198 S.-Afr.Tydskr.Plantk .• 1991. 57(4): 198 - 206 The syntaxonomy and synecology of the forests in the eastern Orange , South Africa. I. The Podocarpetalia latifolii

P.J. du Preez,*t G.J. Bredenkamp* and H.J.T. Venter§ tNational Museum. P.O. Box 266. Bloemfontein. 9300 Republic of South Africa *Oepartment of Botany, University of Pretoria, Pretoria, 0002 Republic of South Africa §Department of Botany, University of the Orange Free State, Bloemfontein, 9300 Republic of South Africa

Accepted 24 April 1991

A formal syntaxonomical classification, synecological interpretation and description of the moist forests of the mountainous area of the eastern Orange Free State are presented. Releves were compiled in 31 sample plots in remaining natural forest confined to specific niches in deep valleys, protected gorges and ravines . The releves were compiled from two separate but related studies. The data were initially classified by TWINSPAN numerical analysis and then refined by Braun-Blanquet procedures. In order to determine a prob­ able vegetation gradient, an ordination algorithm, namely DECORANA, was applied to the floristic data set. A formal syntaxonomical classification was possible, as this investigation forms part of a comprehensive syne­ cological and syntaxonomical study of the vegetation of the southern and eastern Orange Free State. This study contributes significantly to the syntaxonomical knowledge of the Afromontane forests of southern Africa.

'n Formele sintaksonomiese klassifikasie, sinekologiese interpretasie en beskrywing van die vogtige woude van die bergagtige oostelike Oranje-Vrystaat word aangebied. Releves is saamgestel uit 31 monsterpersele. Hierdie monsterpersele is in die oorblywende natuurlike woud , wat in spesifieke nisse in diep en beskermde valleie in die studiegebied voorkom, uitgeplaas. Die saamgestelde data is uit twee afsonderlike maar verwan­ te studies verkry. Die data is aanvanklik deur die TWINS PAN numeriese analise geklassifiseer en die resultaat is deur middel van Braun-Blanquet prosedures verfyn. Om te bepaal of 'n gradient in die plantegroeiversprei­ ding voorkom, is 'n ordeningsalgoritme, naamlik DECORANA, op die floristiese data toegepas. 'n Formele sintaksonomiese klassifikasie kon gedoen word omdat hierdie studie deel uitmaak van 'n omvattende sineko­ logiese en sintaksonomiese projek oor die plantegroei van die suidelike en oostelike Oranje-Vrystaat. Hierdie studie dra sinvol by tot die kennis van die sintaksonomie van die Afro-montaanse plantegroei van Suid-Afrika.

Keywords: Afromontane forests, Braun-Blanquet method, DECORANA, Syntaxonomy, TWINSPAN

'To whom correspondence should be addressed.

Introduction in South Africa was that of Van Zinderen-Bakker (1973) on Many vegetation studies have been undertaken in forests in forest communities in the eastern Orange Free State, while South Africa (White 1978). In the Afromontane forest belL Everard (1986) gave a phytosociological classification of these studies were mostly of a descriptive and floristic some Podocarpus communities from th e Royal Natal nature (Bews 1917; Schelpe 1943; Killick 1963; Edwards National Park. 1967; Moll 1972; Cooper 1982, 1985; Acocks 1988). In Within a comprehensive phyLOsociological research pro­ spite of all these investigations, little is known aoout the gramme in South Africa (Bredenkamp et at. 1989), a syne­ phytosociology, and especially the syntaxonomy of these cological and syntaxonomical study of the vegetation of the forests. south-eastern Orange Free State was initiated (Du Preez, in Most Afromontane tree species have wide geographical prep.). One of the projects within this programme was to distributions and wide ecological amplitudes (White 1983). identify, characterize and describe the communities of Consequently, the Afromontane vegetation is remarkably the Afromontane forests as extrazonal vegetation in the uniform and it is considered as a continuum without conspi­ grassland of the study area. The approach is one of synthe­ cuous discontinuity, which is difficult to classify (White sis, i.e. to incorporate all compatible existing and new 1978). However, White (1978) recognized that South Afri­ phytosociological data and information in a comprehensive can Afromontane forests are more complicated than other synecological and syntaxonomical synopsis. A formal African Afromontane forests (Zimbabwe, Malawi and tropi­ classification and syntaxonomy of the Afromontane forests, cal Africa) because in southern Africa the individual and according to the Ziirich-Montpellier methodology (Mueller­ isolated islands of Afromontane vegetation are surrounded Domoois & Ellenberg 1974), are presented in this report by a great diversity of lowland vegetation types, and also since the climate on the eastern and western slopes of the The study area Drakensberg mountain range, where these forests occur, The study is confined to a broad zone of mountainous area differs. from Danger Point (Korannaberg) in the west to Harrismith One of the first synecological and syntaxonomical studies in the east. It is bound by longitudes 27°00' and 29°30' S.AfrJ.Bot..1991, 57(4) 199

28" 30'

Natal

~ < 1600m ~ 1600m 1·9 Weather Stations o 1800m _ 2000m 10 Kilometres > 2200m Figure 1 The study area in relation to towns and nature reserves.

east and latitudes 28°00' and 29°00' south. The northern to east and a decrease of average daily temperatures from border of Lesotho forms the southern boundary of the study west to east across the study area (Van der Wall 1976). area (Figure 1). Geology Climate The study area is underlain by the Karoo Sequence. The According to the Koppen classification the study area lies in stratigraphic units of this sequence from the bottom are the a mesothermic climatic zone. This Cwb-climate zone re­ Beaufort, Molteno, Elliot and Clarens Formations. On top is ceives approximately 550 mm or more rain per annum, the Drakensberg Formation. The Karoo Sequence is intruded mainly during summer and autumn. This zone experiences by numerous dolorite dykes and sills. Recent alluvium and cold, dry winter periods. The average temperature of the scree covers the valley floors while scree also covers the warmest month is less than 22°C, but an average mountain slopes (Groenewald 1986). temperature of more than 10°C is maintained for at least The first three formations consist of alternate layers of four months per year (Van der Wall 1976). siIt- and mudstone and medium- to coarse-grained sand­ The average annual precipitation increases from the west stone. These layers are of fluvial origin. The next formation to the east across the study area (Weather Bureau 1954) is the characteristic yellowish sandstone cliffs of the Clarens (Figure 2). This is caused by the increasing relief from west Formation and is of aeolian origin. It consists of fine- to medium-grained sandstone. Narrow gorges in this formation 1400 were formed by the relatively rapid weathering of dolorite

] 1200 dykes. During the intrusion phase (Jurassic Period), hot E igneous material penetrated the sandstone and baked it to g 1000 ..c: hard solid rock, which is resistant to weathering. The narrow c: 800 gorges were formed by the subsequent more rapid weather­ .!! :! ing of the dolorite dykes (Bonnett 1977; Groenewald 1986) . 'n. 600 u The highest peaks (1800 m above sea-level) are capped by •0. 400 numerous layers of amygdaloidal and massive varieties of • basaltic lava of the Drakensberg Formation (Groenewald '"!! 200 ~ 1986). o 2 3 s 7 8 9 Methods Wealhe, alallon. Releves were obtained from representative sites over the Figure 2 The average annual precipitation (mm) of nine weather entire range of Afromontane forests in the eastern Orange stations from west to east across the study area (Weather Bureau 1954). Free State and also from adjacent, ecologically and floristically related areas in Natal. No. Station Station No. No. Station Station No. By using the two-step methodology developed by Breden­ I. Danger Point (295/413) 6. Harrismith (333/317) kamp and Bezuidenhout (1991), these releves were com­ 2. Mequatlingsnek (295/539) 7. Tandjiesberg (333/643) pared to and distinguished from 3286 releves representing 3. Gocolan (296/11SA) 8. Gifford Chambers (299/100) other vegetation types in the area. Diagnostic species were 9. Cathedral Peak (299/357) 4. Ficksburg (296/682A) identified for the Afromontane forests and these forests are 5. Fouriesburg (297/337A) considered to represent a vegetation class, newly described 200 S.-Afr.Tydskr.Plantk., 1991,57(4)

in this report. Table 1 (continued) Most releves representing Afromontane forests were com­ Order' piled by authors of two separate, but related studies. In a study of the vegetation of the Afromontane forest in the 2

Harrismith region, Du Preez and Venter (in prep.) compiled, Association' ABCDEF GHIJKL amoogst others, 9 releves in this vegetation type. A further 32 releves were oOOiined from Van Zinderen-Bakker (1973), giv­ Sireplocarpus pusillus 2 AgapanJhus campanuialus 2 ing a total of 31 releves in the eastern part of the study area. In accordance with Van Zinderen-Bakker (1973), the plot Species Group C Asplenium molUlnJhes 15 2 size chosen was 10 x 10 m (100 m ). All species Rubus rigidus 15 recognizable at the time of the surveys were noted. For Cyslopleris fragilis 4 cover the Braun-Blanquet cover-abundance scale as given Galopina eireaeoides 15 by Mueller-Dombois and Ellenberg (1974) was used. Siaehys eaffra 4 Senecio oxyriifolius Environmental infonnation included rock type, altitude, 25 Rhus penJheri 4 aspect, slope and published climatological data. Scadoxus punieeus 24 All releves representing this class were initially classified Dioscorea sylvalica 24 by Two-way Indicator Species Analysis (TwINSPAN) (Hill Pleiopeltis schraderi 4 2 1979a), and this first approximation of vegetation types was AdianJum poiretii 4 Zehneria scabra then refined by the application of Braun-Blanquet proce­ 3 TOOlielrum rhynehocarpum 3 dures (Bredenkamp et al. 1989; Bredenkamp & Bezuiden­ Slachys grandifolia 3 hout 1991). A synoptic table was extracted from -the phyto­ Pseudobromus sylvalieus 3 sociological table. The synoptic table summerizes species Species Group D fidelity to communities within the vegetation units (Cowling Dais eOlinifolia 55 1983). From the synoptic table floristic relationships and Erylhroxylwn emarginalwn 44 differences among the various recognized syntaxa may be Oplismenus hirteilus I 255 Dieles iridioides 222 observed while higher syntaxa than the association may be Begonia sUlherlandii 23 I characterized. The constancy of all species within each Andrachne ovalis 2 I community is rated on the following 20%-interval scale: Species Group E 1: <20%; 2: 21-40%; 3: 41-60%; 4: 61-80% and 5: >80%. Cryplocarya woodii 4 In this study the synoptic table (Table 1) is used to Buddleja auriculala 3 I indicate diagnostic and general species of the class, and to Hypoesles Irijlora 3 distinguish orders, while the phytosociological table (Table Species Group F 2) is used to describe the syntaxa of one of the orders, Seolopia zeyheri 3 Cyperus alboslrialus namely the Podocarpetalia latifolii. Taxon names confonn 3 to those of Gibbs Russell et al. (1985 & 1987). Species Group G Peperomia re/usa 3254 In order to detennine a probable vegetation gradient, an Carissa bispinosa 1 245 1 1 ordination algorithm, namely the Detrended Correspondence Polyslichum pungens 4553 11 Analysis (DECORANA) (Hill 1979b), was applied to the Sireplocarpus hayganhii 5442 floristic data set. Protasparagus setaceus 2235 1 AdianJum capillus-veneris 3 52 Impatiens hochstetteri 2 24 Results PleclranJhus grallalus 5531 2 Classification Species Group H The analysis revealed 12 plant communities under the class Podocarpus latifolius 355 5 Olinia emarginala 123454 CanJhium cilialwn 344313 I Table 1 Synoptic table of the Sc%pietea mundii Amaranlhus sp. 55 5 3 Pleiopeilis macrocarpa 353 I I 2 Order' Species Group I 2 /lex milis 223 454 Association' ABCDEI' GH I} K L Maylenus aeuminala 44 353 Species Group J Species Group A Blechnum auslrale ClauselUl anisala 24 5 Dryopleris ilUlequalis 35 Species Group K Crassula pellucida 43 Crassula exilis 12 Diclis replans 34 Crassula lanceolala subsp. transvaalensis 1 3 Pleris crelica 33 221 Cheilanlhes eckloniana 2 Coccinia palmala 23 Species Group L Galium capense 22 Cineraria Iyrala 2 151 Bowkeria verlicillala 32 Pavonia burchellii 51 Species Group B Plumbago zeylanica 5 Asplenium adianJum-nigrum 3 Leonalis ocymifolia 12 5 Thamnocalamus lessellalus 2 Zanledesehia aelhiopiea 5 S.AfrJ.Bot., 1991,57(4) 201

Table 1 (continued) Scolopietea mundii, six under the order Podocarpetalia lati/olii, described in this report, and six under the order Order' Pittosporetalea viridijlorwn, described by Du Preez and 2 Bredenkamp (1991). The hierarchical classification of the

Association" ABCDEF GHIJKL Podocarpetalia lati/olii is as follows: Class l. Scolopietea mundii Class nov. Solanum retroflexum 12 5 Tagetes minuta 51 Order 1. P odocarpetalia lati/olii Order nov. Bidens bipinnata 5 Alliance 1. Peperomio-Polystichion pungenlis All. Coccinia hirtella 15 nov. Cyathula uncinata 5 Association l. Clauseno-Podocarpetum lali/o­ Datura stramonium 5 lii Ass. nov. Hermannia geniculata 5 Withania somnifera 5 Sub-association l. Clauseno-Podocarpe­ tum lati/olii bowkerietosum verticil/atae Species Group M Stipa dregeana 2 2 2 sub-ass. nov. Rhus lancea 2 2 Sub-association 2. Clauseno-Podocarpe­ Species Group N tum lati/olii-asplenietosum monanthis sub­ Sculia myrtina 12 5454 1 ass. nov. Rhoicissus Iridentala 5453 Association 2. Peperomio-Carissetum Van Osyris lanceolata 233 2 Zinderen-Bakker (1973). Omlis depressa 12 1 13 2 Sub-association 1. Peperomio-Carissetum­ Mohria caffrorum 2 3434 Cetarach cordatum 3 1 5352 cryptocaryetosum woodii sub-ass. nov. Achyranthes sicula 1 3 52 1 Sub-association 2. Peperomio-Carissetum­ Rhus burchellii 252 scolopietosum zeyheri sub-ass. nov. Pellaea calomelanos 12 Alliance 2. Unspecified. Species Group 0 Association 1. Ehrharto-Eucleetum crispae Pittosporum viridiflorum 32 443 3 Van Zinderen-Bakker (1973). Olea europaea subsp. africana 2 2 455554 Sub-association 1. Ehrharto-Eucleetwn Celtis africana 22 23251 1 Rhus dentata 222 1 1 222 33 crispae podocarpetosum Van Zinderen­ Heteromorpha Irifoliata 3 2 1 3 2 2332 Bakker (1973). Grewia occidentalis 32 2 233532 Asplenium aelhiopicum 23 21 124 3 1 Description of the plant communities Haemanthus humilis 1 212 1 Class I. Scoiopietea mundii Class nov. Species Group P Scolopia mundii 144412 455551 This class represents all typical Afromontane forests in the Cassinopsis ilicifolia 23234 434334 study area. According to Cooper (1982) the highest in­ digenous forest (2134 m above sea-level) in South Africa Species Group Q Diospyros whyleana 455554 555554 occurs in the study area. These forests occur in specific May tenus unciata 2 2545 555544 niches in deep valleys, protected gorges, crevices and Myrsine africana 53 33 555544 ravines along the eastern and western slopes of the Ehrharta erecla 24 225 555534 Drakensberg mountain range. They frequently occur imme­ Euclea crispa subsp. crispa 222 55 425 5 4 diately below waterfalls spilling over sandstone or basalt May tenus heterophylla 333344 343543 Clutia pulchella 22325 545 4 3 cliffs, from where they spread laterally along the foot of the Halleria lucida 54251 1 253531 cliffs and downward along stream banks and rivers (Killick Kiggelaria africana 55 43 441532 1963). The Afromontane forests have an archipelago-like Rhus pyroides var. gracilis 52 1 3 354313 distribution pattern (White 1983). Afromontane grasslands Buddleja salviifolia 542 12 223531 are found on the exposed slopes among the patches of Myrsiphyllum asperagoides 44 1 3 352 1 Rhamnus prinoides 35 2 243 1 1 forest, and on the mountain summits above the forests. Leucosicka sericea 33 2 223 22 The majority of the tree species in these forests are very Schoenoxiphium sparteum 122 4 1 34 2 widespread (White 1978, 1983). Among them are Hal/eria Clematis brachiata 44 1 1 5 1 lucida, /lex mitis, Kiggelaria a/deana, Podocarpus lati/olius Cheilanthes hirta 12 1 12 1 and Rapanea melanophloeos. Species endemic to South " I. PODOCARPETAllA LATIFOLll Africa, occurring in this study area, are Cryptocarya woodii A&B Clauseno·Podocarpelum tatifolii and Seolopia mundii (White 1978), and Olinia emarginata C&D Peperomio·Carissetum (Van Zinderen-Bakker 1973). This assemblage of species E&F Ehrharlo·Eucleetum crispae could almost be used to defme the Afromontane region as a

2. pmOSPORETAll!.A VIRlDIFWRUM [described by Du whole. Not one species occurs throughout, but most species Preez and Bredenkamp (1991») . of the assemblage are represented on virtually every 'island' G, H & I llici·Oleelum europaeae of Afromontane forest (White 1978, 1983). In southern 1 Plumbago.Kiggelarielum africanae Africa the species of Afromontane forests diminish from K ROOo·Slipelum dregeanae north to south, resulting in somewhat impoverished commu­ L Olea europaea·Euclea crispa forest community nities in the southern Cape (White 1978, Von Breitenbach 202 S.-Afr.Tydskr.Plantk., 1991, 57(4)

Table 2 Podocarpetalia latifolii

00000 0000 0000 00000 00000 00000000 RELEVE HUMBER 00222 3333 3334 33344 22223 20112121 12567 1234 4572 68901 67891 46919557

PHYTOSOCIOLOGIST VVVVV VVVV ZZZZ ZZZZZ ZZZZZ ZZZZVZZZ TOTAL HUMBER SPECIES/RELEVE 22323 3333 2221 21212 11111 11112111 84164 2378 1219 18261 40250 21160152

SPECIES GROUP A CLAUSENO-PODOCARPETUM LATIFOLII CHARACTER SPECIES Clausena anisata I R RI++ 11 Dryopteris inaequalis I +++1++++1 Crassula pellucida 1++ ++1 R RI Dicl is reptans 1++ +1 R++I Pteris cretica I +++1++ I Coccinia palmata I R +1 +R I Gal ium capense I R+ I + I SPECIES GROUP B DIFFERENTIAL SPECIES Buddle)a salviifolia IR2+2+1 21RI + I + I + + I May tenus acuminata '11 R++IR 2RI I + I + I Myrsiphyllum asparagoides IRR++ I +R+I I I + I Rhamnus prinoides I 2+1 1+11+1 I I + + I Clematis brachiata I R+++ I ++R I I I + I Leucosidea sericea I 232 I +1 I I I + + I SPECIES GROUP C CLAUSENO-POOOCARPETUM LATIFOLII­ BOHKERIETOSUM VERTICILLATAE CHARACTER AND DIFFERENTIAL SPECIES Bowkeria verticillata 11 + 1 I RI I Asplenium adiantum-nigrum IR ++1 I 1+ Thamnocalamus tessellatus I + 11 I I Streptocarpus pusillus I +11 I I Agapanthus campanulatus 111 I I SPECIES GROUP 0 CLAUSENO-PODOCARPETUM LATIFOLII­ ASPLENIUM MONANTHIS CHARACTER AND DIFFERENTIAL SPECIES Asplenium monanthes R 1++++1 Rubus rigidus RIR+RRI Cystopteris fragi I is I +++1 Galopina circaeoides RI+++RI Stachys caff'ra 1+ ++1 Senecio oxyri ifol ius + +11+++ I Rhus pentheri IR R+I R+ I ++11 Scadoxus punlceus + Dioscorea sylvatica ++ I +11 I Pleiopeltis schraderi I +R+I Adiantum poiretii I +++1 Zehneria scabra I RRI Thalictrum rhynchocarpum I + R I Stachys grandit'ol ia I R RI Pseudobromus sylvaticus I + 21 SPEC I ES GROU P E PEPEROMIO-CARISSETUM CHARACTER SPECIES Dais cotinifol ia I I 1++++1+222+1 Erythroxylum emarginatum I I 1+ ++1 ++++1 Opl ismenus hirtellus IR IR 1++++1+++++1 Dletes iridloides I 11 1+ I + +1 Begonia sutherlandii I 1+ I ++ I + I Andrachne oval is I I I 21 +1 SPECIES GROUP F PEPEROMIO-CARISSETUM­ CRYPTOCARYETOSUM HOODII CHARACTER SPECIES Cryptocarya woodii 3+21 Buddle)a auriculata ++ I + Hypoestes triflora ++ I SPECIES GROUP G PEPEROMIO-CARISSETUM­ SCOLOPETOSUM ZEYHERI CHARACTER SPECIES Scolopia zeyheri 1++ + + Cyperus albostriatus 1+++ SPECIES GROUP H PEPEROMIO-POLYSTCHION PUNGENTIS CHARACTER SPECIES Peperomia retusa R+RI+ 1++++1+ +++1 I Carissa bispinosa I +12 12+ +123+++1 + I R Polystichum pungens 1++ +111+1+1++++1+ + +1 I S.AfrJ.Bot., 1991, 57(4) 203

Table 2 (continued)

00000 0000 0000 00000 00000 00000000 RELEVE NUMBER 00222 3333 3334 33344 22223 20112121 12567 1234 4572 68901 67891 46919557 PHYTOSOCIOLOGIST VVVVV VVVV ZZZZ ZZZZZ ZZZZZ ZZZZVZZZ TOTAL NUMBER SPECIES/RELEVE 22323 3333 2221 21212 111 11 11112111 84164 2378 1219 18261 40250 21160152

Streptocarpus haygarthii I+R++11+ ++1+ ++1 ++ I I Prot asparagus setaceus 11 + 1+ I + +1+++++1+ I Adiantum capillus-veneris 1+ + +1 1++++1+ +1 I Impatiens hochstetteri I + 11 I + I ++++1 I Plectranthus grallatus 132+2+1+++RI ++ 1+ I 1+ 2 SPECIES GROUP I PEPEROMIO POLYSTICHION PUNGENT IS DIFFERENTIAL SPECIES Halleria lucida 1+++++1 R+RI + 1+3+++1 + I R I Ilex mitis I R R 13 I 22 I I I + I

SPECIES GROUP J ALLIANCE UNSPECIFIED DIFFERENTIAL SPECIES Euclea crispa subsp. crispa RR R 1+ 1+++++1+3++++++1 Pittosporum viridirlorum I I +2+1 ++ I SPECIES GROUP K EIIRIIARTO-EUCLEETUM CRISPAE­ PODOCARPETOSUM DIFFERENTIAL SPECIES Schoenoxiphium sparteum R I R 1+ I ++++1 R SPECIES GROUP L PODOCARPETALIA LATIFOLII CHARACTER AND DIFFERENTIAL SPECIES Podocarpus latirolius 14+ 3133341+22+1 1221221 I Olinia emarginata IR I +13 + 1++2 212242+12+ 3+ + I CanthiulII ciliatum 11+ 211 2+1++ +1+ ++1 + I + +++1 Amaranthus sp. I I I ++++ I +++++ I I ++++ +++ I Pleiopeltis macrocarpa I +++11+R+1 ++ I +1 +1+ ++ I SPECIES GROUP M SCOLOPIETEA MUNDII CHARACTER SPECIES Scolopia mundii 1+ 11 R+I++ +1++2 +1 +1 +1 + I Cassinopsis ilicirolia I I RI ++ I 2+ I +++122 + + +1 SPECIES GROUP N COMPANION SPECIES Diospyros whyteana 13 ++2IRR2+1+2+41332231++++31+2++1 + I May tenus undata I 1 RI I +1+++++1+22+ 1++2+R+++1 Myrsine arricana 12R++RI+ 1 I I 1+ ++ I +1 ++1 [hl'harta erecta I ++IR++ I I + + 1+ + 1+2++R+ +1 May tenus heterophylla I+R+ 1+ RI+ +1 +++ 1++++ 1++ +++21 Clutia pulche'lla I 11 I + 1+ ++ 1++ 12+++ ++21 Kiggelaria arricana 11+1211+2R+1 I 1+++ 21 2 ++31 Rhus pyriodes var. gracilis 1+111+1 + I I 1+ 1+ ++ +1

SPECIES GROUP 0 PITTOSPORETALEA VIRIDIFLORUM COMPANION SPECIES Heteroillorpha triroliata I +lR I I + 1+ I I + R+ +1 Rhus dentata I R+ I 1+ I +1 I + I Grewia occidental is I I 1+ +1+ +1 I + + I Asplenium aethiopicum I + RI ++ I I 1+ + I + I

v = DU PREEZ & VENTER (IN PREP.) Z = VAN ZINDEREN-BAKKER (1973)

1974) and also to the west (Du Preez & Bredenkamp 1991, in prep.). White 1978). The Scoiopietea mundii is characterized by two character White (l978) classified the vegetation of the study area as species, the evergreen phanerophytes, Scolopia mundii and an Afromontane/Highveld transition zone. In South Africa Cassinopsis i1icifoiia, both occurring in almost all syntaxa the transitional zones between Afromontane forests and classified under the class (species group P, Table I and usually much drier, lowland phytochoria are complex, with species group M, Table 2). Species of wider occurrence, some Afromontane species intruding into these phytochoria often present abundantly in the Scolopietea mundii, are (White 1973). Forest-like bushclump communities from listed under species group Q (Table 1) and species group N these areas contain many diagnostic taxa and are therefore (Table 2). classified under a different class (Du Preez & Bredenkamp, A definite moisture gradient can be observed over the 204 S.-Afr.Tydskr.Plantk., 1991, 57(4)

east-west distribution range of these forests (Figure 2). The include the woody shrub C/ausena anisata, the herbaceous DECORANA scatter diagram (Figure 3) shows a distinct Diclis rep tans, Coccinia palmata and Galium capense, the floristic discontinuity in the class. The moist eastern succulent Crassula pellucida and the ferns Dryopteris communities represent the order Podocarpetalia la tifo Iii inaequalis and Pteris cretica. The differential species (fable 1 and Figure 3). The order Pittosporetalea viridi­ (species group B, Table 2) include the tree May tenus florum (Table 1 and Figure 3), which represents the drier acuminata, the shrubs Buddleja salviifolia, Leucosidea western communities of the study area, is described by Du sericea and Rhamnus prinoides. The liana Clematis Preez and Bredenkamp (1991). brachiata and the herbaceous Myrsiplryllum asparagoides are also diagnostic. Order 1. Podocarpetalia lati/olii Order nov. The tree stratum (12-15 m) is dominated by the tall The Podocarpetalia latifolii represents the moist Afro­ Podocarpus latifolius, Kiggelaria africana, !lex mitis, May­ montane forest communities within the study area. This tenus acuminata and Scolopia mundii. Shorter trees or tall order has a wide distribution on the moist, seaward eastern shrubs (2-8 m) include Canthium ciliatum, Clausena ani­ slopes of the Drakensberg mountain range. Many studies sata, Diospyros wlryteana, Halleria lucida, May tenus have been conducted in this vegetation type (Killick 1963; heterophylla, Rhamnus prinoides and Rhus pyroides var. Edwards 1967; Geldenhuys 1983; Moll 1971, 1972; Van gracilis. Zinderen-Bakker 1973; Cooper 1982, 1985; Everard 1986; Dense layers of shorter shrubs, scrubby species or semi­ Cawe & McKenzie 1989a, b, c). Acocks (1988) classified woody herbaceous species «3 m) are represented by the this area as Highland Sourveld (Veld Type 44). These dominant Plectranthus grallatus and also Buddleja salvii­ forests have been described as 'Podocarpus latifolius climax folia, Leucosidea sericea and Myrsine africana. forest' by Killick (1963) and as 'Mountain Podocarpus The ground layer is mostly sparse or well covered with forest' by Edwards (1967). ferns such as Adianthum capillus-veneris, Asplenium aethio­ Diagnostic character species include the very prominent picum and Polystichum pungens and other herbaceous evergreen mesophanerophytes Podocarpus latifolius and species such as Impatiens hochstelteri, Peperomia retusa, Olinia emarginata and also the thorny shrub Canthium Protasparagus setaceus, StreplOcarpus haygarthii, and the ciliatum. Diagnostic herbaceous species are Amaranthus grass species Ehrharta ereeta. species and the epiphytic or lithophytic fern Pleiopeltis A few epiphytic or lithophytic species may be encounter­ macrocarpa (species group H, Table 1 and species group L, ed, including the ferns Adianthum capillus-veneris, P/eio­ Table 2). peltis macrocarpa, and also the forb Peperomia retusa. At the most hydric localities, on stream banks, the C/au­ Association 1. Clauseno-Podocarpetum lati/olii Ass. nov. seno-Podocarpetum latifolii-bowkerietosum verticillatae Type reI eve: 34 (Du Preez & Venter, in prep.) sub-ass. nov. [type releve 27 of Du Preez and Venter (in This association is hygrophylous and most typical of prep.)] can be found. At these sites the lower shrub and Afromontane forest vegetation in the study area. The ground layers are well developed and dense. The diagnostic Clauseno-Podocarpetum latifolii occurs in sheltered, moist species Agapanthus campanulatus, Asplenium adiantum­ nigrum, Bowkeria verticillate and Streptocarpus pusillus and ravines, along streams on cool south- to east-facing slopes in the endemic bamboo Thamnocalamus tessellatus (species the eastern part of the study area, on the Platberg Mountain group C, Table 2) are conspicuously present. Species group and Rensburgskop near Harrismith. The association is B (Table 1) indicates that these species can be considered as associated with coarse, sandy soils derived from sandstone character species. of the Clarens Formation. The Clauseno-Podocarpetum latifolii-asplenietosum mon­ Species groups A and B (fable 2) are diagnostic for this anthis sub-ass. nov. [type releve 32 of Du Preez and Venter association. Species group A (Table 1) indicates character (in prep.)] occurs away from the hydric sites, in typical species. The character species in group A (Tables 1 and 2), Podocarpus latifolius dominated forest and forest fringes. The many diagnostic species given in species group D •• (Table 2) and species group C (fable I) characterize this I • I • '\ sub-association. The most conspicuous of these are the , '.\ . • 295 . shrubs Rubus rigidus and Rhus pentheri, while the lianas ",.A, \ • I.". .. , ...... I \ \ •• Dioscorea sylvatica and Zehneria scabra and epiphytes ,., "'. t..... -.\ \. • ••• • 195 Adiantum poiretii and Pleiopeltis schraderi are also x .... :A .. ~.. \ \_. • •• •• o .. " \ \ .. .. • prominent. Forbs, namely Ga/opina circaeoides, Senecio 1\, \ . oxyriifolius, Stachys caffra, and the fern CyslOpteris fragilis 95 .: \. are also dominant. I \ I , ,, . , Association 2. Peperomio-Carissetum Van Zinderen­ -5+-----~----~----_+--~·L--r----~~ -5 95 195 295 395 495 Bakker (1973) moIlrt axis 1 dry Van Zinderen-Bakker (1973) described this aSSOCIatIon without designating a type. Releve 35 is chosen as lectotype. Figure 3 A DECORANA scatter diagram to show the moisture gradient and the discontinuity in the class Scolopietea mundii. This association occurs above the Oliviershoek Pass in Legend: Order 1: Podocarpetalia latifolii (.); Order 2: Pitto­ Natal on the eastern Orange Free State border. These forest sporetalea viridiflorum (e). patches extend from 1300 - 1820 m above sea-level and S.Afr.1.Bot., 1991, 57(4) 205

occur on stream banks in deep ravines and gorges on the bispinosa, Impatiens hochstetteri, Peperomio retusa, southern, south-eastern and eastern slopes in the study area. Protasparagus setaeeus and Streptocarpus haygarthii. The association occurs under humid conditions on acid Red Bed soils (Elliot Formation) with a finer texture and less Alliance unspecified organic matter and lower calcium and magnesium content Association 1. Ehrharto-Eucleetum crispae Van Zin­ than the western forest of the Orange Free State (Van deren-Bakker (1973) Zinderen-Bakker 1973). Van Zinderen-Bakker (1973) con­ Van Zinderen-Bakker (1973) did not designate a type, there­ siders this association as typical of the 'Mountain Podo­ fore releve 31 is chosen as lectotype. carpus forest' (Edwards 1967), and Killick (1963) considers In his account of forest communities of the eastern this vegetation as the climax community of the Montane Orange Free State, Van Zinderen-Bakker (1973) described Belt. the Ehrharto-Eucleetum crispae, an association of the drier The association is characterized by the species of group E ravines with coarse, pH-neutral, sandy soils (66% sand) (Table 2) and species group D (Table 1), which include the derived from Sandstone of the Clarens Formation (Cave small trees Dais cotinifolia, Erythroxylum emarginatum and Sandstone). Andrachne ovalis, the grass Oplismenus hirtellus, the The stands of forest vegetation classified under this geophyte Dietes iridioides and the herbaceous Begonia association clearly represent floristically impoverished sutherlandii. examples of the Podocarpetalia lalifolii occurring on the The tall-tree stratum (12-21 m) has a discontinuous limit of the distribution range of this order. In some cases canopy and is dominated by Podocarpus lalifolius, OUnia the vegetation is also severely damaged by grazing or utili­ emarginata, /lex mitis and Scolopia mundii. Shorter trees or zation by man (Cawe & McKenzie 1989a, b, c; King 1941; tall shrubs (3--8 m) include Canthium cilialum, Cassinopsis Van Zinderen-Bakker 1973). Within the Podoearpetalia ilicifolia, Diospyros why leana, Halleria lucida, May tenus latifolii the association has two diagnostic species, namely heterophylla and M. undala. Euclea crispa subsp. crispa and Pittosporum viridiflorum, Dense layers of shorter shrubs, scrubby species or semi­ and is also characterized by the presence of the diagnostic woody species «3 m) are represented by Carissa bispinosa species of this order and the class Scolopielea mundii. Many and Clutia pulchella. species with a wider distribution (species group Q, Table 1) The ground layer consists of ferns such as Adianthum are present in this association. These indicate a relationship capillus-veneris and Polystichum pungens and other herba­ with the Rhoelalea dentatae (Du Preez & Bredenkamp, in ceous species such as Amaranthus species, Impatiens hoeh­ prep.). stetteri and Streptocarpus haygarthii. Some of the releves classified by Van Zinderen-Bakker A conspicuous epiphyte/lithophyte is Peperomia retusa. (1973) under the Ehrharto-Eucleetum crispae, and especial­ The Peperomio-Carissetum-eryptoearyetosum woodii ly the Ehrharto-Eucleetum erispae buddlejetosum, clearly sub-ass. nov. [Type: releve 35 of Van Zinderen-Bakker belong to the Pittosporetalea viridijlorum, a new order (1973)] , occurs in typical climax forest situations. Here the described by Du Preez and Bredenkamp (1991). diagnostic species of the sub-association Cryplocarya The tree stratum is relatively low, not continuous and woodii, Buddleja aurieulata and Hypoesles trijlora (species often not higher than 8-10 m, and is dominated by Kiggel­ group F, Table 2 and species group E, Table 1) are conspic­ aria ajricana, May tenus heterophylla, and M. undata, with uously present. Olinia emarginala and Scolopia mundii also present. The The Peperomio-Carisselum-seolopielOsum zeyheri sub­ scattered shrubby layer is up to 3 m tall, with Canlhium ass. nov. [Type: reIeve 36 of Van Zinderen-Bakker (1973)J ciliatum, Cassinopsis ilicifolia, Diospyros whyteana, Euclea usually occurs at forest fringes. The diagnostic species are crispa subsp. erispa, Myrsine ajricana and the semi-woody given in species group G (Table 2) and species group F ClUlia pulehella prominent. (Table 1) . The forest margin species Seolopia zeyheri and The ground layer is poorly developed, with individuals of Cyperus albostriatus are present in this association. An the grass Ehrharta erecta, the sedge Sehoenoxiphium important additional characteristic of these forest margins is sparteum and a forb Amaranthus species scattered about. the prominence of Dais cotinifolia and Diospyros whyteana Releves compiled in stands of undamaged vegetation of this and the absence of Podocarpus latijolius, indicating the rela­ association from an inaccessible, steep ravine at Wonder­ tively drier conditions at these sites. hoek (Golden Gate) show a dominance of Podocarpus lati­ The Clauseno-Podocarpetum lalifolii and the Peperomio­ jolius and Olinia emarginata. Van Zinderen-Bakker (1973) Carissetum are floristically and ecologically related, as classified these releves under a sub-association, the Ehr­ indicated by species group H (Table 2) and species group G harto-Eucleetum erispae podocarpetosum. (Table 1), and these associations are therefore incorporated in the Peperomio-Polystichion pungentis All. nov. The Conclusions Peperomio-Carissetum is considered as the typical associa­ One class and one order are newly described. In this order, tion of the alliance. Diagnostic species of the alliance are one alliance, one association and four sub-associations are listed in species group H (Table 2). Five of the species newly described, while two associations and one sub-asso­ mentioned by Van Zinderen-Bakker (1973) as diagnostic ciation from previous studies are incorporated in this classi­ species of the Peperomio-Carissetum have a wider fication. distribution and are now considered as diagnostic of The sequence of the syntaxa from left to right in the syn­ Peperomio-Polystichion pungentis. These are Carissa optic table (Table 1) represents a gradient from moist to dry 206 S.-Afr.Tydskr.Plantk., 1991, 57(4)

forest. The DECOR ANA ordination (Figure 3) shows an and related areas with special reference to Korannaberg. Ph.D. interval between the sample plots of the two orders. The dissertation. University of the Orange Free State, Bloem­ order Pittosporetalea viridiflorwn is restricted to the area fontein. between Korannaberg (Danger Point) and Golden Gate DU PREEZ, P J. & BREDENKAMP, G.J. 1991. The syntaxono­ my and synecology of the forests in the eastern Orange Free (Figure 1), while the order Podocarpetalia lali/olii occurs State, South Africa. II. The Pittosporetalea viridiflorwn. S. Afr. from Golden Gate eastwards (Figure 1). J. Bot. 57: 207 - 212. The distribution of the two orders corresponds with the DU PREEZ, P.I. & BREDENKAMP, G.I. (in prep.) The syntax­ west to east geographical gradient and the associated gra­ onomy and synecology of the open woodland in the southern dient in rainfall (Figure 2). Although rock type (geology) and eastern Orange Free State, South Africa. III . The Rhoelaiea and soil type do influence the distribution pattern and extent d.enlatae. DU PREEZ, P.J. & VENTER, H.1.T. (in prep.). The phyto­ of these forest comm unities (Van Zinderen-Bakker 1973), sociology of the Platberg Mountain near Harrismith in the climate is considered the major environmental determinant eastern Orange Free State. regarding the floristic composition and distribution of Afro­ EDWARDS, D. 1967. A plant ecological survey of the Tugela montane forest communities in the eastern Orange Free River Basin, Natal. Mem. bOI . Surv. S. Afr. 36: 1 - 285. State. EVERARD, D.A. 1986. The effects of fIre on the Podocarpus This study represents the first comprehensive synecologi­ latifolius forest of the Royal Natal National Park, Natal Drakensberg. S. Afr. J. Bot. 52(1): 60 - 66. cal and syntaxonomical report on the Afromontane forests in GELDENHUYS, C.J. 1983. Distribution and classification of the southern Africa. It also contributes significantly to the syn­ indigenous evergreen forests. In: South African Forestry taxonomical knowledge of this vegetation type. Handbook, ed. Odendaal, P.B. pp. 262 - 268. South African Institute of Forestry, Pretoria. Acknowledgements GmBS RUSSELL, G.E., REID, C., VAN Rooy, J. & SMOOK, L. 1985. List of species of Southern African . 2nd edn. The authors are indebted to the Directors and Council of the Part 1. Mem. bot. Surv. S. Afr. 51: 1 - 152. National Museum, Bloemfontein, for financing this re­ GIDBS RUSSELL, G.E., WELMAN, W.G., RETIEF, E., search, and the financial support of the Free State branch of IMMELMAN, K.L., GERMISHUISEN, G., PIENAAR, B.J., the Wildlife Society of Southern Africa is also gratefully VAN WYK, M., NICOHOLAS, A., DE WET, C., MOG­ acknowledged. FORD, I.e. & MULVENA, 1. 1987. List of species of South­ ern African Plants. 2nd edn. Part II. Mem. bot. Surv. S. Afr. 56: 1 - 270. References GROENEWALD, G.H. 1986. Geology of the Golden Gate ACOCKS, I.P.H. 1988. Veld types of South Africa. 3rd edn. Highlands National Park. Koedoe 29: 165 - 181. Mem. bOI . Surv. S. Afr. 57: 1 - 146. HILL, M.O. 1979a. TwINSPAN - a Fortran program for arranging BEWS,l.W. 1917. The plant ecology of the Drakensberg range. multivariate data in an ordered two-way table by classification Ann. Nal. Mus. 3(3): 511 - 565 . of individuals and attributes. Dept. of Ecology and System · BEZUIDENHOUT, H. & BREDENKAMP, G.J. 1990. A recon­ atics, Cornell University, Ithaca, New York. naissance survey of the vegetation of the dolomite region in the HILL, M.O. 1979b. DECORANA - a Fortran program for Potchefstroom-Ventersdorp-Randfontein area, South Africa. de trended correspondence analysis and reciprocal averaging. Phylocoenologia 18: 387 - 403. Dept. of Ecology and Systematics, Cornell University, Ithaca, BONNETT, L.T. 1977. Die geologie van die gebied 2827C suid­ New York. oos van Winburg, Oranje-Vrystaat. M.Sc. thesis, University of KILLlCK, DJ.B. 1963. An account of the plant ecology of the the Orange Free State, Bloemfontein. Cathedral Peak Area of the Natal Drakensberg. Mem. bot. Surv. BREDENKAMP, G.I. & BEZUIDENHOUT, H. (1991). A pro­ S. Afr. 34: 1 - 146. posed procedure for the analysis of large data sets in the classi­ KING, N.L. 1941. The exploitation of the indigenous forests of fication of South African grasslands. Vegelatio (in press). South Africa. J. S. Afr. For. Ass. 6: 26 - 48. BREDENKAMP, G.I., JOUBERT, A.F. & BEZUIDENHOUT, H. MOLL, E.I. 1971. Vegetation studies in Three Rivers Region, 1989. A reconnaissance survey of the vegetation of the plains Natal. Ph.D. dissertation, University of Natal, Pietermaritzburg. in the Potchefstroom-Fochville-Parys area. S. Afr. J. BOI. 55: MOLL, E.J. 1972. The current status of mistbelt mixed Podo­ 199 - 206. carpus [atifolius forests in Natal. Bothalia 10: 596 - 598. CAWE, S.G. & MCKENZIE, B. 1989a. The afromontane forests MUELLER-OOMBOIS, D. & ELLENBERG, H. 1974. Aims and of Transkei, southern Africa. I: The importance of phyto­ Methods of Vegetation Ecology. Wiley, New York. geography and past utilization to the study of forest patches SCHELPE, E.A.C.L.E. 1943. The plant ecology of the Cathedral and a description of a sampling strategy. S. Afr. J. BOI. 55(1): Peak area. J. Nat. Univ. Coli. Sci. Soc. 3: 21 - 27. 22 - 30. VAN DER WALL, R.W.E. 1976. Die neerslagklimaat van die CAWE, S.G. & MCKENZIE, B. 1989b. The afromontane forests Oranje-Vrystaat. M.Sc. thesis, University of the Orange Free of Transkei, southern Africa. II. A floristic classification. State, Bloemfontein. S. Afr. J. Bot. 55(1): 31 - 39. VAN ZINDEREN-BAKKER, E.M. (jun.) 1973. Ecological inves­ CAWE, S.G. & MCKENZIE, B. 1989c. The afromontane forests tigations of the forest communities in the eastern Orange Free of Transkei, southern Africa. III. A structural classification. State and adjacent Natal Drakensberg. Vegelalio 28: 299 - 334. S. Afr. J. Bot. 55(1): 40 - 44 . VON BREITENBACH, F. 1974. Southern Cape Forests and COOPER, K.H. 1982. Nelson's Kop Forest: Highest altitude Trees: a Guide. Government Printer, Pretoria. indigenous forest in South Africa. Afr. Wi/dl. 36(2): 101 - 103 . WEATHER BUREAU, 1954. Klimaat van Suid-AfTika. Decl I. COOPER, K.H. 1985. The conservation status of indigenous Klimaatstatistieke. WB 19, Staatsdrukker, Pretoria. forests in Transvaal, Natal and the Orange Free State, South WHITE, F. 1978. The Afromontane Region. In: Biogeography and Africa. Wildlife Society of South Africa, Durban. Ecology of Southern Africa, ed. Werger, M.J.A. W. Junk, COWUNG, R.M. 1983. Phytochorology and vegetation history in The Hague. the south-, South Africa. J. Biogeo. 10: 393 - 419. WIDTE, F. 1983. The vegetation of Africa: a descriptive memoir DU PREEZ, P.l. (in prep.). 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