1 Further evidence for bow hunting and its implications more than 60 000 2 years ago: results of a use-trace analysis of the bone point from Klasies River 3 Main site, South Africa.

4

5 Justin Bradfield1*, Marlize Lombard1, Jerome Reynard2, Sarah Wurz2,3

6 1. Palaeo-Research Institute, University of Johannesburg, P.O. Box 524, Auckland Park, 2006, 7 Johannesburg, South Africa

8 2. School of Geography, Archaeology and Environmental Science, University of the 9 Witwatersrand

10 3. SFF Centre for Early Sapiens Behaviour (SapiensCE), University of Bergen, Bergen, Norway

11 * [email protected] 12 13 14 15 Abstract

16 The bone point (SAM 42160) from >60 ka deposits at Klasies River Main Site, South Africa, is 17 reassessed. We clarify the stratigraphic integrity of SAM 42160 and confirm its Middle Stone 18 Age provenience. We find evidence that indicates the point was hafted and partially coated 19 in an adhesive substance. Internal fractures are consistent with stresses occasioned by high- 20 velocity, longitudinal impact. SAM 42160, like its roughly contemporaneous counterpart, 21 farther north at Sibudu Cave, likely functioned as a hafted arrowhead. We highlight a 22 growing body of evidence for bow hunting at this early period and explore what the 23 implications of bow-and-arrow technology might reveal about the cognition of Middle Stone 24 Age people who were able to conceive, construct and use it.

25

26 Keywords: arrowhead; bone point; bow hunting; traceology; cognition, palaeo-neurology; 27 symbiotic technology

28

29

30 31 1. Introduction

32 Multiple lines of evidence have been mounting over the last decade to support bow hunting 33 in South Africa before 60 ka. Use-wear, micro-residues, macro-fractures indicative of 34 impact, and their distribution patterns have been cited as evidence that some stone tools 35 were hafted and used as arrow tips or barbs at sites like Sibudu Cave and Umhlatuzana Rock 36 Shelter in KwaZulu-Natal, South Africa (Fig.1; e.g., Pargeter, 2007; Lombard and Pargeter, 37 2008; Wadley and Mohapi, 2008; Lombard and Phillipson, 2010; Lombard, 2011; de la Peña 38 et al., 2018; but see Villa et al., 2012 for a counter argument), and experimental work 39 continues to assess best-fit scenarios (e.g., Pargeter et al., 2016; Schoville et al., 2017; 40 Pargeter et al., 2018). Bow hunting is further corroborated by the discovery of a bone point 41 from a Howiesons Poort context at Sibudu Cave dating to 61.7±1.5 ka with macro- and 42 micro-fractures that suggest it experienced longitudinal impact usually associated with such 43 use (Backwell et al., 2008, 2018; Bradfield and Lombard, 2011). Similar bone artefacts are 44 known from ethno-historical sources to have been used as arrowheads or link-shafts by 45 southern African hunter-gatherers (e.g., Vinnicombe, 1971; Wiessner, 1983; Deacon, 1992; 46 Bradfield, 2012), and are still used in this manner today (see Wadley et al., 2015).

47 48 49 Figure 1. Map showing sites mentioned in the text. 50 51 At Sibudu Cave there is evidence of a local tradition of bone tool manufacture from at least 52 77 ka, where a variety of specialised tool forms have been recognised (d’Errico et al., 53 2012a). Bone points also occur during Middle Stone Age occupations at Bushman Rock 54 Shelter (Plug, 1982), Peers Cave (d’Errico and Henshilwood, 2007), and Blombos Cave dating 55 to between ~85 ka and ~73 ka (Henshilwood et al., 2001). In addition to these artefacts, 56 bone points are directly associated with bow hunting 39 ka ago at Border Cave, South Africa 57 (Beaumont, 1978), 36 ka ago at White Paintings Shelter, Botswana (Robbins et al., 2012), 58 and 21 ka ago at Boomplaas Cave, South Africa (Deacon, 1984), most of which are thought 59 to have been used as poisoned arrowheads (d’Errico et al., 2012; Robbins et al., 2012; 60 Bradfield et al., 2015). By at least 15 ka bone tools were firmly embedded within culturally 61 mediated technological strategies in parts of North Africa (Desmond et al., 2018), and were 62 probably being used to hunt with the aid of poison by 13 ka in parts of East Africa (Langley 63 et al., 2016a).

64 65 In 1968 John Wymer excavated the Howiesons Poort layers at Klasies River Main site (KRM) 66 on the Tsitsikamma Coast in the Eastern Cape, South Africa (Fig. 1). Among the cultural 67 artefacts recovered was a single bone point (SAM 42160), almost identical to the one later 68 found at Sibudu; but at the time, a unique find. The KRM point came from a compressed 69 layer in an undisturbed, laminated stratigraphic context, and was associated with many 70 silcrete artefacts and distinctive trapezes, characteristic of the Howiesons Poort 71 technocomplex (Singer and Wymer, 1982: 115-116; Wurz, 2013), for which most dated 72 assemblages have ages of between ~66 ka and ~58 ka (see Lombard et al., 2012). Despite 73 the presence of engraved bone artefacts from the same context at KRM, the remaining 74 faunal assemblage reported by Singer and Wymer (1982) did not appear to have been 75 deliberately fractured or otherwise modified for tool making. In most respects SAM 42160 76 resembles twentieth century ethno-historical arrowheads; a technology previously assumed 77 not to predate the Later Stone Age, starting from ~40 ka in southern Africa.

78 Elsewhere, use-trace evidence suggests bone points were hafted as part of complex 79 mechanisms at Matja Kuru 2, Timor Island, 35 ka ago (O’Connor et al., 2014), and bow 80 hunting of arboreal fauna with bone-tipped arrows is also implicated at Niah Cave, Borneo, 81 from Terminal Pleistocene deposits (e.g., Barton et al., 2009; Wedage et al., 2019). Currently 82 the oldest purported bone arrow point from Eurasia dates to ~32 ka at Potočka Zijalka, 83 Slovenia (Odar, 2011), while mechanically delivered projectile hunting in Europe has 84 recently been pushed back to 40-45 ka (Sano et al., 2019).

85 A full technological description of the bone artefacts from some South African Middle Stone 86 Age sites, including SAM 42160, was published by d’Errico and Henshilwood (2007). They 87 suggested that the KRM point was made from a long bone shaft of a medium-sized bovid, 88 and that it was fashioned by scraping with a sharp retouched lithic edge. The reduction in 89 width near the current proximal end suggested to them that the piece was originally 90 pointed at both ends, but that one of the tips snapped off after deposition. They highlighted 91 the morphological and technological similarities of SAM 42160 to that of twentieth century 92 San hunter-gatherer arrow components, implying that the KRM artefact was similarly used. 93 But, based on its morphological traits, they questioned its association with the Middle Stone 94 Age (d’Errico and Henshilwood, 2007: 154-155). 95 If the Howiesons Poort context for SAM 42160 is accepted, it represents one of the oldest 96 known examples of bone tool technology worldwide. Such an age would link it intimately 97 with discussions on the emergence of behavioural complexity (see Galway-Witham et al., 98 2019 for a recent synthesis). Through their association with bow hunting, bone points also 99 have the potential to inform on the evolution of complex of cognition (see discussion 100 below).

101 We therefore revisit the context of SAM 42160 within the site’s sequence and why it is 102 associated with the Howiesons Poort occupational layers, and not the Later Stone Age. We 103 present new use-trace and computed tomographic evidence, indicating that SAM 42160 was 104 hafted and used in a manner consistent with a modular arrow component, thus 105 corroborating the functional interpretation proposed by d’Errico and Henshilwood (2007). 106 Based on these outcomes, we provide an up-to-date discussion about what this may mean 107 in terms of human cognition at KRM during its Howiesons Poort occupation.

108

109 2. The bone point in the context of Klasies River Main site

110 Klasies River Main site consists of Caves 1, 1A, 1B and 2, which are cut into a Table Mountain 111 sandstone cliff facing the Indian Ocean. The vegetation in the immediate vicinity of KRM 112 consists of a complex mosaic of densely interdigitated thicket, forest and coastal vegetation 113 (Van Wijk et al., 2017), forming part of the greater Cape Floristic Region’s Fynbos biome 114 (Mucina and Rutherford, 2006). Precipitation at KRM generally occurs throughout the year 115 (Chase and Meadows, 2007) and the warm Agulhas Current from the south-east results in a 116 relatively mild climate and sea surface temperatures (Carr et al., 2007).

117 KRM was inhabited during the Middle Stone Age, from approximately 120 ka to 48 ka, with 118 the four cave recesses forming a single depository. The overhang of Cave 1A, contains more 119 than 15 metres of steeply sloped midden deposits that formed between 100 ka and 48 ka. 120 The 1.8 metres of Howiesons Poort deposits occur high on the slope. These layers were first 121 excavated by John Wymer in 1968, in the ‘Initial Cutting’ and thereafter in several transects 122 within the Top Cutting (Singer and Wymer, 1982: figure 6.1, page 88). Singer and Wymer 123 (1982) identified 11 layers (layers 10-21) within the Howiesons Poort strata in Cave 1A. The 124 bone point was reported to have come from layer 19 (Singer and Wymer, 1982: 115). In the 125 1980s Hilary Deacon excavated the Howiesons Poort in squares E50, H51 and J51 to the 126 south east of Wymer’s excavation (Deacon and Geleijnse, 1988). Singer and Wymer’s layer 127 19 corresponds broadly to the layers between YS6 in square H51 and CPx1 in square J51 128 (Fig. 2; Villa et al., 2010; Deacon excavation notes). Initially, Deacon (1989) suggested an age 129 centred around 70 ka for the technocomplex at KRM. Subsequent dating of the middle 130 Howiesons Poort layers at Cave 1 provided age ranges between ~70 ka and 60 ka. For 131 example, Vogel (2001) obtained an age of 65.6±5.3 ka based on uranium series for Layer 14 132 (Fig. 2), and Jacobs and colleagues reported three OSL determinations of respectively 133 63.4±2.6, 65.5±2.3 from Cave 2 and 64.1±2.6 from Cave 1A (Jacobs et al., 2008). The age for 134 Cave 1A, from square E50 layer CP18, equivalent to Singer and Wymer’s layer 14 (Fig. 2, see 135 also Fig. S13, sample ZKR6, Jacobs et al. 2008), has recently been revised to 63.2 ± 2.7 136 (Jacobs and Roberts, 2017). These ages are consonant with the interpretation that the 137 Howiesons Poort of southern Africa is generally associated with the penultimate glacial 138 maximum, Marine Isotope Stage (MIS) 4 (Wurz, 2002; Lombard et al., 2012; Langejans et al., 139 2017).

140 141 142 Figure 2. Stratigraphic context of the relevant excavation profiles showing the location of 143 SAM 42160 in SW layer 19 relative to the dated SW layer 14 of the Howiesons Poort at KRM. 144 Note that the CP and other stratigraphic units in E50 and H51 are not continuous, which, 145 apart from the natural slope of the sediment deposit, accounts for the repetition of 146 stratigraphic unit names at different levels. The grey polygons seen in the inset represent 147 the location of Singer & Wymer’s original excavation transects, with the approximate 148 location of SAM 42160 indicated by a star. 149

150 Palaeoclimatic models suggest that this period in the southern Cape, where KRM is located, 151 was likely cooler than the preceding MSA II phase (Thackeray, 1987; Simon et al., 2015; 152 Langejans et al., 2017). Chase and colleagues argue that in the southern Cape MIS 4 was 153 probably a humid period due to westerly storm fronts and the effects of warmer Indian 154 Ocean sea surface temperatures (Chase and Meadows, 2007; Chase, 2010; Carr et al., 2016). 155 Micromammal indices also show an increase in moisture during the Howiesons Poort period 156 at KRM (Avery, 1987).

157 Given the links between glacial conditions and marine regression, shorelines were likely 158 farther away from the site during the Howiesons Poort. However, the relative abundance of 159 seal remains suggests that the shoreline may have fluctuated during this time (Van Pletzen- 160 Vos et al., 2019). Jacobs and Roberts (2008) note that the Howiesons Poort included both a 161 cooling and warming phase, which may account for the relatively close sea levels implied at 162 times throughout this period.

163 The dietary preference of some of the ungulate communities at KRM suggests that 164 grassland habitats prevailed at certain times during the Howiesons Poort. The associated 165 faunal assemblage is dominated by grazers such as equids (Equuus quagga), wildebeest 166 (Connochaetes gnou), and buffalo (Syncerus caffer and S. antiquus), particularly in the upper 167 part of the Howiesons Poort (Klein, 1976; Van Pletzen-Vos et al., 2019). The earlier 168 Howiesons Poort, where the bone point occurs, also includes Raphicerus and kudu 169 (Tragelaphus strepsiceros) with eland (Tragelaphus oryx) present in the basal layers (SW 170 layers 17-21), thus implying that the lower Howiesons Poort may have been bushier and 171 more closed compared to the upper period.

172 From layer 19, transect B2, which is below the dated Howiesons Poort layers mentioned 173 above, Singer and Wymer (1982: 115, fig 8.1 no 5) found an “elegant, delicate ground bone 174 point” from compressed, laminated layers, where “not the slightest sign of disturbance” was 175 evident. They noted that the point was stained and in a similar condition to the other bone 176 material from the Howiesons Poort context – i.e., different from the typical yellow, greasy 177 bones found in several Later Stone Age middens (cf. Schweitzer and Wilson, 1978; 178 Schweitzer, 1979). SAM 42160 is not the only worked bone artefact from the Howiesons 179 Poort levels at the site. An incised bone artefact (SAM-AA 26733) from layer 20, Cave 1A, 180 was found and its Middle Stone Age context is uncontested (Singer and Wymer, 1982; 181 d’Errico and Henshilwood, 2007). 182 A taphonomic assessment of bone fragments from Layer YS6 in H51, to CPx1 in J51 generally 183 shows a distinct colour difference between bone from the CP (Carbonized Parting) layers 184 (CPx1 and CP8-12), and those from the YS (Yellow Sand) layers (YSx1, YS6 and YS7). Given its 185 similarity to bone in the YS layers, we infer that the bone point most likely originates from a 186 YS, rather than a CP layer. Most bone fragments in the CP layers exhibit sheen or a reflective 187 surface usually linked to abrasion, polish or burning (Bromage, 1984: Nicholson, 1993). They 188 are generally dark brown or black in colour and a significant proportion are grey or white. 189 While manganese staining is evident on some fragments, the abundance of calcined bone 190 suggests that most of the bones in these layers were burnt in hearths (Stiner et al., 2011). In 191 contrast, bone in the YS layers are generally light brown or beige in colour and there is little 192 evidence that these bones were modified by heat. While the edges of most YS bone 193 fragments tend to be ‘rounded’ or smoothed, their surfaces are generally chalky and sheen 194 is not common. This suggests that it is unlikely the polished surfaces of any worked bone in 195 these layers was a result of natural abrasion.

196 Thus far, the preliminary results of the ongoing taphonomic analysis of the KRM faunal 197 remains from the Howiesons Poort layers has identified several bone flakes with minimal 198 use-wear, the characteristics and placement of which are dissimilar from natural abrasion 199 and suggestive of ancient human handling (cf., Reynard, 2014; Reynard et al., 2016, in prep). 200 Based on the above context, the notes by the excavators, the presence and condition of 201 other worked bone in the Middle Stone Age of KRM and its Howiesons Poort layers, as well 202 as the fact that the stone tool component of Cave 1 Layer 19 where the point was found is 203 of bona fide Howiesons Poort character (Singer and Wymer 1982), we consider it to have 204 been found in situ – similar to the Middle Stone Age bone points from Blombos Cave that 205 were initially thought to be intrusive from the Later Stone Age (see discussion in 206 Henshilwood et al., 2001). The undisturbed layer from which SAM 42160 was excavated is 207 overlain by the dated layers, so that its age is at least older than 63.4±2.6 ka, the youngest 208 estimate obtained for the Howiesons Poort at KRM by Jacobs and colleagues (2008).

209

210 3. New analysis of the SAM 42160 bone point

211 3.1 Methods 212 Reflected light microscopy and high-resolution computed tomography (CT) were used to 213 analyse the bone point. We followed standard analytical protocols for each method (see 214 Evora, 2015; Griffin et al., 2015; Backwell et al., 2018; Bradfield, 2015a, 2018a). Residues 215 were analysed as far as possible in situ on the artefact surface and were not removed. The 216 micro-wear and residue analysis were undertaken using an Olympus BX51M light 217 microscope, mounted with a DP72 digital camera. The surface of the bone point was 218 analysed at 10x to 500x magnification under florescent light using a combination of filters 219 and polarising lenses. The interpretation of use-wear and residues was likewise based on 220 the literature, comparative reference collections (e.g., Bradfield, 2014) and our own 221 experience (e.g., Legrand and Sidéra, 2007; Lombard and Wadley, 2007; Langejans, 2011; 222 Bradfield, 2015a, b; Antonites et al., 2016).

223 The high-resolution computed tomography was carried out on a Nikon Metrology XTH 224 225/320 LC dual source industrial CT system housed in the Microfocus X-ray Computed 225 Tomography Facility in the Evolutionary Studies Institute at the University of the 226 Witwatersrand. The bone point was scanned in eight overlapping acquisitions at 70kV and 227 120μA energy settings using 3142 projections and 6μm voxel size. All raw data were 228 reconstructed in CT Pro 3D software and the multiple volumes stitched together and 229 analysed on VG Studio Max 2.3 software. Interpretations of fatigue fractures and histology 230 are based on existing literature, summarised elsewhere (see Bradfield 2015a, 2018a; 231 Bradfield et al., 2016).

232 Initial non-destructive analyses were performed to provide a rough characterisation of the 233 black residue, pending permission to conduct destructive sampling for Liquid 234 Chromatography – Mass Spectrometry (LC-MS). Raman spectra were acquired with a WITec 235 alpha300R Confocal Laser Raman Microscope using a 532 nm strength laser. XRF analysis 236 was conducted with a handheld Vanta machine, with a fixed aluminium filter and silicon PIN 237 detector. This device has a 2-watt X-ray tube with 35 kV tungsten anode excitation source.

238

239 3.2 Results 240 The SAM 42160 bone point is slenderer than other Middle Stone Age bone points from 241 southern Africa, plotting closer to arrowheads from the Iron Age and historic periods than to 242 bone points from older contexts (Fig. 3). As mentioned previously (d’Errico and 243 Henshilwood, 2007), the bone point was fashioned by scraping with a lithic edge. There is no 244 sign of abrasive grinding, which is a common finishing technique on Later Stone Age bone 245 points, although its absence should not be taken as an indicator of age (Bradfield, 2014, 246 2015c).

247

248

249 Fig. 3. Scatter plot showing SAM 42160 relative to other similar MSA bone points and a 250 random sample of bone points from Early Later Stone Age, Robberg, Iron Age and 251 ethnological contexts. Sites not mentioned in the key from which bone points are 252 represented include: Boomplaas shelter, Sehonghong, White Paintings Shelter, Nelson Bay 253 Cave, Kwagandaganda, Mapungubwe and the Fourie and Pitt Rivers Museum collections of 254 historical material.

255

256 A hard, black residue is spattered over the surface of the bone point, but concentrated in a 257 band near the proximal end (Fig. 4). This substance has a circumferential distribution and 258 adheres firmly to the surface of the bone. Surprisingly, neither d’Errico and Henshilwood 259 (2007) nor Singer and Wymer (1982) remarked on the presence of this residue. Immediately 260 below the concentrated band of residue the bone appears slightly discoloured, a feature 261 consistent with bone that has been covered for an extended period with a plant-based 262 substance, such as when hafted (see Bradfield, 2015b, 2018b). Under high magnification the 263 heterogenous consistency of this residue is clear, with many inclusions visible (Fig. 4E, H). 264 No chemical characterisation of this residue is attempted here, but is planned for the future. 265 Thus far, we have not been granted permission for such destructive analysis. The entire 266 surface of the bone point is covered in contaminant residues, probably flecks of skin, which 267 fluoresce brightly under ultraviolet light (Fig. 4A-B). These flecks are more abundant 268 overlaying the black residue, perhaps indicating that the latter substance has greater static 269 properties. This kind of contamination occurs easily when artefacts are handled without 270 latex gloves (Bradfield, 2016a), as is the case in the long curatorial history of SAM 42160. 271 Clumps of starch grains are also present immediately overlaying the hard, black residue over 272 most of the point’s length (Fig. 4C). Based on the shape and size of starch grains at least two 273 plant species are represented (see Torrence et al., 2004; Piperno, 2006). In some instances, 274 it is unclear whether the starch forms part of the black substance, but in other instances it 275 clearly overlays the black material.

276 Consistent with previous observations, very little use-wear is present on the bone point. 277 Use-wear is, however, present overlaying the black residue in places where the latter is still 278 well represented. Together with the heterogenous consistency of the residue, this indicates 279 that the bone point was used after the black substance was applied. Two distinct polish and 280 striation characteristics occur. Towards the proximal section the striations are long, deep 281 and consistent with contact against a hard, woody material (Fig. 4G; St-Pierre, 2007; Van 282 Gijn, 2007; Bradfield, 2015a). Towards the apical end the use-wear overlying the black 283 substance becomes more rounded and polished, with shorter, finer striations consistent 284 with contact against a soft, malleable material, like skin (Fig. 4F; Buc and Loponte, 2007; 285 Buc, 2011; Bradfield, 2015a). That this use-wear is present only overlying the black residue 286 and not directly on the bone surface indicates it did not arise from curatorial handling. Faint 287 striations do occur on the bone itself, but these are confined to the darkened area at the 288 base of the point (Fig. 4D). These circumferential striations run perpendicular to the axis of 289 the bone point and are consistent with contact against a soft plant-like material (Becker, 290 2001; Griffitts and Bonsall, 2001; Buc, 2011; Stone, 2013 Bradfield, 2015a). No post- 291 manufacturing traces are present directly on the bone above this section.

292

293 Fig. 4. Use-wear and residue indicators identified on the KRM bone point (SAM 42160). A-B) 294 contaminant flecks fluorescing under ultra-violet light; C) concentration of starch granules 295 seen under cross-polarised light; D) concentration of faint perpendicular striations at base of 296 piece; E) hard, black residue see under normal light and blue penetrating light; F) minimal 297 polish and wear confined to high point topography characteristic of the medial and apical 298 sections of the bone point; G) use-wear overlying hard, black residue; H) showing 299 heterogenous consistency of hard, black residue. 300

301 The results of the high-resolution CT scan are presented in Fig. 5. Five two-dimensional thin 302 sections taken along the length of the bone point are illustrated. Micro-cracks occur 303 abundantly in the apical region and exhibit the typical tri-directional formation indicative of 304 desiccation (Fig. 5A; Backwell et al., 2018). Internal separation of the bone lamellae is also 305 evident, particularly in the apical half of the bone point and likewise indicates desiccation 306 (Fig. 5A-C). A set of two, partially overlapping, unidirectional micro-cracks run from near the 307 base of the piece to about three-quarters of the way up. These micro-cracks are consistent 308 with stress-related damage occasioned by bending forces seen in bone arrowheads (Fig. 5B, 309 C, E; Bradfield, 2013; Bradfield et al., 2016; Backwell et al., 2018). These micro-cracks appear 310 to propagate from the lower extremity towards the tip, congruous with the placement of 311 macro-fracture damage on hafted projectiles (Guthrie, 1983; Bradfield and Lombard, 2011). 312 The extent and nature of the micro-cracks may be seen in the supplemental video file 313 (Supplemental material).

314 The bone histology comprises a thin outer band of plexiform tissue (visible in Fig. 5B) with 315 the rest of the bone matrix exhibiting a primary reticular formation. No secondary osteons 316 are present. Although histological identification at this resolution is not conclusive, the 317 tissue structure appears most similar to perissodactyla bone (Enlow, 1966; Francillon-Vieillot 318 et al., 1990; Martiniaková et al., 2007). Perissodactyla bone was also used to make tools 319 during the Howiesons Poort and older periods at Sibudu Cave in KwaZulu-Natal (Bradfield, 320 2018a).

321 The CT scans provide confirmatory data on the hard, black residue coating the bone point. 322 Figure 5D shows that the residue has a higher density than the bone itself, similar to 323 mineralised deposits. Unlike mineralisation, however, this substance clearly sits on the 324 surface of the bone and has not infiltrated into the bone tissue (cf. Backwell et al., 2018: 325 figure 6). This, together with the circumferential placement, the overlaying use-wear and 326 heterogenous consistency of the residue, supports our interpretation that it was 327 purposefully applied to the bone and did not accumulate incidentally after burial. 328

329

330

331

332

333

334

335

336

337

338

339

340

341

342

343

344 345

346 Fig. 5. High resolution computed tomographs showing two-dimensional cross-sections 347 through the SAM 42160 at various stages. A) showing tri-directional desiccative microcracks; 348 B-C) parallel and overlapping microcracks, also showing plexiform structure at the periosteal 349 region and reticular canals in the medial and endosteal regions; D) high density substance 350 adhering to the surface of the bone; E) parallel microcracks possibly occasioned by bending 351 strain.

352

353 354 The Raman results clearly indicate the heterogenous nature of the residue, with peaks 355 indicative of mineral and organic ingredients (Fig. 6). The haematite on the surface is of 356 interest given the importance of ochre during the Middle Stone Age and the frequency with 357 which people would have handled it (e.g., Henshilwood et al., 2011; Wadley 2012; Hodgkiss 358 2013). Whilst its presence here could result from ochre in the deposit or from transfer from 359 the hands of one of the hunters, there is clear evidence for the use of ochre in adhesive 360 recipes during the Howiesons Poort of South Africa, and it was applied to the surfaces of 361 mastic objects to facilitate handling during Holocene (Lombard, 2007). In the spectra taken

362 at 1nm depth there is evidence of a sugar ring and methylene (CH2), pointing to the 363 presence of organic molecules in the black substance. Peaks indicative of manganese oxide, 364 however, dominate the same spectra. These Raman results suggest that the presence of 365 manganese may be more complex than simple mineral accretion or absorption, as is the 366 case with the Sibudu bone arrowhead (see Backwell et al., 2008).

367 These results highlight the complex nature of the black residue. Given the wide variety of 368 ingredients, including rock minerals, known to have been used in hafting recipes and arrow 369 poisons (Lombard 2007; Bradfield et al., 2015; Wadley et al., 2015; Chaboo et al., 2019), it is 370 possible that manganese, or something containing manganese, was introduced into the 371 residue by human action. Such an interpretation is corroborated by evidence of use 372 subsequent to the application of the residue in the form of overlaying micro-striations. If the 373 black residue was a post-depositional taphonomic phenomenon, one would not expect to 374 see superimposed use-traces (see Lombard 2005 and Wadley & Lombard 2007 on 375 distribution patterns of related use-traces on archaeological tools). Additional XRF results 376 reveal manganese atoms in higher concentrations on the ‘clean’ bone surface, away from 377 the black residue, than in the black residue itself. Manganese traces on the tool are thus not 378 limited to the black residue only, indicating the mineral was probably introduced 379 incidentally. The composition of the black residue itself can only be resolved with detailed 380 LC-MS studies, which falls outside of the scope of this study, but will be conducted pending 381 permission for destructive analysis.

382 383

384 Fig. 6. Raman spectra obtained of the black residue. The upper two spectra were obtained 385 from the surface and the lower three were obtained at a 1nm depth. The slight florescence 386 of the surface spectra is expected given the curatorial history of SAM 42160.

387

388 389

390 Fig. 7. Bar graph showing the results of the XRF analysis taken at two points along SAM 391 421160, namely, on the proximal band of black residue, and on the clean bone, away from 392 any visible black residue.

393

394 4. Discussion 395 4.1. Our results in context

396 The dimensional dissimilarity of SAM 42160 to other Middle Stone Age bone points lead 397 d’Errico and Henshilwood (2007: 160) to suppose that the bone point may have infiltrated 398 into the Howiesons Poort layers from younger sediments. However, as discussed above, 399 Singer and Wymer (1982:115) carefully considered this hypothesis and rejected it, as the 400 bone point was found in an undisturbed context. Moreover, no LSA deposits occur in Cave 401 1A, only within Cave 1, much lower down in the sequence. We find no reason to doubt the 402 integrity of the original context. Although SAM 42160 is slightly thinner than bone points of 403 comparable age, e.g., Sibudu Cave and Peers Cave, all of these Middle Stone Age pieces 404 overlap dimensionally with Later Stone Age, Iron Age and twentieth century arrowheads, 405 suggesting that the small size differences are not meaningful (Fig. 3). 406 Bone points are not always associated with bow hunting. Use-trace evidence from twelve 407 southern African sites indicates that during the Holocene, bone points – morphologically 408 identical to arrowheads – were used, usually hand-held, additionally for wood-working, 409 basketry and hide processing (Bradfield, 2015b, 2016a). The use-wear evidence for these 410 activities, though, is distinctive, and notably absent on both the Sibudu and KRM bone 411 points. Unlike these domestic tasks, hunting does not typically impart diagnostic polishes or 412 micro-striations to bone weapons that would allow one to directly infer function (cf. Stone, 413 2013; Bradfield, 2012, 2015a, b). In the case of the Klasies River bone point, longitudinal 414 striations, however, overlay the softer black residue, which would be consistent with any 415 longitudinal action – including arrow use.

416 Isolated techniques are inadequate to make robust interpretations for a hunting function 417 (Lombard, 2005; Rots and Plisson, 2014; Rots et al., 2016), and confidently identifying 418 arrows or components thereof in the Stone Age record, and distinguishing them from other 419 forms of projectile technology, is difficult. At present, the best approach is to record full 420 suites of use-traces, including hafting traces and stress fractures that form during high- 421 velocity impact (Bradfield, 2016b; Iovita and Sano, 2016). For this reason, we used a 422 combined approach that looked at use-wear, in situ micro-residues and internal stress 423 cracks during our study of the KRM bone point.

424 Despite the weak development of use-wear over much of SAM 42160 the distribution and 425 orientation of micro-striations suggest the proximal 9 mm of the artefact was hafted, 426 probably in a reed shaft. The darker discolouration of this section, a common feature on 427 ethno-historic arrowheads, lends credence to this interpretation (Fig. 8). Socketing a bone 428 arrowhead into a reed or hollow wooden shaft was the preferred method of hafting 429 employed by southern African hunters in historic times (Sparrman, 1977), but was also used 430 in other parts of the world perhaps as early as 32 ka (Odar, 2011).

431 The black residue, which coats SAM 42160, appears concentrated in a circumferential band 432 close to the proximal end of the bone and immediately above the discoloured section. This 433 substance may have been used as a mastic to secure the bone in the reed shaft, or it may be 434 a poison that was applied to the bone after it was hafted. It may, of course, have served 435 both purposes (see Wadley et al., 2015). Micro-morphologically, this substance is similar to 436 poisons used on twentieth century arrows and poison applicators (Fig. 8A), and to adhesives 437 found on quartz arrow tips or barbs from the Howiesons Poort at Sibudu Cave (Lombard, 438 2007; Lombard and Phillipson, 2010; Lombard, 2011), and for adhesives molecularly 439 identified on Howiesons Poort artefacts from Diepkloof in the Western Cape (Charrié- 440 Duhaut et al., 2013).

441 The results of our preliminary Raman and XRF analyses reveal that the black substance is of 442 a heterogenous nature, containing mineral and organic compounds. Although rich in 443 manganese, the presence of this element does not seem to be the result only of normal 444 post-depositional mineral absorption. There is no infiltration into the bone as can be seen in 445 the case of the Sibudu bone point (see Francillon-Vieillot et al., 1990; Backwell et al., 2018); 446 there is evidence for organic molecules in the residue; and use-wear clearly overlies the 447 residue. More detailed organic chemistry of the residue is required to identify all the 448 ingredients. The current distribution pattern of the black residue on the artefact is 449 consistent with poison applications on bone arrowheads during the last few centuries in 450 southern Africa (Fig. 8b), and it is probable that the residue was initially applied to the entire 451 exposed surface of SAM 42160 but has flacked off with time.

452 453 Fig. 8. Bone tools from Holocene contexts showing the same use-traces present on SAM 454 42160. A) black tar-like poison on a poison applicator (MM-40-69-2607) from the Fourie 455 collection, Museum Africa; B) nineteenth century poisoned arrowhead (ET 6593) from the 456 Kalahari showing typical distribution of poison over the point and the natural limit where 457 the bone is inserted into the reed link-shaft; and C) proximal discolouration on hafted bone 458 arrow component from Mapungubwe. 459 460 Two varieties of starch granules were recorded over the black residue. Poisons and mastics 461 are known to have incorporated numerous ingredients, some of which are rich in 462 polysaccharides (Bradfield et al., 2015; Wadley et al., 2015; Chaboo et al., 2019), which 463 could account for the starch granules (also see Rots et al., 2017 for possible starch-rich 464 poison on a Sibudu artefact older than 77ka), and may reflect the sugar ring observed during 465 Raman spectroscopy of the residue. However, given that most of the contaminant residues 466 also occur on or in the vicinity of the black substance, a parsimonious interpretation would 467 be that the starch granules too are contaminants from the soil. The black substance appears 468 to have static properties that attract and adhere other residues.

469 The high-resolution CT scans reveal internal stress fractures that result from bending forces 470 as well as fractures resulting from desiccation. Although not conclusive evidence, the 471 parallel, grouped micro-cracks in the proximal half on SAM 42160 are consistent with 472 bending forces that occur during longitudinal impact (Cotterell and Kamminga, 1992; 473 Bradfield and Lombard, 2011; Bradfield et al., 2016).

474 Arrows, of course, are not the only type of projectile weapon that would experience 475 longitudinal impact. The impact velocity of a modern commercial javelin is comparable to 476 the impact velocity of an arrow shot from a traditional San hunting bow (cf. Hitchcock and 477 Bleed, 1997; Milks et al., 2019), and we ought to expect similar fatigue stresses and 478 fractures in javelin tips as we do in arrowheads. To date there have been no published 479 studies providing reliable use-trace criteria with which to distinguish hand thrown javelins 480 from arrows. There is, however, circumstantial evidence why we discount the possibility 481 that SAM 42160 was anything other than an arrowhead.

482 1. The distribution of hafting traces only 9 mm from the proximal end of the bone 483 indicates that the bone was not hafted in a manner conducive to withstand repeated 484 use as a javelin (see Cotterell & Kaminga 1992; Knecht 1997).

485 2. The morphological and traceological characteristics of SAM 42160 is similar to 486 thousands of Holocene and ethno-historic examples of bone arrowheads found 487 throughout southern Africa. 488 3. There is a total absence of ethno-historically recorded bone-tipped javelins used in 489 southern Africa. When these weapons are at play they are either single-component 490 sharpened wooden or metal-tipped implements.

491 These points caution against an inferential leap ascribing SAM 42160 to anything other than 492 an arrowhead – unless clear evidence can be provided to the contrary.

493 Considered together, the morphology, use-wear, residue, and micro-fracture evidence, as 494 well as the absence of evidence of possible alternative uses, support a scenario in which 495 SAM 42160 was hafted, coated in an adhesive (and possibly poisonous) substance, and used 496 as part of an impact weapon such as an arrowhead, in a similar manner as the Sibudu bone 497 point is thought to have been used and as twentieth century bone points were used by local 498 hunter-gatherers.

499

500 4.2. Bow hunting in the southern Cape before 60 000 years ago

501 Thus far, evidence for bow hunting technology preceding 60 ka has only been reported from 502 the KwaZulu-Natal region of South Africa (Backwell et al., 2008; Lombard and Phillipson, 503 2010). During the Howiesons Poort occupation at both Sibudu Cave and KRM backed stone 504 geometrics were found that were hafted in innovative arrangements and used as weapon 505 components (Villa et al., 2010), most likely as arrow tips, arrowhead insets or barbs 506 (Lombard, 2011; de la Peña et al., 2018). Similar techno-behaviour is hinted at by 71 ka at 507 Pinnacle Point 5-6 along the southern coast of South Africa, based on the presence of 508 Howiesons Poort-like artefacts (Brown et al., 2012; McBrearty, 2012). However, the Pinnacle 509 Point interpretation is speculative as no functional studies were conducted. The oldest 510 supported evidence for bow hunting in the broader Cape region of South Africa during the 511 early Howiesons Poort is therefore the KRM bone point.

512 Bow hunting has several advantages over other hunting strategies. An arrow travels along a 513 straighter trajectory than a javelin, which has a large parabolic arc, thus increasing the 514 accuracy of the hunter (Cotterell & Kaminga, 1992; Knecht, 1997; Hughes, 1998). This is 515 particularly advantageous in closed, thicketed vegetation. The environment during the early 516 Howiesons Poort in the southern Cape was cooler and probably wetter than today. The area 517 around KRM was characterised by a mixed environment, with closed, bushy vegetation 518 (Klein, 1976; Van-Pletzen-Vos et al., 2019), a situation similar to Sibudu Cave, 15 km from 519 the east coast farther north (Clark, 2013, 2017). Bone-tipped arrowheads were used during 520 the Late Pleistocene in southeast Asia to hunt arboreal fauna in densely thicketed 521 vegetation (Rabet and Piper, 2012). Another advantage of bow hunting is that a hunter may 522 carry many arrows in a quiver, thus allowing for a higher success rate and mitigating against 523 potential misses. The possible addition of backed stone geometrics to bone arrowheads (cf. 524 Clark, 1959; Lombard and Parsons, 2008), would have added stabilising weight to the arrow 525 and increased its lethality (Ellis, 1997).

526 Our results do not, however, automatically imply the persistence of bow-and-arrow 527 technology for more than 60 ka on the southern African landscape (see Kuhn, 2006; 528 Lombard and Parsons, 2010; Parsons and Lombard 2011; Lombard, 2016), nor does it imply 529 cultural continuity (see Mitchell, 2012; Pargeter et al., 2016). It does, however, suggest that 530 bow hunting was practised not only in KwaZulu-Natal during the Howiesons Poort, but also 531 along the southern shores of South Africa.

532

533 4.3. Inferences about human cognition at Klasies River Main Site during the 534 Howiesons Poort based on evidence for bow hunting

535 The Howiesons Poort was a period in the Middle Stone Age of southern Africa which saw a 536 florescence of innovations and the technological manifestation of enhanced cognition 537 (Henshilwood and Dubreuil, 2011; McCall and Thomas, 2012; Wadley, 2015). From at least 538 100 ka people in southern Africa were also combining multiple ingredients to form coloured 539 pastes, possibly for decoration or skin protection (Henshilwood et al., 2011; Rifkin et al., 540 2015); and by 70 ka they were making glues and other compound adhesives using multiple 541 ingredients combined in a series of complicated steps (Wadley et al., 2009, 2015; Wadley, 542 2010b). These glues were then used, among other things, to haft small stone segments in 543 varying arrangements, probably as insets for hunting weapons (Lombard, 2007, 2011; 544 Lombard and Pargeter, 2008). For their part, the insets facilitated easier maintenance of 545 weapons and allowed for greater flexibility in how a weapon could be used (see Bousman, 546 1993; Pargeter, 2007). 547 The presence of these technical elements in the southern African Middle Stone Age is seen 548 to represent cognitive complexity that includes notions of abstract thought, analogical 549 reasoning, multitasking and cognitive fluidity (see Wadley, 2015).

550 Bow hunting has been discussed as the adoption of symbiotic or complementary technical 551 systems (e.g., Haidle & Lombard, 2012; Haidle, et al. 2015), which is a manifestation of 552 human cognitive flexibility (Lombard, 2016, 2019). For example, it involves episodic 553 foresight (e.g., Williams et al., 2014; Coolidge et al., 2016) and causal network reasoning, 554 indicating minds that are able to apply abstract engineering concepts across different 555 knowledge domains (e.g., Lombard & Gärdenfors 2017). An even more complex form of 556 reasoning about a ‘force’ that operates for an extended period of time and over long 557 distances (often out of sight of the hunter), is the use of poison (Gärdenfors & Lombard 558 2018; also see Wadley 2010a on the use of snares). Once wounded with a poisoned arrow 559 an animal may be tracked for many hours or even days before finally killed and harvested 560 (see Marshall-Thomas, 2006). Poison is not a physical force; rather it functions chemically, 561 adding yet another domain-specific causal node set to that of bow hunting when it is used.

562 Direct evidence of poisoned arrows dates to 13 ka in East Africa (Langley et al., 2016b), but 563 may be considerably older at 24 ka at Border Cave, South Africa (d’Errico et al., 2012b). 564 Beyond that, the evidence is equivocal. Hunting poisons and complex hafting adhesives 565 could only be made by individuals possessing a basic understanding of 1) the physical and 566 pharmacological properties of each ingredient, 2) the appropriate methods of preparation 567 that would enhance the desired properties and not denature the toxicity or adhesive 568 properties, and 3) the exact sequence and quantities in which ingredients must be added to 569 achieve a successful adhesive or poison recipe (Wadley et al., 2015). Similar to bow hunting, 570 the application of compound adhesives or poisons to hunting weapons requires a level of 571 foresight, anticipation, planning, analogical thought and multi-tasking (e.g., Bradfield et al., 572 2015; Wadley et al., 2015; Wooding et al., 2017; Chaboo et al., 2019) – which is similar to 573 the ways in which we think today.

574 KRM is the sub-Saharan site with the most prolific assemblage of H. sapiens remains 575 spanning the last 100 ka (see Dusseldorp et al., 2013; Grine et al., 2017). Its archaeological 576 record also sparked the first discussions raising the probability that complex human 577 behaviour and cognition were represented in sub-Saharan Africa long before its appearance 578 in Europe (e.g., Deacon, 1989, 1992, 1995, 2001; Wurz, 1999; Deacon and Wurz, 2005). 579 Subsequently, the revolution in dating methods and the excavation of other long-sequence 580 Middle Stone Age sites, such as Sibudu Cave and Blombos Cave, have demonstrated the 581 development of complex cognition for humans well before the suggested 50-45 ka date 582 (e.g., Klein 2019). These discussions are best summarized in papers, amongst others, by 583 Henshilwood (2012), Lombard (2012), McCall and Thomas (2012), and Wadley (2015), and is 584 further supported by the provenience and use-trace evidence of SAM 42160 presented 585 here.

586

587 5. Conclusion

588 We have presented here use-trace evidence that supports the interpretation of a bone 589 point from KRM as a hafted arrowhead. Furthermore, we hope to have dispelled any doubt 590 as to the provenience of SAM 42160 and its attribution to the Howiesons Poort 591 technocomplex. SAM 42160 is part of a growing number of bone and stone artefacts found 592 in Middle Stone Age contexts that are being recognised as arrow armature components. Our 593 findings indicate that SAM 42160 was certainly hafted, maybe poisoned, and used in a 594 manner that is not technologically dissimilar to bone points from more recent contexts. This 595 implies a symbiotic technology that recurred several times with minimal modification 596 throughout the last ~70 ka in southern Africa (see Högberg & Lombard 2020 on socio- 597 technical dynamics of introduction, acceptance and destabilisation). Symbiotic technologies, 598 such as the bow and arrow, tell us something about human cognition and its development 599 (Lombard and Haidle, 2012; Lombard, 2019). Thus far, evidence to support bow hunting 600 before 60 ka period has come from only a few sites, mainly in KwaZulu-Natal. The bone 601 point from KRM extends this geographical distribution farther south and shows that bow 602 hunting was a widely adopted technological innovation in southern Africa during this period.

603

604 Acknowledgements 605 Justin Bradfield is funded by the National Research Foundation (NRF) through the Worked 606 Bone Technology Programme (RDYR180505326418). The authors thank the IZIKO museums 607 of South Africa for the loan of the material. Dr Kuda Jakarta of the Evolutionary Studies 608 Institute, Wits University, is thanked for running the CT scans. Mr Jaganmoy Jodder is 609 thanked for his help in running and interpreting the Raman spectra and XRF analysis. Sarah 610 Wurz acknowledges the support of a DST-NRF Centre of Excellence in Palaeosciences grant. 611 Jerome Reynard is funded by an NRF Thuthuka grant (grant number 107082). Opinions 612 expressed and conclusions arrived at, are those of the authors and are not necessarily to be 613 attributed to the funding agencies.

614

615 References

616 Antonites, A., Bradfield, J., Forssman, T., 2016. Technological, functional and contextual aspects of 617 the K2 and Mapungubwe bone industries. African Archaeological Review 33, 437-463.

618 Avery, D.M., 1987. Late Pleistocene coastal environment of the Southern Cape province of South 619 Africa: Micromammals from Klasies River Mouth. Journal of Archaeological Science 14(4), 405-421.

620 Backwell, L., Bradfield, J., Carlson, K., Jashavilli, T., Wadley, L., d’Errico, F., 2018. The antiquity of 621 bow-and-arrow technology: evidence from Middle Stone Age layers at Sibudu Cave. Antiquity 92, 622 289-303.

623 Backwell, L., d’Errico, F., Wadley, L., 2008. Middle Stone Age bone tools from the Howiesons Poort 624 layers, Sibudu Cave, South Africa. Journal of Archaeological Science 35, 1566-1580.

625 Barton, H., Piper, P., Rabett, R., Reeds, I., 2009. Composite hunting technologies from the Terminal 626 Pleistocene and Early Holocene, Niah Cave, Borneo. Journal of Archaeological Science 36, 1708-1714.

627 Beaumont, P., 1978. Border Cave. Unpublishd MSc thesis. University of Cape Town.

628 Becker, C., 2001. Bone points – no longer a mystery? Evidence from the Slavic urban fortification of 629 Berlin-Spandau. In: Choyke, A., Bartosiewicz, L., (Eds), Crafting Bone: Skeletal Technologies through 630 Time and Space. B.A.R. International Series 937, Archaeopress, Oxford, pp. 129-146.

631 Bousman, B., 1993. Hunter-gatherer adaptation, economic risk and tool design. Lithic Technology 18, 632 59-86.

633 Bradfield, J., 2012. Macrofractures on bone-tipped arrows: analysis of hunter-gatherer arrows in the 634 Fourie Collection from Namibia. Antiquity 86, 1179-1191. 635 Bradfield, J., 2013. Investigating the potential of micro-focus computed tomography in the study of 636 ancient bone tool function: results from actualistic experiments. Journal of Archaeological Science 637 40, 2060-2613.

638 Bradfield, J., 2014. Pointed bone tool technology in southern Africa. Unpublished doctoral thesis. 639 University of Johannesburg.

640 Bradfield, J., 2015a. Use-trace analysis on bone tools: a brief overview of four methodological 641 approaches. South African Archaeological Bulletin 70, 3-14.

642 Bradfield, J., 2015b. Pointed bone tool technology in southern Africa: results of use-trace analyses. 643 Southern African Humanities 27, 1-27.

644 Bradfield, J., 2015c. Identifying bone-tipped arrow types in the archaeological record of southern 645 Africa: the contribution of use-wear studies. Journal of African Archaeology 13, 135-147.

646 Bradfield, J., 2016a. Bone point functional diversity: a cautionary tale from southern Africa. In: 647 Langley, M., (Ed.), Osseous projectile technology. Springer, Doordrecht, pp. 31-40.

648 Bradfield, J., 2016b. Use-trace epistemology and the logic of inference. Lithic Technology 41, 293- 649 303.

650 Bradfield, J., 2018a. Some thoughts of bone discolouration at archaeological sites. Journal of 651 Archaeological Science: Reports 17, 500-509.

652 Bradfield, J., 2018b. A look at the worked bone assemblage from Sibudu Cave, South Africa: 653 Identifying species using CT-rendered bone histographs. PLoS ONE 13, 1-26.

654 Bradfield, J., Lombard, M., 2011. Initial macrofracture study of bone points used in experimental 655 hunting with reference to the South African Middle Stone Age. South African Archaeological Bulletin 656 66, 67 – 76.

657 Bradfield, J., Hoffman, K., De Beer, F., 2016. Verifying the potential of Micro-focus CT in the study of 658 ancient bone tool function. Journal of Archaeological Science: Reports 5, 80-84.

659 Bradfield, J., Wadley, L., Lombard, M., 2015. Southern African arrow poison recipes, their ingredients 660 and implications for Stone Age archaeology. Southern African Humanities 27, 29-66.

661 Bromage, T.G., 1984. Interpretation of scanning electron microscopic images of abraded forming 662 bone surfaces. American Journal of Physical Anthropology 64, 161-178.

663 Brown, K.S., Marean, C.W., Jacobs, Z., Schoville, B.J., Oestmo, S., Fisher, E.C., Bernatchez, J., 664 Karkanas, P., Matthews, T., 2012. An early and enduring advanced technology originating 71,000 665 years ago in South Africa. Nature, 491(7425), 590.

666 Buc, N., 2011. Experimental series and use-wear in bone tools. Journal of Archaeological Science 38, 667 546-557.

668 Buc, N., Loponte, D., 2007. Bone tool types and microwear patterns: some examples from the Pampa 669 region, South America. In: St-Pierre, G., Walker, R., (Eds), Bones as Tools: Current Methods and 670 Interpretations in Worked Bone Studies. British Archaeological Reports International Series 1622, 671 Archaeopress, Oxford, pp. 143-157.

672 Carignani, G., 2016. On the Origin of Technologies: The Invention and Evolution of the Bow-and- 673 Arrow. In: Panebianco, F., Serrelli, E., (Eds), Understanding Cultural Traits. Springer, Switzerland, pp. 674 315-339.

675 Carr, A.S., Thomas, D.S.G., Bateman, M.D., Meadows, M.D., Chase, B., 2007. Late Quaternary 676 palaeoenvironments of the winter-rainfall zone of southern Africa: palynological and 677 sedimentological evidence from the Agulhas plain. Palaeogeography, Palaeoclimatology, 678 Palaeoecology 239, 147-165.

679 Carr, A.S., Chase, B.M., Mackay, A., 2016. Mid to Late Quaternary landscape and environmental 680 dynamics in the Middle Stone Age of southern South Africa. In: Jones, S.C, Steward, B., (Eds), Africa 681 from MIS 6-2: Population Dynamics and Paleoenvironments. Springer, Dordrecht, pp. 23-47.

682 Chaboo, C., Hitchcock, R., Bradfield, J., Wadley, L., 2019. Beetle and plant arrow poisons of the San 683 people of southern Africa. In: Wexler, P., (Ed.), Toxicology in Antiquity. Elsevier, pp.11-72.

684 Charrié-Duhaut, A., Porraz, G., Cartwright, C.R., Igreja, M., Connan, J., Poggenpoel, C., Texier, P.J., 685 2013. First molecular identification of a hafting adhesive in the late Howiesons Poort at Diepkloof 686 Rock Shelter (Western Cape, South Africa). Journal of Archaeological Science 40(9), 3506-3518.

687 Chase, B.M., 2010. South African palaeoenvironments during marine Oxygen Isotope Stage 4: a 688 context for the Howiesons Poort and Still Bay industries. Journal of Archaeological Science 37, 1359- 689 1366.

690 Chase, B.M., Meadows, M.E., 2007. Late Quaternary dynamics of southern Africa’s winter rainfall 691 zone. Earth-Science Reviews 84, 103-138.

692 Clark, J.D., 1959. The prehistory of southern Africa. Harmondsworth. Penguin.

693 Clark, J.L., 2013. Exploring the relationship between climate change and the decline of the 694 Howieson’s Poort at Sibudu Cave (South Africa). In: Clark, J.L., Speth, J.D., (Eds), Zooarchaeology and 695 Modern Human Origins: Human Hunting Behavior during the Late Pleistocene. New York, Springer, 696 pp. 9-18.

697 Clark, J.L., 2017. The Howieson's Poort fauna from Sibudu Cave: Documenting continuity and change 698 within Middle Stone Age industries. Journal of Human Evolution 107, 49-70.

699 Cole, J., 2019. Knapping in the dark: Stone tools and theory of mind. In: Overmann, K., Coolidge, F.L., 700 (Eds), Squeezing Minds from Stones: Cognitive Archaeology and the Evolution of the Human Mind. 701 Oxford, Oxford University Press, pp. 355-375.

702 Coolidge, F.L., 2019. The enhanced working memory model: Its origin and development. In: 703 Overmann, K., Coolidge, F.L., (Eds), Squeezing Minds from Stones: Cognitive Archaeology and the 704 Evolution of the Human Mind. Oxford, Oxford University Press, pp. 406-431. 705 Coolidge, F.L., Haidle, M.N., Lombard, M., Wynn, T., 2016. Bridging theory and bow hunting: human 706 cognitive evolution and archaeology. Antiquity 90, 219-228.

707 Cotterell, B., Kamminga, J., 1992. Mechanics of Pre-industrial Technology. Cambridge University 708 Press, Cambridge.

709 d’Errico, F., Henshilwood, C., 2007. Additional evidence for bone technology in the southern African 710 Middle Stone Age. Journal of Human Evolution 52, 142-163

711 d’Errico, F., Backwell, L., Wadley, L., 2012a. Identifying regional variability in Middle Stone Age bone 712 technology. The case of Sibudu Cave. Journal of Archaeological Science 39, 2479-2495.

713 d’Errico, F., Backwell, L., Villa, P., Degano, I., Lucejko, J., Bamford, M., Higham, T., Colombini, M., 714 Beaumont, P., 2012b. Early evidence of San material culture represented by organic artifacts from 715 Border Cave, South Africa. Proceedings of the National Academy of Sciences 109, 13214-13219.

716 d’Errico, F., Giacobini, G., Puech, P.-F., 1984. Varnish replicas: a new method for the study of worked 717 bone surfaces. Ossa 9 -11, 29-51.

718 de la Peña, P., Taipale, N., Wadley, L., Rots, V., 2018. A techno-functional perspective on quartz 719 micro-notches in Sibudu's Howiesons Poort indicates the use of barbs in hunting technology. Journal 720 of Archaeological Science 93, 166-195.

721 Deacon, H.J., 1989. Late Pleistocene palaeoecology and archaeology in the southern Cape, South 722 Africa. In: Mellars, P., Stringer, C., (Eds), The human revolution: behavioural and biological 723 perspectives on the origins of modern humans. Edinburgh, Edinburgh University Press, pp. 547-564.

724 Deacon, H.J., 1992. Southern Africa and modern human origins. Philosophical Transactions of the 725 Royal Society, London B 337, 177-183.

726 Deacon, H.J., 1995. Two Late Pleistocene-Holocene archaeological depositories from the southern 727 Cape, South Africa. South African Archaeological Bulletin 50, 121-131

728 Deacon, H.J., 2001. Modern Human Emergence: An African Archaeological Perspective. In: Tobias, 729 P.V., Raath, M.A., Moggi-Cecchi, J., Doyle, G., (Eds), Humanity from African Naissance to Coming 730 Millennia – Colloquia in Human Biology and Palaeoanthropology. Florence, Firenze University Press, 731 pp. 217-226.

732 Deacon, J., 1984. The Later Stone Age of southernmost Africa. Cambridge Monographs in African 733 Archaeology.

734 Deacon, H.J., 1989. Late Pleistocene palaeoecology and archaeology in the southern Cape, South 735 Africa. In: Mellars, P., Stringer, C., (Eds), The Human Revolution: Behavioural and Biological 736 Perspectives on the Origins of Modern Humans. Edinburgh University Press, Edinburgh, pp.547-564.

737 Deacon, H.J., Geleijnse, V.B., 1988. The stratigraphy and sedimentology of the main site sequence, 738 Klasies River, South Africa. South African Archaeological Bulletin 43, 5–14.

739 Deacon, H.J., Wurz, S., 2005. A Late Pleistocene archive of life at the coast, Klasies River. In: Stahl, 740 A.B., (Ed.), African Archaeology: a Critical Introduction. Oxford, Blackwell Publishing, pp. 130–149. 741 Desmond, A., Barton, N., Bouzouggar, A., Douka, K., Fernandez, P., Humphrey, L., Morales, J., 742 Buckley, M., Doyan, L., 2018. ZooMS identification of bone tools from the North African Later Stone 743 Age. Journal of Archaeological Science 98, 149-157.

744 Dusseldorp, G., Lombard, M., Wurz, S., 2013. Pleistocene Homo and the updated Stone Age 745 sequence of South Africa. South African Journal of Science 109 (5/6), 01e07.

746 Ellis, C. 1997. Factors influencing the use of stone projectile tips: an ethnographic perspective. In: 747 Knecht, H. (Ed.), Projectile Technology. Plenum Press, New York, pp. 37-78.

748 Enlow, D., 1966. An evaluation of the use of bone histology in forensic medicine and anthropology. 749 In: Evans, F.G., (Ed.), Symposium on Bone and Joints. Springer, Berlin, pp. 92-112.

750 Evora, M., 2015. Use-wear methodology on the analysis of osseous industries. In: Marreiros, J., 751 Bicho, N., Gibaja, J., (Eds), Use-Wear and Residue Analysis in Archaeology, Manuals in Archaeological 752 Method, Theory and Technique. Springer, Switzerland, pp. 159-170.

753 Francillon-Vieillot, H., de Buffrénil, V., Castanet, J., Géraudie, J., Meunier, F., Sire, J., Zylberberg, L., 754 de Ricqlès, A., 1990. Microstructure and mineralization of vertebrate skeletal tissues. In: Carter, J.G., 755 (Ed.), Skeletal biomineralization: patterns, processes and evolutionary trends. Van Nostrand 756 Reinhold, New York, pp. 471-530.

757 Galway‐Witham, J., Cole, J. and Stringer, C., 2019. Aspects of human physical and behavioural 758 evolution during the last 1 million years. Journal of Quaternary Science 759 https://doi.org/10.1002/jqs.3137

760 Gärdenfors, P., Lombard, M., 2018. Causal cognition, force dynamics and early hunting technologies. 761 Frontiers in Psychology 9, 87.

762 Griffin, J., Neaves, S., Claxton, N., Davidson, M., 2015. Optical Microscopy Primer. 763 https://micro.magnet.fsu.edu/primer/index.html. Accessed 2 June 2019.

764 Griffitts, J., Bonsal, C., 2001. Experimental determination of the function of antler and bone ‘bevel- 765 ended tools’ from prehistoric shell middens in western Scotland. In: Choyke, A., Bartosiewicz, L., 766 (Eds), Crafting Bone: Skeletal Technologies through Time and Space. BAR International Series 937, 767 Oxford, Archaeopress, pp. 207-220.

768 Grine, F.E., Wurz, S., Marean, C.W., 2017. The middle stone age human fossil record from Klasies 769 River main site. J. Hum. Evol. 103, 53–78.

770 Guthrie, R.D., 1983. Osseous projectile points: biological considerations affecting raw material 771 selection and design among Paleolithic and Paleoindian peoples. In: Clutton-Brock, J., Grigson, C., 772 (Eds), Animals and Archaeology 1: Hunters and Their Prey. BAR International Series 163. Oxford, 773 Archaeopress, pp. 273-294.

774 Haidle, M.N., Bolus, M., Collard, M., Conard, N.J., Garofoli, D., Lombard, M., Nowell, A., Tennie, C., 775 Whiten, A., 2015. The nature of culture: an eight-grade model for the evolution and expansion of 776 cultural capacities in hominins and other animals. Journal of Anthropological Sciences 93, 43-70. 777 Henshilwood, C.S., 2012. Late Pleistocene techno-traditions in southern Africa: a review of the Still 778 Bay and Howiesons Poort, c. 75–59 ka. Journal of World Prehistory 25, 205-237.

779 Henshilwood, C., d’Errico, F., Marean, C., Milo, R., Yates, R., 2001. An early bone tool industry from 780 the Middle Stone Age at Blombos Cave, South Africa: implications for the origin of modern human 781 behaviour, symbolism and language. Journal of Human Evolution 41, 631-678.

782 Henshilwood, C., Dubreuil, B., 2011. The Still Bay and Howiesons Poort, 77–59 ka: Symbolic material 783 culture and the evolution of the mind during the African Middle Stone Age. Current Anthropology 784 52, 361-400.

785 Henshilwood, C.S., d’Errico, F., Van Niekerk, K.L., Coquinot, Y., Jacobs, Z., Lauritzen, S.E., Menu, M., 786 García-Moreno, R., 2011. A 100,000-year-old ochre-processing workshop at Blombos Cave, South 787 Africa. Science 334(6053), 219-222.

788 Hitchcock, R., Bleed, P., 1997. Each according to need and fashion: spear and arrow use among the 789 San hunters of the Kalahari. In: Knecht, H., (Ed.), Projectile Technology. London, Plenum Press, pp. 790 345-368.

791 Hodgskiss, T., 2013. Ochre use in the Middle Stone Age at Sibudu, South Africa: grinding, rubbing, 792 scoring and engraving. Journal of African Archaeology, 11(1), pp.75-95.

793 Högberg, A. and Lombard, M., in press. ‘I Can Do It’Becomes ‘We Do It’: Kimberley (Australia) and 794 Still Bay (South Africa) Points Through a Socio-technical Framework Lens. Journal of Paleolithic 795 Archaeology Online, 1-31, https://doi.org/10.1007/s41982-019-00042-4

796 Hughes, S., 1998. Getting to the point: evolutionary change in prehistoric weaponry. Journal of 797 Archaeological Method and Theory 5, 345–407.

798 Iovita, R., Sano, K., 2016. Multidisciplinary Approaches to the Study of Stone Age Weaponry. 799 Springer, Dordrecht.

800 Jacobs, Z., Roberts, R.G., 2008. Testing times: old and new chronologies for the Howieson’s Poort 801 and Still Bay industries in environmental context. South African Archaeological Society Goodwin 802 Series 10, 9 – 34.

803 Jacobs, Z., Roberts, R.G., 2017. Single-grain OSL chronologies for the Still Bay and Howieson's Poort 804 industries and the transition between them: Further analyses and statistical modelling. Journal of 805 Human Evolution 107, 1-13.

806 Jacobs, Z., Roberts, R.G., Galbraith, R.F., Deacon, H.J., Grün, R., Mackay, A., Mitchell, P., Vogelsang, 807 R., Wadley, L., 2008. Ages for the Middle Stone Age of southern Africa: implications for human 808 behavior and dispersal. Science 322(5902), 733-735.

809 Klein, R.G., 1976. The mammalian fauna of the Klasies River Mouth sites, southern Cape Province, 810 South Africa. South African Archaeological Bulletin 32, 14-27.

811 Klein, R.G., 2019. Population structure and the evolution of Homo sapiens in Africa. Evolutionary 812 Anthropology: Issues, News, and Reviews, 28(4), 179-188. 813 Knecht, H. 1997. Projectile points of bone, antler, and stone: experimental explorations of 814 manufacture and use. In: Knecht, H. (Ed.), Projectile Technology. Plenum Press, New York, pp. 191- 815 212.

816 Kuhn, S., 2006. Trajectories of change in the Middle Paleolithic in Italy. In: Hovers, E., Kuhn, S., (Eds), 817 Transitions before the transition: Evolution and stability in the Middle Paleolithic and Middle Stone 818 Age. New York, Springer, pp. 109–120.

819 Langejans, G., 2011. Discerning use-related micro-residues on tools: testing the multi-stranded 820 approach for archaeological studies. Journal of Archaeological Science 38, 985-1000.

821 Langejans, G. Dusseldorp, G.L., Thackeray, J.F., 2017. Pleistocene molluscs from Klasies River (South 822 Africa): Reconstructing the local coastal environment. Quaternary International 427, 59-84.

823 Langley, M., 2016. (ed), Osseous Projectile Weaponry: Towards and Understanding of Pleistocene 824 Cultural variability. Springer. Dordrecht.

825 Langley, M., Prendergast, M., Shipton, C., Morales, E., Crowther, A., Boivin, N., 2016a. Poison arrows 826 and bone utensils in late Pleistocene eastern Africa: evidence from Kuumbi Cave, Zanzibar, Azania: 827 Archaeological Research in Africa 51, 155-177.

828 Langley, M., O’Connor, S., Aplin, K., 2016b. A >46,000-year-old kangaroo bone implement from 829 Carpenter's Gap 1 (Kimberley, northwest Australia). Quaternary Science Reviews 154, 199-213.

830 Legrand, A., Sidéra, I., 2007. Methods, means and results when studying European bone industries. 831 In: St-Pierre, C., Walker, R., (Eds), Bones as Tools: Current Methods and Interpretations in Worked 832 Bone Studies. BAR International Series 1622, Oxford, Archaeopress, pp. 67-78.

833 Lombard, M., 2005. A method for identifying Stone Age hunting tools. South African Archaeological 834 Bulletin 60, 115–120.

835 Lombard, M., 2007. The gripping nature of ochre: the association of ochre with Howiesons Poort 836 adhesives and Later Stone Age mastics from South Africa. Journal of Human Evolution 53, 406-419.

837 Lombard, M., 2008. Finding resolution for the Howiesons Poort through the microscope: micro- 838 residue analysis of segments from Sibudu Cave, South Africa. Journal of Archaeological Science 35, 839 26-41.

840 Lombard, M., 2011. Quartz-tipped arrows older than 60 ka: further use-trace evidence from Sibudu, 841 KwaZulu-Natal, South Africa. Journal of Archaeological Science 38, 1918-1930.

842 Lombard, M., 2012. Thinking through the Middle Stone Age of sub-Saharan Africa. Quaternary 843 International 270, 140-155.

844 Lombard, M., 2016. Mountaineering or ratcheting? Stone Age hunting weapons as proxy for thee of 845 human technological, behavioral and cognitive flexibility In: Haidle, M., Conard, N., Bolus, M., (eds), 846 The Nature of Culture. Springer, Berlin, pp. 135-146. 847 Lombard, M., 2019. On the minds of bow hunters. In: Overmann, K., Coolidge, F.L., (Eds), Squeezing 848 Minds from Stones: Cognitive Archaeology and the Evolution of the Human Mind. Oxford, Oxford 849 University Press, pp. 473-496.

850 Lombard, M., Haidle, M., 2012. Thinking a bow-and-arrow set: cognitive implications of Middle 851 Stone Age bow and stone-tipped arrow technology. Cambridge Archaeological Journal 22, 237-264.

852 Lombard, M., Pargeter, J., 2008. Hunting with Howiesons Poort segments: pilot experimental study 853 and the functional interpretation of archaeological tools. Journal of Archaeological Science 35, 26- 854 41.

855 Lombard, M., Parsons, I., 2008. Blade and bladelet function and variability in risk management 856 during the last 2000 years in the Northern Cape. The South African Archaeological Bulletin, 18-27.

857 Lombard, M., Parsons, I., 2010. Fact or fiction? Behavioural and technological reversal after 60 ka in 858 southern Africa. The South African Archaeological Bulletin 65, 224-228.

859 Lombard, M., Parsons, I., 2011. What happened to the human mind after the Howiesons Poort? 860 Antiquity 85, 1433-1443.

861 Lombard, M., Phillipson, L., 2010. Indications of bow and stone-tipped arrow use 64 000 years ago in 862 KwaZulu-Natal, South Africa. Antiquity 84, 635-648.

863 Lombard, M., Wadley, L., 2007. The morphological identification of micro-residues on stone tools 864 using light microscopy: progress and difficulties based on blind tests. Journal of Archaeological 865 Science 34, 155-165.

866 Lombard, M., Wadley, L., Deacon, J., Wurz, S., Parsons, I., Mohapi, L., Swart, J., Mitchell, P., 2012. 867 South African and Lesotho Stone Age sequence updated (i). South African Archaeological Bulletin 67, 868 123-144.

869 Marshall-Thomas, E., 2006. The Old Way. New York. Picador.

870 Martiniaková, M., Grosskopf, B., Omelka, R., Dammers, K., Vondra´ková, M., Bauerová, M., 2007. 871 Histological study of compact bone tissue in some mammals: a method for species determination. 872 International Journal of Osteoarchaeology 17, 82–90.

873 McBrearty, S., 2012. Palaeoanthropology: sharpening the mind. Nature 491(7425), 531.

874 McBrearty, S., Brooks, A., 2000. The Revolution that wasn’t: a new interpretation of the origin of 875 modern behaviour. Journal of Human Evolution 39, 453-563.

876 McCall, G., Thomas, J., 2012. Still Bay and Howiesons Poort foraging strategies: recent research and 877 models of culture change. African Archaeological Review 29, 7-50.

878 Milks, A., Parker, D., Pope, M., 2019. External ballistics of Pleistocene hand-thrown spears: 879 experimental performance data and implications for human evolution. Scientific Reports 9, 820.

880 Mitchell, P., 2012. San origins and transitions to the Later Stone Age: new research from Border 881 Cave, South Africa. South African Journal of Science 108, 11–12. 882 Mucina, L., Rutherford, M.C., 2006. The Vegetation of South Africa, Lesotho and Swaziland. Pretoria. 883 South African National Biodiversity Institute.

884 Nicholson, R.A., 1993. A morphological investigation of burnt animal bone and an evaluation of its 885 utility in archaeology. Journal of Archaeological Science 20, 411-428.

886 O’Connor, S., Robertson, G., Aplin, K., 2014. Are osseous artefacts a window to perishable material 887 culture? Implications of an unusually complex bone tool from the Late Pleistocene of East Timor. 888 Journal of Human Evolution 67, 108-119.

889 Odar, B., 2011. Archers at Potočka zijalka? Arheološki vestnik 62, 433-456.

890 Pargeter, J., 2007. Howiesons Poort segments as hunting weapons: experiments with replicated 891 projectiles. South African Archaeological Bulletin 62, 147-153.

892 Pargeter, J., Bam, L., De Beer, F., Lombard, M., 2017. Microfocus X-ray tomography as a method for 893 characterising macro-fractures on quartz backed tools. The South African Archaeological Bulletin, 72, 894 148-155.

895 Pargeter, J., MacKay, A., Mitchell, P., Shea, J., Stewart, B., 2016. Primordialism and the ‘Pleistocene 896 San’ of southern Africa. Antiquity 90, 1072-1079.

897 Pargeter, J., Shea, J., Utting, B., 2016. Quartz backed tools as arrowheads and hand-cast spearheads: 898 Hunting experiments and macro-fracture analysis. Journal of Archaeological Science 73, 145-157.

899 Parsons, I., Lombard, M., 2011. The multi-disciplinary landscape of mid-late Upper Pleistocene 900 archaeology. South African Archaeological Bulletin 66, 178-181.

901 Piperno, D., 2006. Phytoliths: A Comprehensive Guide for Archaeologists and Paleoecologists. 902 Altamira Press. Lanham.

903 Plug, I., 1982. Bone tools and shell, bone and OES beads from Bushman Rock Shelter, eastern 904 Transvaal. South African Archaeological Bulletin 37, 57-62.

905 Rabett, R., Piper, P., 2012. The emergence of bone technologies at the end of the Pleistocene in 906 southeast Asia: Regional and evolutionary implications. Cambridge Archaeological Journal 22, 37–56.

907 Reynard, J.P., 2014. Trampling in coastal sites: An experimental study on the effects of shell on bone 908 in coastal sediment. Quaternary International 330, 156-170.

909 Reynard, J.P., Discamps, E., Wurz, S., Badenhorst, S., van Niekerk, K.L., Henshilwood, C.S., 2016. 910 Occupation intensity and environmental change at Klipdrift Shelter, southern Cape, South Africa. 911 Palaeogeography, Palaeoclimatology, Palaeoecology 449, 349-364.

912 Rifkin, R., Dayet, L., Queffelec, A., Summers, B., Lategan, M., d’Errico, F. 2015. Evaluating the 913 photoprotective effects of ochre on human skin by in vivo SPF assessment: implications for human 914 evolution, adaptation and dispersal. PLoS ONE 10(9): e0136090. 915 https://doi.org/10.1371/journal.pone.0136090 916 Robbins, L., Campbell, A., Brook, G., Murphy, M., Hitchcock, R., 2012. The antiquity of the bow and 917 arrow in the Kalahari Desert: bone points from White Paintings Rock Shelter, Botswana. Journal of 918 African Archaeology 10, 7-20.

919 Rots, V., Plisson, H., 2014. Projectiles and the abuse of the use-wear method in a search for impact. 920 Journal of Archaeological Science 48, 154-165.

921 Rots, V., Lentfer, C., Schmid, V.C., Porraz, G., Conard, N.J., 2017. Pressure flaking to serrate bifacial 922 points for the hunt during the MIS5 at Sibudu Cave (South Africa). PloS ONE 12(5), p.e0175151.

923 Rots, V., Hayes, E., Cnuts, D., Lepers, C., Fullagar, R., 2016. Making sense of residues on flaked stone 924 artifacts: learning from blind tests. PLoS ONE 12(4), e017831.

925 Sano, K., Arrighi, S., Stani, C., Aureli, D., Boschin, F., Fiore, I., Spagnolo, V., Ricci, S., Crezzini, J., 926 Boscato, P., Gala, M., Tagliacozzo, A., Birarda, G., Vaccari, L., Ronchitelli, A., Moroni, A., Benazzi, S., 927 2019. The earliest evidence for mechanically delivered projectile weapons in Europe. Nature Ecology 928 and Evolution 3, 1409-1414.

929 Schoville, B.J., Wilkins, J., Ritzman, T., Oestmo, S., Brown, K.S., 2017. The performance of heat- 930 treated silcrete backed pieces in actualistic and controlled complex projectile experiments. Journal 931 of Archaeological Science: Reports 14, 302-317.

932 Schweitzer, F., 1979. Excavations at Die Kelders, Cape Province, South Africa: The Holocene deposits. 933 Annals of the South African Museum 78, 101-233.

934 Schweitzer, F., Wilson, M., 1978. A preliminary report on excavations at Byneskranskop, Bredasdorp 935 district, Cape Province. South African Archaeological Bulletin 33, 134-140.

936 Shea, J., Sisk, M., 2010. Complex projectile technology and Homo sapiens dispersal into Western 937 Eurasia. PalaeoAnthropology 2010, 100-122.

938 Simon, M.H., Ziegler, M., Bosmans, J., Barker, S., Reason, C.J., Hall, I.R., 2015. Eastern South African 939 hydroclimate over the past 270,000 years. Scientific Reports 5, 18153.

940 Singer, R., Wymer, J., 1982. The Middle Stone Age at Klasies River Mouth in South Africa. University 941 of Chicago Press, Chicago.

942 Sparrman, A., 1977 [1786]. A Voyage to the Cape of Good Hope 1772 to 1776. Cape Town. Van 943 Riebeek Society.

944 Stiner, M.C., Gopher, A., Barkai, R., 2011. Hearth-side socioeconomics, hunting and paleoecology 945 during the late Lower Paleolithic at Qesem Cave, Israel. Journal of Human Evolution 60, 213-233.

946 Stone, E., 2013. The identification of perishable technologies through usewear on osseous tools: 947 wear patterns on historic and contemporary tools as a standard for identifying raw materials worked 948 in the Late Upper Palaeolithic. In: Choyke, A., O’Connor, S., (Eds), From these bare bones: Raw 949 materials and the study of worked osseous objects. Oxford, Oxbow Books, pp. 28–35. 950 St-Pierre, C., 2007. Bone awls of the St. Lawrence Iroquoians: a microwear analysis. In: St-Pierre, C., 951 Walker, R., (Eds), Bones as Tools: Current Methods and Interpretations in Worked Bone Studies. 952 British Archaeological Reports International Series 1622, Oxford, 107-118.

953 Thackeray, J.F., 1987. Late Quaternary environmental changes inferred from small mammalian 954 fauna, southern Africa. Climatic Change 10(3), 285-305.

955 Torrence, R., Wright, R., Conway, R., 2004. Identification of starch granules using image analysis and 956 multivariate techniques. Journal of Archaeological Science 31, 519-532.

957 van Gijn, A., 2007. The use of bone and antler tools: two examples from the Late Mesolithic in the 958 Dutch Coastal Zone. In: St-Pierre, C., Walker, R., (Eds), Bones as Tools: Current Methods and 959 Interpretations in Worked Bone Studies. British Archaeological Reports International Series 1622, 960 Oxford, 81-92.

961 van Pletzen-Vos, L. Brink, J. Reynard, J.P., Wurz, S. 2019. Revisiting Klasies River: A report on the 962 large mammal remains from the Deacon excavations of Klasies River main site, South Africa. South 963 African Archaeological Bulletin 74: 127-137.

964 Van Wijk, Y., Tusenius, M.L., Rust, R., Cowling, R.M., Wurz, S., 2017. Modern vegetation at the Klasies 965 River archaeological sites, Tsitsikamma coast, south-eastern Cape, South Africa: a reference 966 collection. Plant Ecology and Evolution 150 (1), 13–34.

967 Villa, P., Lenoir, M., 2006. Hunting weapons of the Middle Stone Age and the Middle Palaeolithic: 968 spear points from Sibudu, Rose Cottage and Bouheben. Southern African Humanities 18, 89-122.

969 Villa, P., Soriano, S., Teyssandier, N., Wurz, S., 2010. The Howiesons Poort and MSA III at Klasies River 970 main site, cave 1A. Journal of Archaeological Science 37, 630-655.

971 Villa, P., Soriano, S., Tsanova, T., Degano, I., Higham, T.F., d’Errico, F., Backwell, L., Lucejko, J.J., 972 Colombini, M.P., Beaumont, P.B., 2012. Border cave and the beginning of the later stone age in 973 South Africa. Proceedings of the National Academy of Sciences 109(33), 13208-13213.

974 Vinnicombe, P., 1971. A Bushman hunting kit from the Natal Drakensberg. Annals of the Natal 975 Museum 20, 611-625.

976 Vogel, J.C., 2001. Radiometric dates for the Middle Stone age in South Africa In: Tobias, P., Raath, 977 M., Maggi-Cecchi, J., Doyle, G., (Eds), Humanity from African Naissance to Coming Millennia- 978 Colloquia in Human Biology and Palaeoanthropology. Florence, Florence University Press, pp. 261– 979 268.

980 Wadley, L., 2010a. Were snares and traps used in the Middle Stone Age and does it matter? A review 981 and a case study from Sibudu, South Africa. Journal of Human Evolution 58, 179-192.

982 Wadley, L., 2010b. Compound-adhesive manufacture as a behavioral proxy for complex cognition in 983 the Middle Stone Age. Current Anthropology 51, S111-S119.

984 Wadley, L., 2015. Those marvellous millennia: the Middle Stone Age of southern Africa. Azania: 985 Archaeological Research in Africa 50, 155-226. 986 Wadley, L. and Lombard, M., 2007. Small things in perspective: the contribution of our blind tests to 987 micro-residue studies on archaeological stone tools. Journal of Archaeological Science, 34(6), 988 pp.1001-1010.

989 Wadley, L., Mohapi, M., 2008. A segment is not a monolith: evidence from the Howiesons Poort of 990 Sibudu, South Africa. Journal of Archaeological Science 35, 2594-2605.

991 Wadley, L., Trower, G., Backwell, L., d’Errico, F., 2015. Traditional glue, adhesive and poison used for 992 composite weapons by Ju/’hoan San in Nyae Nyae, Namibia. Implications for the evolution of 993 hunting equipment in prehistory. PLoS ONE 10(10), e0140269.

994 Wadley, L., Hogskiss, T., Grant, M., 2009. Implications for complex cognition from the hafting of tools 995 with compound adhesives in the Middle Stone Age, South Africa. Proceedings of the National 996 Academy of Sciences 106, 9590-9594.

997 Wedage, O., Amano, N., Langley, M., Douka, K., Blinkhorn, J., Crowher, A., Deraniyagala, S., 998 Kourampas, N., Simpson, I., Perera, N., Boivin, N., Petraglia, M., Roberts, P., 2019. Specialized 999 rainforest hunting by Homo sapiens ~45,000 years ago. Nature Communications 10, 739.

1000 Wiessner, P., 1983. Style and social information in Kalahari San projectile points. American Antiquity 1001 48, 253-276.

1002 Williams, V.M., Burke, A., Lombard, M., 2014. Throwing spears and shooting arrows: preliminary 1003 results of a pilot neuroarchaeological study. The South African Archaeological Bulletin 69, 199-207.

1004 Wooding, M., Bradfield, J., Maharaj, V., Koot, D., Wadley, L., Prinsloo, L., Lombard, M., 2017. Report 1005 on biochemical detection methods of some plant-based arrow poisons used by San hunter-gatherers 1006 from southern Africa. South African Journal of Science 113, 1-10.

1007 Wurz, S., 1999. The Howiesons Poort backed artefacts from Klasies River: an argument for symbolic 1008 behaviour. South African Archaeological Bulletin 54, 38–50

1009 Wurz, S., 2002. Variability in the Middle Stone Age lithic sequence, 115 000–60 000 years ago at 1010 Klasies River, South Africa. Journal of Archaeological Science 29, 1001–1015.

1011 Wurz, S., 2013. Technological trends in the Middle Stone Age of South Africa between MIS 7 and MIS 1012 3. Current Anthropology 54, 305–319.

1013 Wurz, S., Lombard, M., 2007. 70 000-year-old geometric backed tools from the Howiesons Poort at 1014 Klasies River, South Africa: Were they used for hunting? Southern African Humanities 19, 1-16.

1015