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Abstract

Middle Stone Age cooking Early evidence of cooked starchy plant food is sparse, yet the consumption of starchy roots is likely to have been a key innovation in the human diet. Wadley et al. report the identification of whole, charred rhizomes of plants of the genus Hypoxis from Border Cave, South Africa, dated up to 170,000 years ago. These archaeobotanical remains represent the earliest direct evidence for the cooking of underground storage organs. The edible Hypoxis rhizomes appear to have been cooked and consumed in the cave by the Middle Stone Age humans at the site. Hypoxis has a wide geographical distribution, suggesting that the rhizomes could have been a ready and reliable carbohydrate source for Homo sapiens in Africa, perhaps facilitating the mobility of human populations. Science , this issue p. 87

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... The cultural material signals that can thus be obtained from the study of plants are broad, and the study of botanical remains from the southern African archaeological record has been crucial in reconstructing many of the daily activities performed by past hunter-gatherer societies (e.g. Deacon, 1993;d'Errico et al., 2012;Bentsen, 2014;Wadley et al., 2020aWadley et al., , 2020b. ...
... Archaeobotanical evidence has also been used to propose complex cultural behaviours of South African Stone Age populations (Deacon, 1993;Wadley et al., 2011Wadley et al., , 2020aWadley et al., , 2020b, such as during the post-Howiesons Poort (post-HP) of the Middle Stone Age (MSA). The post-HP dates from around 60 ka to 25 ka years ago in southern Africa (Wadley, 2015). ...
... It is plausible that other grass subfamilies were collected but were unrecognised. The high representation of graminoids and monocots at Border Cave might be mostly due to their use by past inhabitants for a variety of reasons such as bedding construction (Backwell et al., 2018;Wadley et al., 2020a;Sievers et al., 2022). However, based on the practices of modern peoples of South Africa (van Wyk and Gericke, 2000;Gebashe et al., 2019;Nortje and van Wyk, 2019), grasses and other graminoids could have been used at Border Cave for other reasons also, such as for crafts (basketry), construction (e.g. ...
Article
Border Cave is a well-known South African Middle and Early Later Stone Age site located in KwaZulu-Natal. The site has exceptional plant preservation, unparalleled in the African Middle Stone Age archaeological record. This study focuses on the phytolith and FTIR analysis of two Members (2 BS and 2 WA) of the under-documented post-Howiesons Poort occupations dating to~60 ka. These members contain complex successions of vertically overlapping, interdigitating light brown sediments, plant bedding and combustion features of various sizes. The complexity and distinctiveness of these deposits provide an excellent opportunity for the study of plant exploitation strategies and their associated human behaviour. Our taphonomic assessment inferred, through the variability of phytolith properties and minerals composing archaeological layers, that specific occupations suffered more physical weathering than others, for example in the form of trampling. The preservation of fragile and highly soluble phy-toliths (eudicot leaf phytoliths) and the high frequencies of articulated phytoliths indicates that some bedding deposits experienced little disturbance after their deposition. Not all bedding layers dating to ⁓60 ka show, from a phytolith perspective, the same plant composition, which could be explained in terms of changes in human preference for the use of plants over time to construct bedding or because distinct types of living floors are represented. Finally, the systematic application of phytoliths and FTIR to the complex archaeological sequence of Border Cave confirm these analyses can be used in the future to identify bedding deposits not visible to the naked eye, and behavioural patterns obscured by diagenetic or biased processes during sampling.
... The cultural material signals that can thus be obtained from the study of plants are broad, and the study of botanical remains from the southern African archaeological record has been crucial in reconstructing many of the daily activities performed by past hunter-gatherer societies (e.g. Deacon, 1993;d'Errico et al., 2012;Bentsen, 2014;Wadley et al., 2020aWadley et al., , 2020b. ...
... Archaeobotanical evidence has also been used to propose complex cultural behaviours of South African Stone Age populations (Deacon, 1993;Wadley et al., 2011Wadley et al., , 2020aWadley et al., , 2020b, such as during the post-Howiesons Poort (post-HP) of the Middle Stone Age (MSA). The post-HP dates from around 60 ka to 25 ka years ago in southern Africa (Wadley, 2015). ...
... It is plausible that other grass subfamilies were collected but were unrecognised. The high representation of graminoids and monocots at Border Cave might be mostly due to their use by past inhabitants for a variety of reasons such as bedding construction (Backwell et al., 2018;Wadley et al., 2020a;Sievers et al., 2022). However, based on the practices of modern peoples of South Africa (van Wyk and Gericke, 2000;Gebashe et al., 2019;Nortje and van Wyk, 2019), grasses and other graminoids could have been used at Border Cave for other reasons also, such as for crafts (basketry), construction (e.g. ...
Article
Border Cave is a well-known South African Middle and Early Later Stone Age site located in KwaZulu-Natal. The site has exceptional plant preservation, unparalleled in the African Middle Stone Age archaeological record. This study focuses on the phytolith and FTIR analysis of two Members (2 BS and 2 WA) of the under-documented post-Howiesons Poort occupations dating to~60 ka. These members contain complex successions of vertically overlapping, interdigitating light brown sediments, plant bedding and combustion features of various sizes. The complexity and distinctiveness of these deposits provide an excellent opportunity for the study of plant exploitation strategies and their associated human behaviour. Our taphonomic assessment inferred, through the variability of phytolith properties and minerals composing archaeological layers, that specific occupations suffered more physical weathering than others, for example in the form of trampling. The preservation of fragile and highly soluble phy-toliths (eudicot leaf phytoliths) and the high frequencies of articulated phytoliths indicates that some bedding deposits experienced little disturbance after their deposition. Not all bedding layers dating to ⁓60 ka show, from a phytolith perspective, the same plant composition, which could be explained in terms of changes in human preference for the use of plants over time to construct bedding or because distinct types of living floors are represented. Finally, the systematic application of phytoliths and FTIR to the complex archaeological sequence of Border Cave confirm these analyses can be used in the future to identify bedding deposits not visible to the naked eye, and behavioural patterns obscured by diagenetic or biased processes during sampling.
... Accumulation and preservation of various types of biological material in the deposits is influenced by many regional and local factors including human activity and variations in temperature, wind, or rainfall patterns, which are controlled by changes in the regional palaeoclimate, soil conditions and hydrological changes. Past local conditions at Border Cave are reflected by numerous finds that include wood (charcoal), grass remains (bedding), phytoliths, seeds, and mammalian macro-and microfauna, which are indicative of the different vegetation communities that occupied the Lebombo escarpment slopes around the cave (Klein, 1977;Beaumont, 1978;Avery, 1982Avery, , 1992Backwell et al., 2018;Wadley et al., 2020aWadley et al., , 2020bZwane and Bamford, 2021;Esteban et al., 2022;Lennox et al., 2022;Sievers et al., 2022). ...
... I. MD69-2048 d 13 C (‰) (Castañeda et al., 2016) 2.1. Border cave local environmental history A regional palaeoclimatic overview that can be interpreted from the available biological materials provides a broad picture of a Savanna Biome woodland environment that showed changes over time ( Fig. S1) (Backwell et al., 2018;Wadley et al., 2020aWadley et al., , 2020bZwane and Bamford, 2021;Sievers et al., 2022;Klein, 1977;Avery, 1982Avery, , 1992. The wood carried into the cave supports the contention that the Savanna Biome existed throughout most of the period of occupation preserved in the sediment sequence (Zwane and Bamford, 2021;Lennox et al., 2022). ...
... Since wood of miombo vegetation, e.g., Brachystegia spiciformis, has many uses including as firewood, the lack of miombo tree wood might indeed signal absence of the taxon (Oyen and Louppe, 2012). Several species of seeds and bedding material provide further evidence of plants that grew in the vicinity of the cave but do not necessarily point to climatic or vegetation changes as they were probably selectively imported into the cave (Wadley et al., 2020a(Wadley et al., , 2020bSievers et al., 2022). ...
Article
As a result of selective anthropogenic accumulation of plant and faunal remains in the sedimentary record at Border Cave, palaeoclimatological records at the site can only be broadly interpreted and cannot be reconstructed with any precision. To aid environmental reconstructions spanning the sedimentary record, we review published climate change proxy records from both marine and terrestrial archives within 500 km in the surrounding the summer rainfall region of the site to derive the history of environmental change. These published records, which show supporting evidence between the marine and terrestrial sequences, of which the former is more continuous, suggest that frequent environmental changes on a millennial scale involved fluctuations in temperature, seasonality, and moisture conditions to which the human occupants of the cave must have had to adapt continuously, e.g., cooler, windy climates, and more open vegetation during Marine Isotope Stage 2. Pollen previously extracted from the Border Cave deposits showed limited potential for climate reconstruction, but further research and comprehensive sampling of the sedimentary succession may potentially provide additional information on local environments to complement the existing palaeobotanical data.
... Border Cave records all of these lines of evidence in older and younger deposits. With an intermittent Stone Age sequence that spans~227 thousand years ago (ka) to 24 ka Beaumont et al., 1978;Beaumont, 1980;Vogel et al., 1986;Miller and Beaumont, 1989;Miller et al., 1999;Beaumont et al., 1992;Grün and Beaumont, 2001;Bird et al., 2003;Grün et al., 2003;Millard, 2006;d'Errico et al., 2012a;Villa et al., 2012;Backwell et al., 2018), the site has recently added entries to the list of criteria that can be considered as evidence of behavioural complexity: grass and ash bedding construction at~200 ka (Wadley et al., 2020a) and transport and sharing of cooked starchy rhizomes at 170 ka (Wadley et al., 2020b). Our research at the site supplements previous work carried out since 1934 by Dart, Malan, Beaumont and their colleagues. ...
... Charcoal from Member 1 RGBS shows that Tarchonanthus sp. was the most abundant taxon collected at the site at 74 ka, possibly for medicinal purposes, followed by Euphorbia species that were perhaps collected for their latex (Zwane and Bamford, 2021). Nine species identified by these authors have known medicinal properties (e.g., Tarchonanthus sp., Euphorbia sp., Fifty-five whole charred underground storage organs, identified as Hypoxis angustifolia, were described (Wadley et al., 2020b). Forty-four come from 4 WA and 11 come from 5 BS, making them the oldest known examples of cooked starchy rhizomes at~170 thousand years old. ...
... Bedding at Border Cave and Sibudu is occasionally burnt, and while it may have been accidental, it may also have been intentional, as a means of ridding the living area of pests (Goldberg et al., 2009;Wadley et al., 2011). The discovery of 55 whole cooked Hypoxis angustifolia rhizomes in layers aged 170 ka at Border Cave (Wadley et al., 2020b) provides the earliest known evidence for the consumption of underground storage organs. Only three other MSA sites have yielded geophytes: Strathalan Cave B (Opperman and Heydenrych, 1990), Boomplaas and Klasies River (Deacon, 1995). ...
Article
In 2015, which marked 35 years since Beaumont had worked at the site, we renewed excavations at Border Cave. Our primary aims were to reassess the stratigraphic context of the sedimentary and cultural sequence, gain insight into site formation processes, make a detailed study of organic remains, identify long term cultural trends, and characterize expressions of complex behaviour and innovation. This contribution serves as an update on activities conducted in 2018 and 2019 and provides an overview of our research findings to date, placing them in the broader context of the Middle Stone Age in southern Africa. New luminescence ages based on feldspar grains in the sedimentary sequence are in broad agreement with the previous chronology established for the site. Geoarchaeology and faunal taphonomy have started to elucidate site formation processes, showing that the members should not be considered as homogeneous units, and that associated formation interpretations established by Beaumont are simplifications that are not representative of the diverse site formation processes active in the shelter. This finding is supported by lithic analysis of the Member 2 WA assemblage that shows differences in technology between artefacts from the top, middle, and lower part of the same member. In addition, the lithic artefacts from the middle and lower part of Member 2 WA show continuities with the lithics from the underlying Members 3 BS and 1 RGBS, which were attributed by Beaumont to a different industry. Grass mats/bedding layers are preserved throughout the sequence, the oldest of which dates to~200 ka. The use of ash and leaves with insecticidal properties in the bedding construction reflects complex cognition, as does the cooking of starchy rhizomes that come from layers dated to 170 ka. In addition to a rich mammal fauna found in all of the deposits, the remains of a new individual, a 3e4-year-old child, were recovered from Member 1 BS.LR C that has an ESR date of 42.6 ka.
... One long-considered exception to this is Border Cave, in the Lebombo mountains overlooking Eswatini, with much older dates associated with a characteristic LSA lithic assemblage [~43 kya BP], prompting debate on whether it represents a singular origin of the LSA, or if multi-regional or multi-temporal hypotheses can be considered 68,69 . More recently, cultural and technological innovations typical of the LSA have been found in clear association with MSA-aged deposits at other sites including, but not limited to, Blombos Cave, Diepkloof, and Sibhudu 68,[70][71][72][73] . Similarly, our research on lithics and ochre has shown evidence consistent with a more complex view on the timing and geographic distribution of this transition. ...
... The procedure bracketed ten unknown samples with a set of standards and quality control samples at the beginning and end of each to monitor instrument stability and drift throughout analytical runs (approximately every 30 mins). The ablated sample vapor was transported to the ICP-MS using He carrier gas and mixed with argon gas at the plasma torch, where the sample was ionized and passed through two detectors that measured the signal intensity in counts per second for 60 isotopes: 7 Li, 9 Be, 11 B, 23 Na, 24 Mg, 27 Al, 29 Si, 31 P, 34 S, 35 Cl, 39 K, 44 Ca, 45 Sc, 47 Ti, 51 V, 52 Cr, 55 Mn, 57 Fe, 59 Co, 60 N, 63 Cu, 66 Zn, 71 Ga, 75 As, 77 The results of trace element analyses were analyzed using multivariate statistics, detailed in Supplementary Notes 4.0 and 5.0. ...
Article
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Our species and other hominins have used earth mineral pigments since at least ~500,000 years ago, if not earlier. Its preservation and ubiquity within archaeological records across sub-Saharan Africa are well documented, but regional-scale networks of mineral selection, mining, transport, and use is an underdeveloped field. Here, we present a framework for interpreting regional variations within an overarching ochre-behavioral community of practice. Deep-time records of ochre provisioning span the final Middle Stone Age and Late Stone Age in modern day Eswatini, revealing longstanding cultural continuities in the intergenerational transmission of shared knowledge on landscapes, geology, and the desired physicochemical properties of mineral pigments. These communities of practice did not develop in isolation, and were part of a wider system of relations that were influenced and mediated by social interactions, such as technological learning, seasonal traveling, material culture exchange, and symbolic expression. We use compositional analyses to determine localized ochre procurement strategies and long-distance transport across a network of fifteen archaeological sites and mineral resources. Newly refined chronologies from Lion Cavern at Ngwenya using optically stimulated luminescence dating also reaffirm its antiquity as the oldest known evidence for intensive ochre mining worldwide (~48,000 years ago).
... The relative abundance of USOs on the Hollow Rock Shelter foraging landscape may have been one of the factors that contributed to the site's strategic importance in the past. Singels et al. (2016) demonstrated how modern foragers along South Africa's south coast can collect edible USOs to satisfy a person's daily caloric requirements within two hours, so that such foods may have been valuable staples (Wadley et al., 2020), which can be stored and/or used as travel foods on hunting and other longer trips. Deacon's (1993) argument that USOs were obvious plant staples in the relatively treeless biomes of southern Africa stimulated geophyte-centred research, often to the exclusion of all other plant foods (e.g., Singels et al., 2016;Botha et al., 2022; (1976) hierarchical analysis of plant foods used by the San huntergatherers of ∕ = Kade Pan in the Central Kalahari of southern Africa demonstrates a more nuanced balance in major-minor (staple) plant foods wherein fruit (including berries and melons) constitute 32 % (5major, 3 = minor), USOs 28 % (6 = major, 2 = minor), stems 20 % (3 = major, 2 = minor), and seeds (2 = major) and leaves (2 = minor) 8 % each (Tanaka, 1976: 117-118). ...
... Thus far, there are no records for cooking food at Hollow Rock Shelter, but there is evidence of hearths (Högberg and Larsson, 2011), and Schmidt and Högberg (2018) demonstrated the heat treatment of silcrete for knapping purposes, probably by placing the nodules in an open fire. Elsewhere in South Africa, people roasted corms at Border Cave in KwaZulu-Natal by ~ 170 ka (Wadley et al., 2020), and at Klasies River in the Eastern Cape since ~ 120 ka (Larbey et al., 2019). It is therefore reasonable to accept that the people who used Hollow Rock Shelter during the Middle Stone Age knew how to make, sustain and control fires, and that they probably also cooked some of their food. ...
Article
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Studying Stone Age foraging behaviours in terms of plant foods is difficult because of preservation, sampling and identification biases. Current foodplant populations, knowledge about their use in recent times, and how they are distributed across the landscape, provide valuable middle-range proxies from which work on archaeological landscapes and material can benefit. With this contribution we provide foodplant lists for three different foraging ranges (radii of ~ 12.5 km, ~35 km, and ~ 70 km) around Hollow Rock Shelter. By comparing data for each of the foraging ranges directly, we discuss proportional increases in foodplant resources when moving further away from the site. We demonstrate that the ~ 35 km foraging range is the most efficient. This implies that people staying at the site (for shorter or longer periods) may have regularly employed a strategy of temporary camping for a night or two away from the site to forage especially preservable foods that could be brought to the site. Our data highlight potential plant-food staples, and show that under-surface storage organs (USOs) of plants, followed by fruit and leaves are the most abundant edible plant parts available on the Hollow Rock Shelter landscape within all three of the foraging ranges, and that most of these could be eaten raw.
... The latter ranged from the intensified occupation of coastal landscapes and exploitation of marine resources in both northern and southern Africa (summarized in Will et al., 2019b) to a broad range of prey animals, including large and dangerous species marking MSA people as capable, sophisticated hunters (Faith, 2008;Dusseldorp, 2010;Clark and Kandel, 2013). Plant use is more widespread in the MSA than previously believed, with the exceptionally well-preserved record from southern Africa demonstrating the dietary use of grass seeds (Mercader, 2009), cooking and consumption of geophytes (Wadley et al., 2020a), and the use of sedges for bedding and as insect repellents by 200 ka, and continuing well into the Late Pleistocene (Goldberg et al., 2009;Wadley et al., 2011Wadley et al., , 2020bSievers et al., 2022). New raw materials also become more frequently incorporated into people's technological repertoire. ...
... Increasing the niche breadth of MSA people (Kandel et al., 2016;d'Errico et al., 2017;Roberts and Stewart, 2018) might have allowed them to expand to new habitats. In connection with more varied habitats, more diverse diets (Clark and Kandel, 2013;Will et al., 2019b;Wadley et al., 2020a) may have buffered against food shortages, decreasing mortality and extinction rates that boosted population sizes. New ways of social organization that increased the interconnectivity between groups can already be seen at the beginning of the MSA in long-distance transport of raw materials but likely increased toward the end of the MSA, such as long-term exchange and/or migration networks suggested by OES beads reaching back to the Late Pleistocene (Stewart et al., 2020;Miller and Wang, 2022). ...
Article
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The behavioral origins of Homo sapiens can be traced back to the first material culture produced by our species in Africa, the Middle Stone Age (MSA). Beyond this broad consensus, the origins, patterns, and causes of behavioral complexity in modern humans remain debated. Here, we consider whether recent findings continue to support popular scenarios of: (1) a modern human 'package,' (2) a gradual and 'pan-African' emergence of behavioral complexity, and (3) a direct connection to changes in the human brain. Our geographically structured review shows that decades of scientific research have continuously failed to find a discrete threshold for a complete 'modernity package' and that the concept is theoretically obsolete. Instead of a continent-wide, gradual accumulation of complex material culture, the record exhibits a predominantly asynchronous presence and duration of many innovations across different regions of Africa. The emerging pattern of behavioral complexity from the MSA conforms to an intricate mosaic characterized by spatially discrete, temporally variable, and historically contingent trajectories. This archaeological record bears no direct relation to a simplistic shift in the human brain but rather reflects similar cognitive capacities that are variably manifested. The interaction of multiple causal factors constitutes the most parsimonious explanation driving the variable expression of complex behaviors , with demographic processes such as population structure, size, and connectivity playing a key role. While much emphasis has been given to innovation and variability in the MSA record, long periods of stasis and a lack of cumulative developments argue further against a strictly gradualistic nature in the record. Instead, we are confronted with humanity's deep, variegated roots in Africa, and a dynamic metapopulation that took many millennia to reach the critical mass capable of producing the ratchet effect commonly used to define contemporary human culture. Finally, we note a weakening link between 'modern' human biology and behavior from around 300 ka ago.
... Grass bedding in Member 5 BS dated at about 170 ka, and Member 5 WA, dated to >227 ka makes it the oldest known (Wadley et al., 2020a;Esteban et al., 2023). The same is true for whole cooked starchy rhizomes, identified as Hypoxis angustifolia, from layers dated to c. 170 ka ago (Wadley et al., 2020b). Charcoal is preserved throughout the sequence, providing a rare and comprehensive record of climate change in the region . ...
... Quaternary Science Reviews 306 (2023) 108030 reducing the availability of this resource during periods of increased aridity, or cultural change. It may be no coincidence that between about 170 and 100 ka ago, we also found many charred whole rhizomes and fragments of the edible Hypoxis angustifolia in Border Cave (Wadley et al., 2020b). The implications of this discovery reach far beyond the interesting fact that people so far back in time cooked starchy foods. ...
Article
Fragments of land snail (Achatinidae) shell were found at Border Cave in varying proportions in all archaeological members, with the exception of the oldest members 5 WA and 6 BS (>227,000 years ago). They were recovered in relatively high frequencies in Members 4 WA, 4 BS, 1 RGBS and 3 WA. The shell fragments present a range of colours from lustrous beige to brown and matt grey. The colour variability can occur when shell is heated. This possibility was explored here through experimental heating of giant land snail shell fragments (Achatinidae, Metachatina kraussi - brown lipped agate snail) in a muffle furnace from 200 to 550 °C for different lengths of time. Colour change, weight loss, and shattering of the heated samples were recorded. Transformation of aragonite into calcite and the occurrence of organic material was investigated by means of Infrared and Raman spectroscopy. Scanning electron microscopy was also used on selected specimens to help identify heat-induced transformation as opposed to taphonomic alteration. The identification on archaeological fragments of features produced by experimentally heating shells at high temperatures or for long periods has led us, after discarding alternative hypotheses, to conclude that large African land snails were systematically brought to the site by humans, roasted and consumed, starting from 170,000 years ago and, more intensively between 160,000 and 70,000 years ago. Border Cave is at present the earliest known site at which this subsistence strategy is recorded. Previous research has shown that charred whole rhizomes and fragments of edible Hypoxis angustifolia were also brought to Border Cave to be roasted and shared at the site. Thus, evidence from both the rhizomes and snails in Border Cave supports an interpretation of members of the group provisioning others at a home base, which gives us a glimpse into the complex social life of early Homo sapiens.
... The close proximity of the shelter mouth to the edge of the cliff and the challenging access route may have encouraged occupants to conduct a wider range of activities inside this shelter than at other sites. Activities that took place inside the shelter include the burial of an infant with associated ornamentation (Cooke et al., 1945;d'Errico and Backwell, 2016), plant food preparation (Wadley et al., 2020a), and the construction of grass bedding and other forms of site management (Wadley et al., 2020b). ...
... These components contribute tangibly, but variably, to the matrix and clast fractions and to the chemical composition of the deposits. Each component of the archaeological record (e.g., seeds, phytoliths, charcoal) is analysed individually as part of a broad multi-proxy research programme (e.g., Wadley et al., 2020a;Zwane and Bamford, 2021;Sievers et al., 2022;Esteban et al., this volume;Lennox et al., this volume). Significant anthropogenic contributions to the sediments include diffuse and dense deposits of burnt and unburnt vegetation, charcoal, bone and wood ash, and aggregated soil particles of various shapes and sizes. ...
Article
Border Cave (BC) has accumulated over 200,000 years of archaeological deposits that document remarkable evidence of human behaviour during the Middle and Later Stone Age. For nearly fifty years, researchers have relied on the stratigraphic framework established by Peter Beaumont in 1973, in which the deposits are lithostratigraphically categorized into a sequence of alternating ‘Brown Sand’ (BS) and ‘White Ash’ (WA) members. Geoarchaeological work in the 1970s focused on stratigraphic sequencing of the anthropogenic assemblages, and proposed broad correlations between autogenic contributions and environmental conditions. The research presented here was undertaken as part of a new excavation campaign at Border Cave started in 2015 under the direction of Backwell at al. Re-examining the stratigraphic context of the deposits and assessing site formation processes are among the key goals of this project; this will enable finer-scale intra- and inter-member comparative analyses of the artefacts and ecofacts recovered at the site. In this paper, we apply a facies and allostratigraphic approach to assess the stratigraphic sequence exposed through the Backwell et al. excavations. We also provide an initial assessment of the prevailing site formation processes active in the deposition and modification of the sediments. The geoarchaeological data are integrated with new zooarchaeological and taphonomic evidence in order to explore inter- and intra-unit patterns throughout the sequence. Results of this work are: (1) exposed sediments can be broadly correlated to members of the Beaumont sequence; (2) we clearly define member boundaries, reassess member stratigraphic complexity and recognise finer intra-member layering; (3) geoarchaeological and taphonomic studies demonstrate that the sediments have been subjected to greater post-depositional disturbance than was previously recognised and affect all levels of the sequence; (4) overall, faunal density at BC appears to be much lower than that at other Middle Stone Age sites such as Blombos and Sibudu; (5) multiproxy analysis suggests that WA and BS members have distinctive taphonomic histories that cross-cut the identified archaeological industries. As such, caution is warranted when combining BS and WA members for analysis of artefacts and ecofacts.
... Plant foods, however, remained a staple of the human diet. Learning how to effectively forage for, extract, and/or prepare such resources from the landscape would have been as challenging and rewarding in terms of behavioral and cognitive evolution as hunting (e.g., Hardy et al., 2015;Henry et al., 2014;Lombard & Kyriacou, 2020;O'Connell et al., 2002;Schnorr et al., 2016;Wadley et al., 2020). ...
... Along the Cape coastline and in the Fynbos Biome geophytes (plants with growth forms that include specifically modified stem or root systems for the storage of energy or water in the form of tubers, corms, rhizomes, or bulbs [see Beentjie, 2016]) may have provided a key carbohydrate source for pre-colonial populations as far back as ~100 ka (e.g., de Vynck et al., 2016;Deacon, 1993;Marean, 2010;Opperman & Heydenrych, 1990). For Early H. sapiens on the inland Savanna Biome, Wadley et al. (2020) reported geophyte preparation by ~170 ka at Border Cave, KwaZulu-Natal, in the form of charred rhizomes-likely that of roasted Hypoxis angustifolia Lam. Another early record of plant-food processing by Pleistocene H. sapiens on the Savanna Biome may be the residues found on stone tools from Lake Niassa, Mozambique, dated to ~105 ka that include prolific starch granules of grass seeds such as sorghum (Mercader, 2009). ...
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Despite a century’s work in the UNESCO Cradle of Humankind World Heritage Site in South Africa, there has been no systematic consideration of the area’s full foodplant regime when palaeo-scientists try to reconstruct past hominin dietary ecologies. Here we present the first inventory of human-foraged foodplants currently growing in the Cradle of Humankind and discuss the time depth of the relative biomes, showing that some of the plant regimes may well have been available for hominin foraging throughout the Quaternary. We list 223 taxa, highlighting the most species-rich foodplant families in the Cradle, such as the Poaceae, Apocynaceae, and Fabaceae. Our results show an almost equal availability of edible fruits, leaves, and underground/underwater storage organs (USOs) in the Cradle’s foodplant population, supporting work suggesting diet specialization or flexible foraging strategies for the Cradle’s hominins. Whereas it was thought that human geophyte and/or USO consumption was particularly high on the Cape Fynbos landscape during the middle-late Pleistocene, the richness of USO plant foods growing in the Cradle—most of them non-geophytic in their growth forms (thus without specialized/modified storage systems such as tubers, corms, rhizomes or bulbs)—demonstrates that such resources were probably also abundantly available to inland hominin populations. We suggest that future dietary reconstructions for the Cradle’s hominins may benefit from working with the known foodplant population presented here.
... Plants have always played an important role in human diet. They are our primary source of carbohydrates and therefore provide us with readily available energy (Hardy et al., 2022;Larbey et al., 2019;Wadley et al., 2020). In addition to collecting plants that can be eaten raw, there are also references to the use of plants that are inedible or even poisonous in their raw state. ...
Article
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Plants are a crucial part of the human diet, serving as a primary source of micronutrients, fiber, and carbohydrates, providing readily available energy. Beyond the consumption of cooked and raw edible plants, early humans also developed methods for plant processing for delayed consumption, to de-toxify/improve bioavailability, and perhaps for flavor. In later prehistory delayed consumption includes preservation processes for storage. The processing of plants through sprouting, fermentation, cooking, and roasting has not only expanded the variety of consumable plant species but also enhanced their nutritional value. There are few methods for detecting fermented foods in the early prehistoric diet. Based on the hypothesis that the microbes active in fermentation change the nitrogen and carbon isotope ratios of plant foods, our pilot study lays the groundwork for further research, offering a promising direction for understanding the complexities of human-plant interactions throughout history. This study focuses on these four plant treatment methods, which have both contemporary relevance and archaeobotanical evidence of use. We aimed to explore how these treatments might affect stable isotope values, such as carbon (δ¹³C) and nitrogen (δ¹⁵N), which are crucial for reconstructing ancient diets. Through an experimental approach involving 12 plant species, our findings suggest that while sprouting and roasting had minimal impact on isotopic values, fermentation and cooking showed more variable effects depending on the plant species. These preliminary results provide new insights into the influence of plant processing on isotopic compositions and underscore the importance of considering these factors in dietary reconstructions.
... For instance, the consumption of starchy, geophytic forbs with USOs by early hominids is key to understanding human evolution (Landen & Wrangham 2005). More recent evidence confirmed USO consumption by plant gatherers in Africa ∼170,000 years ago (Wadley et al. 2020), roughly contemporaneous with the origin of modern Homo sapiens. Ever since, human livelihoods in tropical grasslands have been supported by forbs as a source of food and/or medicinal items for people, livestock, and game. ...
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Forbs are important contributors to species diversity and ecosystem functions in low-latitude grasslands, where they support diverse herbivore communities and millions of people. Native forb assemblages tolerate disturbances and physiological stressors (fire, herbivory, drought, and frost) that together have shaped their exceptional functional diversity. Yet, compared to trees and grasses, forbs have received much less attention in grassland studies until recently. Here, we review forb-centric literature to illustrate that land conversion and responsible management of fire and herbivory are crucial to maintaining forb diversity. Management practices promoting forb diversity offer (a) high-quality food items and medicinal resources that support rural livelihoods and animal diversity (from wild ungulates and livestock to fossorial rodents and insects), including their adaptive foraging patterns, and (b) carbon and nutrient inputs that regulate belowground processes. Improved understanding of the above- and belowground regeneration strategies of forbs is critical for restoration and conservation to secure their services in future old-growth tropical and subtropical grasslands.
... For instance, the consumption of starchy, geophytic forbs with USOs by early hominids is key to understanding human evolution (Landen & Wrangham 2005). More recent evidence confirmed USO consumption by plant gatherers in Africa ∼170,000 years ago (Wadley et al. 2020), roughly contemporaneous with the origin of modern Homo sapiens. Ever since, human livelihoods in tropical grasslands have been supported by forbs as a source of food and/or medicinal items for people, livestock, and game. ...
Article
Full-text available
Forbs are important contributors to species diversity and ecosystem functions in low-latitude grasslands, where they support diverse herbivore communities and millions of people. Native forb assemblages tolerate disturbances and physiological stressors (fire, herbivory, drought, and frost) that together have shaped their exceptional functional diversity. Yet, compared to trees and grasses, forbs have received much less attention in grassland studies until recently. Here, we review forb-centric literature to illustrate that land conversion and responsible management of fire and herbivory are crucial to maintaining forb diversity. Management practices promoting forb diversity offer (a) high-quality food items and medicinal resources that support rural livelihoods and animal diversity (from wild ungulates and livestock to fossorial rodents and insects), including their adaptive foraging patterns, and (b) carbon and nutrient inputs that regulate belowground processes. Improved understanding of the above- and belowground regeneration strategies of forbs is critical for restoration and conservation to secure their services in future old-growth tropical and subtropical grasslands.
... For example, at Border Cave, Wadley et al. (2020a) recorded plant bedding at 200 ka, with variation in bedding construction between 60 and 40 ka when the use of sedges decreased in favor of grasses, and a small piece of woven grass dating to 42.6 ka represents the earliest evidence of simple weaving (Sievers et al., 2022). Since ~ 170 ka people at Border Cave cooked starchy Hypoxis geophytes (Wadley et al., 2020b), and some of the taxa identified through charcoal analysis are known to be good fuelwoods (e.g., Protea spp., Acacia spp., and Leucosidea sericea), and others such as Tarchonanthus spp., Spirostachys africana, and Cryptocarya spp. could have been used for insecticidal or medicinal purposes (Lennox et al., 2022). ...
Article
Full-text available
Current phyto-scapes (plant populations in their geo-spatial context) are not exact replicas of past foraging potential, yet they provide valuable data about the carrying capacity or potential of a foraging-scape. Knowledge about contemporary micro-ecologies and ethno-historical plant use can inform on behavioral aspects, should such plants be found in archaeological deposits. It is in this context that we explore existing information (data and literature) to establish the current vegetation types and micro-ecologies around Holley Shelter, KwaZulu-Natal, South Africa, that contains Middle and Later Stone Age occupations. We present the first plant species inventory consisting of > 1500 taxa growing approximately a day’s foraging distance from the site, compiled from records provided by the South African National Biodiversity Institute and other sources. From this list, we generated separate checklists for foodplants (n = 450) and plants that have other uses (n = 337), to gain insight into the site’s current phyto-fitness potential. The resulting database is intended as a modern proxy for future work on the site’s archaeo-botany and palaeo-climatic reconstructions. The data is also applicable to other sites on the Savanna/Grassland Biomes of southern Africa with records of the same species. Here we use the foodplant checklist and what is known about the edible plant parts, their seasonality and the distribution of some species to speculate about land-use patterns. These hypotheses can be tested for the past with future archaeo-botanical work. We demonstrate that, compared to archaeological sites in the Eastern and Western Cape for which comparable data exist, Holley Shelter is rich in foodplants, and especially so in plant foods that are known to buffer against famine during the periodical droughts of inland South Africa.
... Recent excavations at Umbeli Belli (Bader et al., 2016(Bader et al., , 2018, a comparative work on the final MSA assemblages from Sibhudu (Bader et al., 2022c), and the reexamination of Sibebe in the Highveld of Eswatini (Bader et al., 2022a) have all shed further light on the technical dynamics of the very end of the MSA. Important work on the final MSA and LSA transition has also been carried out at Border Cave (Backwell et al., 2018;d'Errico and Backwell, 2016;Timbrell et al., 2022;Wadley et al., 2020;Zwane and Bamford, 2021), Umhlatuzana (Lombard et al., 2010, Sifogeoarki et al., 2020 and on specific technological aspects such as the regional signals of point morphology (Mohapi, 2009(Mohapi, , 2012(Mohapi, , 2013. Notwithstanding these efforts, there are still few well-documented sites and a lack of key data necessary to understand human behavioral patterns during MIS 3, particularly considering the large area and diverse ecology of the region. ...
... Recent excavations at Umbeli Belli (Bader et al., 2016(Bader et al., , 2018, a comparative work on the final MSA assemblages from Sibhudu (Bader et al., 2022c), and the reexamination of Sibebe in the Highveld of Eswatini (Bader et al., 2022a) have all shed further light on the technical dynamics of the very end of the MSA. Important work on the final MSA and LSA transition has also been carried out at Border Cave (Backwell et al., 2018;d'Errico and Backwell, 2016;Timbrell et al., 2022;Wadley et al., 2020;Zwane and Bamford, 2021), Umhlatuzana (Lombard et al., 2010, Sifogeoarki et al., 2020 and on specific technological aspects such as the regional signals of point morphology (Mohapi, 2009(Mohapi, , 2012(Mohapi, , 2013. Notwithstanding these efforts, there are still few well-documented sites and a lack of key data necessary to understand human behavioral patterns during MIS 3, particularly considering the large area and diverse ecology of the region. ...
... 181 Their subsistence behavior not only included hunting, most likely with handheld stabbing or thrusting spears, 182 but also roasting of starchy rhizomes at 170 ka at that Border Cave. 183 Bedding construction and fire at the Border Cave at 200 ka 184 and early ochre use 185 also reflect complexity. There is only one coastal site that represents the EMSA, Pinnacle Point Cave 13B. ...
Chapter
Currently the concept of the Middle Stone Age (MSA) denotes the period between c . 300 and 25 ka. It is a phase marked by prepared core reduction methods used to knap predetermined flakes and blades that are occasionally retouched into various types of tools. Denticulates, notches, and scrapers occur regularly, and bifacial and unifacial points and backed geometrics are sometimes linked to time-restricted regional patterns, especially for the Still Bay and Howiesons Poort technocomplexes. An uneven geographical representation of data and insufficient dating resolution preclude a coherent consensus chrono-culture stratigraphic framework for the southern African region, the area south of the Kunene and Zambezi Rivers encompassing the modern political entities of Namibia, Botswana, Zimbabwe, southern Mozambique, Swaziland (eSwatini), Lesotho, and South Africa. Therefore many assemblages are described in relation to the marine isotope stages and local industries. Perhaps the most radical development in MSA research during the 20th century relates to the characterization of culture and behavior. In the formation years, when mostly surface collections of stone tools, organized into industries and variants were available, MSA “cultures” of the region were seen as the product of waves of immigrants that entered dark Africa from Europe, in increasingly “advanced” forms. In the latter part of the 20th century, the prevailing Eurocentric paradigm suggested that it was only with the Upper Paleolithic–like Later Stone Age that “modern” culture developed in southern Africa. Although Eurocentric thinking prevails, “modernity” is now linked to the MSA especially after 100 ka. Fluctuating complexity in behavior may relate to various degrees of social interaction within dynamic landscapes. Paleoenvironmental data is growing and, combined with cutting-edge geoarchaeological and digital methods, allow a deeper understanding of past habitats and ecological contexts. Studies on the MSA from southern Africa are expanding rapidly. This growth would be most productive and ethical if research is integrated with African socio-political realities, engaging with decoloniality and inclusivity.
... Our millimeter scale approach to excavations and use of sophisticated analytical instruments is yielding illuminating results. The new excavations have yielded the oldest evidence of cooked starchy rhizomes (Hypoxis angustifolia) at 170 ka (Wadley et al., 2020a). Fifty-five rhizomes were recovered from Members 5 BS and 4 WA. ...
Chapter
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Border Cave in KwaZulu-Natal, South Africa, preserves a long and continuous archaeological record from 227 ka to 24 ka years ago, rendering it a key Middle Stone Age site in southern Africa. It has yielded the skeletal remains of eight anatomically modern Homo sapiens individuals, a lithic sequence that includes MSA 1, MSA 2, MSA 3, and Early Later Stone Age Industries, ochre, marine shells, the oldest burial associated with a personal ornament, and early expressions of complex cognition and innovation. Organic preservation is remarkable, with grass bedding that contains aromatic leaves with insecticidal properties found throughout the sequence. The bedding layers show that it was systematically placed on a layer of ash to deter crawling insects. Comprehending that aromatic leaves and ash deter pests, and using them as tools for delayed gratification following planning and strategizing, and organization that entailed a sequence of events, implies that complex cognition was in place from 200,000 years ago. Charred underground storage organs come from layers dated to 170,000 years ago, making them the oldest known examples of cooked starchy rhizomes. The fact that the rhizomes were cooked implies that the inhabitants of the shelter were able to make fire at will, another indication of complex cognition in early modern humans. A range of organic remains such as ostrich eggshell beads and bone points used as poisoned arrowheads are found in Early Later Stone Age layers starting at around 44,000 years ago, and they represent the earliest examples of modern human behavior as we know it.
... More secure evidence for cooking dates to ~400,000 ya with the earliest known hearth, at Qesem Cave, Israel (Shahack-Gross et al., 2014;Stiner et al., 2011). Charred rhizomes have been identi ed from Border Cave, South Africa, suggesting that humans cooked tubers, bulbs, and other edible geophytes at least by ~170,000 ya (Wadley et al., 2020). There is evidence of Neanderthals preserving meats and sh by drying but also, more controversially, cooking (Hardy & Moncel, 2011;Henry, 2017;Henry et al., 2017;Patou-Mathis, 2000;Sørensen, 2011;Zilhão, 2021). ...
Chapter
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... seeds), the complexity of identification methods, as well as lower archaeobotanical sampling in the regions from which many of these crops originate. Despite these issues, the consumption, management and cultivation of asexually propagated crop species have been shown to be extremely ancient practices (Clarkson et al., 2017;Denham et al., 2003;Florin et al., 2020;Wadley et al., 2020), which have been overlooked archaeologically for many decades creating an incomplete view of ancient subsistence strategies and likely inflating the importance of seed-based agriculture for populations reliant on mixed subsistence practices. ...
Article
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The potential applications of microCT scanning in the field of archaeobotany are only just beginning to be explored. The imaging technique can extract new archaeobotanical information from existing archaeobotanical collections as well as create new archaeobotanical assemblages within ancient ceramics and other artefact types. The technique could aid in answering archaeobotanical questions about the early histories of some of the world’s most important food crops from geographical regions with amongst the poorest rates of archaeobotanical preservation and where ancient plant exploitation remains poorly understood. This paper reviews current uses of microCT imaging in the investigation of archaeobotanical questions, as well as in cognate fields of geosciences, geoarchaeology, botany and palaeobotany. The technique has to date been used in a small number of novel methodological studies to extract internal anatomical morphologies and three-dimensional quantitative data from a range of food crops, which includes sexually-propagated cereals and legumes, and asexually-propagated underground storage organs (USOs). The large three-dimensional, digital datasets produced by microCT scanning have been shown to aid in taxonomic identification of archaeobotanical specimens, as well as robustly assess domestication status. In the future, as scanning technology, computer processing power and data storage capacities continue to improve, the possible applications of microCT scanning to archaeobotanical studies will only increase with the development of machine and deep learning networks enabling the automation of analyses of large archaeobotanical assemblages.
... These plant foods serve/d as staple carbohydrate source for many foraging and farming groups across the globe and in southern Africa, with local wild USOs containing between 40 and 228 calories per 100 g(Singels et al. 2016; also seeLombard 2022). From historical records we know that such 'uintjies' were a prized 'veldkos' of local hunter-gatherers (e.g., Smit 1851;Bleek and Lloyd 1911;Smith 1966), and their archaeological remnants provide some of the earliest evidence for cooking plant foods in South Africa since ~170 ka(Larbey et al. 2019;Wadley et al. 2020). What is more, geophytic USOs are less perishable than other plant foods so that they can be collected, transported and stored for future use. ...
... This protracted adaptive process required human genetic evolution (Mathieson et al. 2015) as well as the development of technologies such as fire and stone tools, the latter for grinding seeds and digging up roots (Hillman 1989;Hillman and Wollstonecroft 2014). Humans were cooking tubers at least 170,000 years ago (Wadley et al. 2020) and grass seeds (including the future domesticate sorghum) at least 105,000 years ago (Mercader 2009). Fifty thousand years ago, Neanderthals were cooking and consuming plants that would subsequently be domesticated by Homo sapiens Piperno 2011, 2014). ...
Chapter
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Contributors explore common elements in the evolutionary histories of both human and insect agriculture resulting from convergent evolution. During the past 12,000 years, agriculture originated in humans as many as twenty-three times, and during the past 65 million years, agriculture also originated in nonhuman animals at least twenty times and in insects at least fifteen times. It is much more likely that these independent origins represent similar solutions to the challenge of growing food than that they are due purely to chance. This volume seeks to identify common elements in the evolutionary histories of both human and insect agriculture that are the results of convergent evolution. The goal is to create a new, synthetic field that characterizes, quantifies, and empirically documents the evolutionary and ecological mechanisms that drive both human and nonhuman agriculture. The contributors report on the results of quantitative analyses comparing human and nonhuman agriculture; discuss evolutionary conflicts of interest between and among farmers and cultivars and how they interfere with efficiencies of agricultural symbiosis; describe in detail agriculture in termites, ambrosia beetles, and ants; and consider patterns of evolutionary convergence in different aspects of agriculture, comparing fungal parasites of ant agriculture with fungal parasites of human agriculture, analyzing the effects of agriculture on human anatomy, and tracing the similarities and differences between the evolution of agriculture in humans and in a single, relatively well-studied insect group, fungus-farming ants. Contributors Duur K. Aanen, Niels P. R. Anten, Peter H. W. Biedermann, Jacobus J. Boomsma, Laura T. Buck, Guillaume Chomicki, Tim Denham, R. Ford Denison, Dorian Q. Fuller, Richard Gawne, Nicole M. Gerardo, Thomas C. Harrington, Ana Ješovnik, Judith Korb, Chase G. Mayers, George R. McGhee, Kenneth Z. McKenna, Lumila P. Menéndez, Peter N. Peregrine, Ted R. Schultz
... The list of commercial products containing African potato plant extracts seem to be expanding in horticultural practices [78]. This, together with increased market value and global trade, has negative long-term impacts on plant preservation [79]. ...
Chapter
South Africa, a country considered affluent in nature, ranks third in global biodiversity and encompasses approximately 9%of higher plants on planet Earth.Many indigenous plants have been utilised as herbal medicine, proving successful in treating numerous ailments. From the common cold to pandemic maladies such as COVID-19 in the 21st century and the treatment of incurable diseases, South African inhabitants have found great promise in the healing properties of these plants. Phytomedicine is a rapidly evolving topic, with in-depth bioactive composition analysis, identifying therapeutic actionmechanisms, and disease prevention.Whilewe are nowpoised to take advantage of nature’s medicine cabinet with greater scientific vigour, it remains critical that these practises are done with caution. Overharvesting significantly impacts biodiversity and cultivation practices amidst the beautiful nature of these nutraceuticals. This book chapter focuses on the therapeutic potential of commonly used South Africanmedicinal plants, their ethnopharmacological properties, and how we can conserve this treasure cove we call home for future generations.
... Border Cave, near Eswatini, has yielded important archaeological evidence that may underscore early human subsistence behavior. Burnt bedding from the 200,000-year-old layers and cooked starchy rhizomes dated to 170,000 years ago show that people during this period used fire for food preparation (Wadley et al. 2020a(Wadley et al. , 2020b. This emphasizes the Middle Stone Age exploitation of varied food sources. ...
Preprint
Analysis of Late Pleistocene fauna exploitation (~130,000–12,000 years ago) in southern Africa is of global academic relevance. Faunal analyses from southern African sites have led to the development of influential hypotheses on the evolution of modern human hunting methods and subsistence economies.In the 1970s and 1980s, analysis of faunal remains from the Middle Stone Age site Klasies River informed the hypothesis that Middle Stone Age humans were less effective hunters than ethnographically documented hunter-gatherers. This was based on the underrepresentation of dangerous prey species in the bone assemblages. The development of detailed taphonomic research in the 1990s and 2000s demonstrated that the accumulation of faunal assemblages was the result of complex processes involving both human and nonhuman agents. These studies helped establish that Middle Stone Age hunters were as capable as those in ethnographically documented societies. Since then, important progress has been made in the identification of the weapons systems that were used to hunt animals. Analyses of lithic implements indicate bow-and-arrow use in southern Africa going back to at least 65,000 years ago.Animal exploitation strategies do change over time. Hunting strategies probably focused on large antelope during the Middle Pleistocene, and the importance of smaller animals increased This change was likely caused by a shift in prey populations that stemmed from a combination of environmental change and perhaps human population pressure.Late Pleistocene archaeological sites show increasing evidence for intensification; that is, an increase in the amount of food extracted from the environment by more thorough processing of prey, exploitation of new prey types, and development of new exploitation strategies. This pattern is usually linked to animal overexploitation and may be a result of human population expansion or environmental change if decreasing productivity limits the supply of animal prey. Notable examples of this are shellfish middens at coastal sites, the abundance of tortoises, and the presence of large numbers of small mammals that were likely snared instead of pursued.
... However, evidence regarding the use of heat specifically for food preparation by H. erectus is inconclusive and controversial 16,17 . While it is likely that early fire-using hominins had already cooked their food, definitive evidence of this practice has only been demonstrated to date for early Homo sapiens and Neanderthals, in association with vegetal material, with the earliest date of 170 thousand years ago (ka) 18,19 . ...
Article
Full-text available
Although cooking is regarded as a key element in the evolutionary success of the genus Homo, impacting various biological and social aspects, when intentional cooking first began remains unknown. The early Middle Pleistocene site of Gesher Benot Ya’aqov, Israel (marine isotope stages 18–20; ~0.78 million years ago), has preserved evidence of hearth-related hominin activities and large numbers of freshwater fish remains (>40,000). A taphonomic study and isotopic analyses revealed significant differences between the characteristics of the fish bone assemblages recovered in eight sequential archaeological horizons of Area B (Layer II-6 levels 1–7) and natural fish bone assemblages (identified in Area A). Gesher Benot Ya’aqov archaeological horizons II-6 L1–7 exhibited low fish species richness, with a clear preference for two species of large Cyprinidae (Luciobarbus longiceps and Carasobarbus canis) and the almost total absence of fish bones in contrast to the richness of pharyngeal teeth (>95%). Most of the pharyngeal teeth recovered in archaeological horizons II-6 L1–7 were spatially associated with ‘phantom’ hearths (clusters of burnt flint microartifacts). Size–strain analysis using X-ray powder diffraction provided evidence that these teeth had been exposed to low temperature (<500 °C), suggesting, together with the archaeological and taphonomic data, that the fish from the archaeological horizons of Area B had been cooked and consumed on site. This is the earliest evidence of cooking by hominins.
... Besides this unusual characteristic, Border Cave has also yielded hominin remains (Beaumont, 1980;Beaudet et al., 2022), including a burial with an associated ornament in Member 1 RGBS, putatively associated with the Howiesons Poort technological tradition (d' Errico and Backwell, 2016). Other outstanding finds include an incised bone (Beaumont, 1978) interpreted as the earliest known system of notation (d 'Errico et al., 2018), and the oldest evidence of bedding documented in prehistory (Wadley et al., 2020a;Sievers et al., 2022) and cooked starchy rhizomes at 170 ka (Wadley et al., 2020b). ...
Article
Full-text available
Lithic assemblages immediately following the Howiesons Poort, often loosely referred to as the 'post-Howiesons Poort' or MSA III, have attracted relatively little attention when compared to other well-known phases of the South African Middle Stone Age (MSA) sequence. Current evidence from sites occurring in widely-differing environments suggests that these assemblages are marked by temporal and technological variability, with few features in common other than the presence of unifacial points. Here we present a technological and geometric morphometric analysis of 'points' from the new excavations of Members 2 BS, 2 WA and the top of 3 BS members at Border Cave, KwaZulu-Natal, one of the key sites for studying modern human cultural evolution. Our complementary methodologies demonstrate that, at this site, hominins adopted a knapping strategy that primarily produced non-standardised unretouched points. Triangular morphologies were manufactured using a variety of reduction strategies, of which the discoidal and Levallois recurrent centripetal methods produced distinctive morphologies. We find technological and morphological variability increases throughout the post-Howiesons Poort sequence, with clear differences between and within chrono-stratigraphic groups. Finally, we assess the suitability of the 'Sibudan' cultural-technological typology proposed for post-Howiesons Poort assemblages at Sibhudu, another KwaZulu-Natal site, and find similarities in the morphological axes characterising the samples, despite differences in the shaping strategies adopted. Overall, our work contributes to the growing body of research that is helping to address historical research biases that have slanted our understanding of cultural evolution during the MSA of southern Africa towards the Still Bay and Howiesons Poort technocomplexes.
... Border Cave, near Eswatini, has yielded important archaeological evidence that may underscore early human subsistence behavior. Burnt bedding from the 200,000-year-old layers and cooked starchy rhizomes dated to 170,000 years ago show that people during this period used fire for food preparation (Wadley et al. 2020a(Wadley et al. , 2020b. This emphasizes the Middle Stone Age exploitation of varied food sources. ...
Chapter
Full-text available
Analysis of Late Pleistocene fauna exploitation (~130,000–12,000 years ago) in southern Africa is of global academic relevance. Faunal analyses from southern African sites have led to the development of influential hypotheses on the evolution of modern human hunting methods and subsistence economies. In the 1970s and 1980s, analysis of faunal remains from the Middle Stone Age site Klasies River informed the hypothesis that Middle Stone Age humans were less effective hunters than ethnographically documented hunter- gatherers. This was based on the underrepresentation of dangerous prey species in the bone assemblages. The development of detailed taphonomic research in the 1990s and 2000s demonstrated that the accumulation of faunal assemblages was the result of complex processes involving both human and nonhuman agents. These studies helped establish that Middle Stone Age hunters were as capable as those in ethnographically documented societies. Since then, important progress has been made in the identification of the weapons systems that were used to hunt animals. Analyses of lithic implements indicate bow-and-arrow use in southern Africa going back to at least 65,000 years ago. Animal exploitation strategies do change over time. Hunting strategies probably focused on large antelope during the Middle Pleistocene, and the importance of smaller animals increased This change was likely caused by a shift in prey populations that stemmed from a combination of environmental change and perhaps human population pressure. Late Pleistocene archaeological sites show increasing evidence for intensification; that is, an increase in the amount of food extracted from the environment by more thorough processing of prey, exploitation of new prey types, and development of new exploitation strategies. This pattern is usually linked to animal overexploitation and may be a result of human population expansion or environmental change if decreasing productivity limits the supply of animal prey. Notable examples of this are shellfish middens at coastal sites, the abundance of tortoises, and the presence of large numbers of small mammals that were likely snared instead of pursued.
... Parenchymatous underground storage organs (USOs) have a long history of exploitation by humans dating back at least 170000 years ago (Wadley et al., 2020) and are important staple crops for billions of people globally today (FAO, 2021). These organs include anatomically distinct bulbs, corms, rhizomes, roots and tubers Hather, 2000). ...
Article
Full-text available
Archaeobotanical evidence for the exploitation of vegetatively propagated underground storage organs (USOs) in the tropical regions of Australia, Southeast Asia and the Pacific is currently limited. Although there have been several key studies of archaeological parenchyma published in the past two decades, systematic application of identification methods for vegetatively propagated crop species utilising charred, desiccated or waterlogged remains of parenchymatous tissue is not undertaken on a regular basis. Here, microCT imaging technology is used to compile a three‐dimensional virtual reference collection of parenchymatous tissues for five key USO species known to have been extensively cultivated by people in these regions. The five species are Dioscorea alata , Dioscorea esculenta , Colocasia esculenta , Alocasia macrorrhiza and Ipomoea batatas . These reference samples are used to illustrate the character of the virtual, microCT derived reference collection, and they also capture inter‐species differentiation and intra‐species morphological variation characteristic of many tuberous root crops.
... The list of commercial products containing African potato plant extracts seem to be expanding in horticultural practices [78]. This, together with increased market value and global trade, has negative long-term impacts on plant preservation [79]. ...
Article
South Africa, a country considered affluent in nature, ranks third in global biodiversity and encompasses approximately 9% of higher plants on planet Earth. Many indigenous plants have been utilised as herbal medicine, proving successful in treating numerous ailments. From the common cold to pandemic maladies such as COVID-19 in the 21st century and the treatment of incurable diseases, South African inhabitants have found great promise in the healing properties of these plants. Phytomedicine is a rapidly evolving topic, with in-depth bioactive composition analysis, identifying therapeutic action mechanisms, and disease prevention. While we are now poised to take advantage of nature’s medicine cabinet with greater scientific vigour, it remains critical that these practises are done with caution. Overharvesting significantly impacts biodiversity and cultivation practices amidst the beautiful nature of these nutraceuticals. This book chapter focuses on the therapeutic potential of commonly used South African medicinal plants, their ethnopharmacological properties, and how we can conserve this treasure cove we call home for future generations.
... Analyses of preserved botanical remains yield excitingly detailed insights into the integral role that plants played in prehistoric human life, as plants provide food, medicine, raw materials, and fuel (Shipley and Kindscher, 2016;Hardy, 2018Hardy, , 2019. Evidence for the Paleolithic human diet gleaned from plant fragments, phytoliths, and microfossils, as well as biomarkers from food preparation tools and dental calculus, has shifted the assumption of a largely animal-based diet to one that included a range of plants (Revedin et al., 2010;Power et al., 2018;Wadley et al., 2020). Specifically, DNA, chemical biomarkers, and starch grains extracted from dental calculus at several Neanderthal sites indicate ingestion of plant foods (Hardy et al., 2012;Salazar-García et al., 2013;Weyrich et al., 2017;Power et al., 2018). ...
Article
Current knowledge about Paleolithic human plant use is limited by the rare survival of identifiable plant remains as well as the availability of methods for plant detection and identification. By analyzing DNA preserved in cave sediments, we can identify organisms in the absence of any visible remains, opening up new ways to study details of past human behavior, including plant use. Aghitu-3 Cave contains a 15,000-yearlong record (from ∼39,000 to 24,000 cal BP) of Upper Paleolithic human settlement and environmental variability in the Armenian Highlands. Finds from this cave include stone artifacts, faunal remains, bone tools, shell beads, charcoal, and pollen, among others. We applied sedimentary ancient DNA (sedaDNA) metabarcoding to the Aghitu-3 sedimentary sequence and combined this with pollen data to obtain a temporal reconstruction of plant assemblages. Our results reveal a stratification of plant abundance and diversity where sedaDNA reflects periods of human occupation, showing higher diversity in layers with increased human activity. Low pollen concentrations combined with high sedaDNA abundance indicate plant remains may have been brought into the cave by animals or humans during the deposition of the lower two archaeological horizons. Most of the recovered plants are reported to be useful for food, flavor, medicine, and/or technical purposes, demonstrating the potential of the environment around Aghitu-3 Cave to support humans during the Upper Paleolithic. Moreover, we identified several specific plant taxa that strengthen previous findings about Upper Paleolithic plant use in this region (i.e., for medicine and the manufacturing and dyeing of textiles). This study represents the first application of plant sedaDNA analysis of cave sediments for the investigation of potential plant use by prehistoric humans.
... Past cognition can be inferred by analysing the material culture prehistoric populations have left behind, under the assumption that behavioural patterns reflect cognitive processes. A wide range of past behaviours have been investigated in this perspective, such as subsistence strategies [5,6], stone and bone tool-making [7][8][9][10][11][12][13][14][15], containers [16], pigments [17][18][19][20][21], tool hafting [22,23], mortuary practices [24,25], ornamental objects [26][27][28], engraving and painting of cave walls and objects [29,30]. More recently, past cognition has become the subject of interdisciplinary research combining archaeological data with methods and concepts from neuroscience [31][32][33]. ...
Article
Full-text available
It has been suggested that engraved abstract patterns dating from the Middle and Lower Palaeolithic served as means of representation and communication. Identifying the brain regions involved in visual processing of these engravings can provide insights into their function. In this study, brain activity was measured during perception of the earliest known Palaeolithic engraved patterns and compared to natural patterns mimicking human-made engravings. Participants were asked to categorise marks as being intentionally made by humans or due to natural processes (e.g. erosion, root etching). To simulate the putative familiarity of our ancestors with the marks, the responses of expert archaeologists and control participants were compared, allowing characterisation of the effect of previous knowledge on both behaviour and brain activity in perception of the marks. Besides a set of regions common to both groups and involved in visual analysis and decision-making, the experts exhibited greater activity in the inferior part of the lateral occipital cortex, ventral occipitotemporal cortex, and medial thalamic regions. These results are consistent with those reported in visual expertise studies, and confirm the importance of the integrative visual areas in the perception of the earliest abstract engravings. The attribution of a natural rather than human origin to the marks elicited greater activity in the salience network in both groups, reflecting the uncertainty and ambiguity in the perception of, and decision-making for, natural patterns. The activation of the salience network might also be related to the process at work in the attribution of an intention to the marks. The primary visual area was not specifically involved in the visual processing of engravings, which argued against its central role in the emergence of engraving production.
... MSA research in southern Africa is often governed by attempts to reconstruct human cultural evolution on a broad scale by detecting significant changes in material culture and tracing their earliest appearance (e.g. Henshilwood et al. 2001aHenshilwood et al. , 2011Marean et al. 2007;Backwell et al. 2008;Brown et al. 2009Brown et al. , 2012Vanhaeren et al. 2013;Schmidt and Mackay 2016;Brooks et al. 2018;Wadley et al. 2020). Since the Still Bay and Howiesons Poort industries have provided the vast majority of the cultural innovations of the MSA mentioned previously, these technocomplexes have received considerable attention. ...
Article
The end of the Middle Stone Age in southern Africa, often called the final MSA (∼40–28 ka), represents one of the most understudied technocomplexes in this part of the world. Researchers have often focused on earlier time periods associated with Marine Isotope Stage 4 or have emphasised the transition between the Middle and the Later Stone Age. Thus, the final MSA has been poorly understood and, at least in KwaZulu-Natal, only a few chrono-cultural markers called hollow-based points are known for it. Since 2016, excavations at Umbeli Belli rock shelter have produced new insights into this period. The site provides one of the most accurately dated sequences for the final MSA, spanning four geological horizons, respectively GH7, GH8, GH9 and GH10, that date to between 29.9 ± 2.3 and 40.3 ± 3.5 ka. Significant technological and typological variations are evident between those horizons, raising questions about the mechanisms behind them. A direct comparative analysis with the final MSA layers Coffee – Espresso at Sibhudu, which date to ∼38 ka, places these results in the regional archaeological context. The analysis shows first that the final MSA encompasses diachronic variability within relatively short time frames at Umbeli Belli. Secondly, it reveals several distinct chronological discrepancies between Sibhudu and Umbeli Belli. A detailed review of the environmental setting of the research area helps to explain these changes.
... The past two decades of intense research have shown that many of the innovations and behavioral changes once thought to originate in the LSA appeared already several millennia before, associated with clear MSA technologies such as at Blombos Cave (d'Errico et al., 2005;Henshilwood et al., 2001a, b), Diepkloof (Texier et al., 2010), Sibhudu (Conard et al., 2012;d'Errico et al., 2008;Wadley, 2005a;Wadley and Mohapi, 2008), Panga Ya Seidi ), or Border Cave (d'Errico et al., 2012Grün and Beaumont, 2001;Wadley et al., 2020). These finds suggest that populations of Late Pleistocene Homo sapiens living on the African continent were not only biologically (Dusseldorp et al., 2013;Rightmire and Deacon, 1991;Wang et al., 2009) but also from a cultural and cognitive perspective similar to modern-day hunter gatherers (Bradfield et al., 2020;Haidle, 2010;Henshilwood et al., 2011;Kandel et al., 2016;Lombard and Haidle, 2012;Wadley et al., 2009). ...
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The MSA/LSA transition is a major shift in the African archaeological record, but questions on its beginning remain debated. In southern Africa, most sites suggest an origin of LSA technology after about 30.000 years BP. The single exception is Border Cave situated at the border between South Africa and Eswatini, with surprisingly old dates of ∼43.000 BP associated with an LSA-like bipolar quartz assemblage. While many researchers now consider Border Cave to represent the origin of the LSA in southern Africa, these findings lack proper contextualization with regional lithic and chronometric data. Here we pursue the question whether Border Cave provides firm evidence for the source of LSA technology that later spread to the rest of southern Africa. To test between different hypotheses, we provide new chronometric and lithic data from the site of Sibebe, situated in the highveld of Eswatini only 100 km distant to Border Cave, and contextualize these results with nearby localities. Eswatini represents an ideal study area as it features many excavated sites but remains heavily understudied, rarely appearing in comparative MSA/LSA research. Our analyses at Sibebe identify two distinct groups of MSA lithic assemblages dated to between 43.000 and 27.000 cal BP by C14, overlain by a late Holocene LSA industry. The latest MSA expression at Sibebe features finely shaped unifacial and bifacial points without trends towards LSA technology. A comparative review of sites in Eswatini, South Africa, and Mozambique confirms similar MSA industries post-dating 30.000 BP but finds no evidence for LSA technologies before ∼27.000 BP. These findings suggest that Border Cave cannot mark the origin of the much later LSA in southern Africa, but rather signifies an early set of locally specific behavioral innovations that appeared and disappeared again. These findings have important implications for the spread of new technologies and our understanding of cultural evolutionary trajectories in southern Africa and beyond.
... Whilst some of the technologies required for processing plant foods were unique to northern Sahul for much of the Pleistocene (Hiscock et al., 2016), the evidence for the long-term use of a broadspectrum plant diet at Madjedbebe is consistent with a growing body of research indicating that broad-based plant exploitation, including the cooking and processing of seeds and underground storage organs, was part of the adaptive toolkit of early humans (Barker et al., 2007;Henry et al., 2014;Inchley et al., 2016;Larbey et al., 2019;Madella et al., 2002;Mercader, 2009;Summerhayes et al., 2010;Wadley et al., 2020;Weiss et al., 2004). This evidence significantly predates the expectations of Kent V. Flannery's (1969) 'Broad Spectrum Revolution', which hypothesises the incorporation of hard-to-process plant foods into hunter-gatherer diets only in the millennia immediately prior to the advent of agriculture. ...
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The plant macrofossil assemblage from Madjedbebe, Mirarr Country, northern Australia, provides insight into human-plant relationships for the ∼65,000 years of Aboriginal occupation at the site. Here we show that a diverse diet of fruits, nuts, seeds, palm and underground storage organs was consumed from the earliest occupation, with intensive plant food processing in evidence. The diet varied through time as foraging strategies were altered in response to changes in environment and demography. This included a broadening of the diet during drier glacial stages, as well as changes in the seasonal round and incorporation of new foods with the formation of freshwater wetlands following sea level rise in the late Holocene. The foundations of the economy evidenced at Madjedbebe include seasonal mobility, a broad diet and requisite plant processing and grinding technologies, all of which are maintained throughout the entire timespan of occupation. This points to a resilient economic system in the face of pronounced environmental change.
... For later periods, macrorests attributed to legumes, seeds, and fruits are reported from the Middle Palaeolithic layers of Kebara Cave, Mt. Carmel in the Levant (Lev et al., 2005), whereby in Mozambique, sorghum starch grains might have been already part of the diet of modern humans settled at Ngalue cave (105-42 ka BP, Mercader, 2009), and at Klasies River and Border cave, South Africa, charred rhizomes are reported in the ashes of hearths starting with the MIS 5 (120-65 ka BP; Larbey et al., 2019;Wadley et al., 2020). If we consider the Australian continent, at the Madjedbebe rockshelter, charred rhizomes have been recognized in layers 65,000 years old, by comparing these with modern-day plants still present near the site (Florin et al., 2020), as already observed for the South African sites. ...
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A selection of five ground stones from Pontic Steppe sites dating back to the Early Upper Palaeolithic (EUP) was used as test-cases to be analysed by combining wear-traces and use-related biogenic residues (U-RBR). The artifacts studied can be termed “legacy” objects, excavated even many decades ago and kept in museum storage facilities. This type of storage might be considered putatively prone to contamination. The multidimensional contextual approach we designed integrates the structural analysis of biogenic residues by means of visual light optical and electronic beams microscopy (OM/VLM and SEM) coupled with FTIR microspectroscopy and imaging (using both conventional and synchrotron infrared sources). SEM and FTIR are meant to provide high resolution morphological and chemical profiles and their coupled analysis revealed the presence of starch grains from the used areas of the stone tools. The goal of this paper is to present a reasoned streamlined procedure to collect appropriate samples suitable to detect the presence of ancient starches from ground stones tools recovered in museum collections.
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Lucy Lloyd's personal copy of Bleek's 1875 "A Brief Account of Bushman Folk-Lore and Other Texts" which is now housed at the University of Texas is described and hand-written notes transcribed.
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Underground storage organs are poorly preserved in the archaeological record, and as a result their contribution to the diet of ancient societies is poorly understood. Starch grain analysis is a well-established methodology used in archaeology to reveal the use of various plants, including tubers, rhizomes, and roots. This paper presents the results of a study of starch grains recovered from millstones from various archaeological sites located in the Paris Basin. Our results highlight the use of tubers by these first agricultural populations at the beginning of the Neolithic (Linearbandkeramik and Blicquy-Villeneuve-Saint-Germain; 5200-4700 BC), providing new data on their contribution to the diet of agro-pastoral societies.
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Thus far, most researchers have focused on the cognition of fire use, but few have explored the cognition of firemaking. With this contribution we analyse aspects of the two main hunter-gatherer firemaking techniques—the strike-a-light and the manual fire-drill—in terms of causal, social and prospective reasoning. Based on geographic distribution, archaeological and ethnographic information, as well as our cognitive interpretation of strike-a-light firemaking, we suggest that this technique may well have been invented by Neanderthal populations in Eurasia. Fire-drills, on the other hand, represent a rudimentary form of a symbiotic technology, which requires more elaborate prospective and causal reasoning skills. This firemaking technology may have been invented by different Homo sapiens groups roaming the African savanna before populating the rest of the globe, where fire-drills remain the most-used hunter-gatherer firemaking technique.
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The Middle Pleistocene is associated with a new level of technology, Mode 2, commonly called the Acheulean Culture. This appears in Africa and Western Asia, but only later in Europe and not at all in Asia. It appears to be an indicator of communication, if not actual migration, among these areas. Climate swings, particularly in Europe, suggest that some forms of shelter and clothing had been invented, but more direct evidence is scarce. Controlled fire appears about a million years ago, but what traces are found suggest it was not widely used. Hunting of larger animals is well documented in the Middle Pleistocene employing weaponry such as lances and throwing spears. Meat consumption appears to have increased in importance, especially in temperate Europe. Strangely, Asian populations did not follow the same behavioral trends.
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This contribution focuses on the recently excavated lithic assemblage from Border Cave Members 1 RGBS, 3 BS, 2 WA and 2 BS. These members were attributed by Beaumont to the Howiesons Poort and post-Howiesons Poort Industries of the southern African Middle Stone Age. Here we consider lithics as indicators of cultural behaviour, site formation processes, and occupation intensity. As such, the assemblage is explored in depth through lithic technology attribute analysis, particle size distribution, and spatial analysis. These lines of inquiry follow the new allostratigraphic divisions proposed for the deposits by Stratford and colleagues. Results show that the lower members share a degree of similarity in terms of flaking strategies and raw material selection, whereas the upper members record a dissimilar set of features, with differentiation between them. The technological and spatial analyses suggest the sporadic presence of different groups at the site, each characterized by distinct sets of knapping techniques and methods but employing related knapping strategies and using similar raw materials. The interpretation of these members as a palimpsest of sporadic occupation is supported by the low number of archaeological finds by sediment volume. The evidence at hand does not support a Howiesons Poort attribution for the lowest members. The data show that lithics from part of Member 2 WA and 2 BS are in accordance with those of the recently defined Sibudan technocomplex.
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Large graminoid species, which often dominate wetland ecosystems with extensive and dense formations, are among the most indicative plants from the first human settlements, where they have been used (even transformed) for various functions ranging from food, cordage, weaving and other utilities. Wetland large graminoid foraging today represents one of the rarest and most archaic customs still in existence, as they have frequently disappeared following changes in society or the disappearance of marshes. These customs have (almost) disappeared in Europe, especially in Italy, following socio-economic changes and wetland reclamation; remaining uses can generally only be found in prehistoric traces. This research in Agro Peligno documents and describes for the first time the remains of these prehistoric uses, which are related to the ancient Peligni (or Paeligni) people. The data collected in the current field study were later compared with food uses of graminoids arising from a large spectrum of archaeological, ethnobotanical, and folkloric literature from other European areas, in a large sense. Problems and outlook regarding the loss of this traditional knowledge are also briefly discussed.
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The African Middle Stone Age (MSA) is the period in human history spanning roughly from 300,000 until 30,000 years ago. Here, we focus on the archaeological record of South Africa, with occasional glimpses at neighboring countries (Eswatini, Lesotho, Namibia). During this time, modern humans evolved in Africa and brought forth a number of key innovations, including art and symbolism, personal ornaments, burial practices, and advanced methods of tool production using different raw materials such as stone, wood, or bone. The MSA is subdivided into several substages based on regional chrono-cultural differences, such as MSA II or Mossel Bay, Still Bay, Howiesons Poort, Sibudan, and the final MSA. Previous research has tended to concentrate on just two of those stages, namely, the Still Bay and Howiesons Poort, as they were considered to be pinnacles of innovation. In the past years, however, assemblages from other periods have gained increasing attention. Some of the major research questions include the nature and timing of both the onset and end of the MSA. The focus on diachronic cultural dynamics not only related to the Still Bay and Howiesons Poort techno-complexes and the increasing awareness of regional diversification during different phases, especially during Marine Isotope Stage 3 (57,000–29,000 years ago), but also to the inherent problems arising from them.
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Neanderthals ate plants, they self-medicated using a range of medicinal plants, and they used complex material processing methods to construct composite objects with plant materials. While the evidence for the consumption of plants as food, medicine, and raw materials by Neanderthals is not abundant, by using a combination of direct and indirect methods, including recovery and analysis of macrobotanical and microfossil remains, biomolecular and genetic evidence and use wear patterns on teeth and tools, reconstruction of the basic premises of their plant use can be partially reconstructed. This provides nuanced insights into their cognitive and technological abilities and the deep ecological knowledge that was the foundation of their existence.
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Globally, fire is a primary agent for modifying environments through the long-term coupling of human and natural systems. In southern Africa, control of fire by humans has been documented since the late Middle Pleistocene, though it is unclear when or if anthropogenic burning led to fundamental shifts in the region's fire regimes. To identify potential periods of broad-scale anthropogenic burning, we analyze aggregated Holocene charcoal sequences across southern Africa, which we compare to paleoclimate records and archaeological data. We show climate-concordant variability in mid-Holocene fire across much of the subcontinent. However, increased regional fire activity during the late Holocene (∼2000 BP) coincides with archaeological change, especially the introduction and intensification of food production across the region. This increase in fire is not readily explained by climate changes, but rather reflects a novel way of using fire as a tool to manage past landscapes, with outcomes conditioned by regional ecosystem characteristics.
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It has been suggested that engraved abstract patterns dating from the Middle and Lower Palaeolithic served as means of representation and communication. Identifying the brain regions involved in visual processing of these engravings can provide insights into their function. In this study, brain activity was measured during perception of the earliest known Palaeolithic engraved patterns and compared to natural patterns mimicking human-made engravings. Participants were asked to categorise marks as being intentionally made by humans or due to natural processes (e.g. erosion, root etching). To simulate the putative familiarity of our ancestors with the marks, the responses of expert archaeologists and control participants were compared, allowing characterisation of the effect of previous knowledge on both behaviour and brain activity in perception of the marks. Besides a set of regions common to both groups and involved in visual analysis and decision-making, the experts exhibited greater activity in the inferior part of the lateral occipital cortex, ventral occipitotemporal cortex, and medial thalamic regions. These results are consistent with those reported in visual expertise studies, and confirm the importance of the integrative visual areas in the perception of the earliest abstract engravings. The attribution of a natural rather than human origin to the marks elicited greater activity in the salience network in both groups, reflecting the uncertainty and ambiguity in the perception of, and decision-making for, natural patterns. The activation of the salience network might also be related to the process at work in the attribution of an intention to the marks. The primary visual area was not specifically involved in the visual processing of engravings, which argued against its central role in the emergence of engraving production.
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RESUMEN Border Cave es una cueva ubicada en las montañas de Lebombo, en la frontera entre la región de KwaZulu-Natal (Sudáfrica) y Esuatini. Las excavaciones arqueológicas se iniciaron en 1934, se retomaron en los 70’ y el sitio vuelve a investigarse en la actualidad. Border Cave es una ventana al conocimiento de la prehistoria sudafricana debido a la excelente preservación del material orgánico. El objetivo del presente trabajo es conocer este sitio arqueológico desde la experiencia de las autoras que han participado de la campaña del año 2019 haciendo énfasis en la relación entre el equipo de investigación y la comunidad local. ABSTRACT Border Cave is a cave located in the Lebombo Mountains, on the border between KwaZulu-Natal region (South Africa) and Eswatini. The archaeological excavations in around 1934, resumed during the 70’s, and the site is currently being investigated. Border Cave, due to its excellent preservation of the material, is a window to the knowledge of South African prehistory. The aim of this work is to observe this archaeological site from the experience of the authors after their participation in the 2019 field work, with an emphasis on the relationship between the research team and the local community.
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Commelina diffusa Burn. f. and Commelina erecta L. (Commelinaceae) are known from tropical and subtropical regions of the world. In the present work, the leaf epidermal characters, midrib and stem anatomy were studied in relation to their taxonomic values. Voucher specimens collected from different parts of Abia, Rivers, Bayelsa and Delta States were analysed. The samples were fixed in FAA, dehydrated in series of ethanol (50%, 70% and 90%), peeled/sectioned, stained in 2% aqueous solution of Safranin O, counter stained in Alcian blue for about 3-5 minutes, mounted in glycerine, viewed and micro-photographed using Leica WILD MPS 52 microscope camera on Leitz Diaplan microscope. Comparative foliar epidermal features, midrib and stem anatomical characters of the two Commelina species indicated that the leaf epidermal characters showed close similarity among the species though with few distinguishing features while the anatomical features of the midrib and lamina could be used to distinguish these species.
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This paper presents an integrated methodology for the analysis of archaeological remains of cereal meals, based on scanning electronic microscopic analyses of microstructures of charred food fragments from Neolithic Çatalhöyük (Turkey). The remains of cereal foods as ‘bread-like’ or ‘porridge-like’ small charred lumps of various amalgamated plant materials are frequently recovered from Neolithic and later archaeological sites in southwest Asia and Europe. Cereal food remains have recently attracted interest because the identification of their plant contents, the forms of food that they represent and the methods used in their creation can provide unique information about ancient culinary traditions and routine food processing, preparation and cooking techniques. Here, we focus on three methodological aspects: (1) the analysis of their composition; (2) the analysis of their microstructure to determine preparation and cooking processes; (3) the comparison with experimental reference materials. Preliminary results are presented on the botanical composition and cooking processes represented by the charred cereal preparations found at Neolithic Çatalhöyük (Turkey), for example cereals processed into bread, dough and/or porridge.
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Recent genomic data have revealed multiple interactions between Neanderthals and modern humans, but there is currently little genetic evidence regarding Neanderthal behaviour, diet, or disease. Here we describe the shotgun-sequencing of ancient DNA from five specimens of Neanderthal calcified dental plaque (calculus) and the characterization of regional differences in Neanderthal ecology. At Spy cave, Belgium, Neanderthal diet was heavily meat based and included woolly rhinoceros and wild sheep (mouflon), characteristic of a steppe environment. In contrast, no meat was detected in the diet of Neanderthals from El Sidrón cave, Spain, and dietary components of mushrooms, pine nuts, and moss reflected forest gathering. Differences in diet were also linked to an overall shift in the oral bacterial community (microbiota) and suggested that meat consumption contributed to substantial variation within Neanderthal microbiota. Evidence for self-medication was detected in an El Sidrón Neanderthal with a dental abscess and a chronic gastrointestinal pathogen (Enterocytozoon bieneusi). Metagenomic data from this individual also contained a nearly complete genome of the archaeal commensal Methanobrevibacter oralis (10.2× depth of coverage)-the oldest draft microbial genome generated to date, at around 48,000 years old. DNA preserved within dental calculus represents a notable source of information about the behaviour and health of ancient hominin specimens, as well as a unique system that is useful for the study of long-term microbial evolution.
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Significance Our knowledge of the diet of early hominins derives mainly from animal skeletal remains found in archaeological sites, leading to a bias toward a protein-based diet. We report on the earliest known archive of food plants found in the superimposed Acheulian sites excavated at Gesher Benot Ya‘aqov, Israel. These remains, some 780,000 y old, comprise 55 taxa, including nuts, fruits, seeds, vegetables, and plants producing underground storage organs. They reflect a varied plant diet, staple plant foods, seasonality, and hominins’ environmental knowledge and use of fire in food processing. Our results change previous notions of paleo diet and shed light on hominin abilities to adjust to new environments and exploit different flora, facilitating population diffusion, survival, and colonization beyond Africa.
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Humans have more copies of amylase genes than other primates. It is still poorly understood, however, when the copy number expansion occurred and whether its spread was enhanced by selection. Here we assess amylase copy numbers in a global sample of 480 high coverage genomes and find that regions flanking the amylase locus show notable depression of genetic diversity both in African and non-African populations. Analysis of genetic variation in these regions supports the model of an early selective sweep in the human lineage after the split of humans from Neanderthals which led to the fixation of multiple copies of AMY1 in place of a single copy. We find evidence of multiple secondary losses of copy number with the highest frequency (52%) of a deletion of AMY2A and associated low copy number of AMY1 in Northeast Siberian populations whose diet has been low in starch content.
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The leaf and corm morphology and anatomy of representatives of the genera Spiloxene, Pauridia and Empodium were studied. The corms are annual and tunicated, except in the group Aquaticae of Spiloxene. They are swollen stems consisting of a number of internodes. In Spiloxene and Pauridia the roots grow from the base of the corm, while in Empodium they develop from the sides. The epidermis of the older corms is replaced by several layers of thin-walled cork. Characters of the corm coverings are used to divide Spiloxene into six groups. Four leaf forms are recognised namely carinate, terete, canaliculate and plicate. Multicellular processes and unicellular hairs occur occasionally.The leaf stomata are paracytic. Most species have mucilage canals containing pectic compounds of mucopoly­saccharides. The vascular bundles have complete or incomplete bundle sheaths and larger bundles have sclerenchyma caps.
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We propose that plant foods containing high quantities of starch were essential for the evolution of the human phenotype during the Pleistocene. Although previous studies have highlighted a stone tool-mediated shift from primarily plant-based to primarily meat-based diets as critical in the development of the brain and other human traits, we argue that digestible carbohydrates were also necessary to accommodate the increased metabolic demands of a growing brain. Furthermore, we acknowledge the adaptive role cooking played in improving the digestibility and palatability of key carbohydrates. We provide evidence that cooked starch, a source of preformed glucose, greatly increased energy availability to human tissues with high glucose demands, such as the brain, red blood cells, and the developing fetus. We also highlight the auxiliary role copy number variation in the salivary amylase genes may have played in increasing the importance of starch in human evolution following the origins of cooking. Salivary amylases are largely ineffective on raw crystalline starch, but cooking substantially increases both their energy-yielding potential and glycemia. Although uncertainties remain regarding the antiquity of cooking and the origins of salivary amylase gene copy number variation, the hypothesis we present makes a testable prediction that these events are correlated.
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The genus Hypoxis L. (Hypoxidaceae) in the East Tropical A complete key with full descriptions and distributions of all known Hypoxis taxa found in the East Tropical Africa is presented in the monograph. The morphology of all species, subspecies and varieties is described, including such important taxonomic characters for this genus like tuber flesh color, tunic type, indumentum and seed testa sculpture. A succulent leaf structure is described for H. kilimanjarica var. prostrata. The anatomical studies were conducted as a part of taxonomical analysis. They have positively evaluated a taxonomic significance of leaf anatomy characters, such as succulent structure, occurrence of bulliform cells in epidermis outside the keel zone, type and distribution of trichomes. The studies of the Hypoxis leaf anatomy added new data concerning anatomical differentiation of the cataphylls and the inner leaves. Also differentiated mesophyll and simultaneous presence of different types of stomata on one leaf are reported. It has been shown that in some species mucilage canals are present in the inner leaves and that this character is not constant. The number of vascular bundles, which can be determined only on the basis of a leaf section, is useful only in species with a small number of veins, not increasing with a plant age. Because of lack of constancy in distribution, number of stomata accessory cells cannot be used as a diversifying character for the East African species of Hypoxis. The wax crystals are revealed to exist in many species of Hypoxis. The anatomical characters of scapes were also studied in a taxonomic context. A sclerenchyma distribution, as well as number of vascular bundles can be used for a species determination. The presence of sclerenchyma prevents the scapes from bending down after anthesis. The studies of phenology revealed that there are two groups of taxa, one with a resting period and the other without it. It is connected with a climate in which the species occurs. The study of distribution maps of the species occurring in the East Africa are provided for this area, as well as for their entire range. This new knowledge, along with a revision of literature data, led to a new conclusion as to a number of all Hypoxis species in Africa, which is now estimated to be 55. The revision demonstrates that distribution of many of the Hypoxis species is connected with White's phytochoria. It proves that not only South Africa, but also the Zambesian Region is a very important center of diversity of this genus. The number of endemic taxa of Hypoxis for the East Tropical Africa is very low, including only one species and one subspecies. Additionally, a study of vertical ranges of Hypoxis is presented. It reveals that most of the species in East Africa grow in the mountains and they show preferences of dispersal in particular altitudinal levels. The analysis of the vertical distribution within the entire ranges of different taxa has showed differences in the altitudinal position depending on the geographic location. The human influence on Hypoxis is studied in terms of their use in folk medicine and believes. Most of the species of Hypoxis survive quite well in East Africa, being a visible component of various types of grasslands. Some species however are under threat of extinction. This is due to their incapability of surviving in changed habitats, especially in shade of cultivated plants. Another threat is a large-scale collection of species believed to cure the HIV, or sold as a substitute of similar taxa, assumed to possess such qualities. The IUCN categories are proposed for the East African taxa of Hypoxis.
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A seco ndary thickening meristem is record ed for the first time in some herbaceous taxa of Asparagales (Herreria montevide nsis and Thysanotus sp inige r), and the new records are assessed in a systematic co ntext. All monocotyl edons lack a vascular cambium, which is typicall y a single per sistent row of cells producing phloem centrifugally and xylem ce ntripetally. However, some monocotyled ons achieve stem thicken ing by means of a different type of later al meristern, either a primary thickening meristem (PTM) near the apex, together with diffuse secondary growth (as in palm s). or a secondary thicken ing meristern (STM) further away from the apex (as in some Asparagales; see Rudall 1991, for review). The PTM and STM are probabl y developmenlally related, although there is complex tissue involvement. In taxa with an STM, the PTM and STM may someti mes be longitudi­ nally co ntinuous, at least at some stage in the life cycle (Steven son 1980). Virtually all monocotyledons have a PTM , but among tree-forming or woody taxa this has developed along different lines, probabl y more than once, either as an exten­ sive apical PTM, as in palms, or as an STM, in some Asparagales (see below). The PTM and STM are not homologous with the vascular cambium, as they are tiered (etagen) men stems which produce distinct vascular bundles (of both xylem and phloem) in a parench ymatou s ground tis sue (Fig. 1-3), mainly ce ntripe tally. There are record s of a PTM in some dicotyledon s and other plant groups (DeMaso n 1983), although the hom ology of these requ ires testing . All unequi vocal records of an STM are in the asparagoid clade (sensn Cha se et al. 1995), which compri ses Dahlgren et al.'s (1985) order Asparagales, together with a few membe rs of their Liliales, such as Iridaceae (Rudall 1991). A STM has been re­ ported in several tree-forming and shru bby asparagoid s: Aga ve, Aloe, Beaucarnea, Calibanus, Cordyiine, Dasylirion, Dra caena, Furcra ea, Klatti a, Nive nia, No lina, Pleomel e, Sansevieria, Witsenia, Xanth orrh oea and Yucca, and also in some more her­ baceou s taxa with a thick ' woody' rhizome or stem, such as Aphyllanthes, Gasteria.
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Cyperaceae are usually perennial, with underground stems mainly rhizomatous, however, other stem types may also occur, such as corms and tubers. The underground stems of five Cyperaceae species were examined. Cyperus rotundus and Fuirena umbellata have plagiotropic rhizomes, while C. esculentus, C. odoratus, Hypolytrum schraderianum and Bulbostylis paradoxa have orthotropic rhizomes. Corms occur in C. rotundus and C. esculentus, and stolons in C. esculentus. The primary body originates from the activity of the apical meristem and later, from the primary thickening meristem (PTM). Secondary growth results from secondary thickening meristem (STM) activity, and occurs in rhizomes of H. schraderianum, B. paradoxa, C. odotarus and F. umbellata. The procambium and the PTM give rise to collateral bundles in H. schraderianum, and amphivasal bundles in the remaining species. The STM gives rise to the vascular system with the associated phloem and xylem. According to our results, the concept of stem type in Cyperaceae depends on external morphology, function, life phase, activity of the thickening meristems and the relative amount of parenchyma.
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Rhizome structure was observed in most rhizomatous genera of the Iridaceae to determine its taxonomic applications within the family and to determine the evolution of the secondary thickening meristem (STM) within the group. An STM occurs in all species of Patersonia examined, similar to that described in Nivenia, Witsenia, and Klattia, but in no other genera of Iridaceae. Most genera have a primary thickening meristem (PTM) near the stem apex, which produces some primary stem vasculature, adventitious roots, and in some species, notably in Orthrosanthus chimboracensis, an extensive pericyclic vascular plexus. The results show that the two subtribes Aristeineae and Sisyrinchiineae are closely allied, and the STM may have evolved from a more extensive PTM or may be retained as a primitive character.
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Four varieties within H. angustifolia Lamarck. the most common species of the genus Hypoxis L. in Africa. are recognized. based in part oil difference, of seed testa features between populations ill Mauritius. Madagascar. and continental Africa. SEM micrographs of' seeds of H. angustifolia var. angustifola, variety luzuloides (Robyns & Tournay) Wiland, and variety madagascariensis Wiland are included, and full descriptions of the above varieties as well as H. angustifolia var. buchananii Baker are given.
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Background: It had long been thought that a lateral meristem, the so-called primary thickening meristem (PTM) was responsible for stem thickening in monocotyledons. Recent work has shown that primary thickening in the stems of monocotyledons is due to the meristematic activity of both the endodermis and the pericycle. Aims: The aim of this work is to answer a set of questions about the developmental anatomy of monocotyledonous plants: (1) Do the stem apices of monocots have a special meristematic tissue, the PTM? (2) Are the primary tissues of the stem the same as those of the root? (3) Is there good evidence for the formation of both the cortex and the vascular tissue from a single meristem, the PTM, in the shoot and from two distinguishable meristems in the root? (4) If the PTM forms only the cortex, what kind of meristem forms the vascular tissue? Methods: Light microscopy was used to examine stem and root anatomy in 16 species from 10 monocotyledonous families. Results: It was observed that radially aligned cortical cells extend outwards from endodermal initial cells in the cortex of the roots and the stems in all the species. The radial gradation in size observed indicates that the cortical cells are derivatives of a meristematic endodermis. In addition, perfect continuity was observed between the endodermis of the root and that of the stem. Meristematic activity in the pericycle gives rise to cauline vascular bundles composed of metaxylem and metaphloem. Conclusion: No evidence was obtained for the existence in monocotyledons of a PTM. Monocotyledons appear to resemble other vascular plants in this respect.