# Climatic controls on Later Stone Age human adaptation in Africa's southern Cape

Brian Chase, J. Tyler Faith, Alex Mackay, Manuel Chevalier, Andrew S. Carr, Arnoud Boom, Sophak Lim, Paula J. Reimer

<figure><img src="image_1.png" /></figure>

► To cite this version:

Brian Chase, J. Tyler Faith, Alex Mackay, Manuel Chevalier, Andrew S. Carr, et al. Climatic controls on Later Stone Age human adaptation in Africa's southern Cape. Journal of Human Evolution, 2018, 114, pp.35 - 44. 10.1016/j.jhevol.2017.09.006. hal-01817664

HAL Id: hal-01817664

https://hal.umontpellier.fr/hal-01817664

Submitted on 12 Jul 2022

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_2.png" /></figure>

Distributed under a Creative Commons Attribution 4.0 International License
Climatic controls on Later Stone Age human
adaptation in Africa's southern Cape

The authors do not recommend the distribution of this
version of this article.

The article is freely available upon request.

To receive a copy, please send a request to Brian Chase at:
**brian.chase@umontpellier.fr**

Brian M. Chase*<fnref n="1" />, J. Tyler Faith<fnref n="2" />, Alex Mackay<fnref n="3" />, Manuel Chevalier<fnref n="1" />, Andrew S. Carr<fnref n="4" />,

Arnoud Boom<fnref n="4" />, Sophak Lim<fnref n="1" />, Paula J. Reimer<fnref n="5" />

<footnote n="1">¹Centre National de la Recherche Scientifique, UMR 5554, Institut des Sciences de l'Evolution de Montpellier, Université Montpellier, Bat.22, CC061, Place Eugène Bataillon, 34095 Montpellier, cedex5, France;</footnote>

<footnote n="2">²Archaeology Program, School of Social Science, University of Queensland, Brisbane, QLD 4072, Australia;</footnote>

<footnote n="3">³Centre for Archaeological Science, School of Earth and Environmental Sciences, Northfields Avenue, Building 41, University of Wollongong, NSW 2522, Australia;</footnote>

<footnote n="4">⁴Department of Geography, University of Leicester, Leicester, LE1 7RH, UK;</footnote>

<footnote n="5">⁵School of Natural and Built Environment, Queen's University Belfast, Belfast, BT7 1NN, Northern Ireland, UK</footnote>

<footnote>*Correspondence to:</footnote>

<footnote>Email: [brian.chase@umontpellier.fr](mailto:brian.chase@umontpellier.fr)</footnote>
22

Abstract

23

24

25

26

27

28

29

30

31

32

33

34

35

36

37

38

39

40

Africa's southern Cape is a key region for the evolution of our species, with early symbolic systems, marine faunal exploitation, and episodic production of microlithic stone tools taken as evidence for the appearance of distinctively complex human behavior. However, the temporally discontinuous nature of this evidence precludes ready assumptions of intrinsic adaptive benefit, and has encouraged diverse explanations for the occurrence of these behaviors, in terms of regional demographic, social and ecological conditions. Here, we present a new high-resolution multi-proxy record of environmental change that indicates that faunal exploitation patterns and lithic technologies track climatic variation across the last 22,300 years in the southern Cape. Conditions during the Last Glacial Maximum and deglaciation were humid, and zooarchaeological data indicate high foraging returns. By contrast, the Holocene is characterized by much drier conditions and a degraded resource base. Critically, we demonstrate that systems for technological delivery – or provisioning – were responsive to changing humidity and environmental productivity. However, in contrast to prevailing models, bladelet-rich microlithic technologies were deployed under conditions of high foraging returns and abandoned in response to increased aridity and less productive subsistence environments. This suggests that posited links between microlithic technologies and subsistence risk are not universal, and the behavioral sophistication of human populations is reflected in their adaptive flexibility rather than in the use of specific technological systems.

41

42

Keywords: paleoclimate; paleoecology; rock hyrax middens; microlithic; macrofauna; Boomplaas Cave

43
44 **Introduction**

South Africa's southern coastal margin is a key region for the evolution and development of our species (Ambrose, 2002; Ambrose and Lorenz, 1990; Brown et al., 2012; Henshilwood et al., 2004a; Henshilwood et al., 2002; Marean, 2010; Powell et al., 2009). The southern Cape archaeological record has reframed the debate about the evolution of human behavior, providing early examples of engravings, ornaments, heat treatment of tool-stone and the focussed consumption of marine resources (Delagnes et al., 2016; Henshilwood et al., 2004b; Henshilwood et al., 2002; Henshilwood et al., 2014; Marean, 2014). The region also exhibits regular technological turnover through the last 100,000 years, with the intermittent production of bladelets, bifacial points and backed artefacts and the use of fine-grained rock, interspersed with periods lacking regular retouched flake forms and dominated by locally available rocks such as quartzite and quartz (Deacon, 1984; Wilkins et al., 2017). The links between these variable technological and subsistence records and their environmental context – necessary to arguments about the evolution of human adaptation – remain surprisingly unclear (Deacon, 1982; Roberts et al., 2016). This reflects the region's particular climatic dynamism (Chase and Meadows, 2007) coupled with disagreement concerning the interpretation of its paleoenvironmental archives (e.g. Chase and Meadows, 2007; Deacon and Lancaster, 1988; Faith, 2013b; Marean et al., 2014).

In this paper, we focus on the Later Stone Age record in the southern Cape, for which
- in contrast with the Middle Stone Age - high resolution environmental and archaeological
data are now available. We explore the strength of coupling between environments, subsistence
behavior and lithic technology over the last 22,300 years to understand whether, and how
closely, human behavior tracked environmental change. Spanning the transition from the Last
Glacial Maximum (LGM; 26.5-19 ka; Clark et al., 2009) to the Holocene (11.7 ka to present
Lowe et al., 2008), and episodes of the use of bladelet-rich technological systems, our data also
have a bearing on broader debates about the role of what are often termed ‘microlithic’
technologies in issues of human adaptation and expansion.

71 Later Stone Age environments and archaeology in the southern Cape

Influenced by both temperate and tropical climate systems (Figure 1), long-term climate change in the southern Cape is characterized by significant and often abrupt fluctuations (Bard and Rickaby, 2009; Chase et al., 2013; Chase and Meadows, 2007; Heaton et al., 1986; Quick et
al., 2015; Quick et al., 2016; Talma and Vogel, 1992). Existing evidence indicates that during the Holocene the relative influences of the two dominant synoptic scale moisture-bearing systems – 1) the southern westerly storm track, which expands/shifts northward in the winter, and 2) the tropical easterly flow, which transports moisture from the Indian Ocean during the summer – have varied significantly (Chase et al., 2013; Chase et al., 2015b). However, there is little detailed paleoenvironmental evidence pre-dating the Holocene (Carr et al., 2016b; Chase and Meadows, 2007), and as a result there are contradictory opinions concerning conditions since the LGM (Chase and Meadows, 2007; Deacon and Lancaster, 1988; Faith, 2013b; Kohfeld et al., 2013; Partridge et al., 1999; Partridge et al., 2004; Sime et al., 2013; Stone, 2014), to the extent that some studies conclude that the region was exceptionally “harsh” and arid during the LGM (Deacon and Lancaster, 1988; Scholtz, 1986), while others infer greater humidity and highly productive terrestrial environments (e.g. Faith, 2013b; Parkington et al., 2000). This uncertainty has fundamentally hindered our understanding of past climate dynamics in the region, and, by extension, the impact of past climate change on hunter-gatherer adaptive and subsistence strategies during both the Later and Middle Stone Age.

In the southern Cape, the Later Stone Age archaeological sequence is typically divided into several industries or technocomplexes: early Later Stone Age (ELSA ~<40-24 cal kBP), Robberg (~24-12 cal kBP), Oakhurst (~12-8 cal kBP) and Wilton (~8-2 cal kBP), followed by the arrival of Khoikhoi herders in the last 2000 years (Deacon et al., 1984; Deacon, 1978; Lombard et al., 2012; Mitchell, 1988). The ELSA is associated with the production of small flakes, often through bipolar reduction of cores, though it otherwise lacks unifying characteristics and has been described as a period of technological heterogeneity (Mitchell, 1988; Wadley, 1993). The Robberg presents more coherent characteristics, including the production of large numbers of bladelets (small, elongate flakes usually less than 24 mm long) produced both from dedicated bladelet cores and from those worked by bipolar reduction (Mitchell, 1988). The Robberg also sees more concentrated, if episodic, use of fine-grained rocks such as a silcrete and chert than the preceding or subsequent phases (Deacon, 1978; Deacon, 1982). The Oakhurst (or Albany) is typified by fewer bladelets, larger flakes, a range of scraper forms and declining use of fine-grained rock, while the Wilton features both scrapers and backed artefacts and highly variable patterns of raw material use (Deacon, 1972; Deacon, 1978; Lombard et al., 2012). While these units are coarse and mask considerable variation,
106
107

they provide a useful heuristic for discussing broad patterns in technological change across the
later LSA.

108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131

Consistent with the imprecise meaning of the term (Pargeter, 2016), the ELSA, Robberg and Wilton have all been described as ‘microlithic’ (Bousman, 2005; Deacon, 1984; Mitchell, 1988; Wadley, 1993), but based on different characteristics – small flakes in the case of the ELSA, bladelets in the case of the Robberg and backed artefacts in the Wilton (Lombard et al., 2012). The advent of dedicated bladelet production in particular – as characterizes the Robberg – is argued to have presented humans with a significant adaptive advantage during our evolution and dispersal (Ambrose, 2002; Bar-Yosef and Kuhn, 1999; Clarkson et al., 2009; Foley and Lahr, 2003). Some researchers have linked an emphasis on bladelet production with responses to heightened subsistence risk associated with low or declining subsistence resource productivity (Elston and Brantingham, 2002; Petraglia et al., 2009) (for discussion of the risk concept used here see Bamforth and Bleed, 1997). Others have suggested that bladelet production provided benefits under conditions of high residential mobility (Goebel, 2002; Neeley, 2002). Both explanations – increased subsistence risk and increased mobility – have been posited for bladelet-rich systems in southern Africa during globally cooler conditions (Ambrose, 2002; Grosjean et al., 2003; McCall, 2007; McCall and Thomas, 2012; Mellars, 2006; Mitchell, 2000). The Robberg specifically has been associated with increased residential mobility in response to inferred diminishing resource density (Ambrose, 2002; Mitchell, 2000), and has been explained as a risk-dampening response to resource stress (Mackay, 2009). Other researchers, however, have suggested that any tracking between LSA technological systems and paleoenvironmental variation was relatively weak, and occurred only at the broadest scale of environmental change (e.g. Deacon, 1982). The reality of coupling between technology, subsistence conditions and environmental change in this period is thus contested, and with it the viability of high-order explanations for the behavioral significance of artefacts such as bladelets.

132

## Sites and regional setting

133
134
135
136
137

To explore the relationship between environmental change and human activities and technology, we focus on sites from the Swartberg mountains of South Africa's southern Cape, one of the major ranges in the east-west axis of the Cape Fold Mountains (Figure 1). From Seweweekspoort, a deep transversal valley in the central Swartberg mountains, a series of rock hyrax (*Procavia capensis*) middens – stratified accumulations of dried urine and fecal pellets
138

(see Chase et al., 2012) – were identified, and six middens from two sites, Seweweekspoort-1 (SWP-1; 33.3668°S, 21.4144°E) and Seweweekspoort-3 (SWP-3; 33.4092°S, 21.4031°), were analysed for this study. SWP-1 is located on a west-facing cliff on the northern slope of the pass (Figure 1). The SWP-1 middens were taken from several locations within the same larger shelter, formed by a ~100 m overhanging cliff. SWP-3 is located on a low eastern cliff in the central section of the valley near the valley bottom, and experiences a more humid microclimate relative to the exposed position of SWP-1.

139

140

141

142

143

144

145

146

<caption><insert Figure 1></caption>

147

148

The SWP-1 site is located in the North Swartberg Sandstone Fynbos, but less than a kilometre to the north is the Matjiesfontein Shale Renosterveld (Mucina and Rutherford, 2006). The former, depending on altitude and aspect, is predominantly asteraceous, proteoid and restioid fynbos, while the latter is dominated by asteraceous elements, particularly *Elytropappus rhinocerotis*, *Eriocephalus* sp. and *Euryops* sp., and by an increasing number of succulents, primarily from the Crassulaceae family (Mucina and Rutherford, 2006). At SWP-1, these vegetation types inter-digitate to some extent, with the broad west-facing drainage just north of the site supporting more hydrophilic taxa, such as *Protea*. The north-facing rock shelves directly adjacent to the site maintain only shallow soils and a dominance of succulents. Grasses in the region are a mixture of C₃ (e.g., *Erharta*) and C₄ (e.g., *Aristida*, *Stipagrostis*) varieties (http://sibis.sanbi.org/faces/DataSources.jsp; Rutherford et al., 2012; Rutherford et al., 2003; SANBI, 2003). SWP-3 is located in the South Swartberg Sandstone Fynbos, which at the site is primarily proteoid in character. In addition, there are numerous arboreal/Cape thicket elements (e.g., *Podocarpus*, *Celastraceae*, *Dodonaea*, *Searsia*, *Euclea*, *Oleaceae*) associated with drainages and the nearby riparian zones of the valley bottom.

149

150

151

152

153

154

155

156

157

158

159

160

161

162

163

The Seweweekspoort sites are located 70 km west of the well-stratified late Pleistocene / Holocene archaeological site of Boomplaas Cave. This site is also situated on the flanks of the Swartberg and occupies a very similar climatic regime, making it ideal for exploring linkages between climate, environment, subsistence and technology at high temporal resolution. Located on the southern flanks of the Swartberg range overlooking the Cango Valley, Boomplaas Cave was excavated by Hilary Deacon in the 1970s (Deacon, 1979). The

164

165

166

167

168
169 excavated sediments were divided into a series of members, some of which were subdivided
170 into units. The site is positioned within a transitional shrubland whose component species vary
171 as a function of temperature and moisture gradients moving upslope from the floor of the valley
172 (Vlok and Schutte-Vlok, 2010). The lowlands to the south are characterized by renosterveld
173 habitats, which give way to fynbos vegetation along the slopes of the Swartberg. Along
174 watercourses and ravines in the Cango Valley are more densely wooded habitats that include
175 sweet thorn trees (*Vachellia (Acacia) karroo*) and ironwood (*Olea* spp.) among others (Moffett
176 and Deacon, 1977). As is also the case at Seweweekspoort, Boomplaas today receives
177 contributions of both winter and summer rainfall (Figure 1). Thus, the limited grasses that do
178 occur in the area today include a mixture of both C₃ and C₄ species.

179 **Materials and methods**

180 Rock hyrax middens accumulate over thousands of years and preserve continuous records of
181 past climate change (Chase et al., 2012). The six middens from Seweweekspoort sites SWP-1
182 and SWP-3 were selected for analysis because they are composed almost entirely of hyraceum
183 (no visible fecal pellets). Our experience suggests that such middens have superior stratigraphic
184 integrity compared to more pellet-rich middens. Representative portions of the middens were
185 processed as described in Chase et al. (2013; 2012). Radiocarbon age determinations (n=36)
186 were processed at the ¹⁴CHRONO Centre, Queen’s University Belfast using accelerator mass
187 spectrometry (AMS) (Supplementary Online Material [SOM] Figure S1; Table S1). The
188 radiocarbon ages were corrected for isotope fractionation using the AMS measured δ¹³C and
189 calibrated using the SHCal13 calibration data (Hogg et al., 2013). The Bacon 3.0.3 software
190 package (Blaauw and Christen, 2011) was used to generate all age-depth models (SOM Figure
191 S1). Results indicate that these sequences continuously span the last 22,300 years.

192 The fossil pollen content of 82 adjacent, contiguous pollen samples were prepared with
193 standard physical (600 µm sieving and decanting) and chemical (HCl, KOH, HF and acetolysis)
194 methods (Moore et al., 1991). *Lycopodium* tablets were added to the weighed sample to
195 estimate pollen concentrations (Stockmarr, 1971). A minimum pollen sum of 400 grains was
196 counted at a magnification of ×400 under a light microscope, and identified with the help of
197 the literature (Scott, 1982; van Zinderen Bakker, 1953, 1956; van Zinderen Bakker and
198 Coetzee, 1959), and photographic and slide reference collections at the Universities of the Free
199 State, Cape Town, and Montpellier.
The bulk stable nitrogen (¹⁵N) and carbon (¹³C) isotope contents of 767 overlapping hyraceum samples were measured at the Department of Archaeology, University of Cape Town following Chase et al. (2010; 2009; 2011; 2012), with contiguous/overlapping samples obtained from two series of offset 1 mm holes. For the stable isotope analyses, the standard deviation derived from replicate analyses of homogeneous material was better than 0.2‰ for both nitrogen and carbon. Nitrogen isotope results are expressed relative to atmospheric nitrogen (SOM Figure S2). Carbon isotope results are expressed relative to Vienna PDB (SOM Figure S3).

Stable isotope results from the different Seweweekspoort rock hyrax middens were combined into a single aggregate record using Local Regression (LOESS) curve fitting of the combined datasets (SOM Figures S2 and S3). As individual middens under the same climate regime may exhibit differences in their isotopic records due to microclimatic influences on individual foraging ranges (i.e., baseline δ¹⁵N variability), we have adjusted the δ¹⁵N to account for these differences prior to LOESS curve fitting. Using the SWP-1-1 and SWP-1-4b records as a datum, an estimated offset of 1.5‰ was added to the δ¹⁵N data from the SWP-3-1 to compensate for the more humid microclimate in which the midden was found, and 0.5‰ and 1‰ were added to SWP-1-5 and SWP-1-2a respectively to account for their more exposed positions.

The carbon isotopic composition of the hyraceum is representative of vegetation around a midden site (Carr et al., 2016a) and provides information on 1) the relative contribution of C₃, C₄ and CAM plants (Smith, 1972) to the animals' diet, and 2) variations in plant water-use efficiency (WUE) as a function of climate (Ehleringer and Cooper, 1988; Farquhar et al., 1989; Farquhar and Richards, 1984; Pate, 2001). Throughout the broader region, the distribution of C₃ and C₄ grasses tracks the proportion of winter versus summer rainfall (Vogel, 1978). As mentioned, at Seweweekspoort today, grasses are a mosaic of C₃ and C₄ varieties (Rutherford et al., 2012; Rutherford et al., 2003; SANBI, 2003), and where aspect and soil depth limit soil water content, CAM plants become increasingly abundant. As C₃ plants are depleted in ¹³C compared with most CAM and all C₄ plants, higher δ¹³C values indicate more abundant warm season (C₄) grasses and/or succulent plants (CAM), and generally warmer/more arid conditions.

Hyraceum δ¹⁵N is an indicator of changes in ecosystem water-availability (Carr et al., 2016a; Chase et al., 2013; Chase et al., 2015b; Chase et al., 2009; Chase et al., 2011). A positive
relationship exists between aridity and δ¹⁵N in soils, plants and herbivores, with drier conditions correlating with enriched δ¹⁵N (Carr et al., 2016a), most likely as a result of denitrification processes in arid/semi-arid soils (Handley et al., 1999; Handley et al., 1994; Hartman, 2011; Heaton, 1987; Murphy and Bowman, 2006, 2009; Wang et al., 2010). In the hyraceum samples, the narrowly defined feeding range of the hyraxes (<60 m; Sale, 1965), and the accumulation rates of the middens (~20-60 years/sample) enforce a spatio-temporal averaging that reduces the δ¹⁵N variability observed in modern ecosystem studies (Carr et al., 2016a), and provides a more reliable index of past water variability (Carr et al., 2016a; Chase et al., 2012).

## 241 *Boomplaas faunal and archaeological archives*

The Boomplaas sequence spans much of the last >65,000 years (Deacon, 1982), though we focus here on the fauna and flaked stone artifacts from the upper stratigraphic units corresponding in age with the Seweweekspoort record (SOM Table S2). We use these data to explore the relationship between the paleoenvironmental changes documented at Seweweekspoort and mammal community composition, foraging efficiency, and technological organization. Ages for Boomplaas follow Deacon (1982), calibrated using SHCal13 (Hogg et al., 2013). These published data do present limitations, as they do not adequately bracket each stratigraphic unit. In an effort to maximize their utility, and estimate likely intervals of time that each unit may represent, we derived depths from the published stratigraphic diagrams and calculated a general age-depth model for the sequence. While apparently quasi-continuous, with a relatively constant depositional rate, the nature of the sequence, in terms of lithology, suggests more sporadic deposition. In plotting each unit, we have included both minimum and maximum weighted mean ages as well as potential minimum and maximum ages of the units considering potential sources of error related to radiocarbon calibration and assumptions of accumulation rates (SOM Figure S4). This highlights the clear need to initiate a systematic revision of the chronologies of many archaeological sites in the region (e.g. Loftus et al., 2016; Sealy et al., 2016) to enable more robust inter-site and inter-regional comparisons.

Taxonomic abundances (number of identified specimens: NISP) of ungulates from Boomplaas Cave were derived from Faith (2013a), with Klein's (1983) specimen counts used for the uppermost member DGL. Comparable data (minimum number of individuals: MNI) for the Boomplaas microfauna were derived from Avery (1982). To facilitate comparison with the SWP record, we conducted a detrended correspondence analysis (DCA) for both data sets,
264

using the first axis (rescaled from 0 to 100) to broadly summarize faunal composition (Figure 3). The linear trends in both taxonomic groups closely parallel a decline in ungulate grazers (Figure 3), suggesting that the DCA axis 1 scores are related to the replacement of grasslands by more shrubby habitats (see also Faith, 2013a). Broadly parallel changes are also observed in the south coast faunal sequences from Nelson Bay Cave (Klein, 1983) and Byneskranskop 1 (Schweitzer and Wilson, 1983) (Figure 1), implying regional shifts in habitat structure.

265

266

267

268

269

270

Humans are the primary accumulator of the Boomplaas Cave macrofauna from stratum CL4 and above (Faith, 2013a; see also Faith, 2011). This allows us to explore how the environmental changes documented at SWP translate to changes in foraging efficiency. Based on models grounded in optimal foraging theory, we examine two zooarchaeological indicators of foraging efficiency. These include the relative abundance of small-bodied and presumably low-ranked prey (leporids and tortoises) and the average food utility index (FUI; Metcalfe and Jones, 1988) – a proxy for energetic returns (e.g., meat, fat, marrow) of various ungulate body parts – of large mammal (size classes 3 and 4: 84 to 900 kg) skeletal elements (data from Faith, 2011b). Given the evidence for attrition at Boomplaas Cave (Faith and Thompson, in press), we follow Cleghorn and Marean's (2004) taphonomic model of bone survivorship and restrict analysis to the long-bones (femur, tibia, humerus, radius, and metapodials) and skull elements (crania and mandibles). Building on previous studies conducted elsewhere (e.g., Grayson, 1991, 2005; Broughton, 1994, 1999; Cannon, 2003; Faith, 2007; Munro 2004; ), we predict that declining foraging efficiency will be characterized by (1) increasing frequencies of low-ranked prey, reflecting declining availability of high-ranked prey (e.g., large game) on the landscape, and (2) increasing mean FUI values, reflecting intensified field processing due to an increase in carcass transport distances and/or search times. We recognize that because tortoises are slow-moving and easily captured, they might be considered a high-ranked prey type that should be collected whenever encountered (e.g. Stiner et al., 2000). However, the significant positive correlation between their abundances and those of leporids (Spearman's ρ: r_s = 0.814, p ≤ 0.001), which are unequivocally a low-ranked prey type, suggests that we can treat tortoises as low-ranked prey in this context.

271

272

273

274

275

276

277

278

279

280

281

282

283

284

285

286

287

288

289

290

291

292

293

<insert Table 1>

294
All lithic data are taken from Deacon (1982), standardized to site mean (standardized value = (layer value - site mean) / site standard deviation). We focus on the abundance of bladelets (Figure 3d), and three indicators of technological delivery: ratio of cores to retouched flakes (Figure 3g), total proportions of retouched flakes (Figure 3e), and artefact density measured as total number of artefacts per bucket of excavated sediment following Deacon (1982) (Figure 3h). These last three values function, respectively, as proxies for the transport of retouched flakes vs tool-making potential (mode of technological 'provisioning' (Kuhn, 1995)), frequency of flake curation, and intensity of site-use (Barton and Riel-Salvatore, 2014; Kuhn and Clark, 2015; Parry and Kelly, 1987; Riel-Salvatore and Barton, 2004). Provisioning, which we define as the systems by which stone artefact technologies are delivered in anticipation of future needs, has been argued to mediate the response of mobility to environmental change (Mackay et al., 2014; Wilkins et al., 2017). Under conditions of diminished residential mobility, we expect increases in core transport and artefact density, and lower rates of curation. The inverse is expected when mobility increases and the scheduling of movements becomes harder to predict. If bladelets are a response to diminishing subsistence returns, then their abundance should track humidity and resource productivity inversely. Similarly, if bladelets are positively associated with increasing residential mobility and declining durations of site occupancy then we expect an inverse relationship with artefact density. We also consider the relative abundance of spatially-rare, fine-grained rocks such as silcrete and crypto-crystalline silicates (CCS, subsuming chert and chalcedony) (Figure 3f); it has recently been shown that the abundance of rocks such as silcrete is responsive to increases in overall artefact abundance (Will and Mackay, 2016), and may thus reflect diminished residential mobility and improved scheduling of movements.

## 318 Results

### 319 Climate change since the Last Glacial Maximum at Seweweekspoort

The Seweweekspoort record shows substantial changes in both δ¹³C (range 5.1‰) and δ¹⁵N (range 9.5‰) over the last 22,300 years, implying significant changes in vegetation and climate (Figure 2). These changes are coherent with the pollen data from the same material. Across this period, a strong first-order trend is apparent, with cool, humid glacial conditions (indicated by increased cryophilic Fynbos Biome vegetation pollen and lower δ¹³C and δ¹⁵N) transitioning into warmer, but substantially drier conditions during the Holocene (declining fynbos pollen and higher δ¹³C and δ¹⁵N; Figure 2). This aridification coincides with the deglacial decline in
327

Antarctic sea-ice extent (using sea-salt sodium as a proxy) between 19-11 ka (Fischer et al., 2007; Levine et al., 2014; Wolff et al., 2010) (Figure 2), which is considered to be a strong determinant on the position of the westerly storm track (Bard and Rickaby, 2009; Stuut et al., 2004). The elevated grass pollen percentages during the last glacial period, coupled with low δ¹³C suggest the increased prevalence of cool growing season C₃ grasses (Vogel, 1978), consistent with the model that much of the precipitation during this period fell during the winter months as a result of increased westerly influence. While recent work has indicated that variability in summer rainfall may have had some significant impact in the winter rainfall zone (Chase et al., 2015a; Chase et al., 2015b), long-term (i.e., glacial-interglacial timescales) precipitation trends in the summer rainfall zone (Chevalier and Chase, 2015) exhibit a clear antiphase relationship with humidity at Seweweekspoort (Figure 2), indicating that tropical systems played a limited role in the region at these timescales.

328

329

330

331

332

333

334

335

336

337

338

339

340

<insert Figure 2>

341

342

343

344

345

346

347

348

349

350

351

352

353

354

Within this broad first-order trend of deglacial aridification at Seweweekspoort, significant second-order abrupt episodes of wetter conditions (centred at 14.5, 11 and 4 cal kBP, and the last millennium) indicate major reorganisations of regional climate dynamics. In southern Africa, where rainfall regimes are defined by their strong seasonality, the varying contribution of the non-dominant moisture-bearing system can have a substantial impact on regional environments, shortening or attenuating the impact of often pronounced drought seasons (Chase et al., 2015a). In this context, humid episodes within the last glacial-interglacial transition (LGIT; 18.5-11.7 ka) – previously identified as being a period of exceptionally high effective precipitation in the region (Scholtz, 1986) – can be linked to the warming of both high southern latitudes (Stocker, 1998; Stocker and Johnsen, 2003) and the oceans surrounding southwestern Africa (Barker et al., 2009; Farmer et al., 2005; Kim and Schneider, 2003), including a response to the slow-down of Atlantic Meridional Overturning Circulation (AMOC) during Heinrich stadial 1 (HS1; ~18-14.6 ka) (McManus et al., 2004) (Figure 2).

355

356

357

While the influence of the westerly storm track may have diminished as the Subtropical Front shifted poleward (Barker et al., 2009), increased evaporation from warmer oceans and the invigoration of the southern African monsoon system would have augmented the summer
358

359

360

361

362

363

364

365

366

367

368

rain component in what was then primarily a winter rainfall regime, reducing rainfall seasonality and drought stress. At Seweweekspoort, peaks in humidity at 14.5 and 11 cal kBP typify this, with reductions in fynbos vegetation under slightly warmer conditions, and with increased grass cover as a function of more regular rains promoting shallow rooting vegetation. With the onset of the Holocene, as warming continued, the combination of tropical and temperate systems that resulted in these phases of LGIT humidity broke down. Changes in global boundary conditions resulted in 1) a more permanent southerly position of the westerly storm track, and less winter rain, and 2) strong regional warming that intensified potential evapotranspiration, enhancing drought stress (Chevalier and Chase, 2016). Combined, these factors are interpreted to have driven the marked aridification exhibited in the Seweweekspoort records (Figure 2).

369

*Changing resources and technology*

370

371

372

373

374

375

376

377

378

379

380

The climatic changes robustly identified at Seweweekspoort are strongly reflected in our newly synthesised faunal and archaeological records (Figure 3). Large mammals and microfauna from Boomplaas Cave indicate open and grassy environments during the LGM, giving way to shrublands across the Pleistocene-Holocene transition; a phase marked by large mammal extinctions and shifts in faunal community composition throughout the region (Faith and Behrensmeyer, 2013). These changes are evident in the DCA axis 1 scores (Figure 3). The abundance of ungulate grazers and axis 1 scores at the site closely tracks δ¹³C, δ¹⁵N (Figure 3), and changes observed in the pollen record at Seweweekspoort (Figure 2). Likewise, the carbon isotope composition of tooth enamel from Boomplaas grazers parallels the δ¹³C shifts at Seweweekspoort, with predominantly C₃ grasses consumed during the LGM giving way to increased C₄ grasses during the LGIT (Sealy et al., 2016).

381

382

<insert Figure 3>

383

384

385

386

387

388

These findings contrast with earlier interpretations from floral (Deacon et al., 1984; Scholtz, 1986) and faunal assemblages (Avery, 1982; Klein, 1972; Klein, 1980, 1983) from southern Cape archaeological sequences, wherein open grassland environments – inferred primarily from a predominance of grazers in large mammal fossil records – were interpreted as signs of increased aridity. Our results support inferences that evidence for grassier vegetation
389

indicates the influence of more/more regular precipitation on the richer soils of the valleys and plains of the southern Cape (Chase, 2010; Faith, 2013b), which currently support relatively xeric karroid vegetation (Cowling, 1983). In contemporary African ecosystems, elevated precipitation is typically associated with increased biomass of large herbivores (Coe et al., 1976; East, 1984), a phenomenon likely to have been enhanced by increased plant nutrient content under reduced atmospheric CO2 concentrations (Faith, 2011a). The implication for human populations is that cooler, more humid late Pleistocene conditions presented a more productive resource base, including the proliferation of large grazing ungulates.

390

391

392

393

394

395

396

397

Zooarchaeological evidence from the human-accumulated vertebrate assemblages at Boomplaas Cave (CL4 and above) indicates that aridification through the LGIT is associated with a decline in foraging efficiency. Across the LGIT (CL4 to BRL5), tortoises and leporids increase in abundance relative to ungulates (tortoises: $\chi^2_{trend} = 90.332, p < 0.001$; leporids: $\chi^2_{trend} = 64.854, p < 0.001$). Because ungulates are larger and provide greater energetic returns than tortoises or hares – in which case they should be pursued whenever encountered – these trends imply a decline in their abundances on the landscape. This is also reflected in changes in carcass transport strategies. The average food utility index of large mammal high-survival skeletal elements increases steadily across the LGIT (Spearman's rho = 0.964, p < 0.001), indicating a greater emphasis on skeletal parts providing the highest energetic returns. Within an optimal foraging framework, this trend is consistent with an increase in carcass transport distances and/or search times stemming from diminished availability of large game (Cannon, 2003; Faith, 2007). These parallel trends imply that the cooler and more humid conditions of the late Pleistocene provided a more productive vertebrate resource base with higher average energetic returns than did the more arid conditions of the early Holocene.

398

399

400

401

402

403

404

405

406

407

408

409

410

411

412

The decline in foraging efficiency through the deglaciation reverses through the Holocene. Barring an increase observed in the two upper units (Table 1), leporids decline in abundance relative to ungulates after the peak in BRL5 (~11-12 cal kBP) ($\chi^2_{trend} = 15.854, p < 0.001$). This trend, which is consistent with an increase in ungulate abundances on the landscape, complements a decline in the mean FUI of large mammal skeletal parts (Spearman's rho = 0.821, p = 0.034).

413

414

415

416

417

418

From the LGM through the Holocene, there are marked temporal trends in lithic indicators of technological systems at Boomplaas Cave (Figure 3). Though not without some variation, there is a general decline through time in the abundance of bladelets ($\chi^2_{trend} =$

419

420
1323.693, *p* < 0.001), cores relative to retouched flakes (*χ*²_trend = 921.328, *p* < 0.001), and fine-
grained raw materials (*χ*²_trend = 775.322, *p* < 0.001), coupled with an increase in the frequency
of retouched pieces (*χ*²_trend = 1510.282, *p* < 0.001). A relationship between technological
systems and the environment is suggested by strong correlations between these indicators and
the DCA axis 1 scores derived for both ungulates and microfauna (Figure 3b, c; Table 2).
Indeed, all the measures of technological change we employ for the Boomplaas assemblage are
significantly correlated with changes in ungulate community composition, and most with the
composition of micromammal communities (Table 2). Both sets of axis 1 scores track changing
frequencies of grassland indicators at the site, as well as the δ¹⁵N and especially δ¹³C values
from Seweweekspoort (Figure 3). It follows that technological change is tracking
environmental change, including the patterns of climate-driven environmental change
documented in the Seweweekspoort records.

433

434

<insert Table 2>

435

436

437

438

439

440

441

442

443

444

During the LGM and early LGIT bladelets are abundant and artefact indicators are
consistent with low residential mobility (Figure 3). With the exception of the earliest LGM
members (GWA/HCA), for which taphonomic data suggest limited human occupation (Faith,
2013a), artefact densities are well above the overall mean at Boomplaas, flake curation is
uncommon, and all core to retouched flake values pre-13 ka (CL member units) are higher than
all those that follow. Associated high frequencies of rock such as silcrete and CCS before 12
cal kBP (CL + BRL 7) imply regular and predictable access to fine-grained rock through this
period (Figure 3). These factors combined imply an emphasis on the transport of tool-making
potential to sites.

445

446

447

448

449

450

451

Coincident with the onset of the first-order aridification trend after ~14 ka (BRL
member units), artefact densities decrease, as do proportions of fine-grained rock and bladelets.
The period is broadly characterised by more common acquisition and reduction of readily
available local rocks (quartz and quartzite), with diminishing intensity of site use and little
transport of cores or curation of flakes. Investment in technological costs in the later LGIT
appears minimal and may reflect greater allocation of energy to search and handling of
subsistence packages in response to diminishing ungulate abundance (cf., Hames, 1992;
452

Mackay and Marwick, 2011). From the beginning of the Holocene (BRL3 through to DGL member), and tracking diminishing humidity, artefact densities are low and locally abundant rock continues to dominate, but flake curation becomes markedly more common – technological systems show a much greater emphasis on transportation and maintenance of implements in this period than in the Pleistocene.

453

454

455

456

457

## Conclusions

458

459

460

461

462

463

464

465

466

467

468

469

470

471

472

473

474

475

476

477

478

479

480

481

482

483

The findings presented here overturn prevailing models of environmental and behavioral change in Africa's southern Cape. A continuous and high-resolution environmental base-line is provided for the first time, indicating a trend from relative humidity during the LGM to increased aridity during the Holocene, with marked shifts in moisture across the LGIT. Rather than being characterised by 'harsh,' conditions (Deacon and Lancaster, 1988; Scholtz, 1986), the mesic environments of the late Pleistocene were highly productive, with more extensive grasslands existing in areas now dominated by drought resistant succulent shrublands. Zooarchaeological data indicate proliferation of a diverse ungulate grassland community during this time, suggesting greater resource availability for humans living in the area, and reduced search and handling times for large game. While lithic technologies track these changes, we found no evidence to support an association between the production of bladelets during the LGM/early LGIT (Robberg) and diminished subsistence conditions. Indeed, bladelets seem to have flourished in a period of relative resource abundance. The period of lowest subsistence productivity inferred from the Seweweekspoort data probably occurred during the Holocene, associated with aridification and the concomitant loss of large ungulates and faunal diversity recorded at Boomplaas. This change led to increases in carcass processing at kill sites, as evidenced by more selective transport of high utility body parts, increased reliance on low-ranked prey, and a technological response in which flakes from locally acquired rocks were curated, core transportation was relatively rare, and bladelets were uncommon. In documenting the strong coupling of environmental, subsistence and technological behavior in Later Stone Age foragers, our data reflect the simple observation that all lithic technologies can be adaptive solutions, not only those often assumed to provide particular adaptive benefits. The findings afforded by high resolution analysis of late Pleistocene and Holocene climate imply more generally that the lack of certain kinds of technologies - such as bladelets, backed artefacts and bifacial points - in the earlier stages of human evolution need not carry inherent meaning. The ability of foragers to track rapid climatic
484 and environmental changes with adaptive cultural responses is a better arbiter of cognitive
485 complexity than the deployment of any specific technological system.
486 Acknowledgements

487 Funding was received from the European Research Council (ERC) under the European Union's
488 Seventh Framework Programme (FP7/2007-2013)/ERC Starting Grant "HYRAX", grant
489 agreement no. 258657, and the International Union for Quaternary Research (INQUA), project
490 no. 1205P. We would like to thank Matthew Britton and Ian Newton for assistance in collecting
491 and analysing the material.

492 References

493 Ambrose, S.H., 2002. Small things remembered: origins of early Microlithic industries in Sub-
494 Saharan Africa, in: Elston, R.G., Kuhn, S.L. (Eds.), Thinking Small: Global Perspectives on
495 Microlithization. American Anthropological Association, pp. 9-30.

496 Ambrose, S.H., Lorenz, K.G., 1990. Social and ecological models for the Middle Stone Age in
497 Southern Africa, in: Mellars, P. (Ed.), The Emergence of Modern Humans. Edinburgh
498 University Press, Edinburgh, pp. 3-33.

499 Avery, D.M., 1982. Micromammals as palaeoenvironmental indicators and an interpretation of
500 the late Quaternary in the southern Cape Province, South Africa. Ann. S. Afr. Mus. 85, 183-
501 377.

502 Bamforth, D.B., Bleed, P., 1997. Technology, flaked stone technology, and risk, in: Clark, G.A.
503 (Ed.), Rediscovering Darwin: Evolutionary Theory in Archaeology. American
504 Anthropological Association, Washington, pp. 109-140.

505 Bar-Yosef, O., Kuhn, S.L., 1999. The big deal about blades: Laminar technologies and human
506 evolution. Amer. Anthrop. 101, 322-338.

507 Bard, E., Rickaby, R.E.M., 2009. Migration of the subtropical front as a modulator of glacial
508 climate. Nature 460, 380-383.

509 Barker, S., Diz, P., Vautravers, M.J., Pike, J., Knorr, G., Hall, I.R., Broecker, W.S., 2009.
510 Interhemispheric Atlantic seesaw response during the last deglaciation. Nature 457, 1097-
511 1102.

512 Barton, C.M., Riel-Salvatore, J., 2014. The formation of lithic assemblages. J. Archaeol. Sci.
513 46, 334-352.

514 Blaauw, M., Christen, J.A., 2011. Flexible Paleoclimate Age-Depth Models Using an
515 Autoregressive Gamma Process. Bayesian Analysis 6, 457-474.

516 Bousman, C.B., 2005. Coping with risk: Later stone age technological strategies at
517 Blydefontein Rock Shelter, South Africa. J. Anthrop. Archaeol. 24, 193-226.

518 Brown, K.S., Marean, C.W., Jacobs, Z., Schoville, B.J., Oestmo, S., Fisher, E.C., Bernatchez,
519 J., Karkanas, P., Matthews, T., 2012. An early and enduring advanced technology originating
520 71,000 years ago in South Africa. Nature 491, 590-593.

521 Cannon, M.D., 2003. A model of central place forager prey choice and an application to faunal
522 remains from the Mimbres Valley, New Mexico. J. Anthrop. Archaeol. 22, 1-25.
523

524

525

Carr, A.S., Chase, B.M., Boom, A., Medina-Sanchez, J., 2016a. Stable isotope analyses of rock hyrax faecal pellets, hyraceum and associated vegetation in southern Africa: Implications for dietary ecology and palaeoenvironmental reconstructions. J. Arid Environ. 134, 33-48.

526

527

528

529

Carr, A.S., Chase, B.M., Mackay, A., 2016b. Mid to Late Quaternary Landscape and Environmental Dynamics in the Middle Stone Age of Southern South Africa, in: Stewart, B.A., Jones, S. (Eds.), Africa from Stages 6-2: Population Dynamics an Palaeoenvironments. Springer.

530

531

Chase, B.M., 2010. South African palaeoenvironments during marine oxygen isotope stage 4: a context for the Howiesons Poort and Still Bay industries. J. Archaeol. Sci. 37, 1359-1366.

532

533

534

Chase, B.M., Boom, A., Carr, A.S., Carré, M., Chevalier, M., Meadows, M.E., Pedro, J.B., Stager, J.C., Reimer, P.J., 2015a. Evolving southwest African response to abrupt deglacial North Atlantic climate change events. QSRv 121, 132-136.

535

536

537

Chase, B.M., Boom, A., Carr, A.S., Meadows, M.E., Reimer, P.J., 2013. Holocene climate change in southernmost South Africa: rock hyrax middens record shifts in the southern westerlies. QSRv 82, 199-205.

538

539

540

Chase, B.M., Lim, S., Chevalier, M., Boom, A., Carr, A.S., Meadows, M.E., Reimer, P.J., 2015b. Influence of tropical easterlies in southern Africa's winter rainfall zone during the Holocene. QSRv 107, 138-148.

541

542

Chase, B.M., Meadows, M.E., 2007. Late Quaternary dynamics of southern Africa's winter rainfall zone. Earth-Sci. Rev. 84, 103-138.

543

544

545

Chase, B.M., Meadows, M.E., Carr, A.S., Reimer, P.J., 2010. Evidence for progressive Holocene aridification in southern Africa recorded in Namibian hyrax middens: implications for African Monsoon dynamics and the "African Humid Period". QuRes 74, 36-45.

546

547

548

Chase, B.M., Meadows, M.E., Scott, L., Thomas, D.S.G., Marais, E., Sealy, J., Reimer, P.J., 2009. A record of rapid Holocene climate change preserved in hyrax middens from southwestern Africa. Geo 37, 703-706.

549

550

551

Chase, B.M., Quick, L.J., Meadows, M.E., Scott, L., Thomas, D.S.G., Reimer, P.J., 2011. Late glacial interhemispheric climate dynamics revealed in South African hyrax middens. Geo 39, 19-22.

552

553

554

Chase, B.M., Scott, L., Meadows, M.E., Gil-Romera, G., Boom, A., Carr, A.S., Reimer, P.J., Truc, L., Valsecchi, V., Quick, L.J., 2012. Rock hyrax middens: a palaeoenvironmental archive for southern African drylands. QSRv 56, 107-125.

555

556

557

Chevalier, M., Chase, B.M., 2015. Southeast African records reveal a coherent shift from high- to low-latitude forcing mechanisms along the east African margin across last glacial-interglacial transition. QSRv 125, 117-130.

558

559

Chevalier, M., Chase, B.M., 2016. Determining the drivers of long-term aridity variability: a southern African case study. JQS 31, 143-151.

560

561

Clark, P.U., Dyke, A.S., Shakun, J.D., Carlson, A.E., Clark, J., Wohlfarth, B., Mitrovica, J.X., Hostetler, S.W., McCabe, A.M., 2009. The Last Glacial Maximum. Sci 325, 710-714.

562

563

564

565

Clarkson, C., Petraglia, M., Korisettar, R., Haslam, M., Boivin, N., Crowther, A., Ditchfield, P., Fuller, D., Miracle, P., Harris, C., Connell, K., James, H., Koshy, J., 2009. The oldest and longest enduring microlithic sequence in India: 35 000 years of modern human occupation and change at the Jwalapuram Locality 9 rockshelter. Antiquity 83, 326-348.
566 Cleghorn, N., Marean, C.W., 2004. Distinguishing selective transport and in situ attrition: a
critical review of analytical approaches. *J Taphon* 2, 43-67.

568 Coe, M.J., Cumming, D.H., Phillipson, J., 1976. Biomass and production of large African
569 herbivores in relation to rainfall and primary production. *Oecologia* 22, 341-354.

570 Cowling, R., 1983. Phytochorology and vegetation history in the south-eastern Cape, South
571 Africa. *J. Biogeogr.* 10, 393-419.

572 Deacon, H.J., 1979. Excavations at Boomplaas Cave : a sequence through the Upper
573 Pleistocene and Holocene in South Africa. *Wld. Archaeol.* 10, 241-257.

574 Deacon, H.J., Deacon, J., Scholtz, A., Thackeray, J.F., Brink, J.S., 1984. Correlation of
575 palaeoenvironmental data from the Late Pleistocene and Holocene deposits at Boomplaas
576 Cave, southern Cape, in: Vogel, J.C. (Ed.), Late Cainozoic Palaeoclimates of the Southern
577 Hemisphere. *Balkema, Rotterdam*, pp. 339-360.

578 Deacon, J., 1972. Wilton: An assessment after fifty years. S. Afr. Archaeol. Bull. 27, 10-48.

579 Deacon, J., 1978. Changing patterns in late Pleistocene / early Holocene prehistory in southern
580 Africa as seen from the Nelson Bay Cave stone artifact sequence. *QuRes* 10, 84-111.

581 Deacon, J., 1982. The Later Stone Age in the southern Cape, South Africa. University of Cape
582 Town, Cape Town, South Africa.

583 Deacon, J., 1984. The Later Stone Age in southernmost Africa. *Archaeopress, Oxford*.

584 Deacon, J., Lancaster, N., 1988. Late Quaternary palaeoenvironments of southern Africa.
585 Clarendon Press, Oxford.

586 Delagnes, A., Schmidt, P., Douze, K., Wurz, S., Bellot-Gurlet, L., Conard, N.J., Nickel, K.G.,
587 van Niekerk, K.L., Henshilwood, C.S., 2016. Early Evidence for the Extensive Heat Treatment
588 of Silcrete in the Howiesons Poort at Klipdrift Shelter (Layer PBD, 65 ka), South Africa. *PLoS
589 One* 11, e0163874.

590 East, R., 1984. Rainfall, soil nutrient status and biomass of large African savanna mammals.
591 Afr. *J. Ecol.* 22, 245-270.

592 Ehleringer, J.R., Cooper, T.A., 1988. Correlations between carbon isotope ratio and
593 microhabitat of desert plants. *Oecologia* 76, 562-566.

594 Elston, R.G.-, Brantingham, P.J., 2002. Microlithic technology in Northern Asia: A risk-
595 minimizing strategy of the Late Paleolithic and Early Holocene, in: Elston, R.G., Kuhn, S.L.
596 (Eds.), *Thinking Small: Global Perspectives on Microlithization*.

597 Faith, J.T., 2007. Changes in reindeer body part representation at Grotte XVI, Dordogne,
598 France. *J. Archaeol. Sci.* 34, 2003-2011.

599 Faith, J.T., 2011a. Late Pleistocene climate change, nutrient cycling, and the megafaunal
600 extinctions in North America. *QSRv* 30, 1675-1680.

601 Faith, J.T., 2011b. Late Quaternary Megafaunal Extinctions in Southern Africa's Cape Floral
602 Region, Department of Anthropology. The George Washington University.

603 Faith, J.T., 2013a. Taphonomic and paleoecological change in the large mammal sequence
604 from Boomplaas Cave, Western Cape, South Africa. *J. Hum. Evol.* 65, 715-730.

605 Faith, J.T., 2013b. Ungulate diversity and precipitation history since the Last Glacial Maximum
606 in the Western Cape, South Africa. *QSRv* 68, 191-199.
607
608

Faith, J.T., Behrensmeyer, A.K., 2013. Climate change and faunal turnover: testing the mechanics of the turnover-pulse hypothesis with South African fossil data. Palb 39, 609-627.

609
610

611
612

Faith, J.T., Thompson, J.C., in press. Low-survival skeletal elements track attrition, not carcass transport behavior in Quaternary large mammal assemblages, in: Giovas, C.M., LeFebre, M. (Eds.), Zooarchaeology in Practice: Case Studies in Methodology and Interpretation in Archaeofaunal Analysis. Springer.

613
614
615

616
617

Farmer, E.C., deMenocal, P.B., Marchitto, T.M., 2005. Holocene and deglacial ocean temperature variability in the Benguela upwelling region: implications for low-latitude atmospheric circulation. PalOc 20, doi:10.1029/2004PA001049.

618
619

620
621
622
623
624
625
626

Farquhar, G.D., Ehleringer, J.R., Hubick, K.T., 1989. Carbon isotope discrimination and photosynthesis. Annu. Rev. Plant Physiol. Plant Mol. Biol. 40, 503-537.

Farquhar, G.D., Richards, R.A., 1984. Isotopic composition of plant carbon correlates with wtaer-use efficiency of wheat genotypes. Aust. J. Plant Physiol. 11, 539-552.

627
628

629
630
631

632
633
634

Fischer, H., Fundel, F., Ruth, U., Twarloh, B., Wegner, A., Udisti, R., Becagli, S., Castellano, E., Morganti, A., Severi, M., Wolff, E., Littot, G., Röthlisberger, R., Mulvaney, R., Hutterli, M.A., Kaufmann, P., Federer, U., Lambert, F., Bigler, M., Hansson, M., Jonsell, U., de Angelis, M., Boutron, C., Siggaard-Andersen, M.-L., Steffensen, J.P., Barbante, C., Gaspari, V., Gabrielli, P., Wagenbach, D., 2007. Reconstruction of millennial changes in dust emission, transport and regional sea ice coverage using the deep EPICA ice cores from the Atlantic and Indian Ocean sector of Antarctica. E&PSL 260, 340-354.

627
628

629
630
631

632
633
634

Foley, R.A., Lahr, M.M., 2003. On stony ground: Lithic technology, human evolution, and the emergence of culture. Evol Anthropol 12, 109-122.

Goebel, T., 2002. The “Microblade Adaptation” and recolonization of Siberia during the Late Upper Pleistocene, in: Elston, R.G., Kuhn, S.L. (Eds.), Thinking Small: Global Perspectives on Microlithization.

635
636

637
638
639

Grosjean, M., Cartajena, I., Geyh, M.A., Nunez, L., 2003. From proxy data to paleoclimate interpretation: the mid-Holocene paradox of the Atacama Desert, northern Chile. Palaeogeogr., Palaeoclimatol., Palaeoecol. 194, 247-258.

Hames, R.B., 1992. Time allocation, in: Smith, E.A., Winterhalder, B. (Eds.), Evolutionary Ecology and Human Behavior. Aldine de Gruyter, New York, pp. 203-235.

635
636

637
638
639

Handley, L.L., Austin, A.T., Stewart, G.R., Robinson, D., Scrimgeour, C.M., Raven, J.A., Heaton, T.H.E., Schmidt, S., 1999. The ¹⁵N natural abundance (δ¹⁵N) of ecosystem samples reflects measures of water availability. Funct. Plant Biol. 26, 185-199.

640
641

642
643

Handley, L.L., Odee, D., Scrimgeour, C.M., 1994. δ¹⁵N and δ¹³C patterns in savanna vegetation: dependence on water availability and disturbance. Funct. Ecol. 8, 306-314.

Hartman, G., 2011. Are elevated δ¹⁵N values in herbivores in hot and arid environments caused by diet or animal physiology? Funct. Ecol. 25, 122-131.

644
645

646
647

Heaton, T.H.E., 1987. The ¹⁵N/¹⁴N ratios of plants in South Africa and Namibia: relationship to climate and coastal/saline environments. Oecologia 74, 236-246.

Heaton, T.H.E., Talma, A.S., Vogel, J.C., 1986. Dissolved gas paleotemperatures and ¹⁸O variations derived from groundwater near Uitenhage, South Africa. QuRes 25, 79-88.
648

Henshilwood, C., d'Errico, F., Vanhaeren, M., Van Niekerk, K., Jacobs, Z., 2004a. Middle

649

Stone Age shell beads from South Africa. *Sci* 304, 404.

650

Henshilwood, C.S., d'Errico, F., Vanhaeren, M., van Niekerk, K., Jacobs, Z., 2004b. Middle

651

Stone Age shell beads from South Africa. *Sci* 384, 404.

652

Henshilwood, C.S., d'Errico, F., Yates, R., Jacobs, Z., Tribolo, C., Duller, G.A.T., Mercier, N.,

653

Sealy, J.C., Valladas, H., Watts, I., Wintle, A.G., 2002. Emergence of modern human behavior:

654

Middle Stone Age engravings from South Africa. *Sci* 295, 1278-1280.

655

Henshilwood, C.S., van Niekerk, K.L., Wurz, S., Delagnes, A., Armitage, S.J., Rifkin, R.F.,

656

Douze, K., Keene, P., Haaland, M.M., Reynard, J., Discamps, E., Mienies, S.S., 2014. Klipdrift

657

Shelter, southern Cape, South Africa: Preliminary Report on the Howiesons Poort layers. *J.*

658

Archaeol. Sci. 45, 284-303.

659

Hogg, A.G., Hua, Q., Blackwell, P.G., Niu, M., Buck, C.E., Guilderson, T.P., Heaton, T.J.,

660

Palmer, J.G., Reimer, P.J., Reimer, R.W., Turney, C.S.M., Zimmerman, S.R.H., 2013.

661

SHCal13 Southern Hemisphere Calibration, 0–50,000 Years cal BP.

662

Kim, J.-H., Schneider, R.R., 2003. Low-latitude control of interhemispheric sea-surface

663

temperature contrast in the tropical Atlantic over the past 21 kyrs: the possible role of SE trade

664

winds. *Cldy* 23, 337-347.

665

Klein, R.G., 1972. The late Quaternary mammalian fauna of Nelson Bay Cave (Cape Province,

666

South Africa): its implications for megafaunal extinctions and environmental and cultural

667

change. *QuRes* 2, 135-142.

668

Klein, R.G., 1980. Environmental and ecological implications of large mammals from Upper

669

Pleistocene and Holocene sites in southern Africa. *Ann. S. Afr. Mus.*, 223-283.

670

Klein, R.G., 1983. Palaeoenvironmental implications of Quaternary large mammals in the

671

Fynbos region, in: Deacon, H.J., Hendey, Q.B. and Lambrechts, J.J.N. (Ed.), Fynbos

672

Palaeoecology: a preliminary synthesis, 75 ed, pp. 116-138.

673

Kohfeld, K.E., Graham, R.M., de Boer, A.M., Sime, L.C., Wolff, E.W., Le Quéré, C., Bopp,

674

L., 2013. Southern Hemisphere westerly wind changes during the Last Glacial Maximum:

675

paleo-data synthesis. *QSRv* 68, 76-95.

676

Kuhn, S.L., 1995. Mousterian Lithic Technology. Princeton University Press, Princeton.

677

Kuhn, S.L., Clark, A.E., 2015. Artifact densities and assemblage formation: Evidence from

678

Tabun Cave. *J. Anthrop. Archaeol.* 38, 8-16.

679

Levine, J.G., Yang, X., Jones, A.E., Wolff, E.W., 2014. Sea salt as an ice core proxy for past

680

sea ice extent: a process-based model study. *Journal of Geophysical Research: Atmospheres*

681

119, 5737-5756.

682

Loftus, E., Sealy, J., Lee-Thorp, J., 2016. New Radiocarbon Dates and Bayesian Models for

683

Nelson Bay Cave and Byneskranskop 1: Implications for the South African Later Stone Age

684

Sequence. *Radiocarbon FirstView*, 1-17.

685

Lombard, M., Wadley, L., Deacon, J., Wurz, S., Parsons, I., Mohapi, M., Swart, J., Mitchell,

686

P., 2012. South African and Lesotho Stone Age sequence updated. *S. Afr. Archaeol. Bull.* 67,

687

120-144.

688

Lowe, J.J., Rasmussen, S.O., Björck, S., Hoek, W.Z., Steffensen, J.P., Walker, M.J.C., Yu,

689

Z.C., 2008. Synchronisation of palaeoenvironmental events in the North Atlantic region during
690 the Last Termination: a revised protocol recommended by the INTIMATE group. QSRv 27, 6-691 17.

692 Mackay, A., 2009. History and Selection in the Late Pleistocene Archaeology of the Western 693 Cape, South Africa, School of Archaeology and Anrthopology. Australian National University.

694 Mackay, A., Marwick, B., 2011. Costs and benefits of technological decision making under 695 variable conditions: Examples from the late Pleistocene in southern Africa, in: Marwick, B., 696 Mackay, A. (Eds.), Keeping your Edge: Recent Approaches to the Organisation of Stone 697 Artefact Technology. Archaeopress, Oxford, pp. 119-134.

698 Mackay, A., Stewart, B.A., Chase, B.M., 2014. Coalescence and fragmentation in the late 699 Pleistocene archaeology of southernmost Africa. J. Hum. Evol. 72, 26-51.

700 Marean, C.W., 2010. Pinnacle Point Cave 13B (Western Cape Province, South Africa) in 701 context: The Cape Floral kingdom, shellfish, and modern human origins. J. Hum. Evol. 59, 702 425-443.

703 Marean, C.W., 2014. The origins and significance of coastal resource use in Africa and Western 704 Eurasia. J. Hum. Evol. 77, 17-40.

705 Marean, C.W., Cawthra, H.C., Cowling, R.M., Esler, K.J., Fisher, E., Milewski, A., Potts, A.J., 706 Singels, E., De Vynck, J., 2014. Stone Age people in a changing South African Greater Cape 707 Floristic Region, in: N. Allsopp, Colville, J.F., Verboom, G.A. (Eds.), Fynbos: Ecology, 708 Evolution, and Conservation of a Megadiverse Region. Oxford University Press, Oxford, UK, 709 pp. 164-199.

710 McCall, G.S., 2007. Behavioral ecological models of lithic technological change during the 711 later Middle Stone Age of South Africa. J. Archaeol. Sci. 34, 1738-1751.

712 McCall, G.S., Thomas, J.T., 2012. Still Bay and Howiesons Poort Foraging Strategies: Recent 713 Research and Models of Culture Change. Afr. Archaeol. Rev. 29, 7-50.

714 McManus, J.F., Francois, R., Gherardi, J.-M., Keigwin, L.D., Brown-Leger, S., 2004. Collapse 715 and rapid resumption of Atlantic meridional circulation linked to deglacial climate changes. 716 Nature 428, 834-837.

717 Mellars, P., 2006. Why did modern human populations disperse from Africa ca. 60,000 years 718 ago? A new model. Proc. Natl. Acad. Sci. U.S.A. 103, 9381-9386.

719 Metcalfe, D., Jones, K.T., 1988. A Reconsideration of Animal Body-Part Utility Indices. Amer. 720 Antiq. 53, 486-504.

721 Mitchell, P., 2000. The organisation of Later Stone Age lithic technology in the Caledon 722 Valley, Southern Africa. Afr. Archaeol. Rev. 17, 141-176.

723 Mitchell, P.J., 1988. The early microlithic assemblages of southern Africa. BAR, Oxford.

724 Moffett, R.O., Deacon, H.J., 1977. The Flora and Vegetation in the Surrounds of Boomplaas 725 Cave: Cango Valley. The South African Archaeological Bulletin 32, 127-145.

726 Moore, P.D., Webb, J.A., Collinson, M.E., 1991. Pollen Analysis, 2nd ed. Blackwell Scientific 727 Publications, Oxford.

728 Mucina, L., Rutherford, M.C., 2006. The vegetation of South Africa, Lesotho and Swaziland, 729 Strelitzia. South African National Biodiversity Institute, Pretoria.
730

Murphy, B.P., Bowman, D.M.J.S., 2006. Kangaroo metabolism does not cause the relationship between bone collagen δ¹⁵N and water availability. Funct. Ecol. 20, 1062-1069.

731

732

Murphy, B.P., Bowman, D.M.J.S., 2009. The carbon and nitrogen isotope composition of Australian grasses in relation to climate. Funct. Ecol. 23, 1040-1049.

733

734

Neeley, M.P., 2002. Going microlithic: A Levantine perspective on the adoption of microlithic technologies, in: Elston, R.G., Kuhn, S.L. (Eds.), Thinking Small: Global Perspectives on Microlithization.

735

736

737

Pargeter, J., 2016. Lithic miniaturization in Late Pleistocene southern Africa. Journal of Archaeological Science: Reports 10, 221-236.

738

739

Parkington, J., Cartwright, C., Cowling, R.M., Baxter, A., Meadows, M., 2000. Palaeovegetation at the Last Glacial Maximum in the Western Cape, South Africa: wood charcoal and pollen evidence from Elands Bay Cave. SAJSc 96, 543-546.

740

741

742

Parry, W.J., Kelly, R.L., 1987. Expedient core technology and sedimentism, in: Johnson, J.K., Morrow, C.A. (Eds.), The organization of core technology. Westview Press, pp. 285-304.

743

744

Partridge, T.C., Scott, L., Hamilton, J.E., 1999. Synthetic reconstructions of southern African environments during the Last Glacial Maximum (21-18 kyr) and the Holocene Altithermal (8-6 kyr). Quat Int 57-8, 207-214.

745

746

747

Partridge, T.C., Scott, L., Schneider, R.R., 2004. Between Agulhas and Benguela: responses of southern African climates of the late Pleistocene to current fluxes, orbital precession and the extent of the circum-Antarctic vortex, Past Climate Variability through Europe and Africa, pp. 45-68.

748

749

750

751

Pate, J.S., 2001. Carbon isotope discrimination and plant water-use efficiency: case scenarios for C₃ plants, in: Unkovich, M., Pate, J., McNeill, A., Gibbs, D.J. (Eds.), Stable Isotope Techniques in the Study of Biological Processes and Functioning of Ecosystems. Kluwer Academic Publishers, Dordrecht, pp. 19-37.

752

753

754

755

Petraglia, M., Clarkson, C., Boivin, N., Haslam, M., Korisettar, R., Chaubey, G., Ditchfield, P., Fuller, D., James, H., Jones, S., Kivisild, T., Koshy, J., Lahr, M.M., Metspalu, M., Roberts, R., Arnold, L., 2009. Population increase and environmental deterioration correspond with microlithic innovations in South Asia ca. 35,000 years ago. Proc Natl Acad Sci U S A 106, 12261-12266.

756

757

758

759

760

Powell, A., Shennan, S., Thomas, M.G., 2009. Late Pleistocene demography and the appearance of modern human behavior. Sci 324, 1298-1301.

761

762

Quick, L.J., Carr, A.S., Meadows, M.E., Boom, A., Bateman, M.D., Roberts, D.L., Reimer, P.J., Chase, B.M., 2015. A late Pleistocene-Holocene multi-proxy record of palaeoenvironmental change from Still Bay, southern Cape Coast, South Africa. JQS 30, 870-885.

763

764

765

766

Quick, L.J., Meadows, M.E., Bateman, M.D., Kirsten, K.L., Mäusbacher, R., Haberzettl, T., Chase, B.M., 2016. Vegetation and climate dynamics during the last glacial period in the fynbos-afrotemperate forest ecotone, southern Cape, South Africa. Quat Int 404, Part B, 136-149.

767

768

769

770

Riel-Salvatore, J., Barton, M., 2004. Late Pleistocene Technology, Economic Behavior, and Land-Use Dynamics in Southern Italy. Amer. Antiq. 69, 257-274.

771
772 Roberts, P., Henshilwood, C.S., van Niekerk, K.L., Keene, P., Gledhill, A., Reynard, J.,
773 Badenhorst, S., Lee-Thorp, J., 2016. Climate, environment and early human innovation: stable
774 isotope and faunal proxy evidence from archaeological Sites (98-59ka) in the Southern Cape,
775 South Africa. PLoS One 11, e0157408.

776 Rutherford, M.C., Mucina, L., Powrie, L.W., 2012. The South African National Vegetation
777 Database: History, development, applications, problems and future. SAJSc 108.

778 Rutherford, M.C., Powrie, L.W., Midgley, G.F., 2003. ACKDAT: A digital spatial database of
779 distributions of South African plant species and species assemblages. S. Afr. J. Bot. 69, 99-
780 104.

781 Sale, J.B., 1965. The feeding behaviour of rock hyraces (genera *Procavia* and *Heterohyrax*) in
782 Kenya. Afr. J. Ecol. 3, 1-18.

783 SANBI, 2003. PRECIS (National Herbarium Pretoria (PRE) Computerized Information
784 System) database.

785 Scholtz, A., 1986. Palynological and Palaeobotanical Studies in the Southern Cape. University
786 of Stellenbosch, Stellenbosch, South Africa.

787 Schweitzer, F.R., Wilson, M.L., 1983. Byneskranskop 1: A late Quaternary living site in the
788 southern Cape province, South Africa. The Rustica Press, Cape Town.

789 Scott, L., 1982. Late Quaternary fossil pollen grains from the Transvaal, South Africa. Rev.
790 Palaeobot. Palynol. 36, 241-278.

791 Sealy, J., Lee-Thorp, J., Loftus, E., Faith, J.T., Marean, C.W., 2016. Late Quaternary
792 environmental change in the Southern Cape, South Africa, from stable carbon and oxygen
793 isotopes in faunal tooth enamel from Boomplaas Cave. JQS 31, 919-927.

794 Sime, L.C., Kohfeld, K.E., Le Quéré, C., Wolff, E.W., de Boer, A.M., Graham, R.M., Bopp,
795 L., 2013. Southern Hemisphere westerly wind changes during the Last Glacial Maximum:
796 model-data comparison. QSRv 64, 104-120.

797 Smith, B.N., 1972. Natural abundance of the stable isotopes of carbon in biological systems.
798 BioSc 22, 226-231.

799 Stiner, M.C., Munro, N.D., Surovell, T.A., 2000. The tortoise and the hare: small-game use,
800 the broad spectrum revolution, and Paleolithic demography. Curr. Anthrop. 41, 39-73.

801 Stocker, T.F., 1998. The seesaw effect. Sci 282, 61-62.

802 Stocker, T.F., Johnsen, S.J., 2003. A minimum thermodynamic model for the bipolar seesaw.
803 PalOc 18.

804 Stockmarr, J., 1971. Tablets with spores used in absolute pollen analysis. Pollen et Spores 13,
805 614-621.

806 Stone, A.E.C., 2014. Last Glacial Maximum conditions in southern Africa: are we any closer
807 to understanding the climate of this time period? Progress in Physical Geography.

808 Stuut, J.-B.W., Crosta, X., Van der Borg, K., Schneider, R.R., 2004. On the relationship
809 between Antarctic sea ice and southwestern African climate during the late Quaternary. Geo
810 32, 909-912.

811 Talma, A.S., Vogel, J.C., 1992. Late Quaternary paleotemperatures derived from a speleothem
812 from Cango Caves, Cape Province, South Africa. QuRes 37, 203-213.
813

van Zinderen Bakker, E.M., 1953. South African pollen grains and spores. Part I. Balkema, Amsterdam/Cape Town.

814

815

van Zinderen Bakker, E.M., 1956. South African pollen grains and spores. Part II. Balkema, Amsterdam/Cape Town.

816

817

van Zinderen Bakker, E.M., Coetzee, J.A., 1959. South African pollen grains and spores. Part III. Balkema, Amsterdam/Cape Town.

818

819

Vogel, J.C., 1978. Isotopic assessment of the dietary habits of ungulates. SAJSc 74, 298-301.

820

Wadley, L., 1993. The Pleistocene Later Stone Age south of the Limpopo River. Journal of World Prehistory 7, 243-296.

821

822

Wang, L., D'Odorico, P., Ries, L., Macko, S.A., 2010. Patterns and implications of plant-soil δ¹³C and δ¹⁵N values in African savanna ecosystems. QuRes 73, 77-83.

823

824

Wilkins, J., Brown, K.S., Oestmo, S., Pereira, T., Ranhorn, K.L., Schoville, B.J., Marean, C.W., 2017. Lithic technological responses to Late Pleistocene glacial cycling at Pinnacle Point Site 5-6, South Africa. PLoS One 12, e0174051.

825

826

827

Will, M., Mackay, A., 2016. What factors govern the procurement and use of silcrete during the Stone Age of South Africa? Journal of Archaeological Science: Reports.

828

829

Wolff, E.W., Barbante, C., Becagli, S., Bigler, M., Boutron, C.F., Castellano, E., de Angelis, M., Federer, U., Fischer, H., Fundel, F., Hansson, M., Hutterli, M., Jonsell, U., Karlin, T., Kaufmann, P., Lambert, F., Littot, G.C., Mulvaney, R., Röthlisberger, R., Ruth, U., Severi, M., Siggaard-Andersen, M.L., Sime, L.C., Steffensen, J.P., Stocker, T.F., Traversi, R., Twarloh, B., Udisti, R., Wagenbach, D., Wegner, A., 2010. Changes in environment over the last 800,000 years from chemical analysis of the EPICA Dome C ice core. QSRv 29, 285-295.

830

831

832

833

834

835

836

Table Captions

837

**Table 1:** Zooarchaeological indicators of foraging efficiency at Boomplaas Cave. These include the relative abundance of tortoises (%), leporids (%), and mean food utility index (FUI) of high-survival body parts belonging to size 3-4 mammals.

838

839

840

**Table 2:** The correlation (Spearman's ρ) between lithic technological indicators and faunal community composition (DCA Axis 1 scores) for ungulates and microfauna at Boomplaas Cave. Significant values in bold.

841

842

843
<figure><img src="image_3.png" /></figure>

845

846

847

848

849

850

851

852

853

854

855

**Figure 1:** (A) Map of southern Africa showing seasonality of rainfall and climatic gradients dictated by the zones of summer/tropical (orange) and winter/temperate (blue) rainfall dominance. Winter rainfall is primarily a result of frontal systems embedded in the westerly storm track. Major atmospheric (white arrows) and oceanic (blue arrows) circulation systems and the austral summer positions of the Inter-Tropical Convergence Zone (ITCZ) and the Congo Air Boundary (CAB) are indicated. The location of the study site in the transitional southern Cape region is shown. (B) Map of southwest African coastal region with the Seweweekspoort sites and other key palaeoenvironmental and archaeological sites indicated (shading as for panel 'A'). (C) Topographical map of Seweweekspoort, with the SWP-1 and SWP-3 sites indicated.
<figure><img src="image_4.png" /></figure>

**Figure 2:** Comparison of δ¹⁵N (e), δ¹³C values (b), fynbos (c) and grass (d) pollen percentages from the Seweweekspoort hyrax middens with relevant palaeoenvironmental records including the northern summer rainfall zone wet season precipitation reconstruction (a; Chevalier and Chase, 2015), the Bermuda Rise record of Atlantic Meridional Overturning Circulation (AMOC) strength and the northward oceanic transport of heat (f; McManus et al., 2004), foraminifera records indicating conditions in the ocean to the south of the Africa (g, h; Barker et al., 2009) and sea salt sodium concentrations from the EPICA DML ice core in Antarctica (i; Fischer et al., 2007).
<figure><img src="image_5.png" /><figcaption>**Figure 3:** Comparison of δ¹⁵N (j) and δ¹³C (i) values from the Seweweekspoort hyrax middens with zooarchaeological records (a-c) of macrofauna (Faith, 2013a; Klein, 1983) and microfauna (Avery, 1982), and (d-h) lithic data (Deacon, 1979; Deacon et al., 1984) from Boomplaas Cave. Error bars on the data from Boomplaas reflect potential age ranges (2σ) of each stratigraphic unit.</figcaption></figure>
883 Supplementary Information

884 *Material and methods*

885 The Seweweekspoort middens were selected for analysis because they are composed almost
886 entirely of hyraceum (no visible pellets). Our experience suggests that such middens have
887 superior stratigraphic integrity compared to more pellet-rich middens. Representative portions
888 of the middens were processed according to the methods described by Chase et al. (Chase et
889 al., 2013; Chase et al., 2012). Radiocarbon age determinations were processed at the
890 ¹⁴CHRONO Centre, Queen’s University Belfast using accelerator mass spectrometry (AMS)
891 (Figure S1; Table S1). The radiocarbon ages were corrected for isotope fractionation using the
892 AMS measured δ¹³C and calibrated using the SHCal13 calibration data (Hogg et al., 2013).
893 The Bacon 3.0.3 software package (Blaauw and Christen, 2011) was used to generate all age-
894 depth models (Figure S1). Stable isotope analysis of bulk midden samples were performed at
895 the Department of Archaeology, University of Cape Town according to the methods described
896 in Chase et al. (2010; 2009; 2011; 2012), with a contiguous/overlapping samples obtained from
897 two series of offset 1 mm holes. For the stable isotope analyses, the standard deviation derived
898 from replicate analyses of homogeneous material was better than 0.2 ‰ for both carbon and
899 nitrogen. Carbon isotope results are expressed relative to Vienna PDB. Nitrogen isotope results
900 are expressed relative to atmospheric nitrogen.

901
<table><caption>**Table S1:** Radiocarbon ages were calibrated using the SHCal13 calibration data (Hogg et al., 2013), except for the surface sample from SWP-1-4b, which was not calibrated, and UBA-19558, which was calibrated with CALIBomb using SHCal13 and the SHZ1_2 dataset of Hua et al. (Hua et al., 2013).</caption><thead><tr><th rowspan="2">Sample</th><th rowspan="2">¹⁴C age BP</th><th rowspan="2">F¹⁴C</th><th rowspan="2">1 sigma error</th><th colspan="2">95.4 % (2σ) cal age ranges</th><th rowspan="2">relative area under distribution</th><th rowspan="2">median probability (cal BP)</th></tr><tr><th>lower cal range BP</th><th>upper cal range BP</th></tr></thead><tbody><tr><td colspan="8"><strong>SWP-1-1</strong></td></tr><tr><td rowspan="3">UBA-16698</td><td rowspan="3">3067</td><td rowspan="3">32</td><td>3078</td><td>3094</td><td>0.02947</td><td rowspan="3">3222</td></tr><tr><td>3108</td><td>3129</td><td>0.031819</td></tr><tr><td>3138</td><td>3349</td><td>0.938712</td></tr><tr><td rowspan="3">UBA-18638</td><td rowspan="3">4509</td><td rowspan="3">40</td><td>4892</td><td>4900</td><td>0.006407</td><td rowspan="3">5131</td></tr><tr><td>4915</td><td>4927</td><td>0.009555</td></tr><tr><td>4959</td><td>5299</td><td>0.984038</td></tr><tr><td rowspan="2">UBA-18639</td><td rowspan="2">5322</td><td rowspan="2">35</td><td>5940</td><td>5977</td><td>0.111729</td><td rowspan="2">6068</td></tr><tr><td>5981</td><td>6183</td><td>0.888271</td></tr><tr><td rowspan="2">UBA-18640</td><td rowspan="2">6226</td><td rowspan="2">33</td><td>6954</td><td>7178</td><td>0.982768</td><td rowspan="2">7082</td></tr><tr><td>7219</td><td>7236</td><td>0.017232</td></tr><tr><td>UBA-18641</td><td>8535</td><td>39</td><td>9447</td><td>9540</td><td>1</td><td>9502</td></tr><tr><td colspan="8"><strong>SWP-1-2a</strong></td></tr><tr><td>UBA-22927</td><td>10243</td><td>39</td><td>11721</td><td>11733</td><td>0.009155</td><td>11886</td></tr><tr><td></td><td></td><td></td><td>11748</td><td>12040</td><td>0.990845</td><td></td></tr><tr><td>UBA-18645</td><td>13621</td><td>60</td><td>16145</td><td>16620</td><td>1</td><td>16359</td></tr><tr><td>UBA-18646</td><td>17143</td><td>83</td><td>20377</td><td>20894</td><td>1</td><td>20627</td></tr><tr><td>UBA-18647</td><td>18746</td><td>95</td><td>22372</td><td>22837</td><td>1</td><td>22564</td></tr><tr><td colspan="8"><strong>SWP-1-4b top</strong></td></tr><tr><td>surface</td><td>modern</td><td>1</td><td></td><td></td><td></td><td></td></tr><tr><td>UBA-19558</td><td></td><td>1.0181</td><td>0.0043</td><td>-7</td><td>-6</td><td>1</td><td></td></tr><tr><td rowspan="2">UBA-19559</td><td rowspan="2">975</td><td rowspan="2">27</td><td>775</td><td>782</td><td>0.020244</td><td rowspan="2">853</td></tr><tr><td>787</td><td>919</td><td>0.979756</td></tr><tr><td>UBA-22921</td><td>1543</td><td>25</td><td>1314</td><td>1426</td><td>1</td><td>1378</td></tr><tr><td rowspan="2">UBA-22922</td><td rowspan="2">2238</td><td rowspan="2">25</td><td>2117</td><td>2123</td><td>0.008228</td><td rowspan="2">2231</td></tr><tr><td>2148</td><td>2319</td><td>0.991772</td></tr><tr><td colspan="8"><strong>SWP-1-4b bottom</strong></td></tr><tr><td>UBA-19560</td><td>11138</td><td>47</td><td>12811</td><td>13076</td><td>1</td><td>12961</td></tr><tr><td>UBA-22924</td><td>12190</td><td>41</td><td>13836</td><td>14166</td><td>1</td><td>14030</td></tr><tr><td>UBA-19561</td><td>12626</td><td>55</td><td>14547</td><td>15185</td><td>1</td><td>14923</td></tr></tbody></table>
<table><tr><td>UBA-22925</td><td>12618</td><td>46</td><td>14586</td><td>15164</td><td>1</td><td>14917</td></tr><tr><td>UBA-19562</td><td>13425</td><td>67</td><td>15855</td><td>16313</td><td>1</td><td>16102</td></tr></table>

SWP-1-5

<table><tr><td>UBA-19565</td><td>410</td><td>21</td><td>328</td><td>378</td><td>0.344762</td><td>456</td></tr><tr><td></td><td></td><td></td><td>391</td><td>401</td><td>0.018871</td><td></td></tr><tr><td></td><td></td><td></td><td>439</td><td>497</td><td>0.636367</td><td></td></tr><tr><td>UBA-22742</td><td>1657</td><td>35</td><td>1415</td><td>1582</td><td>1</td><td>1504</td></tr><tr><td>UBA-22743</td><td>3428</td><td>28</td><td>3512</td><td>3528</td><td>0.013505</td><td>3629</td></tr><tr><td></td><td></td><td></td><td>3558</td><td>3719</td><td>0.986495</td><td></td></tr><tr><td>UBA-22744</td><td>4973</td><td>29</td><td>5596</td><td>5728</td><td>1</td><td>5651</td></tr><tr><td>UBA-22745</td><td>6216</td><td>29</td><td>6959</td><td>7168</td><td>1</td><td>7077</td></tr><tr><td>UBA-22746</td><td>7559</td><td>32</td><td>8206</td><td>8263</td><td>0.154807</td><td>8348</td></tr><tr><td></td><td></td><td></td><td>8290</td><td>8403</td><td>0.845193</td><td></td></tr><tr><td>UBA-22747</td><td>8292</td><td>35</td><td>9035</td><td>9047</td><td>0.011584</td><td>9224</td></tr><tr><td></td><td></td><td></td><td>9088</td><td>9329</td><td>0.882282</td><td></td></tr><tr><td></td><td></td><td></td><td>9342</td><td>9403</td><td>0.106134</td><td></td></tr><tr><td>UBA-22748</td><td>8609</td><td>35</td><td>9480</td><td>9564</td><td>0.955713</td><td>9535</td></tr><tr><td></td><td></td><td></td><td>9567</td><td>9597</td><td>0.044287</td><td></td></tr><tr><td>UBA-19566</td><td>9341</td><td>42</td><td>10296</td><td>10360</td><td>0.074017</td><td>10496</td></tr><tr><td></td><td></td><td></td><td>10366</td><td>10597</td><td>0.897515</td><td></td></tr><tr><td></td><td></td><td></td><td>10624</td><td>10650</td><td>0.028468</td><td></td></tr></table>

SWP-3-1

<table><tr><td>UBA-19320</td><td>3405</td><td>32</td><td>3482</td><td>3535</td><td>0.122002</td><td>3604</td></tr><tr><td></td><td></td><td></td><td>3547</td><td>3694</td><td>0.877998</td><td></td></tr><tr><td>UBA-19321</td><td>3509</td><td>40</td><td>3612</td><td>3848</td><td>1</td><td>3741</td></tr><tr><td>UBA-19322</td><td>4661</td><td>65</td><td>5051</td><td>5191</td><td>0.201609</td><td>5354</td></tr><tr><td></td><td></td><td></td><td>5213</td><td>5481</td><td>0.745676</td><td></td></tr><tr><td></td><td></td><td></td><td>5532</td><td>5578</td><td>0.052715</td><td></td></tr><tr><td>UBA-19323</td><td>5454</td><td>46</td><td>6014</td><td>6081</td><td>0.097688</td><td>6221</td></tr><tr><td></td><td></td><td></td><td>6104</td><td>6158</td><td>0.115457</td><td></td></tr><tr><td></td><td></td><td></td><td>6171</td><td>6301</td><td>0.786855</td><td></td></tr><tr><td>UBA-21381</td><td>6212</td><td>30</td><td>6956</td><td>7167</td><td>1</td><td>7074</td></tr><tr><td>UBA-21382</td><td>7840</td><td>32</td><td>8455</td><td>8502</td><td>0.081029</td><td>8574</td></tr><tr><td></td><td></td><td></td><td>8506</td><td>8642</td><td>0.918971</td><td></td></tr><tr><td>UBA-19324</td><td>9354</td><td>52</td><td>10296</td><td>10359</td><td>0.069114</td><td>10512</td></tr><tr><td></td><td></td><td></td><td>10367</td><td>10668</td><td>0.930886</td><td></td></tr><tr><td>UBA-19325</td><td>10717</td><td>52</td><td>12553</td><td>12718</td><td>1</td><td>12651</td></tr><tr><td>UBA-19326</td><td>12874</td><td>84</td><td>15060</td><td>15648</td><td>1</td><td>15310</td></tr></table>
<figure><img src="image_6.png" /></figure>

**Figure S1:** Stable nitrogen isotope results from the Seweweekspoort rock hyrax midden considered in this study with associated age models. Age-depth models were generated using the Bacon 3.0.3 software package (Blaauw and Christen, 2011).
<figure><img src="image_7.png" /></figure>

**Figure S2:** Stable nitrogen isotope results from the Seweweekspoort rock hyrax midden shown individually in relation to the aggregate record obtained using Local Regression (LOESS) curve fitting of the combined datasets. For comparability, 1.5‰ was added to the δ¹⁵N data from the SWP-3-1 to compensate for the more humid microclimate, relative to the SWP-1 site, in which the midden was found.
<figure><img src="image_8.png" /></figure>

**Figure S3:** Stable carbon isotope results from the Seweweekspoort rock hyrax midden shown individually in relation to the aggregate record obtained using Local Regression (LOESS) curve fitting of the combined datasets.
Taxonomic abundances of ungulates from Boomplaas Cave were derived from Faith (Faith, 2013), with Klein's (Klein, 1983) data used for the uppermost member DGL. Comparable data for the Boomplaas microfauna were derived from Avery (Avery, 1982). We conducted a detrended correspondence analysis (DCA) for both data sets, using the first axis (rescaled from 0 to 100) to broadly summarize faunal composition (Figure 3). The linear trends in both taxonomic groups closely parallel a decline in ungulate grazers (Figure 3), suggesting that the DCA Axis 1 scores are related to the replacement of grasslands by more shrubby habitats (see also Faith, 2013). Parallel changes are also observed at Nelson Bay Cave (Klein, 1983) and Byneskranskop 1 (Schweitzer and Wilson, 1983), implying regional shifts in habitat structure.

Zooarchaeological indicators of foraging efficiency at Boomplaas Cave (Table S2), including the relative abundance of small-bodied prey (leporids and tortoises) and the average utility of high-survival body parts (following the taphonomic model of bone survivorship outlined in Cleghorn and Marean (Cleghorn and Marean, 2004)), were derived from Faith (Faith, 2011). Despite small body size, because tortoises are slow-moving and easy to capture, they might be considered a high-ranked prey type that should be collected whenever encountered (e.g. Stiner et al., 2000). While this makes tortoises a somewhat ambiguous indicator, the parallel increase in leporids – unequivocally a low-ranked prey type – across the LGIT more definitively suggests a decline in foraging efficiency that parallels changing body part representation for large-bodied prey.

<table><caption>**Table S2.** Zooarchaeological indicators of foraging efficiency at Boomplaas Cave. These include the relative abundance of tortoises (%), leporids (%), and mean food utility index of high-survival body parts belonging to size 3-4 mammals (FUI).</caption><thead><tr><th>Stratum</th><th>% Tortoises</th><th>% Leporids</th><th>FUI</th></tr></thead><tbody><tr><td>BLD3</td><td>74.6</td><td>30.9</td><td>1405</td></tr><tr><td>FBL/BLA</td><td>79.1</td><td>28.7</td><td>1737</td></tr><tr><td>BRL/BRL1</td><td>61.8</td><td>10.4</td><td>1465</td></tr><tr><td>BRL2</td><td>71.7</td><td>13.5</td><td>1910</td></tr><tr><td>BRL3</td><td>78.1</td><td>24.2</td><td>1610</td></tr><tr><td>BRL4</td><td>74.1</td><td>24.2</td><td>2245</td></tr><tr><td>BRL5</td><td>80.3</td><td>36.2</td><td>2432</td></tr><tr><td>BRL6</td><td>68.2</td><td>15.2</td><td>2246</td></tr><tr><td>BRL7</td><td>70.9</td><td>22.3</td><td>2188</td></tr><tr><td>CL1</td><td>68.9</td><td>8.5</td><td>2028</td></tr><tr><td>CL2</td><td>71.9</td><td>3.6</td><td>2037</td></tr></tbody></table>
<table><tr><td>CL3</td><td>47.1</td><td>4.4</td><td>1840</td></tr><tr><td>CL4</td><td>54.5</td><td>0.0</td><td>1458</td></tr></table>

932

933

934

935

936

937

938

939

940

Ages for Boomplaas follow Deacon (1982), calibrated using SHCal13 (Hogg et al., 2013). All lithic data are taken from Deacon (1982), standardised to site mean (standardised value = (layer value – site mean) / site standard deviation). We use core to tool ratios to track changes in the way in which technology was delivered; alternative measures such as core to flake ratios may increase or decrease with changing intensity of core reduction independent of delivery systems. All the measures of technological change we employ for the Boomplaas assemblage are significantly correlated with changes in ungulate community composition, and most with the composition of micromammal communities (Table S3).

941

942

943

944

945

946

947

948

949

950

951

As with the faunal data, the technological trends noted for Boomplaas are replicated at Byneskranskop (Schweitzer and Wilson, 1983) (Table S4) and Nelson Bay Cave (Deacon, 1982) (Table S5) (methods as per Boomplaas), with the exception of changes in artefact density. Given the coastal location of these two sites, changes in artefact density – effectively number of artefacts per unit sediment – are likely to have been influenced by changes in sedimentation associated with sea level fluctuations (Chase and Thomas, 2007). Microlithic technologies in the forms of bladelets do reappear at Byneskranskop and Nelson Bay Cave during the more arid conditions of the mid-Holocene, along with backed artefacts, reinforcing arguments for the diversity of contexts in microlithic systems were deployed (Hiscock et al., 2011).
<table><caption>**Table S3:** The correlation between lithic technological indicators and faunal community composition (DCA Axis 1 scores) for ungulates and microfauna at Boomplaas Cave. Significant values in bold.</caption><thead><tr><th rowspan="2"></th><th colspan="2">Ungulates</th><th colspan="2">Microfauna</th></tr><tr><th>r<sub>s</sub></th><th>p</th><th>r<sub>s</sub></th><th>p</th></tr></thead><tbody><tr><td>%Bladelets</td><td><strong>-0.696</strong></td><td><strong>0.004</strong></td><td><strong>0.878</strong></td><td><strong>&lt;0.001</strong></td></tr><tr><td>Core/tool</td><td><strong>-0.842</strong></td><td><strong>&lt;0.001</strong></td><td><strong>0.680</strong></td><td><strong>0.015</strong></td></tr><tr><td>Tool/aft</td><td><strong>0.732</strong></td><td><strong>0.002</strong></td><td><strong>-0.674</strong></td><td><strong>0.012</strong></td></tr><tr><td>CCS+Silcrete</td><td><strong>-0.604</strong></td><td><strong>0.017</strong></td><td>0.484</td><td>0.094</td></tr><tr><td>Density</td><td><strong>-0.618</strong></td><td><strong>0.014</strong></td><td>0.355</td><td>0.234</td></tr></tbody></table>

<table><caption>**Table S4:** Technological measures of the assemblage from Byneskranskop. Ages following (Loftus et al., 2016).</caption><thead><tr><th>Layer</th><th>Age (cal yr BP)</th><th>Core/tool</th><th>Tool/aft</th><th>CCS+Silcrete</th><th>Density</th><th>Bladelets</th></tr></thead><tbody><tr><td>1</td><td>1715-1870</td><td>-0.568</td><td>-0.232</td><td>-0.954</td><td>-0.834</td><td>-1.084</td></tr><tr><td>2</td><td>3450-3818</td><td>-0.616</td><td>1.608</td><td>-0.142</td><td>-1.253</td><td>-0.868</td></tr><tr><td>3</td><td></td><td>-0.589</td><td>1.579</td><td>0.325</td><td>-0.581</td><td>0.740</td></tr><tr><td>4</td><td>3720-3970</td><td>-0.605</td><td>1.380</td><td>0.387</td><td>-0.682</td><td>-0.696</td></tr><tr><td>5</td><td>6015-6290</td><td>-0.665</td><td>1.591</td><td>1.947</td><td>0.031</td><td>0.845</td></tr><tr><td>6</td><td>6315-6495</td><td>-0.617</td><td>1.223</td><td>1.542</td><td>0.090</td><td>1.119</td></tr><tr><td>7</td><td>5905-6180</td><td>-0.564</td><td>0.650</td><td>0.828</td><td>0.218</td><td>1.665</td></tr><tr><td>8</td><td>6285-6410</td><td>-0.578</td><td>0.314</td><td>1.339</td><td>1.017</td><td>0.540</td></tr><tr><td>9</td><td>6740-6945</td><td>-0.593</td><td>0.162</td><td>0.845</td><td>0.634</td><td>0.752</td></tr><tr><td>10</td><td></td><td>-0.511</td><td>-0.350</td><td>0.537</td><td>0.194</td><td>-0.475</td></tr><tr><td>11</td><td>6505-6740</td><td>-0.522</td><td>-0.708</td><td>-0.010</td><td>2.304</td><td>1.288</td></tr><tr><td>12</td><td>8342-8725</td><td>-0.457</td><td>-0.509</td><td>0.413</td><td>1.936</td><td>-0.548</td></tr><tr><td>13</td><td></td><td>-0.196</td><td>-0.711</td><td>-0.557</td><td>-0.448</td><td>-0.514</td></tr><tr><td>14</td><td>10796-11245</td><td>0.053</td><td>-0.988</td><td>-1.174</td><td>-0.939</td><td>-1.372</td></tr><tr><td>15</td><td>11245-11695</td><td>1.926</td><td>-1.004</td><td>-1.439</td><td>-1.315</td><td>-1.359</td></tr><tr><td>16</td><td></td><td>0.963</td><td>-1.002</td><td>-1.359</td><td>-1.013</td><td>-1.046</td></tr><tr><td>17</td><td>13855-14320</td><td>0.168</td><td>-0.992</td><td>-0.689</td><td>-0.728</td><td>0.290</td></tr><tr><td>18</td><td></td><td>0.886</td><td>-0.986</td><td>-1.254</td><td>0.534</td><td>-0.827</td></tr><tr><td>19</td><td>16060-16535<br>16550-17105</td><td>3.086</td><td>-1.025</td><td>-0.583</td><td>0.835</td><td>1.549</td></tr></tbody></table>

<caption>**Table S5:** Technological measures of the assemblage from Nelson Bay Cave. Ages following (Loftus et al., 2016).</caption>
<table><thead><tr><th>Layer</th><th>Age (cal yr BP)</th><th>Core/tool</th><th>Tool/aft</th><th>CCS+Silcrete</th><th>Density</th><th>Bladelets</th></tr></thead><tbody><tr><td>Ivan</td><td>5330-5655<br>5590-5730</td><td>-1.037</td><td>3.073</td><td>1.597</td><td>1.370</td><td>-0.369</td></tr><tr><td>BSC</td><td></td><td>-0.891</td><td>0.163</td><td>-0.034</td><td>1.920</td><td>-0.499</td></tr><tr><td>Rice A</td><td>9030-9400</td><td>-0.849</td><td>0.006</td><td>-1.443</td><td>-0.611</td><td>-0.650</td></tr><tr><td>Rice B</td><td>9305-9520<br>9460-9545</td><td>-0.610</td><td>-0.039</td><td>-1.404</td><td>-0.707</td><td>-0.636</td></tr><tr><td>Jake</td><td>10275-10590</td><td>0.082</td><td>-0.301</td><td>-1.536</td><td>-0.541</td><td>-0.650</td></tr><tr><td>BSBJ</td><td>11405-11975</td><td>0.260</td><td>-0.476</td><td>-1.572</td><td>-0.576</td><td>-0.634</td></tr><tr><td>CS</td><td>11825-12400</td><td>2.730</td><td>-0.590</td><td>-1.295</td><td>-0.742</td><td>-0.612</td></tr><tr><td>BSL</td><td>12020-12515<br>13775-14135</td><td>-0.501</td><td>-0.334</td><td>1.464</td><td>-0.532</td><td>0.000</td></tr><tr><td>GSL</td><td></td><td>0.133</td><td>-0.525</td><td>1.266</td><td>-0.794</td><td>0.440</td></tr><tr><td>YSL</td><td>17645-18050</td><td>0.412</td><td>-0.499</td><td>3.105</td><td>1.213</td><td>2.775</td></tr><tr><td>YGL</td><td>21960-22485<br>22615-23355</td><td>0.271</td><td>-0.477</td><td>-0.150</td><td>1.370</td><td>0.834</td></tr></tbody></table>
956 References

957 Avery, D.M., 1982. Micromammals as palaeoenvironmental indicators and an interpretation of
958 the late Quaternary in the southern Cape Province, South Africa. Annals of the South African
959 Museum 85, 183-377.

960 Blaauw, M., Christen, J.A., 2011. Flexible Paleoclimate Age-Depth Models Using an
961 Autoregressive Gamma Process. Bayesian Analysis 6, 457-474.

962 Chase, B.M., Boom, A., Carr, A.S., Meadows, M.E., Reimer, P.J., 2013. Holocene climate
963 change in southernmost South Africa: rock hyrax middens record shifts in the southern
964 westerlies. Quaternary Science Reviews 82, 199-205.

965 Chase, B.M., Meadows, M.E., Carr, A.S., Reimer, P.J., 2010. Evidence for progressive
966 Holocene aridification in southern Africa recorded in Namibian hyrax middens: implications
967 for African Monsoon dynamics and the "African Humid Period". Quaternary Research 74, 36-
968 45.

969 Chase, B.M., Meadows, M.E., Scott, L., Thomas, D.S.G., Marais, E., Sealy, J., Reimer, P.J.,
970 2009. A record of rapid Holocene climate change preserved in hyrax middens from
971 southwestern Africa. Geology 37, 703-706.

972 Chase, B.M., Quick, L.J., Meadows, M.E., Scott, L., Thomas, D.S.G., Reimer, P.J., 2011. Late
973 glacial interhemispheric climate dynamics revealed in South African hyrax middens. Geology
974 39, 19-22.

975 Chase, B.M., Scott, L., Meadows, M.E., Gil-Romera, G., Boom, A., Carr, A.S., Reimer, P.J.,
976 Truc, L., Valsecchi, V., Quick, L.J., 2012. Rock hyrax middens: A palaeoenvironmental
977 archive for southern African drylands. Quaternary Science Reviews 56, 107-125.

978 Chase, B.M., Thomas, D.S.G., 2007. Multiphase late Quaternary aeolian sediment
979 accumulation in western South Africa: Timing and relationship to palaeoclimatic changes
980 inferred from the marine record. Quaternary International 166, 29-41.

981 Cleghorn, N., Marean, C.W., 2004. Distinguishing selective transport and in situ attrition: a
critical review of analytical approaches. Journal of Taphonomy 2, 43-67.

983 Deacon, J., 1982. The Later Stone Age in the southern Cape, South Africa. University of
984 Stellenbosch.

985 Faith, J.T., 2011. Late Quaternary Megafaunal Extinctions in Southern Africa's Cape Floral
986 Region, Department of Anthropology. The George Washington University.

987 Faith, J.T., 2013. Taphonomic and paleoecological change in the large mammal sequence from
988 Boomplaas Cave, Western Cape, South Africa. Journal of Human Evolution 65, 715-730.

989 Hiscock, P., Clarkson, C., Mackay, A., 2011. Big debates over little tools: ongoing disputes
990 over microliths on three continents. World Archaeology 43, 653-664.

991 Hogg, A.G., Hua, Q., Blackwell, P.G., Niu, M., Buck, C.E., Guilderson, T.P., Heaton, T.J.,
992 Palmer, J.G., Reimer, P.J., Reimer, R.W., Turney, C.S.M., Zimmerman, S.R.H., 2013.
993 SHCal13 Southern Hemisphere Calibration, 0-50,000 Years cal BP.

994 Hua, Q., Barbetti, M., Rakowski, A.Z., 2013. Atmospheric radiocarbon for the period 1950-
995 2010. Radiocarbon 55.
996

Klein, R.G., 1983. Palaeoenvironmental implications of Quaternary large mammals in the

997

Fynbos region, in: Deacon, H.J., Hendey, Q.B. and Lambrechts, J.J.N. (Ed.), Fynbos

998

Palaeoecology: a preliminary synthesis, 75 ed, pp. 116-138.

999

Loftus, E., Sealy, J., Lee-Thorp, J., 2016. New Radiocarbon Dates and Bayesian Models for

1000

Nelson Bay Cave and Byneskranskop 1: Implications for the South African Later Stone Age

1001

Sequence. Radiocarbon 58, 365-381.

1002

Schweitzer, F.R., Wilson, M.L., 1983. Byneskranskop 1: A late Quaternary living site in the

1003

southern Cape province, South Africa. The Rustica Press, Cape Town.

1004

Stiner, M.C., Munro, N.D., Surovell, T.A., 2000. The tortoise and the hare: small-game use,

1005

the broad spectrum revolution, and Paleolithic demography. Current Anthropology 41, 39-73.

1006

1007