Major plant communities of the Marakele National Park

P.J. VAN STADEN and G.J. BREDENKAMP

Van Staden, P.J. & G.J. Bredenkamp. 2005. Major plant communities of the Marakele National Park. Koedoe 48(2): 59-70. Pretoria. ISSN 0075-6458. To manage and conserve any national park efficiently, a profound knowledge of the ecology is a prerequisite, and to achieve that an inventory of the biotic and abiotic com- ponents must be undertaken. As a contribution to such a program this information was collected for Marakele National Park. The study area covers 290.51 km² in the south- western part of the Limpopo Province. The underlying parent rock of the study area is sandstone, shale and mudstone with several diabase dykes. The soils range from shal- low to deep sandy soils on sandstone and clayey soils on diabase and mudstone. The rainfall varies from 556 mm to 630 mm per annum, mainly during the summer months. The study area experiences warm summers with temperatures of up to 32 ºC and cool, dry winters with frost in the low-lying areas. The vegetation of the study area was clas- sified in a hierarchical, plant sociological system by using TWINSPAN and the Braun- Blanquet technique. The floristic data from 130 relevés were classified to identify five major plant communities, namely one forest community, three savanna/grassland com- munities and one wetland community. These plant communities were ecologically inter- preted by habitat.The phytosociological table was condensed to a synoptic table to describe the major plant communities. Key words: Braun-Blanquet, classification, major plant communities, phytosociology, synoptic table.

PJ van Staden, Centre for Wildlife Management, University of Pretoria, 0001 Pretoria; GJ Bredenkamp, Dept of Botany, University of Pretoria, 0001 Pretoria (george.bre- [email protected])

Introduction knowledge to restore the balance of the orig- inal natural system. The influence of man- The primary objective when proclaiming a agement recommendations on the system national park is to conserve parts of pristine must also be regularly monitored to deter- natural ecosystems for future generations mine if the aims that were set have been (Gertenbach 1987). Marakele National Park achieved satisfactorily. Efficient monitoring (MNP) covers an area of 290.51 km² in the systems also depend upon an inventory of the southwestern part of the Limpopo Province biotic and abiotic components (Gertenbach and is managed as a national park since 1988, 1987). but was officially proclaimed a national park on 11 February 1994. The primary aim of this study was to classify and describe the vegetation of the Marakele In order to manage and conserve any conser- National Park in the Waterberg of the vation area, a profound knowledge of the Limpopo Province (Van Staden 2003). The ecology is a prerequisite, and to achieve that purpose of the classification can be described prerequisite, an inventory of the biotic and as to form an inventory as a basis of manage- the abiotic components of that national park ment. must be undertaken (Edwards 1972). The “natural systems” as they occur today cannot be viewed and conserved as “natural” any Study area more, because of the influence of man. Thus, management recommendations can only be The study area covers 29051 ha in the south- made on the basis of interpreted ecological western part of the Limpopo Province of

ISSN 0075-6458 59 Koedoe 48/2 (2005) Fig. 1. A map indicating the location of the study area in relation to towns.

South Africa, between 27°30'E–27°45'E and soils on sandstone and clayey soils on dia- 24°15'–24°30'S. The position of the park in base and mudstone (Van Staden 2003). relation to towns is shown in Fig. 1 (Van The rainfall varies from 556–630 mm per Staden 2003). annum and occurs mainly during the summer The underlying parent rock of the study area months. The study area experiences warm, (SACS 1980) is sandstone of the Matlabas wet summers with average daily tempera- Subgroup, Aasvoëlkop Formation in the tures of 32 ºC and cool, dry winters with south-western and southern parts; with shale frost in the low-lying areas (Van Staden 2003). and mudstone of the Matlabas Subgroup, Aasvoëlkop Formation, Groothoek Mud- MNP is situated mainly in the Waterberg stone Member; a conglomerate outcrop of Moist Mountain Bushveld (Low & Rebelo the Matlabas Subgroup, Aasvoëlkop Forma- 1996) in the Savanna Biome (Rutherford & tion in the west; and with the biggest part of Westfall 1994). The vegetation of the study the study area consisting of sandstone of the area includes Acocks’ (1988) Sour Bushveld Kransberg Subgroup, Sandriviersberg For- (Veld Type 20), Mixed Bushveld (Veld Type mation. 18), Sourish Mixed Bushveld (Veld Type 19) and North-Eastern Mountain Sourveld (Veld The soils that have developed on the parent Type 8). This Sour Bushveld is listed by materials range from shallow to deep sandy Edwards (1972) as one of 52 of South

Koedoe 48/2 (2005) 60 ISSN 0075-6458 African Veld Types that is extremely lacking to each species according to the Braun-Blanquet in conservation. The Sour Bushveld covers cover-abundance scale, as given by Mueller-Dom- 18306 km2, occurring in mountainous areas bois & Ellenberg (1974) and Werger (1974), and adapted by Barkman et al. (1964). in the previous Transvaal Province (Coetzee 1975; Coetzee et al. 1981). Previous plant The following habitat information was recorded at ecological studies in the Sour Bushveld each sample plot—symbols are used in the synoptic (Acocks 1988) include those by Van Vuuren table: land type was read from the land type map (Land Type Survey Staff 1988); soil forms were clas- & Van der Schijff (1970), Coetzee (1975); sified in accordance with MacVicar et al. (1977); the Coetzee et al. (1981), Westfall (1981) and altitude of each sample plot was recorded using an Westfall et al. 1985. Other plant ecological altimeter and is given in metres; the slope of the ter- work of interest in related vegetation types rain of each sample plot was measured in degrees, include those of Theron (1973) who using an optical clinometer. described the vegetation of the Loskopdam The following classification of slope units (Westfall Nature Reserve; Van der Meulen (1979) who 1981) were used in this study: described the vegetation of the bushveld Symbol Description Class south of the Waterberg; Van Rooyen (1983) who described the vegetation of Roodeplaat L level 0.00° -3.49° Dam Nature Reserve; and Brown (1997) G gentle 3.50° - 17.62° M moderate 17.63°- 36.39° who described the vegetation of Borakalalo S steep 36.40° Game Reserve. The aspect of the terrain where each sample plot is situated was determined using a compass. Aspect is given in the eight compass directions, namely: N - Methods North; S - South; NE - Northeast; SW - Southwest; E - East; W - West; SE - Southeast; NW - Northwest. Analysis The surface rock cover in each sample plot was esti- mated as a percentage stones (> 20 mm diameter), Methods applied are described in detail by Van boulders and rocky outcrops. The following five Staden (2003) and are only summarised here. Stereo classes were used, based on its potential influence on aerial photographs on a scale of 1:50 000 (Task 874 mechanical use (ploughing) (Van der Meulen 1979; of 1984) were used to delineate homogeneous units Westfall 1981): on the basis of physiography and physiognomy (Bre- denkamp & Theron 1978; Westfall 1981; Gertenbach Symbol Class Description 1987). O < 1 % No limitation on A total of 130 sample plots were surveyed through- mechanical utilisation out the study area. The sample plot location was L 1–4 % Low limitation on determined by means of stratified-random sampling mechanical utilisation (Westfall 1981). The number of sample plots for M 5–34 % Moderate limitation on each delineated physiographic-physiognomic unit mechanical utilisation was determined according to the size of each delin- H 35–84 % High limitation on eated unit. mechanical utilisation Termitaria and riparian vegetation was not included V 85–100 % No mechanical utilisation in the placing of the sample plots, due to their limit- possible ed size. Additional sample plots were identified for the termitaria and riparian vegetation and sampling was done in these vegetation types. Synthesis A sample plot size of 10 m x 20 m was fixed and The classification is based on the Braun-Blanquet used throughout the study area. This size is consid- method of vegetation classification, discussed in ered adequate for surveys in savanna vegetation by detail by Westhoff & Van der Maarel (1980), Coetzee (1975), Coetzee et al. (1976), Westfall Mueller-Dombois & Ellenberg (1974) and Werger (1981), Van Rooyen (1983) and Gertenbach (1987). (1974). Data were captured on the mainframe com- puter of the University of Pretoria, in the BBNEW At each sample plot a list of all the species present, software package. It was then exported to be used in was compiled. A cover-abundance value was given the software package BBPC (Bezuidenhout et al.

ISSN 0075-6458 61 Koedoe 48/2 (2005) Table 1 Table 1 (continued) Synoptic table of the major plant communities of Community # 1 2 3 4 5 Marakele National Park Land type Ib/Ad Ad Ib/Fa Fa Ib/Fa For symbols see text, Altitude (x 100 = m) 11-17 12-15 15-20 13-16 13-14 constancy values = percentage Slope G,L G,L M M L Aspect All N S,E N N,E Community # 1 2 3 4 5 Surface rock cover O,H O,M H H O Land type Ib/Ad Ad Ib/Fa Fa Ib/Fa Altitude (x 100 = m) 11-17 12-15 15-20 13-16 13-14 Tristachya rehmannii 41 Slope G,L G,L M M L Cheilanthus hirta 39 Aspect All N S,E N N,E Dicoma anomala 39 Surface rock cover O,H O,M H H O Vernonia galpinii 39 Species Group A Xerophyta retinervus 36 Chaetacanthus costatus 36 Pappea capensis 59 Senecio venosus 36 Mimusops zeyheri 59 Rhynchosia nitens 34 Olea europaea Indigofera mollicoma 34 subsp. africana 55 Pentanisia angustifolia 32 Cussonia paniculata 52 33 Parinari capensis 32 Diospyros whyteana 52 Euphorbia ingens 52 Species Group E Rhus leptodictya 45 Eragrostis racemosa 70 82 Cryptolepis transvaalensis 45 Trachypogon spicatus 30 80 Podocarpus latifolius 38 Bulbostylis burchellii 30 73 Maytenus undata 38 Themeda triandra 78 59 Zanthoxylum capense 34 Gnidia capitata 43 34 Myrsine africana 31 Canthium gilfillanii 31 Species Group F Calpurnia aurea 31 Olea capensis 31 Setaria lindenbergiana 100 Ficus sur 31 Rhoicissus revoilii 89 Grewia occidentalis 31 Hypoestes forskaolii 78 Pseudolachnostylis Species Group B maprouneifolia 78 Strychnos pungens 72 Acacia caffra 65 Diplorhynchus Elionurus muticus 61 condylocarpon 67 Faurea saligna 57 Ochna pulchra 67 Aristida congesta 57 Elephantorrhiza burkei 61 Solanum incanum 48 Littonia modesta 61 Dichrostachys cinerea 43 Stylochiton natalense 56 Pterocarpus rotundifolius 43 39 Maytenus tenuispina 56 Eragrostis capensis 39 Lantana rugosa 50 Sida dregei 39 Tephrosia rhodesica 44 Bewsia biflora 39 Cryptolepis oblongifolia 44 Pogonarthria squarrosa 39 Tapiphyllum parvifolium 44 Eragrostis lehmanniana 39 Kalanchoe paniculata 39 Perotis patens 35 Apodytes dimidiata 39 Vernonia oligocephala 39 Aristida scabrivalvus 33 Species Group C Turraea obtusifolia 33 Ancylbotrys capensis 33 Eragrostis curvula 31 65 Talinum caffrum 33 Dombeya rotundifolia 52 65 33 Euclea crispa 41 43 Species Group G Grewia flavescens 34 39 Englerophytum Berchemia zeyheri 48 39 magalismontanum 43 100 Ziziphus mucronata 38 35 Aristida transvaalensis 50 61 Species Group D Rhynchosia totta 52 56 Loudetia simplex 86 50 Panicum natalense 75 Tephrosia longipes 68 50 Anthospermum hispidula 64 Stachys natalensis 36 50 Urelyterum agropyroides 57 Sphenostylis angustifolia 45 33 Thesium utile 52 Rhus dentata 45 33 Rhynchosia monophylla 50 Acalypha angustata 50 Species Group H Protea caffra 48 Burkea africana 43 94 Aeschynomene rehmannii 45 Combretum molle 30 94 Indigofera hedyantha 43 Lannea discolor 43 83 Monocymbium ceresiiforme 43 38 Ozoroa paniculosa 30 50

Koedoe 48/2 (2005) 62 ISSN 0075-6458 Table 1 (continued) Results Community # 1 2 3 4 5 Land type Ib/Ad Ad Ib/Fa Fa Ib/Fa The vegetation composition of the study Altitude (x 100 = m) 11-17 12-15 15-20 13-16 13-14 area is summarised in a synoptic table (Table Slope G,L G,L M M L 1). All species with a constancy value of Aspect All N S,E N N,E Surface rock cover O,H O,M H H O more than 30 % in any community were included in the synoptic table. This table Phyllanthus parvulus 35 78 Raphionachne galpinii 43 56 therefore shows the most frequently found Setaria sphacelata 78 44 species of each major community and sum- Heteropogon contortus 87 44 marises the relationship between the five Vitex rehmannii 39 61 major plant communities recognised. Species Group I Andropogon schirensis 43 80 61 Commelina africana 39 64 100 Classification Diheteropogon amplectans 30 73 67 Schizachyrium sanguineum 30 50 33 The following major communities were Fadogia homblei 39 57 50 identified: Melinis repens 78 36 72 Brachiaria serrata 70 39 44 Species Group J A. Forest Communities: Vangueria infausta 34 74 55 94 1. Olea europaea subsp. africana - Asparagus transvaalensis 38 39 43 56 Diospyros whyteana Major Community Pellaea calomelanos 31 30 50 89 Cheilanthus viridis 38 56 B. Sour Bushveld and North Eastern Mountain Species Group K Sourveld Communities: Xyris capensis 88 Andropogon huilensis 88 2. Acacia caffra-Heteropogon contortus Miscanthus junceus 63 Major Community Fuirena pubescens 56 3. Protea caffra-Loudetia simplex Major Monopsis decipiens 50 Community Helichrysum aureonitens 50 4. Burkea africana-Setaria lindenbergiana Ischaemum fasciculatum 44 Aristida junciformis 44 Major Community Ascolepis capensis 44 Sebaea leiostyla 44 C. Wetland Communities: Hypericum lalandii 44 Syzygium cordatum 38 5. Andropogon huilensis-Xyris capensis Verbena bonariensis 38 Cyperus thorncroftii 38 Major Community. Cliffortia linaerifolia 31 Arundinella nepalensis 31 Drosera madagascariensis 31 Description of the Major Communities

A Forest Communities 1. Olea europaea subsp. africana-Diospyros whyteana Major Community This major community occurs as forests in 1996), After a TWINSPAN analysis (Hill 1979), the the kloofs, as dense bush clumps on south output of the resulting classification was imported and east facing slopes and as bush clumps on into a spreadsheet, for refinement by Braun-Blan- termitaria. The kloofs are the least exposed quet procedures (Behr & Bredenkamp 1988; Bre- of the geomorphology classes found in the denkamp & Brown 2003). The final classification of study area, with water in the spruits. the relevés was then interpreted for identification of major communities, where-after a synoptic table The species composition of the Olea was compiled by calculating the percentage constan- europaea subsp. africana-Diospyros whyte- cy of each species in each of the major communities ana Major Community is given in Table 1. identified. The result is given in Table 1. These forests and dense bush clumps have

ISSN 0075-6458 63 Koedoe 48/2 (2005) the following diagnostic species, all being species similar to the bush clump communi- trees or shrubs (Species Group A, Table 1): ties on termitaria in the Marakele National Pappea capensis, Mimusops zeyheri, Olea Park. europaea subsp. africana, Cussonia panicu- lata, Diospyros whyteana, Euphorbia B Sour Bushveld and North Eastern Mountain ingens, Rhus leptodictya, Cryptolepis trans- Sourveld Communities vaalensis, Podocarpus latifolius, Maytenus 2. Acacia caffra-Heteropogon contortus undata, Zanthoxylum capense, Myrsine Major Community africana, Canthium gilfillanii, Calpurnia aurea, Ficus sur, Olea capensis and Grewia This major community represents a Sour Bushveld (Acocks 1988) with open to dense occidentalis. savanna typically found on the slopes of the Although various different plant communi- Waterberg, and more open woodland or ties can be recognised within these forests grassland on the summits. This open to and bush clumps (Van Staden 2003), species closed woodland vegetation is found on such as Podocarpus latifolius, Mimusops gradual to moderately steep slopes (mainly zeyheri, Diospyros whyteana, Canthium gil- footslopes) of outcrops, hills, ridges and fillanii and Olea capensis are prominent mountains, which are widely distributed throughout the range of this vegetation. over Marakele National Park. It is restricted to shallow and rocky soils with a relatively Grass and forb species are scanty under the high nutritional status (Coetzee 1975), dense forest vegetation, only Eragrostis derived from diabase and also from quartzite curvula, Asparagus transvaalensis, Pellaea (Bezuidenhout et al. 1994). Many of the calomelanos and Cheilanthes viridis occurred sample plots were located on diabase, which with a constancy of more than 30 %. forms the substrate of this major community. The Olea europaea subsp. africana-Diospy- On the lower, less rocky slopes with deeper ros whyteana Major Community is floristi- soils the Acacia caffra-dominated vegetation cally related to the Sour Bushveld communi- often grades into communities of the plains ties (Communities 2, 3 and 4) within the park forming a mixed thornveld, as also described (Species Group J), but shows particular from the Rustenburg Nature Reserve (Coet- affinity to the Acacia caffra-Heteropogon zee 1975). On the other hand, the most mesic contortus Major Community through Acacia caffra-dominated vegetation shows Species Group C. some affinity to higher altitude Protea Coetzee (1975) described a related commu- caffra-dominated vegetation that normally nity from the Rustenburg Nature Reserve as occurs at cooler sites (Species Group E, Hypoestes verticillaris-Mimusops zeyheri Table 1), at altitudes above the Acacia caf- Forests, and Westfall (1981) described a sim- fra-dominated vegetation, as also reported ilar community from the Farm Groothoek in by Coetzee (1975). the Waterberg as Kloof Forest Communities The soils are mainly shallow, of the Mispah on moderately deep soils in moist, sheltered or Glenrosa Form, though deeper soils of the habitats. Du Preez et al. (1991) classified Hutton or Clovelly Form are encountered at this type of forest as typical Afromontane the foot of the mountain slopes (MacVicar et Forests, where these forests occur in specific al. 1977). The soil depth varies from niches in deep valleys, protected gorges, 100 mm to more than 1000 mm (Land Type crevices and ravines along the eastern and Survey Staff 1988). western slopes of the Drakensberg mountain range. The species composition of the Acacia caf- fra-Heteropogon contortus Major Commu- Coetzee et al. (1976) described termitaria nity is given in Table 1. This major commu- bush clump communities from the Nylsvley nity is differentiated from the other major Nature Reserve, which have diagnostic communities in the area by the following

Koedoe 48/2 (2005) 64 ISSN 0075-6458 diagnostic plant species (Species Group B, and ridges in the North-West Province, and Table 1): Acacia caffra, Faurea saligna, Coetzee et al. (1994, 1995) from the Maga- Dichrostachys cinerea, Pterocarpus rotundi- liesberg in the Pretoria area. Coetzee (1974) folius, Elionurus muticus, Aristida congesta, described Acacia caffra Savannas on diabase Eragrostis capensis, Bewsia biflora, Perotis and in sheltered valleys from the Jack Scott patens, Eragrostis lehmanniana, Pogo- Nature Reserve. Westfall (1981) described a narthria squarrosa, Solanum incanum, Sida similar community as Woodland on moder- dregei and Vernonia oligocephala. ately deep soils in moderately exposed habi- tats, from the Farm Groothoek in the Water- Acacia caffra is mostly a dominant woody berg. species, with other prominent species such as Faurea saligna, Burkea africana, 3. Protea caffra-Loudetia simplex Dichrostachys cinerea, Pterocarpus rotundi- Major Community folius, Vitex rehmannii, Vangueria infausta, Ozoroa paniculosa, Lannea discolor and The Protea caffra-Loudetia simplex Major Combretum molle also prominently present Community is representative of Acocks’ in certain plant communities. Quite often the (1988) Sour Bushveld on moderately deep to woody species are grouped in bush clumps, deep soils in moderately exposed habitats. resulting in a savanna with scattered individ- This mountain bushveld is found on gradual ual trees and bush clumps. to steep rocky hills and ridges in the The prominent grass species include Trachy- Marakele National Park. This vegetation pogon spicatus, Themeda triandra, Setaria type is prominent on the higher altitude sphacelata, Schizachyrium sanguineum, slopes and crests. The soils are mainly very Melinis repens, Heteropogon contortus, shallow, of the Mispah and Glenrosa Forms. Eragrostis racemosa, Diheteropogon The soil depth varies from 10 mm to 500 mm amplectens, Brachiaria serrata, Andropogon (Land Type Survey Staff 1988). schirensis and the diagnostic Elionurus muti- The species composition of the Protea caf- cus and Eragrostis lehmanniana. fra-Loudetia simplex Major Community is The most frequently found forbs include Pel- given in Table 1. This major community is laea calomelanos, Gnidia capitata, Fadogia differentiated by the following diagnostic homblei, Bulbostylis burchellii, Asparagus plant species (species group D, Table 1): transvaalensis, Solanum incanum and Ver- Protea caffra, Urelyterum agropyroides, nonia oligocephala. Tristachya rehmannii, Panicum natalense, The Sour Bushveld (Acocks 1988) commu- Monocymbium ceresiiforme, Xerophyta nities that occur in the park (Communities 2, retinervus, Vernonia galpinii, Thesium utile, 3 and 4) are all floristically related, indicated Senecio venosus, Rhynchosia nitens, Rhyn- by Species Group I, but also through Species chosia monophylla, Pentanisia angustifolia, Groups E, G and H. Parinari capensis, Indigofera mollicoma, Indigofera hedyantha, Dicoma anomala, Descriptions of Acacia caffra-dominated Cheilanthus hirta, Chaetacanthus costatus, vegetation are given by Bezuidenhout et al. Anthospermum hispidula, Aeschynomene (1988) from the Vredefort Dome area, rehmannii and Acalypha angustata. Bezuidenhout & Bredenkamp (1991) from the North-West Province grasslands and This major community is dominated by the tree Protea caffra. Other prominent woody Grobler et al. (2002) from the Johannesburg- species include Vangueria infausta, Rhus Pretoria area. Particularly good examples dentata and Englerophytum magalismon- (Eustachys mutica-Acacia caffra Wood- tanum. lands) are provided by Coetzee (1975) from the Rustenburg Nature Reserve, Bezuiden- The most prominent grasses that are often hout et al. (1994) from the slopes of the hills found in this major community are Trachy-

ISSN 0075-6458 65 Koedoe 48/2 (2005) pogon spicatus, Themeda triandra, northeastern aspects with gradual to very Schizachyrium sanguineum, Melinis repens, steep slopes (Bredenkamp & Brown 2003). Loudetia simplex, Eragrostis racemosa, The shallow soils are covered with large Diheteropogon amplectens, Brachiaria ser- boulders and smaller rocks that in some rata, Aristida transvaalensis and Andro- cases provide moister microhabitats than the pogon schirensis. surrounding areas. Many forb species are present in this vegeta- The species composition of the Burkea tion; the most frequently encountered africana-Setaria lindenbergiana Major include Tephrosia longipes, Stachys natalen- Community is given in Table 1. This major sis, Sphenostylis angustifolia, Rhynchosia community is differentiated by the following totta, Pellaea calomelanos, Gnidia capitata, diagnostic plant species (species group D, Fadogia homblei, Commelina africana, Bul- Table 1): bostylis burchellii and Asparagus trans- Tapiphyllum parvifolium, Strychnos pun- vaalensis. gens, Rhoicissus revoilii, Pseudolachnostylis Many authors described Protea caffra-domi- maprouneifolia, Ochna pulchra, Maytenus nated vegetation, mainly from Sour tenuispina, Elephantorrhiza burkei, Dip- Bushveld and Bankenveld, including Coet- lorhynchus condylocarpon, Cryptolepis zee (1974, 1975) from the Magaliesberg oblongifolia, Apodytes dimidiata, Acy- area, Bredenkamp & Theron (1978) from the lobotrys capensis, Setaria lindenbergiana, Suikerbosrand, Behr & Bredenkamp (1988), Aristida scabrivalvus, Turraea obtusifolia, from the Witwatersrand, Bezuidenhout et al. Tephrosia rhodesica, Talinum caffrum, Stylo- (1994) from the Gatsrand area in North-West chiton natalense, Littonia modesta, Lantana Province, Coetzee et al. (1995), Bredenkamp rugosa, Kalanchoe paniculata and & Brown (1998a; 1998b) from the natural Hypoestes forskaolii. areas of the Western Metropolitan Local Apart from the above prominent diagnostic Council and Grobler et al. (2002) within var- species, other prominent woody species ious natural open spaces in Gauteng. This include Vitex rehmannii, Vangueria infausta, emphasises the floristic relationships Rhus dentata, Ozoroa paniculosa, Lannea between Bankenveld, Sour Bushveld and the discolor, Englerophytum magalismontanum, Drakensberg area (Bredenkamp & Brown Combretum molle and Burkea africana. 2003). Grass species that are prominent in this com- munity include Setaria sphacelata, 4. Burkea africana-Setaria lindenbergiana Major Community Schizachyrium sanguineum, Melinis repens, Loudetia simplex, Heteropogon contortus, This major community represents Sour Diheteropogon amplectens, Brachiaria ser- Bushveld (Acocks 1988). It occurs on gentle rata, Aristida transvaalensis and Andro- to moderately steep slopes of rocky sand- pogon schirensis. stone hills, where soils are litholitic and large Forbs include Tephrosia longipes, Stachys rock boulders cover 53 % of the soil surface. natalensis, Sphenostylis angustifolia, Rhyn- Aspects are mostly northerly, and these chosia totta, Pellaea calomelanos, Fadogia slopes are warm and dry. The soils are main- homblei, Cheilanthus viridis and Asparagus ly of the Mispah and Glenrosa Forms. The transvaalensis. soil varies from 10 mm to more than 500 mm in depth (Land Type Survey Staff 1988). Coetzee (1975) described a similar commu- nity as the Barleria bremekampii- This vegetation type is also found at the Diplorhynchus Tree Savanna from Rusten- northernmost areas of the Bankenveld in the burg Nature Reserve, and Westfall (1981) Pretoria and Rustenburg areas and is found described a similar community as a Wood- at high altitudes on warm northern and land, representative of Acocks’ (1988) Sour

Koedoe 48/2 (2005) 66 ISSN 0075-6458 Bushveld, on moderately deep to deep soils leiostyla, Verbena bonariensis and Xyris in moderately exposed habitats from the capensis. Waterberg. Coetzee (1975) described similar communi- Further examples of this vegetation were ties from the Rustenberg Nature Reserve as described from the scarps and slopes of the the Aristida junciformis-Arundinella nepal- Jack Scott (Coetzee 1974), from the warm ensis Grassland and the Pteridium aquil- temperate mountain bushveld within the Pre- inum-Phragmites mauritianus Reedswamp. toria-Witbank-Heidelberg areas by (Coetzee et al. 1993), from the Melville koppies Nature Reserve (Ellery 1994), from the nat- Discussion ural areas of the western local Metropolitan The vegetation of the study area was classi- Council of Gauteng by Bredenkamp & fied in a hierarchical, plant sociological sys- Brown (1998a) and within various natural tem by using the Braun- Blanquet technique. open spaces in Gauteng by Grobler et al. The floristic data from 130 relevés were (2002). classified to identify five major plant com- munities. These plant communities were 5. Andropogon huilensis-Xyris capensis ecologically interpreted by habitats. These Major Community Major major plant communities should form the basis for a management plan, and should also Many streams arise in seepage areas, for be the basis for a more detailed classification instance on mountain slopes within of the vegetation of this national park. Marakele National Park. These are seasonal- According to White (1978) the majority of ly or perennially waterlogged sponges, with the tree species in South African Afromon- vegetation dominated by sedges and other tane forests are widespread. Amongst them hygrophilous angiosperms and mosses are, inter alia, Podocarpus latifolius, Olea (Noble & Hemens 1978). This major wet- capensis and Myrsine africana that occur as land community occurs along streams and diagnostic species in the Olea europaea tributaries of the Matlabas, Mamba and subsp. africana-Diospyros whyteana Major Sterkstroom rivers and shallow submerged Community within the study area. These marshy areas or sponges. forests on the Waterberg have a clear floris- The soils are wet, mainly of the Avalon, Hut- tic affinity to the Afromontane forests found ton, Katspruit, Oakleaf and Westleigh Forms, in the Drakensberg (Du Preez et al. 1991), derived from sandstone of the Sandriviersberg indicating a much more widespread distribu- tion of forests at previous times, with the Formation (De Vries 1968, 1969). The soil Waterberg forests being a western relict depth varies between 300–1 200 mm (Land (Bredenkamp et al. 2002). The above men- Type Survey Staff 1988). tioned assemblage of species could almost The species composition of this wetland be used to define the Afromontane region as Major Community is given in Table 1. This a whole. Not one species occurs throughout, major community is differentiated by the fol- but most species of the assemblage are pre- lowing diagnostic plant species (Species sent on virtually every “island” of Afromon- Group K): Syzygium cordatum, Cliffortia tane forest in the region. linaerifolia, Andropogon huilensis, Aristida It is interesting to note that the vegetation on junciformis, Arundinella nepalensis, the termitaria have species similar to the for- Ischaemum fasciculatum, Miscanthus est communities, and are therefore included junceus, Ascolepis capensis, Cyperus thorn- into this major community. The greater water croftii, Drosera madagascariensis, Fuirena holding capacity of the soils worked over by pubescens, Helichrysum aureonitens, Hyper- the termites renders these habitats suitable icum lalandii, Monopsis decipiens, Sebaea for forest development (Coetzee et al. 1976).

ISSN 0075-6458 67 Koedoe 48/2 (2005) The Acacia caffra-Heteropogon contortus BEHR, C.M. & G.J. BREDENKAMP. 1988. A phytosoci- Major Community on footslopes, Protea caf- ological classification of the Witwatersrand fra-Loudetia simplex Major Community on National Botanical Garden. South African Jour- nal of Botany 54: 525-533. cooler southerly facing slopes and the BEZUIDENHOUT, H. 1992. Verslag oor die abiotiese Burkea africana-Setaria lindenbergiana komponent van die Kransberg Nasionale Park. Major Community on the warmer northerly Unpublished internal report. facing slopes have a clear floristic relation- BEZUIDENHOUT, H., H.C. BIGGS & G.J. BREDENKAMP. ship with similar plant communities in the 1996. A process supported by the utility BBPC Bankenveld. The relationships between Sour for analysing Braun-Blanquet data on a personal Bushveld, Sourish Mixed Bushveld and computer. Koedoe 39(1): 107-112. BEZUIDENHOUT, H., & G.J. BREDENKAMP. 1991. The Bankenveld woodland communities (Acocks vegetation of the Bc Land Type in the western 1988) are clearly indicated by Bredenkamp Transvaal grassland, South Africa. Phyto- & Brown (2003) and can further be demon- coenologia 19: 497-518. strated from studies by Van Vuuren & Van BEZUIDENHOUT, H., G.J. BREDENKAMP & J.H. ELSEN- der Schijff (1970), Du Plessis (1972), BROEK. 1988. Die plantegroei van die alka- Theron (1973), Coetzee (1975), Van der ligraniet en aangrensende kwartsiet in die Vre- defortkoepel noordwes van Parys. Suid- Meulen (1979), Westfall (1981) and Coetzee Afrikaanse Tydskrif vir Natuurwetenskap en (1993). Furthermore, Coetzee (1993), Tegnologie 7: 4-9. O’Connor & Bredenkamp (1997) and Bre- BEZUIDENHOUT, H., G.J. BREDENKAMP & G.K. denkamp & Brown (2003) indicated that the THERON.1994. Syntaxonomy of the vegetation of Bankenveld vegetation also shows an affini- the Fb land type in the western Transvaal grass- ty to the Drakensberg flora. This relationship land, South Africa. South African Journal of may also be seen in the presence of the Botany 60: 72-80. BREDENKAMP, G.J. & L.R. BROWN. 1998a. A vegeta- Afromontane forests in the Drakensberg, tion assessment of the open spaces in the West- Bankenveld and Waterberg. ern Metropolitan local Council area. Pretoria: Ekotrust CC. In the cooler, southern parts of the Banken- BREDENKAMP, G.J. & L.R. BROWN.1998b. A vegeta- veld, Acacia caffra-dominated vegetation is tion assessment of open spaces in the northern usually found on lower and warmer north- areas of the Northern Metropolitan Local Coun- facing slopes, e.g., in the Vredefort Dome cil. Pretoria: Ekotrust CC. area, though in the warmer northern parts BREDENKAMP, G.J. & L.R. BROWN. 2003. A reap- e.g. the Marakele National Park, it may be praisal of Acocks’ Bankenveld: Origin and found at higher altitudes, on crests or south- diversity of vegetation types. South African Journal of Botany 69(1): 7-26. facing slopes (Coetzee 1975; Bezuidenhout BREDENKAMP, G.J., F. SPADA & E. KAZMIERCZAK. et al. 1994; Coetzee et al. 1995). 2002. On the origin of northern and southern hemisphere grasslands. Plant Ecology 163(2): Due to the complex mosaic distribution pat- 209-229. tern of the sub-communities that are includ- BREDENKAMP, G.J. & G.K. THERON. 1978. A syneco- ed under the major communities, a vegeta- logical account of the Suikerbosrand Nature tion map is more appropriate when the Reserve. 1. The phytosociology of the Witwater- details of the sub-communities are pub- srand geological system. Bothalia 12: 513-529. lished. BROWN, L.R. 1997. A plant ecological study and wildlife management plan of the Borakalalo Nature Reserve, North-west Province. Ph.D. the- sis. University of Pretoria, Pretoria. References COETZEE, B.J. 1974. A phytosociological classifica- tion of the vegetation of the Jack Scott Nature ACOCKS, J.P.H. 1988. Veld types of South Africa. Reserve. Bothalia 11: 329 - 347. Memoirs of the botanical Survey of South Africa COETZEE, B.J. 1975. A phytosociological classifica- 57: 1-146. tion of the Rustenburg Nature Reserve. Bothalia BARKMAN, J.J., J. MORAVEC & S. RAUCHERT. 1964. 11: 561-580. Kritische Bemerkungen und Vorschläge zur COETZEE, B.J. & M.J.A. WERGER. 1975. On associa- quantitativen Vegetationanalyse. Acta Botanica tion—Analysis and the classification of plant Neerlandia 13: 394-449. communities. Vegetatio 30: 201-206.

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