South African Journal of Botany 2003, 69(1): 7–26 Copyright © NISC Pty Ltd Printed in South Africa — All rights reserved SOUTH AFRICAN JOURNAL OF BOTANY ISSN 0254–6299

A reappraisal of Acocks’ Bankenveld: origin and diversity of vegetation types

GJ Bredenkamp1* and LR Brown2

1 African Vegetation and Plant Diversity Research Centre, Department of Botany, University of Pretoria, Pretoria 0002, South Africa 2 Applied Natural Sciences, Technikon SA, Private Bag X6, Florida 1710, South Africa * Corresponding author, e-mail: [email protected]

Received 2 October 2002, accepted in revised form 15 November 2002

The great variety of vegetation types found in the sent 16 major vegetation types found in the Bankenveld. Bankenveld was assessed by randomly selecting 220 An assessment was made to relate the vegetation types relevés, representing the major vegetation types, from to various phytochoria and it is concluded that the 23 phytosociological studies from data stored in the Bankenveld has definite floristic affinities to the grass- TURBOVEG database at the University of Pretoria. land of the interior plateaux of South Africa, the savan- These relevés were classified by using TWINSPAN and nas of the Sour Bushveld and Sourish Mixed Bushveld, then refining the result in MEGATAB. The phytosocio- the Drakensberg Afromontane vegetation and the logical table was reduced to 120 relevés which repre- Kalahari vegetation.

Introduction

The term ‘Bankenveld’ was probably coined by Pole-Evans belt of almost 400km, along the 26°S latitude, between 26°E (1920, 1936) to describe the vegetation of the mountainous and 30°E longitude (Figure 1). Low and Rebelo (1996) clas- and hilly area between Rustenburg and the Botswana bor- sify the largest portion of Bankenveld as Rocky Highveld der. Wellington (1955) applied the term ‘Bankeveld’ (without Grassland while this area falls under the Moist Cool the ‘n’) in a physiographic sense to more or less the same Temperate Grassveld (Werger and Coetzee 1978). area to describe the series of cuestas formed by the rocks of Although mapped as a single Veld Type, Acocks recog- the Timeball Hill, Daspoort and Magaliesberg Formations, of nised three variations namely the Western Variation on dry the Pretoria Group in the Transvaal Sequence (SACS 1980). sandy plains, (probably the grasslands of the dolomitic These north-dipping cuestas form the southern rim of the areas), the Central Variation of the Witwatersrand area, Bushveld Basin (Haughton 1969). The vegetation of this largely stony country with rolling topography and the Eastern area is represented by Acocks’ Sourish Mixed Bushveld Variation on wetter sandy plains. The Low and Rebelo (Acocks 1953, 1975, 1988). Acocks’ Bankenveld, however, (1996) map shows most of the Bankenveld area as Rocky occurs south of the Pole-Evans/Wellington Banke(n)veld Highveld Grassland (34) but the area of Eastern Bankenveld and represents the extreme northern parts of the extensive is reduced because the southern part of the Eastern grassland area on the high interior plateaux of South Africa. Bankenveld is considered as being Moist Sandy Highveld Acocks (1953, 1975, 1988) describes the Bankenveld as a Grassland (38) and Moist Cool Highveld Grassland (39), False Grassveld Type. The climax vegetation of this Veld which are related to Acocks’ Cymbopogon-Themeda Veld Type should be, according to Acocks, an open savanna, that (48) or Themeda Veld (Turf Highveld) (52). is a bushveld vegetation, but it has been changed to, and Several phytosociological studies indicate that maintained as grassveld by regular veld fires. Bankenveld vegetation consists of a great variety of grass- This area is generally characterised by a complex topo- land and bushveld communities, distributed in a mosaic pat- graphy which includes rocky hills, ridges, plateaux and tern. All these communities are ecologically interpretable, as plains. Rockiness of the soil surface is a further common they are restricted to specific sets of habitat conditions. characteristic shared by most Bankenveld areas. The In spite of the floristic and structural variability of the vege- Bankenveld borders on the Sourish Mixed Bushveld (Acocks tation and the accompanying variability in habitat, a close 1988) of the savanna biome (Rutherford and Westfall 1994) affinity exists between certain grassveld and bushveld com- to the north. The area covers approximately 23 568km2 and munities, especially grassland and woodland on the same occurs as a 60km (on average) wide east-west stretching geological substrate (Coetzee 1974, Bredenkamp 1975, 8 Bredenkamp and Brown

x Limpopo

Gauteng Mpumalanga

North-West Swaziland

Free State KwaZulu-Natal

Lesotho Northern Cape

Eastern Cape

Western Cape f—nkenveld €rovin™es

PHH H PHH RHH THH VHH Kilometres

Figure 1: The locality of the Bankenveld

Werger and Coetzee 1978, Coetzee 1993, Grobler 2000). altitude, though rockiness of the soil surface, soil texture and Furthermore, the relationships between Bankenveld wood- soil depth are also important factors (Coetzee 1974, land communities and Sourish Mixed Bushveld and Sour Bredenkamp 1975, Bezuidenhout 1993, Coetzee 1993). In Bushveld (Acocks 1953, 1975, 1988) can be demonstrated this area the geology has a major influence on topography, from studies by Van Vuuren and Van der Schijff (1970), Du whereas differences in the land types and soil types are also Plessis (1972), Theron (1973), Coetzee (1975), Van der strongly linked to differences in geological substrata Meulen (1979), Westfall (1981) and Coetzee (1993). On the (Bezuidenhout 1993, Bezuidenhout et al. 1994e). From the other hand the grassveld communities of the Bankenveld mentioned studies it is also apparent that the broad, struc- are obviously related to the Pure Grassveld Types of Acocks turally defined, physiognomic vegetation types are strongly (1953, 1975, 1988), as shown by several authors (Morris linked with geology and topographically defined land forms. 1973, Scheepers 1975, Bezuidenhout 1993, Coetzee 1993). Furthermore, Coetzee (1993) and O’Connor and Geology Bredenkamp (1997) indicated that the Bankenveld vegeta- tion also shows an affinity to the Drakensberg flora. Several geological formations/rock types occur in the The aim of this paper is to identify and delimit the major Bankenveld area (Department of Mineral and Energy Affairs structural and floristic vegetation types within the 1984). A simplified geological map derived from the Bankenveld, and to indicate the origin of Bankenveld from Environmental Potential Atlas (ENPAT 2001) (Figure 2) indi- vegetation and floristic data. Wetland types are not included cates only main types (Du Toit 1954, Haughton 1969, SACS in this reappraisal. 1980), which are briefly mentioned here.

Physical environment Granite and gneiss Old granites and/or gneiss are exposed in three main areas, The main habitat variables that are correlated with differ- namely the Johannesburg Dome, the Vredefort Dome and a ences in floristically defined plant communities, include few small patches west of Potchefstroom. The granites of geology land type, topography (landform, aspect, slope) and the Johannesburg Dome are intensively weathered with South African Journal of Botany 2003, 69: 7–26 9

PT° PU° PV° PW° QH°

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”@ gities282„owns PU° PU° uvi‚uƒhy‚€ x ƒimplified2qeology ”@ ”@ u—roo2ƒediments holomite €e‚‰ƒ q—˜˜ro qr—nite PT° PU° PV° PW° u—rtzite QH° holerite282xorite endesite IHH H IHH PHH Kilometres ƒh—le

Figure 2: A simplified geological map of the Bankenveld (Modified from ENPAT 2001)

deep drainage lines resulting in a gently rolling topography, vegetation may be present where rocky outcrops occur. with shallow, coarse, nutrient poor, well drained soils. The granite landscapes of the Vredefort Dome and an area west Ventersdorp lava of Potchefstroom are flat plains with deeper soils, often used These andesitic lavas mostly form flat plains with deep, for maize cultivation, though some rocky outcrops (insel- nutrient-rich soils and with Cymbopogon-Themeda bergs) with woody vegetation do occur scattered in these Grassveld, often utilised for maize cultivation. In the landscapes. The granite areas are mostly covered by grass- Heidelberg, Alberton and Gatsrand areas, however, the land vegetation. lavas form smooth hills and ridges with woodland communi- ties at sheltered sites, with grassland on exposed high alti- Quartzite and shale tude slopes and plateaux. The quartzites and shales of the Witwatersrand Supergroup and Transvaal Sequence are exposed at various localities, Dolomite forming some of the ridges of the Magaliesberg, Daspoort, Extensive dolomite plains of the Transvaal Sequence, with Timeball Hill, Witwatersrand, Suikerbosrand, Gatsrand and shallow lithosols in the western parts of the Bankenveld are Vredefort Dome (Figure 2). The Magaliesberg, Daspoort and covered with grassland. Chert ridges in the north-western Timeball Hill quartzites and shales form rocky ridges with parts near the escarpment form the habitat for woody plant alternating plains, covered with woody vegetation at shel- communities, while sinkholes often carry a forest-like vege- tered sites and grassland on exposed sites in the Pretoria tation. and Potchefstroom areas. Alluvial plains (Moot, Mooi River and Loopspruit areas) with grassland may occur between Karoo sediments the ridges. Pediments and intrusions of diabase and dolerite In the eastern part of the Bankenveld, coal-bearing Karoo are often covered with Microphyllous Thornveld (Acacia- sediments form extensive flat plains with deep soils which dominated). The older Witwatersrand quartzites normally are often ploughed for maize cultivation. The natural vege- form rugged, rocky hills and ridges, with very coarse sandy, tation on these plains is grassland. leached soils covered with woody vegetation. Exposed high altitude sites carry grassland. Shale areas are well weath- Dolerite, norite and tillite ered, often forming valleys or slightly undulating plains with Sills and sheets of dolerite, norite and tillite are exposed in clayey soils. The vegetation is mostly grassland, but woody the northern parts of the Karoo sediments in the eastern part 10 Bredenkamp and Brown of the Bankenveld. The topography is undulating plains with the northern rim (Kruger 1983, ENPAT 2001), transitional to low hills forming shallow rocky soils, mostly covered with the savanna biome. The complex topography results in grassland but with scanty woody vegetation at sheltered mosaics of different habitats with different vegetation types. sites. Climate Gabbro Hills underlain by gabbro-norite occur in the far north-east- The rainfall of the dry western parts is 500–600mm per ern parts of the Bankenveld. These rocks are rich in heavy annum, the central parts receive 600–700mm while the metals and weather to form ultramafic soils, covered by moister eastern parts receive >700mm per annum (Weather grassland with bushveld on sheltered slopes and in valleys. Bureau 1986, ENPAT 2001). This vegetation contains many endemic species. Average daily maximum temperatures vary from 20–21°C in the southern parts, in the grasslands of the high altitude Land types inland plateaux, to 23–24°C in the central areas of the Bankenveld, to 25–28°C in the northern parts, transitional to Various land types are found within the Bankenveld area the savanna biome. The average daily minimum tempera- (Land Type Survey Staff 1978, 1979a, 1979b, 1979c, 1984, tures vary from 8°C in the southern parts, to 9–10°C in the 1985, 1987, ENPAT 2001), though these are mostly related central areas of the Bankenveld, to 11–12°C in the northern to geology and are therefore only mentioned briefly. parts transitional to the savanna biome (Weather Bureau 1986, ENPAT 2001). A and B land types The A and B land types generally represent flat or slightly Methods undulating landscapes, on granite, shale and Karoo sedi- ments which mostly give rise to deep, freely drained soils or Two hundred and twenty relevés representing the major soils with a plinthic catena. These land types are widespread vegetation types from 23 phytosociological studies through- in the eastern and southern parts of the Bankenveld and are out the Bankenveld region were randomly selected from the mostly covered with grassland, though microphyllous thorn TURBOVEG database (Hennekens 1996a) of the University savanna (Acacia karroo) may occur locally. of Pretoria. These relevés were subjected to a default TWINSPAN classification (Hill 1979), included in the TUR- E land types BOVEG algorithm, to identify vegetation types. The result The Ea land type is restricted to the gabbro-norite hills in the was refined by using the MEGATAB table editing program far north-eastern parts of the Bankenveld, where the vertic Hennekens 1996b), to obtain a phytosociological table or near vertic clayey soils derived from the ultramafic rocks which indicates the classification of the vegetation types, are rich in heavy metals. with their species composition. The table was reduced for practical reasons (to fit journal pages) by randomly discard- F land types ing 100 relevés selected from all vegetation types repre- Shallow Glenrosa or Mispah soils are characteristic of these sented by more than four releves. As this study is a re- land types. The Fa land type is restricted to the dolomite appraisal of major vegetation types found in the Bankenveld areas covered with grassland and with patches of micro- and not a formal syntaxonomical study where plant commu- phyllous Acacia karroo thorn savanna, while the Fb land nities are described, all non-diagnostic species with low type represents the low andesite or quartzite ridges in the constancy at the bottom of the table were also discarded. southern and central parts of the Bankenveld. These ridges A total of 16 major vegetation types were identified; the have grassland on cooler exposed sites and bush on classification and species composition being presented in warmer sheltered sites. Table 1, and a hierarchical habitat interpretation is given in Figure 3. Physiognomic types were identified from the I land types descriptions of the vegetation types in the relevant literature. The Ib land type is characteristic of the very rocky quartzite hills and ridges of the Magaliesberg, Witwatersrand and Results Vredefort Dome areas, with very little, shallow soil. These ridges have grassland on cooler exposed sites and bush on Major physiognomic types warmer sheltered sites. The following six major physiognomic types were identified, Topography and the relationships between these types and the floristi- cally determined vegetation types are indicated in Table 1. Bankenveld represents the extreme northern parts of the 1 Mixed, Short Grassland occurs on the flat to slightly extensive grassland biome area on the high interior plateaux undulating plains of the high altitude interior plateaux, on of South Africa, at altitudes of approximately 1 500–2 000m rounded summits or in broad, shallow alluvial valleys. above lea level. The complex topography of the central part 2 Tall Grassland occurs on well-drained, nutrient poor of the Bankenveld is a result of stony hills, ridges, plateaux soils, or areas with strong anthropogenic influences. and plains while western and eastern parts are flat to slight- 3 Cool Temperate Bushveld occurs throughout the area on ly undulating plains, with conspicuous rocky ridges forming cooler south-facing slopes of hills and ridges or in moder- South African Journal of Botany 2003, 69: 7–26 11

ately dissected, high altitude landscapes where a mosaic rugulosum and Felicia muricata are often present. of bush patches occur locally on rocky outcrops or slopes. Examples of this type of grassland in the Bankenveld were 4 Warm Temperate Bushveld occurs on warmer north-fac- described from the Jack Scott Nature Reserve (Coetzee ing slopes of hills and ridges, especially on the northern 1975), Suikerbosrand Nature Reserve (Bredenkamp and rim of the Bankenveld, in the transition area to the Theron 1980), the eastern Bankenveld (Myburgh 1993) from savanna biome. the Pretoria-Heidelberg-Witbank area (Coetzee et al. 5 Temperate Microphyllous Bushveld occurs on clayey 1995b), Witbank Nature Reserve (Smit et al. 1997), Boskop deeper soils along river banks and on alluvial valley Dam Nature Reserve (Bredenkamp et al. 1994) and from the floors, or at the foot of hills and ridges on deeper collu- Lichtenburg area (Bezuidenhout et al. 1994d). vial soils. 6 Temperate Forest occurs along streams in deep shel- 2. Eragrostis plana Moist Grassland tered kloofs and valleys. Locality and habitat Floristic vegetation types The Eragrostis plana Grassland is a mixed short grassland that occurs scattered over the Bankenveld area, though is 1. Hyparrhenia hirta Anthropogenic Grassland well represented in the eastern and southern parts where the rainfall is relatively high. This grassland type is a moist grass- Locality and habitat land, often found on the A and B land types and usually This tall grassland occurs over vast areas, usually on shal- restricted to flat plains or bottomlands, mostly on moist, deep, low, leached soils on the Johannesburg granite dome, and clayey and poorly drained, seasonally wet soils, adjacent to on undulating north-facing warm andesitic lava slopes of the wetlands such as vleis, spruits, and rivers. These habitats are Suikerbosrand, Witwatersrand and Klipriviersberg areas. often fairly unstable due to seasonal flooding and drying, Disturbed grassland or other disturbed areas such as road which, together with frequent overgrazing, cause degradation reserves or old fields, not cultivated for some years, are also of the vegetation (Bezuidenhout and Bredenkamp 1990). usually Hyparrhenia-dominated. Although some of these tall grasslands appear to be quite natural, they are mostly asso- Vegetation ciated with an anthropogenic influence from recent or even This grassland community is characterised by species group iron-age times. Very often ‘natural’ Hyparrhenia-dominated B, with the diagnostic Eragrostis plana conspicuous, often grasslands occur on ancient lands in the Central Variation of dominant. Paspalum dilatatum, and the rhizomatous the Bankenveld (Acocks 1988) and in the surroundings of Cynodon dactylon, often present in degraded sites, are also archaeological sites (Brown and Bredenkamp 2002), where diagnostic, as well as the forbs Crabbea acaulis, Berkheya the inhabitants had a mosaic of cultivated lands and grazing radula and B. pinnatifida. of domestic stock. It seems that these degraded sites devel- Grass species such as Eragrostis curvula, Themeda trian- oped into to a Hyparrhenia-dominated grassland, which dra, Setaria sphacelata and Digitaria eriantha are often tends to be stable for a very long time. Bredenkamp and abundantly present, and may be locally dominant, while Brown (1998d) found a few relict sites which indicate that the forbs are generally quite rare. original vegetation on the shallow granitic soils of the Good examples of this moist type of grassland are Johannesberg Dome could have been a variant of the described from the moister south-eastern and central parts Monocymbium ceresiiforme-Loudetia simplex Grassland of the Bankenveld (Breytenbach et al. 1993, Coetzee et al. (Vegetation type 7). Hyparrhenia-dominated tall grasslands 1995b). However, the Eragrostis plana Grasslands also are also widely distributed in the midland areas of KwaZulu- occur scattered in bottomland or floodplain situations on the Natal (Moll 1965) and the northern parts of the Eastern Cape drier dolomitic plains of the western Bankenveld (former Transkei area) (Smits et al. 1999), where former and (Bezuidenhout and Bredenkamp 1990, Bredenkamp and present land-uses enhance the prominence of Hyparrhenia Bezuidenhout 1990, Bezuidenhout et al. 1994b). hirta (McKenzie 1984). Eragrostis plana-vegetation is more widely distributed in the wetter eastern escarpment area of southern Africa, e.g. Vegetation footslopes, plains and along drainage lines in eastern Free This grassland is characterised by species group A, which State Drakensberg area (Eckhardt et al. 1993a, 1993b, Fuls includes the tall growing dominant grass Hyparrhenia hirta et al. 1993, Smit et al. 1992, 1993). Eragrostis plana and the invader dwarf shrub Stoebe vulgaris, indicating its becomes prominent in over-utilised and degraded areas, low successional status or degraded condition. and is the dominant grass over large areas in the former Hyparrhenia-dominated grassland mostly has low species Transkei (Smits et al. 1999). This grass became a serious richness, with only a few other species able to establish or invader in the (moist) grasslands of southern Brazil (Pillar survive in the shade of the dense sward of tall grass. Most and Quadros 1997). of these species are relict pioneers or early seral species. The most prominent species include the grasses Cynodon 3. Anthephora pubescens-Schizachyrium sanguineum dactylon, Eragrostis plana, E. racemosa, E. curvula and Sandy Dolomitic Grassland Aristida congesta. 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| |...... |...... |.++..+..+++.|...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |3b+.....++..|...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |.++....+.++.|...... ++|....|...... |..+...... |..|...... |...... |...... |.....|....|...... | |...... |...... |..+..++.+..+|...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |+..+.+..++..|.....+..|....|...... |...... |..|...... |...... |+....+..+..+|.....|....|...... | |...... |...... |+++...+..+..|...... |....|...... |...... |..|.....+..|...... |...... |.....|....|...... | |...... |...... |.++..+.....+|...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |+...+.+...++|...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |++...+..++..|...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |..++.++.....|...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |..+..+++....|...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...++.++....|...... +.|....|...... |...... |..|...... |...... |+.+...... |.....|....|...... | |...... |...... |+..b+...... |...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... +...|...... |+.++++ |...... |...... | |..++...... |...... | |...... |...... | |...... |...... |++.+..+.+++.|.1+1.+++|....|...... |..+...... |..|...... |...... |..+.++...... |.....|....|...... | |...... |...... | |...... |...... |+...+++.+++.|.+..+..+|....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |....+.+.|+++..+..++..|...+..++|....|...... |+++...... +.+...+.|..|...... |...... |...... |.....|....|...... | |...... +...|..1.....|.+...... ++..|....1+.a|+1+1|...... |...... +1...... |..|...... |.++...+..|.1.b.1...... |.....|....|...... | |...... |...... |...... |...... |++++|...... |...... |.| |...... |...... |...... |...... |++++|...... |...... |..|...... |...... |...... |.....|....|...... | |...... +...|...... |...... |.++.+...|++++|...... |++...... +...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |.+++|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |++.+|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |+..+|...... |...... |..|...... |...... |...... |.....|....|...... | |.....+....|...... |...... |.....+..|.++.|...... |...... +....|..|...... |..+++....|..+....+....|.....|....|...... | |...... |...... |...... |...... |.+.+|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |.++.|...... |...... 83547|3166454445|335611|808061123693 4028|2012666983|77|078|999900| 1611233469933| |40052288|122001 2|616266918838|77778| |8333524477|33615666|455544444455|33333566|1111|888078|006 |7134180112|10767035|613475893402|26789908|1234|456393|562891228734637889|5|33203834|447787145|594069982261|56781|7080|025619|2 m a Phytosociological table of the major vegetation types of the Bankenveld of the major vegetation Phytosociological table e e s a m s a m m a s s a a s s s a a m m a s a m a s a a a a s a a a a s a s a a a s a a a s a a a m a s s a a i a a a a s s n a s s a i A a e s e a s i r s a a i a a x s s a i a e r a s egetation typ nthospermum rigidu ristida diffus nthephora pubescen nthospermum hispidulu calypha angustat calypha punctat ster peglera ristea woodi lloteropsis semialat ristida junciformi Table 1: Table Chrysanthemoides monilifer Species group Diplachne fusc Triraphis andropogonoide V Hyparrhenia hirt A A A A A A A A A A Species group B Eragrostis plan Cynodon dactylo Crabbea acauli Berkheya radul Paspalum dilatatu Berkheya pinnatifid Species group C Dicoma anomala Chamaesyce inaequilater Kohautia amatymbic Gazania krebsian Indigofera daleoide Eragrostis stapfi Oxygonum dregeanu Blepharis integrifoli Corchorus asplenifoliu Dicoma macrocephal Helichrysum specie Thesium magalismont Hermannia tomentos Cyperus marginatu Zornia milnean Blepharis angust Eragrostis trichophor Species group D Justicia anagalloide Pogonarthria squarros Barleria macrostegi Helichrysum caespititiu Chamaecrista biensi Crassula lanceolat Ursinia nana Species group E Cymbopogon plurinodi Helichrysum nudifoliu Ledebouria marginat Raphionachme hirsutu Ipomoea crassipe Trichoneura grandiglumi Rhynchosia venulos Hibiscus pusillu Lactuca seriol Dicoma zeyher Hypoxis hemerocallide Species group F Berkheya setifer Rhus discolo Helichrysum miconiifolium Pentanisia prunelloide Setaria nigrirostri Harpochloa fal Lotononis calycin Koeleria capensi Schistostephium crataegifoliu Conyza podocephal Senecio isatideu Wahlenbergia undulat Rubia petiolari Rubus rigidu Rhus rigida Species group G Monocymbium ceresiiform Cyanotis specios Digitaria monodactyl Sporobolus pectinatu Pentanisia angustifoli Bewsia biflor Lopholaena coriifoli Protea welwitschi Sphenostylis angustifoli Species group H Protea roupellia Eragrostis micranth Graderia subintegr Indigofera melanadeni Tritonia nelsoni Crassula nodulos Gnidia sericocephal Indigofera hilari Lotononis erianth Nemesia frutican Tephrosia rhodesic Selago tenuifoli Stoebe vulgari Cyphia angustifoli Stipagrostis uniplumi Chascanum hederaceu Hermannia depress Major physiognomic typ Releve numbe South African Journal of Botany 2003, 69: 7–26 13 ....| . . . . | ....| . . . . | ....| . . . . | ....| . . . . | ....| . . . . | ....| . . . . | .1 ....1| . . . | ....| . 1 . . | ....| . . . . | ....| . . . . | ....| . . . . | ....| . . . . | ....| . 1 . . | .+ ....+| ...... | . . | .....| . . . | . ....+| . . | ...... | . . | .....| . . . | .....| . . . | .....| . . . | .....| . . . | . . . | ....| ...... | . . . | .....| . . . | . . . | ....| ...... | . . . | . . . | ....| ...... | . . . | .....| . . . | .....| . . . | .....| . 1 . | ...... | . . | .....| . . . | . . . | ....1| ...... | . . | .....| . . . | .....| . . . | .....| . . . | . . . | ....| ...... | . . . | 1 . . | ....| ...... | . . . | . . . | ....| 1 . . 1| ....| ...... | . . . | .....| . 1 . | . . . | ....| . . . . | ....| . . . . | ....| ...... | . . . | .....| . . . | .....| . . . | . . . | ....| ...... | . . . | . . . | ....| . 1 . . | ....| . . . . | ....| . . . . | ....| . . . . | ....| . . . . | ....| ...... | . . . | .....| . . . | .....| . . . | . ....1| . . | ...... +| . | 1 ...... | | . . . . | ....| . + . . | ....| . . . . | ....| . . . . | ....| ...... | . . . | . . . | ....| . . . . | ....| . . . . | ....| . . . . | ....| . . . . | 3...| . . . . | +. .+| . . . . | +...| . . . . | +...| . . . . | +...| . . . . | ...... |.....|....|13.31.| ...... |.....|....|...... | ...... |.....|....|...... | a4a..+...... |. .| aa14aa|.+..+|...... |.....|....|...... | ...... 1..+|.....|....|...... |...3|....| ....|...... |.....|....|...+..| ...... |...... |.....|....|...... |...b|....| ...... |..+...... |..1..|....|...... | ...... |...... +...|.....|.+..|...... |...+|....| ...... |...... |.....|....|...... | ++. | . . +|. .++++1.+ ...... |. ++++++. . . .. +++++ ....|...... |...1++++....|.....|....|...... | ....|...... |...... +|.....|....|...... | ....|.....+...|....+1.++.+.|.....|...+|...... | ....|.....+...|...... |.....|....|...... | 3b1b| ++++| +++|1b3+. ....b+11 1+1...... +++| .|. . . . +1|+.1+.|... ..+. . .|b1+1111b ++++|++|b11b33++|.1 ...... |..|...... |...... |....+...+1++|.....|...a|...... | 3b+|+.1. . ..+. . .|.+1 .| 331b1+.|1+|.+1b...... ++|..|3....1..|..1..+...|+133.1a.+.b+|.....|....|...... | ++++ . + . .|. + ..+. . .| +++++ . 1 . . ++|b+|baa1. ++..| . | .+.+.++.1+.| . +1 ...... |. +1+ . ..++..++ 1++|..|. ++++|....|. ++..+++|++...... |...... +1+ . . . +|..|. . +.+++.|++.+|. ++++|. ++++++ 1..|+...+1+..b.+ ++|+b+++++.. . .|. . . .. b . +1 . . . . ++|++|+++b++++|++++++. .|++1+++.+. +1|....|.+.. ++++b+|+++1+...... |. ... +.. . . + . .|+.++|. ++++++|++.++. +++.++|+.++++|1.b++a1+ ++.++|+1.++| + . 1+1+ . +|++|b++1+. ++++|+++.++++|+.++|+1..+b|+++1a+1++. +++++++. ++++++..1+ +++|+.....++1+.+...1 |... . + .1 ...... |. 11 ++++1.+ . .|..|. ++|+++1|.+.. . 1++. ++++++++++++|. 13+++.b+|+...31|bb.+ +++.1 .+. .++++++. 11a3|+++b++.+b++.|1++.1.+.|3211|+b1bb.|+a |...... |...... |...... |...... |....|...... |...... |..|...... |...... |...... |+..++|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... |...... |...... |.....|....|1..++.| |+a3a3+11..|+.+3a.11|.+..+++.++.+|1+++.+1.|211+|....++|..++..a...1+...1..|..|...... +.|.++...... |1.+....b++b1|b+++1|.11.|...... | |...... |...... |...... |...... |....|.....+|...... |..|.....1.a|...... |...1..3.....|4bb.4|b4a1|...... | |...... |...... |...... |...... +|....|.++...|...... +...+.+....|..|.+..++..|...... |...... |.....|....|...... | |...... |...... +.|...... |...... |....|...... |...... |..|...... |.11++1++a|...... |.....|..+3|.....+| |+...... |+...... 1| |.+...+.+..|...... |...... |...... |+.++|...... |....a+.++...... |..|..+.....|...... |+.+.+...++.+|.....|+...|...... | |...... |.....+..|.+..++.+..++|....++..|....|...... |.....+++...... |+.|...... |+.++.+++.|...... ++..|++.++|+.++|...... | |...... |...... |...... |...... |....|...... |...... |..|...... +.|...... |.1...... 1.+|..1.1|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... |...... |...... |.+1.+|....|...... | |...... |...... |...... |...... |....|....1b|...... |a+|43a1 |...... |...... |...... |...... |....|...... |..+...... |..|.++.++.1|...... |...... |.....|....|...... | |...... |...+....|...... |...... |....|+b1...|...... |..|+++..1+.|...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |+...... |..|.33.11..|...... |...... |.....|....|...... | |...... |...... |...... |...... |....|+b+.++|...... |..|.1+..+..|...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...+...... |..|...1+..+|...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...... 1..+++|..|..+b.+..|...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...... |..|.....+++|...... +|...... |.....|....|...... | |...... |...... |...... |...... |....|.++...|.....+...... 1.....|..|+.....+.|...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |++...... +...... |..|1...... +|.....+...|...... +.....|.....|....|...... | |...... |...... |....b.++..+.|...... +.|....|...... |1+1+++3 |...... |...... |...... |+...... |....|...... |...++++++.+....++.|+.|b1+b |...... |...... |....+.+.....|...... |....|...... |.++.++.++..++.....|++|1++.++..|...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |+.1..+.++.+.1+...+|..|...1....|.....+...|...... |.....|....|...... | |.....+....|...... |...... |...... |....|...... |...... +1....+|..|.++.....|...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...... 1.....|..|...... |+a++.b++3|...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...... +...... |..|...... |+.+a1+b1+|...... |.....|....|1b13ba| |.1...... |...... |...... |...... |....|...... |...... |..|...... |ab1..3++b|...... 1.|.....|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... |a1+..1++1|...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... | |...... |...... |...... |...... |....|...... |...... |..|...... |..aa1.bb.|...... |.....|....|...... | . +| |...... |...... |...... |...... |....|...... |...... |..|...... +.|.+1...a.+|...... |.1.1.|....|13.1.1| |...... |...... |...... |...... |....|...... |...... |..|...... |+a....++a|...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... |..+++a...|...... |.....|....|.+.++.| . ++|++|b++b+. . .|++.+++1+++++b+b . +|+1|+... |+. . .+.+ . . 1b|. ++.++++++|++++1+ + ..+. .|++. |+...... |...... |....+.++..b+|1a1b1.+1|....|4b1+34|4411..1 |...... |.....1a.|...... |+..+.+..|....|1....b|..+.+.1..+.+.+....|b.|...... |...... |...... |.....|....|...... | |...+...... |...... |+...b.bb+.+b|.++++.+.|....|...... |..+++1.+b3+.bb....|.+|1..b..++| |...... |...... |.+..b. .b ... . .|+...... |...... |+...++++..++|..++++..|....|.+..+.|++..++.++1++1++1+.|+.|...b....| b . .|. . .1+++1 +.. . . ++|.+|1. |.....+....|...... |....+.+....b|+++++++.|....|.+....|+.++ab.++b.+1...++|.+|b++b....|.+++.+...|...... |.....|....|...... | |...... 3|+1++|. + . . . .|. . . .a. 11|.+ b+++++++.|1aa++. |...... |...... |...... |...... |....|...... |...... |..|...... |...... |b1+111 |...... +..|...... |...... |...... |....|...... |...... |..|.+...+..|.....+...|+1.1.++++1..|+.+.+|++..|...... | |...... |...... |...+...... |...... |....|...... |...... |..|...... |...... a|+...1.+.+++1|b11..|.+.+|+.1...| |...... |...... |...... |...+....|....|...... |...... |..|...... |...... |11+1+...... |.....|....|...... | |...... |...... +.|...... |...... |+.++|...... |...... |..|...... |...... |.1.1+...+..+|.....|....|...... | |...... |...... |...... |...... |....|.....+|...... |.+|...... ++|...... +++|...... ++.+.|.11b1|.a+1|+1b...| |...... |...... |...... |...... |....|....+.|...... |.+|...... |...... |...... +..+.|1+..1|.+..|...... | |...... |...... |++...... +...|..+..b+.|....|...... |...... |..|...... |...... |...1.13.....|.++++|....|...... | |...... |...... |...... |...... |....|...... |...... +...... |..|...... |.....++..|...... |1....|.++.|..1.+.| |...... |...... |...... |..+.....|++++|...... |...... |..|...... +|...... |.1...+..+..+|+.+..|...+|...... | |...... |...... |...... |. |...... +..|...... |...... |...... |....|...... |...... |..|...... |...... |+1+1++...b++|.....|.+..|...... | a s m s s s i s s m s a a s a s i a a a s a i a x a a s a s s a a a s s e a i s a a a i s s a s K Q R s i a x i a a a a r a a a a a a s i i o a a i thrixia elat ncylobotrys capensi ndropogon schirensi cacia karro sparagus suaveolen sparagus laricinu cacia caffra loe marlothi ristida congest ristida transvaalen loe greatheadii v. davyan Table 1 cont. Table Panicum natalens Tristachya leucothri Eustachys paspaloide Dioscorea sylvatica Opuntia ficus-indic Kalanchoe paniculat Setaria sphacelat Maerua cafra Tapiphyllum parvifoliu Vernonia oligocephal Senecio venosu Themeda triandr Species group I Protea caffr A Ochna pulchr Englerophytum magalismontanum A A A A A A A Eragrostis curvula A Brachylaena rotundat A A Rhus leptodicty Species group Burkea african Lippia javanic Table 1 continued Species group O Digitaria diagonali Tristachya biseriat Vernonia natalensi Pearsonia sessilifoli Tristachya rehmanni Rhus zeyher Senecio inornatu Nidorella hottentotic Species group J Loudetia simple Parinari capensi Indigofera hedyanth Hypoxis rigidula Combretum molle Strychnos pungen Croton gratissimu Canthium gilfillani Lannea discolo Crassula swaziensi Species group L Eragrostis racemos Schizachyrium sanguineu Diheteropogon amplecten Bulbostylis burchelli Brachiaria serrata Elionurus muticu Ziziphus mucronat Felicia muricata Ziziphus zeyherian Species group N Commelina africana Vangueria infaust Sida drege Ehretia rigida Isoglossa granti Buddleja salign Felicia filifolia Scolopia zeyher Cassine aethiopic Chlorophytum bowker Pavonia burchelli Ruellia cordat Species group P Setaria lindenbergian Species group Heteropogon contortu Crabbea angustifoli Species group Nuxia congest Clutia pulchell Cymbopogon marginatu Cheilanthes hirt Indigofera comos Trachypogon spicatu Species group M Teucrium trifidum Scabiosa columbari Digitaria erianth Melinis repen Urelytrum agropyroide 14 Bredenkamp and Brown .+b+| 1 b 5 . | .1b.| . . . ..++.| | . . . . | ....| 1 1aa| ....| 1 1+a| ....| 1 1 . a| ....| . 1+ . | ....+| . . . | 1..+.| 1 . 1| 1+1 . | . . . . | b131| . . + . | 1+bb| . . . . | 133 . | . . . . | b3bb| 1 1 . a | +13b| . . . + | +1 . . | . . . . | +1b1| .+1b1| . . .++++| . . . | .++++| . . . | .+11 . | . . . | .+b+ . | . . . | +++ . . | . . . | . . . | . .+++ . | | .+++ . | . . . . . | . .+1 . | . | . . . . | ++1+| . . + + | ++ . . | . 1 . . | .....+...... |.+...|++..|b.++++|++1+|11..| ++++| |...... |...... |...... |...... |....|....+.|...... |..|...... |...... |...... |.....|....|4..1..| |...... |...... |...... |...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |....|....+.|...... |..|...... |...... +..|...... +.|.+..+|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... |...... |...... +.....|.....|.+..|...... | |...... |...... |...... |...... |....|.....1|...... |.+|.....+1+|...... ++|...... +1|.b+.1|+.+.|3+1++.| |...... |...... |...... |...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... +...|...... |...... |...... |....|...... |...... |..|...... |...... |...... |..1..|....|...... | |...... |...... |...... |...... |....|....+.|...... |..|...... +|...... |.1.1...... |.....|+...|...... | |...... |...... |...... |...... |....|....+.|...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |....|.....1|...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |....|....+.|...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |....+...++..|...... +.|....|...... |...... |..|...... +|...... |...... +|..+..|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... +|...... |...... |.....|....|...... | |...... |...... |...... |...... |....|...... |...... |..|...... |...... |...... |.....|....|...... | |...... |...... |...... |...... |....|....+.|...... |..|...... |...... |...... |b1+b+|+...|1.1...| |...... |.....1..|...... |...... |....|....++|...... |.+|.....+..|..+.. |...... |...... |...... |...... |....|.....1|...... |..|...... |...... |...... +....+|11.1.|1a+1|+.1.1.| |...... |...... |...... |...... |....|...... |...... |..|...... |...++...+|...... |+...1|+.++|+.11++| |...... |...... |...... |...... |....|.....+|...... |..|.....+..|..+..+...|...... |.+1..|....|+.....| |...... |...... |...... |...... |....|...... |+....+.....+b.....|+.|...1....|+..++.+..|...... +|++...|++++|++++++| |...... |...... |...... |...... |....|....+1|...... +...... |++|...... |...... |....++....+.|.....|+...|...... | |...... |...... |...... |...... |....|...... |...... |.+|...... +|...... |...... +.|..++.|....|...... | s m m a m e e a s s a a a s s s s a s a a T a a s yrsine africana aytenus undat alopina circaeoide lea europa s. african rewia occidentali tholobium polyphyllu ombretum erythrophyllu hamnus prinoide assinopsis ilicifolia olichos falciformi hoicissus tridentat ombeya rotundifoli ussonia paniculat eltis africana hus pyroide eteromorpha arborescen lematis brachiat iospyros lycioide pecies group S chyranthes asper rgyrolobium rupestr rtemisia afra romus leptoclado splenium aethiopicu pecies group pecies group U uclea crispa uddleja salviifoli hrharta erect ellaea calomelano anthoxylum capens eucosidea serice Table 1 cont. Table S M G A A A A C O R C D B S M S E R D Z C C R G H B E O L C D P South African Journal of Botany 2003, 69: 7–26 15

Bankenveld vegetation

Grassland Woody vegetation

Plains Ridges High altitude Low altitude & hills 7

Short Tall Cool Warm Microphyllous Temperate grass grass temperate temperate bush Forests

Plains High Andesite High altitude Hills & ridges Warm Rocky Foot Mid Sheltered Riverine altitude Granite South facing South facing slopes crests slopes slopes Kloofs communities plateau Anthropogenic Plains 6 1 8 9 10 14 11 15 16

Moist Dry Warm xeric Cool mesic soils soils rocky not rocky 2 12 13

Not rocky Rocky deep soils shallow soils 5

Dolomite Sand over dolomite 4 3

Figure 3: A hierarchical habitat interpretation of the 16 vegetation types of the Bankenveld (numbers refer to the 16 vegetation types)

Sandy Dolomitic Grassland is restricted to the far western Heteropogon contortus and Aristida congesta. Conspicuous drier parts of the Bankenveld, on the Fa land type in the is the absence of Tristachya leucothrix (species group L), Lichtenburg area, North-West Province (Bezuidenhout et al. also mentioned by Acocks (1988) as absent from the west- 1994d). This mixed short grassland occurs on dolomite cov- ern variation of the Bankenveld. ered with an aeolian sand layer of variable depth, and scat- Though seldom dominant, many forbs are found in these tered rock fragments of dolomite and chert. Sink holes occur grasslands, notably Gazania krebsiana, Dicoma anomala, in the dolomite area, and the older sink holes are filled with Senecio venosus, Crabbea angustifolia, Bulbostylis burchel- sand of Kalahari origin. lii, Helichrysum caespititium, Crassula lanceolata, Chamaecrista biensis, Barleria macrostegia, Ursinia nana, Vegetation Chascanum hederaceum and Justicia anagalloides. The analysis indicated several grass and forb species being Dense, patchy stands of Acacia erioloba may be locally diagnostic (species group C). The most prominent diagnos- present in the older sink holes that have been filled with tic species are the grasses Anthephora pubescens, Aristida sand of Kalahari origin. diffusa, Diplachne fusca, Eragrostis stapfii, E. trichophora This vegetation was described by Morris (1976) and and Stipagrostis uniplumis. These grasses, as well as the Bezuidenhout et al. (1994d). perennial forbs Hermannia tomentosa and Indigofera dale- oides indicate a definite Kalahari affinity (Acocks 1988, Smit 4. Triraphis andropogonoides-Trachypogon spicatus 2000). Dolomitic Grassland Other prominent species typical of Bankenveld are the grasses Sporobolus pectinatus, Bewsia biflora, Locality and habitat Trachypogon spicatus, Schizachyrium sanguineum, This mixed short grassland occurs on relatively flat or slight- Diheteropogon amplectens, Brachiaria serrata, Elionurus ly undulating, rugged, stony dolomite plains from Pretoria in muticus, Themeda triandra, Eragrostis curvula, the east to Potchefstroom and Ventersdorp in the west 16 Bredenkamp and Brown

(Bezuidenhout and Bredenkamp 1990). This area repre- (south of Witbank and Middelberg, Coetzee et al. (1995b)) sents the Fa land type (Bezuidenhout et al. 1994a). The parts, where it mainly occurs on the A and B land types, on soils are of variable depth, though mostly shallow and gen- flat or undulating plains with deep, non-rocky soils erally very rocky, leading Louw (1951, 1970) to describe the (Bezuidenhout et al. 1994b). Although mapped as area as ‘Klipveld’. Dolomite and/or chert rock outcrops are Bankenveld these Themeda-dominated grasslands rather widely distributed and sinkholes are frequently found in this represent Acocks’ (1988) Cymbopogon-Themeda Veld (48). area. Large areas are not arable, and therefore much of the These grasslands are relatively poor in plant species, area is presently still covered with natural vegetation, which though, due to the arable soils, much has been destroyed for is used for grazing of livestock and game. agricultural purposes. This type of grassland is very widely distributed over the interior plateaux of South Africa Vegetation (O’Connor and Bredenkamp 1997). Although no characteristic species could be identified, this vegetation type can be recognised by the presence of Vegetation species group D, though with the absence of species group This mixed short grassland is characterised by species C, the latter species group being restricted to the more group E, with the grasses Cymbopogon plurinodis and sandy Anthephora pubescens-Schizachyrium sanguineum Trichoneura grandiglumis diagnostic. Although there are Sandy Dolomitic Grassland (Vegetation Type 3) in the west- many diagnostic forbs, they are never dominant and are ern parts of the dolomite area. mostly inconspicuous and hidden in the dense grass layer. Due to a mosaic of varied rockiness and associated vari- Of these Helichrysum nudifolium, Anthospermum hispidu- ation in soil depth, the vegetation likewise forms a complex lum, Acalypha angustata, Ipomoea crassipes, Hermannia mosaic without specific dominants (Louw 1951, 1970). depressa and the geophyte Hypoxis hemerocallidea are the Prominent species include the grasses Triraphis andro- most prominent. pogonoides, Pogonarthria squarrosa, while individuals of the The dominant grass is mostly Themeda triandra, with forbs Justicia anagalloides, Chascanum hederaceum, Eragrostis curvula, Heteropogon contortus, Setaria sphace- Ursinea nana, Barleria macrostegia, Chamaecrista biensis, lata and Aristida congesta also conspicuous. Crassula lanceolata and Helichrysum caespititium are often Plant communities representative of this grassland from present. the Bankenveld area were described by inter alia The most prominent species include the grasses Bredenkamp et al. (1989), Bredenkamp and Theron (1980), Eragrostis racemosa, E. curvula, Trachypogon spicatus, Bezuidenhout and Bredenkamp (1991), Bezuidenhout et al. Schizachyrium sanguineum, Diheteropogon amplectens, (1994b), Coetzee et al. (1995b). Several phytosociological Andropogon schirensis, Elionurus muticus, Brachiaria serra- studies of similar Cymbopogon plurinodis-Themeda triandra ta, Setaria sphacelata, Heteropogon contortus, Themeda Grasslands from outside the Bankenveld emphasise the triandra, Aristida congesta and Melinis repens. Forbs often affinity to Acocks’ (1988) Cymbopogon-Themeda Grassland encountered are Bulbostylis burchellii, Vernonia oligocepha- (Kooij et al. 1992, Smit et al. 1992, Eckhardt et al. 1993b, la, Crabbea angustifolia and Senecio venosus. Fuls et al. 1993). In some of the older and larger sinkholes a forest-like vegetation has established, with species such as Diospyros 6. Berkheya setifera-Trachypogon spicatus Plateaux lycioides, Celtis africana, Olea europaea subsp. africana, Grassland Heteromorpha arborescens and Xanthoxylum capense and undergrowth which includes Ehrharta erecta, Commelina Locality and habitat benghalensis, Ceterach cordatum, Plectranthus madagas- This grassland is restricted to cool, moist, high altitude cariensis, Cheilanthes hirta, Scadoxus puniceus and (>1 600m) rocky plateaux, crests and higher slopes of the Setaria verticillata (Van Wyk and Bredenkamp 1986). Witwatersrand and Suikerbosrand mountain ranges, mostly Detailed phytosociological classifications and plant com- on andesitic lavas, on the Fb or Ib land types, weathering to munity descriptions for this area are given by Bezuidenhout a clayey soil (Bredenkamp and Theron 1980). and Bredenkamp (1990), Bezuidenhout et al. (1994a). Representative vegetation descriptions from Boskopdam Vegetation Nature Reserve near Potchefstroom (Bredenkamp et al. This mixed short grassland is characterised by many diag- 1994) and the Abe Bailey Nature Reserve near Carletonville nostic species, listed in species group F, including many (Van Wyk and Bredenkamp 1986) contribute to the knowl- forbs. The most prominent of these are Berkheya setifera, edge of this vegetation. Rhus discolor, Acalypha punctata, Helichrysum miconiifoli- um and Pentanisia prunelloides. Diagnostic grasses include 5. Cymbopogon plurinodis-Themeda triandra Grassland Setaria nigrirostris and Koeleria capensis, while diagnostic, small shrubby species such as Chrysanthemoides monilif- Locality and habitat era, Rubus rigidus and Rhus rigida are dwarfed by a dense Patches of the Cymbopogon plurinodis-Themeda triandra grassy layer. Grassland are distributed throughout the Bankenveld, The dominant grasses are Trachypogon spicatus, though it is more widely spread in the southern Themeda triandra, Harpochloa falx and Digitaria diagonalis (Potchefstroom, Johannesburg and Heidelberg, while Vernonia natalensis, Becium obovatum and Crabbea Bezuidenhout and Bredenkamp (1991)) and south-eastern acaulis are often present. This grassland, which is classified South African Journal of Botany 2003, 69: 7–26 17 as Moist Cool-temperate Grassland by Werger and Coetzee alata, Panicum natalense, Urelytrum agropyroides, (1978), shows a Drakensberg affinity (Smit et al. 1995, Tristachya leucothrix, Andropogon schirensis, Themeda Eckhardt et al. 1996). triandra and the forbs Cyanotis speciosa and Senecio veno- An account of plant communities from the Suikerbosrand, sus indicates an affinity with the Drakensberg Highveld representative of this high altitude Bankenveld grassland, is Sourveld vegetation (Acocks 1988, Smit et al. 1992, 1995, given by Bredenkamp and Theron (1980). Eckhardt et al. 1996).

7. Monocymbium ceresiiforme-Loudetia simplex 8. Protea roupelliae Cool Temperate Mountain Bushveld Grassland Locality and habitat Locality and habitat This cool temperate bushveld is mainly found at high alti- This high altitude mixed short grassland is found throughout tudes on relatively steep southern mid-slopes of rocky the study area on rocky mid-slopes of quartzite or andesite quartzite ridges (Ib land type) and is restricted to the west- ridges and hills, often on the Fb and Ib land types. Examples ern and southern parts of the Gauteng region (Bredenkamp are the higher-lying grassland areas in the Suikerbosrand and Brown 1998a, 1998c, Grobler et al. 2002a). The slopes Nature Reserve (Bredenkamp and Theron 1980), normally have a high rock cover with shallow sandy soils. Rustenburg Nature Reserve (Coetzee 1975) and Jack Scott These areas represent the relatively moist and cool habitats. Nature Reserve (Coetzee 1974) and also the grasslands of Mpumalanga (Myburgh 1993). The soils are often shallow Vegetation with high rock cover (up to 60% in some cases). This vege- The diagnostic species include the tree Protea roupelliae, tation is found mostly on cooler aspects, but also occurs on the grass Eragrostis micrantha and the forbs Crassula nodu- the warmer north-facing aspects where scattered individuals losa, Gnidia sericocephala, Graderia subintegra, Indigofera of dwarf shrubs are present. In some degraded areas hilaris, I. melanadenia, Lotononis eriantha, Nemesia fruti- Aristida junciformis, A. congesta and Cynodon dactylon may cans, Tephrosia rhodesica, Tritonia nelsonii and Selago become more prominent. tenuifolia. The vegetation is dominated by the trees Protea roupelli- Vegetation ae (species group H), Protea caffra (species group I) and the This mixed short grassland is characterised by the presence grasses Loudetia simplex (species group J), Trachypogon of species from species group G and include the dwarf spicatus and Tristachya leucothrix (species group L). shrubs Protea welwitschii, Lopholaena coriifolia, the grasses Other woody species also present include the trees Rhus Monocymbium ceresiiforme, Digitaria monodactyla, leptodictya, R. lancea, R. pyroides, Euclea crispa, the Sporobolus pectinatus, Alloteropsis semialata, Bewsia biflo- shrubs Grewia occidentalis, Lippia javanica and the grasses ra and the forbs Cyanotis speciosa, Sphenostylis angustifo- Monocymbium ceresiiforme, Panicum natalense, Urelytrum lia and Pentanisia angustifolia. agropyroides, Themeda triandra and the forb Vernonia This grassland is dominated by the grasses oligocephala. Monocymbium ceresiiforme, Digitaria monodactyla, Examples of this vegetation type were described by Loudetia simplex, Trachypogon spicatus, Eragrostis race- Bredenkamp and Brown (1998a, 1998c) and Grobler et al. mosa, Andropogon schirensis, Schizachyrium sanguineum, (2002a). In the Bankenveld relatively little has been pub- Brachiaria serrata and Themeda triandra. lished on the distribution of this relatively rare community The woody layer consists mainly of a few scattered indi- and it has been included as part of the larger Protea caffra viduals of the dwarf shrubs Protea welwitschii, Lopholaena plant community descriptions. coriifolia, and the geoxylophyte Parinari capensis, that are The presence of the trees Protea roupelliae, Protea caffra locally prominent. The grasses Alloteropsis semialata, and the grasses Loudetia simplex, Trachypogon spicatus, Panicum natalense, Urelytrum agropyroides, Tristachya leu- Tristachya leucothrix, Monocymbium ceresiiforme, Themeda cothrix and Elionurus muticus are also abundant together triandra, Diheteropogon amplectens and Hyparrhenia hirta with the forbs Cyanotis speciosa, Bulbostylis burchellii and indicates a definite affinity to the Drakensberg vegetation Senecio venosus. (Afro-montane phytochorium) (Acocks 1988, Smit et al. This grassland is classified as Moist Cool Temperate 1992, 1995, Eckhardt et al. 1996). Grassland by Werger and Coetzee (1978). Examples of this vegetation type was described by Coetzee (1974) from the 9. Tristachya biseriata-Protea caffra Cool Temperate Jack Scott Nature Reserve, Coetzee (1975) from the Mountain Bushveld Rustenburg Nature Reserve, Bredenkamp and Theron (1980) from the grasslands of the Suikerbosrand Nature Locality and habitat Reserve, Bezuidenhout et al. (1994b) from the grasslands of This cool temperate bushveld is found on gradual to steep the North-West Province, Grobler et al. (2002b) from the rocky quartzite and andesite hills and ridges, on the Fb and urban open space areas of Gauteng and Myburgh (1993) Ib land types, in the Witwatersrand, Magaliesberg, from the high altitude areas in the eastern Bankenveld of Suikerbosrand, Gatsrand and Vredefort Dome areas Mpumalanga. (Coetzee 1974, 1975, Bredenkamp and Theron 1978, The presence of the grasses Monocymbium ceresiiforme, Bezuidenhout et al. 1994c). This vegetation type is promi- Loudetia simplex, Trachypogon spicatus, Alloteropsis semi- nent on the higher altitude slopes and crests. At the northern 18 Bredenkamp and Brown

-most latitudes it is mainly restricted to the cooler south-fac- The woody layer is well developed and can cover up to ing slopes, though at higher latitudes to the south, and espe- 60% of the area. The herbaceous layer has a lower cover cially at high altitudes where it is generally cooler, it has and is comprised of the grasses Schizachyrium san- been recorded on north-facing slopes too (Behr and guineum, Diheteropogon amplectens, Andropogon schiren- Bredenkamp 1988). Here the Protea roupelliae Cool sis and the forbs Bulbostylis burchellii and Commelina Temperate Mountain Bushveld (Vegetation type 8) occupies africana. The grass Setaria lindenbergiana is also locally the south-facing slopes (Bredenkamp and Brown 1998a, prominent. 1998b, 1998c). Examples of this vegetation were described from the scarps and slopes of the Jack Scott and Rustenburg Nature Vegetation Reserves by Coetzee (1974, 1975), from the Suikerbosrand The presence of the tree Protea caffra and the forbs Athrixia Nature Reserve by Bredenkamp and Theron (1978), the elata, Vernonia natalensis and Pearsonia sessillifolia warm temperate mountain bushveld within the Pretoria- (species group I) is characteristic of this vegetation type. Witbank-Heidelberg areas by Coetzee (1993), Melville The vegetation is dominated by the tree Protea caffra Koppies Nature Reserve by Ellery (1994), from the natural (species group I) and the grasses Tristachya biseriata, areas of the western local Metropolitan Council of Gauteng Loudetia simplex and Themeda triandra. The tree Acacia by Bredenkamp and Brown (1998a) and within various natu- caffra is locally prominent, while the grasses Panicum natal- ral urban open spaces in Gauteng by Grobler et al. (2002a). ense, Eragrostis racemosa, Diheteropogon amplectens and Brachiaria serrata are also abundant. The forb layer com- 11. Themeda triandra-Acacia karroo Microphyllous prises mainly Vernonia natalense, Sphenostylis angustifolia Woodland and Pentanisia angustifolia. Acocks (1988) classified this as a false grassveld type Locality and habitat belonging to the central variation of the Bankenveld, while Acacia karroo-dominated woodland is a temperate micro- Werger and Coetzee (1978) classified it as Upland phyllous bushveld that occurs widespread throughout the (Temperate) Sub-humid Mountain Bushveld. Many authors grassland biome and is very prominent in the Eastern Cape described this vegetation, including Coetzee (1974, 1975) (Lubke and Bredenkamp 1996) and KwaZulu-Natal from the Magaliesberg area, Bredenkamp and Theron (1978) (Granger 1996). In the Bankenveld area this type of wood- from the Suikerbosrand, Behr and Bredenkamp (1988) from land is found on colluvial soils on foot-slopes, in bottomland the Witwatersrand, Hattingh (1991) from the hills of the plains and as riparian vegetation along streams and rivers Voortrekker Monument in Pretoria, Bezuidenhout et al. (Bezuidenhout and Bredenkamp 1991). This vegetation type (1994c) from the Gatsrand area in North-West Province, occurs over a wide range of geology, land types, soils and Coetzee et al. (1995a) and Bredenkamp and Brown (1998a, terrain types with low rock cover, but is mostly associated 1998b, 1998c) from the natural areas of the Northern and with moderately deep and often clayey, high nutrient, alluvial Western Metropolitan Local Council and Grobler et al. soils derived from andesite shale, karoo sediments or (2002a) within various natural open spaces in Gauteng. dolomite. The areas in which it occurs are in many cases overgrazed, due to the presence of palatable grass species. 10. Ochna pulchra-Englerophytum magalismontanum Overgrazed or poorly managed sites in the grassland are Warm Temperate Mountain Bushveld often invaded by Acacia karroo and associated species, resulting in bush encroachment and degradation of the Locality and habitat herbaceous layer (Friedel 1987, Bredenkamp and This warm temperate bushveld is restricted to the northern Bezuidenhout 1990). most areas of the Bankenveld in the Pretoria and Rustenburg areas and is found on quartzite, on the Ib land Vegetation type, at high altitudes on warm northern and north-eastern This woodland may be open to quite dense and is charac- aspects with gradual to very steep (up to 55°) slopes terised by the presence of species belonging to species (Coetzee 1974, 1975, Bredenkamp and Theron 1978, group M, with the diagnostic trees Acacia karroo and Coetzee 1993). The shallow soils are covered with large Ziziphus mucronata dominating the woody layer. The diag- boulders and smaller rocks that in some cases provide nostic multi-stemmed shrubs Asparagus suaveolens and moister microhabitats than the surrounding areas. This veg- Asparagus laricinus and the forb Teucrium trifidum are etation represents a transition to the savanna biome and is almost always associated with this vegetation type. classified as Upland (Temperate) Sub-humid Mountain The herbaceous layer is dominated by the grasses Bushveld by Werger and Coetzee (1978). Themeda triandra, Setaria sphacelata while the grasses Eragrostis curvula, Heteropogon contortus, Digitaria eri- Vegetation antha and Elionurus muticus are also abundant locally. The trees Ochna pulchra, Burkea africana, Strychnos pun- On lower mountain slopes this vegetation type and Acacia gens, Croton gratissimus, Combretum molle, Canthium gil- caffra-dominated vegetation (Community 13) often merge to fillanii, the small tree Englerophytum magalismontanum, the form an ecotonal mixed microphyllous woodland. shrub Ancylobotrys capensis, dwarf shrub Lannea discolor Examples of this vegetation were described by inter alia and succulent Crassula swaziensis (species group K) are Coetzee (1975) from the Rustenburg Nature Reserve, not only characteristic, but also the dominant species. Bredenkamp and Theron (1980) from Suikerbosrand, Van South African Journal of Botany 2003, 69: 7–26 19

Wyk and Bredenkamp (1986) on the Abe Bailey Nature karroo-dominated communities (Vegetation type 11), form- Reserve, Bezuidenhout et al. (1988) from the Vredefort ing a mixed thornveld, e.g. in the Faan Meintjes Nature Dome area, Bredenkamp and Bezuidenhout (1990) on the Reserve near Klerksdorp (Bredenkamp and Bezuidenhout Faan Meintjies Reserve in the North-West Province, 1990) and other thornveld communities in the North-West Bezuidenhout and Bredenkamp (1991) from the Bc land Province (Bezuidenhout and Bredenkamp 1991), and also in types of the North-West Province. the Rustenburg Nature Reserve (Coetzee 1975). On the other hand the most mesic Acacia caffra-dominated vegeta- 12. Acacia caffra-Aloe marlothii Mountain Bushveld tion shows some affinity to higher altitude Protea caffra- dominated vegetation (Vegetation type 9) that normally Locality and habitat occurs at cooler sites, above the Acacia caffra-dominated Aloe marlothii-dominated plant communities are found in vegetation (Coetzee 1975). isolated patches on the warmer and xeric, steep and rocky north-facing slopes of the Magaliesberg, Witwatersrand, Vegetation Gatsrand and Suikerbosrand mountain ranges The vegetation is characterised by species group P, with (Bredenkamp and Theron 1980), on quartzite or andesite Setaria lindenbergiana the only diagnostic species. In con- and on the Fb and Ib land types. text to related and surrounding vegetation, this community type seems to be poor in diagnostic species, but is rather Vegetation characterised by the specific species combination, associat- This temperate microphyllous bushveld forms part of the ed with the dominance of Acacia caffra in this particular Acacia caffra-dominated vegetation and may vary from habitat type (Bezuidenhout et al. 1988, Bezuidenhout et al. dense bush to a more open savanna, characterised by the 1994d, Grobler et al. 2002a). prominence of the tall Aloe marlothii. Diagnostic species fur- Acacia caffra is mostly the only dominant woody species, ther include the woody Buddleja saligna, Cassine aethiopica with other prominent species such as Euclea crispa, and Scolopia zeyheri, the shrubby Isoglossa grantii, Felicia Combretum molle, Rhus pyroides, R. leptodictya, filifolia and Pavonia burchellii, the forbs Chlorophytum bowk- Xanthoxylum capense, Dombeya rotundifolia, Diospyros eri and Ruellia cordata, the liana Dioscorea sylvatica and the lycioides, Ehretia rigida and Ziziphus mucronata also pres- succulents Kalanchoe paniculata and the exotic Opuntia ent. Quite often the woody species are grouped in bush ficus-indica (species group O). clumps, resulting in a savanna with scattered individual trees Other prominent though more widespread woody species and bush clumps, with Euclea crispa very prominent. are the dominant Acacia caffra and also Ziziphus mucrona- The most prominent grass species include the diagnostic ta, Rhus leptodictya, Euclea crispa, Grewia occidentalis, Setaria lindenbergiana, and also Eragrostic curvula, while Dombeya rotundifolia and the scandent liana Clematis forbs include Pellaea calomelanos and Commelina africana. brachiata. Although the herbaceous layer is often poorly Descriptions of this community are given by Bezuidenhout developed due to the dense woody layers, grasses such as et al. (1988) from the Vredefort Dome area, Bezuidenhout Eustachys paspaloides and Eragrostis curvula are often and Bredenkamp (1991) from the North-West Province present. grasslands and Grobler et al. (2002a) from the Bredenkamp and Theron (1980) provided a description of Johannesburg-Pretoria area. Particularly good examples this vegetation from the Suikerbosrand Nature Reserve. are provided by Coetzee (1975) from the Rustenburg Nature Reserve, Bezuidenhout et al. (1994d) from the slopes of the 13. Setaria lindenbergiana-Acacia caffra Mountain hills and ridges (Fb land type) in the North-West Province, Bushveld and Coetzee et al. (1995a) from the Magaliesberg in the Pretoria area. Locality and habitat This open- to closed-temperate microphyllous bushveld, 14. Nuxia congesta-Brachylaena rotundata Rocky classified as Upland (Temperate) Sub-humid Mountain Summit Bushveld Bushveld by Werger and Coetzee (1978), is found on grad- ual to moderately steep slopes of outcrops, hills, ridges and Locality and habitat mountains (Fb and Ib land types), which are widely distrib- This cool temperate bushveld is restricted to the extremely uted over the central and western parts of the Bankenveld. rocky crest summits of the quartzite hills and ridges on the It is restricted to shallow and rocky soils with a relatively high Fb and Ib land types of the Magaliesberg, Suikerbosrand, nutritional status (Coetzee 1975), derived from andesite or Witwatersrand, Gatsrand and Vredefort Dome area dolerite, and also from quartzite (Bezuidenhout et al. (Bredenkamp and Theron 1978, Bezuidenhout et al. 1988, 1994d). In the cooler, southern parts of the Bankenveld, this Coetzee 1975, Coetzee et al. 1995). Large quartzite rock vegetation is usually found on lower and warmer north-fac- boulders cover most of the soil surface and the soil is ing slopes, e.g. in the Vredefort Dome area, though in the restricted to cracks and spaces between the rocks. warmer northern parts it may be found at higher altitudes, on crests or south-facing slopes (Coetzee 1975, Bezuidenhout Vegetation et al. 1994d, Coetzee et al. 1995a). This bushveld is characterised by species group R, which On the lower, less rocky slopes with deeper soils the includes the diagnostic trees Brachylaena rotundata, Nuxia Acacia caffra-dominated vegetation often grades into Acacia congesta, Maerua cafra and Tapiphyllum parvifolium, as well 20 Bredenkamp and Brown as the shrubby Indigofera comosa and Clutia pulchella, the 16. Combretum erythrophyllum Riparian Forest grasses Aristida transvaalensis and Cymbopogon margina- tus, the succulent Aloe greatheadii var. davyana and the Locality and habitat xerophytic fern Cheilanthes hirta. Most of the riparian vegetation along larger rivers in the Prominent species include Englerophytum magalismon- Gauteng area of the Bankenveld belongs to this temperate tanum, Canthium gilfilllanii and Ancylobotrys capensis forest vegetation type (Coetzee 1975, Behr and (species group K), indicating a relationship with the Ochna Bredenkamp 1988, Bredenkamp and Brown 1998b). pulchra-Englerophytum magalismontanum Warm Temperate Mountain Bushveld (Vegetation type 10), which occurs on Vegetation the same mountain ranges, though is restricted to the warm This riparian forest is characterised by species group T, with north-facing slopes below the summits. the trees Combretum erythrophyllum and Maytenus undata Euclea crispa, Diospyros lycioides and Zanthoxylum the diagnostic species. capense are other prominent trees, and the xerophytic fern Prominent species include the woody Euclea crispa, Pellaea calomelanos is constantly present. Grewia occidentalis, Zanthoxylum capense, Dombeya This vegetation type was described from the rotundifolia, Cussonia paniculata, Diospyros lycioides, Rhus Suikerbosrand (Bredenkamp and Theron 1978), from the pyroides, R. leptodictya and Celtis africana. Exotic invader Witwatersrand (Behr and Bredenkamp 1988), from the species such as Populus spp., Morus nigra, Salix babyloni- Vredefort Dome area (Bezuidenhout et al. 1988) and from ca, Melia azedarach and Sesbania punicea are often part of the Magaliesberg area (Coetzee 1975, Coetzee et al. 1995). the vegetation. Due to the dense woody layer and due to dis- turbances along rivers, e.g. flooding and trampling, the 15. Myrsine africana-Euclea crispa Temperate Forest herbaceous layer is poorly developed, and mostly consists of annual pioneer weedy species. Locality and habitat Examples of this vegetation were described from the Scattered patches of forest-like bush-clumps represent tem- Magaliesberg area by Coetzee (1975), from the perate forests which occur on the cool south-facing slopes Witwatersrand area by Behr and Bredenkamp (1988) and and along small seasonally dry drainage lines found on the from the Johannesburg Northern Metropolitan area by Fb and Ib land types of the Magaliesberg, Suikerbosrand, Bredenkamp and Brown (1998b). Witwatersrand, Gatsrand and Vredefort Dome area (Bredenkamp and Theron 1980, Bezuidenhout et al. 1994c, Affinities of the Bankenveld Coetzee 1994, Coetzee et al. 1995a). This vegetation is mostly restricted to the clayey, though rocky soils derived Plant community level from andesitic Ventersdorp lavas, shales of the Plant communities indicating the grassland affinity are the Witwatersrand Supergroup or diabase dykes, though may Cymbopogon plurinodis-Themeda triandra Grassland also be found on the more sandy and rocky soils from (Vegetation type 5), and the Hyparrhenia hirta quartzite of the Witwatersrand Supergroup. Anthropogenic Grassland (Vegetation type 1). Many phyto- sociological studies from the grassland biome in the south- Vegetation ern Gauteng area (Bredenkamp and Theron 1980, Grobler The vegetation comprises of short, dense bush-clumps, and et al. 2002b), Free State (Kooij et al. 1990, 1992, Fuls et al. locally the bush-clumps may merge to form taller forest 1992, 1993, Eckhardt et al. 1993b), North-West Province patches. Species group T is characteristic and prominent (Bredenkamp et al. 1989, Bezuidenhout and Bredenkamp woody diagnostic species include Buddleja salvifolia, 1991) and Mpumalanga (Breytenbach et al. 1992, Coetzee Rhamnus prinoides, Heteromorpha arborescens, et al. 1995b) demonstrate this floristic relationship. We sug- Leucosidea sericea, Myrsine africana and Olea europaea gest that the Cymbopogon plurinodis-Themeda triandra subsp. africana. The dominant woody species are, however, Grassland (Vegetation type 5) identified from this study the more widespread Euclea crispa, Rhus pyroides, though mapped by Acocks (1988) as Bankenveld, should Diospyros lycioides, Celtis africana, Rhoicissus tridentata rather be regarded as representative of Acocks’ (1988) and sometimes Cussonia paniculata. Cymbopogon-Themeda Veld (Veld Type 48). Particularly the Where the woody layer of the bush-clumps is very dense, southern parts of the Eastern Variation of the Bankenveld, is the herbaceous layer is poorly developed, often with bare now regarded as Cymbopogon-Themeda Veld (48) or ground and only a few scattered individuals of the diagnos- Themeda Veld (52). tic shade loving grasses Ehrharta erecta and Bromus lepto- The Ochna pulchra-Englerophytum magalismontanum clados and forbs such as Galopina circaeoides, Warm Temperate Mountain Bushveld (Vegetation type 10), Argyrolobium rupestre, Achyranthes aspera, Dolichos falci- the Acacia caffra-Aloe marlothii Mountain Bushveld and the formis, Asplenium aethiopicum and Artemisia afra. Setaria lindenbergiana-Acacia caffra Mountain Bushveld Excellent examples of this vegetation type were described (Vegetation types 12 and 13) show the affinity to the savan- by Bredenkamp and Theron (1980) from the Suikerbosrand, na biome, especially to the Sour Bushveld (Acocks Veld Behr and Bredenkamp (1988) from the Witwatersrand, Type 20) and the Sourish Mixed Bushveld (Acocks Veld Bezuidenhout et al. (1994c) from the Gatsrand area, Coetzee Type 19). This affinity becomes clear when comparing the (1994) and Coetzee et al. (1995a) from the Magaliesberg species composition of these vegetation types to the plant area and Grobler et al. (2002a) from the Gauteng area. communities described by Theron (1973) from the Loskop South African Journal of Botany 2003, 69: 7–26 21

Dam Nature Reserve, Coetzee et al. (1976) from the roides, Aristida stipitata, Andropogon chinensis, Digitaria Nylsvlei Nature Reserve, Van der Meulen (1979) from the brazzae, Brachiaria nigropedata, Enneapogon cenchroides, savanna northwest of Pretoria, Van Rooyen (1983) from the Eragrostis nindensis, Tricholaena monachne and Aloe mar- Roodeplaat Dam Nature Reserve and Westfall et al. (1985) lothii. from the Waterberg. The following Kalahari species also occur in the western Furthermore, many plant species and vegetation types parts of the Bankenveld: Anthephora pubescens, Aristida from the Bankenveld show an affinity to the Drakensberg meriodionalis, Schmidtia pappophoroides, Eragrostis vegetation. The Berkheya setifera-Trachypogon spicatus lehmanniana, Stipagrostis uniplumis, Aristida diffusa, Plateaux Grassland (Vegetation type 6), the Monocymbium Hermannia tomentosa and Indigofera daleoides. ceresiiforme-Loudetia simplex Grassland (Vegetation type 7), the Protea roupelliae Cool Temperate Mountain Bushveld Origin of Bankenveld vegetation (Vegetation type 8), the Tristachya biseriata-Protea caffra Under present climatic conditions the Bankenveld is situated Cool Temperate Mountain Bushveld (Vegetation type 9) and in the climatic transitional zone between temperate grass- the Myrsine africana-Euclea crispa Temperate Forest land and subtropical savanna. Altitudinal gradients often (Vegetation type 15) all represent vegetation types within the show gradients from savanna on the low-lying areas to Bankenveld with floristic similarities with the Drakensberg. grasslands on the high altitude summits of hills, ridges and Acocks (1988) mentioned the migration of vegetation and mountains (Bredenkamp et al. 2002). The grassland biome, specifically emphasised mountain ranges and broken coun- situated on the cool high altitude inland plateaux, is separat- try as particularly favourable migration routes for a wide vari- ed from the lower-lying, and therefore warmer, savanna ety of plants. The east-west mini-escarpment (eastern biome by the mini-escarpment formed by the Magaliesberg extension of the Magaliesberg) forms a direct link between and its eastern and western extensions. Generally situated the Bankenveld and the Drakensberg. However, the at high altitudes, the Bankenveld area is cold, particularly on Drakensberg floristic elements in the Bankenveld (and the exposed summits and crests, and is presently suitable for Waterberg) are now interpreted as relicts of a former wide- the development and maintenance of grassland. Though, spread Drakensberg flora, which shrank to its present distri- large parts of the area are generally characterised by a bution when western inland climatic conditions became drier rugged topography with rocky hills and ridges, with shel- and unfavourable for this Afro-montane vegetation tered, warmer valleys and slopes, which are adequately (Bredenkamp et al. 2002). warm habitats for savanna vegetation. Being situated in a The Anthephora pubescens-Schizachyrium sanguineum climatic transitional zone, the scattered and complex mosa- Sandy Dolomitic Grassland in the far western parts of the ic of a great variety of meso- and micro-habitats, some Bankenveld shows affinity to Kalahari Thornveld (Acocks exposed and cool and some sheltered and warm, results in Veld Type 16). similar patchy mosaics of grassland and savanna. Pollen data (Scott et al. 1997) suggest that although mod- Plant species level ern South African biomes were well established during the The distributions of some species emphasise the floristic Quaternary glacial events, marked cycles of vegetation affinities of the Bankenveld to the grassland, savanna, the change occurred in response to glacial-interglacial changes Drakensberg and the Kalahari (Figure 4). in climate. During the last glacial maximum period (18 000 Some species that occur in the Bankenveld with years BP) vegetation belts were generally lowered by up to Drakensberg affinity are Premna mooiensis, Protea roupelli- 1 000m in altitude, responding to a drop of 5°C. During the ae, Cassinopsis illicifolia, Psydrax obovata, Canthium cold glacial periods grasslands expanded regularly at the mundianum, Trema orientalis, Clutia pulchella, Rhamnus expense of savanna vegetation, with grassy vegetation prinoides, Ochna serrulata, Chrysanthemoides monilifera, occupying a much greater area to the north (Scott 1982a, Diospyros lycioides, Buddleja salviifolia, Cyathea dregei, 1982b, 1987). On the contrary, during the warmer climatic Cussonia paniculata, Rhus dentata, Leucosidea sericea, optimum in the Holocene (7 000 years BP) the savanna veg- Heteromorpha arborescens, Pennisetum sphacelatum, etation spread southwards over the edge of the plateau into Ctenium concinnum, Harpochloa falx, Heteropogon filifolius, the Highveld grassland area (Scott and Vogel 1983). The Andropogon appendiculatis, Sporobolus pectinatus, Aristida data, therefore, suggest that in the Bankenveld area the sciurus, Eulalia villosa, Helichrysum callicomum, H. aure- vegetation often changed, ranging from savanna during onitens, H. polycladum, H. rugulosum, H. lepidissimum, H. warm interglacial phases, to cool open grassland during gla- acutatum, H. coriaceum, H. miconiifolium, H. mundtii and H. cial maxima and with Podocarpus forests during intermedi- cephaloideum. ate wetter phases (Scott 1982a, 1982b). Certain interglacial Savanna species also found in the Bankenveld include periods of favourable climate with increased rainfall in the Psydrax livida, Vangueria infausta, Dombeya rotundifolia, Late and Middle Pleistocene provided opportunity for Ochna pulchra, Pavetta gardeniifolia, P. zeyheri, Strychnos Afromontane (Drakensberg) forests to establish on the pungens, Cassine transvaalensis, Mimusops zeyheri, Magaliesberg and Waterberg. During the last Glacial Englerophytum magalismontanum, Croton gratissimus, Maximum the forests became restricted to the moister Ficus abutilifolia, Cussonia transvaalensis, C. spicata, Drakensberg area and due to aridification they survived only Combretum erythrophyllum, C. molle, C. zeyheri, Ozoroa in protected mountain valleys in the interior of South Africa, paniculosa, Rhus leptodictya, Mundulea sericea, Acacia including the Bankenveld (Scott 1982b, Partridge et al. robusta, Burkea africana, Perotis patens, Uryletrum agropy- 1993). 22 Bredenkamp and Brown

Figure 4: Distribution of some Bankenveld species: Englerophytum magalismontanum in savanna and Bankenveld, Sporobolus discosporus in grassland and Bankenveld, Canthium mundianum in the Drakensberg region and Bankenveld, Panicum coloratum in grassland, Bankenveld and savanna

From the above it becomes clear that the Bankenveld are not arable, and the grazing value is low, due to the pre- originated from an interplay of savanna and grassland alter- dominance of sour grasses. It seems therefore that Acocks’ nating with changes in climatic conditions, the present veg- delimitation of the Bankenveld Veld Type as a mapping unit etation being the relicts of these interactions, where both can be broadly justified as having a similar (low) grazing temperate grassland and (sub)tropical savanna contributed. potential. However, from the variety of vegetation types iden- The Drakensberg floristic elements are relicts from the cool, tified, it is clear that the Bankenveld is quite heterogeneous wetter periods when Drakensberg Afromontane vegetation in terms of floristic and plant community composition. Over was more widespread. such a large area with great variety in climate (rainfall and temperature), geology, land type and topography, this diver- Discussion sity in flora and vegetation is expected. From a conservation or urban development viewpoint, the range of structural and A single important common feature in the habitat over the floristic variation is considered too large to define range of the Bankenveld is shallow, leached, acid lithosols, Bankenveld as a single unit, even on the reconnaissance this probably leading Acocks (1988) to suggest that the veg- level of vegetation investigation. etation of the area has small enough variation to permit the The Acocks (1953, 1975, 1988) interpretation of whole of it to have the same farming potential. Deeper soils Bankenveld being a False Grassveld (fire origin and main- are limited and restricted to local depressions, pediments or tained by fire), is rejected by Bredenkamp and Van Vuuren alluvial valleys. From an agricultural viewpoint, the lithosols (1987), Coetzee (1993), O’Connor and Bredenkamp (1997) South African Journal of Botany 2003, 69: 7–26 23 and Bredenkamp (1999), as there is essentially no differ- Vredefort Dome. An alternative, probably more accept- ence in the fire regimes of the grassland and savanna bio- able interpretation could be to restrict the Bankenveld in mes. These authors interpret Bankenveld as a mosaic of these western areas to the dolomite further south, while grassland and woodland communities controlled by (micro-) the areas north of the dolomite area are included in climatic conditions that exist in the topographically hetero- Acocks’ (1953) Sourish Mixed Bushveld or Sour geneous landscape in the transition zone between these two Bushveld. biomes. Woodland communities occur on relatively warm • Furthermore, the southern Bankenveld boundary in the sites in sheltered valleys and on slopes, while grassland area east of Bronkhorstpruit could be moved northwards communities occur on relatively cold, exposed high altitude to the Witbank-Middelburg area, since the southern plateaux and plains. Woody species are often associated areas are presently mapped by Low and Rebelo (1996) with rocks, also rock walls of archaeological sites as Moist Cool Highveld Grassland. (Bredenkamp and Theron 1980), where they are better pro- • The far north-eastern areas on gabbro-norite should be tected against fire (Acocks 1988, Pillar and Quadros 1997) included in the unique vegetation of Sekhukhuneland and harsh climatic conditions, while these sites are also (Siebert et al. 2001). more moist than rockless plains (Coetzee et al. 1993, • The Cymbopogon plurinodis-Themeda triandra Bredenkamp and Deutschlander 1995). Acocks (1953, Grassland (Vegetation type 5) is not considered as rep- 1975, 1988) indeed mentioned that bushveld vegetation still resentative of Bankenveld, and rather represents occurs along the northern margins of the Bankenveld, on Cymbopogon-Themeda Veld of Acocks (1988). rocky outcrops and hills. He also stated that the western • Parts of the Hyparrhenia hirta Anthropogenic Grassland variation of Bankenveld is clearly transitional from (Vegetation type 1) may not be representative of Cymbopogon-Themeda Veld, a ‘pure’ grassveld type, to Bankenveld, often being ploughed or degraded Sourish Mixed Bushveld. Cymbopogon plurinodis-Themeda triandra Grassland, Although the separation on a structural basis between though ‘typical’ Bankenveld grassland vegetation on grassland and bushveld is ecologically sound, close floristic rocky, non-arable soils, may also be replaced by relationships exist between certain grassland and bushveld Hyparrhenia hirta Anthropogenic Grassland due to communities that occur on similar geological substrates, as human induced impacts. suggested by Werger and Coetzee (1978) for the Moist Cool • The Eragrostis plana Moist Grassland (Vegetation type Temperate Grassland and the Upland Sub-humid Mountain 2), which occupies special moist habitats scattered over Bushveld, and clearly demonstrated by Bredenkamp (1975), the area of the Bankenveld, could be considered as Coetzee (1974) and Coetzee (1993). intra-zonal patches of the Aristida junciformis-Eragrostis Being a transitional area between the grassland and plana grassland from the moist eastern inland plateaux savanna biomes, it is evident that both these biomes con- of southern Africa (O’Connor and Bredenkamp 1997). tribute to the floristics of the Bankenveld. 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By following the geological Phytocoenologia 19: 497–518 lithology it would be possible to delineate the boundaries Bezuidenhout H, Bredenkamp GJ, Elsenbroek JH (1988) Die of the Bankenveld more accurately. plantegroei van die alkaligraniet en aangrensende kwartsiet in die • The present Bankenveld boundary in the Pretoria- Vredefortkoepel noordwes van Parys. SA Tydskrif vir Bronkhorstspruit area westwards to Rustenburg, could Natuurwetenskap en Tegnologie 7: 4–9 be further north to include the Magaliesberg area Bezuidenhout H, Bredenkamp GJ, Theron GK (1994a) A classifica- (Coetzee et al. 1995a, 1995b), as the grassland and tion of the vegetation of the western Transvaal dolomite and chert woodland vegetation of the Magaliesberg are essentially grassland South Africa. South African Journal of Botany 60: 152–161 similar to that of the Witwatersrand, Suikerbosrand and Bezuidenhout H, Bredenkamp GJ, Theron GK (1994b) The vegeta- 24 Bredenkamp and Brown

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