Acocks' Valley Bushveld 50 Years On

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Acocks’ Valley Bushveld 50 years on: new perspectives on the delimitation, characterisation and origin of subtropical thicket vegetation

JHJ Vlok1*, DIW Euston-Brown1 and RM Cowling1, 2

1 Department of Botany, Terrestrial Ecology Research Unit, University of Port Elizabeth, PO Box 1600, Port Elizabeth 6000, South Africa 2 Southern African Hotspots Programme, Conservation International, Terrestrial Ecology Research Unit, University of Port Elizabeth, PO Box 1600, Port Elizabeth 6000, South Africa * Corresponding author, e-mail: janvlok@mweb.co.za

Received 15 October 2002, accepted in revised form 26 November 2002

We present the approach and results of an intuitive, ments. We also compiled a list of plant species that expert-based mapping exercise to identify subtropical yielded a rich flora of 1 558 species, 20% of which are thicket (including Acocks’ (1953) Valley Bushveld, endemic to our expanded thicket biome. Consistent Noorsveld and Spekboomveld) vegetation types as fea- with previous studies, endemics were strongly associ- tures for conservation planning. The study area com- ated with succulent members of the Aizoaceae, prised 105 500km2 in southern and south-eastern South Asphodelaceae, Euphorbiaceae, Apocynaceae and Africa, the planning domain for the Subtropical Thicket Crassulaceae. We discuss our results in terms of Ecosystem Planning (STEP) Project. We developed a Acocks’ (1953) typology as well as those of more recent four-tier typological hierarchy based on geography, treatments, and comment on the evolution of subtropi- floristics, structure and grain. This yielded 112 unique cal thicket vegetation. Although some confusion regard- thicket vegetation types, 78 of which comprised thicket ing the delimitation and characterisation of thicket was clumps in a matrix of non-thicket vegetation (mosaics). resolved by this study, much more research is required By identifying mosaics, we expanded the subtropical to develop and test hypotheses on the determinants of thicket concept and increased its extent in the study thicket boundaries and the origins and evolution of area by between 1.8 and 2.8 times that of earlier assess- thicket species.

Introduction

John Acocks (1911–1979) made an extraordinary contribu- tation ‘...whose range of variation is small enough to permit tion to vegetation science in South Africa (Cowling 1999, the whole of it to have the same farming potentialities’ Hoffman and Cowling 2003). His work significantly (Acocks 1953), attracted some criticism. His criteria for dis- advanced our knowledge in three major fields: vegetation tinguishing veld types was never fully defined and often description and inter-relationships (Killick 1980), large-scale incorporated vague and untested statements on history, util- vegetation dynamics (Bond et al. 2003, Meadows 2003) and isation and dynamics (Martin and Noel 1960). Based on livestock grazing systems (Hoffman 2003). Acocks’ seminal assumed and essentially Clementsian successional relation- contribution, published in 1953, was Veld types of South ships, structurally and floristically unrelated types were Africa, a richly worded memoir that remains a standard text grouped into a single veld type (Cowling 1984). to this day. Although firmly rooted in the Clementsian para- Furthermore, the top tier of his vegetation hierarchy is prob- digm of the time, Acocks provided a uniquely South African lematic: Knysna Forest, of Afromontane affinity (White perspective of vegetation pattern and dynamics, stressing in 1978), is included as a Coastal Tropical Forest Type; Valley particular the role of grazing and fire in shaping the plant Bushveld, of subtropical affinity (Bews 1925, Moll and White cover and composition of the country. His knowledge of the 1978, Cowling 1983, Palmer 1990, Hoffman and Cowling flora, one of the richest in the world (Goldblatt 1978), was 1991), is treated as a Karroid Type; and False Fynbos extraordinary and will probably never again reside in any (Macchia) and Fynbos are placed in different units owing to one individual. the formers presumed derivation from mountain grassland, Acocks was a maverick and pioneering botanist — a man thicket and forest (Cowling 1984). His firm belief (Acocks of wide interests and strong views (Cowling 1999). His veld 1953) that contemporary vegetation patterns and dynamics type concept, described as an agro-ecological unit of vege- over much of western South Africa were underpinned by the 28 Vlok, Euston-Brown and Cowling anthropogenic demise of ‘climax’ grasslands and forests of Bushveld, especially west of the Kei River, as a savanna is tropical origin, and their subsequent replacement by shrub- problematic, given the absence of a field layer of C4 grasses lands of karroid and fynbos affinity, confused a generation of and the lack of fire as a source of defoliation. range managers and plant ecologists. Acocks (1979) did Influenced by Tinley (Heydorn and Tinley 1980), Cowling make attempts to deal with these and other problems with (1984) was the first to formalise the thicket concept in the veld types, and it is a great pity that he never lived long South African phytosociological literature. From his research enough to complete a revision of what he regarded as a pre- in the south-eastern fynbos biome, he identified Subtropical liminary and hastily compiled work (Hoffman and Cowling Transitional Thicket as a vegetation class that extended from 2003). the Kei River to the south-western Cape, and defined it as Acocks’ (1953) Valley Bushveld has been a particularly follows: (i) dominance of species of Tongoland-Pondoland problematic veld type in terms of its delimitation, origins, affinity with strong links to the Karoo-Namib (drier forms) affinities and dynamics. Acocks placed it in his Karoo and and Afromontane (wetter forms) Regions; (ii) relatively low Karroid Types, along with Succulent Karoo of the winter- regional endemism (at least in comparison to elsewhere in rainfall region, and the many karroid veld types of the semi- the fynbos biome), comprising mainly succulent species of arid to arid Great, Upper and Eastern Karoos that experi- karroid affinity; (iii) dominated by broad-leaved sclerophyl- ence warm-season rainfall. In doing this, he may have been lous shrubs, many of which have spines, and having a con- influenced by Bews (1925) who suggested that the Karoo spicuous woody vine and succulent component, especially flora was derived from the ‘xerophytic scrub’ of the eastern in drier forms; and (iv) associated with deepish, well-drained seaboard river valleys. Bews (1925) acknowledged the sub- and relatively fertile soils. Additional characteristics include tropical affinity of the ‘succulent and thorny scrub of the east- an association with fire-protected sites and a rainfall regime ern river-valleys’ (p 10) and grouped it with the savanna veg- of between 200 and 800mm yr -1, at least 20% of which falls etation of the South Africa’s subtropical summer rainfall in the winter (May–September) months. In his study area, region. Interestingly, and in contradiction to the presumed Cowling (1984) recognised two vegetation orders: Kaffrarian karroid affinities of Valley Bushveld, Acocks (1953) suggest- Thicket (weakly succulent with stronger Afromontane forest ed that the Addo Bush of the lower Sundays River Basin was affinities) and Kaffrarian Succulent Thicket (strongly succu- derived from Alexandria Forest, a Coastal Tropical Forest lent with stronger Karoo-Namib affinities). The Subtropical Type. Clearly, he was confused about the origin and affini- Transitional Thicket concept includes Acocks’ (1953) Valley ties of Valley Bushveld. Bushveld south of the Kei River, Noorsveld, Spekboomveld Tinley (1975) was the first to recognise Valley Bushveld (Figure 1), as well as the thicket vegetation in the Southern and allied types (Spekboomveld and Noorsveld) as part of a Form of Eastern Province Thornveld, False Thornveld of the ‘thicket biome’, characterised by a closed-canopy vegetation Eastern Cape, Dense Strandveld Scrub, Coastal consisting of an impenetrable tangle of shrubs and low trees Renosterveld, Coastal Fynbos and Mountain Fynbos. It is usually interwoven by woody climbers and occurring exten- distinguished from thicket to the north of the Kei by its strong sively or mosaic-like on coastal dunes, clayey soils, rock evergreeness, greater spinescence and grass cover, and outcrops and termitaria throughout subtropical and tropical higher diversity and cover of succulents (Acocks 1953, cf. Africa. He later extended this concept to include the thicket- Edwards 1967). like vegetation of temperate South Africa, including the win- Subsequent phytosociological studies in the Eastern Cape ter-rainfall region (Heydorn and Tinley 1980). Acocks (1979) have adopted Cowling’s (1984) concepts. Everard (1987) was later to refer to the latter as thicket (e.g. ‘thicket clumps recognised xeric and mesic forms of Kaffrarian Thicket and in Coastal Renosterveld’ (p 677) and ‘thicket species of the Kaffrarian Succulent Thicket in the coastal belt between Strandveld’ (p 681)), although it is not clear whether this was Humansdorp and East London. Palmer (1991a) identified merely a physiognomic term or reference to a wider concept Camdeboo Thicket from the Escarpment foothills between such as Tinley’s. Biogeographically, Moll and White (1978) Graaff Reinet and Pearston. However, despite a general included the thicket vegetation of the eastern seaboard in acceptance of the thicket concept, and an acknowledgement the Pondoland-Tongaland Region of the Indian Ocean of its subtropical affinities, its broad ecological context Coastal belt, stressing its close relationship to the subtropi- remained obscure (Midgley and Stuart-Hill 1991). In a recent cal forests that grew there in wetter parts. Ecologically, volume on the vegetation of southern Africa, subtropical Boucher and Moll (1980) included the thicket vegetation of thicket was included in a chapter on forests on the basis that the winter-rainfall fynbos biome (including the western out- ‘...(it) is not fire-prone and is functionally similar to forest, for liers of Acocks’ (1953) Valley Bushveld and Spekboomveld) example in nutrient-cycling processes and the high inci- in a ‘mediterranean shrubland’ concept (along with dence of species with vertebrate-dispersed fruits.’ (Midgley Californian chaparral and Mediterranean Basin maquis), et al. 1997 p 294). However, thicket differs from forest in that associated with base- and calcium-rich soils, and charac- (i) large herbivores (Kerley et al. 1995) and not tree falls are terised by an open-scrub overstorey of evergreen sclero- the major source of disturbance; (ii) most canopy species phyllous shrubs and an understorey of seasonal grasses regenerate by ramet recruitment (Midgley and Cowling and herbs. Huntley (1984) included Valley Bushveld in the (1993); dominant canopy species are relatively shade-intol- savanna biome, interpreting it as the xeric end of eutrophic erant (Holmes and Cowling (1993); and it grows where savanna continuum. Later, Rutherford and Westfall (1986) annual rainfall may be as low as 200mm (Acocks 1953). also included Valley Bushveld (and Spekboomveld but not The recognition by Low and Rebelo (1996) of a thicket Noorsveld) in a savanna biome. The conception of Valley biome in southern Africa was, therefore, a major break- South African Journal of Botany 2003, 69: 27–51 29 through in providing an appropriate ecological context for LANDSAT Thematic Mapper (TM) imagery (December thicket. They included in this biome, Acocks’ Valley Bushveld 1997–February 1998) for field mapping. (including its Northern Variation), Noorsveld and Prior to field work, we stratified the study area as follows. Spekboomveld, as well as the dune thickets of the eastern Firstly, we divided the area into a coastal sector (on dunes) seaboard and the fynbos/thicket mosaics and strandveld of and a mainland sector, consistent with the evidence that the coastal dunes in the Cape Floristic Region (Figure 1). dune thickets are floristically distinct from those on other They distinguished thicket from forest on the presence of a substrata (Acocks 1979, Cowling 1984, Low and Rebelo single stratum of woody plants and lower overall canopy 1996). Secondly, we divided the coastal sector into western, height, and from savanna on the absence of a well-devel- central and eastern regions and the mainland sector into oped grass component. Low and Rebelo (1996) recognised arid, valley and thicket regions, based on our intuition and only five thicket types in southern Africa (Figure 2), whose experience as well as the work of Cowling (1984) and delimitation corresponded closely with Acocks’ (1953) types, Everard (1987). Thirdly, we divided each of the coastal and did not identify thicket mosaics. Both Moll et al. (1984) regions into estuarine and dune, and each of the mainland and Cowling and Heijnis (2001) recognised fynbos/thicket regions according to the major drainage systems in the mosaic types in the Cape Floristic Region, but only on study area, namely Gouritz, Gamtoos, Sundays, Albany coastal dunes. forelands, Fish, Keiskamma, Buffels and Kei. Bews (1925) Some fifty years after the publication of Acocks’ (1953) and Acocks (1953) provide some floristic evidence for the monograph, substantial progress has been made in our floristic integrity of these drainage basins, as do the distribu- understanding of the delimitation and relationships of thicket tion patterns for selected taxa (e.g. Haworthia (Bayer 1999)). vegetation in South Africa. However, the classification and Fourthly, we allocated each of these hypothetical units into mapping of thicket was at a scale inadequate for conserva- structural types, depending on whether they were solid tion planning. The launch of the Subtropical Thicket thicket or whether thicket clumps occurred in a mosaic with Ecosystem Planning (STEP) Project in 2001 required the other vegetation types. Finally we subdivided the solid types identification of biodiversity features, including land classes, into those with and without prominent spekboom as targets for systematic conservation planning (Pressey (Portulacaria afra) cover. We subdivided the mosaics and Cowling 2001). In this paper we present the outcomes according to the possible biome-level affinity of the matrix of a vegetation mapping exercise aimed at identifying thick- vegetation. This stratification yielded a total of 180 hypothet- et land classes for the STEP Project. In doing so, we discuss ical vegetation units at the lowest rank. our vegetation typology, and the dynamics, origins and affini- The data sheet included entries on polygon number, the ties of thicket vegetation in relation to Acocks’ (1953) semi- hierarchical groups described above, floristics (including nal work, as well as research subsequent to this. We con- rare and endemic species), and categories (high, medium, clude that our research has greatly expanded and enriched low) of anthropogenic transformation and threats (grazing, the thicket concept but much more work is required to under- agriculture, alien plant infestation, mining and impound- stand the complex dynamics of thicket in ecological and evo- ments). We sampled the study area systematically between lutionary time, especially with regard to its relationships with February and December 2001 and mapped the pre-transfor- adjacent vegetation types. mation extent of each of the mapping units. We used fence- line contrasts, vegetation remnants, physiographic corre- Methods lates and the expert knowledge of landowners to extrapolate mapping units to transformed areas. These vegetation data Study area were captured in a GIS (ArcView 3.2 Redlands CA). We assessed the environmental correlates of the 112 veg- The study area comprises the planning domain for the STEP etation types by intersecting them with a digital elevation Project (Figure 1). It comprises 105 500km2 and includes model (100m x 100m scale, Computing Centre for Water that part of the thicket biome (sensu Low and Rebelo 1996) Research (CCWR)), extrapolated mean annual rainfall data (1 that coincides with the Little Karoo and Albany Centres. x 1 minute scale, CCWR) and a geology map (1:1 000 000 These are centres of diversity and endemism for succulents scale, Council for Geoscience). We also intersected our and other groups, many of which grow in thicket (Van Wyk thicket map with digitised maps of Acocks (1953) and Low and Smith 2001). The study area is one of great topograph- and Rebelo (1996) in order to assess synonymy between ical and climatic diversity and, in addition to thicket, includes these and our mapping units. vegetation associated with the fynbos, forest, Nama-karoo, succulent karoo and savanna biomes (Low and Rebelo Vegetation hierarchy and nomenclature 1996). We modified our thicket hierarchy during the course of field Field survey and data capture work. While we retained the first-tier of our hierarchy (dune vs mainland), we recognised four and not three dune units Our approach was to derive an expert-based vegetation (Figures 3a and 4a). Owing to floristic relationships, we map through direct observations in the field. This was likely included the Keiskamma region with the Fish region and to be more accurate and therefore more useful for conser- recognised a new region in the escarpment zone of the vation planning than a map based on remote sensing Sneeuwberg and Winterberg Mountains. The dune units (Ferrier 2001). We used geometrically rectified (1:100 000) were arranged along a west-east geographical gradient 30 Vlok, Euston-Brown and Cowling

Figure 1: Map of the study area (grey) showing Acocks’ (1953) Noorsveld, Spekboomveld and Valley Bushveld, Low and Rebelo’s (1996) Thicket Biome (see also Figure 2), and the extent of subtropical thicket vegetation according to this study

Figure 2: Map of the study area showing Low and Rebelo’s (1996) thicket vegetation types

associated with the increase in the proportion of summer entiated these units on the basis of structural characteristics, rain and increasing thicket shrub richness (Celastraceae, namely the relative cover of woody, grass and succulent Ebenaceae, Anacardiaceae etc.) (Heydorn and Tinley 1980, species, and the incidence of spinescence and woody lianas Cowling 1983). The mainland regions were less geographi- (see Vlok and Euston-Brown (2002) for further details). cally discrete and we recorded disjunct occurrences of some These three main structural types were further divided on of the units (Figure 4a). Our observations upheld the original the basis of vegetation grain, i.e. as solid types (unbroken second-tier of our hierarchy (arid, valley and thicket) within thicket canopy) or as mosaic types (isolated thicket clumps the mainland regional units (Figures 3b and 4b). We differ- in a matrix of a vegetation of different floristics, structure and South African Journal of Botany 2003, 69: 27–51 31

(a) THICKET (iv) Fynbos Thicket, Forest Thicket, Grassland Thicket etc. to denote the biome affinity of the matrix vegetation in Dune Mainland mosaic types. (v) Bontveld to describe mosaic types where thicket clumps are associated with matrix vegetation compris- Gouritz Algoa Albany Transfish ing an admixture of the floras of more than one biome (cf. Archibald 1955). Gouritz Gamka Groot Gamtoos Sundays Albany Fish Buffels Kei Escarpment (vi) Strandveld to describe a fragmented dune thicket in a matrix of vegetation with fynbos and succulent karoo

(b) affinities (cf. Acocks 1953). (vii) Gwarrieveld to describe highly fragmented thicket, REGIONAL UNIT (e.g. Sundays) dominated by Euclea undulata, in a matrix of succulent karoo. THICKET VALLEY THICKET ARID THICKET (viii) Doringveld to describe highly fragmented thicket with matrix of prominent Acacia karroo (doringboom) and Solid Mosaic Solid Mosaic Solid Mosaic Nama-karoo dwarf shrubs.

Floristics

With spekboom Without spekboom We compiled a preliminary checklist of the flora of thicket Grassland Savanna Forest Nama-Karoo Succulent Karoo Fynbos vegetation in the study area. This was no simple task since thicket is highly fragmented and decisions had to be made Figure 3: Typological hierarchy for the thicket vegetation classifica- about whether matrix species in mosaic types were compo- tion. (a) First two tiers (geography and biogeograpy); (b) third and nents of a thicket flora or components of the floras of the fourth tiers (structure and grain) matrix biome. First we compiled a preliminary list of species occurring in the study area from the latest version of the PRECIS database located in the National Herbarium (PRE) of the National Botanical Institute (Gibbs Russell 1985). function) (Figures 3b and 4c). We were compelled to map Next, we eliminated all species not associated with thicket only >500ha tracts of unbroken vegetation as solid thicket. vegetation and added those not included in the list. All of the The thicket bush-clumps in our mosaic units varied in densi- added species were checked for vouchers lodged in at least ty (sparse to abundant) and extent (total surface area from one of the herbaria administered by the National Botanical 1–50%). Institute. Finally, we applied the following rules to compile We noted two types of mosaic thicket (see (iv) and (v) the checklist: below for names). At certain sites the thicket occurred in a (i) We did not list sub-specific taxa; in cases where fewer coarse-grained patchwork (10–500ha stands of solid thick- than all subspecies grow in thicket vegetation, we et), but were usually restricted to fire protected sites in the included the corresponding species-level taxon (e.g. matrix vegetation. At other sites the thicket frequently Cotyledon orbicularis). occurred as small, but abundant, bush-clumps (<10ha (ii) We included only those hydrophytes associated with patches) in the matrix vegetation. We classified the mosaic areas covered in solid thicket vegetation (e.g. units on the basis of the biome-level affinity of the matrix Phragmites australis). vegetation. Finally, we subdivided the solid types according (iii) We listed all species that were dominant and common to whether spekboom was a prominent feature or not (Figure components of solid thicket vegetation, irrespective 3b); this was not required for the mosaic types since whether they were also common in adjacent biomes spekboom was very seldom dominant in isolated thicket (e.g. Pappea capensis). clumps. In those cases where spekboom had been severe- (iv) We included only those mosaic species that were either ly reduced in cover or eliminated by browsing animals, we locally abundant or specifically associated with the cor- used indicator species and fence-line contrasts to assess responding mosaic type (e.g. we included Themeda the original abundance and cover of this plant. triandra which is a dominant component of many grass- We developed a nomenclature for the thicket types that land thicket and savanna thicket types (Appendix 1) but comprised a prefix referring to a geographical or place name excluded Protea repens which is found occasionally in where the type occurred, and a suffix referring to the floristic some fynbos thicket types). or structural characteristics of the type. We used the follow- We also compiled a list of thicket biome endemics. These ing rules in allocating the suffix: included species in the list that were endemic to our study (i) Thicket indicating solid thicket and its structural form in area, and also those that extended beyond it to the east and the case of Arid-, and Valley-. west, but were nonetheless always associated with subtrop- (ii) Spekboom Thicket implying a high abundance of ical thicket vegetation (e.g. Portulacaria afra). spekboom in Valley Thicket types. We assessed the taxonomic correlates of endemism in the (iii) Spekboomveld implying a high abundance of 10 largest families by testing the null hypothesis that the fre- spekboom in Arid Thicket types. quency of endemics in a particular family would not differ 32 Vlok, Euston-Brown and Cowling

Figure 4: Map of the study area showing (a) biogeographical regions, (b) structural types, and (c) grain types South African Journal of Botany 2003, 69: 27–51 33 significantly from the frequency in the overall flora, minus solid subtropical thicket is largely replaced by savanna that family (independent sample). We used Fisher’s exact (Edwards 1967) and extensive savanna thicket mosaics. test to assess the significance of the frequency data in the 2 The absence of pronounced rainfall seasonality is a major x 2 contingency tables. correlate of subtropical thicket distribution. Overall, the temperature regime is not extreme, except for Results and Discussion parts of the Arid Thicket where absolute maximum summer temperatures may exceed 50°C (Hoffman 1989). Frost is Thicket concept rare and lowest mean minima range between 1°C Escarpment region) and 8°C (Arid Thicket) (Table 1). We concur with Tinley (1975) and Low and Rebelo (1996) in Highest mean maxima range from 26°C (Dune Thicket) to recognising a thicket biome in South Africa, based on growth 30°C in the mainland types. Subtropical thicket is seldom form and climate. Growth form characteristics include a found above 1 000m. dense canopy of largely evergreen shrubs and low trees We observed subtropical thicket vegetation growing on all (0.5–3.0m) often straddled by woody lianas, and a sparse of the geologies encountered in the study area, including the understorey of shade-tolerant herbs, mostly comprising geo- quartzitic sandstones of the Cape system that generally phytes and succulents, but also C3 and C4 grasses. Large yield highly infertile soils (Appendix 1). While subtropical succulent shrubs may dominate the canopy (e.g. thicket is generally associated with nutrient-rich soils Portulacaria afra, Crassula ovata) or emerge from it (Aloe (Cowling 1984, Palmer et al. 1988, Palmer 1991b), this may spp., Euphorbia spp.). Members of the Celastraceae, be more a consequence of plant-induced soil enrichment — Ebeneaceae and Anarcadiaceae dominate the evergreen a product of nutritious foliage and the absence of fire — than shrub component. Overall, growth form diversity is very high parent material (Cowling 1984). In the absence of fire, and (Cowling et al. 1994). This thicket characterisation is consis- given adequate moisture, subtropical thicket will colonise tent with that of Cowling (1984) for Subtropical Transitional any substratum (Euston-Brown 1995). Thicket. Hereafter, we refer to it as subtropical thicket. The spatial geography of our subtropical thicket concept dif- Our subtropical thicket biome has clear climatic correlates. fers significantly in the study area from that proposed by Low Annual rainfall varies from about 200mm (on the inland and Rebelo (1996) (Figure 1). In our study, subtropical thicket fringes of Arid Thicket) to a maximum of 1 050mm (in Dune covered some 47 465km2 (44.9% of the study area), almost Forest Thicket Mosaics in the south east) (Appendix 1). All twice that of Low and Rebelo (1996), who recognised only parts of the biome in our study area receive at least 20% 26 061km2 (24.7%). Similarly, the three Acocks (1953) veld winter rain (April–August) (Appendix 1). We suspect that in types regarded by Cowling (1984) as the core Subtropical the river basins of the strongly summer rainfall parts of the Transitional Thicket types in the study area, namely Valley eastern seaboard (cf. Acocks 1953, Low and Rebelo 1996), Bushveld, Spekboomveld and Noorsveld, covered 17 257km2

Table 1: Environmental and biological charactersitics of thicket structural types in the planning domain for the Subtropical Thicket Ecosystem Planning (STEP) Project (Figure 1). Environmental data from Appendix 1

Structural type Area No. vegetation MARc (SD)d Mean (SD)d Temperature Character species typesb (mm) modal altitide (m) regime (°C)e (ha) (%)a Min Max Dune 195 819 4 16 649 (143) 59 (52) 7.7 26.3 Sideroxylon inerme, Ischyrolepis eleocharis.

Thicket 882 769 19 25 586 (129) 293 (184) 3.9 31.2 Euphorbia triangularis, Olea europaea subsp. africana, Scutia myrtina, Rhus pallens.

Valley 2 144 565 45 49 456 (100) 380 (254) 3.3 32.3 Pappea capensis, Azima tetracantha, Rhus longispina, Digitaria eriantha, Euphorbia spp., Aloe africana, Crassula spp., Haworthia spp. Arid 1 504 744 32 22 296 (94) 576 (218) 0.9 32.6 Portulacaria afra, Euclea undulata, Euphorbia spp. (non-arborescent forms) a Of total thicket area in the planning domain b Units described in Appendix 1 c MAR = mean annual rainfall d SD = standard deviation e Lowest mean minimum and highest mean maximum temperature per annum recorded in the areas in which major Thicket vegetation units are found 34 Vlok, Euston-Brown and Cowling The distribution of the 112 thicket vegetation types in the study area thicket vegetation The distribution of the 112 Figure 5: South African Journal of Botany 2003, 69: 27–51 35

(16.4%), just over a third of our thicket extent. subtropical thicket (e.g. Olea europae ssp. africana, Euclea There are two major reasons explaining why we include a undulata) (Acocks 1953, 1979). What is clear is the strong greater proportion of the study area into subtropical thicket. climatic and floristic contrast between subtropical thicket and Firstly, we identified mosaic types not mapped as thicket by the thickets of the frost-affected Escarpment plateau and Low and Rebelo (1996). These comprised 56% of our thicket Upper Karoo. The latter have conspicuous species that are area. The remainder — solid thicket — covered 20 885km2,a seldom, if ever, found in subtropical thicket, namely Rhus figure not dissimilar to Low and Rebelo’s thicket area. Areas erosa, Diospyros austro-africana, Cussonia paniculata, mapped by Low and Rebelo as fynbos, forest, coastal Euclea coriacea, and Rhus burchellii. Palmer (1991a) renosterveld, thornveld (savanna) and karroid shrublands describes this formation in the Sneeuwberg Escarpment were often included in our mosaic types (Appendix 1). (adjacent to the western sector of our Escarpment region) as Secondly, we mapped a greater area of thicket with promi- a ‘Karoo shrubland’, implying Karoo-Namib affinities. nent spekboom (Spekboom Thicket and Spekboomveld) Werger (1980) has documented very similar vegetation (31 492km2) than the area mapped as Spekboomveld by growing on Karoo kopjes in the vicinity of the upper Orange Acocks (8 792km2) and that mapped as Spekboom Succulent River. Low and Rebelo (1996) mapped this type of thicket as Thicket by Low and Rebelo (5 013km2). Both of these treat- Valley Thicket in the upper Kei River valley (Figure 2). Our ments mapped much of our spekboom-dominated vegeta- opinion is that this formation is best described as ‘temperate tion as karroid, fynbos, savanna and renosterveld types thicket’ and should not be included in the subtropical thicket (Appendix 1). We believe that we provided a more accurate biome. estimation of the pre-transformation extent of spekboom- dominated vegetation than the earlier work (see also Mapping units and synonomy Robertson and Palmer 2002), through comprehensive field work based on the presence of remnants and indicator We mapped a total of 112 subtropical thicket vegetation species (e.g. Aloe speciosa is an excellent indicator for past types, associated with dune and mainland locations, 14 bio- spekboom dominance (Vlok and Euston-Brown 2002)). geographical regions, three structural types and two grain Owing to the high palatability of most forms of Portulacaria types (solid vs mosaic) (Appendix 1, Figure 5). We do not afra (Aucamp et al. 1978), and its slow recovery after defoli- provide detailed descriptions of our types and their environ- ation (Aucamp and Tainton 1984), spekboom-dominated mental correlates here: summary data are shown in Table 1 vegetation has suffered the greatest extent of transformation and Appendix 1, and detailed descriptions are given in Vlok in the subtropical thicket biome (Acocks 1953, Hoffman and and Euston-Brown (2002). Cowling 1990, Stuart-Hill 1992, Lloyd et al. 2002). Our mapping units corresponded in many respects to Solid thicket was most extensive in the central parts of the those of other workers, but there were also some important study area, especially in the Sundays, Groot, western differences. Acocks’ (1953) Valley Bushveld included the Escarpment and Fish regions. Mosaic types were most bulk of our Valley and Thicket structural types, especially in widespread on the western and eastern margins (Figure 4a the east (Figure 1 cf. 4b), while his Noorsveld corresponded cf. 4c). A possible reason for this pattern is the increased broadly with our Arid Thicket (Figure 1 cf. 4b), especially rainfall seasonality to the west and east, and the influence Sundays Noorsveld (Figure 5). Spekboomveld (Acocks this has on the fire regime. The relatively low-frequency but 1953) straddled both Arid and Valley Thicket (Figure 1 cf. high-intensity fires associated with fynbos and renosterveld 4b), consistent with our recognition of spekboom-dominated vegetation in the more strongly winter-rainfall west may limit vegetation in both of these two structural types the development of solid thicket (Cowling et al. 1997). In the (Spekboomveld and Spekboom Thicket, respectively). more strongly summer rainfall eastern areas, the combined Our Valley Thicket and Thicket concepts correlated well effect of regular grass fires and the impacts of larger num- with Cowling’s (1984) Kaffrarian Succulent Thicket and bers of indigenous herbivores, including megaherbivores, Kaffrarian Thicket, respectively (see also Everard 1987). may also have limited thicket expansion (Trollope 1974). A However, we recognised Dune Thicket as a separate unit further factor is the high incidence of succulence in the cen- whereas Cowling (1984) and Everard (1987) included it with tral, non-seasonal rainfall area where solid thicket domi- Kaffrarian Thicket. In this sense, we agree with Low and nates. We believe that large succulents, especially Rebelo (1996) who also recognised Dune Thicket (Figure 2). spekboom, can retard recurrent fire and may promote thick- However, Low and Rebelo (1996) included in this concept all et consolidation. of our dune mosaic types. Otherwise, there was very little We suggest that there is more than one thicket biome in correspondence between Low and Rebelo’s four other thick- South Africa. The thicket units we mapped are best et types in the study area and our structural units (Figure 2 described as falling into a subtropical thicket biome. This cf. Figure 4b). They appeared to adopt Acocks’ (1953) con- biome extends westwards, deep into the fynbos biome, as cept of Spekboomveld in their delimitation of Spekboom small patches of thicket on coastal dunes, clayey soils Succulent Thicket. (including termitaria) and lower mountain slopes (Acocks Spekboom-dominated thicket requires some elaboration. 1979, Heydorn and Tinley 1980, Cowling 1984). We are Spekboom is, in many respects, a remarkable plant capable uncertain of the affinities of the thickets of the strongly win- of dominating the thicket biomass over a wide range of cli- ter-rainfall Namaqualand uplands, these being dominated by matic and edaphic conditions. We recorded it as a dominant Ozoroa spp., Ficus spp. and other elements endemic to the component, in both Spekboom Thicket and Spekboomveld, western seaboard, in addition to species commonly found in on a wide range of geologies from sea level to 1 500m altitude, 36 Vlok, Euston-Brown and Cowling and experiencing rainfall ranging from about 100mm yr -1 to poral separation of grassland, fynbos and ‘scrub forest’ 650mm yr -1 (Appendix 1). Physiologically, spekboom is states. Our data suggest that spatial separation of subtropi- extremely drought tolerant, capable of shifting from C3 to cal thicket and fynbos in a mosaic structure is widespread CAM photosynthetic mode in response to water and NaCl along the lower slopes of the Cape Folded Belt, something stress (Ting and Hanscom 1977), increasing daylength that has not hitherto been documented. (Guralnick et al. 1984a), and increasing temperature, irrespective of moisture status (Guralnick et al. 1984b). Boundaries Moreover, owing to an unusual ability to retain foliage and conserve enzymes when severely water stressed for up to Our study was descriptive and we did not carry out any 140 days, it is capable of responding rapidly to improved soil quantitative analyses of vegetation-environment relation- moisture after prolonged droughts (Guralnick and Ting ships. However, several studies have quantified the deter- 1987). While spekboom is vulnerable to goat browsing minants of community boundaries within subtropical thicket, (Aucamp and Tainton 1984, Hoffman and Cowling 1990, and between it and adjacent biomes (Cowling and Campbell Stuart-Hill 1992, Moolman and Cowling 1994), and has been 1983, Cowling 1984, Hoffman 1989, Everard 1987, Palmer eliminated from vast areas of the subtropical thicket biome 1991b, Euston-Brown 1995). The generalisations that (Acocks 1953, Lloyd et al. 2002), it is tolerant of heavy emerge from these studies is that the boundary between solid browsing by African elephant (Loxodonta africana) (Stuart- thicket and adjacent vegetation is controlled mainly by com- Hill 1992). Little is known about its reproductive biology. plex interactions between soil depth, soil fertility, frost regime, Extensive lateral vegetative expansion, by using creeping rainfall and exposure to recurrent fire. Subtropical thicket is shoots, has successfully covered large areas in some habi- best developed in areas where annual rainfall is >200mm tats. Anecdotal information suggests that Spekboom is capa- (otherwise karroid shrublands prevail) but < ca. 800mm (oth- ble of good recruitment from wind-dispersed seed in open erwise forest prevails) provided the area is subject to little microsites where protected from domestic livestock (Midgley frost and is protected from recurrent fire. Thicket appears to and Von Maltitz 1991). We suspect that spekboom may establish more rapidly on deep, fertile soils (and hence, be have achieved local dominance after the elimination of the capable of becoming fire-proof more rapidly) but there are non-succulent component as result of dry phases during the no data to corroborate this. Thicket-clad river valleys also Holocene, periods of heavy browsing by indigenous herbi- experience a high frequency of radiation fog during the vores, followed by population collapse, and catastrophic autumn and winter months (Cowling 1984, Hoffman 1989) fires, especially where subtropical thicket abuts fire-prone but there are no data to indicate how this may benefit the fynbos. Furthermore, spekboom may be an important pio- component species. neer for subtropical thicket establishment at the thicket-fyn- Indigenous herbivores may have played a crucial role in bos biome interface, being capable of invading fynbos and determining vegetation boundaries, as do domestic livestock renosterveld under certain circumstances. Although highly today under certain management regimes (Trollope 1974, speculative at this stage, these ideas have many testable Hoffman and Cowling 1990). Several studies have shown predictions that warrant further investigation. that African elephant has a substantial impact on subtropical Of our 112 vegetation types, 78 were mosaics. This thicket composition (Stuart-Hill 1992, Moolman and Cowling reflects the high diversity of matrix communities whose com- 1994, Lombard et al. 2001, Cowling and Kerley 2002). position is under greater geological and climatic controls Indeed, these animals as well as black rhinoceros (Dicornis than solid types. Moll et al. (1984) and Cowling and Heijnis (2001) are the only other studies to map thicket mosaics, and then only on coastal dunes. Acocks (1953) described bushclump veld — thicket clumps in a grassy matrix, or our Table 2: Species endemism in the ten largest families in the thick- grassland thicket — as occupying a band of vegetation et flora of the STEP planning domain along the rims of the major river valleys in our study area, above Valley Bushveld and Noorsveld. He acknowledged Family No. spp. (%) Sig.a the existence of thicket clumps in coastal renosterveld and Non-endemic Endemic on coastal dunes throughout the fynbos biome (Acocks Total flora 1 236 (79.3) 322 (20.7) – 1953, 1979) but did not map any thicket mosaics in these Asteraceae 115 (89.8) 13 (10.2) ** areas. Acocks (1953) emphasised the karroid affinities of Apocynaceae 64 (63.4) 37 (36.4) *** Karroid Broken Veld, much of which we mapped as Aizoaceae 59 (62.8) 35 (47.3) *** Gwarrieveld, a karoo thicket mosaic. Only much later did he Crassulaceae 54 (66.7) 27 (33.3) ** emphasise strong relationship between this vegetation and Euphorbiaceae 51 (65.4) 27 (34.6) ** Noorsveld (MT Hoffman, pers. comm.). Asphodelaceae 35 (47.3) 39 (52.7) *** Interestingly, Acocks (1979) did not entertain the notion of Fabaceae 57 (85.1) 10 (14.9) ns Poaceae 63 (100) 0 (0) *** fynbos thicket mosiacs. He regarded all of the mountain fyn- Scrophulariaceae 44 (69.8) 19 (30.2) ns bos in our study area as false, in the sense that it had been Hyacinthaceae 37 (77.1) 11 (22.9) ns relatively recently derived from forest and grassland (Acocks 1953). However, in order to accommodate the huge number a Significance: Fisher Exact Test (two-tailed) of localised endemics in fynbos vegetation, he invoked a *** = P < 0.001 ‘rotating climax’ where different fire regimes enable the tem- ** = P < 0.01 ns = not significant South African Journal of Botany 2003, 69: 27–51 37 bicornis) may well have been responsible for limiting tree Floristics, affinities and origin euphorbias and large aloes to herbivore-free sites such as steep slopes and areas lacking surface water (A Kamineth, We recorded 1 558 plant species strictly associated with unpubl. data). However, evidence suggests that even under subtropical thicket vegetation in our study area. This is a rel- exceptionally high population density, elephants, unlike atively rich flora, by any standards. When standardised for goats, do not convert solid thicket into a mosaic structure area, thicket floras vary greatly in size, the largest being (Stuart-Hill 1992). We believe that megaherbivores may recorded in areas dominated by our Valley Thicket (Johnson have fragmented solid stands of our Arid Thicket, but most 1998, Johnson et al. 1999). of our Valley Thicket and Thicket types are probably much We recorded 322 endemic species in our flora, comprising more resilient to the impacts of indigenous herbivores. about 20% of the total. This is an unexpectedly high figure, Very little is known about how subtropical thicket mosaics elevated by the fact that only strict subtropical thicket species are established and maintained. Cowling et al. (1997) were included in our list. There are few other sources of data showed that thicket pioneers (mainly Rhus species) estab- on endemism in subtropical thicket. Cowling (1983) recorded lish preferentially under emergent fynbos shrubs (principally between 4% and 15% Albany Centre endemics in floras from Metalasia muricata) in our St. Francis Dune Thicket (a mosa- six phytosociologically characterised communities in Valley ic with fynbos), and that thicket invasion was an inexorable Thicket and Thicket in the Gamtoos region. Highest values process that could only be curtailed by occasional high were recorded in Valley Thicket. Hoffman and Cowling (1991) intensity fires that caused the mortality of established thick- recorded a total of 18 Albany Centre endemics (7.3% of total et individuals. In other mosaics, the distribution of thicket species) in Valley Thicket, Arid Thicket and karroid shrub- clumps is non-random, being associated with termitaria lands in the Sundays region. Endemism was again highest in (Palmer et al. 1988) or fractures in hardpans (e.g. in Grass Valley Thicket. Of the 581 species in the original Addo Ridge Bontveld on the Alexandria limestones of the Algoa Elephant National Park, 12.4% are Albany Centre endemics Basin). However, in these and other cases, nothing is known or Red Data List species (Johnson et al. 1999). Palmer about the biology of subtropical thicket invasion. We suspect (1990) recorded no Albany Centre endemics in the Arid that matrix species such as Pteronia incana, may well play Thicket of the Escarpment region. a keystone role in promoting thicket invasion by outcompet- These published data and our own observations indicate ing grasses and hence reducing fire incidence. This biology that endemism is highest in Valley Thicket, declining in of thicket invasion is a fertile topic for future research. Thicket (see also Cowling and Campbell 1983) and Arid Over much of its distribution, subtropical thicket (especial- Thicket. We suggest that, along with some of the Valley ly spekboom-dominated vegetation) occupies the interface Thicket types of the Gamtoos, Sundays and Fish regions, between the Cape fynbos and subtropical grassland and endemism would be highest in some of the mosaic types, savanna floras. Acocks (1953) suggested that the anthro- especially where the matrix vegetation is fynbos, renoster- pogenic transformation of vegetation in the warm temperate, veld or succulent karoo. These communities are especially southern parts of South Africa was causing ‘...an artificial rich in locally endemic succulents, notably species of reversal of the evolutionary replacement of the southern Haworthia (Bayer 1999) and Gasteria (Van Jaarsveld 1994). scrub vegetation (fynbos) by a sward of grass of tropical ori- Endemism is expected to be lowest in the savanna thicket gin.’ (p 4). Owing to its closed canopy, thicket would have and grassland thicket mosaics of the east, where competi- helped, in addition to low soil fertility and cool-season pre- tion from grasses and frequent fires would restrict the estab- cipitation, to maintain the island-like nature of fynbos vege- lishment of small succulent plants. These hypotheses tation. Fire proof thicket barriers may have stifled the migra- remain to be tested. tion of taxa from other fire biomes into fynbos, especially Thicket endemics are not a random assemblage taxo- those that require open microsites for recruitment (i.e. most nomically and biologically. Our results (Table 2) are consis- grasses and savanna trees). This barrier function may have tent with other studies (Cowling 1983, Hoffman and Cowling been especially important during the generally warmer and 1991, Cowling and Holmes 1991, Johnson et al. 1999) in wetter conditions of the Holocene optimum, about 4 000BP showing that endemics are strongly associated with a limit- to 5 000BP when tropical vegetation expanded throughout ed number of families, namely Aizoaceae, Asphodelaceae, the subcontinent (Scott et al. 1997). In this sense, thicket Crassulaceae, Euphorbiaceae and Apocynaceae. As most acted as a ‘semi-permeable membrane’, incorporating a of the component species in these families are succulents, subset of tropical species but preventing the expansion of there is little surprise that succulents, and to a lesser extent others. The breakdown of this barrier effect, as a conse- bulbs, dominate thicket endemics (Cowling 1983, Hoffman quence of overgrazing, has had catastrophic consequences and Cowling 1991, Cowling and Holmes 1991, Johnson et for thicket; Nama-karoo elements have invaded Arid types al. 1999). Indeed, the Albany and Little Karoo Centres, (Hoffman and Cowling 1990) and subtropical grasses have which incorporate most of our study area and the bulk of massively increased in cover in some Valley Thicket types. subtropical thicket vegetation, is renowned globally for its The latter have created savanna-like vegetation that burns high diversity of endemic succulents (Van Wyk and Smith at regular intervals, eliminating succulents and fire-sensitive 2001). While Cowling (1983) has emphasised the Karoo- shrubs (R Lechmere-Oertel, unpubl. data). Are fynbos Namib affinity of these endemics, Hoffman and Cowling islands that have lost their thicket buffer, now more threat- (1991) suggest that many are derived from subtropical line- ened by advancing savanna or karoo taxa? How recently ages. Explanations on the origin and affinity of thicket formed is grassy fynbos? endemics will require phytogeographical data for a wide 38 Vlok, Euston-Brown and Cowling range of lineages. We suspect that the endemic flora will Conclusions include ancient taxa, basal to clades that have diversified in adjacent vegetation types; as well as clusters of terminal Much progress has been made in our understanding of the taxa derived from karroid, fynbos, and forest relatives. delimitation, characterisation and origins of subtropical thick- The subtropical affinity of the thicket vegetation that we et vegetation since Acocks published his monograph in mapped is widely accepted, as is its transitional nature, 1953. Thicket confused Acocks and it still confuses us today. including components with Cape, Karoo-Namib (especially There is much work to be done, especially on the dynamics Arid Thicket) and Afromontane (especially Thicket) affinities of thicket vegetation, its ecological and evolutionary rela- (Cowling 1983, Hoffman and Cowling 1991, Palmer 1990). tionships with adjacent vegetation types, the role of indige- While Acocks (1953) proposed a Karoo origin for Valley nous herbivores — especially megaherbivores — as deter- Bushveld and related veld types, he was clearly uncertain of minants of pattern and process, and the origin and evolution this, stating that these have a ‘bushveld affinity’ and sug- of thicket plants and animals. gested the term ‘Succulent Bushveld’ to describe them (p We have identified and mapped 112 thicket vegetation 66). He was later to include them with Tropical Bushveld types in what is the core of the subtropical thicket biome. We types, along with Lowveld, a savanna of the coastal plain of are confident that these types are adequate features for con- northern KwaZulu-Natal, Swaziland and Mpumulanga (MT servation planning since each is characterised by its own Hoffman, pers. comm.). While Lowveld does include patch- floristic composition and environmental selective regime. es of succulent vegetation not dissimilar from some of our The inclusion of mosaic communities means that important

Thicket types, it differs fundamentally in the high cover of C4 processes associated with ecotones will not be ignored in grasses and its fire-driven dynamics. conservation planning (Smith et al. 2001). While our We suggest that subtropical thicket is an ancient vegeta- approach was intuitive and expert-based, and therefore not tion type derived from the Tertiary forest flora of southern readily repeatable, we believe that it has several advantages Africa after the establishment of cooler and drier climatic over rapid assessments based on remotely sensed data or conditions in the mid to late Miocene (Willis and McElwain the classification of digitised environmental themes. Such 2002). Included in the thicket region, and in outliers of sub- maps would not be able to capture patterns associated with tropical thicket vegetation elsewhere in southern Africa, are floristic changes that are unrelated to spectral characteris- a number of monotypic genera of tropical origin (e.g. tics or contemporary ecological environments. Most impor- Hippobromus, Loxostylis, Pappea, Ptaeroxylon, tantly, remote approaches do not provide the opportunity to Smellophyllum) that may have grown in Tertiary xerophytic learn more about the ecology and evolution of the biological forest and thicket, and have persisted to this day. These feature and thereby generate new information and ideas early xerophytic formations may have included arborescent about its composition, functioning and evolution. Acocks succulents (Euphorbia, Aloe), in the same way that contem- used the field as his laboratory. He knew his plants and porary Madagascan dry forest does (Koechlin et al. 1974). could therefore identify patterns that none had seen before. With increasing aridity in the Pliocene (Willis and McElwain We are proud to follow in his footsteps. 2002), and the evolution of semi-arid karoo vegetation (Scott et al. 1997), various karroid and fynbos elements would Acknowledgements — This study was funded by the Global have invaded thicket and diversified there. In mosaic com- Environment Facility through a grant administered by the University munities, where the matrix is succulent karoo, renosterveld of Port Elizabeth. Chris Berens and Trevor Wolft carried out the GIS or arid fynbos, the margins of thicket clumps support a high analyses and produced the figures. Ashley Richardson edited the figures. We thank other STEP participants, namely Andre Boshoff, diversity of shade-tolerant and locally endemic succulents in Graham Kerley, Andrew Knight, Wendy Lloyd, Tony Palmer and the Apocynanceae, Asphodelaceae and Crassulaceae Sharon Wilson, for useful discussion and insights. Tony Dold helped (pers. obs.). Tropical lineages such as certain clades in with floristics. We also thank Richard Lechmere-Oertel and Ayanda stapeliads and succulent Euphorbieae, have also diversified Sigwela for their insights. extensively in subtropical thicket vegetation (Hoffman and Cowling 1991). References Why did the tropical grass and savanna flora that also evolved in the late Tertiary (Willis and McElwain 2002, Bond Acocks JPH (1953) Veld types of South Africa. Memoirs of the et al. 2003) not overwhelm subtropical thicket? One possi- Botanical Survey of South Africa 28: 1–128 bility is that thicket pre-dated the evolution of savanna and Acocks JPH (1979) The flora that matched the fauna. Bothalia 12: grassland, and was able to resist their invasion, even when 673–709 climatic conditions favoured them. Another possibility is that Archibald EEA (1955) An ecological survey of the Addo Elephant the climatic patterns in the subtropical thicket biome have National Park. Journal of South African Botany 20: 137–154 Aucamp AJ, Howe LG, Smith DWWQ, Mostert JM (1978) The never been suitable for grassland and savanna establish- browse value of the Eastern Cape Valley Bushveld. Proceedings ment, except at the eastern margins where relatively depau- of the Grassland Society of Southern Africa 13: 91–93 perate mosaics of savanna thicket and grassland thicket are Aucamp AJ, Tainton NM (1984) Veld Management in the Valley found today. The high levels of endemism and diversity in Bushveld of the Eastern Cape. Government Printer, Pretoria, the Valley Thicket types of the Gamtoos and Sundays South Africa regions attest to a relatively long history in the area. What is Bayer B (1999) Haworthia Revisited. A Revision of a Genus. true is that much more research is required for us to com- Umdaus Press, Hatfield prehend the origins of this intriguing vegetation. Bews JW (1925) Plant Forms and their Evolution in South Africa. Longmans, London South African Journal of Botany 2003, 69: 27–51 39

Bond WJ, Midgley GF, Woodward FI (2003) What controls South Guralnick LJ, Ting IP (1987) Physiological changes in Portulacaria African vegetation — climate or fire? South African Journal of afra (L.) Jacq. during a summer drought and rewatering. Plant Botany 69: 79–91 Physiology 85: 481–486 Boucher C, Moll EJ (1980) South African mediterranean shrublands. Heydorn AEF, Tinley KL (1980) Estuaries of the Cape Part 1. In: Di Castri F, Goodall DW, Specht RL (eds) Mediterranean-Type Synopsis of the Cape coast: natural features, dynamics and uti- Shrublands. Elsevier, Amsterdam, pp 233–248 lization. CSIR Research Report 380. CSIR, Pretoria Cowling RM (1983) Phytochorology and vegetation history in the Hoffman MT (1989) Vegetation Studies and the Impact of Grazing in south-eastern Cape, South Africa. Journal of Biogeography 10: the Semi-arid Eastern Cape. PhD thesis, University of Cape 393–419 Town, Cape Town, South Africa Cowling RM (1984) A syntaxonomic and synecological study in the Hoffman MT (2003) ‘Nature’s method of grazing’. Non-Selective Humansdorp region of the Fynbos Biome. Bothalia 15: 175–227 Grazing (NSG) as a means of veld reclamation in South Africa. Cowling RM (1999) Classics in physical geography revisited. South African Journal of Botany 69: 92–98 Acocks JPH 1953: Veld types of South Africa. Memoirs of the Hoffman MT, Cowling RM (1990) Desertification in the lower Botanical Survey of South Africa 28, 1–128. Progress in Sundays River Valley, South Africa. Journal of Arid Physical Geography 23: 251–255 Environments 19: 105–117 Cowling RM, Campbell BM (1983) A comparison of fynbos and non- Hoffman MT, Cowling RM (1991) Phytochorology and endemism fynbos coenoclines in the lower Gamtoos River Valley, south- long aridity and grazing gradients in the lower Sundays River eastern Cape, South Africa. Vegetatio 53: 161–178 Valley, South Africa: implications for vegetation history. Journal Cowling RM, Heijnis CE (2001) Broad Habitat Units as biodiversity of Biogeography 18: 189–201 entities for conservation planning in the Cape Floristic Region. Hoffman MT, Cowling RM (2003) ‘Enough to be considered useful’: South African Journal of Botany 67: 15–38 John Acocks’ contribution to South African botany. South African Cowling RM, Holmes PM (1991) Subtropical thicket in the south Journal of Botany 69: 1–6 eastern Cape: a biogeographical perspective. In: Zacharias PJK, Holmes PM, Cowling RM (1993) Effects of shade on seedling Stuart-Hill GC, Midgley JJ (eds) Proceedings of the First Valley growth, morphology and leaf photosynthesis in six subtropical Bushveld/Subtropical Thicket Symposium. Grassland Society of thicket species from the eastern Cape, South Africa. Forest Southern Africa, Pietermaritzburg, pp 3–4 Ecology and Management 61: 199–220 Cowling RM, Kerley GIH (2002) Impacts of elephants on the flora Huntley BJ (1984) Characteristics of South African biomes. In: and vegetation of subtropical thicket in the Eastern Cape. In: Booysen P de V, Tainton NM (eds) Ecological Effects of Fire in Kerley G, Wilson S, Massey A (eds), Elephant Conservation and South African Ecosystems. Springer, Berlin, pp 1–17 Management in the Eastern Cape. Workshop proceedings, Johnson CF (1998) Vulnerability, Irreplaceability and Reserve selec- Terrestrial Ecological Research Unit. Report No. 35, University of tion for the Elephant-impacted Flora of the Addo Elephant Port Elizabeth, pp 55–72 National Park, Eastern Cape, South Africa. MSc thesis, Rhodes Cowling RM, Kirkwood D, Midgley JJ, Pierce SM (1997) Invasion University, Grahamstown, South Africa and persistence of bird-dispersed, subtropical thicket and forest Johnson CF, Cowling RM, Phillipson PB (1999) The flora of the species in fire-prone fynbos. Journal of Vegetation Science 8: Addo Elephant National Park, South Africa: are threatened 475–488 species vulnerable to elephant damage? Biodiversity and Cowling RM, Mustart PJ, Laurie H, Richards MB (1994) Species Conservation 8: 1447–1456 diversity, functional diversity and functional redundancy in fynbos Kerley GIH, Knight MH, De Kock M (1995) Desertification of sub- communities. South African Journal of Science 90: 333–337 tropical thicket in the Eastern Cape, South Africa: Are there alter- Edwards D (1967) A plant ecological survey of the Tugela River natives? Environmental Monitoring and Assessment 37: Basin. Memoirs of the Botanical Survey of South Africa 36: 211–230 1–285 Killick DJB (1980) John Phillip Harrison Acocks (1911–1979). ERDAS Imagine (1997) ERDAS Field Guide (4th edn). Fourth Bothalia 13: 239–244 Edition. ERDAS Inc., Atlanta, Georgia Koechlin J, Guillaumet J-L, Morat P (1974) Flore et Végétation de Euston-Brown DIW (1995) Environmental and Dynamic Madagascar. Cramer, Liechtenstein Determinants of Vegetation Distribution in the Kouga and Lloyd JW, Van den Berg EC, Palmer AR (2002) STEP Project — Baviaanskloof Mountains, Eastern Cape. MSc Thesis, University Patterns of transformation and degradation in the Thicket Biome. of Cape Town, Cape Town, South Africa ARC Report No. GW/A/2002/30 Everard DA (1987) A classification of the subtropical transitional Lombard AT, Johnson CF, Cowling RM, Pressey RL (2001) thicket in the eastern Cape, based on syntaxonomic and structur- Protecting plants from elephants: botanical reserve scenarios al attributes. South African Journal of Botany 53: 329–340 within Addo Elephant National Park. Biological Conservation Ferrier S (2001) Mapping spatial pattern in biodiversity for regional 102: 191–203 conservation planning: where to from here? Systematic Biology Low AB, Rebelo AG (1996) Vegetation of South Africa, Lesotho and 51: 331–363 Swaziland. Department of Environmental Affairs and Tourism, Gibbs Russell GE (1985) PRECIS: The National Herbarium’s Pretoria Computerised Information System. South African Journal of Martin ARH, Noel ARA (1960) The Flora of Albany and Bathurst. Science 81: 62–65 Rhodes University, Grahamstown Goldblatt P (1978) An analysis of the flora of southern Africa: its Meadows ME (2003) John Acocks and the expanding Karoo hypoth- characteristics, relationships and origins. Annals of the Missouri esis. South African Journal of Botany 69: 62–67 Botanical Garden 65: 369–436 Midgley JJ, Cowling RM (1993) Regeneration patterns in Cape sub- Guralnick LJ, Rorabaugh PA, Hanscom Z (1984a) Influence of pho- tropical thicket: where are all the seedlings? South African toperiod and leaf age on Crassulacean Acid Metabolism in Journal of Botany 59: 496–499 Portulacaria afra (L.) Jacq. Plant Physiology 75: 454–457 Midgley JJ, Cowling RM, Seydack AHW, Van Wyk GF (1997) Guralnick LJ, Rorabaugh PA, Hanscom Z (1984b) Seasonal shifts of Forest. In: Cowling RM, Richardson DM, Pierce SM (eds) photosynthesis in Portulacaria afra (L) Jacq. Plant Physiology Vegetation of Southern Africa. Cambridge University Press, 76: 643–646 Cambridge, pp 278–295 40 Vlok, Euston-Brown and Cowling

Midgley JJ, Stuart-Hill GC (1991) Spekboom, megaherbivores, objective characterisation. Memoirs of the Botanical Survey of Acocks and goats: The first valley bushveld symposium. South South Africa 54: 1–98 African Journal of Science 87: 510–552 Scott L, Anderson HM, Anderson JM (1997) Vegetation history. In: Midgley JJ, Von Maltitz G (1991) Life and death in valley bushveld. Cowling RM, Richardson DM, Pierce SM (eds) Vegetation of Veld & Flora 77: 12–13 Southern Africa. Cambridge University Press, Cambridge, pp Moolman J, Cowling RM (1994) The impact of elephant and goat 62–84 grazing on the endemic flora of South African succulent thicket. Smith TB, Kark S, Schneider CJ, Wayne RK (2001) Biodiversity Biological Conservation 68: 53–61 hotspots and beyond: the need for preserving environmental tran- Moll EJ and White F (1978) The Indian Ocean Belt. In: Werger MJA sitions. Trends in Ecology and Evolution 16: 431 (ed) Biogeography and Ecology of Southern Africa. Junk, The Stuart-Hill GC (1992) Effects of elephants and goats on the Hague, pp 561–598 Kaffrarian succulent thicket of the eastern Cape, South Africa. Moll EJ, Campbell BM, Cowling RM, Bossi L, Jarman ML, Boucher Journal of Applied Ecology 29: 699–710 C (1984) A description of the major vegetation categories in and Ting IP, Hanscom Z (1977) Induction of acid metabolism in adjacent to the fynbos biome. South African National Scientific Portulacaria afra. Plant Physiology 59: 511–514 Programmes report 83, CSIR, Pretoria Tinley KL (1975) Habitat physiognomy, structure and relationships. Palmer AR (1990) A qualitative model of vegetation history in the University of Pretoria Publications New Series 97: 67–77 eastern Cape midlands, South Africa. Journal of Biogeography Trollope WSW (1974) Role of fire in preventing bush encroachment 17: 35–46 in the eastern Cape. Proceedings of the Grassland Society of Palmer AR (1991a) A syntaxonomic and synecological account of South Africa 9: 67–72 the vegetation of the eastern Cape midlands. South African Van Jaarsveld EJ (1994) Gasterias of South Africa. A New Revision Journal of Botany 57: 76–94 of a Major Succulent Group. Fernwood Press, Newlands Palmer AR (1991b) Vegetation/environment relationships in the cen- Van Wyk AE, Smith GF (2001) Regions of Floristic Endemism in tral area of the Cape midlands, South Africa. Coenoces 6: 29–38 Southern Africa. Umdaus Press, Hatfield Palmer AR, Crook BJS, Lubke RA (1988) Aspects of the vegetation Vlok JHJ, Euston-Brown DIW (2002) Report by biological survey and soil relationships in the Andries Vosloo Kudu Reserve, Cape component for conservation planning for biodiversity in the thick- Province. South African Journal of Botany 54: 309–314 et biome. Unpublished report, Regalis Environmental Services, Pressey RL, Cowling RM (2001) Systematic conservation planning Oudtshoorn for the Subtropical Thicket Ecosystem Planning (STEP) Project: Werger MJA (1980) A phytosociological study of the Upper Orange a conceptual and methodological basis. Terrestrial Ecological River Valley. Memoirs of the Botanical Survey of South Africa Research Unit. Report No. 31, University of Port Elizabeth, Port 46: 1–98 Elizabeth White F (1978) The Afromontane Region. In: Werger MJA (ed) Robertson MP, Palmer AR (2002) Predicting the extent of succulent Biogeography and Ecology of Southern Africa. Junk, The Hague, thicket under current and future climate scenarions. African pp 465–510 Journal of Range and Forage Science 19: 21–28 Willis KJ, McElwain JC (2002) The Evolution of Plants. Oxford Rutherford MC, Westfall RH (1986) Biomes of southern Africa. An University Press, Oxford

Edited by MT Hoffman and RM Cowling South African Journal of Botany 2003, 69: 27–51 41 f Synonomy Coastal Macchia, Coastal Renosterbos-veld, Knysna Macchia, Coastal Coastal Forest, Dune Afromontane Forest (Acocks 1953); Fynbos, South Thicket, Limestone Fynbos, Mountain and South-west Coast Renosterveld (Low Rebelo 1996) Alexandria Forest, False Macchia, Succulent Karoo, Forest, Afromontane Bushveld (Acocks 1953); Valley Thicket, Grassy Fynbos, Mesic Succulent Dune Thicket, South and South-west Coast Renoster-veld, Thicket (Low and Rebelo 1996) Valley Thornveld, Alexandria Forest, Eastern Province Bushveld (Acocks 1953); Succulent Karoo, Valley Thicket, Grassland, Dune Forest, Coastal Coastal Thicket, Thorn Bushveld, Mesic Succulent Eastern Thicket (Low and Rebelo 1996) Valley Thornveld, Forest and Alexandria Forest, Coastal Grassland, Bushveld (Acocks 1953); Coastal Valley Thorn Bushveld (Low and Thicket, Eastern Dune Rebelo 1996) Renosterbos-veld (Acocks Macchia, Coastal Coastal Thicket, Limestone Fynbos, South and 1953); Dune South-west Coast Renosterveld (Low and Rebelo 1996) Renosterbos-veld, Valley Macchia, Coastal Coastal Thicket, South and Bushveld (Acocks 1953); Dune South-west Coast Renosterveld (Low and Rebelo 1996) Renosterbosveld, Knysna Forest (Acocks Coastal Thicket (Low and Forest, Dune Afromontane 1953); Rebelo 1996) Macchia, Knysna Forest (Acocks 1953); Coastal Thicket, South and South- Forest, Dune Afromontane west Coast Renosterveld (Low and Rebelo 1996) Alexandria Forest, False Macchia (Acocks 1953); Thicket, Grassy Fynbos (Low and Rebelo 1996) Dune Forest, Afromontane Alexandria Forest (Acocks 1953); Thicket, Grassy Fynbos, South and South-west Dune Coast Renosterveld (Low and Rebelo 1996) Bushveld Alexandria Forest, Succulent Karoo, Valley (Acocks 1953); Grassy Fynbos, Mesic Succulent Thicket, South and South-west Coast Renosterveld (Low and Rebelo 1996) Thornveld (Acocks Alexandria Forest, Eastern Province Thicket, Mesic Succulent Forest, Dune 1953); Coastal Thicket (Low and Rebelo 1996) Thicket, Valley Thornveld, Valley Alexandria Forest, Eastern Province Forest, Coastal Bushveld (Acocks 1953); Coastal Thorn Bushveld, Thicket, Eastern Grassland, Dune Thicket (Low and Thicket, Valley Mesic Succulent Rebelo 1996) nis pidatus eleocharis occidentalis Rhus lucida Ischyrolepis Mystroxylon Aloe africana Rhus crenata Mimusops caffra, Mimusops Lycium cinereum, Lycium Themeda triandra Cynodon dactylon Dominant species Sideroxylon inerme Sideroxylon inerme Sideroxylon inerme Sideroxylon inerme, Sideroxylon inerme, Sideroxylon inerme, Sideroxylon inerme, Sideroxylon inerme, Sideroxylon inerme, Sideroxylon inerme, aethiopicum, Grewia Brachylaena discolor Pterocelastrus tricus- Agathosma apiculata, Thamnochortus insig- e spp. , lia fourcadei Schotia afra, Cordia caffra Acacia karroo Strelitzia nicolai Strelitzia Diosma aristata, Erica chloroloma Lycium cinereum Lycium Euphorbia bayeri Mimusops caffra, Mimusops Rapanea gilliana, Rapanea Agathosma muirii Lycium cinereum, Lycium Nylandtia spinosa Nylandtia Cotyledon adscen- Jordaaniella dubia, Brunsvigia orientalis Ekebergia capensis, Celtis africana, Erica Leucospermum Brachylaena discolor, Allophyllus natalensis dens, Morella cordifo- Gymnosporia capitata 10 30 10 10 20 10 70 50 20 70 140 170 140 (0–80) (0–180) (0–230) (0–190) (0–100) (0–260) (0–210) (0–200) (0–200) (0–130) (0–220) (30–150) (120–290) (min-max)(m) d L L L L L L L L H M M M M Summer Modal altitude Character species ) -1 c 510 712 647 775 503 548 762 457 745 829 385 586 671 (mmyr (min-max) rainfall (400–640) (381–964) (406–801) (669–987) (365–692) (406–730) (638–887) (348–858) (627–909) (329–467) (558–639) (534–860) (487–1044) b (%) Geology MAR Q (28) TL (75) TL (25) FS (34) PS (41) PS (68) PS (63) PS (68) PS (98) PS (56) RS (48) RS (19) RS (75) CE (18) RS (97) RS (33) FQ (14) RS (81) RS (16) RS (26) RS (84) FM (39) RS (100) a 0.07 0.15 0.12 0.11 1.06 0.09 0.18 0.11 0.19 0.05 0.10 0.05 0.48 4 118 9 110 8 419 5 280 2 404 4 697 2 260 3 171 7 295 5 839 5 075 22 796 49 935 Type Area (ha) % 1.Gouritz Dune Thicket 2.Algoa Dune Thicket 3.Albany Dune Thicket 4.Transfish Dune Thicket Bay Dune 5.Still Thicket 6.Hartenbos Strandveld 7.Goukamma Dune Thicket 8.Robberg Dune Thicket Francis 9.St. Dune Thicket 10.Sardinia Forest Thicket 11.Colchester Strandveld 12.Zuney Strandveld 13.Kasouga Dune Thicket Thicket vegetation types in the planning domain for the Subtropical Thicket Ecosystem Planning (STEP) Project (Figure 1) showing environmental and biological characteris- Thicket Ecosystem Planning (STEP) Project (Figure 1) showing environmental types in the planning domain for Subtropical Thicket vegetation Region Appendix 1: tics. Types are listed according to their location along an east-west axis (Figure 5). See text and Figures 3a b for explanation of the hierarchy Types tics. DUNE TYPES Dune solid Gouritz Algoa Albany Transfish Dune mosaic Gouritz Gouritz Gouritz Gouritz Algoa Algoa Algoa Albany Albany 42 Vlok, Euston-Brown and Cowling f Synonomy Alexandria Forest, Coastal Forest and Thornveld, Forest and Alexandria Forest, Coastal Bushveld (Acocks Thornveld, Valley Eastern Province Grassland, Dune Forest, Coastal 1953); Coastal Thicket (Low Thorn Bushveld, Valley Thicket, Eastern and Rebelo 1996) Thornveld, Forest and Alexandria Forest, Coastal Forest, Bushveld (Acocks 1953); Coastal Valley Thicket (Low Thicket, Valley Grassland, Dune Coastal and Rebelo 1996) Bushveld Thornveld, Valley Forest and Coastal Grassland, Forest, Coastal (Acocks 1953); Coastal Thicket Thorn Bushveld, Valley Thicket, Eastern Dune (Low and Rebelo 1996) Bushveld Alexandria Forest, False Macchia, Valley Thicket, Forest, Dune Afromontane (Acocks 1953); Grassy Fynbos, South and South-west Coast Thicket (Low and Rebelo 1996) Renosterveld, Valley Thornveld, False Macchia, Succulent Eastern Province Afromontane Bushveld (Acocks 1953); Karoo, Valley Thorn Bushveld, Grassy Fynbos, Forest, Eastern Thicket, South and South-west Coast Mesic Succulent Thicket (Low and Renosterveld, Xeric Succulent Rebelo 1996) Eastern Province Grassveld, False Central Lower Thornveld of False Karoo, False Karroid Broken Veld, Eastern Province, Highland Sourveld and Dohne Veld, Sourveld, Karroid Danthonia Mountain Bushveld (Acocks 1953); Valley Spekboomveld, Forest, Central Nama Karoo, Dune Afromontane Thorn Thicket, Eastern Mixed Nama Karoo, Bushveld, Moist Upland Grassland, South-eastern Thicket, Succulent Grassland, Spekboom Mountain Thicket, Xeric Subarid Thorn Bushveld, Valley Thicket (Low and Rebelo 1996) Succulent Thornveld, False Alexandria Forest, Eastern Province Afromontane Bushveld (Acocks 1953); Macchia, Valley Grassland, Dune Forest, Coastal Forest, Coastal Thorn Bushveld, Grassy Fynbos, Thicket, Eastern Thicket, Xeric Thicket, Valley Mesic Succulent Thicket (Low and Rebelo 1996) Succulent Thornveld, False Alexandria Forest, Eastern Province Thornveld of Eastern Province, Valley Macchia, False Forest, Coastal Afromontane Bushveld (Acocks 1953); Thicket, Eastern Grassland, Dune Forest, Coastal Thicket, Xeric Thorn Bushveld, Grassy Fynbos, Valley Thicket (Low and Rebelo 1996) Succulent Thornveld, Eastern Province Forest and Coastal Thornveld, Highland Sourveld and Dohne Sourveld, Forest, Afromontane Bushveld (Acocks 1953); Valley Thicket, Grassland, Dune Forest, Coastal Coastal Thicket (Low and Thorn Bushveld, Valley Eastern Rebelo 1996) africana africana, africana, ens nicolai gularis phoratus Acacia karroo, Olea europaea Olea europaea Anthospermum subsp. subsp. subsp. Harpephyllum caf- Eugenia capensis, Euphorbia triangu- Dominant species Grewia occidentalis Hippobromus pauci- laris, Olea europaea Tarchonanthus cam- Tarchonanthus Plumbago auriculata Gymnosporia buxifolia, Allophylus decipi- aethiopicum, Strelizia frum, Euphorbia trian- florus, Schotia latifolia Harpephyllum caffrum e africana, inerme mucronata Scutia myrtina Olea europaea laris, Canthium ceum, Ziziphus Erythrina caffra subsp. Bobartia gracilis Cussonia spicata, Phoenix reclinata, Watsonia pillansii, Watsonia laris, Acacia caffra Euphorbia triangu- Elaeodendron cro- Euphorbia triangu- Euphorbia triangularis, Schotia latifolia Acmadenia kiwanen- sis, Phoenix reclinata Euphorbia triangularis 60 60 90 50 20 190 750 180 200 (0–190) (0–230) (0–260) (0–300) (20–740) (10–840) (10–510) (10–610) (460–1650) (min-max)(m) d L H H H H H M M M Summer Modal altitude Character species ) -1 c 693 656 913 641 486 576 555 539 719 (mmyr (min-max) rainfall (559–894) (581–770) (432–886) (257–704) (375–696) (394–709) (609–876) (810–1048) (280–1069) b (%) Geology MAR FS (26) FS (32) FS (67) FS (20) KS (51) PS (16) KS (66) KS (64) KS (87) PS (18) PS (16) KS (62) KS (82) RS (15) CE (33) CE (14) KD (13) KD (14) KM (16) KM (29) KM (16) a 0.75 0.27 0.36 0.19 1.00 2.23 1.52 0.51 0.78 8 940 35 635 12 746 17 039 47 490 71 893 24 072 36 748 105 522 Type Area (ha) % 14.Hamburg Dune Thicket 15.Kiwane Dune Thicket Dune 16.Cintsa Thicket 17.Gamtoos Thicket 18.Sundays Thicket 19.Escarpment Thicket 20.Albany Thicket 21.Fish Thicket 22.Buffels Thicket Appendix 1 cont. Region Transfish Transfish Transfish Thicket solid Gamtoos Sundays Escarpment Albany Fish Buffels MAINLAND SOLID TYPES South African Journal of Botany 2003, 69: 27–51 43 f Synonomy Coastal Forest and Thornveld, Valley Bushveld Thornveld, Valley Forest and Coastal Thicket (Low Forest, Valley (Acocks 1953); Coastal and Rebelo 1996) Renosterbos-veld, False Macchia, Coastal Coastal Bushveld (Acocks 1953); Limestone Macchia, Valley Fynbos, South and South-west Coast Renosterveld (Low and Rebelo 1996) (Acocks 1953); Grassy False Macchia, Spekboomveld Thicket (Low and Fynbos, Valley Fynbos, Mountain Thicket Rebelo 1996); Gamtoos Mesic Succulent (Cowling and Heijnis 2001) Alexandria Forest, False Macchia, Spekboomveld, Forest, Afromontane Bushveld (Acocks 1953); Valley Thicket, Grassy Fynbos, Mesic Succulent Dune Fynbos, South and South-west Thicket, Mountain Thicket (Low and Rebelo Coast Renosterveld, Valley 1996) Thornveld, False Macchia, Eastern Province Bushveld (Acocks 1953); Central Noorsveld, Valley Nama Karoo, Grassy Fynbos, Mesic Succulent Thicket, South and South-west Coast Renosterveld, Thicket (Low and Rebelo 1996) Xeric Succulent Thornveld, False Karroid Broken Eastern Province Bushveld Valley False Macchia, Spekboomveld, Veld, Forest, Forest, Coastal Afromontane (Acocks 1953); Grassland, Eastern Mixed Nama Karoo, Coastal Thorn Bushveld, Grassy Fynbos, Mesic Eastern Thicket, Xeric Succulent Thicket, Valley Succulent Thicket (Low and Rebelo 1996) Thornveld of Eastern Province, Highland False Bushveld Sourveld and Dohne Sourveld, Valley Thorn Forest, Eastern Afromontane (Acocks 1953); Thorn Bushveld, Moist Upland Grassland, Subarid Thicket (Low Thicket, Xeric Succulent Bushveld, Valley and Rebelo 1996) Central Lower Karoo, False Karroid Danthonia False Karroid Broken Veld, (Acocks 1953); Central Spekboomveld Veld, Mountain Grassland, Nama Karoo, South-eastern Mountain Thicket (Low and Thicket, Valley Succulent Spekboom Rebelo 1996) Thornveld, Eastern Province Forest and Coastal Thornveld of Eastern Province, Thornveld, False Highland Sourveld and Dohne Sourveld, Valley Forest, Eastern Afromontane Bushveld (Acocks 1953); Thorn Bushveld, Moist Upland Grassland, Valley Thicket (Low and Rebelo 1996) Renosterbosveld, Mountain Karroid Broken Veld, (Acocks 1953); Little Succulent Karoo, Spekboomveld Thicket (Low and Rebelo 1996) Succulent Spekboom africana undulata phoratus Aloe speciosa, Grewia robusta subsp. Schotia latifolia, Euclea undulata Portulacaria afra Rhus longispina, Portulacaria afra, Pappea capensis Pappea capensis Pappea capensis, Pappea capensis, Euphorbia triangu- Dominant species Gloveria integrifolia Tarchonanhus cam- Tarchonanhus Sideroxylon inerme, Schotia afra, Euclea laris, Olea europaea Euphorbia curvirama Euphorbia tetragona, e sis didens rotundifolia Schotia afra Rhus glauca, Aloe africana Aloe speciosa Boscia oleoides Schotia afra, Aloe Schotia afra, Euphorbia grandi- Pappea capensis, Pappea capensis, Harpephyllum caf- Euphorbia ledienii, Ozoroa mucronata, Cussonia thyrsiflora africana, Kalanchoe sis, Euphorbia gran- frum, Aloe barberiae Schotia brachypetala dens, Atalaya dens, Atalaya capen- Lycium ferocissimum, Lycium Cussonia gamtoosen- 120 130 400 290 150 260 670 780 370 450 (0–630) (10–370) (10–300) (10–550) (20–910) (210–940) (340–670) (170–1100) (120–1080) (690–1010) (min-max)(m) d L L L L H H H M M M Summer Modal altitude Character species ) -1 c 777 413 378 530 440 513 534 332 608 300 (mmyr (min-max) rainfall (689–843) (315–533) (254–550) (372–730) (276–697) (382–652) (391–750) (224–429) (522–758) (230–458) b (%) Geology MAR AL (50) AL (13) FS (19) FS (24) FS (88) FS (51) VS (24) KS (95) KS (99) KS (95) FQ (33) CE (26) CE (50) KD (97) CE (21) FM (50) CM (53) a 0.04 0.56 0.37 0.70 1.23 1.97 0.91 1.34 0.46 0.34 2 084 26 305 17 634 33 281 58 099 92 918 43 053 63 564 21 551 15 929 Type Area (ha) % 23.Kei Thicket 24.Gouritz Thicket Valley 25.Baviaans Thicket Valley 26.Gamtoos Thicket Valley 27.Sundays Thicket Valley 28.Albany Valley Thicket 29.Fish Valley Thicket 30.Escarpment Thicket Valley Valley 31.Buffels Thicket 32.Gamka Spekboom Thicket Valley solid with spekboom Valley Kei solid Valley Gouritz Groot Gamtoos Sundays Albany Fish Escarpment Buffels Gamka Appendix 1 cont. Region 44 Vlok, Euston-Brown and Cowling f Synonomy False Macchia, Karroid Broken Veld, Mountain False Macchia, Karroid Broken Veld, Renosterbosveld, Noorsveld, Succulent Karoo, Bushveld (Acocks 1953); Valley Spekboomveld, Central Nama Karoo, Grassy Fynbos, Mountain Fynbos, South and South-west Coast Renosterveld, Thicket, Xeric Thicket, Valley Succulent Spekboom Thicket (Low and Rebelo 1996) Succulent False Macchia, Karroid False Karroid Broken Veld, Valley Noorsveld, Spekboomveld, Broken Veld, Forest, Central Afromontane Bushveld (Acocks 1953); Succulent Nama Karoo, Grassy Fynbos, Spekboom Thicket (Low Thicket, Xeric Succulent Thicket, Valley and Rebelo 1996) Thornveld, False Macchia, Karroid Eastern Province Noorsveld, Succulent Karoo, Broken Veld, Bushveld (Acocks 1953); Valley Spekboomveld, Thorn Bushveld, Mesic Central Nama Karoo, Eastern Thicket, Xeric Succulent Thicket, Spekboom Succulent Thicket (Low and Rebelo 1996) Succulent Central Lower Karoo, False Karroid Danthonia False Karroid Broken Veld, (Acocks 1953); Central Spekboomveld Veld, Mountain Grassland, Nama Karoo, South-eastern Mountain Thicket (Low and Thicket, Valley Succulent Spekboom Rebelo 1996) False Central Lower Karoo, Karroid Broken Veld, Bushveld (Acocks 1953); False Macchia, Valley Forest, Eastern Mixed Nama Karoo, Afromontane Thicket (Low and Rebelo 1996) Xeric Succulent Alexandria Forest, Central Lower Karoo, Eastern Thornveld, Province Grassveld, Eastern False Central Lower Karoo, Karroid Broken Veld, Thornveld of Eastern Province, False Macchia, Highland Sourveld and Dohne Sourveld, Karroid Veld, Karroid Danthonia Mountain Broken Veld, Bushveld (Acocks Valley Noorsveld, Spekboomveld, Forest, Central Nama Karoo, Afromontane 1953); Thicket, Grassland, Dune Forest, Coastal Coastal Thorn Bushveld, Eastern Mixed Nama Karoo, Grassy Fynbos, Moist Upland Grassland, South-east- Succulent Grassland, Spekboom ern Mountain Thicket, Xeric Thorn Bushveld, Valley Thicket, Subarid Thicket (Low and Rebelo 1996) Succulent False Central Lower Karoo, False Karroid Broken Veld, Bushveld Valley Macchia, Noorsveld, Spekboomveld, (Acocks 1953); Eastern Mixed Nama Karoo, Valley Thicket (Low and Rebelo Thicket, Xeric Succulent 1996) Succulent Karoo, False Macchia, Karroid Broken Veld, (Acocks 1953); Central Mountain Spekboomveld Renosterveld, Central Nama Karoo, Great Karoo, Little Succulent Lowland Fynbos, South and South-west Coast Karoo, Mountain cantha oleoides Euphorbia Schotia afra Aloe speciosa Portulacaria afra Portulacaria afra, Portulacaria afra, Portulacaria afra, Portulacaria afra, Portulacaria afra, Portulacaria afra, Pappea capensis Pappea capensis Pappea capensis, Crassula rupestris Dominant species Gymnosporia polya- coerulescens, Boscia e spp. , cantha lowiczii Euphorbia trispinosus Schotia afra coerulescens Encephalartos Aloe speciosa, Haworthia Cadaba aphylla, Portulacaria afra Portulacaria afra, Euphorbia ledienii, Ozoroa mucronata Gloveria integrifolia, Gymnosporia polya- Rhigozum obovatum Euphorbia tetragona, Rhigozum obovatum, Gymnosporia szyszy- 860 300 150 950 540 720 320 950 (10–670) (110–640) (130–910) (290–870) (10–1740) (240–1390) (770–1560) (230–1310) (min-max)(m) d H H H H H M M M Summer Modal altitude Character species ) -1 c 336 335 367 396 406 414 259 223 (mmyr (min-max) rainfall (174–593) (216–645) (166–534) (183–566) (327–520) (222–673) (175–483) (102–436) b (%) Geology MAR KT (37) KT (22) FS (27) FS (18) FS (43) FS (78) KS (90) KS (28) KS (83) KS (85) VS (22) KS (19) CE (20) FM (35) CM (41) a 1.36 0.55 2.21 0.76 0.49 6.72 2.88 1.04 64 191 25 954 35 712 23 399 49 023 104 704 317 558 136 359 Type Area (ha) % 33.Baviaans Spekboom Thicket 34.Paardepoort Spekboom Thicket 35.Sundays Spekboom Thicket 36.Escarpment Spekboom Thicket 37.Albany Spekboom Thicket 38.Fish Spekboom Thicket 39.Sundays Noorsveld 40.Gamka Arid Spekboomveld Groot Groot Sundays Escarpment Albany Fish Arid solid Sundays Gamka Appendix 1 cont. Region spekboom with solid Arid South African Journal of Botany 2003, 69: 27–51 45 f Synonomy Renosterveld, Spekboom Succulent Thicket (Low and Succulent Renosterveld, Spekboom Rebelo 1996) Central Lower Karoo, False Macchia, Karroid Broken Renosterbosveld, Noorsveld, Succulent Mountain Veld, (Acocks 1953); Central Nama Karoo, Spekboomveld Karoo, Grassy Fynbos, Great Nama Mountain Fynbos, South and South-west Coast Renosterveld, Thicket, Xeric Thicket, Valley Succulent Spekboom Thicket (Low and Rebelo 1996) Succulent Veld Central Lower Karoo, Danthonia Mountain replaced by Karoo, False Central Lower Karroid Karroid Broken Veld, Karroid Broken Veld, Succulent Karoo, Veld, Danthonia Mountain (Acocks 1953); Central Nama Karoo, Spekboomveld Eastern Mixed Nama Karoo, South-eastern Mountain Thicket, Valley Succulent Grassland, Spekboom Thicket (Low and Rebelo 1996) Thornveld, False Macchia, Karroid Eastern Province Valley Succulent Karoo, Spekboomveld, Broken Veld, Bushveld (Acocks 1953); Central Nama Karoo, Grassy Thicket (Low and Succulent Fynbos, Spekboom Rebelo 1996) Central Lower Karoo, False False Macchia, Karroid Broken Karroid Broken Veld, Noorsveld, Succulent Karoo, Spekboomveld, Veld, Bushveld (Acocks 1953); Central Nama Karoo, Valley Eastern Mixed Nama Karoo, Grassy Fynbos, Thicket, Xeric Thicket, Valley Succulent Spekboom Thicket (Low and Rebelo 1996) Succulent Forest, Afromontane Bushveld (Acocks 1953); Valley Thicket (Low and Thicket, Xeric Succulent Valley Rebelo 1996) Bushveld Thornveld of Eastern Province, Valley False Thorn Forest, Eastern Afromontane (Acocks 1953); Thicket (Low Thicket, Xeric Succulent Bushveld, Valley and Rebelo 1996) (Acocks Renosterbosveld, Spekboom-veld Mountain 1953); South and South-west Coast Renosterveld (Low and Rebelo 1996) Alexandria Forest, False Macchia (Acocks 1953); Grassy Fynbos, South and South-west Coast Renosterveld (Low and Rebelo 1996); Humansdorp Grassy Fynbos (Cowling and Heijnis 2001) Bushveld (Acocks 1953); False Macchia, Valley Thicket (Low and Grassy Fynbos, Mesic Succulent Rebelo 1996) Alexandria Forest, False Macchia (Acocks 1953); South and South-west Coast Renosterveld, Valley Thicket (Low and Rebelo 1996) Bushveld Valley False Macchia, Spekboomveld, Forest, Grassy Fynbos, Afromontane (Acocks 1953); Fynbos, South and Thicket, Mountain Mesic Succulent subsp. subsp. tis africana, Pentzia incana Pentzia incana Portulacaria afra Portulacaria afra Euclea undulata, Euclea undulata, Digitaria eriantha Digitaria Portulacaria afra, Portulacaria afra, Portulacaria afra, Pappea capensis, Euphorbia bothae, Rhus chirendensis datus, Rhus lucida Ekebergia capensis Dominant species Afrocarpus falcatus, Pterocelstrus tricuspi- Olea europaea Olea europaea Olea europaea Olea europaea Enneapogon desvauxii Calodendrum capense, africana, Scutia myrtina Elytropappus rhinocero- Elytropappus e salignum Euphorbia Euphorbia coerulescens coerulescens, Leucadendron Crassula ovata Restio triticeus, Boscia oleoides Euphorbia ferox Strelitzia reginae Strelitzia Cussonia spicata Cussonia spicata Cussonia spicata Calpurnea intrusa, Euphorbia bothae, Euphorbia bothae, Sterculia alexandri Sterculia Selago geniculata, Euphorbia atrispina, Rhigozum obovatum Rhigozum obovatum, Rhoicissus tomentosa Euphorbia triangularis, Euphorbia triangularis, Euphorbia triangularis, 90 670 960 500 530 450 340 690 220 380 380 (40–600) (50–500) (40–190) (90–770) (310–810) (250–520) (550–840) (140–590) (430–1460) (570–1560) (170–1060) (min-max)(m) d L L L L H H H H H M M Summer Modal altitude Character species ) -1 c 247 320 230 261 523 420 334 666 585 735 534 (mmyr (min-max) rainfall (147–548) (167–531) (172–360) (153–474) (462–548) (350–514) (263–400) (580–810) (519–716) (692–775) (366–769) b (%) Geology MAR KS (72) KD (20) KS (37) KT (13) KT (63) FS (70) FS (33) KS (61) KS (88) VS (68) FM (11) FQ (33) FQ (31) FQ (42) FM (59) FS (100) VS (100) VS (100) a 2.54 2.80 0.38 6.07 0.03 1.43 0.04 0.24 0.06 0.00 0.30 235 1 460 2 079 3 049 11 571 11 17 866 67 403 14 392 120 122 132 154 286 979 Type Area (ha) % 41.Groot Arid Spekboomveld 42.Escarpment Spekboomveld 43.Gamtoos Arid Spekboomveld 44.Sundays Spekboomveld 45.Albany Spekboomveld 46.Fish Noorsveld 47.Cango Renoster Thicket 48.Klipfontein Fynbos Thicket 49.Otterford Forest Thicket 50.Vanstadens Forest Thicket 51.Zuurberg Forest Thicket Groot Escarpment Gamtoos Sundays Albany Fish Thicket mosaic Gamka Gamtoos Gamtoos Gamtoos Sundays MAINLAND MOSAIC TYPES Appendix 1 cont. Region 46 Vlok, Euston-Brown and Cowling f Synonomy South-west Coast Renosterveld, Xeric Succulent Thicket (Low and Rebelo 1996) Bushveld (Acocks 1953); Xeric False Macchia, Valley Thicket (Low and Rebelo 1996) Succulent Thorn Bushveld (Acocks 1953); Eastern Valley Thicket (Low and Rebelo Bushveld, Xeric Succulent 1996) Thornveld, Valley Alexandria Forest, Eastern Province Forest, Eastern Bushveld (Acocks 1953); Coastal Thicket, Xeric Thorn Bushveld, Mesic Succulent Thicket (Low and Rebelo 1996) Succulent Thornveld (Acocks Alexandria Forest, Eastern Province Thorn Bushveld, Mesic Forest, Eastern 1953); Coastal Thicket (Low and Rebelo 1996) Succulent Thornveld, Valley Alexandria Forest, Eastern Province Forest, Eastern Bushveld (Acocks 1953); Coastal Thicket, Valley Thorn Bushveld, Mesic Succulent Thicket (Low and Rebelo 1996) Thornveld, False Alexandria Forest, Eastern Province Bushveld False Macchia, Valley Karroid Broken Veld, Forest, Forest, Coastal Afromontane (Acocks 1953); Grassland, Eastern Mixed Nama Karoo, Coastal Thorn Bushveld, Grassy Fynbos, Valley Eastern Thicket (Low and Rebelo 1996) Thicket, Xeric Succulent Bushveld (Acocks Thornveld, Valley Eastern Province Thorn Bushveld, Valley Forest, Eastern 1953); Coastal Thicket (Low and Rebelo 1996) Thornveld, False Alexandria Forest, Eastern Province Thornveld of Eastern False Karroid Broken Veld, Bushveld (Acocks 1953); Province, Valley Thorn Forest, Eastern Forest, Coastal Afromontane Thicket (Low Thicket, Xeric Succulent Bushveld, Valley and Rebelo 1996) Thornveld, Eastern Province Forest and Coastal Bushveld (Acocks 1953); Thornveld, Valley Forest, Coastal Forest, Coastal Afromontane Thicket Thorn Bushveld, Valley Grassland, Eastern (Low and Rebelo 1996) Thornveld, Eastern Province Forest and Coastal Thornveld of Eastern Province, Thornveld, False Forest, Afromontane Bushveld (Acocks 1953); Valley Thorn Grassland, Eastern Forest, Coastal Coastal Thicket (Low and Rebelo 1996) Bushveld, Valley Thornveld, Eastern Province Forest and Coastal Thornveld of Eastern Province, Thornveld, False Highland Sourveld and Dohne Sourveld, Valley Forest, Coastal Afromontane Bushveld (Acocks 1953); Thicket, Eastern Grassland, Dune Forest, Coastal Thorn Bushveld, Moist Upland Grassland, Valley Thicket (Low and Rebelo 1996) Bushveld Thornveld, Valley Forest and Coastal Thicket, Eastern Forest, Dune (Acocks 1953); Coastal Thicket (Low and Rebelo 1996) Thorn Bushveld, Valley subsp. sis um datus africana Rhus pallens, Acacia karroo Scutia myrtina Olea woodiana, Erythrina caffra, Loxostylis alata, europaea europaea Hyparrhenia hirta Hyparrhenia Digitaria eriantha, Digitaria Canthium inerme, Themeda triandra Dominant species Themeda triandra, Elaeodendron cro- Sideroxylon inerme Pteroxylon obliquum Phyllanthus verruco- Pittosporum viridifoli- ceum, Erythrina caffra Ptaeroxylon obliquum, Ptaeroxylon sus, Ekebergia capen- Harpephyllum caffrum, Diospyros dichrophylla Pterocelastrus tricuspi- Digitaria eriantha, Olea Digitaria e spp. , spp. subsp. briata gularis africana obliquum altensteinii cuneiforme Faucaria Acacia karoo, Encephalartos Erica simulans, Restio triticeus, Leucospermum europaea europaea laris, Ptaeroxylon Umtiza listeriana, Buxus macowanii Cussonia spicata, Themeda triandra Schotia afra, Olea Erica cerinthoides, Euphorbia triangu- Euphorbia triangu- Brachystelma Hypodiscus striatus Cliffortia linearifolia, Brachylaena ilicifolia Phyllanthus verruco- sus, Euphorbia trian- laris, Sporobolus fim- Brunsvigia grandiflora 350 130 320 340 320 270 280 250 370 210 510 130 (0–200) (90–200) (40–730) (90–680) (10–580) (110–440) (180–660) (260–410) (190–410) (230–410) (130–350) (10–1070) (min-max)(m) d H H H H H M M M M M M M Summer Modal altitude Character species ) -1 c 305 382 494 599 607 589 538 519 586 612 708 833 (mmyr (min-max) rainfall (299–310) (382–382) (426–625) (545–662) (530–656) (462–733) (490–602) (430–633) (529–629) (488–790) (792–917) (529–1054) b (%) Geology MAR FS (11) FS (12) FS (73) FS (76) FS (89) KD (11) PS (96) PS (98) PS (83) PS (12) PS (24) KS (87) KS (86) KS (51) KD (48) KM (10) KS (100) KS (100) CM (100) a 0.01 0.02 0.47 0.09 0.06 2.68 0.04 0.57 0.09 1.08 5.32 0.07 588 1 174 4 360 3 015 2 013 4 264 3 161 22 349 26 879 50 843 126 909 251 460 Type Area (ha) % 52.Zuurberg Fynbos Thicket 53.Kremlin Grassland Thicket 54.Paterson Savanna Thicket 55.Thorndale Forest Thicket 56.Geluk Grassland Thicket 57.Grahams- town Grassland Thicket 58.Narraway Fynbos Thicket 59.Crossroads Grassland Thicket 60.Mountvale Grassland Thicket 61.Mountcoke Grassland Thicket 62.Berlin Savanna Thicket 63.Umtiza Forest Thicket Appendix 1 cont. Region Sundays Sundays Albany Albany Albany Albany Albany Fish Fish Buffels Buffels Buffels South African Journal of Botany 2003, 69: 27–51 47 f Synonomy Coastal Forest and Thornveld, Eastern Province Forest and Coastal Thornveld, Highland Sourveld and Dohne Sourveld, Forest, Afromontane Bushveld (Acocks 1953); Valley Thorn Bushveld, Moist Upland Grassland, Eastern Thicket (Low and Rebelo 1996) Valley Renosterbosveld, False Macchia, Coastal Coastal Afromontane Bushveld (Acocks 1953); Macchia, Valley Thicket, Limestone Fynbos, Mountain Forest, Dune Fynbos, South and South-west Coast Renosterveld (Low and Rebelo 1996) Mountain False Macchia, Karroid Broken Veld, (Acocks 1953); Little Renosterbosveld, Spekboomveld Fynbos, South and South- Succulent Karoo, Mountain Succulent west Coast Renosterveld, Spekboom Thicket (Low and Rebelo 1996) Mountain False Macchia, Karroid Broken Veld, (Acocks 1953); Renosterbosveld, Spekboom-veld Fynbos, South and Little Succulent Karoo, Mountain Succulent South-west Coast Renosterveld, Spekboom Thicket (Low and Rebelo 1996) Succulent Karoo, False Macchia, Karroid Broken Veld, (Acocks 1953); Central Nama Karoo, Spekboomveld Great Nama Karoo, Little Succulent Mountain Fynbos, South and South-west Coast Renosterveld, Thicket (Low and Rebelo 1996) Succulent Spekboom Mountain False Macchia, Karroid Broken Veld, (Acocks 1953); Renosterbosveld, Spekboom-veld Thicket (Low and Rebelo 1996) Grassy Fynbos, Valley (Acocks 1953); Grassy False Macchia, Spekboomveld Fynbos (Low and Rebelo 1996) Spekboomveld False Macchia, Karroid Broken Veld, Thicket (Low (Acocks 1953); Grassy Fynbos, Valley and Rebelo 1996) (Acocks 1953); Grassy False Macchia, Spekboomveld Thicket (Low and Rebelo 1996) Fynbos, Valley Bushveld Alexandria Forest, False Macchia, Valley (Acocks 1953); Grassy Fynbos, Mesic Succulent Thicket, South and South-west Coast Renosterveld (Low and Rebelo 1996) Spekboomveld, False Macchia, Karroid Broken Veld, Bushveld (Acocks 1953); Central Nama Karoo, Valley Thicket, Xeric Succulent Grassy Fynbos, Spekboom Thicket (Low and Rebelo 1996) Succulent Bushveld (Acocks 1953); False Macchia, Valley Thicket, South and Grassy Fynbos, Mesic Succulent Thicket (Low South-west Coast Renosterveld, Valley and Rebelo 1996) Bushveld Valley False Macchia, Spekboomveld, (Acocks 1953); Grassy Fynbos, Mesic Succulent Thicket, South and South-west Coast Renosterveld, Thicket (Low and Rebelo 1996) Valley Bushveld Valley False Macchia, Spekboomveld, (Acocks 1953); Grassy Fynbos, Mesic Succulent Thicket, South and South-west Coast Renosterveld, Thicket (Low and Rebelo 1996) Valley africana ta alata ferox pidatus falcatus africana undulata Aloe ferox Aloe ferox, rhinocerotis Elytropppus Erica demissa Protea repens Celtis africana Olea europaea subsp. Pteronia incana Euclea undulata, Themeda triandra, Themeda triandra, pidatus, Loxostylis Dominant species Bechium burchellia- Afrocarpus falcatus, Hypodiscus striatus, Hypodiscus striatus, Schotia afra, Euclea Ficus sur, Afrocarpus Ficus sur, Pterocelastrus tricus- Pterocelastrus tricus- Pteronia incana, Aloe Pteronia incana, Pteronia incana, Aloe Pteronia incana, Passerina obtusifolia, num, Cussonia spica- e lia latum coides incana incana tenuior fasciata longifolius Rhus lucida, Widdringtonia Felicia filifolia, Acacia karroo, Rhus undulata Smellophyllum Erica simulans Pentzia incana Eriocephalus eri- Turraea pulchella Turraea Cussonia spicata, Aloe humilis, Aloe Aloe humilis, Aloe microstigma, Haworthia cooperi sis, Encephalartos Crassula rupestris, schwarzii, Pteronia Euphorbia atrispina Haworthia longiana, capense, Haworthia Aspalathus kougaen- Aspalathus Pelargonium denticu- Agathosma recurvifo- Passerina obtusifolia, Passerina obtusifolia, Acacia karoo, Pentzia 90 10 110 470 560 140 610 540 710 440 540 950 560 510 (0–270) (0–430) (50–700) (10–310) (50–240) (110–980) (340–880) (340–950) (170–990) (270–790) (250–940) (380–1160) (420–1310) (260–1010) (min-max)(m) d L L L L L L L L L L L H H M Summer Modal altitude Character species ) -1 c 735 432 359 336 252 324 484 364 424 594 325 427 407 525 (mmyr (min-max) rainfall (548–957) (207–638) (203–538) (179–545) (133–530) (266–412) (388–641) (206–564) (300–536) (537–625) (184–489) (284–551) (318–516) (355–674) b (%) Geology MAR Q (66) AL (24) FS (11) KT (18) FS (49) FS (28) FS (75) FS (21) FS (43) VS (15) KS (60) VS (78) FQ (38) FQ (14) FQ (49) FQ (43) CE (50) CE (32) CE (29) FQ (20) CE (25) FQ (17) CE (73) KD (34) CE (24) FM (21) FM (38) FM (43) FM (58) FM (85) FM (49) CM (30) a 1.22 2.49 0.38 0.79 1.66 0.09 0.35 0.24 0.12 0.07 0.82 0.15 0.18 0.19 4 355 5 592 3 351 6 931 8 549 8 831 11 527 11 57 679 17 905 37 435 78 549 16 480 3 8810 117 496 117 Type Area (ha) % 64.Butterworth Savanna Thicket 65.Herbertsdale Renoster Thicket 66.Meirings- poort Fynbos Thicket 67.Mons Ruber Fynbos Thicket 68.Barandas Gwarrieveld 69.Baviaans Doringveld 70.Kouga Fynbos Thicket 71.Baviaans Renoster Thicket 72.Baviaans Fynbos Thicket 73.Bethelsdorp Bontveld 74.Kleinpoort Karroid Thicket 75.Gamtoos Bontveld 76.Gamtoos Doringveld 77.Andrieskraal Fynbos Thicket Appendix 1 cont. Region Kei mosaic Valley Gouritz Gamka Gamka Gamka Groot Groot Groot Groot Groot Groot Gamtoos Gamtoos Gamtoos 48 Vlok, Euston-Brown and Cowling f Synonomy False Macchia, Valley Bushveld (Acocks 1953); False Macchia, Valley Grassy Fynbos, South and South-west Coast Thicket (Low and Rebelo 1996) Renosterveld, Valley Bushveld Alexandria Forest, False Macchia, Valley Thicket, Grassy Fynbos, South (Acocks 1953); Dune Thicket and South-west Coast Renosterveld, Valley (Low and Rebelo 1996) Bushveld (Acocks 1953); False Macchia, Valley Thicket, Mountain Grassy Fynbos, Mesic Succulent Fynbos, South and South-west Coast Renosterveld, Thicket (Low and Rebelo 1996) Valley Thicket, South and False Macchia (Acocks 1953); Dune South-west Coast Renosterveld (Low and Rebelo 1996) Bushveld (Acocks 1953); Grassy False Macchia, Valley Thicket, South and South- Fynbos, Mesic Succulent west Coast Renosterveld (Low and Rebelo 1996) Thornveld, False Macchia, Karroid Eastern Province Bushveld (Acocks Noorsveld, Valley Broken Veld, Thorn Bushveld, Mesic Succulent 1953); Eastern Thicket (Low and Rebelo 1996) Thicket, Xeric Succulent Bushveld (Acocks 1953); Grassy False Macchia, Valley Thicket, South and South- Fynbos, Mesic Succulent west Coast Renosterveld (Low and Rebelo 1996) Thornveld, False Alexandria Forest, Eastern Province Bushveld (Acocks 1953); Mesic Macchia, Valley Thicket, South and South-west Coast Succulent Renosterveld (Low and Rebelo 1996) Bushveld (Acocks Thornveld, Valley Eastern Province Thicket (Low and Rebelo 1996) 1953); Xeric Succulent Bushveld (Acocks 1953); False Macchia, Valley Forest, Grassy Fynbos, Mesic Succulent Afromontane Thicket (Low and Rebelo 1996) Bushveld (Acocks Thornveld, Valley Eastern Province Thorn Bushveld, Mesic Succulent 1953); Eastern Thicket (Low and Rebelo 1996) Thicket, Xeric Succulent Thornveld, Valley Alexandria Forest, Eastern Province Forest, Mesic Bushveld (Acocks 1953); Coastal Thicket (Low and Rebelo 1996) Succulent Thornveld, Valley Alexandria Forest, Eastern Province Thorn Bushveld, Bushveld (Acocks 1953); Eastern Thicket (Low and Rebelo 1996) Mesic Succulent Bushveld (Acocks 1953); False Macchia, Valley Forest, Eastern Mixed Nama Karoo, Afromontane Thorn Bushveld, Grassy Fynbos, Xeric Eastern Thicket (Low and Rebelo 1996) Succulent Thornveld, False Alexandria Forest, Eastern Province Afromontane Bushveld (Acocks 1953); Macchia, Valley Thorn Bushveld, Grassy Forest, Eastern Forest, Coastal Thicket, Xeric Thicket, Valley Fynbos, Mesic Succulent Thicket (Low and Rebelo 1996) Succulent False Macchia, Noorsveld, False Karroid Broken Veld, Bushveld (Acocks 1953); Valley Spekboomveld, Forest, Eastern Mixed Nama Karoo, Afromontane Thorn Bushveld, Grassy Fynbos, Xeric Eastern Thicket (Low and Rebelo 1996) Succulent subsp. spp. ferox ferox datus incana triandra africana africana speciosa Aloe ferox tetracantha Felicia filifolia, Scutia myrtina, tis, Aloe africana europaea europaea Digitaria eriantha, Digitaria Themeda triandra Dominant species Aloe ferox, Pteronia Sideroxylon inerme, Sideroxylon inerme, Aloe africana, Azima Aloe africana, Passerina obtusifolia Pteronia incana, Aloe Pteronia incana, Dodonea angustifolia, Acacia karroo, Lycium Euclea undulata, Olea Euclea undulata, Gymnosporia buxifolia Elytropppus rhinocero- Cussonia spicata, Aloe Cussonia spicata, Pterocelastrus tricuspi- Pittosporum viridifolium Diospyros dichrophylla, Erica sparsa, Themeda Erica sparsa, Themeda triandra, Aloe Themeda triandra, Aloe Themeda triandra, e spp. , spp. tum ledienii africanus tetracantha Protea Drimia elata rhinocerotis, rhinocerotis, Elytropappus Elytropappus Aloe africana, Felicia filifolia, Panicum Protea lorifolia, Ficinia truncata, Pteronia incana, Euphorbia melo- Aspalathus nivea Aspalathus Gasteria pulchra, Bechium burchel- formis, Euphorbia Aspalathus nivea, Aspalathus Carissa bispinosa Euphorbia ledienii Haworthia sordida Haworthia cooperi Euryops ericifolius Euryops Euryops algoensis Euryops Euphorbia triangu- Helichrysum refrac- Euphorbia polygona Acacia karoo, Azima Acacia karoo, Passerina obtusifolia lianum, Eriocephalus Acmadenia obtusata, laris, Gasteria bicolor Agathosma puberula, 90 50 90 60 90 110 230 150 280 220 270 130 310 520 260 470 (0–470) (0–210) (90–550) (20–470) (20–310) (10–100) (30–230) (10–410) (80–300) (90–410) (110–580) (140–380) (130–270) (230–350) (290–590) (260–810) (min-max)(m) d L L L L L L L L L M M M M M M M Summer Modal altitude Character species ) -1 c 508 604 552 617 563 335 613 460 398 619 452 565 471 488 513 409 (mmyr (min-max) rainfall (413–581) (507–676) (364–648) (531–671) (529–601) (164–493) (591–654) (350–596) (370–432) (468–783) (364–583) (484–654) (421–551) (426–516) (415–604) (283–614) b (%) Geology MAR TL (20) TL (58) AL (48) AL (20) AL (16) KT (18) FS (24) FS (91) FS (97) FS (78) FS (46) FS (16) FS (44) PS (12) PS (93) PS (86) PS (17) KS (18) FM (11) FQ (53) CE (71) FQ (12) CE (61) CE (99) CE (15) FM (15) FM (22) FM (24) CM (26) CM (51) CM (19) CM (14) CM (82) FS (100) a 0.06 0.14 0.16 0.04 0.05 0.84 0.06 0.35 0.10 0.06 0.52 0.24 0.08 0.05 0.69 1.93 6766 2 644 7 342 1 823 2 134 2 774 4 812 2 871 3 623 2 382 11 359 11 39 602 16 448 24 615 32 397 91 196 Type Area (ha) % 78.Patensie Fynbos Thicket 79.Kabeljous Renoster Thicket 80.Loerie Fynbos Thicket 81.Kromme Forest Thicket 82.Elands Forest Thicket 83.Sundays Doringveld 84.Rocklands Renoster Thicket 85.Motherwell Karroid Thicket 86.Koedoeskloof Karroid Thicket 87.Groendal Fynbos Thicket 88.Grass Ridge Bontveld 89.Nanaga Savanna Thicket 90.Shamwari Grassland Thicket 91.Alicedale Fynbos Thicket 92.Salem Karroid Thicket 93.Saltaire Karroid Thicket Appendix 1 cont. Region Gamtoos Gamtoos Gamtoos Gamtoos Sundays Sundays Sundays Sundays Sundays Sundays Sundays Albany Albany Albany Albany Albany South African Journal of Botany 2003, 69: 27–51 49 , Valley f Acacia karroo Synonomy Bushveld (Acocks 1953); Afromontane Forest, Eastern Afromontane Bushveld (Acocks 1953); Thorn Bushveld, Moist Upland Grassland, Subarid Thicket (Low and Rebelo 1996) Thorn Bushveld, Valley Mountain False Macchia, Karroid Broken Veld, Bushveld Valley Renosterbosveld, Spekboom-veld, Renosterveld, Little (Acocks 1953); Central Mountain Fynbos, Spekboom Succulent Karoo, Mountain Thicket (Low and Rebelo 1996) Succulent Succulent Karoo, False Macchia, Karroid Broken Veld, (Acocks 1953); Central Mountain Spekboomveld Renosterveld, Little Succulent Karoo, Mountain Fynbos, South and South-west Coast Renosterveld, Thicket (Low and Rebelo 1996) Succulent Spekboom Succulent Karoo, False Macchia, Karroid Broken Veld, (Acocks 1953); Central Mountain Spekboomveld Renosterveld, Central Nama Karoo, Little Succulent Fynbos, Karoo, Lowland Succulent Mountain South and South-west Coast Renosterveld, Spekboom Thicket (Low and Rebelo 1996) Succulent Mountain False Macchia, Karroid Broken Veld, Renosterbosveld, Succulent Karoo, Spekboomveld Renosterveld, Little (Acocks 1953); Central Mountain Fynbos, South and South- Succulent Karoo, Mountain False Karroid Broken Veld, False Macchia, Valley False Karroid Broken Veld, Bushveld (Acocks 1953); Eastern Mixed Nama Karoo, Thorn Bushveld, Grassy Fynbos, Valley Eastern Thicket (Low and Rebelo 1996) Eastern Province Grassveld, False Central Lower Thornveld of False Karoo, False Karroid Broken Veld, Eastern Province, Highland Sourveld and Dohne Valley Veld, Sourveld, Karroid Danthonia Mountain Forest, Eastern Afromontane Bushveld (Acocks 1953); Thorn Bushveld, South- Mixed Nama Karoo, Eastern Thorn Bushveld, Grassland, Subarid eastern Mountain Thicket (Low and Thicket, Xeric Succulent Valley Rebelo 1996) Eastern Province Grassveld, Thornveld, False Central Lower Karoo, Karroid Thornveld of Eastern Province, False Broken Veld, Highland Sourveld and Dohne Sourveld, Karroid Bushveld (Acocks Valley Veld, Danthonia Mountain Forest, Eastern Mixed Nama Afromontane 1953); Thorn Bushveld, Moist Upland Karoo, Eastern Thicket, Thorn Bushveld, Valley Grassland, Subarid Thicket (Low and Rebelo 1996) Xeric Succulent Thornveld, Forest and Alexandria Forest, Coastal Thornveld of Thornveld, False Eastern Province Bushveld (Acocks 1953); Eastern Province, Valley Forest, Coastal Forest, Coastal Afromontane Thicket, Thorn Bushveld, Valley Grassland, Eastern Thicket (Low and Rebelo 1996) Xeric Succulent Thornveld, Eastern Province Forest and Coastal Thornveld, Highland Sourveld and Dohne Sourveld, Invasion of Grassveld by incana Schotia afra Acacia karoo, lianum, Pentzia Pteronia incana Pteronia incana Pteronia incana Pteronia incana Portulacaria afra Euclea undulata, Euclea undulata, Agthosma ovata, Portulacaria afra, Portulacaria afra, Portulacaria afra, Aloe microstigma Bechium burchel- Themeda triandra Dominant species Euphorbia tetragona, Euphorbia tetragona, e tum lianum lowiczii guiforme Scotia afra Aloe tenuior rhinocerotis, Elytropappus Acacia karroo Aloe africana, Glottiphyllum lin- Euclea undulata, Laurida reticulata Bechium burchel- Pappea capensis, Glovria integrifolia, Haworthia truncata Diospyros scabrida, Gloveria integrifolia, Tylecodon panicula- Tylecodon Euphorbia tetragona, Euphorbia curvirama, Gymnosporia szyszy- 640 650 480 210 630 420 360 360 500 (0–630) (70–790) (540–680) (360–970) (70–1260) (310–830) (310–1390) (200–1000) (190–1240) (min-max)(m) d L L L L H H H H M Summer Modal altitude Character species ) -1 c 605 401 496 518 616 264 248 254 336 (mmyr (min-max) rainfall (112–494) (116–392) (558–631) (312–556) (337–745) (394–659) (423–892) (156–452) (212–528) b (%) Geology MAR AL (41) KT (45) FS (30) TS (18) FS (75) FS (72) FS (89) KS (97) KS (97) KS (83) KS (50) FM (11) KD (22) CE (57) FM (13) FM (15) KM (27) a 0.07 3.54 3.36 2.83 2.73 1.80 0.83 0.83 1.22 3 082 85 217 39 081 39 203 57 902 167 554 158 906 133 694 128 878 Type Area (ha) % 94.Albany Bontveld 95.Hartebeeste Karroid Thicket 96.Bedford Savanna Thicket 97.Doubledrift Karroid Thicket 98.Bolo Savanna Thicket 99.Vanwyksdorp Gwarrieveld 100.Calitzdorp Karroid Thicket 101.Kandelaars Karroid Thicket 102.Blossoms Karroid Thicket Albany Fish Fish Fish Kei Arid mosaic Gouritz Gamka Gamka Gamka Appendix 1 cont. Region 50 Vlok, Euston-Brown and Cowling f Synonomy west Coast Renosterveld, Spekboom Succulent west Coast Renosterveld, Spekboom Thicket (Low and Rebelo 1996) Succulent Karoo, Spekboomveld Karroid Broken Veld, (Acocks 1953); Little Succulent Karoo, Spekboom Thicket (Low and Rebelo 1996) Succulent Mountain False Macchia, Karroid Broken Veld, Renosterbosveld, Succulent Karoo, Spekboomveld Renosterveld, Little (Acocks 1953); Central Mountain Fynbos, South and South- Succulent Karoo, Mountain Succulent west Coast Renosterveld, Spekboom Thicket (Low and Rebelo 1996) Succulent Karoo, False Macchia, Karroid Broken Veld, (Acocks 1953); Central Nama Karoo, Spekboomveld Little Succulent Karoo, South and South-west Coast Thicket (Low and Succulent Renosterveld, Spekboom Rebelo 1996) Mountain False Macchia, Karroid Broken Veld, Renosterbosveld, Succulent Karoo, Spekboomveld (Acocks 1953); Central Nama Karoo, Great Fynbos, South Karoo, Little Succulent Mountain and South-west Coast Renosterveld, Spekboom Thicket (Low and Rebelo 1996) Succulent Succulent Karoo, Spekboomveld Karroid Broken Veld, (Acocks 1953); Central Nama Karoo, Great Thicket (Low and Rebelo Succulent Karoo, Spekboom 1996) Central Lower Karoo, False Macchia, Karroid Broken Valley Succulent Karoo, Spekboomveld, Veld, Bushveld (Acocks 1953); Central Nama Karoo, Grassy Thicket, Thicket, Valley Succulent Fynbos, Spekboom Thicket (Low and Rebelo 1996) Xeric Succulent Thornveld, False Macchia, Karroid Eastern Province Valley Succulent Karoo, Spekboomveld, Broken Veld, Bushveld (Acocks 1953); Central Nama Karoo, Grassy Fynbos, South and South-west Coast Renosterveld, Thicket Thicket, Xeric Succulent Succulent Spekboom (Low and Rebelo 1996) False Central Lower Karoo, Karroid Broken Veld, Noorsveld (Acocks 1953); Eastern Mixed Nama Karoo, Thicket (Low and Rebelo 1996) Xeric Succulent Central Lower Karoo, False Noorsveld, Spekboomveld Karroid Broken Veld, (Acocks 1953); Central Nama Karoo, Spekboom Thicket, Xeric Succulent Succulent Thicket, Valley Thicket (Low and Rebelo 1996) Bushveld (Acocks 1953); Grassy Fynbos, Xeric Valley Thicket (Low and Rebelo 1996) Succulent incana rhinocerotis Elytropappus Acacia karroo Pentzia incana Pentzia incana, Pentzia incana, Pentzia incana, Eragrostis plana Portulacaria afra Portulacaria afra Euclea undulata, Euclea undulata, Euclea undulata, Euclea undulata, Portulacaria afra, Dominant species Rhigozum obovatum Passerina obtusifolia Digitaria argyrograpta Digitaria Schotia afra, Pteronia e var. spuria var. var. morgsana longispina Euphorbia Euphorbia polyacantha Zygophyllum Elytropappus Acacia karroo, Crassula ovata noorsveldensis, Euphorbia ferox Rhodocoma arida Euphorbia bothae Crassula rupestris, Astroloba foliolosa, coerulescens, Rhus major, Gymnosporia major, orbiculare Euphorbia esculenta, Passerina obtusifolia, Glottiphyllum difforme Haworthia archnoidea Euphorbia horrida rhinocerotis, Cotyledon 320 430 780 890 940 780 440 510 680 540 (260–500) (140–890) (390–970) (280–680) (440–590) (560–1050) (640–1290) (760–1080) (310–1050) (460–1000) (min-max)(m) d L L H H H H H H M M Summer Modal altitude Character species ) -1 c 235 304 258 304 198 242 232 317 267 568 (mmyr (min-max) rainfall (115–459) (167–322) (200–392) (152–508) (136–363) (157–350) (152–353) (273–378) (159–350) (527–599) b (%) Geology MAR AL (86) FS (11) KT (21) KT (69) FS (96) FS (66) FS (10) FS (91) FS (29) KS (78) KS (56) KS (88) KS (94) CE (14) CE (48) FM (23) FM (14) a 0.26 0.76 0.66 1.28 0.25 1.46 2.66 0.56 2.05 0.05 22 95 11 894 11 12 508 35 741 31 072 60 361 68 920 26 513 97 131 12 5540 Type Area (ha) % 103.Oudtshoorn Karroid Thicket 104.De Rust Karroid Thicket 105.Willowmore Gwarrieveld 106.Willowmore Renoster Thicket 107.Beervlei Karroid Thicket 108.Groot Gwarrieveld 109.Gamtoos Gwarrieveld 110.Waterford Doringveld 111.Sundays Gwarrieveld 112.Ecca Bontveld Appendix 1 cont. Region Gamka Gamka Groot Groot Groot Groot Gamtoos Sundays Sundays Albany South African Journal of Botany 2003, 69: 27–51 51

Appendix 1 cont. a Extent of thicket vegetation in STEP planning domain b Underlying geology covering >10% of each type. Simplified codes are (ranked according to age of deposit (Council for Geosciences 2002); RS = Recent marine sand; AL = alluvium and calcrete; PS = Pleistocene marine sands; TL = Tertiary limestone; TS = Tertiary silcrete; CM = Cretaceous mudstone; CE = Cretaceous conglomerate; KD = Karoo dolerite; KM = Karoo mudstone; KS = Karoo shale; KT = Karoo tillite; FS = Cape shales; FM = Cape sandstones/shales/tillite; FQ = Cape quartzitic sandstone; PG = Cape granite; VS = Precambrian sediments (limestone, phyllite etc.) c MAR = mean annual rainfall d Summer = Sept–Mar. L = low (<50% MAR), M = moderate (50–75% MAR), H = high (>75% MAR) e Species that characterise vegetation types within each of the structural units f Synonomy was assessed by digitally comparing overlap between our vegetation map and that of Acocks (1953) and Low and Rebelo (1996) (overlapping types are listed alphabetically in both cases)