Modern vegetation at the Klasies River archaeological sites, Tsitsikamma coast, south-eastern Cape, South Africa: a reference collection Authors: van Wijk, Yvette; Tusenius, Madelon L.; Rust, Renee; Cowling, Richard M.; Wurz, Sarah https://doi.org/10.5091/plecevo.2017.1286

Abstract

Background and aims – The Klasies River cultural landscape, on the Tsitsikamma coast, south-eastern
Cape, South Africa, features prominently in modern human origins research. The archaeobotanical
information for the Klasies River landscape and its immediate environment is sparse. The aim of this
study is the collection of a taxonomically valid and comprehensive reference database of modern botanical
specimens as an aid to identifying macro- and micro-botanicals such as seeds, charcoal, phytoliths,
parenchyma and pollen in the Klasies River archaeological deposits. This is an essential step in providing
context for the identification of past vegetation and its usage by Stone Age populations.
Methods – Herb, shrub, tree, grass, fern and geophyte voucher specimens were collected in 24 areas in
the vicinity of the Klasies River sites, and further inland within a 5km radius, between 2013 and 2015. The
collecting was done at different times of year so that all stages of the flowering, fruiting and seeding cycles
for most plants could be sampled.
Key results – A total of 268 species, in 196 genera and 78 families were collected. Only 69 of these
268 species currently appear on the relevant database grid of the Integrated Biodiversity Information
System (SIBIS), the South African National Biodiversity Institute (SANBI). Our work clearly indicates
the need for thorough and systematic collecting at archaeologically significant sites in the Cape region to
provide further environmental proxies for the interpretation and contextualisation of the development of
anatomically modern human behaviour.
Conclusions – The Klasies River landscape, although located within the broad Fynbos Biome, cannot
be classified as such, as relatively few fynbos species are represented in the core area surrounding the
sites. The vegetation is in fact a complex mosaic of thicket, forest and coastal vegetation. This densely
interdigitated vegetation provides a wide variety of useful resources.
Key words – Klasies River archaeological sites, Tsitsikamma region, southern Cape, south-eastern Cape,
thicket, forest, coastal vegetation, fynbos, medicinal plants, edible plants.

Full text

Plant Ecology and Evolution 150 (1): 13–34, 2017
https://doi.org/10.5091/plecevo.2017.1286
Modern vegetation at the Klasies River archaeological sites, Tsitsikamma
coast, south-eastern Cape, South Africa: a reference collection
Yvette van Wijk1,*, Madelon L. Tusenius2, Renee Rust3, Richard M. Cowling4,5 & Sarah Wurz6
1Botany Department, Rhodes University, Grahamstown, 6140, South Africa
2School of Geography, Archaeology and Environmental Studies, University of the Witwatersrand, Johannesburg, 2050, South Africa
3School of Geography, Archaeology and Environmental Studies, University of the Witwatersrand, Johannesburg, 2050, South Africa
4Botany Department, Nelson Mandela University, Port Elizabeth, South Africa
5Centre for Coastal Palaeosciences, Nelson Mandela Metropolitan University, Port Elizabeth 6031, South Africa
6School of Geography, Archaeology and Environmental Studies, University of the Witwatersrand, Johannesburg, 2050, South Africa
*Author for correspondence: vanwijkyvette@gmail.com
INTRODUCTION
The Klasies River cultural landscape is situated on the Tsi-
tsikamma coast, south-eastern Cape, South Africa, about
90kmwestofPortElizabeth(g.1).Thislandscape,aNa-
tional Heritage site, consists of a 2 km stretch of coast be-
tween the mouth of the Klasies River (known locally as the
Kaapserivier) to the west and Druipkelder Point to the east
andincludesbuerzonesintotheintertidalzoneandcoastal
platform towards the south and north respectively. There are
ve major archaeological features on this landscape: Main
Site(gs1 &2)occurs closest toKlasiesRiver mouth and
consistsofa complexofcaves(Caves 1,1A,1B, 1C &2).
Caves 3 and 4, not yet investigated archaeologically, occur
further to the east, with Cave 5, the eastern most feature,
close to Druipkelder Point. Various parts of the Klasies River
landscapehavebeenexcavatedsincethe1960sbySinger&
Wymer (1982), Deacon (1986, 1989, 1995) and Binneman
(1995), with Sarah Wurz, University of the Witwatersrand,
starting a new project in 2014. The excavations at Main Site
All rights reserved. © 2017 Botanic Garden Meise and Royal Botanical Society of Belgium
ISSN: 2032-3913 (print) – 2032-3921 (online)
REGULAR PAPER
Background and aims – The Klasies River cultural landscape, on the Tsitsikamma coast, south-eastern
Cape, South Africa, features prominently in modern human origins research. The archaeobotanical
information for the Klasies River landscape and its immediate environment is sparse. The aim of this
study is the collection of a taxonomically valid and comprehensive reference database of modern botanical
specimens as an aid to identifying macro- and micro-botanicals such as seeds, charcoal, phytoliths,
parenchyma and pollen in the Klasies River archaeological deposits. This is an essential step in providing
contextfortheidenticationofpastvegetationanditsusagebyStoneAgepopulations.
Methods – Herb, shrub, tree, grass, fern and geophyte voucher specimens were collected in 24 areas in
the vicinity of the Klasies River sites, and further inland within a 5km radius, between 2013 and 2015. The
collectingwasdoneatdierenttimesofyearsothatallstagesoftheowering,fruitingandseedingcycles
for most plants could be sampled.
Key results – A total of 268 species, in 196 genera and 78 families were collected. Only 69 of these
268 species currently appear on the relevant database grid of the Integrated Biodiversity Information
System (SIBIS), the South African National Biodiversity Institute (SANBI). Our work clearly indicates
theneedforthoroughandsystematiccollectingatarchaeologicallysignicantsitesintheCaperegionto
provide further environmental proxies for the interpretation and contextualisation of the development of
anatomically modern human behaviour.
Conclusions – The Klasies River landscape, although located within the broad Fynbos Biome, cannot
beclassied as such, as relatively few fynbos species are represented in the core area surrounding the
sites. The vegetation is in fact a complex mosaic of thicket, forest and coastal vegetation. This densely
interdigitated vegetation provides a wide variety of useful resources.
Key words – Klasies River archaeological sites, Tsitsikamma region, southern Cape, south-eastern Cape,
thicket, forest, coastal vegetation, fynbos, medicinal plants, edible plants.

14
Pl. Ecol. Evol. 150 (1), 2017
and Cave 5 have revealed MSA and LSA occupation of the
sites from the rst part of the Late Pleistocene to the Late
Holocene. Acheulean bifaces and MSA artefacts have been
noted in the deation hollows of the fossilized Geelhout-
boomduneontheplateauabovethecavesites(g.1,Laidler
1947,Deacon&Geleijnse1988).
Main Site was intensively occupied by coastal foragers
between c. 120 000 and 55 000 years ago and thus preserves
an extraordinarily comprehensive record of cultural adapta-
tion within the context of one of the earliest MSA coastal
economies that has been recorded (Singer & Wymer 1982,
Deacon1989,Deacon&Geleijnse1988,Wurz2002,2012).
Cave 5 also preserves evidence of a pulse of occupation
contemporaneous with ca 120 000 year old layers at Main
Site(Singer&Wymer1982).Someoftheearliest andmost
extensive fossil evidence for early anatomically modern hu-
mans occurs at Main Site (Singer & Wymer 1982, Deacon
2008, Grine et al. 2017). From around c. 4800 BP–2300 BP
(Nami et al. 2016), coastal dwellers again occupied Main
Site,andalso Cave5(Singer &Wymer1982), demonstrat-
ing that this landscape has provided a favourable habitat over
a long period of time. Several studies have already been un-
dertaken to provide insight into the palaeo-environmental as-
Figure 1 – Locality of the Klasies River archaeological sites,
Tsitsikamma coast, South Africa. AdaptedfromDeacon&Geleijnse
(1988).
Figure 2 – Panorama of Main Site, situated in the complex Klasies River coastal landscape.
pects of these Stone Age occupations, for example, the anal-
yses of the large mammal fauna, microfauna, shellsh and
isotopes(e.g.Klein1976,Singer&Wymer1982,Deaconet
al. 1986, Avery 1986, 1987, Deacon et al. 1988, Thackeray
1988, Van Pletzen 2000, Langejans et al. 2012). More recent-
ly Late Pleistocene speleothems from Cave 1C have been
studied (Braun 2014). Several on-going research projects are
referred to below as unpublished research (unpubl. res.).
Archaeobotanical information for the Klasies River land-
scape and its immediate environment is sparse, although pre-
liminary studies of small charcoal samples were undertaken
(Deacon et al. 1986, M.L. Tusenius, University of the Wit-
watersrand, South Africa, pers. comm.), and more recently,
therst seeds(Zwane,WitwatersrandUniversity,Johannes-
burg, SA, unpubl.res.), and parenchyma (C. Larbey et al.,
Cambridge University, UK, unpubl. res.) have been identi-
ed.Besidesobtaining dataonpast vegetation, archaeobot-
anical analyses at other sites, for example Sibudu Cave, have
shown that they have much potential to provide unexpected,
nuanced interpretations of populations’ behaviour and ad-
aptation (e.g. Wadley 2015). Plants provide micronutrients,
not available anywhere else, which are essential to human
diet and health – humans cannot live without plants as food
and medicine. Seeds, fruit, leaves, roots, tubers, bulbs, gum,
buds, ower stalks, internodes, nectar (De Vynck et al.
2016a), plus shoots, bark, cambium, sap, pollen, pods, and
galls were, and still are, all utilised. Deacon (1989, 1992,
1993) suggested that a long history of ethnographically
known plant exploitation may have been evident at Klasies
River since the Late Pleistocene. He interpreted carbonised
material observed in blackened layers at the site as altered
plant remains (Deacon 1993, 1995). Deacon (1993) also sug-
gested that when above-ground food resources were limited,
plants with underground storage organs (USOs) would have
been important sources of carbohydrates (see De Vynck et al.
2016a, 2016b, Singels et al. 2016). USOs may not have been
the only carbohydrate resource at Klasies River or elsewhere
as grasses may also have been an important food resource
(e.g. Henry et al. 2014, Eoin 2016).
One of the goals of the current excavation project at
Klasies River is to expand archaeobotanical investigation
for both the LSA and MSA deposits. In archaeological re-
ports, the botanical context provided for the Klasies cave
sites is noted as fynbos, either as a vegetation type or as a

15
van Wijk et al., Modern vegetation in the Klasies River cultural landscape
biome (e.g. Klein 1976, Faith 2011, Henry et al. 2014) or as
a complex forest-scrub and forest-grassveld-macchia mosaic
(Butzer 1978), with some reference to Afromontane Forest
(Deacon 1993). With the exception of specimens collected
for wood samples by Tusenius in 1984 and 1985, no system-
atic sampling of the composition of the vegetation has been
undertaken at the Klasies River cultural landscape and its
immediate environment. An essential step in providing com-
parative material and context for past vegetation and its us-
age by the Stone Age populations is to produce a taxonomi-
cally valid and comprehensive botanical database.
In this paper we report on the collection of 90–95% per-
centage of the modern botanical species in close proximity
to the Klasies River Main Site and Caves 3 and 4, and we
describe and characterise the present vegetation within the
regionalbiophysicalcontext(tables1&2).Wealsoindicate
which of the taxa collected are edible and/or medicinal or
haveotheruses(tables1,3&4),andweemphasisethehigh
prevalence of useful species collected. The latter topic is,
however, to be covered more comprehensively in another pa-
per(Y.vanWijk&R.Rust,RhodesUniversity,SouthAfrica,
unpubl. res.). The vegetation we sampled in detail in close
proximity to the Klasies River sites is a complex mosaic of
predominantly thicket, forest and coastal vegetation, with a
few fynbos elements.
BIOPHYSICAL CONTEXT
Klasies River falls within the Cape Fold Belt geomorphic
province where the dominant bedrock comprises quartzitic
sandstones of the Peninsula Formation of the Table Mountain
Group (TMG). These, together with other formations of the
TMG and some narrow bands of Bokkeveld Group shales,
form the Tsitsikamma coastal plateau, a narrow (50–13 km)
Figure 3 – Greater collection area within 5 kilometres of Main Site:
20 Fynbos remnant, 21 Forest remnant, 22 above cave 5, 23 on
plateau, 24 Kamsedrif.
relatively level plain ranging from c. 150–260 m in altitude
and located between the coast and the Tsitsikamma Moun-
tains. The plateau is massively incised by the narrow gorges
of short rivers such as the Klasies and Tsitsikamma rivers,
locally named Kaapserivier and Kamrivier, hence Kaapse-
drif and Kamsedrif occurring between the mountains and the
coast(g.3).Periodsof lowersealevelsinthepast,for ex-
ample, between 100000 and 55000 BP, would have had sig-
nicantimpactson thevegetationand past resourcesasthe
coast would have been between 2 and 5 km away (Langejans
et al. 2012). At present, the coastal margin, including the c.
100–150mcoastalclis,ismantledbya2–3kmwidecoast-
al band of Cenozoic (Algoa Group) sediments of a coastal
and aeolian origin (Le Roux 2000). These sediments include
the Nanaga Formation –Pliocene to Early Pleistocene aeo-
lian deposits, e.g. those of the Geelhoutboom fossil dune
(Deacon& Geleijnse1988)– as wellasthe unconsolidated
windblown sand of the vegetated dunes of Holocene age (Le
Roux 2000). The sea-land contact zone at present is deep-
ly indented and fractured with ragged rocks forming small
points, tunnels, and narrow bands which are vegetated right
to the edge of the rocks. Rock outcrops alternate with small
sandy bays, often covered in a thick layer of rounded quartz-
ite cobbles of varied size and colour, which were utilised for
tool-makinginthepast(Singer&Wymer1982).
Seasonal temperature variation is low and this equable
climateisaconsequenceofthestabilizingeectofthewarm
Agulhas Current. Between 1959 and 2001, the annual aver-
age minimum and maximum temperatures recorded at Cape
St Francis, some 60 km east of Klasies River, were between
9 and 18°C in winter and between 18 and 23°C in summer
(DuPlessis2015).Becauseof the cooling eect of the on-
shore winds bringing mist and sea-spray, temperatures are
lower in the vicinity of the sites than up on the plateau and
inland (Lubke 1985). As is typical of the Cape south coast,
rainfall is all year round with small peaks during autumn and
spring(Weldon&Reason2014).Theaverageannualrainfall
on the southern slopes of the nearby Tsitsikamma mountains
isbetween875and1,375mm(Hosking&du Preez 1999)
and at Storms River weather station 25 km west of Klasies
River it is 810 mm (Grey et al. 1987). This is optimal for
Afrotemperate forest and more than enough to sustain thick-
et. The prevailing winds are from the west-southwest in win-
ter, but in summer the frequency and strength of east-south-
east winds increases (Elkington 2012, Lubke 1985).
The Klasies River sites are located in the broad Eastern
Fynbos and Renosterveld bioregion of the Cape Floristic
Region’sFynbosbiome(Mucina&Rutherford2006).How-
ever, as is typical of this bioregion, Fynbos and Renosterveld
interdigitate and form mosaic structures with three other
biomes, namely Forest, Subtropical Thicket and Grassland
(Cowling 1982, 1984, Cowling & Potts 2015). There are
species-based botanical assessments relevant to the broader
area which encompasses Klasies River, such as that of the
southern Cape forests (Geldenhuys 1993, 1993), the Tsitsi-
kamma National Park to the west (Hanekom et al. 1989) and
the Oyster Bay dunes to the east (Cowling 1984). Several na-
tional or regional vegetation maps, mainly at relatively crude
scales do exist for the area (e.g.Acocks 1988, Cowling &
Heijnis2001,Mucina&Rutherford2006,Vloketal.2008),

16
Pl. Ecol. Evol. 150 (1), 2017
but all are too coarse to realistically depict vegetation pat-
terns in the immediate Klasies River area.
In general, vegetation of the Algoa Group sediments
along the coastal margin comprises a complex array of veg-
etation types whose distribution is governed by exposure to
salt laden winds, exposure to recurrent re, soil formation
and aspect. However, owing to heavy infestations of alien
trees (Acacia longifolia (Andrews) Willd., A. saligna (La-
bill.) Wendl.), especially on the inland slopes of the dune
cordon,itisdicult todiscern theoriginal vegetationcom-
position. The Nanaga sands support a complex mosaic of for-
est, thicket, grassland and fynbos, as is the case further east
of Klasies River (Cowling 1984, Hoare et al. 2006). In wind-
and re-protected sites such as incised drainage lines lead-
ing to the coast, the vegetation is Southern Coastal Forest, a
subtropical type with a relatively rich tree ora. Conspicu-
ous species include Sideroxylon inerme, Ekebergia capensis,
Olea capensis subsp. capensis, Canthium inerme, Pteroce-
lastrus tricuspidatus, Gymnosporia nemorosa, Cassine pera-
gua, Acokanthera oppositifolia and Searsia chirindensis.
Drier and more exposed sites support patches of dune
thicket, which may occur in large blocks, such as the wind-
exposed slopes along the coast, or as clumps in a matrix of
grassland or fynbos. The composition is similar to the dune
thickets described by Cowling (1984). Dominant species are
Sideroxylon inerme, Pterocelastrus tricuspidatus, Euclea
racemosa, Putterlickia pyracantha, Tarchonanthus littoralis
and Searsia glauca. The younger (Holocene) dunes support
a mosaic of dune fynbos and thicket, a community that has
been well described by Cowling (1984) in the Cape St Fran-
cis area.
Along the coast, either on hummock dunes or on harder
surfaces exposed to salt-laden winds is an azonal community
of low herbs and shrubs, many of which are succulent. This
communityformspartofMucina&Rutherford’sCapeSea-
shore Vegetation (2006). The vegetation of the coastal plateau
Figure 4 – Core collecting areas numbered and described.
1 at entrance to Cave 1a – badly worn and trampled, succulents and grass
2 rubble berm on lower eastern edge of Cave 1c – disturbed and unstable
2a low veg in slight depression and wash-away to east of area 2 – fresh water seep present
3 highnorthernendofbermbesideCave1c–milkwoodforest&thicketfrom20cmto2m
4 concave slope rising to the north – grassy species-rich wind-sheared vegetation
4a Similarto4butincludingwind-sheared‘Lilliputian’forest&thicketspecies
5 steeplyslopingareabetweenshore&clitocave1b–lowgrassywind-shearedthicket
6 immediatelyatentrancetoCave3–densethornythicket&forest
7 immediately at entrance to Cave 4 – thorny thicket, succulents and herbs
8 mixedspecies-richgrassyveg-thicketinleeofwind,slopinguptoclisonnorth-west
8a densevegetation,mostlythicket&forestupto2.5m
9 long valley sloping steeply up west, grassy, succulents, geophytes, thicket in lee of wind
10 grassyatsbetweenclisandseashore–lowgrass,succulent,coastal/littoral
10a similar to 10 but with more low shrubby and succulent species / littoral
11 from the edge of rocks above sea-shore – from low grassy scrub to thicket 1 m
12 along both sides of entrance track sloping south – dense coastal thicket
13 along both sides of entrance track sloping west – thicket, forest, lianas
14 along both sides of entrance track sloping north – thicket, + few fynbos spp.
15 smallcoastalforestpatchbelowcaves3&4–shelteredbyfore-dunewiththicket
16 narrowbandonedgeofbeach&fore-dune–lowcoastalveg.,thicketateasternend
17 small patch of 3–4 m forest and thicket, some disturbed grassy area
18 along footpath above rocky edge to seashore – thicket to 1.5 m+, below forest patch
19 smallbeach&clionwestbankofKlasiesRiver-cli&estuarinevegetation.

17
van Wijk et al., Modern vegetation in the Klasies River cultural landscape
is included in the Tsitsikamma Sandstone Fynbos of Rebelo
et al. (2006). Remnant patches of fynbos on well-drained pla-
teau areas suggest dominance by overstorey proteoid shrubs
with a lower stratum of ericoid shrubs, restiods and grasses.
Geophytesare conspicuousafter re.Almostallofthisear-
lier vegetation on the plateau has, however, been replaced by
pastures, pine plantations and dense stands of alien trees.
MATERIALS AND METHODS
TherstbotanicalreferencematerialwascollectedbyTuse-
nius in 1984/1985 and consisted of woody taxa to be used
asacomparativecollectionfortheidenticationofcharcoal
remains from the Deacon excavations. During the present
excavation project, the existing wood reference material has
been expanded to a comprehensive collection of representa-
tive modern specimens of herbs, shrubs, trees, ferns, grasses
and geophytes by van Wijk, Rust, Tusenius, Larbey, Novello
and Cowling (table 1 & electronic appendix), to compile a
reference database as an aid to identifying macro- and micro-
botanicals such as seeds, charcoal, phytoliths, parenchyma
and pollen in the Klasies River archaeological deposits.
Specimens were collected as widely as possible in the im-
mediatecorearea,closetoMainSiteandtoCaves3&4in
collectingareas 1to19 (g.4).Additionalcollectionswere
made further inland within a 5 km radius of Main Site in ar-
eas20to24(g.3).TheseincludedapatchofFynbosanda
relatively intact forest section on the western slope above the
Klasies River itself (collecting areas 20 & 21 respectively
ing.3).Woodspecimenswerealsocollectedinlandabove
Cave 5 by Tusenius in 1984, and Cowling subsequently col-
lectedinthatareain2015(collectingarea22ing.3).
The ideal is to collect at dierent times of the year at
eachsite,and to collect allstagesofthe owering, fruiting
and seeding cycle for most plants, but unfortunately this is
seldompossibleduetotimeconstraintsandthedicultyof
access to many sites. In this study however, four week-long
collectingseasonsatdierenttimesoftheyearbyVanWijk
between 2013 and 2015 made it possible to collect duplicate
specimensinowerorbudforpollensampling,andinseed
or fruit for identication of charred seed remains. Collect-
ing took place in November, December, February, March and
May with unfortunately no collecting in the winter and early
spring. However, due to year-round rainfall and a temperate
climate, owering and fruiting of woody plants appear to
take place all year round – depending more on rainfall than
on the seasons (Y. van Wijk, Rhodes University, South Af-
rica,pers.comm.,seealsoPierce&Cowling1984).Onthe
other hand, many herbaceous plants, especially geophytes,
showed a more seasonal cycle with most phenological activ-
ity in the cooler months (De Vynck et al. 2016b).
The 2015 collection of underground tubers, bulbs, and
roots specically made for research on parenchyma in ar-
chaeological contexts (C. Larbey et al. Cambridge Univer-
sity, UK, unpubl. res.) in which 34 species were located, was
successful only because these geophytes had been seen and
collectedinoweror seedpreviously.Intermsofthewood
collection, an attempt was made to collect as many of the
woody taxa currently growing in the vicinity of the cave sites
as possible. Emphasis was also placed on getting examples
of the same taxa from dierent ecological micro-niches so
that any microscopic anatomical variability due to environ-
mental conditions may be observed. Sampling for the wood
collection is an on-going project.
Leafy voucher specimens for the woody taxa collected
in the 1980s, as well as charred specimens of the associated
wood samples, are lodged at Iziko:SA Museum, Cape Town.
All ethnobotanical and vegetation survey specimens collect-
ed by Van Wijk, Rust and Tusenius from 2013 to 2015 were
pressed to serve as voucher specimens for reference pur-
poses. Most of these specimens were photographed before
pressing and images have been, or will be, submitted to iSpot
for conrmation of identication, and to help assemble an
image database for the area. Additional special samples were
collectedinowerorbudin2013and2014forfuturepollen
research,and inseedforidenticationof charredseedrem-
nants recovered during excavation. These ower and seed
vouchers were sent to Johannesburg to be deposited in the
Evolutionary Studies Institute herbarium at the University
of the Witwatersrand. A complete set of voucher specimens
will be lodged at the Selmar Schonland Herbarium, Graham-
stown, with any duplicates going to the Ria Olivier Herbar-
ium at Nelson Mandela Metropolitan University (NMMU),
Port Elizabeth.
Identications were made mainly by Yvette van Wijk,
with assistance from Johan Baard, Jan Vlok, Richard Cowl-
ing (Ria Olivier Herbarium), Tony Dold (Selmar Schonland
Herbarium),SandraBurrowsandRodneyMoett,withsome
input from iSpot (2016). Classication and authors are ac-
cording to the Angiosperm Phylogeny Group (2016), and the
International Plant Names Index (IPNI 2016). Naming for
allAngiosperms follows Manning & Goldblatt (2012), and
Pteridophytes follow Crouch et al. (2011). To prevent confu-
sion due to many recent name changes, synonyms are given
in the tables for family, genus and species wherever relevant.
RESULTS
A total of 268 species, in 196 genera and 78 families were
collected and are listed in table 1 (see also electronic appen-
dix). The species were collected in 24 numbered areas shown
ings 3&4, andrepresentpresence andabsencedata.Ac-
cessibilityinsomeofthesiteswashamperedbysheerclis,
impenetrable thorny vegetation or dense stands of the alien
invasive tree Acacia cyclops and some areas have not been
collected. The species tally from these areas is therefore un-
likely to be comprehensive. The number of collecting areas
in which each species was collected is indicated in table 1.
The common names of each species including, where possi-
ble, Khoe-San names are given in the table and are according
toBattenetal.(2001),iSpot (2016), Manning & Goldblatt
(2012),Smith(1966),vanWyketal.(1997)andvanWyk&
Gericke (2000). The uses of these taxa as medicinal and/or
edible plants, as well as those with other uses are also indi-
cated(Fox &Norwood-Young 1982,Hutchingsetal.1996,
Smith 1966, van Wyk et al. 1997, Arnold et al. 2002, van
Wyk & Gericke 2000, Pote et al. 2006,Y. van Wijk & R.
Rust, Rhodes University, South Africa, unpubl. res.)
Where the actual species present at Klasies is not referred
to in the references cited, but the genus is however described

18
Pl. Ecol. Evol. 150 (1), 2017
Taxon No of Areas
present Common name Uses Veg type
Acanthaceae FM
Hypoestes aristata(Vahl)Roem.&Schult.
(verticillaris)19 Ribbonbush, Seeroogblom E FTC
Justicia leptantha (Nees) T.Anderson 3 Kiesieblaar GM –
Isoglossa ciliata Lindau 6 Businessman’s plant GM FT
Adiantaceae (Pteridaceae)
Adiantum capillus-veneris L. 1 Maiden Hair fern M –
Agavaceae (Anthericaceae)
Chlorophytum comosum (Thunb.) Jacques 4 Hen&Chickens E FTC
Aizoaceae
Carpobrotus deliciosus (L.Bolus) L.Bolus 10 Sour Fig, Suurvy, Tandsprooi, Brakvy,
Gaukum, Dikgoena M E TCfy
Conicosia pugioniformis (L.) N.E.Brown 1 Snotwortel, Varkslaai E T
Delosperma litorale (Kensit) L.Bolus 4 Kalkklipvygie GM TC
Delosperma saxicola Lavis 1 Rotsvgie GM –
Drosanthemum oribundum (Haw.) Schwantes 5 Douvygie – T
Lampranthus spectabilis (Haw.) N.E.Br. 2 Rankvygie – –
Mesembryanthemum aitonis Jacq. 7 Brakvygie, Brakslaai M E C
Ruschia lineolata Schwantes 10 Muisvygie – –
Tetragonia decumbens Mill. 3 Dune spinach E TC
Tetragonia fruticosa L. 13 Sea spinach GM TC
Amaranthaceae (Chenopodiaceae)
Chenolea diusa Thunb. 8 Seawrack, Soutbossie M TC
*Chenopodium ambrosioides L. 2 Wormseed, Galsiekbos M E –
*Chenopodium murale L. 5 Misbredie, Varklossie, Tjuana-seep M E –
*Chenopodium vulvaria L. 1 Pisbossie GM –
Exomis microphylla (Thunb.) Aellen 5 Hondepisbossie, Rambos M E TC
Pupalia lappacea (L.) Juss. 3 Klitz, Sweethearts M E FT
Sarcocornia littorea (Moss) A.J.Scott 3 Samphire, Lidjiesbos, Ganna-asbos E TC
Amaryllidaceae FM
Haemanthus albios Jacq. 1 Paintbrush, Poeierkwas, Veldskoenblaar M O FTC
Anacardiaceae
Searsia chirindensis(Bakerf.)Moett(Rhus) 1 Wild Currant, Boskaree M E O F
Searsia crenata(Thunb.)Moett(Rhus) 10 Soettaaibos, Korentebos E O FTCfy
Searsia glauca(Thunb.)Moett(Rhus) 6 Blinkblaartaibos, Krintingbessie, E O FTCfy
Searsia nebulosa(Schönland)Moett(Rhus) 1 Sandtaaibos E T
Searsia pyroides(Burch.)Moett(Rhus) 6 Deurmekaartaaibos, Rivierkaree E O T
Apiaceae FM
Apium decumbensEckl.&Zeyh. 6 Sea Celery M C
*Centella asiatica (L.) Urb. 2 Pennywort, Waternavel M E –
Dasispermum suruticosum (P.J.Bergius) B.L.Burtt
(Heteroptilis)2 Sea Parsley E C
Lichtensteinia interrupta E.Mey. 1 Wild Anise, Kalmoes M –
Table 1 – Klasies River species list 2013 to 2015.
Synonyms are italicised in brackets - aliens marked with *. M = medicinal, E = edible, O = other use. GM = Genus reported in the literature
as medicinal, GE = edible, GO = other use. FM = Family reported in the literature as important medicinally. F = forest, T = thicket, C =
coastal, fy = Fynbos.

19
van Wijk et al., Modern vegetation in the Klasies River cultural landscape
Taxon No of Areas
present Common name Uses Veg type
Apiaceae FM
Notobubon ferulaceum (Thunb.) Magee
(Peucedanum)2Lidjiesbos, Wildeseldery, Gatagaai GM T
Notobubon laevigatum (Aiton) Magee
(Peucedanum)3 Lidjiesbos, Bergseldery, Gatagaai M E FT
Apocynaceae FM
Acokanthera oppositifolia (Lam.) Codd 3 Bushman’s poison, Boesmansgif M FTC
Carissa bispinosa (L.) Desf. ex Brenan
(haematocarpa)12 Num-num, Nam-noempies, Noem-Noem M E O FTC
Cynanchum ellipticum (Harv.) R.A.Dyer 4 Dawidjiewortel, Bobbejaantou, Bokhoring,
Nenta M E FTC
Cynanchum natalitium Schltr. 5 Dawidjiewortel, Bobbejaantou, Bokhoring,
Nenta E GM FTC
Secamone alpini Schult. 3 Melktou, Bobbejaantou GM FTC
Aquifoliaceae
Ilex mitis (L.) Radlk. 2 Wild Holly M E F
Araceae FM
Zantedeschia aethiopica (L.) Spreng. 2 Arum Lily, Hottentotsblare M E F
Araliaceae FM
Cussonia thyrsiora Thunb. 4 Spekbos, Nooiensboom E TC
Asparagaceae (Liliaceae) FM
Asparagus aethiopicus L. 1 Wild Asparagus, Katdoring, Katjang M E FTCfy
Asparagus densiorus (Kunth) Jessop (sprengeri) 10 Wild Asparagus, Katdoring, Kattestert M T
Asphodelaceae (Liliaceae, Xanthorrhoeaceae)
Gasteria acinacifolia (J.Jacq.) Haw. 2 Beestong, Ox Tongue, Hottentot Rice,
Bontkouterie E GM FT
Trachyandra divaricata Kunth 2 Veldkool, Wild Cabbage E FC
Aspleniaceae
Asplenium adiantum-nigrum (Kunze) J.P.Roux 2 Maidenhair Fern M –
Asteraceae FM
Arctotis linearis Thunb. 1 Gousblom GM C
Arctotheca prostrata (Salisb.) Britten 5 Cape weed, Gousblom, Skaapoor M –
Arctotheca populifolia (P.J.Bergius) Norl. 1 Seepampoen GM TC
Artemisia afra Jacq. ex Willd. 1 Wildeals M T
Berkheya decurrens (Thunb.) Willd. 3 Grootdisseldoring M T
Cineraria geifolia (L.) L. 13 Geelkransbossie GM –
Cotula sericea L.f. 10 Buttons, Ganskos M TCfy
*Cotula coronopifolia L. 1 Ganskos GM C
Dimorpotheca fruticosa (L.) Less.
(Osteospermum barberae)2 Cape Daisy GM TC
Felicia echinata (Thunb.) Nees 3 Seebloubossie GM TCfy
Gazania rigens var. leucolaena (DC.) Roessler 5 Strandgousblom M C
Gazania rigens var. uniora (L.f.) Roessler 9 Botterblom, Gousblom M TC
Gazania pectinata (Thunb.) Hartweg 2 Gousblom E –
Table 1 (continued) – Klasies River species list 2013 to 2015.
Synonyms are italicised in brackets; aliens marked with *. M = medicinal, E = edible, O = other use. GM = Genus reported in the literature
as medicinal, GE = edible, GO = other use. FM = Family reported in the literature as important medicinally. F = forest, T = thicket, C =
coastal, fy = Fynbos.

20
Pl. Ecol. Evol. 150 (1), 2017
Taxon No of Areas
present Common name Uses Veg type
Asteraceae FM
Gerbera piloselloides (L.) Cass. 2 Swarteebossie, Pitpursies M F
Haplocarpha nervosa (Thunb.) Beauverd 1 Brandbossie M –
Helichrysum albanense Hilliard 1 Everlasting GM T
Helichrysum anomalum Less. 1 Everlasting, Hotnotstee GM Cfy
Helichrysum aureum (Houtt.) Merr. 1 Everlasting M –
Helichrysum cymosum (L.) D.Don 5 Kooigoed, Hotnotskooigoed, Hotnotstee M FTCfy
Helichrysum herbaceum (Andrews) Sweet 1 Everlasting M –
Helichrysum petiolareHilliard&B.L.Burtt 5Kooigoed, Vaalky, Geita Hotnotskooigoed,
Langbeenghyta, M FCfy
Helichrysum rosum (P.J.Bergius) Less 2 Everlasting GM –
Helichrysum teretifolium (L.) D.Don 5 Verpis GM TCfy
Helichrysum umbraculigerum Less. 5 Kerriebos, Kerriekruie, Hotnotstee – –
Metalasia muricata (L.) D.Don 6 Blombos, Witsteekbossie M TCfy
Oncosiphon africanus (P.J.Bergius) Källersjö 1 Wild Chamomile, Kamella GM –
Osteospermum (Chrysanthemoides) moniliferum L. 1 Bitou, Boetabessie, Bietou, Bilhô M E FTCfy
Othonna carnosa Less. 4 – GM T
Plecostachys serpyllifolia
(P.J.Bergius)Hilliard&B.L.Burtt 1 Vaaltee M E T
Senecio angulatus L.f. 1 Cape Ivy M FTC
Senecio elegans L. 5 Strandblommetjie GM C
Senecio oederiifolius DC. 12 Hongerbos, Ragwort GM FT
Senecio othonniorus DC. 1 Sybos M T
Stoebe plumosa (L.) Thunb. 1Slangbossie, Vaalbossie M Cfy
Syncarpha argentea (Thunb.) B.Nord. 1 Everlasting, Sewejaartjies GM C
Syncarpha striata (Thunb.) B.Nord. 2 Everlasting, Sewejaartjies GM –
Tarchonanthus littoralis P.P.J.Herman 6 Wild Camphor, Seesalie, Siriehout,
Swartsalie, Hottentot Tobacco M O FTCfy
Boraginaceae FM
Cordia cara Sond. 1 Ouhout, Septee M E FT
Brassicaceae FM
Heliophila linearis DC. 3 Bloubekkie GM T
Heliophila subulataBurch.&DC. 2 Blompeperbossie M –
Campanulaceae
Prismatocarpus campanuloides (L.) Sond. 1 Bell Flower M –
Wahlenbergia undulata (L.f.) A.DC. 1 African Bluebell M E –
Capparaceae FM
Capparis sepiaria L. 2 Wild caper, Wag-’n-bietjie M FTC
Maerua racemulosa Pax 1 Without, Bush Cherry E FTC
Caryophyllaceae
Silene (primuliora) crassifolia L. 7 DuneCatchy,S’nama M FTCfy
Silene undulata Aiton 4 Wildetabak, Ubulao M E FT
*Stellaria media (L.) Vill. 7 Chickweed M –
Table 1 (continued) – Klasies River species list 2013 to 2015.
Synonyms are italicised in brackets - aliens marked with *. M = medicinal, E = edible, O = other use. GM = Genus reported in the literature
as medicinal, GE = edible, GO = other use. FM = Family reported in the literature as important medicinally. F = forest, T = thicket, C =
coastal, fy = Fynbos.

21
van Wijk et al., Modern vegetation in the Klasies River cultural landscape
Taxon No of Areas
present Common name Uses Veg type
Celastraceae FM
Cassine peragua L. 2 Bastersafraan, Koeboebessie M O FTCfy
Elaeodendron croceum (Thunb.) DC. 2 Saron,Safraan M O FT
Gymnosporia nemorosa(Eckl.&Zeyh.)Szyszył.
(Maytenus)6 Pendoring, Kamnassiehout M FT
Lauridia tetragona (L.f.) R.H.Archer (Cassine) 3 Droëlewer E FTCfy
Maytenus procumbens (L.f.) Loes. 6 Duinekokoboom O TCfy
Mystroxylon aethiopicum (Thunb.) Loes. 4 Cape Cherry, Safraanbas, Koeboehout M E O FTC
Putterlickia pyracantha (L.) Endl. 1 Wolwedoring, – FTC
Pterocelastrus tricuspidatus Walp. 14 Kershout, Cherry Wood M O FTCfy
Robsonodendron maritimum (Bolus) R.H.Archer 1 Duinesybas – FTC
Commelinaceae
Commelina africana L. 2 Yellow Wandering Jew M FT
Crassulaceae FM
Cotyledon orbiculata L. 8 Plakkies, Varkoor, Kouterie M E FTC
Crassula atropurpurea (Haw.) D.Dietr. 1 Persplakkie GM T
Crassula cultrata L. 4 Plakkiebos GM T
Crassula expansa Aiton 1 Strepies GM TC
Crassula nudicaulis L. 1 Skraalplakkie GM TC
Crassula orbicularis L. 2 Klipblom M FT
Crassula ovata (Mill.) Druce 1 Plakkieblaar, Karky, t’karekey M E T
Crassula pellucida subsp. marginalis
(Dryand) Tolkein 4 – GM FTfy
Crassula tetragona L. 1 Karkai M T
Cucurbitaceae FM
Kedrostis nana Cogn. 5 Stinkpatat, Bospatat, Karu M E FT
Zehneria scabra Sond. 2 CapeZehneria M E FTC
Cyperaceae
Carex aethiopica Schkuhr 6 African Tussock Sedge E FC
*Cyperus rotundus L. 4 Nutgrass, Watergras, Uintjie M E –
Ficinia nodosa
(Rottb.)Goetgh.,Muasya&D.A.Simpson 2 Vleibiesie – TC
Ficinia ramosissima Kunth 2 Biesie – Tfy
Ficinia bergiana Kunth 1 Biesie, Rush – –
Isolepis sororia Kunth 1 Sedge – –
Schoenus nigricans L. 1 Black Bog Rush – –
Tetraria involucrata (Rottb.) C.B.Clarke 1 Rush, Biesie – fy
Dennstaedtiaceae (Aspleniaceae)
Pteridium aquilinum subsp. capense
(Thunb.) C.Chr. 1 Bracken Fern M Ffy
Dipsacaceae
Scabiosa incisa Mill. 1 Wild Scabious, Pisgoedbossie M T
Table 1 (continued) – Klasies River species list 2013 to 2015.
Synonyms are italicised in brackets - aliens marked with *. M = medicinal, E = edible, O = other use. GM = Genus reported in the literature
as medicinal, GE = edible, GO = other use. FM = Family reported in the literature as important medicinally. F = forest, T = thicket, C =
coastal, fy = Fynbos.

22
Pl. Ecol. Evol. 150 (1), 2017
Table 1 (continued) – Klasies River species list 2013 to 2015.
Synonyms are italicised in brackets - aliens marked with *. M = medicinal, E = edible, O = other use. GM = Genus reported in the literature
as medicinal, GE = edible, GO = other use. FM = Family reported in the literature as important medicinally. F = forest, T = thicket, C =
coastal, fy = Fynbos.
Taxon No of Areas
present Common name Uses Veg type
Dryopteridaceae
Rumohra adiantiformis (G.Forst.) Ching 1 Seven Week Fern – FCfy
Ebenaceae FM
Diospyros dichrophylla (Gand.) De Winter 7 Star Apple, Koolhout, Tolbos,
Hotnotstolletjie M E O FTCfy
Euclea racemosa L. 5 Kersiebos, See-ghwarrie M E FTCfy
Ericaceae
Erica glandulosa subsp. fourcadei
(L.Bolus)E.G.H.Oliv.&I.M.Oliv. 2 Erica O fy
Erica pectinifolia Salisb. 1 Erica O fy
Erica peltata Andrews 1 Ker-ker, Raasheide O –
Euphorbiaceae FM
Acalypha ecklonii Baill. 3 Katpisbossie GM FT
Acalypha peduncularis E.Mey. ex Meisn.
(incl. A. zeyheri Baill.) 1 Besembos M T
Adenocline acuta (Thunb.) Baill. 5 Spurge M E FT
Euphorbia erythrina Link 2 Pismelkbos GM T
Fabaceae FM
*Acacia cyclops Cunn. ex Don 14 Port Jackson, Rooipitjie, Makboom M O –
Dipogon lignosus (L.) Verdc. 2 Wild pea M E FTCfy
Erythrina cara Thunb. 1Coral Tree M FT
Indigofera porrectaEckl.&Zeyh. 3 Leeuhoutjie, Louhoud M E –
Indigofera tomentosaEckl.&Zeyh. 1 Louhoud GM –
Psoralea repens P.J.Bergius 1 Creeping Psoralea M TC
Rhynchosia caribaea (Jacq.) DC. 2 Vaalertjie M F
Tephrosia capensis (Jacq.) Pers. 3 Rankbossie, Platertjie M Tfy
Tephrosia grandiora (Aiton) Pers. 1 Rooiertjie M T
Trifolium burchellianum Ser. 5 Wild Clover M –
Vigna vexillata (L.) A.Rich. 1Wild Sweetpea, Wilde-ertjie E –
Gentianaceae FM
Chironia peduncularis Lindl. 1 Christmas Berry, Aambeibossie GM C
Geraniaceae FM
Geranium incanum Burm.f. 1 Ou-meid-op-die-werf, Mickie-Jan-Willem M E Tfy
Pelargonium capitatum (L.) L’Hér. 8 Wild Geranium, Malva M TC
Pelargonium grossularioides (L.) L’Hér. 2 Rooirabasam M –
Pelargonium odoratissimum (L.) L’Hér. 1 Scented Geranium, Malva M –
Hyacinthaceae
Albuca virens(Lindl.)J.C.Manning&Goldblatt
(Ornithogalum tenuifolium)6 Slymuintjie, Jikui, Gambry E T
Drimia unioraJ.C.Manning&Goldblatt 5 Fairy Snowdrop GM –
Ornithogalum graminifolium Thunb. 3 Grass Chink GM T
Veltheimia bracteata Harv. ex Baker 1 Sandlelie, Quarobe, Kwarobe GM FT

23
van Wijk et al., Modern vegetation in the Klasies River cultural landscape
Table 1 (continued) – Klasies River species list 2013 to 2015.
Synonyms are italicised in brackets - aliens marked with *. M = medicinal, E = edible, O = other use. GM = Genus reported in the literature
as medicinal, GE = edible, GO = other use. FM = Family reported in the literature as important medicinally. F = forest, T = thicket, C =
coastal, fy = Fynbos.
Taxon No of Areas
present Common name Uses Veg type
Hypoxidaceae FM
Spiloxene trifurcillata (Nel) Fourc. (Pauridia) 2 Sterretjie GM –
Hypoxis cf. stellipilis Ker Gawl. 2 African Potato, Toevanna M O TC
Iridaceae
Aristea ecklonii Eckl. ex Klatt 1 Blue Stars M –
Bobartia orientalis Gillett 1 Geelblombiesie GE Tfy
Chasmanthe aethiopica (L.) N.E.Br. 2 Suurkanol GE FTC
Watsonia knysnana L.Bolus 1 Watsonia GE –
Juncaginaceae
Triglochin elongata Buchenau 1 Bulbous Arrowgrass M E C
Triglochin striataRuiz&Pav. 1Three-rib Arrowgrass GM TC
Lamiaceae FM
Clerodendrum glabrum E.Mey. 1 Tinderwood M FT
Leonotis leonurus (L.) R.Br. 1 Duiwelstwak, Lion’s Ears, Wildedagga M E FT
Salvia africana-lutea L. 3 Strandsalie, Bruinsalie M TCfy
Stachys aethiopica L. 3 Katpisbossie, Kruie M FTCfy
Linaceae
Linum africanum L. 5 African Flax M Cfy
Lobeliaceae (Campanulaceae) FM
Lobelia anceps L.f. 2 Leafy Lobelia M FC
Lobelia cuneifoliaLink&Otto 5 Wedge Leaved Lobelia M F
Lobelia accida (C.Presl) A.DC. 6 Wild Lobelia M –
Monopsis decipiens (Sond.) Thulin 2Wild Violet M –
Malvaceae
Grewia occidentalis L. 7 Raisin bush, Kruisbessie, Vierpuntjies,
Booghout, Pylhout M E O FTC
*Hibiscus trionum L. 1 Wild Hibiscus M –
Meliaceae
Ekebergia capensis Sparrm. 1 Cape Ash, Essenhout M T
Menispermaceae FM
Cissampelos torulosaE.Mey.exHarv.&Sond. 1 Dawidtjieswortel M FT
Moraceae
Ficus burtt-davyi Hutch. 7 Wild Fig, Rankvy, Hottentotstou E O FTC
Myricaceae FM
Morella cordifolia (L.) Killick 1 Waxberry, Gammabos M E O FTCfy
Morella serrata (Lam.) Killick 1 Waterolier, Bergwasbessie M Ffy
Myrsinaceae
Rapanea gilliana (Sond.) Mez 1 Dwarf Cape Beech, Kleinblaarboekenhout, GM TCfy
Rapanea melanophloeos (L.) Mez 5 Cape Beech, Boekenhout M FC
Nymphaeaceae FM
Nymphaea nouchali var. caerulea Burm.f. 1 Blue Water Lily M E –

24
Pl. Ecol. Evol. 150 (1), 2017
Table 1 (continued) – Klasies River species list 2013 to 2015.
Synonyms are italicised in brackets - aliens marked with *. M = medicinal, E = edible, O = other use. GM = Genus reported in the literature
as medicinal, GE = edible, GO = other use. FM = Family reported in the literature as important medicinally. F = forest, T = thicket, C =
coastal, fy = Fynbos.
Taxon No of Areas
present Common name Uses Veg type
Oleaceae
Chionanthus foveolatus (E.Mey.) Stearn (Linociera) 4 Pock Ironwood M FTC
Jasminum angulare Vahl 1 Wild Jasmine M FT
Olea capensis L. 3 Wild Olive, Ironwood M E O FC
Orchidaceae
Bonatea speciosa (L.f.) Willd. 4 Wood Orchid GM FT
Satyrium parviorum Sw. 3 Orchid M –
Satyrium princeps Bolus 1 Orchid GM TCFy
Oxalidaceae
Oxalis smithianaEckl.&Zeyh. 6 Suuring E –
Penaeaceae
Penaea cneorum Meerb. 1 Brickleaf GM Ffy
Pittosporaceae
Pittosporum viridiorum Sims 2 Cheesewood, Kaarsuur M FC
Plantaginaceae FM
Plantago crassifolia Forssk. 5 Weeblaar GM TC
Plumbaginaceae
Limonium scabrum Kuntze 6 Sea Lavender, Brakbossie – TCfy
Poaceae
Cynodon dactylon (L.) Pers. 1 Bermuda Grass M TCfy
Digitaria eriantha Steud. 2 Fingergrass M O T
Ehrharta calycina Sm. 5 Polgras, Bushman Grass – FTCfy
Ehrharta erecta Lam. 1 Panic Veldtgrass – FT
Eragrostis capensis (Thunb.) Trin. 1 Love Grass, Hartjiegras GM FT
Eragrostis curvula (Schrad.) Nees 1 Weeping Lovegrass E T
Helictotrichon hirtulum (Steud.) Schweick. 1 Oat grass – –
Heteropogon contortus
(L.)P.Beauv.exRoem.&Schult. 1 Tanglehead M Tfy
Pentameris pallida
(Thunb.)Galley&H.P.Linder(Pentaschistis)1 Duinegras – T
Polypogon strictus Nees 1 Baardgras – C
Setaria sp. 1 Foxtail Grass GM T
Sporobolus africanus(Poir.)Robyns&Tournay 1 Paramatta Grass M TCfy
Sporobolus virginicus (L.) Kunth 11 Brakkweek GM TC
Stenotaphrum secundatum (Walter) Kuntze 16 Bualo,Strandkweek M E FTCfy
Themeda triandra Forssk. 2 Rooigras M TFy
Tristachya leucothrix Nees (Apochaete hispida) 1 Trident grass – fy
*Vulpia myuros (L.) C.C.Gmel. 1 Wildegars, Wild Barley – T
Polygalaceae FM
Muraltia squarrosa (L.f.) DC. 1 Skilpadbos M Tfy
Polygala ericaefolia DC. 1 Milkwort GM Tfy

25
van Wijk et al., Modern vegetation in the Klasies River cultural landscape
Table 1 (continued) – Klasies River species list 2013 to 2015.
Synonyms are italicised in brackets - aliens marked with *. M = medicinal, E = edible, O = other use. GM = Genus reported in the literature
as medicinal, GE = edible, GO = other use. FM = Family reported in the literature as important medicinally. F = forest, T = thicket, C =
coastal, fy = Fynbos.
Taxon No of Areas
present Common name Uses Veg type
Polygonaceae FM
Rumex sagittatus Thunb. 1 Climbing Sorrel, Ranksuring M E TC
Proteaceae
Leucadendron salignum R.Br. 2 Tolbos, Geelbos M O Cfy
Leucadendron spissifolium I.J.Williams 1 Tolbos – –
Leucospermum cuneiforme (Burm.f.) Rourke 1 Pincushion – fy
Protea coronata Lam. 1 Green Protea GM –
Protea cynaroides (L.) L. 1 King Protea GM fy
Protea neriifolia R.Br. 1 Protea GM fy
Protea tenax R.Br. 1 Ground Protea GM fy
Rhamnaceae
Phylica litoralis(Eckl.&Zeyh.)D.Dietr. 3 Luisbos – TCfy
Phylica purpurea Sond. 4 Luisbos – Ffy
Rhamnus prinoides L’Hér. 2 Blinkblaar, Hondepishout M E FT
Scutia myrtina (Burm.f.) Kurz 6 Katdooring, Rank M E O FTC
Restionaceae
Restio leptoclados Mast. (Ischyrolepis) 4 Besemgoed O fy
Restio triticeus Rottb.1 Besemgoed O TCfy
Thamnochortus fruticosus P.J.Bergius 2 Thatching reed O –
Rosaceae
Cliortia ilicifolia L. 1 Doringtee, Rysbos M fy
Cliortia linearifoliaEckl.&Zeyh. 1 Glastee GM Tfy
Rubus pinnatus Willd. 1 Wild Blackberry, Braambossie, Vaalbraam M E F
Rubiaceae FM
Burchellia bubalina (L.f.) Sims 1 Wild Pomegranate M FTC
Canthium inerme (L.f.) Kuntze 2 Bokdrol, Hardepeer M E FTC
Canthium spinosum
(KlotzschexEckl.&Zeyh.)Kuntze 2 Doringtou M E FTC
Psydrax obovata
(KlotzschexEckl.&Zeyh.)Bridson 3 Psydrax, Kwar, Quar M FT
Rutaceae FM
Agathosma apiculata
E.Mey.exBartl.&H.L.Wendl. 4AnysBuchu,Knoelbuchu,
Hottentotsboegoe M Cfy
Agathosma ovata (Thunb.) Pillans 2 False Buchu, Valsboegoe, Rondeblaar
boegoe M E FTfy
Agathosma serpyllaceaLicht.exRoem.&Schult. 1 Steenbok Buchu, Steenbokboegoe M O –
Clausena anisata (Willd.) Hook.f. ex Benth. 1 Perdepis M FT
Coleonema pulchellum I.Williams 1 Confetti Bush, Aasbossie, Muishondboegoe M T
Zanthoxylum capense (Thunb.) Harv. 5 Perdepram, Wildekardamom M O FTC
Salicaceae (Flacourtiaceae) FM
Scolopia zeyheri(Nees)Szyszył. 4Doringpeer, Wolwedoring M FT

26
Pl. Ecol. Evol. 150 (1), 2017
Table 1 (continued) – Klasies River species list 2013 to 2015.
Synonyms are italicised in brackets - aliens marked with *. M = medicinal, E = edible, O = other use. GM = Genus reported in the literature
as medicinal, GE = edible, GO = other use. FM = Family reported in the literature as important medicinally. F = forest, T = thicket, C =
coastal, fy = Fynbos.
Taxon No of Areas
present Common name Uses Veg type
Santalaceae
Colpoon compressum P.J.Bergius (Osyris) 7 Bloupruim, Basbessie, Looibos,
’Nantegara, Namtarri, Notchou M E O FTC
Thesidium fragile (Thunb.) Sond. 5 Teringbossie GM TCfy
Sapindaceae FM
Allophylus decipiens (E.Mey.) Radlk. 5 Rooibessie M FTC
Sapotaceae
Sideroxylon inerme L. 10 Milkwood M E FTCfy
Scrophulariaceae
Chaenostoma polyanthum Benth. (Sutera) 3 Sutera, Ruikbossie GM T
Chaenostoma cordatum (Thunb.) Benth. (Sutera) 4 Sutera, Ruikbossie GM FT
Jamesbrittenia microphylla (L.f.) Hilliard (Sutera) 1 – GM Tfy
Selago corymbosa L. 1 Slakkiebos, Aambeibos M T
Selago myrtifolia Rchb. 1 Bitterbos GM T
Teedia lucida (Aiton) Rudolphi 1 M FT
Sinopteridaceae (Pteridaceae, Adiantaceae)
Cheilanthes hirta Sw. 1 Kleinvaring, Oorstokkies M F
Solanaceae FM
Lycium ferocissimum Miers 10 Wolvebessie, Slangbessie, Kariedoring M E O T
Solanum africanum Mill. (americanum) 7 Dronkbessie M TC
Solanum linnaeanumHepper&P.-M.L.Jaeger 5 Gifappel, Bitterappel M T
Solanum retroexum Dunal (Solanum nigrum) 1 Sobo-sobo, Nasgal, Wolwebos M E T
Withania somnifera (L.) Dunal 3 Geneesvelletjie, Geneesblaar M T
Stilbaceae (Loganiaceae, Buddlejaceae)
Nuxia oribunda Benth. 11 Wildevlier, Wild Elder M F
Theophrastaceae (Samolaceae)
Samolus porosus Thunb. 2 Water Pimpernel GM C
Samolus valerandi L. 1 Brookweed M –
Thurniaceae (Prioniaceae)
Prionium serratum (L.f.) Drège 1 Palmiet, E –
Thymelaeaceae FM
Passerina corymbosa Eckl. ex C.H.Wright 1 Bakbossie GM Cfy
Passerina ericoides L. 2 Bakbos, Christmas berry, Dronkbessie,
Gonna M O –
Passerina rigida Wikstr. 7 Bakbos, Gonnabas M O FTC
Struthiola hirsuta Wikstr. 1Roemenaggie, Aand gonna – fy
Urticaceae
Didymodoxa cara(Thunb.)Friis&Wilmot-Dear 2 Nettle – FT
Vitaceae
Rhoicissus digitata(L.f.)Gilg&M.Brandt 17 Wild grape, Wildepatat, Boesmansdruif M E O FTC
Rhoicissus tomentosa(Lam.)Wild&R.B.Drumm. 2 Forest grape, Bobbejaantou M E FTC

27
van Wijk et al., Modern vegetation in the Klasies River cultural landscape
Vegetation Type – references
Total No spp.
in table 1 per
reference
No spp.
collected
per veg type
% of
268 spp.
collected
FOREST
Southern Cape Forests Geldenhuys (1993) 93
FOREST
100 20%
Southern Afrotemperate Forest (FOz1)Mucina&Rutherford(2006) 18
Southern Coastal Forest (FOz6)Mucina&Rutherford(2006) 13
Tsitsikamma National Park Hanekom et al. (1989) 23
THICKET
Subtropical thicket, Subtropical Thicket Ecosystem Project (STEP)
Vlok&Euston-Brown(2002) 148
THICKET
163 33%
Humansdorp region, Kararian thicket Cowling (1982) 27
Subtropical Dune Thicket (AZs3)Mucina&Rutherford(2006) 38
Gamtoos Thicket (AT4)Mucina&Rutherford(2006) 39
COASTAL
Eastern & Southern Cape Coasts Lubke&vanWijk(1998a,1998b) 73
COASTAL
109 22%
Tsitsikamma National Park Hanekom et al. (1989) 52
Cape Seashore Vegetation (AZd3)Mucina&Rutherford(2006) 20
Algoa Dune Strandveld (AZs1)Mucina&Rutherford(2006) 26
FYNBOS
Humansdorp region, S Coast dune fynbos Cowling (1982) 19
FYNBOS
73 15%
Southern Cape Dune Fynbos (FFd11)Mucina&Rutherford(2006) 20
Humansdorp region, Grassy fynbos Cowling (1982) 31
Tsitsikamma National Park Hanekom et al. (1989) 31
Absent in referenced literature 55 55 10%
Table 2 – Collected species grouped into vegetation types according to cited references. Some species are listed in more than one
vegetation type.
as a medicinal genus, it is listed in table 1 and in the elec-
tronic appendix as a ‘medicinal genus’ (GM). Similarly, an
edible genus or a genus with other uses is indicated as GE or
GO respectively. There is often substitution of species within
a genus by users depending on what is locally available (Y.
van Wijk, Rhodes University, South Africa, unpubl. thesis)
which validates our use of these categories included here in
order to show more clearly the overall high level of useful
plants present in the Klasies area.
In addition, the vegetation types in which each taxon
occurslocallyis noted(tables1 & 2).The vegetation types
– forest, thicket, coastal vegetation or fynbos – have been
identiedaccordingto specieschecklists from therelevant
literature (Cowling 1982, Hanekom et al. 1989, Geldenhuys
1993, Lubke & van Wijk 1998a, 1998b, Vlok & Euston-
Brown 2002, Mucina & Rutherford 2006). Mucina & Ru-
therford list taxa for each vegetation type but make it clear
thatthese are onlythe“important taxa”(Mucina& Ruther-
ford 2006: 6) and not every species to be found in that spe-
cicvegetationtype.Many of the species collectedatKla-
sies River occur in more than one vegetation type and are
therefore included in each type.
Tables 1 and 2 show that the vegetation we sampled with-
in the wider 5 km radius at Klasies River is comprised of
thicket (163 spp., 33%), forest (100 spp., 20%) and coastal
species (109 spp., 22%). Of the collected species, 49 co-oc-
cur in forest, thicket and coastal vegetation, 78 in forest and
thicket, but only nineteen occur in all four vegetation types.
The 73 species associated with fynbos vegetation account
for only 15%. The 55 species not included in any reference
lists contribute 10%. Thicket and forest types form a mosaic
structure in much of the study area with many thorny taxa
which form dense, impenetrable stands (g. 5A). Thicket
and forest species are particularly dominant in the areas sam-
pled close to the cave sites – collecting areas 3, 8, 8a, 9, 12,
13, 14 and 17 in the vicinity of Main Site, as well as Areas 6,
7and15nearCaves3and4(g.4).Coastalvegetationoc-
curs mainly in areas 2a, 4a, 5, 10, 10a, 11, 16, 18 and 19 and
is very species rich. The coastal species are associated with
thesea-sprayaectedlittoralandhummockdunevegetation.
Most of the fynbos species listed in table 1 were growing in
asmallpatchoffynbosinarea20(g.3),whilesomefynbos
taxawerealsopresentontheplateauinareas14and23(gs
3&4,electronicappendix).
For purposes of comparison, species lists were obtained
from the Integrated Biodiversity Information System (SIBIS)
of the South African National Biodiversity Institute (SAN-
BI, http://biodiversityadvisor.sanbi.org/online-biodiversity-
data/sabif-3/sibis/). SIBIS check lists for four coastal 1:50
000 grids including and surrounding the Klasies River area
–3424AA,AB, BA&BB – were compared with our spe-
cies list. The results of this study show that the Klasies grid
Clarkson 3424AB is signicantly under-represented on SI-

28
Pl. Ecol. Evol. 150 (1), 2017
BIS, with only 69 of the 268 species we collected (table 1)
appearing on the SIBIS database. The lack of data captured
by SIBIS (now Brahms) for the Klasies grid, and the lack
ofothersurveysundertakenorpublishedspecicallyforthis
grid, precludes the use of this data for analysis. The Cape
St Francis 3424BB grid, representing the area to the east of
Klasies, is however better collected, doubtless due to years
of collecting by Richard Cowling and the local Fourcade
Botanical Club (http://stfranciskrommetrust.co.za/outreach-
fourcade/). Of the 922 species in the SIBIS list for 3424BB,
137 species occur in our Klasies species list; at only 15% of
the SIBIS total, this is still minimal, and again does not allow
for useful data analysis.
A large proportion of the shrubs and trees listed in table 1,
particularly in forest and thicket vegetation types, have me-
dicinal properties and/or bear edible fruits. Table 3 shows that
67% of the 268 species collected by us and listed in table
1, were reported as useful during our ongoing ethnobotani-
calsurveyand in the literaturecitedpreviously(in the rst
paragraph of this section). In addition, 23% of the species
are listed in table 1 as GM (medicinal genera in the literature
cited) which brings the overall useful tally to 90%, while only
10% had no reported uses. Table 4 presents a breakdown of
the use categories – medicinal (M), edible (E) or other uses
(O). Many taxa are reported in more than one use category.
Of the total reported uses for the 179 individual species, 57%
were medicinal, 30% edible and 13% had other uses.
DISCUSSION
In this review and analysis of the vegetation we provide a
new detailed description and characterisation of the Kla-
sies River landscape. Our work extends the current knowl-
edge of vegetation in this area signicantly although there
is still much research and systematic plant collecting to be
done before the composition of its vegetation is understood
more fully. We have shown that the Klasies River landscape,
which forms a part of the SIBIS 3424 AB map vegetation
record, is under-represented and our collection of a restricted
area has added 168 species to this record. The fact that the
Klasies area has not been well collected previously, and that
the coastal strip is privately owned with restricted entrance
bypermissiononly,as wellasthe dicultyincollectingin
impenetrablesectionsandon the steep clis, could explain
why 55 species or 10% of the species we collected are not
present in any of the lists referenced in table 2. Some ex-
otic species recorded such as Chenopodium spp., Withania
somnifera, Solanum nigrum / retroexum, Stellaria media,
Cyperus rotundus and Centella asiatica are not listed in the
literature researched, possibly because they are considered,
perhaps erroneously, as relatively recent and weedy intro-
ductions to the area. It is thought-provoking that most of
these weedy species occur frequently at archaeological sites
throughout the southern Cape (Y. van Wijk, Rhodes Univer-
sity, South Africa, pers. obs.) and all are widely used today.
The collection of a range of modern botanical specimens
has also enabled us to address the perception that the Klasies
sitesarelocatedinfynbosvegetation.Mucina&Rutherford
(2006), for example, characterise the Klasies River area as
Southern Cape Dune Fynbos on the Algoa sediments and
Tsitsikamma Sandstone Fynbos on the plateau. This might
have led to the perception that fynbos comprises “96.83%
of the major habitat types represented in a 10 km radius
around Klasies River” (Marean et al. 2014: 170). However,
as shown in tables 1 & 2, the vegetation in the immediate
vicinity of the sites is a complex mosaic consisting of mainly
thicket, forest and coastal vegetation types, with some fyn-
bos elements.
The great diversity of vegetation types – a feature typical
ofthesouth-eastern Cape(Cowling1984,Cowling &Potts
2015) – is due to a variety of factors which have resulted in
the extremely varied terrain providing a wide range of micro-
habitats and micro-climates often associated with distinc-
tive vegetation types or even biomes. These factors include
theunderlyinggeologyandassociatedsoils,re,aspectand
salt-laden wind exposure. Where the sea winds and salt spray
blows unhindered, the vegetation is no more than 10 to 20
cm tall yet remains species rich and could be called ‘Lillipu-
tian’ (Acocks 1988) coastal thicket-forest. Where the valleys
or slopes are protected from the prevailing winds by clis
and ridges, the thicket-forest comes into its own and trees as
high as 3 to 4 metres are present, often within just a few me-
tres of dwarfed examples of the same species. For example,
isolated dwarfed Sideroxylon inerme (milkwood) trees only
20 cm in height occur in area 2 whereas dwarf milkwood
forest 20 cm high, intertwined with thorny Lycium ferocissi-
# individual species reported as being useful
(ie: medicinal, edible, other, see below)
# species in genera
referenced as useful # nul uses Total species listed
in table 1
179 62 27 268
67% 23% 10% 100%
Table 3 – Quantication of useful species for the 268 species listed in table 1.
Medicinal Edible Other Total reported uses
for 179 spp.
148 77 36 261
57% 30% 13% 100%
Table 4 – Uses allocated to the 179 individual useful species in table 3.
Many species have more than one use category.

29
van Wijk et al., Modern vegetation in the Klasies River cultural landscape
mum, is found higher up the same south facing slope in area
3. This abruptly becomes 1–3 m impenetrable milkwood for-
estasthegroundsuddenlyfallsawaytothenorth(g.5A).
Afrotemperate coastal forest 3–6 m tall survives in shel-
tered areas below Caves 3 and 4 (g. 5B), below Cave 5,
and above area 18. On low lying sections along the coast,
such as areas 10, 10a, and sections of 18, the vegetation con-
sists of salt-tolerant grass, low herbs, geophytes and shrubs,
many of which are succulent. Large tracts of closely packed
Trachyandra divaricata (Veldkool) in this grassy seashore
vegetation (g. 5C) promise good pickings of owerbuds
for a nutritious green bredie (stew) in Spring (van Wyk &
Gericke 2000, and Y. Van Wijk, Rhodes University, South
Africa, pers. comm.). A variety of cli dwelling succulent
plant species populate the cracks and ledges of the weath-
eredanderodedquarziticsandstoneoftheclisaboveareas
8,9 and11(g.5D).Thiscli vegetationincludes treeand
thicket species as well as geophytes such as the rare Saty-
rium princeps and succulents such as Gasteria acinacifolia.
The richness and complexity of the vegetation is not ad-
equately shown by the presence and absence data in table 1.
While fynbos is regarded as more species rich than thicket
andforestvegetationtypes,itisconnedtosmallpatcheson
the inland plateau in the broader study area. The coastal veg-
etation is however very species rich. An example of this is, to
some extent, illustrated by listing the surprisingly numerous
species collected in one measured square metre of what ap-
pearedatrstglancetobesimplylowgrassygroundcover.
This square metre in area 8 contained 21 dierent species,
includingtrees,herbs,creepersandgeophytes(seeg.6).
The soil underlying this small plot is nutrient-rich, spongy
with humus, and dark black-brown in colour. This topsoil, a
fertile loam, covers the fossil dune(s) and is also evident in
areas4,8,9,12, 13 and 14 (g. 4).Thisdarkbrownearth
is the result of millennia of thicket growth, die back, peren-
nial leaf shedding, and capture of wind-blown sands and soil
from the coast and inland (Tinley 1985, Y. van Wijk, Rhodes
University, South Africa, pers. comm.). The 80–90 cm depth
Figure 5 – A, impenetrable vegetation in upper area 3: Lycium ferocissimum, Sideroxylon inerme and Hypoestes aristata; B, forest patch
inarea15 andprotectivefore-duneinarea16,locatedbetweenandbelowcaves3and4;C,area10&10a,largepatchesofTrachyandra
divaricata(veldkool,wildcabbage)intheforeground;D,uppersectionofarea11.Clivegetation.PhotographsbyY.vanWijk.

30
Pl. Ecol. Evol. 150 (1), 2017
of the dark humus-rich loamy soil covering the fossil dune
directlyabovetheKlasiesRivermainsite,(g.7)isanindi-
cation of the ancient origin of the dense thicket community
in this area. Although climate oscillations during the Pleisto-
cenewouldhave aectedthedistribution ofthicket,thicket
extent in the Klasies region has probably changed little since
the end of the Late Pleistocene (Cowling et al. 2005, Potts
et al. 2013). In contrast to the loams, the plateau soils are
mainly infertile, poorly drained, duplex forms (sand overly-
ingclayat0.5–1mdepth)(Strydom&Schafer1997).
Inthecontextof the total SouthAfrican ora of nearly
24,000 species, only 3,000 or 12.5%, have been reported as
medicinal (van Wyk & Gericke 2000, Arnold et al. 2002).
The 57% of medicinal plants out of a total of 268 species
collectednear the sitesatKlasies (tables1& 4)indicatesa
much higher proportion of medicinal species than would be
expected, and the 43% of species which are edible or have
other uses is also impressive. Edible species and those with
otheruseshavenotbeenquantiedforSouthAfricaandthe
dicultyofdoing so isformidable.Ofthe 78 familiescol-
lected (table 1), all except three are reported as useful, while
33 of these families have been shown through chemical and
pharmaceutical research to be particularly active medicinally
(Hutchingsetal.1996,vanWyk&Gericke2000,Zhuetal.
2011).
Of the 163 thicket species present, 56 species (34%) are
reported as edible, lending support to De Vynck et al. (2016a)
who demonstrate that thicket taxa comprise the bulk of ed-
ible species in Cape south coast landscapes. Geophytes or
underground storage organs (USOs), are plentiful and varied
▲Figure 6 – One square metre with 21 species present. Adenocline
acuta, Senecio elegans, Senecio angulatus, Cotula coronipifolia,
Carpobrotus deliciosus, Crassula pellucida subsp. marginalis,
Rumex saggitatus, Hypoestes aristata, Trachyandra divaricata,
Ornithogalum graminifolium, Indigofera porrecta, Cynanchum
natalicum, Putterlickia pyracantha, Ficus burtt-dayvi, Acokanthera
oppositifolia, Cineraria geifolia, Centella asiatica, Stenotaphrum
secundatum, Ehrharta calycina, Polypogon strictus, Sporobolus
virginicus. (Compound Voucher YvWijk 6382).
►Figure 7 – A vertical cutting through the fossil dune in area 13.
Showing depth of loam supporting dense thicket vegetation.

31
van Wijk et al., Modern vegetation in the Klasies River cultural landscape
(table 1), but are well hidden in dense thicket and ground-
covers when not in ower.Year-round collectionof speci-
mens and intimate knowledge of the area is necessary to en-
suretheyareadequatelyquantied. Itisinterestingthattwo
fresh-water plant species, Prionium serratum and Nymphaea
nouchali var. caerulea, were collected within the wider 5 km
radius. Although neither are listed in the vegetation literature
cited, both are important because they could provide carbo-
hydrates in larger quantities year-round than the seasonal
geophytes (Fox & Norwood-Young 1982, Wrangham et al
2009, De Vynck 2014).
The presence of so many useful taxa puts the richness of
the Klasies River area in perspective. The large number of
medicinal taxa and those with edible fruit, leaves, bark and
USOs available, plus the presence of highly nutritious sea-
weeds (Anderson et al. 1989, Buchholz et al. 2012) and the
very important availability of much needed salt (Brigand
&Weller2015),would have meant that coastal areas were
sought-after, resource rich areas because of their abundant
and useful plant wealth, as well as readily available seafood
and faunal prey.
The botanical data collected by us during the present
study, makes it possible to compare the vegetation at Klasies
with that at 74 other archaeological sites in the southern and
south-eastern Cape which form part of a larger study about
the connection through time between humans and the sur-
rounding vegetation at habitation sites, being undertaken by
Y. van Wijk (Rhodes University, South Africa, unpubl. res.).
This connection begs further investigation of the nature of
the areas associated with archaeological sites, for example,
about how ‘pristine’ they actually are. Concepts such as
the synergistic co-evolution of plants and humans, and the
anthropogenic eects on vegetation pattern can be seen as
distinct possibilities. The results of vegetation sampling at
Klasies River show considerable overlap with the vegetation
at other important southern Cape coastal sites, for example,
Nelson’s Bay, Pinnacle Point and Blombos. Klasies River
has however a much higher species diversity than other sites.
Our work clearly indicates the need for thorough and sys-
tematiccollecting atarchaeologicallysignicantsitesinthe
Cape region. This is particularly important considering the
many archaeological sites that occur in the area, and the need
for and increasing interest in comparative material for further
archaeobotanical research. The collections provide data for
theidenticationofandcomprehensiveresearchonthe ar-
chaeobotanical remains in terms of past vegetation and plant
use, and will provide further environmental proxies for the
interpretation and contextualisation of human behaviour in
the past. It will also contribute to the broader debate about the
context in which the development of anatomically modern
humans took place.
SUPPLEMENTARY DATA
SupplementarydataareavailableinExcelleatPlant Ecol-
ogy and Evolution, Supplementary Data Site (http://www.
ingentaconnect.com/content/botbel/plecevo/supp-data) and
consists of all plant species collected in 1984/1985 and from
2013 to 2015 within 5 km of the Klasies River archaeologi-
cal sites in 24 areas. Vegetation types, use categories, and
voucher numbers are included.
ACKNOWLEDGEMENTS
We would like to thank the following individuals and in-
stitutions: Gideon F. Smith and Hugh Glen are thanked for
their supportive and helpful comments; The Struwig Ger-
meshuysen Trust for permission to work within the Klasies
River landscape; Kobus (Bolla) Burger and family for as-
sistance and support; Iziko:SA Museum for access to stored
Klasies River botanical material; John Almond for helping
M.L Tusenius with the collecting and sawing of many of the
wood specimens for charcoal research; The local inhabitants
of Klasies River and Clarkson area for help in general and for
information regarding plant uses past and present. The work
of S. Wurz, and Y. van Wijk (in part) is based on research
supported by the National Research Foundation. Any opin-
ion,ndingandconclusionorrecommendationexpressedin
this material is that of the authors and the National Research
Foundation does not accept any liability in this regard.
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