sign in sign up
<<
Home , Fynbos, Renosterveld

Phenology of , and subtropical thicket in the south

Shirley M. Pierce and A.M. Cowling Department of Botany, University of ,

Qualitative and quantitative phenological observations were made Introduction on 173 species in eight communities in climatically similar sites. Results indicated that in species growing on different substrates, Phenology is the study of the timing of recurring biological soil type had a minimal effect on phenophases. Phenophase pat­ events. The phenology of plant growth and reproduction is terns were analyzed by grouping species into growth form a major focus of the Fynbos Project (Kruger 1978) and classes. Geophytes and annuals grew from autumn to spring. The is the subject of two reviews (Kruger 1981; Pierce 1983). The majority of restioids and C grasses grew most in the cool wet 3 phenology of plants from the south western of the biome seasons. C grass species showed either a summer growth 4 has been reviewed by Kruger (1981) while Bond (1980) presents season or an additional cooler growth season; the former species do not occur westwards in the winter rainfall region while the lat­ data from the southern region. Earlier studies have been car­ ter do. Most succulents grew in autumn and spring while two ried out in Namaqualand (van Rooyen et a/. 1979) and the species also grew in summer. Small leaved shrubs eastern Cape (Palmer 1982) . This study reports on the grew throughout the year and/or showed a summer growth peak. phenology in the south eastern Cape of three types The former pattern is consistent with a 'generalist strategy' but (fynbos, renosterveld, subtropical thicket) in the biome. the latter is not readily explained because of summer drought The south eastern Cape forms the eastern limit of the Fyn­ conditions. Subtropical large leaved sclerophyll shrubs showed ir­ regular growth and reproduction whereas large leaved proteoid bos Biome and comprises the meeting place of four African shrubs grew in summer and autumn. In all shrub growth forms phytochoria, namely the Cape, -Namib, Tongaland­ maximum leaf loss occurred in summer. Phenophase patterns Pondoland and (Goldblatt 1978; Werger were explained in terms of ecophysiological factors but biological 1978; Gibbs Russell & Robinson 1981; White 1983). The area and historical factors were also considered. is rich in types (cf Acocks 1953; Gibbs Russell & S. Afr. J. Bot. 1984, 3: 1- 16 Robinson 1981) most of which are chorologically complex Kwalitatiewe en kwantitatiewe fenologiese waarnemings is op 173 (Werger 1978; Cowling 1983a). We studied the phenology of plantspesies in agt gemeenskappe uit klimatologies-ooreenstem­ eight plant communities, representing three vegetation classes mende gebiede gemaak. Die resultate dui daarop dat in die geval (sensu Cowling 1983a), in a climatically homogeneous area van spesies wat op verskillende substrate groei, die grondsoort 'n minimale invloed op fenofases het. Fenofasepatrone is ontleed within a 10 km radius. Species were grouped into growth deur spesies in groeivormklasse te groepeer. Geofiete en een­ forms, and phenophases within each group were discussed in jarige plante het vanaf die herfs tot die lente gegroei. Die terms of ecophysiological, biological and historical factors. The meerderheid restio-agtige en C3 grasse het meestal in die koel nat data have been used for periodicity comparisons across the seisoene gegroei. C grasspesies het 6f 'n somergroeiseisoen 6f 4 Fynbos Biome (Pierce 1983) to test the overstorey/ understorey 'n addisionele koeler groeiseisoen getoon. Eersgenoemde spesies phenological model for mediterranean (Pierce & kom nie weswaarts tot in die winterreenvalstreek voor nie terwyl laasgenoemde wei daar voorkom. Die meeste sukkulente groei in Cowling 1983) and as a basis for improving the grazing quality die herfs en lente terwyl twee spesies ook in die somer groei. Die of renosterveld (Cowling et a/. 1983). sklerofiliese struike met klein blare groei dwarsdeur die jaar en/of toon 'n groeipiek in die somer. Eersgenoemde patroon is in oor­ Study Area eenstemming met 'n 'algemene strategie' maar vanwee die droe Site descriptions somertoestande, is laasgenoemde patroon nie maklik verklaarbaar nie. Die subtropiese struike met groot blare se groei en voort­ The study area is located near Humansdorp, south eastern planting was onreelmatig terwyl die -agtige struike met Cape (Figure 1) on a level coastal plain which cuts across two groot blare in die somer en herfs groei. By aile struikagtige groei­ geological formations: sandstone of the Group vorme het maksimum blaarval in die somer voorgekom. Die feno­ (TMG) and shales of the Bokkeveld Group. Along the coast fasepatrone is in terme van ekofisiologiese faktore verklaar maar there are deposits of recent calcareous sands. A detailed biologiese en historiese faktore is ook in aanmerking geneem. description of the environment of the study area is given in S.-Afr. Tydskr. Plantk. 1984, 3: 1- 16 Cowling (1983a). Since the principle vegetation types corres­ Keywords: Fynbos, phenology, renosterveld, south eastern Cape, pond closely to geological substrate, it was possible to study subtropical thicket eight communities which all experience similar mesoclimatic Shirley M. Pierce* and R.M. Cowling conditions. Some details of the sites, including dominant Present address: School of Biology, Western Australian Institute of species, structure and soil data are given in Table I. The follow­ Technology, Kent Street, Bentley, Western 6102 ing vegetation concepts are defined and described in detail by *To whom correspondence should be addressed Cowling (1983a). Grassy Fynbos communities are endemic to Accepted 13 October 1983 the south eastern Cape and occur on infertile, sandy soils. 2 S.-Afr. Tydskr. Plantk., 1984, 3(1)

South Coast Fynbos is distributed along the southern South Coast Renosterveld is a shrubland confmed to the south Cape coast and is confined to calcareous coastal dune sands. coastal foreland of the Cape Region (sensu Goldblatt 1978), and in contrast to Cape Fynbos, occurs on loamy, moderately fertile soils (Taylor 1978; Boucher & Moll1980) which, in the study area, are derived from Bokkeveld shale. Kaffrarian Thicket is a subtropical thicket type which penetrates the Cape Region on deepish, well drained fertile soils in the lowlands. In the study area, thicket occurs on deep, well drained soils derived from Bokkeveld shales as well as on dune sands.

Land use The dune , shale grassland, renosterveld and restioid Cape St Francis Skm grassland sites (Table 1) are moderately grazed and mowed or burnt on a 4- 5 year rotation. The last mentioned site was inadvertently burnt in February 1981, at the start of the sam­ pling. Grazing animals were excluded from the grassland sites Figure 1 Map of the study area in the Humansdorp district showing the during the sampling period. The dune fynbos and grassy fyn­ numbered study sites in each community (see Table 1) . bos sites (Table 1) were ungrazed and unburnt for seven and

Table 1 Vegetation and soil data for the study sites

Site numbera

4 6

Vegetation orderb South Coast Kaffrarian Thicket South Coast Grassy Fynbos Grassy Fynbos Kaffrarian Thicket South Coast South Coast Dune Fynbos Dune Fynbos Renosterveld Renosterveld Communityb Restio eleocharis- Cassine aerhiopica- - Thamnochortus Thamnochortus Pterocelastrus Elytropappus rhino- Themeda triandra Agathosma steno- Cussonia thyrsijlora Stenotaphrum g/aber- fruticosus- Tristachya tricuspida!Us-Euclea cerotis-Me!alasia lineari- petala Dune thicket secundatum diaphana /eucothrix undulata muricata folia Dune fynbos Dune grassland Grassy fynbos Rest ioid grassland Kramme River Renosterveld Shale grassland thicket Dominant species Agathosma steno- Sideroxylon inerme, Themeda triandra, Erica diaphana, E. Themeda triandra, Euclea undulata, Themeda triandra, Themeda triandra. petala, A. apiculata, Pterocelastrus tri- Stenotaphrum secun- pectinifolia, Leuca- Tristachya leu- Maytenus acumi- Elytropappus rhi- Sporobolus africam;, Restio eleocharis, cuspidatus, Cassine datum, Passerina dendron salignum, cothrix, Thamno- notus, Olea africana, nocerotis, Ruschia Cynodon dactylon R. /eptoc/ados, aethiopica, Olea vulgaris, Rhus Thamnochortus chortus fruticosus, Sideroxy/on tene/la, Meta/ Pentaschistis Passerina vulgaris exasperara laevigata glaber, Restio E/egia vaginel/ata inerme muricata angustifolia triticeus Vegetation Low closed ericoid Mid-high closed Closed grassland Mid-high closed Closed rest ioid Tall closed Low open grassy C losed grassland structure" sh rubland with large leaved wilh low sparse ericoid and proteoid grassland with low large leaved small leaved with sparse dwarf open Testioid under- shrubland ericoid shrub over- shrubland with open sparse ericoid shrub shrubland shrubland small-leaved shrul storey srorey restioid oversrorey overstorey overstorey Soil propenies Soil description Excessively drained, Excessively drained, Seasonally water- Excessively drained, Seasonally water- Well drained, fertile Poorly drained, Seasonally water- and depth (m) moderately ferti le, moderately fenile, logged, moderately infert ile, acid, logged, infertile, loam (duplex) moderately logged, (duplex) fine calcareous medium, calcareous fertile, fin e, cal- coarse sand acid, medium sand 0,3 - 1,0 fertil e, loam moderately fertile dune sand dune sand careous dune sand 1, 1 I ,5 0,4 loam > 3 >3 1,5 0,5 Soil form & Fernwood Fernwood Fernwood Constantia Longlands Clovelly Glenrosa Swartland seriesd Motopi Motopi Brinley Strombolis Orkney Williamson Bridbach Geology Recem sand Recent sand Recent sand Table Mountain Table Mountain Bokkeveld shale Bokkeveld shale Bokkeveld shale Group sandstone Group sandstone H,O holding capacity' (g) 27,6 48,6 29,8 40,2 69,3 42,1 40,9 pH 7,7 7,6 7,6 4,2 4,3 5,1 5,3 5,3 Na (ppm) 44 193 130 34 119 196 182 379 K (ppm) 40 63 98 31 34 414 240 129 Ca (ppm) 19 18 2031 3039 186 209 2168 756 712 Mg (ppm) 57 246 309 58 172 773 264 427 S values (m-equiv.)' 10,3 13 ,3 18 ,6 1,65 3,09 19, 16 7,38 9,10 Oxidizable C' ("'o) 2,4 4,5 7,4 3,2 3,5 18,6 5,1 5,0 Total Nh (%) 0,075 0, 102 0,305 0,046 0,089 0,443 0, 124 0,195 Available P; (ppm) 35,7 15,1 15,6 1,5 3,9 25,7 2,0 3,3

'Reference to site marked on map (Figure I) tvegetation orders and communities sensu Cowling (1983a); names below binomial in column 3 are those used in the text 'Structural characterization sensu Campbell eta/. (1981) dSoil classification sensu MacVicar eta/. (1977) 'Weight absorbed at saturation by 100 g soil (Piper 1950)

'Sum of exchangeable cations (I mol dm - 3 NH4 acetate leachate) 'Walkley-Black method hKjeldahl method ;Modified Olsen (alkali extraction, pH 8) for I - 3 Bray No. 2 (acid extraction, pH 3) for 4 -8.

\ S. Afr. J. Bot., 1984, 3(1) 3

fourteen years respectively, according to records kept by the growth forms (Table 2) defined in terms of height of peren­ landowners. The thickets on shale and dune sand were pro­ nating bud (Raunkiaer 1934), photosynthetic mode and leaf bably unburnt for fifty years or longer. size. The 'classical' descriptive method involving the qualitative Climate assessment of phenophase by simple observation has been The climate is warm temperate and transitional between Kop­ widely used in phenology (Williams 1971; Dickinson & Dodd pen Csa and Csb climates (Specht & Moll 1983). Most rain 1976; Heinrich 1976). We have similarly used a descriptive is cyclonic and associated with fronts from the circumpolar method in which elements of time and site are preselected and westerly belt. The study area is usually included in the non­ the phenophase then determined. seasonal rainfall region although the three summer months (December- February) are always the driest (Figure 2b). Bond (1980) has shown that semi-arid and subhumid climates in the Table 2 Growth form classes Cape constant rainfall region have an effective moisture regime Herbaceous growth forms that is unequivocally mediterranean as typified by moisture Annuals surpluses in winter and deficits in summer (Figure 2b). The Geophytes warmest month at Cape St. Francis is February (mean monthly Non-geophytic forbs Graminoids temperature of 19,9 °C) and the coolest is July (14,2 °C). Grasses (Poaceae) - C 3 Highest and lowest mean temperatures are 29,7 oc (in April) - c. and 5,0 oc (in August) respectively (Anon 1942). Frost is un­ Restioids () (Campbell eta/. 1983) common. The area is subject to strong winds and occasional Cyperoids (Cyperaceae) (Campbell et a/. 1983) gales at any time of the year. Evaporation and radiation are Succulents clearly highest in the summer months (Nov- Feb) (Figure 2a). CAM species (Mooney, Troughton & Berry 1977) Woody growth forms As all sites are on level ground, the radiation regime was as­ Small leaved shrubs" (Campbell e1 a/. 1983) sumed to be similar for all vegetation types. During the study Large leaved shrubs" (Campbell el at. 1983) (1981), exceptionally heavy precipitation resulted in doubling Proteoidsb the average annual total and rainfall was unequivocally Thicket speciesc bimodal (Figure 2c). "Most species have sclerophyllous leaves blsobilateral sclerophyll leaves (all species are ) coorsiventral sclerophyllleaves (components of Subtropical Transitional

® ~- -•Radiation Thicket (Cowling 1983a)). _.f:vaporation

250 -"', 25 In phenology, measures of different plant processes can give mm , ' ' different growth peaks (Groves 1965). Because detailed mJm-2 day1 ,' ' measures of processes such as translocation of metabolites and 150 15 ,t' cambial growth were beyond the scope of this study, were­ .. @ corded the most obvious manifestations of growth which in­ cluded shoot elongation, leaf initiation and development. Other 50 5 mm conspicuous phenophases noted were: preflowering (bud) and 200 full flowering (open flower); and unripe and ripe fruiting/ @ 100 seeding. Leaf yellowing, prior to and including abscission, was mm 100 noted. In this way we made qualitative observations at monthly 60 30 oc intervals on a total of 173 species in the eight communities shown in Table 1 during the period January 1981 - April1982. Observations were made on 3-5 individuals of each species. 20 10 In addition, successive monthly measures of shoot length in­ crements on 20 of the selected species were made to show peak JA,SONOJFMAMJ uFMAMJJASONDJFM Winter · Sprinq lsummeriAutumn ( 1981 1982 growth periods. To do this we tagged two shoots and measured Figure 2 Climate of the study area. (a) Mean monthly solar radiation their lengths on each of six specimens of each shrub species. (earth's surface) for 1957- 1958 and mean monthly evaporation (American Shoot elongation on two shoots per ten tufts of each restioid Class 'A' Evaporation Pans) for 1957-1961 at (McCallum species was monitored. Twelve shoots of different grass tufts 1972). (b) Walter-Lieth climate diagram for Cape St. Francis (Heydorn were tagged and the number of tillers and/or leaves per shoot & Tinley 1980). (c) Rainfall at Cape St. Francis during the sampling period were counted, depending on the species' growth habit. To (Weather Bureau, unpublished). measure growth of the arborescent rosette succulent, Aloe africana, we recorded the length of the three newest leaves on Methods six tagged specimens. Leaf fall of selected species was measured At each of the eight sites (see Table 1) the most common species by placing traps (200 mm diameter) beneath each of two were selected for study. The 173 species selected were iden­ specimens. Leaves trapped during each sampling interval were tified by one of us (RMC) and voucher specimens are located air-dried to a constant mass and weighed. At each sampling in the Albany Museum Herbarium. we also recorded all geophytes within a set area of 95 m2 in The results were analysed for phenological patterns. The each community. most meaningful patterns emerged when species were grouped Increments were used in growth analysis of shoots and aloe into growth forms, equivalent to guilds or groups of directly leaves, but actual counts of tillers and/or leaves each month competing individuals (Peet 1978). We grouped species into were used for grasses. 4 S.-Afr. Tydskr. Plantk., 1984, 3(1)

The data were not normally distributed and therefore would Themeda triandra Forsk. be poorly represented by means and standard errors. Instead 0 the growth data are presented in the form of five-number sum­ I 1?\'""d maries (sensu Underhill 1981). The method of measuring shoot length increment is com­ 5 pared with an alternative, widely used method of growth deter­ mination. This latter method involves the use of bar graphs 5 Fire Q Q J, $ of relative abundance (percentage) of tagged specimens of a ~ o.!. o ~ ~ ~ ~ ~ species showing the same phenophases at the time of sampling ® (Frankie et a!. 1974; Guy et a!. 1979). , 35 Results and Discussion 6 25 Phenodiagrams of the common species in each community are c: shown in Appendices 1 - 7. The classification of species into ~ growth forms is shown in Appendix 8. Five-number summaries ~ 15 depicting the detailed growth measurements of selected species 5 and including information on other phenophases are shown LJ ~ 8 cp I in Figures 3 - 11 . ~~ ® ~ 15

5 g $ e ~ JFMAMJJAS~ ~ ~~ 0~ ~NDJFMA ~ ~ ~ 1981 1982

Figure 4 Phenophases of Themeda triandra in (a) restioid grassland, (b) dune grassland and (c) shale grassland. Key: see Figure 3.

Tristachya -leucothrix Nees ~0 II n ~5 .:, .1~o..,.:. J,~,~,+"'j -' Fire T l,J T I 8 T 0 0 .1o Sporobolus africanus(Poir.) Robyns & g Tournay 1 I .J, ~ o E3 ~ '*' ~ $ "' 8$ ~ 1' e 8 'i"

ci c: • JLB Gi 5 I i=o- ®- 6 - 6 ~ 6 ~6 ~ ~ - Q 0

1

~ ~ "' t ~ e '*' ~ $ ~ $ $ 8 8 ~ 0 Stenotaphrum secundatu~ (Walt.) 0. Kuntze ~ ······ 1 ~ ~ ®- do ~ . Q ~ ~ 6~ ~ 6 0 g8 O JFMAMJJAS 0 NDJFM 1981 1982

80 Figure 5 Phenophases of other C4 grasses in (a) restioid grassland, (b) shale grassland and (c) dune grassland. Key: see Figure 3.

Evaluation of methods '"'E .§ 40 In Figure 11 the two methods of determining growth are given: .s: bar graphs denote the percentage of observed individuals grow­ i 2o ing at any one sampling time while detailed increments are ex­ (.!)e pressed as five-number summaries. The bar graphs simply in­ dicate that most of the growth of both species occurs from 0 autumn, through winter to spring. The five-number summaries, however, reveal the bimodal nature of this cooler season growth on the loam soil. Low shoot increment in Pterocelastrus Figure 3 Phenophases of restioids in (a) dune fynbos, (b) grassy fynbos and (c) restioid grassland. Key: growth depicted in five-number summaries tricuspidatus in September (Figure 11 a) and in Sideroxylon - see text; pre-flowering - stripes; full flowering - shaded; unripe seeds inerme in October (Figure lid) coincided with a high percen­ - open stipples; mature seeds - dense stipples. tage of shrubs showing growth (Figures 11 b and e) . This ex- S. Afr. J. Bot., 1984, 3(1) 5

Pentaschistis angustifoJia Nees. Stapf 4 0 .rw 2 0 10 '= I F' ,, ~"''"""" ,,,,,,,,,,,, ,:l ~ ~ ~ ~ ~ l 9~ ¢ + • ? * 6o (L.) Less. 40

20

0 0 1981 1982

Figure 6 Phenophases of C3 grasses in (a) shale grassland and (b) dune grassland. Key: see Figure 3. 2 0

60 Agathosma apiculata G. F. W. Meyerl Aloe africana Mill. 40 40

20 '"'30 E .§ 0 ~ 20 @~ "i 0 ~ c, 4 10 " ~ 2 «; 0 0 ~ ~-F~M~A~M~-J~J~A~S~~O~~N ~D~J~F~M~ 1981 1982 e .5 60 Figure 7 Phenophases of Aloe ajricana in Kromme River thicket. Key: see Figure 3. ~ ie 40 (!)

emplifies the point raised earlier that different measures give 20 different peaks. In this case, however, the results of the two ~ methods are not contradictory; a high proportion of shoots 0 could be growing but their growth, expressed as shoot incre­ ® ment, could be minimal. FMAMJJASONDJFM 1981 1982 Effects of substrate on phenophase Figure 8 Phenophases of small leaved sclerophyll shrubs in dune fynbos. Species which occurred on more than one substrate type are Leaf tall is expressed as the mean of two values (bars). Key: see Figure 3. listed in Table 3 where their phenodiagrams are referred to by figure number. In general, the phenophases of these species were synchronous, suggesting that substrate type does not have the dune sand site (Figure 8) resulted in a 'false peak' in August. a marked effect on phenophases. Heyward (1931) analysed the The tendency for earlier initiation of phenophases in the flowering of 554 indigenous genera occurring in Victoria, loam soils might be explained by the higher water-holding

Australia and found that most species' anthesis differed by capacity of the loam (42,1 g per 100 g H 20) relative to the less than a few weeks in spite of substantial environinental dune sand and acid sand (27 ,6 and 29,8 g per 100 g HzO respec­ variation. In our study, loam soil tended to initiate growth (e.g. tively) (Table 1). After a light rainfall, more moisture would Pteroce/astrus tricuspidatus and Sideroxylon inerme, Figure be held in the loam than in the excessively drained sand. 11) and flowering (Helichrysum teretifolium, Appendices 3 and Themeda triandra showed similar growth rhythms in dune 6) in certain species earlier than the dune sand but showed no sand and loam, though early summer growth on the loam other marked difference. preceded that on the sand by about a month (Figure 4). The Meta/asia muricata showed no growth from autumn to late contrasting pattern of ever increasing growth of Themeda trian­ winter in the dune sand (Figure 8) and acid sand (Appendix 3) dra on the acid sand of the restioid grassland is seen as a con­ though some growh was apparent at the loam site (Figure 10) sequence of post-fire recovery. for the same period. However, maximal growth occurred in Differences in fruiting behaviour associated with different November on all three substrate types. Highest leaf fall in the substrate type were apparent in Sideroxylon inerme (Figure 11). loam was recorded in February following maximal growth No fruits were produced at the loam site while unripe fruits (Figure 10). Inclusion of old flowers in the litter sample from were recorded continuously in the dune sand. Flowering was 6 S.-Afr. Tydskr. Plantk., 1984, 3(1)

ucospermum cuneiforme ® Spreng. (Burm.f.) Rourke

40 20 I 10 j ------~ ~ ~ 0 !$ ~ ~ ~ salignum @ @ '"'Cl Berg. 0 v 1

-m Erica pectinifolia Cl> ..J Salis b.

60

e 3o 40 E v 20 20 =3: 0 ~ Q .p o 0 .. l l B - ~ I~ 0 .~ B ~ d ~ M A M J J A S NDJFMA 1981 1982 1981 1982

Figure 9 Phenophases of proteoids (a) and (b) and small leaved shrubs (c) and (d) in grassy fynbos. Leaf fall is expressed as the mean of two values (bars). Key: see Figure 3.

tropical thicket shrubs is common in mesic, coastal en­ vironments (Liversidge 1972; K.L. Tinley pers. comm.). Phenophases of growth forms Geophytes Numbers of geophyte species recorded at each site within a 95 m2 plot were: five in dune grassland; fifteen in renosterveld; 40 none in grassy fynbos; two in dune fynbos and dune thicket; thirteen in restioid grassland and eight in shale grassland (Ap­

20 pendices 1, 2, 3, 4, 6 and 7). These results support the evidence for a tendency towards a greater diversity of geophytes with increasing soil fertility (Kruger 1979). Geophytes tended to in­ 0 itiate leaf growth from autumn (April/May) through to spring (September/October). Flowering in all communities was mainly in spring with occasional flowering in summer (Monadenia bracteata, Micranthus plantagineus, Appendix 4) and autumn/ winter (Oxalis polyphylla, Appendix 6). Our data in­ dicate that flowering behaviour may be a response to rainfall ytropappus rhinocerotis ~ coinciding with higher temperatures. Maximum growth of e geophytes in spring was also noted in Dark Island , E10 v Australia (Specht & Rayson 1957). In the S.W. Cape, Kruger ..<: 5 i Q $ (1981) reported geophyte leaf initiation in autumn/early winter 0 0 8 ~ (~f.) t:· ~ 9~ ~ ~ but anthesis and leaf death varied considerably. Observations ~ FMAMJJAS 0 NDJFM on leaf initiation in our study indicated similar timing to that 1981 1982 in the S.W. Cape.

Figure 10 Phenophases of small leaved shrubs in renosterveld. Leaf fall Annuals is expressed as the mean of two values (bars). Key: see Figure 3. Almost all annuals observed grew and flowered in early autumn, later winter and spring (Appendices 2, 4, 6 and 7). noted at both sites, though to a lesser extent on the loam. Thus growth of annuals in the study area coincided with an However, irregular flowering and fruiting of individuals of sub- essentially winter growing season for agricultural crops. S. Afr. J. Bot., 1984, 3(1) 7

..,,.,,. .. ,.,,,..elastrus tricuspidatus ideroxylon inerme L. (Lam.) Sond. 0 30

1981 1982 1981 1982

Figure 11 Phenophases and growth (five-number summaries and bar graphs- see text) of large leaved sclerophyll shrubs in Kromme River thicket (a) , (b), (d) and (e); and dune thicket (c) and (f). Leaf fall is expressed as the mean of two values (bars). Key: see Figure 3.

Table 3 List of species occurring on more than one substrate type. Their phenodiagrams are shown in corresponding figures (Fig.) and appendices (App.)

Site numbers"

Species occurring on more 2 3 4 6 7 8 than one substrate type Dune sand Dune sand Dune sand Acid sand Acid sand Loam Loam Loam

Pterocelastrus tricuspidatus (Fig. 11) (Fig. 11) Sideroxylon inerme (Fig. 11) (Fig. II) Themeda triandra (Fig. 4) (Fig. 4) Meta/asia muricata (Fig. 8) {App. 3) (Fig. 10) Helichrysum teretifolium (App. 3) (App. 6) lxia orienta/is (App. 2) {App. 7) Briza maxima (App. 4) (App. 6) Babiana patersoniae (App. 4) (App. 7) Sporobolus ajricanus (App. 2) (App. 4) (App. 7) Tephrosia capensis {App. 2) (App. 7) Oxalis polyphylla (App. 4) (App. 6)

"See Table I for detailed soil descriptions

Restioids the restioid grassland showed growth through autumn and Restio leptoclados in dune fynbos and R. triticeus in grassy winter while the soil was waterlogged (Appendix 4). Restio fynbos, both on well drained sand, showed most growth in sieberi in the shale grassland grew most from April to June spring and autumn (Figure 3) when most rain fell (Figure 2). but also grew in the hot months of February/ March 1982 (Ap­ However, Elegia vaginulata behaved differently with growth pendix 7) when above average rainfall was noted. peaking in May (Figure 3), coinciding with the onset of winter Bond (1980) found a similar growth pattern for restioids waterlogging at that site. in the southern Cape, with maximal growth correlating with This winter shoot increment was very high, possibly as a moisture availability, though he also noted some summer result of post-fire regrowth in response to higher soil nutrients growth. Further westwards, in the south , restioids following a burn (Runde! 1983). start growth in spring/early summer (Kruger 1981) while some Qualitative observations on Thamnochortus fruticosus in species grow only in summer (J. Sommerville pers. comm.). 8 S.-Afr. Tydskr. Plantk., 1984, 3(1)

These data suggest that there is a trend for growth of restioids Ellis eta/. 1980). Our study confirms a cool, wet growth season

to occur earlier in the year the further east they occur in the for C 3 species in our area. Cape Region. Apart from rainfall differences, south eastern The study area in the south eastern Cape has equal numbers

Cape regions have higher mean winter temperatures and I - 2 h of species of C3 and C4 grasses (Cowling 1983b) which sug­ more bright sunshine per day relative to the south western Cape gests that this region of non-seasonal rainfall, lying between (Fuggle 1981). It is possible that restioid growth is temperature the winter rainfall area to the west and the summer rainfall controlled (Cowling 1983b). region to the east, is the overlap area for the c3 and c4 photo­ Bond (1980) predicted that winter drought in the summer synthetic modes (see also Vogel eta/. 1978). However, in terms

rainfall region would limit the eastward distribution of restioids of cover abundance, C4 species (particularly T. triandra and and reduce their competitive advantage. He inferred that a other species capable of cool season growth) predominate over

combination of high soil moisture, low temperatures and low C3 species in the study area (Cowling 1983b). We suggest that light conditions are a prerequisite for growth. However, the this high cover abundance is largely the result of the competitive

fact that some restioids grow only in summer during periods advantage of the C4 grass species with more than one growth of high temperature, high light intensity and low moisture, con­ season per year, in a region of mild temperatures and non­ tradicts this theory. Restioid roots appear to be shallow and seasonal rainfall.

able to exploit any chance summer rainfall. The limited The C4 grasses with more than one growth season, the C3 distribution of restioids eastwards of the Cape Region may grasses, the sedges and the restioids, all show some or all be related to factors such as limitations to seed dispersers (cf. growth in the autumn to spring period. All are shallow rooted Bond & Slingsby 1983) or the lack of highly infertile soils (cj. and able to exploit any summer rainfall but most of the growth Campbell 1983). occurs in the moderately warm, wetter months. These data refute the hypothesis that greater grass cover in eastern fyn­ Grasses bos communities is due to a distinct temporal separation of growth activities within grasses and restioids in a given com­ Counts of tillers and/ or leaves per shoot of the C4 grasses Themeda triandra (Figure 4) , Tristachya /eucothrix, munity (Cowling 1983a). Stenotaphrum secundatum and Sporobo/us ajricanus (Figure 5) showed more than one peak of maximum growth. Other Succulents c4 grasses which showed growth interrupted by a period of The succulents observed were: Aloe ajricana (Figure 7), inactivity were Cynodon dacty/on and Eragrostis capensis (Ap­ Crassu/a cu/trata and Sarcostemma viminale (Appendix 5) in pendix 4). In contrast, Elionurus muticus (Appendix 4) and the Kramme River thicket; Ruschia tene//a (Appendix 6) in Diheteropogon fi/ifo/ius (Appendix 3) had only one growth the renosterveld and Crassu/a expansa (Appendix 1) in the dune season, occurring in summer. fynbos. All these succulents grew in spring and autumn while

Growth measures of the C 3 grass species, Pentaschistis A. ajricana and S. vimina/e also grew in summer. The angustijolia and Lasioch/oa /ongifolia indicated mostly phenophases of A. africana (Figure 7) are similar to those of autumn/ early winter growth (Figure 6) while He/ictotrichon A. jerox, another arborescent, rosette succulent (Holland et hirtulum grew in autumn and spring (Appendix 7). These fin­ a/. 1977). dings support the generalized notion of cool season growth Most, if not all the succulents we studied probably have of C3 species and warm season growth of C4 species (e.g. Teeri crassulacean acid metabolism (CAM) (see Mooney eta/. 1977) & Stowe 1976; Ehleringer 1978; Boutton eta/. 1980). and possibly facultative CAM, whereby the mode of carbon

From our study and from analysis of Palmer's (1982) work fixation changes to C3 (Hartstock & Nobell976) and less com­ we were able to classify C4 grass species according to growth monly to C4 (Bartakke & Joshi 1976) under changing condi­ patterns of either one growth season or more than one growth tions of temperature and soil moisture. The autumn and spring season per year. We then compared the distributions (Meredith growth of succulents in our area may be explained by C3 fixa­ 1955) of these grass species in the Cape Region and found a tion during these moist, mild periods with small diurnal fluc­ correlation between growth pattern type and distribution. tuations in temperature. C3 fixation ensures much greater car­ Almost all species which had a continuous, summer growth bon fixation than CAM fixation which has low productivity season do not occur much west of Humansdorp, (e.g. Era­ (Ting & Szarek 1975; Kluge & Ting 1978). The additional sum­ grostis lehmanniana, Elionurus muticus, Diheteropogon fili­ mer growth of two succulent species may be the result of C4 jo/ius), while Panicum deustum does not penetrate the truly fixation. The strategy of CAM, and in particular facultative .winter rainfall area of less than 300Jo summer rain. These single­ CAM, is especially suited to a non-seasonal rainfall regime. growth season species are apparently limited to summer and CAM enables plants to tolerate dry periods in a vigorous, non­ non-seasonal rainfall areas which receive a critical level of sum­ dormant state so that response to any available moisture is mer rain. In contrast, those grass species with more than one rapid (Sutton & Osmond 1972). The shallow roots charac­ growth season all appear to penetrate into the winter rainfall teristic of most CAM succulents (Kluge & Ting 1978) would area of the south western Cape. also enable an almost immediate response to very light

The ability of both growth pattern types of C4 grasses to precipitation. grow during the summer enables them to dominate in the In the Fish River Valley of the eastern Cape, Palmer (1982) eastern, summer rainfall area of southern (cj. Vogel found similar spring and autumn growth patterns in some suc­ eta/. 1978). However, the capacity for growth during more culents (e.g. Crassu/a expansa, Crassu/a muscosa ( = than one season, the second season usually in early/ midwinter, lycopodioides)) while other succulents tended to grow at all clearly gives these species a competitive advantage in the winter times except in spring and autumn (e.g. Euphorbia bothae). rainfall region. C3 species are well adapted to the winter rainfall Optimum growth of Kalanchoe daigremontiniana requires low area, shown by their predominance in the south western Cape night and high day temperatures (Osmond eta/. 1976) which (Vogel eta/. 1978), because they require lower temperatures may explain the winter growth of K. rotundijolia and other associated with periods of low water stress (Boutton eta/. 1980); succulents in the Fish River Valley. S. Afr.). Bot. , 1984, 3(1) 9

Small leaved shrubs even in midsummer when the upper layers (0,3 - 0,6 m deep) Growth patterns showed mainly either summer peaks or lacked moisture. variable growth throughout the year. In some species, sum­ Levyns (1956) noted that the summer growth of the two mer growth was preceded by a spring increment (e.g. Erica shrubs, E. rhinocerotis and M. muricata, was 'not at harmony pectinifolia, Figure 9; litoralis, Appendix 1) or con­ with present climatic conditions'. Also, although seed is set tinued into autumn (e.g. Passerina vulgaris, Figure 8; Stoebe in time for favourable germination conditions, germination is plumosa, Appendix 3). Peak growth in summer was shown delayed for a year. These two factors suggested to her a sum­ by Disparago ericoides (Appendix 1) and Carpacoce vaginellata mer rainfall origin for these species. We suggest that their ability (Appendix 3), both in very well drained sand, and by Heli­ to tap underground water reserves does not limit their growth chrysum teretifo/ium in loam (Appendix 6). These are all small, during summer drought and an historical hypothesis need not shallow rooted species. Other small leaved shrubs which be invoked. showed most growth in summer were the deep rooted shrubs, Elytropappus rhinocerotis (Figure 10) and Meta/asia muricata Large leaved (sclerophyll) shrubs (Figures 8 & 10; Appendix 3), though slight, variable growth Thicket Species. No obvious pattern of growth emerged from was apparent in some individuals throughout the rest of the phenology of the large leaved sclerophyll shrubs in both the year. Intermittent growth throughout the year was apparent dune and Kramme River thickets, though there was a strong in Erica diaphana (Figure 9); Agathosma apiculata (Figure 8); tendency for growth to cease for a short period in midsum­ A . stenopeta/a, Sutera microphylla and Mura/tia squarrosa mer and all leaf fall occurred in summer (Appendices 1 and 5). (Appendix 1). In the southern Cape Mountain Fynbos, Erica In the dune thicket, new leaves and shoots were produced in seriphiifolia showed variable growth throughout the year while spring (e.g. Cussonia thyrsif/ora); winter, spring and autumn Phylica panicu/ata had a summer growth peak (Bond 1980). (e.g. Rapanea gilliana) and throughout most of the year (e.g. Bond (1980) inferred from the aseasonal growth of an ericoid Olea exasperata) (Appendix 1). In the Kramme River thicket, (sensu Campbell eta/. 1981) small leaved shrub in the southern spring and summer growth was most common although Rhus Cape that it had adopted a 'generalist' strategy (cf. Morrow incisa and R . /ongispina grew in autumn and R. g/auca grew & Mooney 1974) of growing whenever soil moisture and mostly from spring through the autumn but ceased growth temperature are suitable. In our study, the phenology of in December (Appendix 5). The 'generalist' strategy of con­ ericoids showing similarly variable, aseasonal growth could also tinuous carbon fixation and growth whenever conditions are be explained by this strategy. Bond (1980) predicted that this suitable is shown by deep rooted, large leaved sclerophyllous year-round growth of ericoids and small leaved sclerophyll shrubs of Californian (Morrow & Mooney 1974; shrubs might be flexible enough to allow subtle seasonal divi­ Mooney eta/. 1975; Mooney eta/. 1974; Mooney 1983). This sion of resources between species which could allow for high description may fit the behaviour in our area of some deep rooted thicket species with relatively long lived leaves (more species diversity within this growth form. His theory would 1 help to explain the coexistence in a small stand of numerous than 1 /z years, pers. obs.). species of Erica, Agathosma, Phylica and other ericoids. Tem­ Certain thicket species were observed growing at any time poral partitioning of resources between the two closely related of the year but when all thicket species were considered, spring Agathosma species, A. apiculata (Figure 8) and A. stenopetala and autumn growth predominated (Pierce 1983). Many species (Appendix 1) may be inferred from their staggered flowering can grow throughout the winter (e.g. Sideroxylon inerme, and fruiting phases. Both have similar sized flowers (Pillans Pteroc/astrus tricuspidatus, Rapanea gilliana) which indicates 1950) and may reduce competition for and that winter temperatures are not an important limiting factor dispersers (ants) by staggered reproductive phenophases. for the growth of thicket species in the mild coastal climate of the south eastern Cape. Leaf loss for all species was highest No simple explanation, such as the 'generalist' strategy, exists in summer. The predominance of midsummer leaf loss if usual­ for species which had growth peaks during the hottest, driest ly associated with low soil moisture (Frankie eta/. 1974; Kum­ summer months when moisture is presumably limiting. In con­ merow 1983). trast, growth rhythms of mediterranean-type shrublands (sensu Cowling (1983c) has argued that subtropical thicket species Di Castri 1980) are clearly related to soil moisture availability, would have migrated into the Cape Region from the north with most of the growth in spring (Rundel1977; Mooney 1983). east with the onset of warmer, wetter Holocene conditions after Productivity gradually decreases as summer drought progresses ± 12 000 B.P. Only those species with wide phenological flex­ (Mooney & Dunn 1970; Mooney 1983) . In the southern Cape, ibility would have been able to penetrate the non-seasonal rain­ Bond (1980) recorded maximal growth of the ericoid Phylica fall area as insufficient summer rainfall would severely restrict panicu/ata during the summer water deficit period. This ability summer growing and winter deciduous thicket species (e.g. of small leaved, fynbos shrubs, possibly all shallow rooted, Acacia caffra and A. schweinjurthil). We suggest that flexi­ to grow during hot periods of high water stress is difficult to bility in the growth phenophases of deep rooted, 'high cost­ explain and needs detailed studies on water budgets, rooting low profit' (Orians & Solbrig 1977) sclerophyll shrubs would depths and water stress tolerances (cf. Kruger 1981; Mooney pre-adapt them for penetration of the non-seasonal and winter 1983) . rainfall regions of the Cape. The ability to exploit bimodal Summer growth by the ericoid shrubs, Meta/asia muricata and winter rains has led to the successful penetration of the and Elytropappus rhinocerotis is more easily explained by their southern and south western Cape by two thicket species in par­ extensive root systems. The depth of the tap root alone of E. ticular. Sideroxylon inerme gains importance as a dune thicket rhinocerotis has been measured as 6,06 m and the laterals element further westwards of our area as far as the Cape Penin­ spread to a diameter of 4- 5 m (Scott & van Breda 1937); the sula and Pteroce/astrus tricuspidatus penetrates the dry (150mm 1 latter are probably effective exploiters of chance summer rains. y- ) area of the Cape west coast as far In the renosterveld community, the roots of these species easily as Elands Bay, where summer rain is less than 200Jo of the total. reached the pedocutanic horizon, 0,65 m deep, which was moist Flowering and fruiting patterns of thicket species were ir- 10 S.-Afr. Tydskr. Plantk., 1984, 3(1)

regular. However, when all species were considered, there were in the moister springtime and not in midsummer (Morrow & reproductive peaks in spring and autumn (Appendices I and 5; Mooney 1974; Runde! 1977; Mooney 1983). We suggest that Liversidge 1972; Siegfried 1982; Pierce 1983; K.L. Tinley pers. deep roots reaching underground water supples explain the comm.). Individuals within a species flower and fruit at dif­ summer growth of proteoids. Thus for example, Fernandez ferent times of the year and also show variability in the quan­ & Caldwell (1975) found that continued root growth to greater tity of their reproductive output (this study; Liversidge 1972; depths allowed a semi-desert shrub to transpire and fix car­ Frost 1976; K.L. Tinley pers. comm.). Of the nine species bon during the driest time of the year. monitored both in this study and in Liversidge's (1972) study Other large leaved proteoid shrubs which grow during sum­ of dune thicket near Port Elizabeth, seven species showed mer in a mediterranean-type climate are the dominant over­ marked differences in flowering and fruiting phenophases be­ storey shrubs (e.g. Banksia in Australian heath (Specht & tween the two study areas. Rayson 1957)). This 'out of phase' growth is explained as the There are major differences between Frost's (1976) data heritage of a tropically evolved flora, which now, by geographic from a south western Cape dune thicket and ours on reproduc­ circumstance and changing world climates, occupies a mediter­ tive phenophases of Euclea racemosa and Pteroce/astrus ranean climate area (Groves 1965). Bond (1980) considers the tricuspidatus, though strict comparison was limited by the short summer growth of proteoids to be better suited to a summer duration of her study (April- August). In the south western rainfall (or even non-seasonal) area, which further supports Cape, P. tricuspidatus has peaks of ripe fruit in May and the theory of a tropical or subtropical origin for the family August and a high proportion of unripe fruits throughout the Proteaceae (Johnson & Briggs 1975) and the genus Leucosper­ study period. Frost (1976) suggests that this staggering fruiting mum (Rourke 1972). Even the genus Leucadendron, for which pattern, even on individual shrubs, is a strategy to reduce com­ Williams (1972) has argued a mediterranean climate origin, petition for dispersers. In the south western Cape E. racemosa exhibits summer growth. set fruit in a short well-defined period (April- June). However, Within the constraints of limiting factors (e.g. light, in our area, E. racemosa produced unripe fruit over a long nutrients, temperature) growth of proteoids may be determined period culminating in a short period of mature fruit (October­ by other phenophases such as periodicity of pollinators e.g. December) (Appendix 1). Irregular timing of reproductive insects (Williams I972), birds (Rourke 1972) and mice (Rourke behaviour has been noted in tropical trees and has been & Wiens 1977), dispersers e.g. ants (Bond & Slingsby 1983), variously explained as strategies for pollination (Stiles 1977) and predators e.g. rodents (Bond 1983) may be the ultimate and possible escape from seed predation (Janzen 1969, 1970). determinants (cf. Mooney 1983). Asynchrony in fruit production and dispersal within a species may be a way of limiting competition (McKey 1975; Grubb Concluding Remarks 1977) and thus account for the relatively high alpha diversity This study is essentially a descriptive account of phenophases and lack of single species dominance in thickets in the south of a wide range of species and growth form in four major eastern Cape (Cowling 1983d). The proportion of viable seed vegetation types. It was not possible to explain adequately the of different species available to recolonize the small disturbance phenophases of all species. However, many hypotheses emerged patches characteristic of thicket will show appreciable temporal from this study and detailed autecological studies are required and spatial variability. In contrast, south western Cape dune to test these. For example, reciprocal transplant and controlled thickets are often dominated by a few species of thicket shrubs environment studies should test hypotheses regarding the com­ (viz. Sideroxylon inerme, Euclea racemosa, Pteroce/astrus petitive interplay between grasses and restioids. Experimental tricuspidatus, Olea exasperata). We argue that this is an his­ evidence is necessary to determine the relative roles of soil torical consequence of the westward depauperization in the moisture and temperature in determining the growth of thicket temperate Cape Region of subtropical thicket (cf. Cowling species. Ecophysiological and root behaviour studies should 1983a). explain the summer growth of certain fynbos species. Much Proteoid species. The proteoid Cape endemics, Leucadendron more work is required on the periodicity of pollinators and sa/ignum and cuneiforme (Figure 9) show sum­ dispersers before we can evaluate their effects in governing mer growth maxima with L. sa lignum continuing growth into phenophases. An experimental approach is likely to yield a autumn. Bond (1980) found similar growth patterns for two fuller understanding of phenophases in a shorter time than proteoids in the southern Cape. Leucadendron u/iginosum and more detailed long term observations (cf. Rutherford & , which grew mostly in the dry summer period. Panagos 1982). Our findings of a summer/autumn peak in L. sa/ignum agrees with the generalized phenology for the genus in the south Acknowledgements western Cape (Kruger 1981). L. cuneiforme, however, ceased We thank Ms J. Sommerville for her advice on methods. F.J. growth earlier than the time generalized for Leucospermum Kruger, J. Midgley and E.J. Moll criticized an earlier draft. in the south western Cape (Kruger 1981). This study forms part of the Fynbos Biome Project and was L. sa/ignum flowered in autumn/winter (Figure 9) and re­ funded by the C.S.I.R. We thank Ms M.L. Jarman and Ms tained its seeds for about a year, releasing them only when P. van Helsdingen (C.S.P.:C.S.I.R.) for liaison and other new seed was present in the next season's cones (Williams support. 1972). Maximum leaf fall followed peak growth of L. cuneiforme, but L. sa/ignum showed variable leaf loss (Figure References 9). Both proteoids studied are widespread in the Cape Region ACOCKS, J.P.H. 1953 . types of . Mem . Bot. Surv. and resprout from persistent woody rootstocks after fire S. Ajr. 28 . (Rourke 1972; Williams 1972). ANON. 1942. Weather on the coast of . Part 3. to East London. Government Printer, Pretoria. The two proteoid shrubs showed maximum growth in mid­ BARTAKKE, S.P. &. JOSHI, G,V. 1976. Crassulacean acid summer when soil water deficits were highest. Even the large metabolism and photosynthesis in Aloe barbadensis Mill. Proc. In­ leaved, evergreen 'generalists' of Californian chaparral grow dian Nat. Sci. A cad. 42: 227-233. S. Afr. J. Bot., 1984, 3(1) 11

BOND, W.J. 1980. Periodicity in fynbos of the non-seasonal rainfall HEYDORN, A.E.F. & TINLEY, K.L. 1980. Estuaries of the Cape. belt. 11. S. Afr. Bar. 46: 343 - 354. Part I. Synopsis of the Cape. CSIR Research Report 380, BOND, W .J. 1983. Seed predators, fire and Cape Proteaceae - Pretoria. limits of the population approach to succession. Vegerario (in HEYWARD, J. 1931. Flowering periods of Victorian plants. Proc. R. press). Soc. Vier. 43: !54 - 165. BOND, W.J. & SLINGSBY, P. 1983. Seed dispersal by ants in Cape HOLLAND, P.G., STEYN, D.G. & FUGGLE, R.F. 1977. shrublands, South Africa. Submitted to S. Afr. 1. Sci. occupation by Mill. (Liliaceae) in relation to BOUCHER, C. & MOLL, E.J. 1980. South African mediterranean topographic variations in direct beam solar radiation income. 1. shrublands. In: Mediterranean-type shrublands, eds. Di Castri, F., Biogeogr. 4: 61 - 72. Goodall, D.W. & Specht, R.L. Elsevier, Amsterdam. JANZEN, D.H. 1969. Seed-eaters versus seed size, number, toxicity BOUTTON, T.W., HARRISON, A.T. & SMITH, B.N. 1980. and dispersal. Evolurion 23: I - 27. Distribution of biomass of species differing in photosynthetic JANZEN, D. H. 1970. Herbivores and the number of tree species in pathway along an altitudinal transect in south-eastern Wyoming tropical forests. Am. Nar. 104: 501 - 528. grassland. Oecologia 45: 287-298. JOHNSON, L.A.S. & BRIGGS, B.G. 1975. On the Proteaceae- the CAMPBELL, B.M. 1983 . Montane plant environments in the Fynbos evolution and classification of a southern family. 1. Linn. Soc. Biome. Borha/ia 14: 283 - 298. (Botany) 70: 83- 182. CAMPBELL, B.M., COWLING, R.M., BOND, W.J. & KRUGER, KLUGE, M. & TJNG, I.P. 1978. Crassulacean acid metabolism. F.J. 1981. Structural characterization of vegetation in the Fynbos Analysis of an ecological adaptation. Springer, Berlin . Biome. S.A. Nat. Sci. Prog. Rep. No. 53. CSIR, Pretoria. KRUGER, F.J. 1978. A description of the Fynbos Biome Project. COWLING, R.M. 1983a. A syntaxonomic and synecological study in South African National Scientific Programmes Report No. 28. the Humansdorp region of the Fynbos Biome. Borha/ia 15. (in CSIR, Pretoria. press). KRUGER, F.J. 1979. South African Heathlands. In: Heathlands and COWLING, R.M. 1983b. The occurrence of CJ and grasses in c. related shrublands of the world. A. Descriptive studies, ed. Specht, fynbos and allied shrublands in the south eastern Cape, South R.L. Elsevier, Amsterdam. Africa. Oecologia 58: 121-127. KRUGER, F.J. 1981. Seasonal growth and flowering rhythms: South COWLING, R.M. 1983c. Phytochorology and vegetation history in African heathlands. In: Heathlands and related shrublands of the the south eastern Cape, South Africa. 1. Biogeogr. 10 (in press). world. B. Analytical studies, ed. Specht, R.L. Elsevier, Amsterdam. COWLING, R.M. 1983d. Diversity relations in Cape shrublands and KUMMEROW, J. 1983. Comparative phenology of mediterranean­ other vegetation in the south eastern Cape, South Africa. type plant communities. In: Mediterranean-type ecosystems, the Vegetatio (in press). role of nutrients, eds. Kruger, F.J., Mitchell, D.T. & Jarvis, COWLING, R.M., PIERCE, S.M. & MOLL, E.J. 1983. Management J.U.M. Springer, New York. and utilization of South Coast Renosterveld. Submitted to 11. S. LIVERSIDGE, R. 1972. A preliminary study on fru it production in Afr. Bot. certain plants. Ann. Cape Prov. Mus. (Nat. His.) 9: 51 - 63. DI CASTRI, F. 1980. Mediterranean-type shrublands of the world. LEVYNS, M.R. 1956. Notes on the biology and distribution of the In: Mediterranean-type shrublands of the world, eds. Di Castri, F., rhenoster bush. S. Afr. 1. Sci. 52: 141 - 143. Goodall, D.W. & Specht, R.L. Elsevier, Amsterdam. MacVICAR, C.N., DE VILLIERS, J.M., LOXTON, R.F., DICKINSON, C.E. & DODD, J.L. 1976. Phenological patterns in the VERSTER, E., LAMBRECHTS, J.J .N., MERRYWEATHER, short-grass prairie. Am. Midi. Nat. 96: 367 - 378. F.R., LEROUX, J., VAN ROOYEN, T.H. & VON M. HARM­ EHLERINGER, J .R. 1978. Implications of quantum yield differences SE, H.J. 1977. Soil classification. A binomial system for South on the distribution of C3 and C4 grasses. Oecologia 31: 255- 267. Africa. Sci. Bull. 390. Department of Agriculture, Pretoria. ELLIS, R.P., VOGEL, J.C. & FULS, A. 1980. Photosynthetic MEREDITH, D. (ed .) 1955. The grasses and pastures of South pathways and the geographical distributions of grasses in South Africa. C.N.A., Johannesburg. /Namibia. S. Afr. 1. Sci. 76: 307-314. McCALLUM, D.M. (ed.) 1972. Meteorology. Environmental Study, FERNANDEZ, O.A. & CALDWELL, M.M. 1975. Phenology and Swartkops River Basin. Hill, Kaplan, Scott and Partners, Cape dynamics of root growth of three cool semi-desert shrubs under Town. field conditions. 1. Ecol. 63: 703-714. McKEY, D. 1975. The ecology of co-evolved seed dispersal systems. FRANKIE, G.W., BAKER, H.G. & OPLER, P .A. 1974. Com­ In: Co-evolution of animals and plants, eds. Gilberts, L.E. & parative phenological studies of trees in tropical wet and dry Raven, P.H. University of Texas Press, Austin. forests in the lowlands of Costa Rica. 1. Ecol. 62: 881-919. MOONEY, H.A. 1983 . Carbon-gaining capacity and allocation pat­ FROST, S. 1976. A study of the fruiting phenology of three coastal terns of mediterranean-climate plants. In: Mediterranean-type dune macchia species. Botany III Project, University of Cape ecosystems. The role of nutrients, eds. Kruger, F.J., Mitchell, Town. D.T-. anc;l Jarvis, J.U.M. Springer, New York. FUGGLE, R.F. 1981. Macro-climatic patterns within the Fynbos MOONEY, H.A. & DUNN, E.L. 1970. Photosynthetic systems of Biome. Final Report, National Programme for Environmental mediterranean-climate shrubs and trees of California and Chile. Sciences. CSIR, Pretoria. Am. Nat. 104: 447 - 453. GIBBS RUSSELL, G.E. & ROBINSON, E.R. 1981. Phytogeography MOONEY, H.A., HARRISON, A.T. & MORROW, P.A. 1975. En­ and speciation in the vegetation of the eastern Cape. Botha/ia 13: vironmental limitations of photosynthesis in a California evergreen 467-472. shrub. Oecologia 19: 293-301. GOLDBLATT, P. 1978. An analysis of the flora of southern Africa; MOONEY, H.A., PARSONS, D.J. & KUMMEROW, J . 1974. Plant its characteristics, relationships and origins. Ann. Mo. bot. Gdn. development in mediterranean climates. In: Phenology and 65: 369-436. seasonality modelling, ed. Lieth, H. Springer, New York. GROVES, R.H. 1965 . Growth of heath vegetation. II. The seasonal MOONEY, H.A., TROUGHTON, J.H. & BERRY, J.A. 1977. Car­ growth of a heath on ground-water podzol at Wilson's promon­ bon isotope ratio measurements of succulent plants in southern tory, Victoria. Aust. 1. Bot. 13: 281-289. Africa. Oeco/ogia 30: 295-305. GRUBB, P.J. 1977. The maintenance of species-richness in plant MORROW, P.A. & MOONEY, H.A. 1974. Drought adaptations in communities: the importance of the regeneration niche. Bioi. Rev. two Californian evergreen sclerophylls. Oecologia 15 : 205-222. 52: 107 - 145. ORIANS, G.H. & SOLBRIG, O.T. 1977. A cost-income model of GUY, P.R., MAHLANGU, Z. & CHARIDZA, H. 1979. Phenology leaves and roots with special reference to arid and semi-arid areas. of some trees and shrubs in the Sengwa Wildlife Research Area, Am. Nat. Ill: 677 - 690. Zimbabwe-. S. Afr. 1. Wild/. Res. 9: 47- 54. OSMOND, C.B., BENDER, M.M. & BURRIS, R.H. 1976. Pathways HARTSTOCK, T.L. & NOBEL, P.S. 1976. Watering converts a of C02 fixation in the CAM plant Kalanchoe daigremontiniana. CAM plant to daytime C02 uptake. Nature 262 : 574- 576. III. Correlation with IJC values during growth and water stress. HEINRICH, B. 1976. Flowering phenologies: bog, woodland and Aust. 1. Plant Physiol. 3: 787-799. disturbed . Ecology 57: 890- 899. PALMER, A.R. 1982. A study of the vegetation of the Andries 12 S.-Afr. Tydskr. Plantk., 1984 , 3(1)

Vosloo Kudu Reserve. M.Sc. thesis, Rhodes University. PEET, R.K. 1978. Forest vegetation of the Colorado Front Range: Patterns of species diversity. Vegetatio 37: 65 - 78. PIERCE, S.M. 1983. A synthesis of plant phenology in the Fynbos Biome. S. Afr. Nat. Sci. Prog. Rep. CSIR, Pretoria (in press) . PIERCE, S.M. & COWLING, R.M. 1984. Seasonal growth of the overstorey and understorey in Mediterranean-type shrublands and heathlands in the south eastern Cape, South Africa. S. Afr. 1. Bot. 3: 17 -21. PILLANS, N.S. 1950. A revision of Agathosma. 11. S. Afr. Bot .. 16: 55 - 183 . PIPER, C.S. 1950. Soil and plant analysis. University of Adelaide, Adelaide. RAUNKIAER, C. 1934. The life forms of plants and statistical plant geography. Oxford University Press, Oxford. ROURKE, J.P. 1972. Taxonomic studies on Leucospermum R. Br. 11. S. Afr. Bot. Suppl. 8: 1- 194. ROURKE, J.P. & WIENS, D. 1977 . Convergent floral evolution in Appendix 1 Phenodiagrams of species in dune fyn­ South African and Australian Proteaceae and its possible bearing on pollination by non-flying mammals. Ann. Mo. bot. Gdn. 64: bos and thicket 1-17. RUNDEL, P. W. 1977. Water balance in mediterranean sclerophyll KEY ecosystems. Proceedings of the symposium on the environmental D Ripe fruit/ seed Ficinia ramosissima Kunth consequences of fire and fuel management in mediterranean D Unripe fruit/ seed ecosystems. USDA Forest Services General Technical Report •Full flower ea Pre-flower !llllilll!l!l I I !!! !) !! rrnmm W0-3. Palo Alto, California. liD Active growth RUNDEL, P.W. 1983. Impact of fire on nutrient cycles. In: D Slight growth ~No growth Mediterranean-type ecosystems. The role of nutrients, eds. Kruger, • Leaf loss F.J., Mitchell, D.T. & Jarvis, J.U.M. Springer, New York. "' Leaf death • Start of sampling RUTHERFORD, M.C. & PANAGOS, M.D. 1982. Seasonal woody Rhus crenata Thunb. plant shoot growth in Burkea africana - Ochna pulchra . S. Afr. 1. Bot. 1: 104-116. ,,,1111111111' SCOTT, J .l. & VAN BREDA, N.G. 1937. Preliminary studies on the Putterlickia pyracantha (L.) Szyszyl. root system of the rhenosterbos (Eiytropappus rhinocerotis) on the Worcester Veld Reserve. S. Afr. 1. Sci. 33 : 560 - 569. i!ll!i!jjll!l I I lll.!llllllll• !ll!lll!lllll ...Wachendorfia thyrsiflora L. SIEGFRIED, W.R. 1982. Trophic structure of some communities of Cassine aethiopica (Thunb.) fynbos birds. In: Proceedings of a symposium on coastal lowlands of the western Cape, ed. Moll, E.J. University of the Western ([II!Ill :::CI:dr'r:J a: Cape, Bellville. SPECHT, R.L. & RAYSON, P. 1957. Dark Island Heath (Ninety­ mile Plain, South Australia). I. Definition of the ecosystem. Aust. 1. Bot. 5: 52-85. SPECHT, R.L. & MOLL, E.J. 1983 . Heathlands and sclerophyllous shrublands - an overview. In: Mediterranean-type ecosystems. The role of nutrients, eds. Kruger, F.J. , Mitchell, D.T. & Jarvis,

J.U.M. Springer, New York. llll!llllllll I I iller: f!ll!!!ll!ll!ll STILES, F.G. 1977 . Co-adapted competitors: the flowering seasons of Olea exasperata Jacq. hummingbird pollinated plants in a tropical forest. Science 198 : 1177 - 1178 . 111111~ I I 111111111111\11111111'"" Rhus schlechteri Diels SUTTON, B.G. & OSMOND, C.B. 1972. Dark fixation of C02 by crassulacean plants - evidence for a single step. Plant Physiol. 50 360-365. TAYLOR, H.C. 1978 . Capensis. In: and ecology of southern Africa, ed. Werger, M.J.A. Junk, The Hague. TEERI, J . & STOWE, L. 1976. Climatic patterns and the distribution of c. grasses in . Oecologia 23 : I- 12. TING, J.P. & SZAREK, S.R. 1975 . Drought adaptation in crassula­ cean acid metabolism plants. In: Environmental physiology of desert organisms, ed. Hadley, N.F. Dowden, Hutchinson & Ross, Stroudsberg. UNDERHILL, L.G. 1981. Introstat. Juta, Cape Town. VAN ROOYEN, M.W., GROBBELAAR, N. & THERON, G.K. 1979. Phenology of the vegetation in the Hester Malan Nature Reserve in the Namaqualand Broken Veld . 11. S. Afr. Bot. 45: 279-293. VOGEL, J.C., FULS, A. & ELLIS, R.P. 1978. The geographical distribution of Kranz grasses in South Africa. S. Afr. 1. Sci. 74: 209-215. WERGER, M.J.A. (ed.) 1978. Biogeography and ecology of southern Africa. Junk, The Hague. WHITE, F. 1983. Vegetation of Africa. UNESCO, Paris (in press). WILLIAMS, O.B. 1971. Phenology of species common to three semi­ arid . Proc. Linn. Soc. N .S. W. 96: 193-203. WILLIAMS, I.J.M. 1972. A revision of the genus Leucadendron (Proteaceae). Contr. Bolus Herb. 3: 1-425. S. Afr. J. Bot. , 1984, 3(1) 13

Appendix 2 Phenodiagrams of species in dune grassland. Key: see Appendix 1

Centella cor iacea Nannfd. Cerastium,,,,,r capense Send. .. MedicaQo hispida Gaertn 57 ~ Torilus africana Spreng. Rhvnchosia caribaea (Jacq.) DC . Appendix 4 Phenodiagrams of species in restioid IIIII;: .1111 !!!!!Ill!!~~ grassland. Key: see Appendix 1 Sebaea minutiflora Schinz. · Tephrosia caoensis (Thunb.) Per s.

LI J I : :U:,Jii angustifolia (Lam.) Psoralea decumbens Ait. Cliffortia ferru!=Jine a l.f. R. Dahlgr. ssp. angustifo!ia

1 I I I I I I I I Ill! IIIII! I Satyrium princeps Bolus I I I I I I 11d I lllnll:!ll Aspa!athus spinosa L. ssp. Spinosa

!IIIII~«: Scirpus antarcticus L. lxia orientalis Ker. I I I ! I IIH I I I Ill! Commelina africana L. I I I I ,.cc D Cerastium capense Sand. J .. •1111 l!illlllll!llllllillll Romulea atrandra Lewis £. Cullumia decurrens Less aliciae Hamet Ill! I~ Icc 1 Spi!oxene ? stellata Oxa!is smithiana E.& Z. CTTTJ ' I I I 1 I i!?L 1 r 1 r n=r-cc lndigofera stricta L.f. Briza maxima L. ~ IIIII !!IIIII Micranthus plantagineus Eckl. r",~::,Ta!um cf. ecklonii lll./11!1!111 ~"' d Briza minor L. J. lndigofera incana Thunb. Monadenia bracteata (Sw.) Dru ~ ~~

Babiana patersoniae L. Cassine tetragona (L.f.) Loes. Manulea obovata Benth. Elionurus muticus (Spreng.) Kunth ...... ~ / '' , '' , '' , '', '', '', '' , '' , '' , ''!!!!!!II!!!!! : I ... IIIII ccc::IIIJ ~Ill c Hesperanth~ata (L.f.) Ker. Cotula sericea L.f. .fu Bu!bosty!is co!!ina Kunth =~<~:~:.~~?.~~:~:L. HII I I II I II,! Ill I I I I I I I I !liB I I I I rr::r::El Phy!ica !itoralis D. Dietr. Vigna unguicu!!ata (L.) Walp Oxa!is sp.

;: ''''' ··· · ·· · ·' ·'' ''' '' 'iitiriri1iirr:: I I [ I [ [ I I a: ITIJ Hu r I I I I I I llll_i.~Li __ b~d~ ; ,, , , , ~~~~~~~~~~~~n~~, Romulea dichotoma (Thunb.) Baker JFMAMJJASONDJFM JFMAMJJASONDJFM 1981 1982 1981 1982 _.p;:tl ~ Arctopus echinatus L. Sporobo!us africanus (Poir.l Robyns & ~- ~ UTournay II~ !ID Oxa!is po!yphy!!a Jacq. Appendix 3 Phenodiagrams of species in grassy Thamnochortus fruticosus Berg. fynbos. Key: see Appendix 1 111111111111 ~ rm Hypoxis sp. Gazania linearis (Thunb.) Druce

Diheteropogon fi!ifo!ius (Nees) Clayton UD Helichrysum cymosum (L.) Less. 1111111111.-: []] ~"' Hypochoeris· radicata L . Wachendorfia paniculata L. . rrrrrrrm Tetraria involucrata (Rottb.) C.B.C!. l!!llllll ~ lll!l • ll!!!!!!!!!! !lll!! ~ !ll AMJJASONDJFMA He!ichrysum teretifo!ium (L.) Less. 1981 1982

llllllllll!ll I !111111111 1-=:/111/1 Metalasia muricata (L.) Less. Se!ago g!omerata Thunb.

FE€1 ::: !111/111,;111 ~ I ! ! ! ! !!!Ill! Stoebe p!umosa Thunb. Carpacoce vaginellata Salter

tllllll~llll!lllrr JFMAMJJASONDJFM 1981 1982 1981 1982 14 S.-Afr. Tydskr. Plantk., 1984, 3(1)

Appendix 5 Phenodiagrams of species in Kromme Appendix 7 Phenodiagrams of species in shale River thicket. Key: see Appendix 1 grassland. Key: see Appendix 1

rMoraea sp. c:::I::::. c:c Olea africana Miller Euclea undulata Thunb. ·· ······ Gladiolus longicollis Bak. rn:n:r.m a: Ill ! Ill! I I l !l!l t l!ll l l!l 1: ~~~~:~~JdJe~f~:suta [CTTTJI:;~==~I~I;1~1 ~~~~~~~-~ Tarch~'Oiii7hJs ~atDPf.D Bulbine narciss/olia -=m 1 1 1 1 1r !!liM - Salm- Dyck ...;t "'''''''"''''''''''l Euclea schimperi (A.DC .) Dandy Aristea pusiiia CThunb.) Ker lxia orientalis Ker I I M : 1=·=·=3 , I m Babiana patersoniae L. 1 1 1Ill 1 1 1 : :1:=:::=4cr: ITIJIII Rhoict~ssus d1g1 !ata.f. llll~~~~~ ~~~~~1 g.m randt • . CD l!ll!ll! !l --:---' Arctopus echinatus L. I!!I!!III!!I!!!!I!M:: ::::=:=:=:) Sarcostemma viminale (L.) R. Br. Senecio erubescens Ait. rn:.l]a: Drosera cistiflora L. 11111 M ::::::Jcc ~ :11:1:11 :11 I ', . 1. :0101. ~ L 4 Linociera foveolata (E.Meye~obl. illJ.lilllL~~~~~~_ulilljjJJ Oxalis purpurea L. rrrnrmw "' Rhus glauca Desf. Roem. & Schult.

, . cr=J nrm::o.D D . § !IIIII III! Crassula ciliata Thunb. P1ttosporum Vlr!dtflorum S1ms. Rhus longispina E. & Z. .11111111111 1~111 Passerina rubra C.H. Wr. c:fill DO Ficinia tristachya (Rottb.l Nees 1 11 1 1 ..,arnn I I I I I Mayte~Tac~m inkt~ t~. t.) L~ • Rhu smc1s~ D mmrm .1111111111~111111111 dichondraefolium DC Tetraria cuspidata (Roll b.) C.B.CI.

I I 1 I r??JIIIIII mun ...1111 I I I I I !!IIIII :I!"I!J Restio sieberi Kunth Asparagus capensis L. El .Fl ' . •, Crassula cultrata L. Helictotrichon hirtulum(Steud) Helichrysum odoratissimum (L.) Less. ; Schweickerdt

lllllllllllllcllllll .11111111111111111~J FMAMJ J ASONDJ FM an an . Cynodon dactylon (L) Pers .~ 1981 1982 1981 1982 Tephrosia capensisCThunb.)Pers. ... •rrmn IIIK:IIIIIIIMJFMAMJJASONDJFM Appendix 6 Phenodiagram of species in 1981 1982 renosterveld. Key: see Appendix 1

Briza maxima L. ~ Lobelia erinus L. ~~~~~~~~--= Appendix 8 Classification of species into growth Sebaea aurea (L.t.) Roem. & Schult. [ I - form classes (see Table 2). Phenophases are given !-{esperantha falcata Ker I I I Sl in appendices and figures where indicated Moraea bellendini N.E.Br. ssp. bellendenii Thunbergiella filifolius (Lam.) Wolff. Micranthus plantagineus Eckl. c::::r:::. Ornithogalum subulatum Bak. o:::::J!I Dune fynbos and thicket (see Appendix I) Homalium sp. c. Moraea algoensis Goldbl. I I M Geophytes Romulea longipes Schltr. ! I I M :::-1 Satyrium princeps Bolus lxia orientalis Ker. o:=- Satyrium membranaceum sw 1 I M Wachendorjia thyrsiflora L. Bulbine narcissifolia Salm - Dyck ~ Oxalis sp. 1 c:::I::::. Restioids Babiana patersoniae L. 11!!1!1 I! p:w :q Restio leptoclados Mast. (Figure 3) Oxalis sp. 2 Oxalis polyphylla Jacq. Cyperoids Spiloxene minuta (L.) Fourc. :--CD::::J Ornithogalum minuatum Jacq. Ficinia ramosissima Kunth MAMJJASONDJFM- Succulents Helichrysum teretifolium (L.) Less Crassula expansa Ait. ssp filicaulis (Haw.) Toelk. Small leaved shrubs lllllll,llm:lllllllll Eu clea crispa (Thunb.) GUrke Muraltia squarrosa (Thunb.) DC Sutera microphylla (L.F.) Hiern mmrm Cotula turbinata L. Phylica litoralis D. Dietr. Lobostemon argenteus Buek. Disparago ericodes Gaertn. Chironia baccifera L. 111111111 :IIIJI111111111111 lndigofera denudata Thunb. Agathosma stenopeta/a Steud. 111111111111111111111~ echinata (Thunb.) Nees Aspalathus spinosa L. Anthospermum aethiopicum L. ssp. Spinosa Meta/asia muricata (L.) Less. (Figure 8) rmnr=n I Ill! I I Agathosma apicu/ata G.F.W. Meyer (Figure 8) Cliffortia linearifolia E. & z. Gnaphalium repens L. Passerina vulgaris Thoday (Figure 8) Large leaved shrubs llillllllilll!l Ill! I I I I I ! I lvl !!IIIII Pelargonium dichondraefolium DC. Ruschia teneua \Haw.} Schw. Thicket Species Euclea racemosa Murr. '"IMAMJJASONDJFM :lllllllln ~11111~11111111~MAMJ JASON OJ FM Rhus crenata Thunb. 1981 1982 1981 1982 Putterlickia pyracantha (L.) Szyszyl. S. Afr. J. Bot., 1984, 3(1) 15

Cassine aethiopica fhunb. Restioid grassland (see Appendix 4) Olea exasperata Jacq. Annuals Rhus schlechteri Diels Scirpus antarcticus L. Cussonia thyrsijlora Thunb. Cerastium capense Sond. Rapanea gi/liana (Sond.) Mez. Drosera a/iciae Hamet Sideroxylon inerme L. (Figure II) Briza maxima L. Pterocelastrus tricuspidatus (Lam.) Sond. (Figure II) Briza minor L. Geophytes Dune grassland (see Appendix 2) Monadenia bracteata (Sw.) Dru. Annuals Spiloxene? stel/ata Cerastium capense Sond. Micranthus plantagineus Eckl. Medicago hispida Gaertn Babiana patersoniae L. Tori/us africana Spreng Hesperantha fa/cata (L. f.) Ker. Sebaea minutiflora Schinz. Oxalis sp. Geophytes Romulea dichotoma (Thunb.) Baker Satyrium princeps Bolus Arctopus echinatus L. Ixia orienta/is Ker. Oxa/is po/yphyl/a Jacq. Romulea attandra Lewis Hypoxis sp. Oxa/is smithiana E. & Z. Wachendorfia panicu/ata L. Ornithogalum cf. ecklonii Non-geophytic forbs Non-geophytic forbs Gazania linearis (Thunb.) Druce Centel/a coriacea Nannfd. Hypochoeris radicata L. Rhynchosia caribaea (Jacq.) DC Commelina africana L. Tephrosia capensis (Thunb.) Pers Grasses Psora/eo decumbens Ait. c. Elionurus muticus (Spreng.) Kunth Sutera campanulata (Benth.) 0. Kze. Cynodon dactylon (L.) Pers. Senecio burchellii DC Eragrostis capensis (Thunb.) Trin. Indigo/era strict a L. f . Sporobolus africanus (Poir.) Robyns & Tournay Indigo/era incana Thunb. Themeda triandra Forsk. (Figure 4) Manu/eo obovata Benth. Tristachya leucothrix Nees (Figure 5) Co tufa sericea L. f. Restioids Vigna unguiculata (L.) Walp Thamnochortus fruticosus Berg. Grasses Elegia vaginu/ata Mast. (Figure 3) Cyperoids c. Sporobolus ajricanus (Poir) Robyns & Tournay Kunth Themeda triandra Forsk. (Figure 4) Bulbostylis col/ina Stenotaphrum secundatum (Walt.) 0 . Kuntze (Figure 5) Small leaved shrubs C/iffortia jerruginea L.f. C 3 Lasiochloa longijolia (Schrad.) Kunth (Figure 6) Small leaved shrubs Aspalathus angustifolia (Lam.) R.Dahlgr. ssp. angustifolia Phylica litoralis D. Dietr. Aspa/athus spinosa L. Cul/umia decurrens Less Kromme River thicket (see Appendix 5) Large leaved shrubs Thicket species Succulents Cassine tetragona (L.f.) Loes Sarcostemma viminale (L.) R.Br. Rhus laevigata L. Crassula cultrata L. Large leaved shrubs Grassy fynbos (see Appendix 3) Thicket species Grasses Olea africana Miller Tarchonanthus camphoratus L. C4 Diheteropogon filifolius (Nees) Clayton Restioids Rhoicissus digitata (L.f.) Gilg. & Brandt. Restio triticeus Rottb. (Figure 3). Linociera joveolata (E. Meyer) Knob!. Cyperoids PiUosporum viridiflorum Sims. Tetraria invo/ucrata (Rottb.) C.B.Cl. Maytenus acuminata (L.f.) Loes Small leaved shrubs Lachnostylis hirta (L.f.) Mull. Arg. Meta/asia muricata (L.) Less. Rhoiacarpos capensis (Harv.) A.DC. Stoebe plumosa Thunb. Euclea undulata Thunb. Helichrysum cymosum (L.) Less. Euclea schimperi (A. DC.) Dandy Helichrysum teretifolium (L.) Less. Rhus glauco Desf. Selago glomerata Thunb. Rhus longispina E. & Z. Carpacoce vaginellata Salter Rhus incisa L. f. Erica diaphana Spreng. (Figure 9) Pterocelastrus tricuspidatus (Lam.) Sond. (Figure II) Erica pectinifolia Salisb. (Figure 9) Sideroxylon inerme L. (Figure II). Large leaved shrubs Renosterveld (see Appendix 6) Proteoids Leucospermum cuneiforme (Burm. f.) Rourke (Figure 9) Annuals Berg. (Figure 9) . Briza maxima L. 16 S.-Afr. Tydskr. Plantk., 1984, 3(1)

Lobelia erinus L. Large leaved shrubs Sebaea aurea (L.f.) Roem. & Schult. Thicket species Geophytes Euclea crispa (Thunb.) Giirke Hesperantha fa/cata Ker. Moraea bellendinii N.E.Br. Shale grassland (see Appendix 7) Thunbergiella filifo/ius (Lam.) Wolff. Annuals Mircranthus plantagineus Eckl. Senecio erubescens Ait. Ornithogalum subu/atum Bak. Drosera cistiflora L. Homalium sp. Sebaea aurea (L.f.) Roem. & Schult. Moraea algoensis Goldbl. Geophytes Romulea longipes Schltr. Moraea sp. Ixia orienta/is Ker . Oxalis purpurea L. Satyrium membranaceum Sw. Galdiolus longicollis Bak. Bulbine narcissifo/ia Salm-Dyck Piloselloides hirsuta (Forsk.) Jeffrey Oxalis sp. I Bulbine narcissifolia Salm-Dyck Babiana patersoniae L. Aristea pusi/la (Thunb.) Ker Oxalis sp. 2. Ixia orienta/is Ker Oxalis polyphylla Jacq. Babiana patersoniae L. Spiloxene minuta (L.) Fourc. Arctopus echinatus L. Ornithogalum minuatum Jacq. Non-geophytic forbs Non-geophytic forbs Pelargonium dichondraejo/ium DC Pelargonium dichondraefolium DC Tephrosia capensis (Thunb.) Pers. Cotula turbinata L. Grasses Succulents C3 Helictotrichon hirtulum (Steud) Schweickerdt Ruschia tenella (Haw.) Schw. C4 Cynodon dactylon (L.) Pers. Small leaved shrubs Restioids Aspa/athus spinosa L. Restio sieberi Kunth. Cliffortia linearifolia E. & Z. Meta/asia muricata (L.) Less (Figure 10) Cyperoids E/ytropappus rhinocerotis (L.f.) Less (Figure 10) Ficinia tristachya (Rottb.) Nees Helichrysum teretifolium (L.) Less Tetraria cuspidata (Rottb.) C.B.Cl. Helichrysum anomalum (Sch. Bip.) Less Succulent Chrysocoma tenuifolia Berg. Crassu/a ciliata Thunb. Se/ago corymbosa L. Small leaved shrubs Gnaphalium repens L. Passerina rubra C.H. Wr. Lobostemon argenteus Buek. Asparagus capensis L. Indigofera denudata Thunb. Helichrysum odoratissimum (L.) Less