Modern vegetation at the Klasies River archaeological sites, Tsitsikamma coast, south-eastern Cape, South Africa: a reference collection

sarah  wurz

The Klasies River caves are situated on the Tsitsikamma coast about 45km West of Cape St Francis. Klasies River main site figures prominently in modern human origins research. Since 2013 a group of archaeologists from South Africa and overseas, led by Sarah Wurz, have worked at the site each year on a wide range of research projects, some of which revolve around finding evidence for past plant use. Myself and the other authors on this paper have collected as comprehensive a selection of botanical specimens as possible given the difficulty of the terrain and dense thorny thicket, to provide a reference collection of modern botanical species growing at present in close proximity to the Klasies River caves. A total of 268 species, in 195 genera and 77 families have been collected. The vegetation immediately surrounding the Klasies River sites is a complex mosaic of predominantly subtropical thicket, coastal dune thicket, Tsitsikamma or South Coast Afrotemperate forest, and littoral vegetation along the coast and cliffs, with fynbos dominating on the inland plateau. At the same time Renee Rust and I have documented ethnobotanical information and stories with the help of local people who know and use local plants for medicine and food. This project is part of a larger research project I (Yvette) have undertaken in the wider southern Cape and western eastern Cape for a PhD through Rhodes University, Grahamstown. The Klasies River is a key site in early modern human origins research, but the area has not previously been closely studied regarding either the vegetation nor ethnobotanically.

Klasies River is a prominent site among a number of sites along the Cape south coast. Our research forms part of the Klasies Landscape Project headed by Sarah Wurz. The unique convergence of natural resources and the relatively narrow continental shelf at Klasies allowed for a unique and continuous occupation spanning the Middle and Late Pleistocene into the Holocene. Research into consumption of seafood has been dominant, but here we consider the under-researched, mutually beneficial evolution between humans and the plants they used. We consider the way local indigenous people interact with their landscape and value their plant knowledge and folklore today. We used “walk in the veld”, discussion, and verbatim recordings, all of which were indicative of their astuteness and storytelling skills. The collection of modern vegetation samples in tandem with local ethnobotanical research, is essential for any understanding of past plant use and their role in the development of modern cognition. Our results make a strong case for the wide use of rich plant resources around the Klasies in the past. Climate and vegetation change have generally been the main drivers of archaeobotanical research to date. Climate change, however, does not tell the whole story - ethnobotany and archaeobotany also act as important windows into the past of mutually beneficial and evolutionarily adaptive human-plant interactions. This survey clearly indicates the need for thorough and systematic collecting at archaeologically significant sites to provide further environmental proxies for the interpretation and contextualisation of the development of anatomically modern human behaviour. (250) KEY WORDS - KLASIES RIVER PROJECT: INDIGENOUS KNOWLEDGE: FOLKLORE: VEGETATION SURVEY: ARCHAEOBOTANY: LANDSCAPE HERITAGE

The importance of South Africa lies in the many caves and rock shelters containing well preserved evidence of human activity, cultural material complexity and a growing number of early modern human fossils dating to the Middle Stone Age (MSA). South Africa also hosts the world's smallest floral kingdom, now called the Greater Cape Floristic Region (GCFR), with high species richness and endemism. We use phytoliths –amorphous silica particles that formed in epidermal cells of plants–in order to study the evolution of plant exploitation strategies by first modern humans and to understand the response of GCFR environments to glacial-interglacial cycles and rainfall shifts and its implication with the evolution of first modern humans that inhabited the south coast of South Africa during the Upper Pleistocene. In paleoanthropological research, improving our capacity to reconstruct past climatic and environmental conditions can help us to shed light on survival strategies of hunter- gatherers. To do this, one must use actualistic studies of modern assemblages from extant habitats to develop analogies for the past and improve paleoenvironmental reconstructions. Accordingly, this thesis contemplates fossil and modern material: i) archaeological sediment samples from Pinnacle Point 5-6 site (PP5-6) located on the south coast of South Africa; ii) modern plants from the GCFR and susceptible to have been exploited by first modern humans inhabiting the south coast; iii) modern surface soil samples from different GCFR vegetation types of the south coast of South Africa. The modern plant reference collection is the first quantitative and morphological study conducted with wood and leaves of trees and shrubs, leaves, bulb scale leaves and edible part of the bulb of geophytes, restios–Restionaceae and grasses–Poaceae from the GCFR on the south coast of South Africa. We observed that grasses are the highest phytolith producers among plant types. We reported through thin sections and SEM that the characteristic restio phytoliths belong to and characterize the Restionaceae family and have been detected in the parenchyma sheath of the culms. Geophytes did not produce diagnostic phytolith morphotypes that can be used for taxonomic identification what might make difficult their identification in the fossil record. The results of the modern surface soil samples showed that phytolith concentration relates mostly to vegetation types and the dominant vegetation rather than to the type of soils. More abundant phytoliths from Restionaceae and woody/shrubby vegetation are also noted from fynbos vegetation and grass phytoliths are a recurrent component in all the vegetation types in spite of being a minor component in the modern vegetation. The grass silica short cells (GSSCs) from these plants, however, suggest a mix of C3 and C4 grasses in most of the vegetation types with a major presence of the rondels ascribed to C3 grasses. The exceptions are riparian, coastal thicket and coastal forest vegetation, which are characterized by the dominance of C4 grass phytoliths. The study of the modern plants and soils from the surrounding areas of Pinnacle Point were used as proxy for the reconstruction of past human foraging strategies and paleoenvironmental reconstruction through the phytolith record from PP5-6 site. The study of the archaeological sediment samples from this site indicated a wide range of plants used by first modern humans inhabiting the area including wood, leaves and fruits of trees and shrubs, grasses and restios. We reported for the first time the presence of restios in the South African archaeological record through the study of phytoliths. From an environmental perspective, the changes observed in the phytolith record from Pinnacle Point deposits are indicative of vegetation movements accordingly to climate changes and sea level fluctuations, in a continuous regional mosaic of habitats.

Plant Ecology and Evolution 150 (1): 13–34, 2017 https://doi.org/10.5091/plecevo.2017.1286 REGULAR PAPER 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 1 Botany Department, Rhodes University, Grahamstown, 6140, South Africa School of Geography, Archaeology and Environmental Studies, University of the Witwatersrand, Johannesburg, 2050, South Africa 3 School of Geography, Archaeology and Environmental Studies, University of the Witwatersrand, Johannesburg, 2050, South Africa 4 Botany Department, Nelson Mandela University, Port Elizabeth, South Africa 5 Centre for Coastal Palaeosciences, Nelson Mandela Metropolitan University, Port Elizabeth 6031, South Africa 6 School of Geography, Archaeology and Environmental Studies, University of the Witwatersrand, Johannesburg, 2050, South Africa *Author for correspondence: vanwijkyvette@gmail.com 2 Background and aims – The Klasies River cultural landscape, on the Tsitsikamma coast, southeastern 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 microbotanicals such as seeds, charcoal, phytoliths, parenchyma and pollen in the Klasies River archaeological deposits. This is an essential step in providing context for the identiication 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 diferent times of year so that all stages of the lowering, 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 signiicant 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 classiied 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, southeastern Cape, thicket, forest, coastal vegetation, fynbos, medicinal plants, edible plants. INTRODUCTION The Klasies River cultural landscape is situated on the Tsi tsikamma coast, southeastern Cape, South Africa, about 90 km west of Port Elizabeth (ig. 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 bufer zones into the intertidal zone and coastal platform towards the south and north respectively. There are ive major archaeological features on this landscape: Main Site (igs 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: 20323913 (print) – 20323921 (online) Pl. Ecol. Evol. 150 (1), 2017 and Cave 5 have revealed MSA and LSA occupation of the sites from the irst part of the Late Pleistocene to the Late Holocene. Acheulean bifaces and MSA artefacts have been noted in the delation hollows of the fossilized Geelhout boom dune on the plateau above the cave sites (ig. 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 palaeoenvironmental as Figure 1 – Locality of the Klasies River archaeological sites, Tsitsikamma coast, South Africa. Adapted from Deacon & Geleijnse (1988). pects of these Stone Age occupations, for example, the anal yses of the large mammal fauna, microfauna, shellish 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 ongoing 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 irst seeds (Zwane, Witwatersrand University, Johannes burg, SA, unpubl.res.), and parenchyma (C. Larbey et al., Cambridge University, UK, unpubl. res.) have been identi ied. 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, lower 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 aboveground 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 Figure 2 – Panorama of Main Site, situated in the complex Klasies River coastal landscape. 14 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 forestscrub and forestgrassveldmacchia 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 (ig. 3). Periods of lower sea levels in the past, for ex ample, between 100000 and 55000 BP, would have had sig niicant 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 clifs, 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 sealand 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 efect 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 efect of the on shore winds bringing mist and seaspray, 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 westsouthwest in win ter, but in summer the frequency and strength of eastsouth 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 speciesbased 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), 15 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 ire, 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 diicult 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 ire-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 lora. Conspicu ous species include Sideroxylon inerme, Ekebergia capensis, Olea capensis subsp. capensis, Canthium inerme, Pteroce- lastrus tricuspidatus, Gymnosporia nemorosa, Cassine peragua, 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 saltladen 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 washaway 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 speciesrich windsheared vegetation 4a Similar to 4 but including wind-sheared ‘Lilliputian’ forest & thicket species 5 steeply sloping area between shore & clif 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 clifs 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 lats between clifs 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 seashore – 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 & clif on west bank of Klasies River - clif & estuarine vegetation. 16 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 welldrained pla teau areas suggest dominance by overstorey proteoid shrubs with a lower stratum of ericoid shrubs, restiods and grasses. Geophytes are conspicuous after ire. 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 irst 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 identiication 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 (ig. 4). Additional collections were made further inland within a 5 km radius of Main Site in ar eas 20 to 24 (ig. 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 ig. 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 ig. 3). The ideal is to collect at diferent times of the year at each site, and to collect all stages of the lowering, fruiting and seeding cycle for most plants, but unfortunately this is seldom possible due to time constraints and the diiculty of access to many sites. In this study however, four weeklong collecting seasons at diferent times of the year by Van Wijk between 2013 and 2015 made it possible to collect duplicate specimens in lower or bud for pollen sampling, and in seed or fruit for identiication 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 yearround rainfall and a temperate climate, lowering 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 speciically 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 lower 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 diferent ecological micro-niches so that any microscopic anatomical variability due to environ mental conditions may be observed. Sampling for the wood collection is an ongoing 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 conirmation of identiication, and to help assemble an image database for the area. Additional special samples were collected in lower or bud in 2013 and 2014 for future pollen research, and in seed for identiication of charred seed rem nants recovered during excavation. These lower 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. Identiications 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 Mofett, with some input from iSpot (2016). Classiication 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 igs 3 & 4, and represent presence and absence data. Ac cessibility in some of the sites was hampered by sheer clifs, 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, KhoeSan 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 17 Pl. Ecol. Evol. 150 (1), 2017 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. Taxon No of Areas present Common name Uses Veg type Acanthaceae Hypoestes aristata (Vahl) Roem. & Schult. (verticillaris) Justicia leptantha (Nees) T.Anderson FM 19 3 Kiesieblaar GM – Isoglossa ciliata Lindau 6 Businessman’s plant GM FT 1 Maiden Hair fern M – 4 Hen & Chickens E FTC ME TCfy E T Ribbonbush, Seeroogblom E FTC Adiantaceae (Pteridaceae) Adiantum capillus-veneris L. Agavaceae (Anthericaceae) Chlorophytum comosum (Thunb.) Jacques Aizoaceae Carpobrotus deliciosus (L.Bolus) L.Bolus 10 Conicosia pugioniformis (L.) N.E.Brown 1 Sour Fig, Suurvy, Tandsprooi, Brakvy, Gaukum, Dikgoena Snotwortel, Varkslaai Delosperma litorale (Kensit) L.Bolus 4 Kalkklipvygie GM TC Delosperma saxicola Lavis 1 Rotsvgie GM – Drosanthemum loribundum (Haw.) Schwantes 5 Douvygie – T Lampranthus spectabilis (Haw.) N.E.Br. 2 Rankvygie – – Mesembryanthemum aitonis Jacq. 7 Brakvygie, Brakslaai ME 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 difusa Thunb. 8 Seawrack, Soutbossie M TC *Chenopodium ambrosioides L. 2 Wormseed, Galsiekbos ME – *Chenopodium murale L. 5 Misbredie, Varklossie, Tjuanaseep ME – *Chenopodium vulvaria L. 1 Pisbossie GM – Exomis microphylla (Thunb.) Aellen 5 Hondepisbossie, Rambos ME TC Pupalia lappacea (L.) Juss. 3 Klitz, Sweethearts ME FT Sarcocornia littorea (Moss) A.J.Scott 3 Samphire, Lidjiesbos, Gannaasbos E TC FM Amaryllidaceae Haemanthus albilos Jacq. 1 Paintbrush, Poeierkwas, Veldskoenblaar MO FTC Searsia chirindensis (Baker f.) Mofett (Rhus) 1 Wild Currant, Boskaree MEO F Searsia crenata (Thunb.) Mofett (Rhus) Searsia glauca (Thunb.) Mofett (Rhus) 10 Soettaaibos, Korentebos EO FTCfy 6 Blinkblaartaibos, Krintingbessie, EO FTCfy Searsia nebulosa (Schönland) Mofett (Rhus) 1 Sandtaaibos E T Searsia pyroides (Burch.) Mofett (Rhus) 6 Deurmekaartaaibos, Rivierkaree EO T Apium decumbens Eckl. & Zeyh. 6 Sea Celery *Centella asiatica (L.) Urb. Dasispermum sufruticosum (P.J.Bergius) B.L.Burtt (Heteroptilis) Lichtensteinia interrupta E.Mey. 2 Pennywort, Waternavel 2 1 Anacardiaceae Apiaceae 18 FM M C ME – Sea Parsley E C Wild Anise, Kalmoes M – 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 Apiaceae Notobubon ferulaceum (Thunb.) Magee (Peucedanum) Notobubon laevigatum (Aiton) Magee (Peucedanum) Apocynaceae Acokanthera oppositifolia (Lam.) Codd Carissa bispinosa (L.) Desf. ex Brenan (haematocarpa) No of Areas present Common name Uses Veg type FM 2 Lidjiesbos, Wildeseldery, Gatagaai GM T 3 Lidjiesbos, Bergseldery, Gatagaai ME FT FM 3 Bushman’s poison, Boesmansgif 12 Numnum, Namnoempies, NoemNoem Cynanchum ellipticum (Harv.) R.A.Dyer 4 Cynanchum natalitium Schltr. 5 Secamone alpini Schult. 3 Dawidjiewortel, Bobbejaantou, Bokhoring, Nenta Dawidjiewortel, Bobbejaantou, Bokhoring, Nenta Melktou, Bobbejaantou 2 Wild Holly M FTC MEO FTC ME FTC E GM FTC GM FTC ME F Aquifoliaceae Ilex mitis (L.) Radlk. Araceae Zantedeschia aethiopica (L.) Spreng. FM 2 Arum Lily, Hottentotsblare Araliaceae Cussonia thyrsilora Thunb. ME F FM 4 Spekbos, Nooiensboom Asparagaceae (Liliaceae) E TC FM Asparagus aethiopicus L. 1 Wild Asparagus, Katdoring, Katjang Asparagus densilorus (Kunth) Jessop (sprengeri) 10 Wild Asparagus, Katdoring, Kattestert ME FTCfy M T E GM FT E FC M – Asphodelaceae (Liliaceae, Xanthorrhoeaceae) Gasteria acinacifolia (J.Jacq.) Haw. 2 Trachyandra divaricata Kunth 2 Beestong, Ox Tongue, Hottentot Rice, Bontkouterie Veldkool, Wild Cabbage 2 Maidenhair Fern Arctotis linearis Thunb. 1 Gousblom Arctotheca prostrata (Salisb.) Britten 5 Cape weed, Gousblom, Skaapoor Arctotheca populifolia (P.J.Bergius) Norl. 1 Seepampoen Aspleniaceae Asplenium adiantum-nigrum (Kunze) J.P.Roux Asteraceae FM GM C M – 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. Dimorpotheca fruticosa (L.) Less. (Osteospermum barberae) Felicia echinata (Thunb.) Nees 1 Ganskos GM C 2 Cape Daisy GM TC 3 Seebloubossie GM TCfy Gazania rigens var. leucolaena (DC.) Roessler 5 Strandgousblom M C Gazania rigens var. unilora (L.f.) Roessler 9 Botterblom, Gousblom M TC Gazania pectinata (Thunb.) Hartweg 2 Gousblom E – 19 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 Asteraceae Uses Veg type 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 M – Helichrysum petiolare Hilliard & B.L.Burtt 5 M FCfy Helichrysum rosum (P.J.Bergius) Less 2 Everlasting Kooigoed, Vaalky, Geita Hotnotskooigoed, Langbeenghyta, 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 ME FTCfy Othonna carnosa Less. Plecostachys serpyllifolia (P.J.Bergius) Hilliard & B.L.Burtt Senecio angulatus L.f. 4 – GM T 1 Vaaltee ME T 1 Cape Ivy M FTC Senecio elegans L. 5 Strandblommetjie GM C Senecio oederiifolius DC. 12 Hongerbos, Ragwort GM FT Senecio othonnilorus DC. 1 Sybos M T Stoebe plumosa (L.) Thunb. 1 Slangbossie, Vaalbossie M Cfy Syncarpha argentea (Thunb.) B.Nord. 1 Everlasting, Sewejaartjies GM C Syncarpha striata (Thunb.) B.Nord. 2 GM – Tarchonanthus littoralis P.P.J.Herman 6 Everlasting, Sewejaartjies Wild Camphor, Seesalie, Siriehout, Swartsalie, Hottentot Tobacco MO FTCfy Boraginaceae Cordia cafra Sond. FM 1 Ouhout, Septee Brassicaceae ME FT FM Heliophila linearis DC. 3 Bloubekkie GM T Heliophila subulata Burch. & DC. 2 Blompeperbossie M – Prismatocarpus campanuloides (L.) Sond. 1 Bell Flower Wahlenbergia undulata (L.f.) A.DC. 1 African Bluebell M – ME – Capparis sepiaria L. 2 Wild caper, Wag’nbietjie M FTC Maerua racemulosa Pax 1 Without, Bush Cherry E FTC Campanulaceae Capparaceae FM Caryophyllaceae Silene (primulilora) crassifolia L. 7 Dune Catchly, S’nama Silene undulata Aiton 4 Wildetabak, Ubulao *Stellaria media (L.) Vill. 7 Chickweed 20 M FTCfy ME FT M – 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 Celastraceae Uses Veg type FM Cassine peragua L. 2 Bastersafraan, Koeboebessie MO FTCfy Elaeodendron croceum (Thunb.) DC. Gymnosporia nemorosa (Eckl. & Zeyh.) Szyszył. (Maytenus) Lauridia tetragona (L.f.) R.H.Archer (Cassine) 2 Safron, Safraan MO FT 6 Pendoring, Kamnassiehout M FT 3 Droëlewer E FTCfy Maytenus procumbens (L.f.) Loes. 6 Duinekokoboom O TCfy Mystroxylon aethiopicum (Thunb.) Loes. 4 Cape Cherry, Safraanbas, Koeboehout Putterlickia pyracantha (L.) Endl. 1 Wolwedoring, Pterocelastrus tricuspidatus Walp. 14 Kershout, Cherry Wood Robsonodendron maritimum (Bolus) R.H.Archer 1 Duinesybas 2 Yellow Wandering Jew MEO FTC – FTC MO FTCfy – FTC M FT Commelinaceae Commelina africana L. Crassulaceae FM Cotyledon orbiculata L. 8 Plakkies, Varkoor, Kouterie ME FTC Crassula atropurpurea (Haw.) D.Dietr. 1 Persplakkie GM T Crassula cultrata L. 4 Plakkiebos GM T Crassula expansa Aiton 1 Strepies GM TC GM TC M FT Crassula nudicaulis L. 1 Skraalplakkie Crassula orbicularis L. 2 Klipblom Crassula ovata (Mill.) Druce Crassula pellucida subsp. marginalis (Dryand) Tolkein Crassula tetragona L. 1 Plakkieblaar, Karky, t’karekey ME T 4 – GM FTfy 1 Karkai M T Kedrostis nana Cogn. 5 Stinkpatat, Bospatat, Karu ME FT Zehneria scabra Sond. 2 Cape Zehneria ME FTC Carex aethiopica Schkuhr 6 African Tussock Sedge *Cyperus rotundus L. Ficinia nodosa (Rottb.) Goetgh., Muasya & D.A.Simpson Ficinia ramosissima Kunth 4 Nutgrass, Watergras, Uintjie 2 Vleibiesie 2 Ficinia bergiana Kunth 1 Cucurbitaceae FM Cyperaceae E FC ME – – TC Biesie – Tfy 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. Dipsacaceae 1 Bracken Fern M Ffy Scabiosa incisa Mill. 1 Wild Scabious, Pisgoedbossie M T 21 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 – FCfy Dryopteridaceae Rumohra adiantiformis (G.Forst.) Ching 1 Seven Week Fern Ebenaceae FM Diospyros dichrophylla (Gand.) De Winter 7 Euclea racemosa L. 5 Star Apple, Koolhout, Tolbos, Hotnotstolletjie Kersiebos, Seeghwarrie Ericaceae Erica glandulosa subsp. fourcadei (L.Bolus) E.G.H.Oliv. & I.M.Oliv. Erica pectinifolia Salisb. 2 Erica peltata Andrews 3 Katpisbossie MEO FTCfy ME FTCfy Erica O fy 1 Erica O fy 1 Kerker, Raasheide O – Euphorbiaceae FM Acalypha ecklonii Baill. Acalypha peduncularis E.Mey. ex Meisn. (incl. A. zeyheri Baill.) Adenocline acuta (Thunb.) Baill. GM FT 1 Besembos M T 5 Euphorbia erythrina Link 2 Spurge ME FT Pismelkbos GM T *Acacia cyclops Cunn. ex Don 14 Port Jackson, Rooipitjie, Makboom MO – ME FTCfy M FT Fabaceae FM Dipogon lignosus (L.) Verdc. 2 Wild pea Erythrina cafra Thunb. 1 Coral Tree Indigofera porrecta Eckl. & Zeyh. 3 Leeuhoutjie, Louhoud ME – 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 grandilora (Aiton) Pers. 1 Rooiertjie M T Trifolium burchellianum Ser. 5 Wild Clover M – Vigna vexillata (L.) A.Rich. 1 Wild Sweetpea, Wildeertjie E – Gentianaceae Chironia peduncularis Lindl. FM 1 Christmas Berry, Aambeibossie Geraniaceae GM C FM Geranium incanum Burm.f. 1 Oumeidopdiewerf, MickieJanWillem Pelargonium capitatum (L.) L’Hér. 8 Wild Geranium, Malva ME Tfy 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) Drimia unilora J.C.Manning & Goldblatt 6 Slymuintjie, Jikui, Gambry E T 5 Fairy Snowdrop GM – Ornithogalum graminifolium Thunb. 3 Grass Chink GM T Veltheimia bracteata Harv. ex Baker 1 Sandlelie, Quarobe, Kwarobe GM FT 22 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 Hypoxidaceae Uses Veg type FM Spiloxene trifurcillata (Nel) Fourc. (Pauridia) 2 Sterretjie GM – Hypoxis cf. stellipilis Ker Gawl. 2 African Potato, Toevanna MO 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 – Triglochin elongata Buchenau 1 Bulbous Arrowgrass ME C Triglochin striata Ruiz & Pav. 1 Threerib Arrowgrass GM TC Juncaginaceae Lamiaceae FM Clerodendrum glabrum E.Mey. 1 Tinderwood Leonotis leonurus (L.) R.Br. 1 Duiwelstwak, Lion’s Ears, Wildedagga M FT ME FT Salvia africana-lutea L. 3 Strandsalie, Bruinsalie M TCfy Stachys aethiopica L. 3 Katpisbossie, Kruie M FTCfy 5 African Flax M Cfy Linaceae Linum africanum L. Lobeliaceae (Campanulaceae) FM Lobelia anceps L.f. 2 Leafy Lobelia M FC Lobelia cuneifolia Link & Otto 5 Wedge Leaved Lobelia M F Lobelia laccida (C.Presl) A.DC. 6 Wild Lobelia M – Monopsis decipiens (Sond.) Thulin 2 Wild Violet M – MEO FTC M – M T Malvaceae Grewia occidentalis L. 7 *Hibiscus trionum L. 1 Raisin bush, Kruisbessie, Vierpuntjies, Booghout, Pylhout Wild Hibiscus 1 Cape Ash, Essenhout 1 Dawidtjieswortel 7 Wild Fig, Rankvy, Hottentotstou Meliaceae Ekebergia capensis Sparrm. Menispermaceae Cissampelos torulosa E.Mey. ex Harv. & Sond. FM M FT EO FTC Moraceae Ficus burtt-davyi Hutch. Myricaceae FM Morella cordifolia (L.) Killick 1 Waxberry, Gammabos Morella serrata (Lam.) Killick 1 Waterolier, Bergwasbessie Rapanea gilliana (Sond.) Mez 1 Dwarf Cape Beech, Kleinblaarboekenhout, Rapanea melanophloeos (L.) Mez 5 Cape Beech, Boekenhout MEO FTCfy M Ffy GM TCfy M FC Myrsinaceae Nymphaeaceae Nymphaea nouchali var. caerulea Burm.f. FM 1 Blue Water Lily ME – 23 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 M FTC Oleaceae Chionanthus foveolatus (E.Mey.) Stearn (Linociera) 4 Pock Ironwood Jasminum angulare Vahl 1 Wild Jasmine Olea capensis L. 3 Wild Olive, Ironwood M FT MEO FC 4 Wood Orchid GM FT Satyrium parvilorum Sw. 3 Orchid M – Satyrium princeps Bolus 1 Orchid GM TCFy 6 Suuring E – 1 Brickleaf GM Ffy 2 Cheesewood, Kaarsuur M FC Orchidaceae Bonatea speciosa (L.f.) Willd. Oxalidaceae Oxalis smithiana Eckl. & Zeyh. Penaeaceae Penaea cneorum Meerb. Pittosporaceae Pittosporum viridilorum Sims Plantaginaceae Plantago crassifolia Forssk. FM 5 Weeblaar 6 Sea Lavender, Brakbossie GM TC – TCfy Plumbaginaceae Limonium scabrum Kuntze Poaceae Cynodon dactylon (L.) Pers. 1 Bermuda Grass Digitaria eriantha Steud. 2 Fingergrass M TCfy MO T Ehrharta calycina Sm. 5 Ehrharta erecta Lam. 1 Polgras, Bushman Grass – FTCfy 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. Heteropogon contortus (L.) P.Beauv. ex Roem. & Schult. Pentameris pallida (Thunb.) Galley & H.P.Linder (Pentaschistis) Polypogon strictus Nees 1 Oat grass – – 1 Tanglehead M Tfy 1 Duinegras – T 1 Baardgras Setaria sp. 1 Foxtail Grass Sporobolus africanus (Poir.) Robyns & Tournay 1 Paramatta Grass – C GM T M TCfy Sporobolus virginicus (L.) Kunth 11 Brakkweek GM TC Stenotaphrum secundatum (Walter) Kuntze 16 Bufalo, Strandkweek ME 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 Polygala ericaefolia DC. 1 Milkwort 24 M Tfy GM Tfy 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 Polygonaceae Rumex sagittatus Thunb. Uses Veg type FM 1 Climbing Sorrel, Ranksuring ME TC Leucadendron salignum R.Br. 2 Tolbos, Geelbos MO Cfy Leucadendron spissifolium I.J.Williams 1 Tolbos – – Leucospermum cuneiforme (Burm.f.) Rourke 1 Pincushion – fy Proteaceae 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 3 Luisbos – TCfy Rhamnaceae Phylica litoralis (Eckl. & Zeyh.) D.Dietr. Phylica purpurea Sond. 4 Luisbos Rhamnus prinoides L’Hér. 2 Blinkblaar, Hondepishout – Ffy ME FT Scutia myrtina (Burm.f.) Kurz 6 Katdooring, Rank MEO FTC Restio leptoclados Mast. (Ischyrolepis) 4 Besemgoed O fy Restio triticeus Rottb. Thamnochortus fruticosus P.J.Bergius 1 Besemgoed O TCfy 2 Thatching reed O – 1 Doringtee, Rysbos M fy Restionaceae Rosaceae Clifortia ilicifolia L. Clifortia linearifolia Eckl. & Zeyh. 1 Glastee GM Tfy Rubus pinnatus Willd. 1 Wild Blackberry, Braambossie, Vaalbraam ME F Burchellia bubalina (L.f.) Sims 1 Wild Pomegranate M FTC Canthium inerme (L.f.) Kuntze Canthium spinosum (Klotzsch ex Eckl. & Zeyh.) Kuntze Psydrax obovata (Klotzsch ex Eckl. & Zeyh.) Bridson Rutaceae Agathosma apiculata E.Mey. ex Bartl. & H.L.Wendl. 2 Bokdrol, Hardepeer ME FTC 2 Doringtou ME FTC 3 Psydrax, Kwar, Quar M FT Rubiaceae FM FM Agathosma ovata (Thunb.) Pillans 2 Agathosma serpyllacea Licht. ex Roem. & Schult. 1 Anys Buchu, Knofelbuchu, Hottentotsboegoe False Buchu, Valsboegoe, Rondeblaar boegoe Steenbok Buchu, Steenbokboegoe Clausena anisata (Willd.) Hook.f. ex Benth. 1 Perdepis 4 Coleonema pulchellum I.Williams 1 Confetti Bush, Aasbossie, Muishondboegoe Zanthoxylum capense (Thunb.) Harv. 5 Perdepram, Wildekardamom Salicaceae (Flacourtiaceae) Scolopia zeyheri (Nees) Szyszył. M Cfy ME FTfy MO – M FT M T MO FTC FM 4 Doringpeer, Wolwedoring M FT 25 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 MEO FTC GM TCfy Santalaceae Colpoon compressum P.J.Bergius (Osyris) 7 Thesidium fragile (Thunb.) Sond. 5 Bloupruim, Basbessie, Looibos, ’Nantegara, Namtarri, Notchou Teringbossie Sapindaceae Allophylus decipiens (E.Mey.) Radlk. FM 5 Rooibessie M FTC 10 Milkwood ME FTCfy Sapotaceae Sideroxylon inerme L. 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 Selago myrtifolia Rchb. 1 Bitterbos Teedia lucida (Aiton) Rudolphi 1 M T GM T M FT M F Sinopteridaceae (Pteridaceae, Adiantaceae) Cheilanthes hirta Sw. 1 Kleinvaring, Oorstokkies Solanaceae FM Lycium ferocissimum Miers 10 Wolvebessie, Slangbessie, Kariedoring Solanum africanum Mill. (americanum) 7 Dronkbessie Solanum linnaeanum Hepper & P.-M.L.Jaeger 5 Gifappel, Bitterappel Solanum retrolexum Dunal (Solanum nigrum) 1 Sobosobo, Nasgal, Wolwebos Withania somnifera (L.) Dunal 3 MEO T M TC M T ME T Geneesvelletjie, Geneesblaar M T 11 Wildevlier, Wild Elder M F Samolus porosus Thunb. 2 Water Pimpernel GM C Samolus valerandi L. 1 Brookweed M – 1 Palmiet, E – Stilbaceae (Loganiaceae, Buddlejaceae) Nuxia loribunda Benth. Theophrastaceae (Samolaceae) Thurniaceae (Prioniaceae) Prionium serratum (L.f.) Drège Thymelaeaceae FM Passerina corymbosa Eckl. ex C.H.Wright 1 Passerina ericoides L. 2 Passerina rigida Wikstr. 7 Bakbossie Bakbos, Christmas berry, Dronkbessie, Gonna Bakbos, Gonnabas GM Cfy MO – MO FTC Struthiola hirsuta Wikstr. 1 Roemenaggie, Aand gonna – fy 2 Nettle – FT Rhoicissus digitata (L.f.) Gilg & M.Brandt 17 Wild grape, Wildepatat, Boesmansdruif MEO FTC Rhoicissus tomentosa (Lam.) Wild & R.B.Drumm. 2 Forest grape, Bobbejaantou ME FTC Urticaceae Didymodoxa cafra (Thunb.) Friis & Wilmot-Dear Vitaceae 26 van Wijk et al., Modern vegetation in the Klasies River cultural landscape Table 2 – Collected species grouped into vegetation types according to cited references. Some species are listed in more than one vegetation type. Total No spp. No spp. in table 1 per collected reference per veg type Vegetation Type – references FOREST THICKET COASTAL FYNBOS Southern Cape Forests Geldenhuys (1993) 93 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 Subtropical thicket, Subtropical Thicket Ecosystem Project (STEP) Vlok & Euston-Brown (2002) 148 Humansdorp region, Kafrarian thicket Cowling (1982) 27 Subtropical Dune Thicket (AZs 3) Mucina & Rutherford (2006) 38 Gamtoos Thicket (AT 4) Mucina & Rutherford (2006) 39 Eastern & Southern Cape Coasts Lubke & van Wijk (1998a, 1998b) 73 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 Humansdorp region, S Coast dune fynbos Cowling (1982) 19 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 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 identiied according to species check lists from the relevant literature (Cowling 1982, Hanekom et al. 1989, Geldenhuys 1993, Lubke & van Wijk 1998a, 1998b, Vlok & EustonBrown 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 ciic 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 cooc % of 268 spp. collected FOREST 100 20% THICKET 163 33% COASTAL 109 22% FYNBOS 73 15% 55 10% 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 (ig. 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 (ig. 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 afected 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 (ig. 3), while some fynbos taxa were also present on the plateau in areas 14 and 23 (igs 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/onlinebiodiversity data/sabif3/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 signiicantly under-represented on SI 27 Pl. Ecol. Evol. 150 (1), 2017 Table 3 – Quantiication of useful species for the 268 species listed in table 1. # 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 4 – Uses allocated to the 179 individual useful species in table 3. Many species have more than one use category. Medicinal Edible Other Total reported uses for 179 spp. 148 77 36 261 57% 30% 13% 100% 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 speciically 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 irst 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 signiicantly 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 underrepresented 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 diiculty in collecting in 28 impenetrable sections and on the steep clifs, 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 / retrolexum, 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 thoughtprovoking 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 microclimates often associated with distinc tive vegetation types or even biomes. These factors include the underlying geology and associated soils, ire, aspect and saltladen 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 thicketforest. Where the valleys or slopes are protected from the prevailing winds by clifs and ridges, the thicketforest 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- 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 (ig. 5A). Afrotemperate coastal forest 3–6 m tall survives in shel tered areas below Caves 3 and 4 (ig. 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 salttolerant grass, low herbs, geophytes and shrubs, many of which are succulent. Large tracts of closely packed Trachyandra divaricata (Veldkool) in this grassy seashore vegetation (ig. 5C) promise good pickings of lowerbuds 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 clif dwelling succulent plant species populate the cracks and ledges of the weath ered and eroded quarzitic sandstone of the clifs above areas 8, 9 and 11 (ig. 5D). This clif vegetation includes tree and thicket species as well as geophytes such as the rare Satyrium 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 conined 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 irst glance to be simply low grassy ground cover. This square metre in area 8 contained 21 diferent species, including trees, herbs, creepers and geophytes (see ig. 6). The soil underlying this small plot is nutrientrich, spongy with humus, and dark blackbrown 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 (ig. 4). This dark brown earth is the result of millennia of thicket growth, die back, peren nial leaf shedding, and capture of windblown 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. Clif vegetation. Photographs by Y. van Wijk. 29 Pl. Ecol. Evol. 150 (1), 2017 of the dark humusrich loamy soil covering the fossil dune directly above the Klasies River main site, (ig. 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 afected 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 lora 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 quantiied for South Africa and the diiculty 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. 30 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 lower. Year-round collection of speci mens and intimate knowledge of the area is necessary to en sure they are adequately quantiied. It is interesting that two freshwater 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 yearround 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 soughtafter, 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 southeastern 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 coevolution of plants and humans, and the anthropogenic efects 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 signiicant 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 identiication 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 ile at Plant Ecology and Evolution, Supplementary Data Site (http://www. ingentaconnect.com/content/botbel/plecevo/suppdata) 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, inding 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. REFERENCES Acocks J.P.H. (1988) Veld types of South Africa. 3rd Ed. Memoirs of the Botanical Survey of South Africa 57: 1–146. 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Communicating Editor: Elmar Robbrecht.

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Modern vegetation at the Klasies River archaeological sites, Tsitsikamma coast, south-eastern Cape, South Africa: a reference collection

Modern vegetation at the Klasies River archaeological sites, Tsitsikamma coast, south-eastern Cape, South Africa: a reference collection

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