Identifying Asteraceae, particularly Tarchonanthus parvicapitulatus, in archaeological charcoal from the Middle Stone Age

Quaternary International xxx (2017) 1e17

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Identifying Asteraceae, particularly Tarchonanthus parvicapitulatus, in archaeological charcoal from the Middle Stone Age Sandra J. Lennox*, Marion K. Bamford Evolutionary Studies Institute and School of Geosciences, University of the Witwatersrand, P. Bag 3, Wits, 2050, Johannesburg, South Africa

a r t i c l e i n f o

a b s t r a c t

Article history: Received 15 June 2016 Received in revised form 15 March 2017 Accepted 31 March 2017 Available online xxx

Sibudu rockshelter, an archaeological site in KwaZulu-Natal, has evidence of the local vegetation, environment and wood use during the Middle Stone Age, from well-preserved seeds and charcoal, approximately 77e38 000 years ago. In order to confidently identify some charcoal taxa, closely related species were studied in detail. Modern wood was charred and examined under the light microscope and a combination of anatomical features was used to distinguish the various taxa. Tarchonanthus parvicapitulatus P.P.J. Herman (syn. in part Tarchonanthus camphoratus L.) is an evergreen, woodland shrub or tree, which is tolerant of hot, dry, salty or icy conditions. Essential oils from the leaves have antimicrobial and insecticidal properties. The camphor smoke is used in traditional African medicine, the aromatic leaves are used in organic camp bedding and the hard, heavy wood is insect resistant. Since the wood anatomy of this shrub is very similar to Brachylaena discolor DC, another woody member of the Asteraceae, the modern reference charcoal has been studied, to distinguish between these and other species. The confirmed presence of aromatic T. parvicapitulatus in hearths probably implies deliberate burning for insect repellent smoke. © 2017 Elsevier Ltd and INQUA. All rights reserved.

Keywords: Anthracology Brachylaena Camphor smoke Hearths

1. Introduction Interest in Asteraceae was stimulated by their putative identification amongst more than 250 species from charcoal specimens from Sibudu Cave in KwaZulu Natal (KZN), South Africa. Sibudu is valued as an archaeological site, with deposits from approximately 77 to 35 000 years ago (ka) from the Middle Stone Age (MSA) (Wadley et al., 2011). Charcoals from Sibudu, studied by Allott (2005, 2006), include some specimens thought to be from Brachylaena and others from unspecified Asteraceae. Members of the Asteraceae are predominantly herbs, with a few woody members occurring in southern Africa and other parts of the world. In all 1535 Asteraceae genera and 25 000 species are cosmopolitan (Boon, 2010). In southern Africa there are 246 genera and 2305 species and amongst these the few woody Asteraceae consist of 14 genera and 34 species (CoatesePalgrave, 2002). Brachylaena sp. was identified from archaeological charcoal in occupational layers of different ages: Grey Sand (GS), Grey Rocky (GR) 2 and GR from >60ka, Ebony (Eb) from ~60ka and Buff (Bu)

* Corresponding author. E-mail addresses: [email protected], [email protected] (S.J. Lennox), [email protected] (M.K. Bamford).

from ~37ka (Allott, 2006; Wadley and Jacobs, 2006; Jacobs et al., 2008a,b). Some members of the Asteraceae family are reported to have medicinal properties, for example Tarchonanthus camphoratus, camphor bush (Beentje, 1999, 2000; Herman, 2002; Omolo et al., 2004, 2005; Matasyoh et al., 2007; Braithwaite et al., 2008; Abimola, 2010; Nanyonga et al., 2013; Hulley et al., 2016) and this further prompted interest in securely identifying the Sibudu specimens. Although the camphor bush is relatively easy to identify the more recent collections and analyses have shown that there are important differences. The Tarchonanthus camphoratus L. complex in southern Africa includes five species depending on distribution, leaf shape and margin, synflorescences and flowering times: T. minor, T. camphoratus, T. obovatus, T. littoralis and T. parvicapitulatus (Herman, 2002). T. parvicapitulatus (small-head camphor bush) occurs near Sibudu Cave (Herman, 2002; Boon, 2010). Older collections do not necessarily have the updated identifications nor separated the species, so the older specimens are still referred to here as “T. camphoratus” as it is beyond the scope of this work to check all earlier identifications. T. camphoratus (narrowly defined as one of five species in the T. camphoratus complex) has foliage with a grey-green appearance and the leaves are narrowly oblong to elliptic. The inflorescences

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(cypsellas) are enveloped by yellow cotton hairs. On the contrary, in T. parvicapitulatus (syn. T. camphoratus in part, one of five species in the T. camphoratus complex) the leaves are usually oblanceolate, with denticulate margins towards the apex and dark green. The inflorescences are smaller and more open. The leaves of both emit a strong smell of camphor when crushed (Herman, 2002; Van Wyk and Van Wyk, 2013: 124). Since several members of the Asteraceae family could occur in the Sibudu charcoal, it is important to be able to distinguish the taxa confidently, based on their wood anatomy. The identifications of Asteraceae were not possible solely from modern charcoal or wood anatomy descriptions that are already published (Carlquist, 1961, 1966, 2001, 2012; Kromhout, 1975; Tusenius, 1986; Esterhuysen and Mitchell, 1996; Allott, 2005, 2006; Cartwright, 2013; Chikumbirike, 2014) or from more general texts (Metcalfe and Chalk, 1950; Carlquist, 1975, 2010, 2012; Ilic, 1991; Neumann et al., 2000; InsideWood, 2004onwards; Richter and Dallwitz, 2000; Wheeler, 2011). It was therefore necessary to create a modern reference collection of selected charred woods and to create our own data base of anatomical attributes after detailed microscopy. Charcoal must be used for the comparative collection because the available literature on modern woods is based on fresh material. Woody tissues shrink and fragment on charring (Bamford and Henderson, 2003; Gonçalves et al., 2012; Chrzazvez et al., 2014) so the quantitative features of charcoals will differ from those of fresh woods (Hubau et al., 2012, 2013, 2014). However, vessel diameter and frequency can be affected by climatic conditions and the position of the branch or stem within the plant, thus qualitative characteristics are probably more reliable than absolute sizes (Wheeler et al., 1989). The archaeological charcoal in hearths from three occupational layers from approximately 58 ka and 49 ka has been identified to test for the selection of wood for specific tasks by populations at Sibudu. Archaeological charcoal specimens from

occupational layers Brown under Yellow Ash (i) (BYA2(i)), Spotty Camel (SPCA) and Mottled Deposit (MOD) were studied. SPCA has not been dated, but lies between layers with ages of approximately 61 300 and 56 000 years ago. MOD has an age of approximately 49 100 ka (Jacobs et al., 2008a,b). The ages of occupational floors were previously obtained by optically stimulated luminescence (OSL) dating (Jacobs et al., 2008a,b). Th anatomical study of the reference material and archaeological charcoal is summarised in Table 2. The use of poisonous wood, Spirostachys africana, burnt in one hearth on each layer implied that this wood was collected and burnt for the use of noxious smoke (Allott, 2006; Wadley, 2012b, 2013b, 2015; Lennox and Bamford, 2015; Lennox et al., 2015). It has been shown that wood with different burning properties imply wood selection for hearths with different purposes at Sibudu (Wadley, 2012a; Bentsen, 2012, 2013, 2014a,b). Asteraceae woods are used in traditional herbal medicine (Watt and BreyerBrandwijk, 1962; Cunningham, 2001; Van Wyk, 2009, 2013; Van Wyk et al., 2009: 286; Hulley et al., 2016); therefore, the detailed anatomical study of reference material was made to identify them. The identification of Tarchonanthus (with camphor) in the archaeological charcoal would imply wood collected and burnt for the use of aromatic camphor smoke.

2. Geographical setting Sibudu Cave is situated in a cliff, above the uThongathi River 15 km inland of on the east coast of South Africa (Fig. 1). The area receives summer rainfall and summers are hot and humid, winters are mild with some precipitation. Mean annual rainfall is about 900 ml. The bioregion is in the KwaZulu-Natal Coastal Belt (Mucina and Rutherford, 2006). The forest is evergreen with deciduous and semi-deciduous trees at the margins. Savanna patches are nearby. A mosaic of vegetation types occurs near Sibudu today with forest

Table 1 The woody Asteraceae selected for charcoal analysis. Scientific name

Author

Artemisia afra Brachylaena discolor B. elliptica B. huillensis B. rotundata B. transvaalensis

Jacq. ex Willd. DC. (Thunb.) DC. O. Hoffm. S. Moore E. Philips & Schweick B. uniflora Harv. Chrysanthemoides monilifera (L.) T.Norl. Distephanus divaricatus (Steetz) H.Rob. & B. Kahn Euryops spathaceus DC. Gymnanthemum coloratum G. coloratum G. coloratum Lopholaena coriifolia

Metalasia densa Seriphium plumosum Tarchonanthus camphoratus T. littoralis T. parvicapitulatus T. trilobus var. galpinii T. trilobus var. trilobus

Synonym

Vernacular name

#

Reference

e e e

wildeals, wormwood coast Silver-oak bitter leaf silver-oak Lowveld silver-oak mountain silver-oak forest Silver Oak

SJL 90 SJL 56/127 SJL 105 SJL 41 Allott 28 Allott 27

Annandale, Vivo, Limpopo Port Edward, KZN/Ex. Hort., Wits Ex. Hort., Kirstenbosch Botanical Gardens, SANBI Annandale, Vivo, Limpopo Allott, 2005 Allott, 2005

SJL 61 Allott 29 SJL 128/129

B. discolor ssp. transvaalensis e tall silver-oak e bush tick-berry Vernonia aurantiaca

(Willd.) H.Rob. & B.Kahn (Willd.) H.Rob. & B.Kahn (Willd.) H.Rob. & B.Kahn (Sond.) E.Phillips & C.A. Sm.

Vernonia colorata

Lowveld bitter-tea

SJL 123

Port Edward, KZN Allott, 2005 MOSS Herbarium, WITS, A.O.D. Mogg, Inhaca Island, Mocambique MOSS Herbarium, Collector no. 1761, Port Elizabeth, Eastern Cape MOSS Herbarium, R.L. Davies, 09/07/194, Limpopo

V. colorata

Lowveld bitter-tea

SJL 124

MOSS Herbarium, C.E. Moss, J. 14 993, Limpopo

V. colorata

Lowveld bitter-tea

SJL 125

(L.) D. Don (L.) Thunb L. P.P.J.Herman P.P.J.Herman (Hutch. & E.Phillips) Paiva DC.

e Stoebe vulgaris e T. camphoratus T. camphoratus e

MOSS Herbarium, G. Hemm 689, Messina, J. 50 098, Limpopo Small-leaved fluffbush SJL 121/122 MOSS Herbarium: M. McPaty, 21432, Balfour, Tweefonten/A.O.D. Mogg, Krugersdorp, J. 78 491, Gauteng bristle-bush SJL 106 Ex. Hort., Kirstenbosch zig-zag bush SJL 126 Ex. Hort., Grassland, WITS camphor bush SJL 108 Callidendron, Limpopo coast camphor bush Ex. Hort., Kirstenbosch small-head camphor bush SJL 92 Mokopane, Limpopo broad-leaf camphor bush Kromhout 163 Kromhout, 1975

e

trident camphor bush

SJL 107

SJL 130

e

Ex. Hort., Kirstenbosch

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occurring in the shaded southern aspect and savanna on the sunny northern slopes of the hill opposite Sibudu. The Celtis mildbraedi growing at the site suggests a remnant of past vegetation. Mosaic vegetation may have occurred there in the past. Today this vegetation is adversely affected by anthropogenic activities (Wadley, 2004, 2006; Allott, 2006; Lennox et al., 2015). The past climate and vegetation have been studied by means of several environmental proxies, including charcoal and seeds (Allott, 2004, 2005, 2006; Bruch et al., 2012; Hall et al., 2014; Lennox, 2016a; Wadley, 2013a), phytoliths (Schiegl et al., 2004; Murungi in prep) and isotopes (Hall et al., 2014). 3. Material and methods The archaeological Tarchonanthus was identified by means of direct comparisons with modern charcoal anatomy. A selection of woody Asteraceae was made, which have the growth habit of low shrubs or small trees and a distribution in KwaZulu-Natal and which could be used as fuel. These include Artemisia, Brachylaena, Chrysanthemoides, Distephanus, Euryops, Gymnanthemum, Lopholaena, Metalasia, Seriphium and Tarchonanthus (Table 1). Artemisia afra was included as it is widely used as traditional medicine (Liu et al., 2009; Van Wyk, 2009). A reference collection of charcoal charred from modern wood was studied. At least three charred fragments of branch wood from each taxon were examined to accommodate natural variation in the woods. The Brachylaena specimens included Brachylaena discolor, B. elliptica, B. huillensis, B. rotundata, B. transvaalensis and B. uniflora. The Tarchonanthus specimens included T. camphoratus, T. littoralis, T. parvicapitulatus

and T. trilobus. Other carbonised wood and voucher specimens, labelled SJL, are housed in the Palaeobotany Herbarium, Evolutionary Studies Institute (ESI) at the University of the Witwatersrand (Wits). The identifications were checked at the Moss Herbarium, University of the Witwatersrand with the details in Table 1. Comparisons were also made from the charcoal reference collection made by Lucy Allott (Archaeology Department, University of the Witwatersrand), and from the published literature. Although the sample sizes were fairly small the authors are confident that they have accounted for some variation in anatomical features due to environmental factors (Table 2). Wood blocks were carbonised in a LENTON 0861 muffle furnace (Lenton, Hope, UK) for 3.5 h at 350
C at the Evolutionary Studies Institute, University of the Witwatersrand. and were studied by means of stereomicroscopy (Olympus SZX16, Munster, Germany) and reflective light microscopy (Olympus BX51) at magnifications of 100, 200 and 500. The anatomical features according to the International Association of Wood Anatomists (Wheeler et al., 1989) were recorded and digitally photographed using an Olympus DP32 camera and Stream Essentials image analysis software with extended focal image capability. Anatomical features of scanned electron microscope images (SEM) of fresh woods in the Allott (2005) charcoal reference collection at the University of the Witwatersrand, Archaeology Department, from the InsideWood database (2004onwards) and from specialised literature (Carlquist, 1961, 1966, 1975, 2001, 2010; 2012; Kromhout, 1975; Tusenius, 1986) were used for comparative identification (Bamford, 2011, 2015a,b) (see Fig. 1).

Table 2 Summarised charcoal anatomy of a selection of woody Asteraceae. The IAWA features are according to Wheeler et al. (1989). Note that (v) means variation. Scientific name

3

IAWA features

Modern charcoal reference material Artemisia afra 1, 4, 6, 11, 9v, 12v, 13, 22, 25, 31, 40 (~50 mm), 49, 58, 61, 63, 69, 86, 97, 109, 115 Brachylaena discolor 2, 5, 7 (1e3 cells), 40 (~50 mm), 49, 69, 75, 79, 97, 106, 116, 118v, Brachylaena discolor 2, 5, 7 (1e3 cells), 10, 11v, 13, 22, 24, 40 (~50 mm), 49, 52, 61, 66, 70v, 75, 79, 83v, 97 (1e2 cells), 104, 116, 118, 136, 138 Brachylaena elliptica 2, 5, 7, 10, 40, 49, 70, 76, 78, 96, 97v, 106 (body cells procumbent, marginal cells square) Brachylaena huilensis 2, 5, 7, 10, 40 (~50 mm), 50, 70, 76, 78, 96v, 97, 106 (body cells procumbent, marginal cells upright or square), 118v Brachylaena huilensis 2, 5, 7, 10, 40, 41, 50, 70, 76, 86v, 78, 96v, 97, 106 (body cells procumbent, marginal upright or square), 118 Brachylaena rotundata 2, 5, 7 (1e3 cells), 9v, 10, 11v, 13, 22, 40, 49, 52, 66, 70, 71, 76, 79, '98, 97 (1-3-5 cells), 104, 115, 118v (almost) Brachylaena transvaalensis 2, 5, 7 (1e3 cells), 11v, 13, 22, 41, 49, 52, 61, 66, 70, 71, 75, 79, 80v, 83v, 97, 98 (2-3-6 cells), 104, 116, 118 Brachylaena uniflora 2, 5, 7, 9, 41, 49, 69, 70, 75, 78, 97, 106 (body cells procumbent, marginal cells upright or square), 116, 136, 138 Chrysanthemoides monilifera 2, 5, 7 (1e3 cells), 10, 13, 22, 25, 31, 41 (~50 mm), 47, 52, 61, 66, 70, 71, 86, 78, 97, 104, 116, Distephanus divaricatus 2, 5, 7 (1e3 cells), 9, 11v, 13, 22, 24, 30, 40, 49, 53, 61, 63, 66, 68, 71, 76, 78, 96, 97, 105, 116 Distephanus garnierianus 1, 4, 11, 13, 22, 24, 30, 36, 41, 48, 52 (v), (53 (v), 54 (v), 61, 63 (v), 66, 69, 71 (v), 72 (v), 73 (v), 86, 78, 79, 97, 98, 106, 116 Euryops spathaceus 1, 4, 7, 10, 13, 22, 24, 31, 40, 50, 53, 61, 66, 69, 71, 76, 78, 96v, 97, 104, 116 Euryops sp. 1, 4, 9, 13, 22, 24, 30, 40, 49, 52, 61, 66, 70, 71, 75, 97, 98 (3e5 cells), 109, 116 Gymnanthemum coloratum 2, 5, 7 (1e3 cells), 13, 22, 24, 30, 36, 37, 40, 48, 52, 61, 63, 66, 70, 71, 76, 78, 97, 102, 109 (procumbent, upright), 116 Lopholaena coriifolia 2, 5, 7 (1e3 cells), 11v, 13, 22, 24, 30, 40, 49, 53, 61, 63, 66, 69, 71, 76, 78, 97, 102, 109 (procumbent, upright), 116 Metalasia densa 2, 4, 7 (1e3 cells), 13, 22, 24, 30, 40, 49, 53, 61, 63, 66, 68, 71, 76, 78, 97, 109 (upright, square), 116 Seriphium plumosum 2, 4, 11, 13, 22, 24, 30, 40, 41, 48, 52, 61, 66, 71, 76, 78, 96, 97, 104 (upright), 116 Stoebe kilimandscharica 1, 4, 11, 13, 22, 24, 30, 36, 37, 40, 48, 52, 61, 66, 71 (v), 72 (v), 85, 97, 105, 106, 116 Tarchonanthus camphoratus 2, 5, 7 (1e3 cells), 11, 13, 22, 24, 30, 40, 48, 52, 61, 66, 69, 71, 76, 78, 97 (1e2 cells), 109 (procumbent, square), 116 Tarchonanthus littoralis 2, 5, 7 (1e3 cells), 13, 22, 24, 30, 40, 49, 52, 61, 66, 70, 71, 75, 78, 96, 97, 109 (procumbent, square), 116 Tarchonanthus 2, 5, 7 (1e3 cells), 13, 22, 24, 30, 40, 50, 52, 61, 66, 69, 71, 75, 78, 97 (1e2 cells), 109 (procumbent, square), 116 parvicapitulatus Tarchonanthus trilobus 2, 5, 7, 10, 13, 22, 24, 30, 41, 49, 52, 61, 66, 69, 71, 75, 97 (1-2-3 cells), 104, 116, 136, 138, 141 Tarchonanthus trilobus 7, 10, 13, 22, 24, 30, 41, 49, 52, 61, 66, 70, 71, 76, 97 (1-2-3 cells), 104, 116 Archaeological charcoal Brachylaena discolor 2, 5, 7, 10, 13, 22, 24, 30, 40, 49, 52, 61, 66, 70, 71, 75, 78, 79, 97 (1e4 cells), 106 (body cells procumbent, marginal cells square), 116, 118 Brachylaena discolor 2, 5, 7, 10, 13, 22, 24, 30, 40, 48, 52, 61, 66, 70, 71, 76, 78, 97 (1e3 cells), 106 (body cells procumbent, marginal cells upright), 116, 118 Brachylaena discolor 2, 5, 7, 10, 13, 22, 24, 30, 40, 48, 52, 61, 66, 69, 71, 76, 78, 97 (1e4 cells), 106 (body cells procumbent, marginal cells upright), 116, 118 Brachylaena discolor 2, 5, 7, 10, 13, 22, 24, 30, 40, 49, 52, 61, 66, 69, 71, 75, 78, 79, 97 (1e4 cells), 104, 116, 118 Tarchonanthus 2, 5, 7 (1e3 cells), 13, 22, 24, 30, 40, 50, 52, 61, 66, 69, 71, 75, 78, 97 (1e2 cells), 109 (procumbent, square), 116, parvicapitulatus Tarchonanthus 2, 5, 7 (1e3 cells), 13, 22, 24, 30, 40, 50, 52, 61, 66, 68, 71, 75, 78, 97 (1e2 cells), 109 (procumbent, square), 116 parvicapitulatus Tarchonanthus 2, 5, 7 (1e3 cells), 13, 22, 24, 30, 40, 50, 52, 61, 66, 68, 71, 75, 78, 97 (1e2 cells), 109 (procumbent, square), 116 parvicapitulatus

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Fig. 1. Geographic location of the Middle Stone Age site of Sibudu Cave overlooking the uThongathi River, KZN (A) map of the locality, and (B) photograph of the archaeological site.

4. Results and discussion 4.1. Descriptions of the modern reference material 4.1.1. Artemisia afra SJL 90 (Fig. 2) The distinct growth rings are traced by axial parenchyma in narrow bands of less than three cells. Vessels are mostly in clusters (>60%) and occasionally solitary. Some vessels are angular in outline. Vessels are small with a mean tangential diameter of less than 50 mm and are abundant with between 40 and 100 vessels/ mm2. Perforation plates are simple and horizontal. Inter-vessel pits and vessel-ray pits are alternate and small, ~6 mm or between 4 and 7 mm wide. Shiny deposits occur in the vessels. Fibres are septate and the fibre walls are thin - thick with fibre lumina less than three times the double wall thickness and distinctly open (Wheeler et al., 1989). Simple pits occur in both tangential and radial walls. A pith of regular hexagonal cells occurs and pith has a diameter of ~12 mm, which is approximately a half of the stem diameter of ~24 mm. The stem diameter is given in relation to the pith and varies between the woody taxa. This is given for this particular specimen. Rays are triseriate and heterocellular with procumbent, square and upright body cells and with one to two rows of marginal upright cells.

4.1.2. Brachylaena discolor SJL 56, B. elliptica SJL 105, B. huilensis SJL 41, B. rotundata Allott 28, B. transvaalensis Allott 27, B. uniflora SJL 61 (Figs. 3e6) Characteristics common to Brachylaena, including those according to Carlquist (1961) are, the wood is diffuse-porous and the vessels are round and commonly arranged in a diagonal pattern, with a low degree of grouping. Vessel diameters are small,  50 mm, but may be medium 50 mm. Vessels are wider than the fibres, 50/

10 mm in a ratio of ~5:1. Perforation plates are simple and oblique. Intervessel pits and vessel-ray pits are alternate, minute, 2e4 mm and polygonal in shape with distinct borders. There may be grooves alongside the pits (Carlquist, 1961), not seen in this material. Fibres are longer than vessels and radially flattened (Carlquist, 1961), not visible in the charcoal. Fibres are thick walled where the fibre lumina are almost completely closed (Wheeler et al., 1989). Fibres are septate with simple to minutely bordered pits in both tangential and radial walls. Axial parenchyma is variable, for example, paratracheal scanty or vasicentric or apotracheal diffuse. There may be paratracheal confluent or apotracheal banded axial parenchyma. Rays are frequently uniseriate and low, < 200 mm. Rays are frequent,  12 rays/mm. Ray cells are procumbent with some species having square marginal cells. Tyloses are absent. B. discolor SJL 56 (Fig. 3). Vessels are commonly arranged in short radials of 2e4, vessel diameters are small to medium between 50 and 100 mm and abundant between 40 and 100/mm2. Fibre walls are thin e thick. Axial parenchyma is paratracheal vasicentric with 1e2 rows of cells around the vessel. Rays are 1e3 (6) cells wide, 10e20 cells high, heterocellular with procumbent body cells and one row of marginal square cells. Rays are stacked in a pattern which tends towards being storied although these are not as regular as storied rays. The mineral inclusions are prismatic crystals which rarely occur and which fill the ray cells. Inter-vessel and vessel-ray pits are alternate and 2e4 mm in diameter in this and the other species of Brachylaena. B. elliptica SJL 105 (Fig. 6 A, C and E). Vessels are arranged in radial lines of four or more, vessels are small,  50 mm and abundant, 40e100/mm2. Fibres are thick walled. Axial parenchyma is paratracheal scanty and apotracheal diffuse. The rays are (1)-3 cells wide and uniseriate rays are always present. Ray cell arrangement is heterocellular procumbent, with marginal square cells.

Please cite this article in press as: Lennox, S.J., Bamford, M.K., Identifying Asteraceae, particularly Tarchonanthus parvicapitulatus, in archaeological charcoal from the Middle Stone Age, Quaternary International (2017), http://dx.doi.org/10.1016/j.quaint.2017.03.074

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Fig. 2. Photomicrographs of Artemisia afra, SJL 90: (A) TS (transverse section) of growth rings traced by thin parenchyma bands, with central pith approximately half the width of the stem. (B) TS thin-thick walled fibres. (C) TLS (tangential longitudinal section) of triseriate rays. (D) TLS of alternate inter-vessel pits small 6 ~m, perforation plates and parenchyma (p). (E) RLS (radial longitudinal section) of rays with procumbent, square and upright cells. (F) RLS of marginal upright ray cells.

B. huilensis SJL 41 (Fig. 4). Vessels are arranged in radial multiples of four or more, small and dense 100 vessels/mm2. Vessel walls with helical thickenings (Essien et al., 2004onwards) were not visible in this material. Fibre walls are thin to thick, but some have very thick walls as a variation. Axial parenchyma is apotracheal diffuse or paratracheal scanty. Rays are 1e3-seriate low and weakly heterocellular with body cells procumbent and occasionally with one row of upright and/or square marginal cells. A shiny deposit is visible in the vessels. B. rotundata Allott 28 (2005). Vessels are arranged in radial multiples of one to three, with rare occurrences of long radial multiples of more than four; vessels are small, approximately 50 mm and abundant, between 40 and 100/mm2. Fibre walls are thick. Axial parenchyma is mostly apotracheal diffuse, but

occasionally paratracheal vasicentric too. Rays are 1-3-5 cells wide. Biseriate rays almost 1 mm high, as reported by Chikumbirike (2014) were not observed in this material. Ray cells are homocellular with procumbent body and marginal cells. Rays tend to storied. B. transvaalensis Allott 27 (2005). Vessels are arranged in radial multiples of 4 or more cells, small to medium sized, mostly 50 mm, with a range of tangential diameter, 50e100 mm; vessel frequency is 20e40/mm2. Fibre walls are thick. Axial parenchyma is paratracheal, vasicentric, with a variation of confluent parenchyma. Rays are 2-3-6 cells wide, homocellular procumbent and tend to storied. B. uniflora SJL 61 (Fig. 5). Vessels are arranged in short radials of 1e3 but many are solitary, small or medium, 50e100 mm and

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Fig. 3. Photomicrographs of Brachylaena discolor, SJL 56. (A) TS of vessels in short radials of two to four, vessels are small to medium and abundant. (B) TS of fibres with thin-thick walls and narrower than vessels. (C) TLS of triseriate rays which tend to storied, with crystals. (D) TLS of alternate intervessel pits minute ~3 mm, and vasicentric parenchyma. (E) RLS of rays with procumbent cells and one row of marginal square cells, (F) RLS of vessel-ray pits equal in size and shape, to inter-vessel pits.

abundant, 40e100 vessels/mm2. Fibre walls are thin e thick to thick, septate with simple pits in both tangential and radial walls. Axial parenchyma is paratracheal scanty. Rays are 1-2-3 cells wide and heterocellular with procumbent body cells and one row of marginal square cells. Rays tend to storied. Mineral inclusions are present as prismatic crystals occurring rarely in procumbent ray cells, completely filling the cells. There are a few distinguishing characteristics between the very similar Brachylaena species: B. discolor, B. huilensis and B. uniflora rays are narrow, triseriate and they are heterocellular consisting of procumbent body cells, with one row of square marginal cells. Uniseriate rays occur in B. elliptica and B. huilensis, being absent from the others. On the contrary, B. transvaalensis and B. rotundata

rays are wider with up to 5e6 cells and they are homocellular comprising only procumbent cells. 4.1.3. Chrysanthemoides monilifera Allott 29 (2005) Growth rings are distinct and consist of compressed parenchyma and fibre cells. The vessels are arranged in short radial multiples of 1e3 cells and some are in long radial multiples of 4 cells. Vessels are medium in diameter 100 mm and abundant 40/mm2. The perforation plates are simple and the inter-vessel pits are alternate and small ~4 mm. Fibres are thick walled and 1 mm in length. Axial parenchyma is apotracheal banded, with bands  3 cells wide, 2 bands/mm and paratracheal scanty. Rays are homocellular with upright cells and there are 12 rays/mm.

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Fig. 4. Photomicrographs of Brachylaena huilensis, SJL 41: (A) TS of vessels in long radial multiples with scanty parenchyma. (B) TS of very thick fibre walls. (C) TLS of uniseriate and triseriate rays. (D) TLS pf perforation plates, intervessel pits distinctly bordered and septate fibres. (E) RLS of procumbent rays. (F) RLS of vessel-ray pits equal to intervessel pits.

4.1.4. Distephanus divaricatus (syn. Vernonia aurantiaca) SJL 128 and SJL 129 (Fig. 6 B, D and F) Growth rings are distinct. Vessels are most commonly solitary, small 50 mm (40), abundant 40e100/mm2 (68 average). Perforation plates are simple and inter-vessel pits are alternate, polygonal and minute 4 mm. Vessel-ray pits are similar in size and shape to inter-vessel pits. Fibres are thin walled. Axial parenchyma is apotracheal diffuse or paratracheal scanty. Ray width is uniseriate and cell arrangement is upright.

4.1.5. Euryops spathecus SJL 130 Growth rings are distinct. Vessels are commonly arranged in long radial multiples 4, vessels are small <50 mm (10), dense  100/mm2 (120). Axial parenchyma is apotracheal diffuse or paratracheal scanty. Rays are narrow, 1e2 cells wide and high

1 mm. Ray cell arrangement is mixed square and upright. Fibre walls are thick. Inter-vessel pits are minute 4 mm and alternate. Vessel walls with helical thickenings (Tusenius, 1986) were not seen in this material.

4.1.6. Gymnanthemum coloratum (syn. Vernonia colorata) SJL 124 (Fig. 7 A, C and E) Vessels are commonly arranged in short radial lines, paired, small in diameter, < 50 mm (40), and a frequency of 20e40/mm2 (24). Fibre walls are thick. Axial parenchyma is apotracheal diffuse or paratracheal scanty. Rays are narrow, 1e3 cells wide and high 1 mm. Rays are heterocellular with procumbent and upright cells. A pith is present comprising regularly shaped, oval cells. The pith has a diameter of ~6 mm, which is approximately a half of the stem diameter of ~12 mm.

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Fig. 5. Photomicrographs of Brachylaena uniflora, SJL 61: (A) TS of vessels in short radials, occasionally solitary and abundant. The axial parenchyma is scanty. (B) TS of thin-thick walled fibres. (C) TLS of bi-to triseriate rays. (D) TLS of crystals in ray cells and of intervessel-pits. (E) RLS of procumbent ray cells (F) TLS of prismatic crystals and of vessel-ray pits similar in size and shape to inter-vessel pits.

4.1.7. Lopholaena coriifolia SJL 122 (Fig. 7 B, D and F) Vessels are commonly arranged in short radial lines, with the variation of clusters. Vessels are small 50 mm, and abundant, 40e100/mm2. Fibre walls are thin - thick. Axial parenchyma is apotracheal diffuse or scanty paratracheal Rays are narrow, 1e3 cells wide and high  1 mm, and heterocellular with procumbent and upright cells. 4.1.8. Metalasia densa SJL 106 (Fig. 8 A, C and E) Vessels are commonly arranged in short radial lines. Vessels are small <50 mm and abundant 40e100/mm2. Fibre walls are thin. Axial parenchyma is apotracheal diffuse and paratracheal scanty. Rays are narrow 1e2 cells wide and low in height, <1 mm. They are heterocellular with upright and square cells.

4.1.9. Seriphium plumosum (syn. Stoebe vulgaris) SJL 126 (Fig. 8 B, D and F) Growth rings are distinct. Vessels are commonly arranged in short radial multiples with the variation of clusters. Vessels are small 50 mm, frequency 20e40/mm2. Perforation plates are simple. Inter-vessel pits are alternate, polygonal and minute. Vessel-ray pits are similar in size and shape, with distinct borders. Fibre walls are thin. Axial parenchyma is apotracheal diffuse and paratracheal scanty. Rays are 1e3 seriate, homocellular with only procumbent cells. A closely related species, Stoebe kilimandscharica (Metcalfe and Chalk, 1950; InsideWood, 2004onwards), has vessel walls with helical thickenings, axial parenchyma arranged in wide bands,  3 bands/mm, and rays heterocellular, with procumbent body cells and 1-2-4 rows of upright or square marginal cells but these do not

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Fig. 6. Photomicrographs of Brachylaena elliptica SJL 105 (A) TS of vessels in long radial lines, axial parenchyma is scanty, fibres are thick walled. (C) TLS of uniseriate rays. (E) RLS of procumbent ray cells. Photomicrographs of Distephanus divaricata (syn. Vernonia aurantiaca) SJL 128, (B) TS of solitary vessels, scanty parenchyma and thin walled fibres, (D) TLS of uniseriate rays and (F) RLS rays of upright cells.

occur in S. plumosum.

heterocellular, procumbent body and marginal cells with intermittent, rare square cells.

4.1.10. Tarchonanthus camphoratus SJL 108, T. parvicapitulatus SJL 92 and T. trilobus SJL 107 (Figs. 9 and 10) Characteristics common to Tarchonanthus, including those according to Carlquist (1961) are wood which is diffuse-porous, perforation plates are simple and oblique, vessel-ray pits and inter-vessel pits are alternate and minute, ~2 mm, with distinct borders, and rays are low and frequent,  12/mm.

4.1.12. T. littoralis Tusenius 324 (1986) Vessels are commonly arranged in low radial multiples of 1e3 cells. Vessels are small,  50 mm, abundant 40e100/mm2. Fibre walls are thin-thick. Axial parenchyma is paratracheal scanty. Rays are uniseriate, triseriate, heterocellular, procumbent body and marginal cells with intermittent rare, square cells.

4.1.11. T. camphoratus SJL 108 (Fig. 10 A, C and E) Vessels are commonly arranged in low radial multiples, clusters are common. Vessels are small 50 mm and 20e40/mm2. Fibre walls are thin - thick. Parenchyma is apotracheal diffuse and paratracheal scanty. Rays are narrow, 1e2 cells wide, weakly

4.1.13. T. parvicapitulatus SJL 92 (Fig. 9) Vessels are commonly arranged in low radial lines of 1-3-5 cells, small, ~29 mm wide, round and dense, 100 vessels/mm2. Fibres are septate with thin to thick walls. Axial parenchyma is paratracheal scanty. Rays are narrow, uni-to biseriate. They are weakly

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Fig. 7. Photomicrographs of Gymnanthemum coloratum (syn. Vernonia colorata) SJL 123 (A) TS of vessels in short radial lines, (C) TLS of narrow, high rays and (E) RLS rays of procumbent and upright cells. Phytomicrographs of Lopholaena coriifolia SJL 121 (B) TS of vessels in short radial lines, (D) TLS of narrow, high rays and (F) RLS rays of procumbent and upright cells.

heterocellular, with long and short procumbent cells and rare, square cells which are infrequently distributed within each ray. Shiny deposits, possibly gum, are present in the vessels and also fill the central pith cells. The vessel to fibre diameter ratio is low in T. parvicapitulatus with 30/20 mm or 1.5:1. 4.1.14. T. trilobus SJL 107 (Fig. 9 B, D and F) Vessels are commonly arranged in long radial multiples of 4 cells. Vessels are medium, 50e100 mm wide and abundant, 40e100/mm2. Fibre walls are thin-thick. Parenchyma is apotracheal diffuse. Rays are 1-2-3 cells wide, homocellular, procumbent. Prismatic crystals are plentiful and fill procumbent ray cells. Prismatic crystals are rare in non-chambered axial parenchyma cells. There are distinguishing characteristics between the very similar Tarchonanthus species. T. parvicapitulatus SJL 92 rays are

heterocellular consisting of procumbent body cells and rare intermittent square cells, whereas T. trilobus (Kromhout, 1975) rays are homocellular consisting of procumbent cells. Prismatic crystals are present in T. trilobus but absent from T. camphoratus, T. littoralis and T. parvicapitulatus. Vessels are commonly arranged in long radial multiples 4 in T. trilobus and in short radial multiples in T. camphoratus, T. parvicapitulatus and T. littoralis. 4.2. Characteristics useful for distinguishing between the genera Characteristics of the woody Asteraceae according to Carlquist (1961) and Metcalfe and Chalk (1950) were observed in this material. Vessels are commonly arranged in radial lines, occasionally in clusters. Vessels are small, 50 mm to medium, 50e100 mm and abundant, 40e100/mm2. Perforation plates are simple, horizontal

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Fig. 8. Photomicrographs of Metalasia densa SJL 106 (A) TS of vessels in short radial lines, (C) TLS of narrow, biseriate, low rays and (E) RLS rays of upright and square cells. Phytomicrographs of Seriphium plumosum (syn. Stoebe vulgaris) SJL 126, (B) TS of vessels in short, radial multiples and clusters, (D) TLS of triseriate rays and (F) RLS rays of homocellular procumbent cells.

or oblique. Fibre pits are small and simple. Axial parenchyma is sparse, for example scanty or diffuse. Inter-vessel pits are alternate and minute. Vessel-ray pits are equal in size and shape. Rays are generally narrow in these genera (not wide as mentioned in the literature), occasionally uniseriate and occasionally storied. Rays are composed of heterocellular, irregular, mixed procumbent and upright cells. Crystals are rare and there are various types including prismatic. The ten genera and 19 species may be separated into two groups on the grounds of growth rings being present in five genera, Artemisia, Chrysanthemoides, Distephanus, Euryops and Seriphium and absent in five genera, Brachylaena, Gymnanthemum, Lopholaena, Metalasia and Tarchonanthus. It should be noted, however, that the presence or absence of distinct growth rings can be influenced by environmental conditions and limited by the size of the piece of

charcoal. Other anatomical features need to be considered. Amongst the five genera with growth rings, Artemisia parenchyma bands are narrow, fibre walls are thin-thick and rays are heterocellular with marginal upright cells. Chrysanthemoides rays are homocellular upright. Distephanus parenchyma bands are narrow and fibre walls are thin. Euryops rays are higher than 1 mm, fibre walls are thick and vessels present helical thickenings (Tusenius, 1986) although not observed in this material. Seriphium parenchyma bands are wide, fibre walls are thick and vessels present helical thickenings (Metcalfe and Chalk, 1950; InsideWood, 2004onwards) although not observed in this material. For example, the Artemisia afra SJL 90 charcoal specimens are distinguished from the Chrysanthemoides monilifera Allott 29 charcoal samples by the banded parenchyma and ray cell arrangement. A. afra parenchyma is arranged in narrow bands of less than three

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Fig. 9. Photomicrographs of Tarchonanthus parvicapitulatus, SJL 92: (A) TS of vessels in short radials of 1-3-5 cells, vessels are small and dense. Vessels are similar to fibres in diameter. (B) TS of scanty parenchyma and fibres with thin-thick walls. (C) TLS of perforation plates and inter-vessel pits (D) TLS of biseriate rays (E) RLS of vessel ray pits equal to inter-vessel pits. (F) RLS rays of procumbent cells with rare square cells.

cells and the rays are heterocellular consisting of mixed procumbent, square and upright cells with marginal upright cells. On the contrary, C. monilifera parenchyma is arranged in wide bands of greater than three cells and the rays are homocellular consisting of upright cells (Table 2). The five other genera present diffuse porous wood and no growth rings. Brachylaena fibre walls are thin-thick to thick, rays are low and wider, 1-3(-6) cells and tend to storied, ray cells are homocellular procumbent or heterocellular procumbent with marginal square cells, prismatic crystals are present in B. discolor. B. huilensis parenchyma bands are narrow and vessels present helical thickenings (Essien et al., 2004onwards) although not observed in this material. Gymnanthemum and Lopholaena rays are high 1 mm and Gymnanthemum fibre walls are thick, whereas

Lopholaena fibre walls are thin-thick. Metalasia fibre walls are thin and rays are low, heterocellular with marginal upright and square cells. Tarchonanthus fibre walls are thin-thick, rays are low, heterocellular, with procumbent long and short cells, rare square cells are intermittent. Similarities between Brachylaena and Tarchonanthus are: the wood is diffuse porous, fibre walls are thin-thick to thick, ray cells are procumbent and square. To distinguish between Brachylaena and Tarchonanthus the differences are: vessels are less grouped and vessels are less dense, fibre walls are thicker in Brachylaena than Tarchonanthus. Fibre diameter is narrower, so the vessels and fibres lumina are varied in Brachylaena but more equable in Tarchonanthus. Fibres are radially flattened in Brachylaena and fibre dimorphism with long and short fibres present in Tarchonanthus

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Fig. 10. Photomicrographs of Tarchonanthus camphoratus SJL 108 (A) TS of vessels in low radial multiples, fibre walls are thin-thick, (C) TLS of narrow, biseriate rays and (E) RLS rays are procumbent with intermittent, rare, square cells. Phytomicrographs of T. trilobus SJL 107 (B) TS vessels are in long radial multiples, fibre walls are thin-thick (D) TLS of rays 1-23 cells wide and (F) RLS rays are procumbent and present prismatic crystals.

(Carlquist, 1961). Parenchyma is sparse, apotracheal diffuse in Brachylaena and paratracheal scanty in Tarchonanthus (Carlquist, 1961), although there are narrow bands in B. huilensis (Essien et al., 2004onwards). Rays are wider in Brachylaena than in Tarchonanthus, although uniseriate rays are present in both, B. huilensis and T. littoralis. Rays tend to storied in Brachylaena and not in Tarchonanthus. Rays are homocellular procumbent or heterocellular procumbent and marginal square cells in Brachylaena, heterocellular procumbent with long and short cells and intermittent square cells in Tarchonanthus. Crystals are rare in Brachylaena and abundant in Tarchonanthus. For example, Brachylaena discolor, SJL 56 vessel diameter, 50 mm is greater than Tarchonanthus parvicapitulatus, SJL 92, 30 mm. B. discolor vessel density, 40e100 vessels/ mm2 is greater than T. parvicapitulatus,  100 vessels/mm2. Axial

parenchyma is scarce, in B. discolor, the arrangement is paratracheal vasicentric and in T. parvicapitulatus parenchyma is paratracheal scanty. In Brachylaena the ray width is 1e6 cells: B. rotundata 1e3e5 (Allott 28, 2005) and B. transvaalensis 2-3-6 (Allott 27, 2005). In Tarchonanthus the width is 1e3: T. parvicapitulatus 1e2 and T. trilobus 1e3 (Kromhout 163, 1975). B. discolor rays are either homocellular with procumbent cells, or heterocellular with procumbent body cells and one row of marginal square cells. T. parvicapitulatus rays are heterocellular, procumbent, long and short body cells, with rare square cells. Rays tend to be storied in Brachylaena but not in Tarchonanthus. Fibres of B. discolor with diameters of 10 mm are narrower than T. parvicapitulatus with diameters of 20 mm. In B. discolor the vessels are wider than the fibres, but similar in T. parvicapitulatus. The vessel to fibre diameter ratio is

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Fig. 11. Photomicrographs of Brachylaena cf. discolor, MOD C6a 32: (A) TS of small and abundant vessels, fibres with thin-thick walls, where vessels are wider than fibres. (C) TLS of wide rays which tend to storied. (E) RLS of rays of procumbent cells with marginal square cells. Photomicrographs of B. discolor SPCA B4c 32: (B) TS of small and abundant vessels and thin-thick fibre walls (D) TLS of wide rays which tend to storied. (F) RLS of procumbent ray cells. The distortion is due to the nature of the archaeological charcoal.

high in B. discolor with 50/10 mm or 5:1 and low in T. parvicapitulatus with 30/20 mm or 1.5:1.

4.3. Identification of Tarchonanthus parvicapitulatus and Brachylaena cf. discolor in archaeological charcoal from Sibudu

parenchyma is paratracheal vasicentric. Rays are one to four cells wide, low and frequent, 12 rays/mm, tending to storied. Rays consist of nine rows of procumbent cells and one row of marginal square cells. Charcoal specimens SPCA B4a 06, SPCA B4a 17 and MOD E4a 54 are conferred B. discolor since some features are distorted (Fig. 11).

4.3.1. Brachylaena cf. discolor. Material studied: MOD C6a 32, SPCA B4c 32, SPCA B4c 11 and 18 (Fig. 11) In the described specimen, MOD C6a 32, the vessels are commonly arranged in long radial multiples of four or more, vessels are small, 20e50 mm to medium, 50e100 mm in diameter and abundant, 50e70 vessels/mm2. The inter-vessel and vessel-ray pits are alternate and minute, ~4 mm. The fibre walls are thin-thick. Vessels are wider than the fibres, in a ratio of ~5:1. The axial

4.3.2. Tarchonanthus parvicapitulatus. Material studied: SPCA B4b 22, SPCA B4c 27 and B4b 08 (Fig. 12) In the described specimen, SPCA B4b 22, the vessels are commonly arranged in short radial lines of 1-3-5, vessels are small, 20e50 mm and dense, 120e150 vessels/mm2. The inter-vessel pits and the vessel-ray pits are alternate and minute, 2e4 mm. Fibre walls are thin. Vessels and fibres are similar in diameter, in a ratio of

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Fig. 12. Photomicrographs of Tarchonanthus parvicapitulatus SPCA B4b 22: (A) TS of vessels in short radials of 1-3-5 cells, vessels are small and dense. Fibre walls are thin. Vessels and fibres are similar in diameter. Parenchyma is scanty; (C) TLS of biseriate rays and irregular cells. Inter-vessel pits are alternate and minute (E) RLS of procumbent ray cells. Photomicrographs of T. parvicapitulatus SPCA B4c 27 (B) TS of vessels in 1-3-5 radial multiples, vessels are small and dense. Fibre walls are thin, (D) TLS of biseriate rays biseriate. Inter-vessel pits are alternate and minute, (F) RLS of rays of procumbent, occasionally short cells.

~1.5:1. The axial parenchyma is paratracheal scanty. The rays are narrow, biseriate, low and frequent, 12 rays/mm, with absent storied structure. Rays consist of procumbent cells and occasionally these are short cells, with rare square cells. 5. Conclusions From the detailed anatomical study of modern reference material and other sources it is possible to distinguish between the selected woody Asteraceae with varying levels of confidence. Artemisia afra and Chrysanthemoides monilifera are distinguished from other woody members in KZN by means of banded parenchyma and ray composition. A. afra rays are heterocellular, mixed cells with marginal upright cells and C. monilifera rays are

homocellular, upright cells. Tarchonanthus parvicapitulatus vessel density is higher than that of Brachylaena discolor. There is less difference between vessel and fibre diameters within T. parvicapitulatus than within B. discolor. T. parvicapitulatus rays are narrower than the B. discolor rays which tend to storied. T. parvicapitulatus ray cells are procumbent, occasionally square. B. discolor ray cells are procumbent with marginal square cells. Woody taxa in charcoal from hearths from SPCA, ~58ka and from MOD, ~49ka are B. discolor SPCA B4c 18 and MOD C6a 32. On the contrary, T. parvicapitulatus charcoal specimens SPCA B4c 27 and SPCA B4b 22 are only from SPCA ~58ka. B. discolor is common in coastal forest and adjacent woodland T. parvicapitulatus occurs in forest and bushveld, mostly inland to 1 850 m.

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Brachylaena discolor may simply have been collected as useful firewood whereas it seems likely that T. parvicapitulatus was deliberately selected for its medicinal properties, due to aromatic oils in the leaves, and perhaps its smoke repelled insects from the camp in Sibudu cave. Insecticidal and larvicidal leaves of Cryptocarya woodii have been recovered from bedding in the 77 ka layers of Sibudu Cave (Wadley et al., 2011) and the charcoals of poisonous wood Spirostachys africana were found in hearths of BYA2(i), SPCA and MOD layers (Lennox and Bamford, 2015; Lennox, 2016b). The confirmed presence of T. parvicapitulatus in one hearth from SPCA layer is another addition to the likely medicinal repertoire at Sibudu in KwaZulu Natal. Acknowledgements SJL would like to express her gratitude to Professor Marion Bamford and Professor Lyn Wadley for enabling this research. The Mellon Foundation (grant to MKB), and Evolutionary Studies Institute (ESI) provided equipment and the research was made possible by funding to Professor Lyn Wadley from the National Research Foundation (NRF) and to SJL from the Palaeontological Scientific Trust - Scatterlings Projects (PAST). Material was collected from farms with permission from the landowner and obtained from the Moss Herbarium, WITS and from Kirstenbosch Botanical Gardens, SANBI. References Abimola, A., 2010. Taxonomy and Pharmacognostic Studies of Tarchonanthus camphoratus Species Complex. MSc Thesis. University of Limpopo, Mankweng. Allott, L.F., 2004. Changing environments in oxygen isotope stage 3: reconstructions using archaeological charcoal from Sibudu Cave. South Afr. J. Sci. 100, 179e184. Allott, L., 2005. Palaeoenvironments of the Middle Stone Age at Sibudu Cave, KwaZulu- Natal, South Africa: an Analysis of Archaeological Charcoal. Unpublished PhD thesis. University of the Witwatersrand, Johannesburg. Allott, L.F., 2006. Archaeological charcoal as a window on palaeovegetation and wood use during the Middle Stone Age at Sibudu Cave. South Afr. Humanit. 18 (1), 173e201. Bamford, M., 2011. Late pliocene woody vegetation of Area 41, Koobi Fora, East Turkana Basin, Kenya. Rev. Palaeobot. Palynol. 164, 191e210. Bamford, M.K., 2015a. Macrobotanical remains from Wonderwerk Cave (Excavation 1), oldowan to late pleistocene (2 Ma to 14 ka BP), South Africa. Afr. Archaeol. Rev. 32, 813e838. Bamford, M.K., 2015b. Charcoal from pre-Holocene Stratum 5, Wonderwerk Cave, South Africa. Palaeoecol. Afr. 33, 153e174. Bamford, M.K., Henderson, Z.L., 2003. A reassessment of the wooden fragment from Florisbad, South Africa. J. Archaeol. Sci. 30, 637e651. Beentje, H.J., 1999. The genus Tarchonanthus (Compositae e Mutisieae). Kew Bull. 54 (1), 81e95. Beentje, H.J., 2000. The genus Brachylaena (Compositae e Mutisieae). Kew Bull. 55 (1), 1e41. Bentsen, S.E., 2012. Size matters: preliminary results from an experimental approach to interpret MSA hearths. Quat. Int. 270, 95e102. Bentsen, S.E., 2013. Controlling the heat: an experimental approach to Middle Stone Age pyrotechnology. South Afr. Archaeol. Bull. 68, 137e145. Bentsen, S.E., 2014a. Using pyrotechnology: fire-related features and activities with a focus on the African Middle Stone Age. J. Archaeol. Res. 22, 141e175. Bentsen, S.E., 2014b. By the Campfire. Pyrotechnology and Middle Stone Age Hearths at Sibudu Cave. PhD Dissertation. University of the Witwatersrand, Johannesburg. Boon, R., 2010. Pooley's Trees of Eastern South Africa. Fauna and Flora Publications Trust, Durban. Braithwaite, M., Van Vuuren, S.F., Viljoen, A.M., 2008. Validation of smoke inhalation therapy to treat microbial infections. J. Ethnopharmacol. 119 (3), 501e506. Bruch, A.A., Sievers, C., Wadley, L., 2012. Quantification of climate and vegetation from southern African Middle Stone Age sites e an application using late Pleistocene plant material from Sibudu, South Africa. Quat. Sci. Rev. 45, 7e17. Carlquist, S., 1961. Wood anatomy of Inuleae (Compositae). Aliso 5, 21e37. Carlquist, S., 1966. Wood anatomy of Compositae: a summary with comments on factors controlling wood evolution. Aliso 6 (2), 25e44. Carlquist, S., 1975. Ecological Strategies of Xylem Evolution. University of California Press, Berkley. Carlquist, S., 2001. Wood anatomy of the endemic woody Asteraceae of St Helena 1: phyletic and ecological aspects. Bot. J. Linn. Soc. 137, 197e210. Carlquist, S., 2010. Caryophyllales: a key group for understanding wood anatomy character states and their evolution. Bot. J. Linn. Soc. 164, 342e393.

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Please cite this article in press as: Lennox, S.J., Bamford, M.K., Identifying Asteraceae, particularly Tarchonanthus parvicapitulatus, in archaeological charcoal from the Middle Stone Age, Quaternary International (2017), http://dx.doi.org/10.1016/j.quaint.2017.03.074