152 S.AfrJ.Bot., 1994, 60(3): 152 - 161 A classification of the vegetation of the western Transvaal dolomite and chert grassland, South Africa

H. Bezuidenhout, * G.J. Bredenkamp and G.K. Theron Department of Botany, University of Pretoria, Pretoria 0002, Republic of South Africa

Received 21 October 1993; revised 1 March 1994

Relatively little is known about the vegetation of the western Grassland Biome in South Africa. The classification of the dolomite and chert grassland in the western Transvaal (Fa land type) forms part of a research programme on the syntaxonomic and synecological synthesis of the vegetation of the western Grassland Biome. Using a numerical classification technique (TWINSPAN) as a first approximation, the classification was refined by applying Braun-Blanquet procedures. The result is a phytosociological table from which two new alliances, six new associa­ tions, two new sub·associations and two new communities without syntaxonomic rank are recognized. The new syntax a are ecologically interpreted as well as described. Associated gradients in habitat were identified by using an ordination algorithm (DECORANA) . This study should contribute to the present knowledge and ecological under­ standing of the vegetation of the western Transvaal.

Relatief min inligting is oor die plantegroei van die westelike Grasveldbioam van Suid-Afrika beskikbaar. Die klassi­ fikasie van die plantegroei van die dolomitiese en chert-grasveld in die Wes-Transvaal (Fa-Iandtipe) vorm deel van die sintaksonomiese en sinekologiese sintese van die plantegroei van die westelike Grasveldbioom. 'n Numeriese tegniek (TWINSPAN) is as eerste klassifikasie van die floristiese data aangewend. Daarna is die Braun-Blanquet­ prosedure gevolg om twee nuwe alliansies, ses nuwe assosiasies, twee nuwe subassosiasies en twee nuwe gemeenskappe sonder range in 'n fitososiologiese tabel te identifiseer. Die nuwe sintaksons word ekologies ge"lnterpreteer en beskryf. Geassosieerde gradiente in habitat is deur toepassing van 'n ordeningsalgoritme (DECORANA) ge"ldentifiseer. Hierdie studie behoort 'n waardevolle bydrae tot die ekologiese kennis oar die plante­ groei van die Wes-Transvaal te lewer.

Keywords: Braun·Blanquet, Fa land type, Grassland Biome, vegetation classification.

* To whom correspondence should be addressed at: National Parks Board, Scientific Services, P.O. Box 110040, Hadison Park, Kimberley 8306, Republic of South Africa.

Introduction (Van Rooyen et al. 1981), in order to identify conservation Mentis and Huntley (1982) as well as Scheepers (1986) stated priorities. the necessity to determine the location and extent of the major This is a comprehensive vegetation classification of the vegetation types and subtypes within the Grassland Biome. As dolomite and chert area which includes the entire Fa land type part of a phytosociological research programme on the syn­ in the western Transvaal. This report forms part of the syn­ taxonomic synthesis of the vegetation of the Grassland Biome thesis of the vegetation of the western Grassland Biome of in South Africa, several studies have been carried out (Bezui­ South Africa and the results should contribute significantly to denhout 1988; Turner 1989; Kooij 1990; Myburgh 1990; Brey­ the ultimate aim of a phytosociological and syntaxonomical tenbach 1991; du Preez 1991; Eckhardt 1993). A mosaic of synthesis of the western Grassland Biome. land types occur in the study area (Figure 1). As land types represent an ecologically based stratification of the study area, Study area each land type is separately used to describe the vegetation of The study area is situated in the western part of the Highveld the western Transvaal grassland (Bezuidenhout, in prep.). In Agricultural Region in the Transvaal and is bounded by lati­ this paper, a description of the vegetation of the Fa land type in tudes 25°45' and 27°15' south and longitudes 24°45' and 28° the study area is presented. A broad, regional account of the 00' east (Figure 1). The Fa land type covers approximately vegetation of the Fa land type and a number of other land types 557320 ha of land, and the Land Type Survey Staff (1984) was given by Acocks (1953, 1988). Vegetation studies have estimated that 35% is unavailable for agronomy. The main been conducted in the Potchefstroom area (Louw 1951), Lich­ rock types which underly the Fa land type are dolomite and ten burg area (Morris 1973), Abe Bailey Nature Reserve (Van Wyk 1983) and the Mooi River catchment area (Bezuidenhout chert of the Chuniespoort Group (Transvaal Sequence). The 1988). A part of the dolomite or Klipveld vegetation was slightly undulating plains are dissected by prominent rocky described by Bezuidenhout and Bredenkamp (1990). In the chert ridges (Land Type Survey Staff 1984). More than 500/0 of Jack Scott Nature Reserve, which is situated adjacent to the the main soil types are relatively shallow (between 50 and 150 north-eastern boundary of the study area, the dolomite vegeta­ mm) and rocky (dolomite and chert) with the dominant soil tion was investigated by Coetzee (1974) and Scogings and forms Mispah, Glenrosa and shallow Hutton (Land Type Sur­ Theron (1990). vey Staff 1984). Proper land use requires a classification of the vegetation The vegetation is distributed in a complex mosaic pattern, and is not dominated by a single or a few species. It rather Some of the data for this research were collected by H.B. while represents a mosaic of many co-dominants (Louw 1951). Ac­ employed by the Department of Agricultural Development, Grass­ cording to the Acocks (1988) classification, the largest part of land Research Centre, Private Bag X05, Lynn East, 0039 Republic the Fa land type vegetation is represented by the western vari­ of South Africa. ations of the Bankenveld (veld type 61a) while the remaining S.AfrJ.Bot.,1994,60(3) 153

parts are represented by Cymbopogon - Themeda veld (veld rockiness of the soil surface, was noted (Figure 3). The soil type 48b). nomenclature follows the classification of MacVicar et ai. (1977). The Fa land type is drained by the Schoonspruit and the The floristic data were classified using TwINS PAN (Hill 1979a). Mooi River and their tributaries. The sources of both river The result from the classification was refined by Braun-Blanquet systems are springs which originate from the dolomite water procedures (Mueller-Dombois & Ellenberg 1974). The final result reservoir in the northern central part of the Fa land type (Du of the classification procedure is represented in a phytosocio­ Toit 1954). logical table (Table 1). An ordination technique, DECORANA (Hill Altitudes vary between 1356 and 1450 m above sea-level. 1979b), was also applied to the floristic data to illustrate floristic According to the Koppen climate classification system, two relationships between communities to detect possible climatic regions occur over the study area, namely a cool dry gradients in and between communities and to detect possible steppe with summer rains and a warm temperate climate with habitat gradients associated with vegetation gradients (Figure 4). summer rains. The rainfall exceeds 600 mm per year (Lichten­ Taxa names conform to those of Gibbs Russell et ai. (1985, 1987). burg 601 mm, Potchefstroom 625 mm and Carletonville 670 New syntaxa are described and formal syntaxonomy, in accord­ mm) which is relatively high for the study area (Bezuidenhout, ance with the Code of Phytosociological Nomenclature (Barkman in prep.). The summer temperatures are high, with the mean et al. 1986), is applied to the classification. maximum monthly temperatures exceeding 32°C during October to January, while the mean minimum monthly temp­ Results eratures are below _1°C during May to September. The Classification winters are severely frosty (Weather Bureau 1988). In the phytosociological table, two alliances, six associations, Methods two sub-associations and two communities without syntaxo­ nomic rank are recognized (Table 1). No syntaxonomic rank is Within the Fa land type, which was used as a first stratification formally assigned to the latter two communities because too unit in the investigation of the western Transvaal grasslands, terrain types were used for subsequent finer stratification. The little information is available about them, and their syntaxo­ nomic position is still uncertain. The hierarchical classification term land type is used in a land·use classification system of the syntaxa is as follows: describing a homogeneous terrain with regard to soil pattern and climate (Land Type Survey Staff 1984). The following terrain 1. Grewio flavae - Rlwion pyroidis types were recognized in the Fa land type: plateau (I), midslope 1.1 Rhoo lanceae - Acacietum eriolobae (3), footslope (4a), floodplain (4b) and drainage line (5) (Figure 1.2 Ziziplw mucronatae - Acacietum karroo 2) . Releves were compiled in 91 stratified sample plots. Plot sizes 1.3 Digitaria eriantha - Rhus pyroides Shrubveld (com­ were fixed at 16 m2 for the grassland vegetation and 100 m2 for munity without syntaxonomic rank) the woody vegetation (Bredenkamp & Theron 1978). For every 2. Trachypogono spicati - Diheteropogonionamplectentis plant species present in the sample plot, a cover-abundance value 2.1 Loudetio simplicis- Diheteropogonetumamplectentis was estimated according to the Braun-Blanquet scale (Mueller­ 2.2 Alloteropsido semialatae - Tristachyetum leucothricis Dombois & Ellenberg 1974). Height and canopy cover for the 2.3 Cymbopogon excavatus - Diheteropogon amplectens tree, shrub and herbaceous layers were additionally recorded in Grassland (community without syntaxonomic rank) each sample plot, and average values calculated for each plant 3. Cymbopogono plurinodis - Eragrostidetumgummifluae community. Environmental information such as rock type, terrain 3.1 Cymbopogono plurinodis - Eragrostidetum gummifluae type and soil type as well as soil depth and an estimation of aristidetosumcanescentis

Armoe.dsv .lakte Vryburg

10

Figure 1 The location of the Fa land type (black areas) in the western Transvaal, South Africa (adapted from the Land Type Series 1979). 154 S.-Afr.Tydskr.Plantk., 1994,60(3)

3.2 Cymbopogono plurinodis - Eragrostidetum gummifluae classes, namely woodland and grassland, can easily be distin­ eragrostidetosumsuperbae guished (Table 1, Figure 4). The hierarchical classification of 4. Paspalo dilatati - Hyparrhenietum hirtae the vegetation emphasizes the correlation between habitat and communities in the Fa land type, as was also noted by Sco­ Description of the syntaxa gings and Theron (1990) in the Jack Scott Nature Reserve, as In the vegetation of the dolomite and chert grassland in the well as the relationships between communities (Figure 3). The western Transvaal (Fa land type), two broad physiognomic vegetation is often overgrazed and burned, as indicated by the

Terrain types 4a. 3 1 3 4a. 5 41:>

Syntaxa 1_1 3_1 1_3 2_1 2_2 2_3 1_2 4 3_2

1 450 m ------I I 1 356 m ------I -

Soil Hu (50) Hu (47) Rock (67) MsjGs (72) MsjGs (88) Hu (50) WejVa (40) Hu (50) MsjGs (33) MsjGs (46) Rock ( 8) MsjGs Hu (40) MsjGs (33) types MsjGs (33) Rock (16) (25) Rock (17) Rock ( 7) Hu (12) Hu ( 4) Rock (25) MsjGs (14) Rock (17)

Figure 2 The location of the syntax a on the topographical terrain types (adapted from Land Type Survey Staff 1984) within the dolomite and chert Grassland in the western Transvaal, South Africa (all abbreviations and numbers are explained in the text).

Vegetation Fa land ty~ I I I Upland stony plains Bottomlands Terrain types 1 & 3 Terrain types 4 & 5 I I I Grasslaoo Shrubland I $

Chert oob:rops IXllomite Ruins sinkholes Footslo~s (4a) Floodplains (4b) Drainage lines (5) aoo ridges Kidslo~ (3) DialOOoo Clay content Clay content Rocks or stones No rocks Soil depth diggings < 10\ > 20% > 5% or stones 250 -1 200 rom Ilidslope (3) Soil depth Soil depth Soil depth Soil depth soil depth 300->1200 IIIlI 250-1200 mm 250-1200 II1II 300-800 mm Very rocky !.ess rocky 2 500->1 200 rom Plateau (1) l!idslo~ (3)

soil depth Soil depth soil depth 50-150 mm 300-900 rom 100-150 IliII

2.1 2.2 2.3 3.1 1.3 1.1 1.2 3.2 Figure 3 A dendrogram to illustrate the habitat relationships of the syntaxa, of the dolomite and chert Grassland in the western Transvaal, South Africa (all abbreviations and numbers are explained in the text). S.AfrJ.Bot.,1994,60(3) 155

500 .,...'"Q) iJ >. .-< Q) .,...> +J .-< 400 '"Q) <>:

r 300 Q) :l N +J'" .,... 0"' X :c 3 200 1 :J..

Q) +J'" 100 +J Q) :J >. .-< Q) .>,... +J -< "'Q) <>: 0 100 200 300 400 500 600 X :J..

Woodland • structure ~ Grassland 250 - > 1 200 DUD .. Soil depth • 50 - BOO mID Figure 4 The relative positions of the major syntaxa (numbers refer to those in the text) along the first two axes of the ordination of the dolomite and chert Grassland in the western Transvaal, South Africa (all abbreviations are explained in the text).

Table 1 A phytosociological table of the vegetation of the dolomite and chert in the western Transvaal grassland, South Africa (all numbers are ex­ plained in the text)

441666242 555451114 lllllll 55555555450144 1121224 4411222 111111 22244550111111411111122224 011222 Sample mmbers 004888405 222338444 2288948 34232323330544 7909013 4323010 222233 32734232335589466668903574 224017 210132931 345461344 0229120 38209584699615 6972109 0317875 578989 09553178560770645678809042 961663

Syntaxa numbers I 1.1 I /2 I 1.3 I 2.1 122.2 I 2.3 I 3.21 3.3

Species group A

Acacia eri oloba ++4+1+5 4 Rhus lancea 22+222 2 Pavonia burchellii +++ + Eragrostis bitlora 2 4 Eragrostis capensis +2 + +++ + + Lantana rugosa + + + setaria verticillata 4

Species group B

Acacia karroo 1221+322+ + ziziplills mucronata +111++++ Sporobolus fimbriatus 21l ++ Tarchonanthus camphoratus +++ Acacia hereroensis 12 Acacia calfra 2+

Species group C

Haytenus heterophy lla tt+ + ++ + + Protasparagus alricanus + +tt + 1 Ehretia rigida 1+ ++++ Brachiaria nigropedata ++ ++ 156 S.-Afr.Tydskr.Plantk., 1994,60(3)

Table 1 Continued

441666242 555451114 1111111 55555555450144 1121224 4411222 111111 22244550111111411111122224 011222 Sample numbers 004888405 222338444 2288948 34232323330544 7909013 43230 10 222233 32734232335589466668903574 224017 210132931 345461344 0229120 38209584699615 6972109 0317875 578989 09553178560770645678809042 961663

Syntaxa mllnbers 1.1 1.2 I 1.3 \ 2.1 I \2 I 2.3 \ 3.21 J.J

Species group D

Rhus pyroides J+ ++ + ++1+++ ++ +++4+22 Diospyros lycioides 1+3++ +++ +++ + 2+ ++ Prot asparagus suaveolens +12 H +++ + + +++1 + ++++ + + Grewia flava + ++1+1+ + + +H + HH + Protasparagus laricinus ++ + + ++++ +++ + + +++ ++ Celtis africana + H+ 12 ++1 H R + +

Species group E

wudetia simplex + + 11 +1 +1 2 + 22 Ophrestia oblongifolia ++++H+ + + tt AIldropogon schirensis + + 1+ + Bewsia biflora + + + + + Digitaria tricholaenoides + + H Parinari cap€nsis 1++ Chaetacanthus bilichellii +++ Heliclirysum miconiifolium +++ Rhus magalismontana + + 2 Indigofera sanguinea H Kohautia amatyl!bica + ++ Monocyl!bium ceresiiforme +1 Gnidia species + + Cucumis hirsutus Hypoxis species + + Indigofera species tt

Species group F

Dianthus mooiensis Panicum coloratum I + ++ ++++++ + +1 ttl Alloteropsis semialata I + I' + '~m2+2 I I

Species group G

Tristachya leucothrix H+ +++ 2H H +2+1 Sporobolus p€ctinatus + + + H + +++++ + Ipomoea ommaneyi + + tt + +++++ Pentanisia angustitolia + + + + + ++ +++ + Pearsonia cajanitolia + + +H

Species group H

Diheteropogon amplectens 1 Hl++ ++ HI + + + 11 ++++ Bulbosty lis burchellii + + + +++ ++++ H +++ + :1 + I + Schizachyrium sanguineum I I I +2+ +1 + + 3+ I

Species group I

Eragrostis gummiflua + + + tt + + +3+ + 4+ ++ + ++ Chamaesyce hirta ++++ ++H + ++ + + +++ ++ Helichrysum nuditolium tt ++++ + ++ H ++ + ++++ HI +++ Dicoma cap€nsis + + tt +++ + + + + + ziziphus zeyheriana tt + + + + ++++tt+

Species group J

Aristida canescens H + ++2 tt Kyphocarpa angustitolia + + + +++ + + + Indigotera comosa + + + + H + +++ + Solanum cap€nse +++ + + + +++H tt Helichrysum caespititium + + + + ++++ + + +

Species group K

Salvia radula + + + + +++ ++ +++ +++++++++ Solanum pandiliitorme ++ + + + + + +++ + ++H Tragus berteronianus + + + + 2 ++ + Helichrysum rugulosum + ++ + + ++ + + +++ ++ Tylosema esculentum ++ tt +++++ Helichrysum coriaceum ++ + + + ++ ++1 + Ipomoea obscura \ +\ + +++ +\ I + S.Afr.J.Bot..1994.60(3) 157

Table 1 Continued

441666242 555451114 1111111 55555555450144 1121224 4411222 111111 22244550111111411111122224 011222 Sample numbers 004888405 222338444 2288948 3423232JJJ0544 7909013 4323010 222233 32734232335589466668903574 224017 210132931 345461344 0229120 38209584699615 6972109 0317875 578989 09553178560770645678809042 961663

1 Syntaxa numbers 1.1 1.2 I 1.3 I 2.1 I 22.2 I 2.3 I 3,2 I 3,3

Speci es group L

Eragrostis racellJOsa + + + ++++++ HH HHH ++++ + +1+ H+ ++++ + Crabbea angustifolia + + ++++HH+ H+ + H H+ H H+ ++++H++++++++ Trichoneura grandiglumis ++ +++++++ + + + m +H+ + +m + 2+ + + + Acalypha angustata H + + +++++ ++++H + ++ + + H ++H Aristida diffusa 1 H + 1 1 m2+ + H H +2 H + H + CYmbopcgon excavatus ++ 2 + ++ +++H + 11++ + HI coronatus + H + H++ m + + mH Hibiscus trionum + H+ H + H +H+ + + Becium grandiflorum +HH+ + +H + ++ H + + Thesium utile H H +H + + + Tephrosia longipes H +++ + H H +

species group M

Brachiaria serrata H + + + +++++++ H H+ 2++++ HH + ++++ H ++++++++ + Heteropcgon contortus 1 + H H + ++++ + 1+ H+ + +++1 H+ + ++++ senecio venosus + +H +++H+ H+++ H ++++H H+ +m cyanotis speciosa + + HH H H+H+ + + +H + +++++ Pygmaeothamnus zeyheri ++++ H+ ++H H ++ ++ +m Felicia muricata H H ++ ++m Dicoma anomala ++ + + + +HHH+H+ + +++ + + Justicia anagalloides H + H + +m ++++++++ H +++++++ +++ +++++ HH+ H +H +m++++ + Eragrostis lehmanniana 213 2+ +2 4 + + + +++ + + 4 + + 2 H Raphionacme hirsuta +H +++ + + H H+ mH H+ + H+ H Eragrostis superba ++H++++ + + + H+ H H2 2 H

Species group N

Chamaecrista millOsoides + + H +++ H H H + H ++++ ++++ ++++++ + ++++ + + ++++ ++++ + Plexipus hederaceus + +++ + H H H + H H +++ + +H H + ++ H m Bar leria macrostegia +HH++ H H + + H + + H ++++++ ++++ m +++++ H m Eustachys paspaloides H + + H + + H 3 + HH 2 m 22+ + H+ H2H+ Elephantorrhiza elephantina H + + +++ + H + + +++++ Hm +HH+ + + Trachypcgon spicatus H + H 11+21 + 1+ 2+ + + H H H + + H Commelina africana + H + + + + H++ ++++ H + H Mariscus indecorus H H++ + + ++ H + H++++ H H

Species group 0

Hyparrhenia hirta + + 125+42 Verbena bonariensis + +++H Eragrostis plana +H + H + 2H Scabiosa columbaria + + + H+ Chloris pycnothrix H+ Paspalum dilatatum +3

Species group P

Eragrostis curvula 11+221+1 1 + +23+ + 2+++ + 1 Hl+ +++11 H2H2 21+++42 3H22+ 2+3211 +J+322 ++++++ 3+++ ++++ + Elionurus muticus H + + + ++++ 2 +2+ + ++++1+1+2 + 22 +2H +2H2 +++++ + +++HH +++ 2H2+3+++ + + Themeda triandra 2 +22+ 3 + + +2+++ H H +++ + 1+ H ++++ + H 2 H+ +1111H5+ 2 2H23224+252 H + Aristida congesta + 21+++ +H + 4+ H + + + 11+ + + + + +2H + H 3H + H11+++++++ + +++++ +++ +4 Digitaria eriantha 2+ + 2 2 lH + HI 4+ 2 1 1++++ + ++++ H 3 + H1Hl++++++++ + +2 lH Anthospermum hispidulum + H + + + H++++ + +++ H + + ++++ ++++ H+H H + + H+ + + H H HH Vernonia oligocephala H + + ++H+ + H + + H + + H m+++++++++++ + + + Triraphis andropcgonoides +Hl H+ H + 1 HH H H H2+32 +2 H H + + Helinis repens + + + + + +++ + + + + HH + + +++ + H 2H + + + + 2 + cymbopcqon plurincdis +1 + 2+ + + 2 + + + +3H2 H H + +H +2H H + 1 H cyncdon dactylon 1 1+++ + + + H + + + + H + + + + + + + +++32 + +H Corchorus asplenifolius + ++++ H+ H H H+HH H+ + ++++ + setaria sphacelata H + 3H 2 H + H + + ++++ + 12 + HI + +2 Lactuca serriola ++ ++H ++ + H+ ++++ + +H H + Crabbea acaulis +H + HH+H + ++ ++++ Sida dregei +++ H + + +++ + H + + ++++ H HerJIannia depressa + H H H+ H+++m ++ H Oxygonum dregeanum + ++++ H+ +++ + H H . +++++ Pcqonarthria sguarrosa H + + + HH 1+ H+ +H H H Schkuhria pinnata ++ 1 + H + H H ++++ stoebe vulgaris R +H H H H Solanum incanum ++ + + ++++ ++ + Turbina oblongata H + + + H ++ H+H Blepharis angusta H++ H + + H ++ + +++ Walafrida densiflora H ++ H + H+ H+ +H 158 S.-Afr.Tydskr.Plantk., 1994, 60(3)

Table 1 Continued

441666242 555451114 lllllll 55555555450144 1121224 4411222 Illlll 22244550111111411111122224 011222 Sa~ple nullbers 004888405 222338444 2288948 34232J23330544 7909013 4J23010 222233 32734232335589466668903574 224017 2101J2931 345461344 0229120 38209584699615 6972109 0317875 578989 09553178560770645678809042 961663

2 Syntaxa nullbers 1.1 I 1.2 I 1.3 I 2.1 1 2.2 I 2.3 I 3.21 J.3 I : I

Rhyncbosia nervosa + + + + + + ++ H + Polygala hottentotta + + + H + + + + H + +H Pollichia campestris ++ + H + + + H H + + Lightfootia denticulata + + ++ H + + + + + + Sphenostylis angustifolia + +++++ ++++ + ++ Gnidia capitata ++ ++ + + + H + + Lippia scaberrima + + H + H + + IpclJO€a bathycolpcs H+ ++ ++ + + + + Mansonia angustif alia ++ + ++ + + + t t H Oral is species t t H Geigeria burkei tH t H +++ t t 8ermannia lancifalia H t t t H t t t Gomphrena celasiaides ++++ t t t 8elichrysum callical11Um t t ++ ++ t

Species with an occurrence af < 10 have been omitted. presence of pioneer grasses such as Aristida congesta, Cyno­ most scenic landscapes in the western Grassland Biome and don dactylon, Eragrostis superba and Melinis repens (Table 1; certainly deserves high conservation priority. species group P). 1.2 Zizipho mucronatae - Acacietum lUl"oo ass. nov. 1. Grevio jUlvae - Rhoion pyroidis all. nov. Nomenclatoral type: releve 523 Nomenclatoral type: releve 189 This association is not prominent in the Fa land type but it This alliance represents the woody vegetation of the Fa land occurs on the footslopes (4a) and sometimes may encroach into type and is characterized by species group D (Table 1). The the floodplains (4b) where it holds a subordinate position in the diagnostic species are the tree Celtis africana and the shrubs floristic composition, as was also mentioned by Bredenkamp Rhus pyroides, Grewia flava and Diospyros Iycioides, and the and Bezuidenhout (1990). This encroachment is usually the result of overgrazing or other forms of disturbance of the two small shrubby species Protasparagus suaveolens and P. vegetation (Friedel 1987). The soils are relatively deep (250 - laricinus. Two associations and one community without syn­ 1200 mm) and clayey (clay content> 20%) with the Hutton taxonomic rank can easily be recognized. A total of 25 releves (Hu) (500/0 of the terrain type), Mispah (Ms) and Glenrosa (Gs) represent this woody vegetation of the Fa land type. (25% of the terrain type) soil forms being dominant. The diag­ nostic species are the three Acacia tree species, namely A. 1.1 Rhoo lanceae - Acacietum eriolobae ass. nov. karroo, A. hereroensis and A. caffra, as well as Ziziphus Nomenclatoral type: releve 140 mucronata, while the shrub Tarchonanthus camphoratus and The Rhoo lanceae - Acacietum eriolobae is usually associated the grass Sporobolus fimbriatus are also diagnostic for this with the slightly undulating bottomlands of the Fa land type association (Table 1, species group B). The species of species (Figure 3) and can be observed while travelling from Potchef­ groups C, D (diagnostic for the alliance), M, Nand P are also stroom to Klerksdorp (Figure 1). This association is situated in present in this association (Table 1). An average of 32 species the south-western area of the Fa land type near Klerksdorp. was noted per sample plot. Sinkholes are characteristically present in this habitat which is The tree stratum, which is dominated by Acacia karroo, is underlain by dolomite. The soil is normally shallow « 300 5.9 m tall and has a canopy cover of 22.8% while the shrub mm) and clayey but the sinkholes, which are mostly filled by stratum is 2.2 m tall with a canopy cover of 26.9%. The herba­ deep (> 1200 mm) aeolian sand have a clay content less than ceous layer is 0.6 m tall and has a canopy cover of 48.9%. 10%. The dominant soil forms are Hutton (Hu) (50% of this Similar communities were described on different geological terrain type), Mispah (Ms) and Glenrosa (Gs) (together 33% of strata (Bezuidenhout et al. 1988; Bredenkamp et al. 1989; the terrain type) and also dolomite outcrops (17% of the terrain Bezuidenhout & Bredenkamp 1991) and also on dolomite (Be­ type) (Figure 2). The trees Acacia erioloba and Rhus lancea, zuidenhout & Bredenkamp 1990), but these communities have the shrub Lantana rugosa, the forb Pavonia burchellii and the never been ranked syntaxonomically. grasses Setaria verticil/ata, Eragrostis biflora and E. capensis are the diagnostic species (Table 1, species group A). Acacia 1.3 Digitaria eriantha - Rhus pyroides Shrubveld (com­ erioloba is normally restricted to the deep sandy soils in the munity without syntaxonomic rank) old sinkholes. Species from species groups C, D (diagnostic This community is usually associated with ruins and debris of for the alliance), N and the more common, widely distributed old diamond diggings situated on the midslope (3) of the Fa species from species group P (Table 1) are present in this asso­ land type, along archaic river beds. Surface rock usually ciation. An average of 31 species was recorded per sample comprises old diamond mine debris. The soils were disturbed plot. and overturned as a result of the diggings and therefore soil The tree stratum is well developed and is 6.5 m tall with a depth varies between 250 and >1200 rn. The clay content canopy cover of 24.7%. The shrub stratum is 1.8 m tall and has varies from 13 to 30%. This habitat leads to a relatively high a canopy cover of 9.6%, while the herbaceous layer is 0.5 m diversity in floristic composition. This community is classified tall with a canopy cover of 54.4%. under the Grewio flavae - Rhoion pyroidis, and although no Aesthetically, this association is associated with one of the syntaxonomic rank is assigned to this shrubveld, it is suggested S.Afr.J.Bot., 1994,60(3) 159 that it may represent an association. There are no diagnostic ropogonetum amplectentis in that the soil is deeper (300 - 900 species for this commWlity but the strong presence of species mm) and has a higher clay content (> 10%). The dominant soil group D (Table 1) (diagnostic for the alliance) characterizes types present in this association are the Mispah (Ms) and Olen­ this commWlity. Species of species groups L, M, Nand P rosa (Os) (72% of the terrain type) and the Hutton (Hu) (12% (Table 1) are also present in this commWlity. An average of 43 of the terrain type) forms (Figure 2). Large rocks cover 10 - species per sample plot was recorded. 70% of the soil surface. The diagnostic species for this asso­ The tree stratum is poorly developed and is 4.4 m tall with a ciation are the grasses Panicum coloratum and Alloteropsis canopy cover of 10%. The shrub stratum is 2.4 m tall with a semialata and the inconspicuous forb Dianthus mooiensis canopy cover of 14.9% while the well-developed herbaceous (Table 1, species group F). The species of species groups 0 layer is 0.8 m tall and has a canopy cover of 56.4%. and H (diagnostic for the aIliance), and widespread species of A similar community was identified by Bezuidenhout and species groups L, M, N and P are also present in this asso­ Bredenkamp (1990). ciation (Table 1). An average of 40 species was noted per sample plot. 2. Trachypogono spicati - Diheteropogonion amplectentis The tree stratum is absent and a scanty shrub stratum may be all. nov. present with a canopy cover of 5% and 1.8 m tall. The herba­ Nomenclatoral type: releve 533 ceous layer, with Tristachya leucothrix and Alloteropsis semia­ This alliance occurs on the relatively flat or slightly undulating lata the prominent grasses, is 0.7 m tall and has a canopy cover stony plains of the upland areas. The soil of this aIliance is of 47.9%. shaIlow (50 - 150 mm) and generally very rocky with the Similar communities were identified by VanWyk and Bre­ exception of the Alloteropsido semialatae - Tristachyetum denkamp (1986) and Bezuidenhout and Bredenkamp (1990), leucothricis whose soil is deeper (300 - 900 mm). Two grass but no formal syntaxonomical rank was assigned. species, namely Diheteropogon amplectens and Schizachyrium sanguineum, and the grass-like forb Bulbostylis burchellii are 2.3 Cymhopogon excavatus - Diheteropogon amp/ectens the diagnostic species for this alliance (Table 1, species group Grassland (community without syntaxonomic rank) H). These species are typical for the relatively dry upland, This grassland community is also associated with the high­ sandy or rocky, weIl-drained areas and do not occur on the lying areas in the Fa land type (Figure 2). It is fOWld on the floodplains or on other relatively wet, clayey, poorly drained midslopes (3) but is situated downslope, below the Alloterop­ bottomlands situations. This was also noted by Van Wyk and sido semialatae - Tristachyetum leucothricis (2.2) (Figure 3). Bredenkamp (1986), Bredenkamp and Bezuidenhout (1990), Mispah (Ms), Olemosa (Os) (88% of the terrain type) and Scogings and Theron (1990), and Bezuidenhout and Breden­ Hutton (Hu) (4% of the terrain type) soil forms are dominant in kamp (1991). This alliance is described by 28 releves and this commWlity. Less than 100/0 of the soil surface is covered comprises two associations. One commWlity without syntaxo­ with rocks and stones. No syntaxonomic rank was assigned to nomic rank is also recognized. this grassland because too little is known about the syntaxo­ nomic position of this commWlity. There are no diagnostic 2.1 Loudetio simplicis - Diheteropogonetum amp/ectentis species for this community but the presence of the diagnostic ass. nov. species for the aIliance (species group H) and the absence of Nomenclatoral type: releve 533 species groups D (diagnostic for Grewio flavae - Rhoion pyroidis) and I (diagnostic for Cymbopogono plurinodis - The Loudetio simplicis - Diheteropogonetum amplectentis is Eragrostidetum gummifluae) suggest that this community strongly associated with the rocky chert outcrops and ridges on should be classified under the Trachypogono spicati - the plateau (1) of the Fa land type's undulating landscape Diheteropogonion amplectentis (Table 1). Widespread species (Figures 2 and 3). The limited shaIlow soil (50 - 150 mm) with from species groups L, M, Nand P (Table 1) are present in a low clay content « 10%) present in this association is pre­ this commWlity. An average of 32 species was recorded per dominated by chert outcrops (67% of the terrain type) and the sample plot. Mispah (Ms) and Olemosa (Os) (33% of the terrain type) soil The tree and shrub strata are absent but the he.baceous layer forms. Less than 10% of the soil surface is covered by scatter­ is well developed, 0.8 m taIl and has a canopy cover of 57.1 %. ed rocks. Diagnostic species of this association include the Old cultivated areas left to recuperate are also fOWld in this prominent grass species Loudetia simplex, Andropogon grassland. schirensis, Bewsia biflora and Monocymbium ceresiiforme, all associated with rocky areas, and the dwarf shrubs Rhus 3. Cymhopogono plurinodis - Eragrostidetum gummijluae magalismontana and Parinari capensis as well as the con­ ass nov. spicuous forbs Ophrestia oblongifolia and Helichrysum Nomenclatoral type: releve 135 miconiifolium (Table 1, species group E). Species from species This association occurs on the midslopes (3) and the bottom­ groups 0 and H (diagnostic for the alliance), and the wide­ land floodplains (4b) of the Fa land type (Figures 2 and 3). The spread species of species groups L, M, Nand P (Table 1) may soils are relatively deep (250 - 1200 mm) and have a clay also occur in this association. An average of 37 species per content of between 13 and 300/0. The pioneer grass species sample plot was recorded. Eragrostis gummiflua and the dwarf shrub 2iziphus zeyheriana The tree stratum is absent and the shrub stratum is poorly as weIl as the conspicuous forbs Helichrysum nudifolium and developed, 1.0 m tall with a canopy cover of only 7.4%. The Dicoma capensis, together with the inconspicuous forb Cha­ shrub stratum, with Protea welwitschii being prominent, is maesyce hirta, are diagnostic for this association (Table 1, spe­ conspicuous in the central northern areas of the Fa land type, cies group I). The vegetation is often subjected to overgrazing, adjacent to the Sourish Mixed Bushveld (veld type 19) (Acocks resulting in degradation and the subsequent presence of many 1988). The herbaceous layer has a canopy cover of 55.7% and pioneer species. Two sub-associations can be identified. is 0.8 m tall. 3.1 Cymhopogono plurinodis - Eragrostidetum gummijluae 2.2 Alloteropsido semwlatae - Tristachyetum leucothricis aristidetosum canescentis subass. nov. a'is. nov. Nomenclatoral type: releve 125 Nomenclatoral type: releve 176 The Cymbopogono plurinodis - Eragrostidetum gummifluae This grassland association occurs on the relatively high altitude aristidetosum canescentis is associated with bottomlands but plateaux (1) and differs from the Loudetio simplicis - Dihete- relatively high-lying, rocky, lower midslopes (3) just above the 160 S.-Afr.Tydskr.Plantk., 1994, 60(3)

floodplains of the Fa land type. There may be outcrops of impaired by the presence of a strong group of common species dolomite rock, and sometimes rock sheets occur just beneath (Table 1, species group P). The presence of these common the soil surface in which case the soil is shallow « 250 mm). species is ascribed to the compilation of the data in a relatively Normally the soil is deeper than 250 mm with the Hutton (Hu) homogenous land type. (47% of the terrain type), Mispah (Ms) and Glenrosa (Gs) Along the first axis the woodland syntaxa (1) occur to the (46% of the terrain type) soil forms predominant in this sub­ left of the diagram while the grassland syntax a (2 and 3) occur association (Figure 2). The diagnostic species are the pioneer in the centre with the wetland syntaxa (4) to the right of the grass species Aristida canescens and the conspicuous pioneer diagram. Along the second axis a gradient is illustrated which forbs Kyphocarpa angustifolia and Solanum capense and the may be related to altitude and moisture (Figure 4). This trend dwarf shrub Indigo/era comosa. The inconspicuous mat­ confirms the result of the classification, and is not discussed forming forb Helichrysum caespititium (Table 1, species group further. J) is also diagnostic. Species from species groups I (diagnostic for the association), L, M and N and the more common species Conclusions from species group P are also present in this sub-association This is the first comprehensive syntaxonomical account of the (Table 1). An average of 44 species was noted per sample plot. vegetation of the dolomite and chert region (Fa land type) in The tree and shrub strata are absent. The herbaceous layer is the western Transvaal Grassland Biome. New syntax a describ­ degraded, 0.7 m tall, with a canopy cover of 35%. ed include two alliances, six associations and two sub-associa­ tions with two communities without syntaxonomical status. 3.2 Cymbopogono plurinodis - Eragrostidetum gummifluae The floristic and habitat diversity of the Fa land type, eragrostidetosumsuperbae subass. nov. especially the Rhoo lanceae - Acacietum eriolobae should Nomenclatoral type: reI eve 135 have high conservation priority. Small nature reserves, such as This sub-association is found in the low-lying floodplains (4b) the Abe Bailey Nature Reserve (1887 ha) (Van Wyk 1983) and of the Fa land type (Figure 2). The soil (250 - 1200 mm deep) a part of the Jack Scott Nature Reserve (2100 ha) (Coetzee has more than 5% rocks and stones on the surface. The domi­ 1974), are at present the only known protected vegetation areas nant soil forms are the Hutton (Hu) (50% of the terrain type), in the Fa land type. It is alarming that only 4000 ha (0.72%) of Mispah (Ms) and Glemosa (Gs) (33% of the terrain type). The this vegetation is being formally conserved. The presence of diagnostic species are the pioneer grass Tragus berteronianus pioneer species in species groups L, M and N (Table 1) indi­ and the forbs Salvia radula, Solanum panduriforme, Heli­ cates that this vegetation is in a degraded state. The main chrysum rugulosum, Tylosema esculentum, Helichrysum coria­ factors seem to be the relatively low rainfall of the area over ceum and Ipomoea obscura (Table 1, species group K). Other the past ten years and the continuous overgrazing of the natural species from species groups I (diagnostic for the association), vegetation by domestic stock. This classification of vegetation L, M, Nand P are also present in this sub-association (Table and associated habitat should form a basis for all vegetation­ 1). An average of 36 species was recorded per sample plot. related management planning as well as for establishing prior­ The tree stratum is absent. A poorly developed shrub ities for conservation of land. This description of the Fa land stratum is locally present, with a canopy cover of 8%, and is type contributes considerably to the understanding and present 1.83 m tall. The well-developed herbaceous layer is 0.77 m tall knowledge of the western Transvaal grassland. and has a canopy cover of 62%. Acknowledgement 4. Paspalo dilatati- Hyparrhenietum hirtae ass nov. Mrs D. Bezuidenhout is thanked for assistance in the prepara­ Nomenclatoral type: releve 141 tion of the manuscript. The Paspalo dilatati - Hyparrhenietum hirtae is strongly associated with the low-lying drainage lines (5) in the Fa land References type. These seasonally wet bottomlands have soils with a ACOCKS, l.P.H. 1953. Veld types of South Africa, 1st edn. Mem. higher clay content (> 25% clay) than those of the upland bot. Surv. S. Afr. 28: 1 - 192. areas. No or very little rock or stone occur on the soil surface. ACOCKS, l.P.H. 1988. Veld types of South Africa, 3rd edn. Mem. The dominant soil forms present in this association are bot. Surv. S. Afr. 57: 1 - 146. Westleigh (We), Valsrivier (Va) (40% of the terrain type) and BARKMAN, J.J., MORAVEC, 1. & RAUSCHERT, S. 1986. Code Hutton (Hu) (40% of the terrain type) (Figure 2). The of Phytosociological Nomenclature, 2nd edn. Vegetalio 67: diagnostic species are the perennial grasses Hyparrhenia hirta, 145 - 195. Chloris pycno thrix, Eragrostis plana and Paspalum dilatatum BEZUIDENHOUT, H. 1988. 'n Plantsosiologiese studie van die and the conspicuous forbs Verbena bonariensis and Scabiosa Mooirivieropvanggebied, Transvaal. M.Sc. thesis, Potchefstroom columbaria (Table 1, species group 0). The only other species University for CHE, Potchefstroom. present are the more common species of species group P BEZUIDENHOUT, H. in prep. Syntaxonomy and synecology of (Table 1). An average of 26 species was noted per sample plot. western Transvaal grasslands, South Africa. Ph.D. dissertation, A tree stratum is absent and the 1.7 m tall shrub layer covers University of Pretoria, Pretoria BEZUIDENHOUT, H. & BREDENKAMP, G.1. 1990. A reconnais­ less than 10% of the area. The herbaceous layer is well devel­ sance survey of the vegetation of the dolomitic region in the oped, with a canopy cover of 56.7%, and is 1.0 m tall. Potchefstroom - Ventersdorp - Randfontein area, South Africa. This habitat is fairly unstable owing to seasonal flooding and Phytocoenologia 18: 387 - 403. subsequent desiccation which, together with the frequent over­ BEZUIDENHOUT, H. & BREDENKAMP, G.l. 1991. The vegetation grazing of these sites, causes the advanced state of degradation of the Bc land type in the western Transvaal Grassland, South of the vegetation. Africa PhYlOcoenologia 19: 497 - 518. A similar community was described by Bezuidenhout and BEZUIDENHOUT, H., BREDENKAMP, GJ. & ELSENBROEK, Bredenkamp (1990). 1.H. 1988. Die plantegroei van die a1kaligraniet en aangrensende kwartsiet in die Vredefortkoepel noordwes van Parys. S. Afr. Ordination Tydskr. Naluurwel. Tegnol. 7: 4 - 9. In the ordination diagram the distribution of the major syntaxa BREDENKAMP, G.l. & BEZUIDENHOUT, H. 1990. The vegetation along the first and second axes of the ordination is given of the Faan Meintjes Nature Reserve, Transvaal. S. Afr. 1. BOI. 56: (Figure 4). The syntaxa are restricted to specific spacial areas 54 - 64. in the ordination diagram. It is possible to circumscribe the BREDENKAMP, GJ., JOUBERT, A.F. & BEZUIDENHOUT, H. different syntaxa but a distinct discontinuity between them is 1989. A reconnaissance survey of the vegetation of the plains in S.Afr.J.Bot., 1994,60(3): 161-167 161

the Potchefstroom-Fochville-Parys area. S. Afr. J. BOI. 55: 199 - 206. West Rand and 2726 Kroonstad. Mem. agric. nal. Resour. S. Afr. BREDENKAMP, GJ. & THERON, G.K. 1978. A synecological 4: 1 - 441. account of the Suikerbosrand Nature Reserve. 1. The phyto­ LOUW, W.J. 1951. An ecological account of the vegetation of the sociology of the Witwatersrand geological system. BOlhalia 12: Potchefstroom area Mem. bot. Surv. S. Afr. 24: 1 - 105. 513 - 529. MACVICAR, C.N., LOXTON, R.F., LAMBRECHTS, J.J.N., LE BREYffiNBACH, P.J.J. 1991. Die fitososiologie van die Villiers - ROUX, J., DE VILLIERS, J.M., VERSTER, E., MERRY­ Grootvlei-omgewing. M.Sc. thesis, University of Pretoria, Pretoria WEATHER, F.R., VAN RooYEN, T.H. & HARMSE, HJ.von COETZEE, BJ. 1974. A phytosociological classification of the vege­ M. 1977. Soil classification - A binomial system for South Africa. tation of the Jack Scott Nature Reserve. BOlhalia 11: 329 - 347. Department Agricultural-technical services, Pretoria DU PREEZ, PJ. 1991. A syntaxonomical and synecological study of MENTIS, M.T. & HUNTLEY, B.J. 1982. A description of the Grass­ the vegetation of the southern and eastern Orange Free State and land Biome Project. S. Afr. Nat. Sci. Prog. 62: 1 - 29. related areas with special reference to Korannaberg. D.Sc. disser­ MORRIS, J.W. 1973. Automatic classification and ecological profiles tation, University of the Orange Free State, Bloemfontein. of south-western Transvaal Highveld grassland. Ph.D. dissertation, DU TOrr, A.c. 1954. The geology of South Africa Oliver & Boyd, University of Natal, Pietermaritzburg. Edinburgh. MUELLER-DOMBOIS, D. & ELLENBERG, H. 1974. Aims and ECKHARDT, H.C. 1993. A synecological study of the vegetation of methods of vegetation ecology. Wiley, New York. the north-eastern Orange Free State. M.Sc. thesis, University of MYBURGH, W.J. 1990. 'n Plantsosiologiese studie van die Pretoria, Pretoria Bankenveld en die vleigebiede in die Grootvlei-distrik, Suid­ FRIEDEL, M.H. 1987. A preliminary investigation of the woody Transvaal. B.Sc.(Hons.) thesis, University of Pretoria, Pretoria plant increase in the western Transvaal and implications for veld SCHEEPERS, J.C. 1986. Grassland Biome Project: Proceedings of assessment. 1. Grassl. Soc. Soulh Afr. 4: 25 - 30. the workshop on the classification and mapping. Ecosystems GIBBS RUSSELL, G.E., REID, C., V AN Rooy, J. & SMOOK, L. Programmes Occasional Report Series No. 16, CSIR, Pretoria 1985. List of species of southern African , 2nd edn., Part 1. SCOGINGS, P.F. & THERON, G.K. 1990. An application of multi- Mem. bOI. Surv. S. Afr. 51: 1 - 152. variate analysis techniques to dry-weight-rank data from a GIBBS RUSSELL, G.E., WELMAN, W.G., RETIEF, E., IMMEL­ Bankenveld nature reserve. S. Afr. J. BOI. 56: 648 - 653. MAN, K.L., GERMISHUIZEN, G., PIENAAR, B.J., VAN WYK, TURNER, BJ. 1989. A phytosociological study of the south-eastern M. & NICHOLAS, A. 1987. List of species of southern African Transvaal Highveld grasslands. M.Sc. thesis, University of Pre­ plants, 2nd edn., Part 2. Mem. bOI. Surv. S. Afr. 56: 1 - 270. toria, Pretoria HILL, M.O. 1979a TwINS PAN - A Fortran program for arranging V AN WYK, S. 1983. 'n Plantekologiese studie van die Abe Bailey­ multivariate data in an ordered two-way table by classification of natuurreservaat. M.Sc. thesis, Potchefstroom University for CHE, the individuals and attributes. Cornell University, New York. Potchefstroom. HILL, M.O. 1979b. DECORANA - A Fortran program for detrended V AN WYK, S. & BREDENKAMP, GJ. 1986. 'n Braun-Blanquet­ correspondence analysis and reciprocal averaging. Cornell Univer­ klassifikasie van die plantegroei van die Abe Bailey-Natuur­ sity, New York. reservaat. S. Afr. 1. Bot. 52: 321 - 331. KOOH, M.S. 1990. A phytosociological study of the vegetation of the VAN RooYEN, N., THERON, G.K. & GROBBELAAR, N. 1981. A north-western Orange Free State. M.Sc. thesis, University of floristic description and structural analysis of the plant commun­ Pretoria, Pretoria ities of the Punda Milia - Pafuri - Wambiya area in the Kruger LAND TYPES SERIES. 1979. 2626 Wes-Rand. Prepared by the Soil National Park. 1. The hygrophilous communities. JI S. Afr. BOI. and Irrigation Research Institute, Pretoria. 1 :250 ODD. Government 47: 213 - 246. Printer, Pretoria WEATHER BUREAU. 1988. Climate of South Africa. WB 40: 1 - LAND TYPE SURVEY STAFF. 1984. Land types of the maps 2626 475. Government Printer, Pretoria

Two new species of Cineraria L. (, )

G.V. Cran Department of Botany, University of the Witwatersrand, Private Bag 3, Wits 2050, Republic of South Africa

Received 7 December 1993; revised 3 March 1994

Two new species of Cineraria L., C. austrotransvaalensis Gron and C. hederifolia Gron, are described from the Transvaal. Their affinities, geographical distribution and conservation status are discussed.

Twee nuwe spesies van Cineraria L., C. austrotransvaalensis Gron en C. hederifolia Gron, vanuit die Transvaal word beskryf. Hulle affiniteite, geografiese verspreiding en bewaringstatus word bespreek.

Keywords: Asteraceae, Cineraria, new species, Senecioneae.

Introduction glabrous or ciliate with thickened margins or wings. Cineraria L. is an African genus with approximately forty-five The two new species described are from the Transvaal: C . species currently described, although a few of these are con­ austrotransvaalensis Cron from the koppies and ridges of the sidered doubtful (Hilliard 1977; Jeffrey 1986). Most species Witwatersrand and adjacent areas, and C. hederifolia Cron are southern African, with a few in east and tropical Africa and from the Blouberg in the north-west and the Drakensberg in one in Madagascar. The genus includes herbs and sub shrubs , the east. Both species are characterized by the tomentose with alternate (or rarely radical), lobed leaves. The capitula are (white-felted) indumentum, which is widespread in the genus mainly heterogamous and radiate, with yellow florets. The and reflected in the name 'Cineraria', meaning grey or ash­ cypselae are obovate, compressed, brown or black, and either coloured.