van staden.qxd 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- denkamp@.up.ac.za) ISSN 0075-6458 59 Koedoe 48/2 (2005) Introduction The primary objective when proclaiming a national park is to conserve parts of pristine natural ecosystems for future generations (Gertenbach 1987). Marakele National Park (MNP) covers an area of 290.51 km² in the southwestern part of the Limpopo Province and is managed as a national park since 1988, but was officially proclaimed a national park on 11 February 1994. In order to manage and conserve any conser- vation area, a profound knowledge of the ecology is a prerequisite, and to achieve that prerequisite, an inventory of the biotic and the abiotic components of that national park must be undertaken (Edwards 1972). The “natural systems” as they occur today cannot be viewed and conserved as “natural” any more, because of the influence of man. Thus, management recommendations can only be made on the basis of interpreted ecological knowledge to restore the balance of the orig- inal natural system. The influence of man- agement recommendations on the system must also be regularly monitored to deter- mine if the aims that were set have been achieved satisfactorily. Efficient monitoring systems also depend upon an inventory of the biotic and abiotic components (Gertenbach 1987). The primary aim of this study was to classify and describe the vegetation of the Marakele National Park in the Waterberg of the Limpopo Province (Van Staden 2003). The purpose of the classification can be described as to form an inventory as a basis of manage- ment. Study area The study area covers 29051 ha in the south- western part of the Limpopo Province of South Africa, between 27°30'E–27°45'E and 24°15'–24°30'S. The position of the park in relation to towns is shown in Fig. 1 (Van Staden 2003). The underlying parent rock of the study area (SACS 1980) is sandstone of the Matlabas Subgroup, Aasvoëlkop Formation in the south-western and southern parts; with shale and mudstone of the Matlabas Subgroup, Aasvoëlkop Formation, Groothoek Mud- stone Member; a conglomerate outcrop of the Matlabas Subgroup, Aasvoëlkop Forma- tion in the west; and with the biggest part of the study area consisting of sandstone of the Kransberg Subgroup, Sandriviersberg For- mation. The soils that have developed on the parent materials range from shallow to deep sandy soils on sandstone and clayey soils on dia- base and mudstone (Van Staden 2003). The rainfall varies from 556–630 mm per annum and occurs mainly during the summer months. The study area experiences warm, wet summers with average daily tempera- tures of 32 ºC and cool, dry winters with frost in the low-lying areas (Van Staden 2003). MNP is situated mainly in the Waterberg Moist Mountain Bushveld (Low & Rebelo 1996) in the Savanna Biome (Rutherford & Westfall 1994). The vegetation of the study area includes Acocks’ (1988) Sour Bushveld (Veld Type 20), Mixed Bushveld (Veld Type 18), Sourish Mixed Bushveld (Veld Type 19) and North-Eastern Mountain Sourveld (Veld Type 8). This Sour Bushveld is listed by Edwards (1972) as one of 52 of South Koedoe 48/2 (2005) 60 ISSN 0075-6458 Fig. 1. A map indicating the location of the study area in relation to towns. African Veld Types that is extremely lacking in conservation. The Sour Bushveld covers 18306 km2, occurring in mountainous areas in the previous Transvaal Province (Coetzee 1975; Coetzee et al. 1981). Previous plant ecological studies in the Sour Bushveld (Acocks 1988) include those by Van Vuuren & Van der Schijff (1970), Coetzee (1975); Coetzee et al. (1981), Westfall (1981) and Westfall et al. 1985. Other plant ecological work of interest in related vegetation types include those of Theron (1973) who described the vegetation of the Loskopdam Nature Reserve; Van der Meulen (1979) who described the vegetation of the bushveld south of the Waterberg; Van Rooyen (1983) who described the vegetation of Roodeplaat Dam Nature Reserve; and Brown (1997) who described the vegetation of Borakalalo Game Reserve. Methods Analysis Methods applied are described in detail by Van Staden (2003) and are only summarised here. Stereo aerial photographs on a scale of 1:50 000 (Task 874 of 1984) were used to delineate homogeneous units on the basis of physiography and physiognomy (Bre- denkamp & Theron 1978; Westfall 1981; Gertenbach 1987). A total of 130 sample plots were surveyed through- out the study area. The sample plot location was determined by means of stratified-random sampling (Westfall 1981). The number of sample plots for each delineated physiographic-physiognomic unit was determined according to the size of each delin- eated unit. Termitaria and riparian vegetation was not included in the placing of the sample plots, due to their limit- ed size. Additional sample plots were identified for the termitaria and riparian vegetation and sampling was done in these vegetation types. A sample plot size of 10 m x 20 m was fixed and used throughout the study area. This size is consid- ered adequate for surveys in savanna vegetation by Coetzee (1975), Coetzee et al. (1976), Westfall (1981), Van Rooyen (1983) and Gertenbach (1987). At each sample plot a list of all the species present, was compiled. A cover-abundance value was given to each species according to the Braun-Blanquet cover-abundance scale, as given by Mueller-Dom- bois & Ellenberg (1974) and Werger (1974), and adapted by Barkman et al. (1964). The following habitat information was recorded at each sample plot—symbols are used in the synoptic table: land type was read from the land type map (Land Type Survey Staff 1988); soil forms were clas- sified in accordance with MacVicar et al. (1977); the altitude of each sample plot was recorded using an altimeter and is given in metres; the slope of the ter- rain of each sample plot was measured in degrees, using an optical clinometer. The following classification of slope units (Westfall 1981) were used in this study: Symbol Description Class L level 0.00° -3.49° G gentle 3.50° - 17.62° M moderate 17.63°- 36.39° S steep 36.40° 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 - North; S - South; NE - Northeast; SW - Southwest; E - East; W - West; SE - Southeast; NW - Northwest. The surface rock cover in each sample plot was esti- mated as a percentage stones (> 20 mm diameter), boulders and rocky outcrops. The following five classes were used, based on its potential influence on mechanical use (ploughing) (Van der Meulen 1979; Westfall 1981): Symbol Class Description O < 1 % No limitation on mechanical utilisation L 1–4 % Low limitation on mechanical utilisation M 5–34 % Moderate limitation on mechanical utilisation H 35–84 % High limitation on mechanical utilisation V 85–100 % No mechanical utilisation possible Synthesis The classification is based on the Braun-Blanquet method of vegetation classification, discussed in detail by Westhoff & Van der Maarel (1980), Mueller-Dombois & Ellenberg (1974) and Werger (1974). Data were captured on the mainframe com- puter of the University of Pretoria, in the BBNEW software package. It was then exported to be used in the software package BBPC (Bezuidenhout et al. ISSN 0075-6458 61 Koedoe 48/2 (2005) Koedoe 48/2 (2005) 62 ISSN 0075-6458 Table 1 Synoptic table of the major plant communities of Marakele National Park For symbols see text, constancy values = percentage Community # 1 2 3 4 5 Land type Ib/Ad Ad Ib/Fa Fa Ib/Fa Altitude (x 100 = m) 11-17 12-15 15-20 13-16 13-14 Slope G,L G,L M M L Aspect All N S,E N N,E Surface rock cover O,H O,M H H O Species Group A Pappea capensis 59 Mimusops zeyheri 59 Olea europaea subsp. africana 55 Cussonia paniculata 52 33 Diospyros whyteana 52 Euphorbia ingens 52 Rhus leptodictya 45 Cryptolepis transvaalensis 45 Podocarpus latifolius 38 Maytenus undata 38 Zanthoxylum capense 34 Myrsine africana 31 Canthium gilfillanii 31 Calpurnia aurea 31 Olea capensis 31 Ficus sur 31 Grewia occidentalis 31 Species Group B Acacia caffra 65 Elionurus muticus 61 Faurea saligna 57 Aristida congesta 57 Solanum incanum 48 Dichrostachys cinerea 43 Pterocarpus rotundifolius 43 39 Eragrostis capensis 39 Sida dregei 39 Bewsia biflora 39 Pogonarthria squarrosa 39 Eragrostis lehmanniana 39 Perotis patens 35 Vernonia oligocephala 39 Species Group C Eragrostis curvula 31 65 Dombeya rotundifolia 52 65 33 Euclea crispa 41 43 Grewia flavescens 34 39 Berchemia zeyheri 48 39 Ziziphus mucronata 38 35 Species Group D Panicum natalense 75 Anthospermum hispidula 64 Urelyterum agropyroides 57 Thesium utile 52 Rhynchosia monophylla 50 Acalypha angustata 50 Protea caffra 48 Aeschynomene rehmannii 45 Indigofera hedyantha 43 Monocymbium ceresiiforme 43 38 Tristachya rehmannii 41 Cheilanthus hirta 39 Dicoma anomala 39 Vernonia galpinii 39 Xerophyta retinervus 36 Chaetacanthus costatus 36 Senecio venosus 36 Rhynchosia nitens 34 Indigofera mollicoma 34 Pentanisia angustifolia 32 Parinari capensis 32 Species Group E Eragrostis racemosa 70 82 Trachypogon spicatus 30 80 Bulbostylis burchellii 30 73 Themeda triandra 78 59 Gnidia capitata 43 34 Species Group F Setaria lindenbergiana 100 Rhoicissus revoilii 89 Hypoestes forskaolii 78 Pseudolachnostylis maprouneifolia 78 Strychnos pungens 72 Diplorhynchus condylocarpon 67 Ochna pulchra 67 Elephantorrhiza burkei 61 Littonia modesta 61 Stylochiton natalense 56 Maytenus tenuispina 56 Lantana rugosa 50 Tephrosia rhodesica 44 Cryptolepis oblongifolia 44 Tapiphyllum parvifolium 44 Kalanchoe paniculata 39 Apodytes dimidiata 39 Aristida scabrivalvus 33 Turraea obtusifolia 33 Ancylbotrys capensis 33 Talinum caffrum 33 Species Group G Englerophytum magalismontanum 43 100 Aristida transvaalensis 50 61 Rhynchosia totta 52 56 Loudetia simplex 86 50 Tephrosia longipes 68 50 Stachys natalensis 36 50 Sphenostylis angustifolia 45 33 Rhus dentata 45 33 Species Group H Burkea africana 43 94 Combretum molle 30 94 Lannea discolor 43 83 Ozoroa paniculosa 30 50 Table 1 (continued) Community # 1 2 3 4 5 Land type Ib/Ad Ad Ib/Fa Fa Ib/Fa Altitude (x 100 = m) 11-17 12-15 15-20 13-16 13-14 Slope G,L G,L M M L Aspect All N S,E N N,E Surface rock cover O,H O,M H H O ISSN 0075-6458 63 Koedoe 48/2 (2005) Phyllanthus parvulus 35 78 Raphionachne galpinii 43 56 Setaria sphacelata 78 44 Heteropogon contortus 87 44 Vitex rehmannii 39 61 Species Group I Andropogon schirensis 43 80 61 Commelina africana 39 64 100 Diheteropogon amplectans 30 73 67 Schizachyrium sanguineum 30 50 33 Fadogia homblei 39 57 50 Melinis repens 78 36 72 Brachiaria serrata 70 39 44 Species Group J Vangueria infausta 34 74 55 94 Asparagus transvaalensis 38 39 43 56 Pellaea calomelanos 31 30 50 89 Cheilanthus viridis 38 56 Species Group K Xyris capensis 88 Andropogon huilensis 88 Miscanthus junceus 63 Fuirena pubescens 56 Monopsis decipiens 50 Helichrysum aureonitens 50 Ischaemum fasciculatum 44 Aristida junciformis 44 Ascolepis capensis 44 Sebaea leiostyla 44 Hypericum lalandii 44 Syzygium cordatum 38 Verbena bonariensis 38 Cyperus thorncroftii 38 Cliffortia linaerifolia 31 Arundinella nepalensis 31 Drosera madagascariensis 31 Results The vegetation composition of the study area is summarised in a synoptic table (Table 1). All species with a constancy value of more than 30 % in any community were included in the synoptic table. This table therefore shows the most frequently found species of each major community and sum- marises the relationship between the five major plant communities recognised. Classification The following major communities were identified: A. Forest Communities: 1. Olea europaea subsp. africana - Diospyros whyteana Major Community B. Sour Bushveld and North Eastern Mountain Sourveld Communities: 2. Acacia caffra-Heteropogon contortus Major Community 3. Protea caffra-Loudetia simplex Major Community 4. Burkea africana-Setaria lindenbergiana Major Community C. Wetland Communities: 5. Andropogon huilensis-Xyris capensis Major Community. Description of the Major Communities A Forest Communities 1. Olea europaea subsp. africana-Diospyros whyteana Major Community This major community occurs as forests in the kloofs, as dense bush clumps on south and east facing slopes and as bush clumps on termitaria. The kloofs are the least exposed of the geomorphology classes found in the study area, with water in the spruits. The species composition of the Olea europaea subsp. africana-Diospyros whyte- ana Major Community is given in Table 1. These forests and dense bush clumps have Table 1 (continued) Community # 1 2 3 4 5 Land type Ib/Ad Ad Ib/Fa Fa Ib/Fa Altitude (x 100 = m) 11-17 12-15 15-20 13-16 13-14 Slope G,L G,L M M L Aspect All N S,E N N,E Surface rock cover O,H O,M H H O 1996), After a TWINSPAN analysis (Hill 1979), the output of the resulting classification was imported into a spreadsheet, for refinement by Braun-Blan- quet procedures (Behr & Bredenkamp 1988; Bre- denkamp & Brown 2003). The final classification of the relevés was then interpreted for identification of major communities, where-after a synoptic table was compiled by calculating the percentage constan- cy of each species in each of the major communities identified. The result is given in Table 1. the following diagnostic species, all being trees or shrubs (Species Group A, Table 1): Pappea capensis, Mimusops zeyheri, Olea europaea subsp. africana, Cussonia panicu- lata, Diospyros whyteana, Euphorbia ingens, Rhus leptodictya, Cryptolepis trans- vaalensis, Podocarpus latifolius, Maytenus undata, Zanthoxylum capense, Myrsine africana, Canthium gilfillanii, Calpurnia aurea, Ficus sur, Olea capensis and Grewia occidentalis. Although various different plant communi- ties can be recognised within these forests and bush clumps (Van Staden 2003), species such as Podocarpus latifolius, Mimusops zeyheri, Diospyros whyteana, Canthium gil- fillanii and Olea capensis are prominent throughout the range of this vegetation. Grass and forb species are scanty under the dense forest vegetation, only Eragrostis curvula, Asparagus transvaalensis, Pellaea calomelanos and Cheilanthes viridis occurred with a constancy of more than 30 %. The Olea europaea subsp. africana-Diospy- ros whyteana Major Community is floristi- cally related to the Sour Bushveld communi- ties (Communities 2, 3 and 4) within the park (Species Group J), but shows particular affinity to the Acacia caffra-Heteropogon contortus Major Community through Species Group C. Coetzee (1975) described a related commu- nity from the Rustenburg Nature Reserve as Hypoestes verticillaris-Mimusops zeyheri Forests, and Westfall (1981) described a sim- ilar community from the Farm Groothoek in the Waterberg as Kloof Forest Communities on moderately deep soils in moist, sheltered habitats. Du Preez et al. (1991) classified this type of forest as typical Afromontane Forests, where these forests occur in specific niches in deep valleys, protected gorges, crevices and ravines along the eastern and western slopes of the Drakensberg mountain range. Coetzee et al. (1976) described termitaria bush clump communities from the Nylsvley Nature Reserve, which have diagnostic species similar to the bush clump communi- ties on termitaria in the Marakele National Park. B Sour Bushveld and North Eastern Mountain Sourveld Communities 2. Acacia caffra-Heteropogon contortus Major Community This major community represents a Sour Bushveld (Acocks 1988) with open to dense savanna typically found on the slopes of the Waterberg, and more open woodland or grassland on the summits. This open to closed woodland vegetation is found on gradual to moderately steep slopes (mainly footslopes) of outcrops, hills, ridges and mountains, which are widely distributed over Marakele National Park. It is restricted to shallow and rocky soils with a relatively high nutritional status (Coetzee 1975), derived from diabase and also from quartzite (Bezuidenhout et al. 1994). Many of the sample plots were located on diabase, which forms the substrate of this major community. On the lower, less rocky slopes with deeper soils the Acacia caffra-dominated vegetation often grades into communities of the plains forming a mixed thornveld, as also described from the Rustenburg Nature Reserve (Coet- zee 1975). On the other hand, the most mesic Acacia caffra-dominated vegetation shows some affinity to higher altitude Protea caffra-dominated vegetation that normally occurs at cooler sites (Species Group E, Table 1), at altitudes above the Acacia caf- fra-dominated vegetation, as also reported by Coetzee (1975). The soils are mainly shallow, of the Mispah or Glenrosa Form, though deeper soils of the Hutton or Clovelly Form are encountered at the foot of the mountain slopes (MacVicar et al. 1977). The soil depth varies from 100 mm to more than 1000 mm (Land Type Survey Staff 1988). The species composition of the Acacia caf- fra-Heteropogon contortus Major Commu- nity is given in Table 1. This major commu- nity is differentiated from the other major communities in the area by the following Koedoe 48/2 (2005) 64 ISSN 0075-6458 diagnostic plant species (Species Group B, Table 1): Acacia caffra, Faurea saligna, Dichrostachys cinerea, Pterocarpus rotundi- folius, Elionurus muticus, Aristida congesta, Eragrostis capensis, Bewsia biflora, Perotis patens, Eragrostis lehmanniana, Pogo- narthria squarrosa, Solanum incanum, Sida dregei and Vernonia oligocephala. Acacia caffra is mostly a dominant woody species, with other prominent species such as Faurea saligna, Burkea africana, Dichrostachys cinerea, Pterocarpus rotundi- folius, Vitex rehmannii, Vangueria infausta, Ozoroa paniculosa, Lannea discolor and Combretum molle also prominently present in certain plant communities. Quite often the woody species are grouped in bush clumps, resulting in a savanna with scattered individ- ual trees and bush clumps. The prominent grass species include Trachy- pogon spicatus, Themeda triandra, Setaria sphacelata, Schizachyrium sanguineum, Melinis repens, Heteropogon contortus, Eragrostis racemosa, Diheteropogon amplectens, Brachiaria serrata, Andropogon schirensis and the diagnostic Elionurus muti- cus and Eragrostis lehmanniana. The most frequently found forbs include Pel- laea calomelanos, Gnidia capitata, Fadogia homblei, Bulbostylis burchellii, Asparagus transvaalensis, Solanum incanum and Ver- nonia oligocephala. The Sour Bushveld (Acocks 1988) commu- nities that occur in the park (Communities 2, 3 and 4) are all floristically related, indicated by Species Group I, but also through Species Groups E, G and H. Descriptions of Acacia caffra-dominated vegetation are given by Bezuidenhout et al. (1988) from the Vredefort Dome area, Bezuidenhout & Bredenkamp (1991) from the North-West Province grasslands and Grobler et al. (2002) from the Johannesburg- Pretoria area. Particularly good examples (Eustachys mutica-Acacia caffra Wood- lands) are provided by Coetzee (1975) from the Rustenburg Nature Reserve, Bezuiden- hout et al. (1994) from the slopes of the hills and ridges in the North-West Province, and Coetzee et al. (1994, 1995) from the Maga- liesberg in the Pretoria area. Coetzee (1974) described Acacia caffra Savannas on diabase and in sheltered valleys from the Jack Scott Nature Reserve. Westfall (1981) described a similar community as Woodland on moder- ately deep soils in moderately exposed habi- tats, from the Farm Groothoek in the Water- berg. 3. Protea caffra-Loudetia simplex Major Community The Protea caffra-Loudetia simplex Major Community is representative of Acocks’ (1988) Sour Bushveld on moderately deep to deep soils in moderately exposed habitats. This mountain bushveld is found on gradual to steep rocky hills and ridges in the Marakele National Park. This vegetation type is prominent on the higher altitude slopes and crests. The soils are mainly very shallow, of the Mispah and Glenrosa Forms. The soil depth varies from 10 mm to 500 mm (Land Type Survey Staff 1988). The species composition of the Protea caf- fra-Loudetia simplex Major Community is given in Table 1. This major community is differentiated by the following diagnostic plant species (species group D, Table 1): Protea caffra, Urelyterum agropyroides, Tristachya rehmannii, Panicum natalense, Monocymbium ceresiiforme, Xerophyta retinervus, Vernonia galpinii, Thesium utile, Senecio venosus, Rhynchosia nitens, Rhyn- chosia monophylla, Pentanisia angustifolia, Parinari capensis, Indigofera mollicoma, Indigofera hedyantha, Dicoma anomala, Cheilanthus hirta, Chaetacanthus costatus, Anthospermum hispidula, Aeschynomene rehmannii and Acalypha angustata. This major community is dominated by the tree Protea caffra. Other prominent woody species include Vangueria infausta, Rhus dentata and Englerophytum magalismon- tanum. The most prominent grasses that are often found in this major community are Trachy- ISSN 0075-6458 65 Koedoe 48/2 (2005) pogon spicatus, Themeda triandra, Schizachyrium sanguineum, Melinis repens, Loudetia simplex, Eragrostis racemosa, Diheteropogon amplectens, Brachiaria ser- rata, Aristida transvaalensis and Andro- pogon schirensis. Many forb species are present in this vegeta- tion; the most frequently encountered include Tephrosia longipes, Stachys natalen- sis, Sphenostylis angustifolia, Rhynchosia totta, Pellaea calomelanos, Gnidia capitata, Fadogia homblei, Commelina africana, Bul- bostylis burchellii and Asparagus trans- vaalensis. Many authors described Protea caffra-domi- nated vegetation, mainly from Sour Bushveld and Bankenveld, including Coet- zee (1974, 1975) from the Magaliesberg area, Bredenkamp & Theron (1978) from the Suikerbosrand, Behr & Bredenkamp (1988), from the Witwatersrand, Bezuidenhout et al. (1994) from the Gatsrand area in North-West Province, Coetzee et al. (1995), Bredenkamp & Brown (1998a; 1998b) from the natural areas of the Western Metropolitan Local Council and Grobler et al. (2002) within var- ious natural open spaces in Gauteng. This emphasises the floristic relationships between Bankenveld, Sour Bushveld and the Drakensberg area (Bredenkamp & Brown 2003). 4. Burkea africana-Setaria lindenbergiana Major Community This major community represents Sour Bushveld (Acocks 1988). It occurs on gentle to moderately steep slopes of rocky sand- stone hills, where soils are litholitic and large rock boulders cover 53 % of the soil surface. Aspects are mostly northerly, and these slopes are warm and dry. The soils are main- ly of the Mispah and Glenrosa Forms. The soil varies from 10 mm to more than 500 mm in depth (Land Type Survey Staff 1988). This vegetation type is also found at the northernmost areas of the Bankenveld in the Pretoria and Rustenburg areas and is found at high altitudes on warm northern and northeastern aspects with gradual to very steep slopes (Bredenkamp & Brown 2003). The shallow soils are covered with large boulders and smaller rocks that in some cases provide moister microhabitats than the surrounding areas. The species composition of the Burkea africana-Setaria lindenbergiana Major Community is given in Table 1. This major community is differentiated by the following diagnostic plant species (species group D, Table 1): Tapiphyllum parvifolium, Strychnos pun- gens, Rhoicissus revoilii, Pseudolachnostylis maprouneifolia, Ochna pulchra, Maytenus tenuispina, Elephantorrhiza burkei, Dip- lorhynchus condylocarpon, Cryptolepis oblongifolia, Apodytes dimidiata, Acy- lobotrys capensis, Setaria lindenbergiana, Aristida scabrivalvus, Turraea obtusifolia, Tephrosia rhodesica, Talinum caffrum, Stylo- chiton natalense, Littonia modesta, Lantana rugosa, Kalanchoe paniculata and Hypoestes forskaolii. Apart from the above prominent diagnostic species, other prominent woody species include Vitex rehmannii, Vangueria infausta, Rhus dentata, Ozoroa paniculosa, Lannea discolor, Englerophytum magalismontanum, Combretum molle and Burkea africana. Grass species that are prominent in this com- munity include Setaria sphacelata, Schizachyrium sanguineum, Melinis repens, Loudetia simplex, Heteropogon contortus, Diheteropogon amplectens, Brachiaria ser- rata, Aristida transvaalensis and Andro- pogon schirensis. Forbs include Tephrosia longipes, Stachys natalensis, Sphenostylis angustifolia, Rhyn- chosia totta, Pellaea calomelanos, Fadogia homblei, Cheilanthus viridis and Asparagus transvaalensis. Coetzee (1975) described a similar commu- nity as the Barleria bremekampii- Diplorhynchus Tree Savanna from Rusten- burg Nature Reserve, and Westfall (1981) described a similar community as a Wood- land, representative of Acocks’ (1988) Sour Koedoe 48/2 (2005) 66 ISSN 0075-6458 Bushveld, on moderately deep to deep soils in moderately exposed habitats from the Waterberg. Further examples of this vegetation were described from the scarps and slopes of the Jack Scott (Coetzee 1974), from the warm temperate mountain bushveld within the Pre- toria-Witbank-Heidelberg areas by (Coetzee et al. 1993), from the Melville koppies Nature Reserve (Ellery 1994), from the nat- ural areas of the western local Metropolitan Council of Gauteng by Bredenkamp & Brown (1998a) and within various natural open spaces in Gauteng by Grobler et al. (2002). 5. Andropogon huilensis-Xyris capensis Major Community Major Many streams arise in seepage areas, for instance on mountain slopes within Marakele National Park. These are seasonal- ly or perennially waterlogged sponges, with vegetation dominated by sedges and other hygrophilous angiosperms and mosses (Noble & Hemens 1978). This major wet- land community occurs along streams and tributaries of the Matlabas, Mamba and Sterkstroom rivers and shallow submerged marshy areas or sponges. The soils are wet, mainly of the Avalon, Hut- ton, Katspruit, Oakleaf and Westleigh Forms, derived from sandstone of the Sandriviersberg Formation (De Vries 1968, 1969). The soil depth varies between 300–1 200 mm (Land Type Survey Staff 1988). The species composition of this wetland Major Community is given in Table 1. This major community is differentiated by the fol- lowing diagnostic plant species (Species Group K): Syzygium cordatum, Cliffortia linaerifolia, Andropogon huilensis, Aristida junciformis, Arundinella nepalensis, Ischaemum fasciculatum, Miscanthus junceus, Ascolepis capensis, Cyperus thorn- croftii, Drosera madagascariensis, Fuirena pubescens, Helichrysum aureonitens, Hyper- icum lalandii, Monopsis decipiens, Sebaea leiostyla, Verbena bonariensis and Xyris capensis. Coetzee (1975) described similar communi- ties from the Rustenberg Nature Reserve as the Aristida junciformis-Arundinella nepal- ensis Grassland and the Pteridium aquil- inum-Phragmites mauritianus Reedswamp. Discussion The vegetation of the study area was classi- fied in a hierarchical, plant sociological sys- tem by using the Braun- Blanquet technique. The floristic data from 130 relevés were classified to identify five major plant com- munities. These plant communities were ecologically interpreted by habitats. These major plant communities should form the basis for a management plan, and should also be the basis for a more detailed classification of the vegetation of this national park. According to White (1978) the majority of the tree species in South African Afromon- tane forests are widespread. Amongst them are, inter alia, Podocarpus latifolius, Olea capensis and Myrsine africana that occur as diagnostic species in the Olea europaea subsp. africana-Diospyros whyteana Major Community within the study area. These forests on the Waterberg have a clear floris- tic affinity to the Afromontane forests found in the Drakensberg (Du Preez et al. 1991), indicating a much more widespread distribu- tion of forests at previous times, with the Waterberg forests being a western relict (Bredenkamp et al. 2002). The above men- tioned assemblage of species could almost be used to define the Afromontane region as a whole. Not one species occurs throughout, but most species of the assemblage are pre- sent on virtually every “island” of Afromon- tane forest in the region. It is interesting to note that the vegetation on the termitaria have species similar to the for- est communities, and are therefore included into this major community. The greater water holding capacity of the soils worked over by the termites renders these habitats suitable for forest development (Coetzee et al. 1976). ISSN 0075-6458 67 Koedoe 48/2 (2005) The Acacia caffra-Heteropogon contortus Major Community on footslopes, Protea caf- fra-Loudetia simplex Major Community on cooler southerly facing slopes and the Burkea africana-Setaria lindenbergiana Major Community on the warmer northerly facing slopes have a clear floristic relation- ship with similar plant communities in the Bankenveld. The relationships between Sour Bushveld, Sourish Mixed Bushveld and Bankenveld woodland communities (Acocks 1988) are clearly indicated by Bredenkamp & Brown (2003) and can further be demon- strated from studies by Van Vuuren & Van der Schijff (1970), Du Plessis (1972), Theron (1973), Coetzee (1975), Van der Meulen (1979), Westfall (1981) and Coetzee (1993). Furthermore, Coetzee (1993), O’Connor & Bredenkamp (1997) and Bre- denkamp & Brown (2003) indicated that the Bankenveld vegetation also shows an affini- ty to the Drakensberg flora. This relationship may also be seen in the presence of the Afromontane forests in the Drakensberg, Bankenveld and Waterberg. In the cooler, southern parts of the Banken- veld, Acacia caffra-dominated vegetation is usually found on lower and warmer north- facing slopes, e.g., in the Vredefort Dome area, though in the warmer northern parts e.g. the Marakele National Park, it may be found at higher altitudes, on crests or south- facing slopes (Coetzee 1975; Bezuidenhout et al. 1994; Coetzee et al. 1995). Due to the complex mosaic distribution pat- tern of the sub-communities that are includ- ed under the major communities, a vegeta- tion map is more appropriate when the details of the sub-communities are pub- lished. References ACOCKS, J.P.H. 1988. Veld types of South Africa. Memoirs of the botanical Survey of South Africa 57: 1-146. BARKMAN, J.J., J. MORAVEC & S. RAUCHERT. 1964. Kritische Bemerkungen und Vorschläge zur quantitativen Vegetationanalyse. Acta Botanica Neerlandia 13: 394-449. BEHR, C.M. & G.J. BREDENKAMP. 1988. A phytosoci- ological classification of the Witwatersrand National Botanical Garden. South African Jour- nal of Botany 54: 525-533. BEZUIDENHOUT, H. 1992. Verslag oor die abiotiese komponent van die Kransberg Nasionale Park. Unpublished internal report. BEZUIDENHOUT, H., H.C. BIGGS & G.J. BREDENKAMP. 1996. A process supported by the utility BBPC for analysing Braun-Blanquet data on a personal computer. Koedoe 39(1): 107-112. BEZUIDENHOUT, H., & G.J. BREDENKAMP. 1991. The vegetation of the Bc Land Type in the western Transvaal grassland, South Africa. Phyto- coenologia 19: 497-518. BEZUIDENHOUT, H., G.J. BREDENKAMP & J.H. ELSEN- BROEK. 1988. Die plantegroei van die alka- ligraniet en aangrensende kwartsiet in die Vre- defortkoepel noordwes van Parys. Suid- Afrikaanse Tydskrif vir Natuurwetenskap en Tegnologie 7: 4-9. BEZUIDENHOUT, H., G.J. BREDENKAMP & G.K. THERON.1994. Syntaxonomy of the vegetation of the Fb land type in the western Transvaal grass- land, South Africa. South African Journal of Botany 60: 72-80. BREDENKAMP, G.J. & L.R. BROWN. 1998a. A vegeta- tion assessment of the open spaces in the West- ern Metropolitan local Council area. Pretoria: Ekotrust CC. BREDENKAMP, G.J. & L.R. BROWN.1998b. A vegeta- tion assessment of open spaces in the northern areas of the Northern Metropolitan Local Coun- cil. Pretoria: Ekotrust CC. BREDENKAMP, G.J. & L.R. BROWN. 2003. A reap- praisal of Acocks’ Bankenveld: Origin and diversity of vegetation types. South African Journal of Botany 69(1): 7-26. BREDENKAMP, G.J., F. SPADA & E. KAZMIERCZAK. 2002. On the origin of northern and southern hemisphere grasslands. Plant Ecology 163(2): 209-229. BREDENKAMP, G.J. & G.K. THERON. 1978. A syneco- logical account of the Suikerbosrand Nature Reserve. 1. The phytosociology of the Witwater- srand geological system. Bothalia 12: 513-529. 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. COETZEE, B.J. 1974. A phytosociological classifica- tion of the vegetation of the Jack Scott Nature Reserve. Bothalia 11: 329 - 347. COETZEE, B.J. 1975. A phytosociological classifica- tion of the Rustenburg Nature Reserve. Bothalia 11: 561-580. COETZEE, B.J. & M.J.A. WERGER. 1975. On associa- tion—Analysis and the classification of plant communities. Vegetatio 30: 201-206. Koedoe 48/2 (2005) 68 ISSN 0075-6458 COETZEE, B.J., F. VAN DER MEULEN, S. ZWANZIGER, P. GONSALVES & P.J. WEISSER. 1976. A phytosocio- logical classification of the Nylsvley Nature Reserve. Bothalia 12: 137-160. COETZEE, B.J., P. VAN WYK, W.P.D. GERTENBACH, A. HALL-MARTIN & S.C.J. JOUBERT. 1981. ‘n Plantekologiese verkenning van die Water- berggebied in die Noord-Transvaalse Bosveld. Koedoe 24:1-23. COETZEE, J.P. 1993. Phytosociology of the Ba and Ib land types in the Pretoria-Witbank-Heidelberg area. MSc thesis, University of Pretoria, Preto- ria. COETZEE, J.P., G.J. BREDENKAMP & N. VAN ROOYEN. 1993. The Sub-humid Warm Temperate Moun- tain Bushveld plant communities of the Pretoria- Witbank-Heidelberg area. South African Journal of Botany 59(6): 623-632. COETZEE, J.P., G.J. BREDENKAMP & N. VAN ROOYEN. 1994. An overview of the physical environment and vegetation units of the Ba and Ib land types of the Pretoria-Witbank-Heidelberg area. South African Journal of Botany 60(1): 49-61. COETZEE, J.P., G.J. BREDENKAMP & N. VAN ROOYEN. 1995. Plant communities of the Sub-humid Cool Temperate Mountain Bushveld in the Pretoria Witbank Heidelberg area. South African Journal of Botany 61: 114-122. DE VRIES, W.C.P. 1968-69. Stratigraphy of the Waterberg System in the Southern Waterberg area, Northwestern Transvaal. Annals of the geo- logical Survey in South Africa 43-56. DU PLESSIS, C.J. 1972. ‘n Floristies-ekologiese studie van die plaas Doornkop in die distrik Middelburg, Transvaal. MSc Thesis, University of Pretoria, Pretoria, DU PREEZ, P.J., G.J. BREDENKAMP & H.J.T. VENTER. 1991. The syntaxonomy and synecology of the forests in the eastern Orange Free State, South Africa. I. The Podocarpetalia latifolii. South African Journal of Botany 57: 198-206. EDWARDS, D. 1972. Botanical survey and agricul- ture. Proceedings of the Grassland Society of South Africa 7: 15-19. ELLERY, W.N. 1994. The vegetation ecology of Melville Koppies Nature reserve and Louw Geldenhuys View Site: Proposals for their Man- agement. Johannesburg: Botanical Society of South Africa. GERTENBACH, W.P.D. 1987. ‘n Ekologiese studie van die Suidelikste Mopanieveld in die Nasionale Krugerwildtuin. D.Sc proefskrif (Ongepub- liseer). Universiteit van Pretoria. GROBLER, C.H., G.J. BREDENKAM. & L.R.BROWN. 2002. Natural woodland vegetation and plant species richness of the urban open spaces in Gauteng, South Africa. Koedoe 45: 19-34. HILL, M.O. 1979. Twinspan - a Fortran program for arranging multivariate data in an ordered two way table by classification of individuals and attributes. Cornell University, Ithaca, New York. LAND TYPE SURVEY STAFF. 1988. Land types of the maps 2426 Thabazimbi, 2428 Nylstroom. Mem- oirs of the agricultural natural Resoures of South Africa 10: 1-431. LOW, A.B. & A.G. REBELO. 1996. Vegetation of South Africa, Lesotho and Swaziland. Pretoria: Department of Environmental Affairs and Tourism. MACVICAR, C.N., R.F. LOXTON, J.J.N. LAMBRECHTS, J. LE ROUX, J.M. DE VILLIERS, E. VERSTER, F.R. MERRY-WEATHER, T.H. VAN ROOYEN & H.J. VON M. HARMSE. 1977. Grondklassifikasie, 'n bi- nomiese sisteem vir Suid Afrika. Pretoria: Departement Lanbou Tegniese Dienste. MUELLER-DOMBOIS, D. & H. ELLENBERG. 1974. Aims and Methods of Vegetation Ecology. New York: Wiley. NOBLE, R.G. & J. HEMENS. 1978. Inland water ecosystems in South Africa a review of research needs. South African Natural Science Progress Report 34: 1-150. O’CONNOR, T.G.. & G.J. BREDENKAMP. 1997. Grass- land. Pp. 215-257. In: COWLING R.M., D.M. RICHARDSON & S.M. PIERCE (eds.). Vegetation of Southern Africa. Cambridge: Cambridge Uni- versity Press. Rutherford, M.C. & R.H. Westfall. 1994. Biomes of southern Africa: An objective categorization. Memoirs of the botanical Survey of South Africa 63: 1-94 SOUTH AFRICAN COMMISSION FOR STRATIGRAPHY (SACS). 1980. Statigraphy of South Africa. Part 1 (Comp. L.E. Kent). Lithostratigraphy of the Republic of South Africa, South West Africa/ Namibië and the Republics of Bophuthatwana, Transkei, and Venda. Pretoria: Government Printer. (Handbook for Geological Survey in South Africa 8.) THERON, G.K. 1973. ‘n Ekologiese studie van die Plantegroei van die Loskopdam - natuurreser- vaat Ongepubl. D.Sc - thesis, University of Pre- toria, Pretoria. VAN DER MEULEN, F. 1979. Plant sociology of the Western Transvaal Bushveld. South Africa: Syn- taxonomic and synecological study. Vaduz: Cramer. (Dissertationes Botanica 49.) VAN ROOYEN, N. 1983. Die Plantegroei van die Roodeplaatdam - natuurreservaat II. Die plant- gemeenskappe. South African Journal of Botany 2: 115 -125. VAN STADEN, P.J. 2002. An ecological study of the plant communities of Marakele National Park. M.Sc. thesis, University of Pretoria, Pretoria. VAN VUUREN, D.R.J. & H.P. VAN DER SCHIJFF. 1970. ‘n Vergelykende ekologiese studie van die plantegroei van ‘n noordelike en suidelike kloof ISSN 0075-6458 69 Koedoe 48/2 (2005) van die Magaliesberg. Tydskrif vir Natuurweten- skappe 10: 16-75. WERGER, M.J.A. 1974. On concepts and techniques applied in the Zürich-Montpellier method of vegetation survey. Bothalia 11(3): 309-323. WESTFALL, R.H. 1981. The plant ecology of the farm Groothoek, Thabazimbi District. M.Sc. thesis, University of Pretoria, Pretoria. WESTFALL, R.H., N. VAN ROOYEN & G.K. THERON. 1985. The plant ecology of the farm Groothoek, Thabazimbi District. 2. Classification. Bothalia 15: 655-688. WESTHOFF, V. & E. VAN DER MAAREL. 1980. The Braun-Blanquet approach. Pp. 287-399. In: R.H. WHITTAKER (ed.). Classification of plant com- munities. The Hague: Junk. WHITE, F. 1978. The Afromontane region. Pp. 465- 513. In: WERGER, M.J.A. (ed.). Biogeography and ecology of Southern Africa. The Hague: Junk. 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