habitat Units (BHUs) within the Cape Floristic Region that are associated with lowland habitats (roughly below 300 m elevation) (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 1996). Lowland Fynbos and Renosterveld comprises all of the Dune Pioneer, Fynbos/Thicket Mosaic, Sand Plain Fynbos, Grassy Fynbos, Fynbos/Renosterveld Mosaic, Coastal Renosterveld, and two Inland Renosterveld (Waveren-Bokkeveld and Kannaland) BHUs.
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Prepared by: Shirley Cowling
Reviewed by: In progress