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Research on Palaeolithic hunter-gatherer diet has focused on the consumption of animals. Evidence for the use of plant foods is comparatively limited but is rapidly expanding. The authors present an analysis of carbonised macro-remains of processed plants from Franchthi Cave in the Aegean Basin and Shanidar Cave in the north-west Zagros Mountains. Microscopic examination of the charred food remains reveals the use of pounded pulses as a common ingredient in cooked plant foods. The results are discussed in the context of the regional archaeobotanical literature, leading the authors to argue that plants with bitter and astringent tastes were key ingredients of Palaeolithic cuisines in South-west Asia and the Eastern Mediterranean.
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Research Article
Cooking in caves: Palaeolithic carbonised plant food
remains from Franchthi and Shanidar
Ceren Kabukcu1,
*
, Chris Hunt2, Evan Hill3, Emma Pomeroy4,
Tim Reynolds5, Graeme Barker4& Eleni Asouti1
1
Department of Archaeology, Classics and Egyptology, University of Liverpool, UK
2
Research Centre in Evolutionary Anthropology and Palaeoecology, Liverpool John Moores University, UK
3
School of Natural and Built Environment, Queens University Belfast, UK
4
Department of Archaeology, University of Cambridge, UK
5
Department of History, Classics and Archaeology, Birkbeck University of London, UK
* Author for correspondence C.Kabukcu@liverpool.ac.uk
Research on Palaeolithic hunter-gatherer diet has
focused on the consumption of animals. Evidence
for the use of plant foods is comparatively limited
but is rapidly expanding. The authors present an ana-
lysis of carbonised macro-remains of processed plants
from Franchthi Cave in the Aegean Basin and Shani-
dar Cave in the north-west Zagros Mountains.
Microscopic examination of the charred food remains
reveals the use of pounded pulses as a common ingre-
dient in cooked plant foods. The results are discussed
in the context of the regional archaeobotanical litera-
ture, leading the authors to argue that plants with bit-
ter and astringent tastes were key ingredients of
Palaeolithic cuisines in South-west Asia and the East-
ern Mediterranean.
Keywords: South-west Asia, Eastern Mediterranean, Palaeolithic diet, prehistoric food preparation, hunter-
gatherers, archaeobotany
Introduction
The dietary choices and food preparation technologies of Palaeolithic hunter-gatherers are the
subject of much debate. Palaeolithic peoples have been portrayed, for example, as specialist
hunters focusing on large mammals (Richards & Trinkhaus 2009) or as generalist foragers
targeting easy-to-gather resources out of necessity, due to pressures on the availability of pre-
ferred animal prey (Stiner et al.2000; Speth 2010). In this article, we focus on the dietary
contribution of plant foods. In a calorie-driven interpretation of Palaeolithic diet, plants
Received: 22 March 2022; Revised: 8 June 2022; Accepted: 17 June 2022
© The Author(s), 2022. Published by Cambridge University Press on behalf of Antiquity Publications Ltd. This is an
Open Access article, distributed under the terms of the Creative Commons Attribution licence (https://
creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, distribution, and reproduction in any
medium, provided the original work is properly cited.
Antiquity 2023 Vol. 97 (391): 1228
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are categorised as low-rankedresources, due to the time- and labour-intensive nature of gath-
ering and processing them. Consequently, scholarly emphasis has focused on the signicance
of the more carbohydrate-rich plant foods that were easy to collect and prepare as potential
dietary staples (Prado-Nóvoa et al.2017; Hardy et al.2022).
Archaeobotanical data from pre-agricultural sites in South-west Asia and the Eastern
Mediterranean, however, indicate a reliance on a much wider range of plant foods than
just starch-rich tubers and grasses (e.g. Martinoli 2004; Weiss et al. 2004; Lev et al.2005;
Asouti et al.2018,2020; Caracuta et al.2021). Almost all sites from these regions dating
to the Middle and Upper Palaeolithic and the Epipalaeolithic/Mesolithic periods, for
example, provide evidence for the use of wild almonds, which contain high levels of cyano-
genic metabolites that can produce hydrogen cyanide (Asouti et al.2020; Caracuta et al.
2021). Several other plants also feature prominently in the regional archaeobotanical record,
including tannin-rich wild pistachios (terebinth), wild pulses (some containing neuro-toxic
compounds) and astringent wild mustards. Most of these plants require several preparation
steps to leach out unpalatable and/or toxic compounds prior to consumption. The long-term
and widespread use of almonds, terebinths and pulses therefore suggests that Palaeolithic
foragers developed processing technologies and associated food preparation practices that
enabled their routine safe consumption.
In this article, we report new evidence concerning the long-term histories of Palaeolithic
plant food use and associated food preparation practices from two multi-period sites:
Franchthi Cave (Greece) and Shanidar Cave (Iraqi Kurdistan). We focus on the analysis of
amorphous, charred plant aggregates retrieved from otation samples from the two sites;
some of these materials represent the earliest remains of their kind discovered to date in
South-west Asia and Europe. Such remains, often representing the charred residues of
food preparation (hereafter food remains), can provide direct evidence for the plant species
consumed, often combined in multi-component foods, as well as preparation methods
(Carretero et al.2017; Heiss et al.2017; Arranz-Otaegui et al.2018; Valamoti et al.
2021). Our results highlight the early exploitation of a diverse range of plant foods that
required specialised processing techniques, bringing to the fore the signicance of food
preparation and cooking practices in ancient human dietary practices.
Materials and methods
Franchthi Cave is located in the Argolid peninsula of southern mainland Greece. It was exca-
vated between 1969 and 1976 by T.W. Jacobsen of Indiana University and M.H. Jameson of
Pennsylvania University, under the auspices of the American School of Classical Studies in
Athens and in collaboration with the Greek Archaeological Service (Farrand 2000). Occupa-
tion at the site spans the Upper and Final Palaeolithic, Mesolithic and Neolithic (c. 38 000
6000 cal BP) (Farrand 2000; Asouti et al.2018). Sampling for archaeobotanical remains was
carried out by machine-assisted water otation. The non-wood charred plant macro-remains
were previously studied and published by Julie Hansen (1991). The presence of charred,
amorphous plant aggregates that potentially represent food remains was noticed during
anthracological analyses conducted by Eleni Asouti and Ceren Kabukcu at the University
of Liverpool in 2017 (Asouti et al. 2018). The four charred plant aggregates from Franchthi
Cooking in caves: Palaeolithic plant food remains from Franchthi and Shanidar
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Table 1. Summary of provenance, phasing and corresponding radiometric dates for the analysed food fragments.
Site
Fragment
context no.
Flotation
sample no. Phase Chronometric date range Illustrated in
Franchthi Cave H1A 167 H1A 167 FGP VI/VII 11.711.4 ka cal BP (Early Holocene onset) Figure 1
Franchthi Cave H1A 168 H1A 168 FGP VI/VII 11.711.4 ka cal BP (Early Holocene onset) Figure 2
Franchthi Cave H1A 172 H1A 172 FGP VI 12.911.7 ka cal BP (Younger Dryas) Figure 4
Franchthi Cave H1A 177 H1A 177 FGP V 13.112.9 ka cal BP (Bølling Allerød, GI-1A warm period) Figure 3
Shanidar Cave 636 4778 Baradostian 40 ka BP (MIS 3) Figure 9
Shanidar Cave 1812 5511 Baradostian >40 ka BP (MIS 3) Figure 5
Shanidar Cave 1823 5541 Baradostian >40 ka BP (MIS 3) Figure 7
Shanidar Cave 1866 5714 Initial Baradostian >40 ka BP (MIS 3) Figures 6 &8
Shanidar Cave 1924 5631 Mousterian 7075 ka BP
(likely MIS 5a)
Figure 10
Ceren Kabukcu et al.
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Cave examined here originate from two chrono-cultural phases: stratum T3, which is assigned
to the later phases of the Upper Palaeolithic (Mediterranean Gravettian), corresponding to
the Bølling-Allerød warm period (Franchthi General Phase (FGP) V, c. 13 10012 900 cal
BP), and the Final Palaeolithic (Epigravettian) strata, which are dated to the Younger
Dryas (FGP VI, c. 12 90011 700 cal BP) and the start of the Holocene (FGP VI/VII,
c. 11 70011 400 cal BP) (Hansen 1991; Farrand 2000; Asouti et al.2018)(Table 1).
Shanidar Cave, located on the western anks of the Zagros Mountains of Iraqi Kurdistan,
was originally excavated between 1951 and 1960 by Ralph and Rose Solecki of Columbia
University and colleagues (Solecki 1971). Since 2015, a team led by Graeme Barker has con-
ducted systematic excavations at the site (Reynolds et al.2015), during which the fragments
analysed in this study were collected. Five charred plant aggregates were recovered from
Upper Palaeolithic (Baradostian) and one further fragment from the Middle Palaeolithic
(Mousterian) deposits. Although the full radiometric dating programme is ongoing, the
Baradostian strata date to c. 42 00035 000 years ago (Reynolds et al.2018), which corre-
sponds to the later part of Marine Isotope Stage 3 (MIS 3). Regionally, the Baradostian
techno-cultural industry is interpreted as coeval with the Aurignacian of the European
Upper Palaeolithic, which is associated with Homo sapiens (Reynolds et al.2018; Shidrang
2018). In Iran, the Baradostian industry has been associated with anatomically modern
humans at the cave site of Eshkaft-e Gavi (Scott & Marean 2009), where it has been
dated to c. 42 00030 000 years agosomewhat later than the Baradostian assemblage of
Kaldar Cave, which is dated to 54 40046 050 cal BP (Bazgir et al.2017). Meanwhile,
the samples recovered from the Mousterian strata at Shanidar Cave probably date to
>70 00075 000 years ago, based on their broad stratigraphic association with the well-
known Neanderthal ower burialand the recently discovered Shanidar Zarticulated skel-
etal remains, dated to c. 73 000 BP (Pomeroy et al.2017,2020).
Charred plant aggregates may take the form of large, recognisable items of food (Heiss
et al.2017), carbonised crusts adhering to the walls of pottery vessels (Kubiak-Martens
et al.2015) or amorphous lumps, some of which could represent accidentally charred
food remains (Valamoti et al.2021; Bates et al.2022); microscopic examination is required
to conrm their interpretation as food remains and to identify their plant components. The
charred amorphous plant aggregates recovered at Franchthi and Shanidar were sorted using a
Leica S8 APO stereo-zoom microscope (magnication ×780). Under the stereo-zoom
microscope, the fragments under study here appeared as discrete, non-friable masses, some-
times with visible seed fragment inclusions.
As carbonised dung is frequently found in archaeobotanical samples, it was important to
exclude the possibility that the charred, amorphous plant aggregates might be faecal
remains. During microscopic examination, charred dung fragments often appear brous,
with matted/layered stems included in a dense matrix that often contains spherulites
(Smith et al.2019; Bates et al.2022). None of the Franchthi and Shanidar fragments con-
tain spherulites, or inclusions of grass stems and leaf fragments and we therefore interpret
these amorphous plant aggregates as likely food remains. In addition, they mostly contain
fragments of seeds and grain-derived plant cells, and are irregular in form and porous in
texture, with voids and cracks of variable sizes. Similar items, interpreted as carbonised
food remains and matched by experimentally reproduced examples, have previously been
Cooking in caves: Palaeolithic plant food remains from Franchthi and Shanidar
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reported in recent archaeobotanical literature (Carretero et al.2017; Valamoti et al.2021;
Bates et al.2022).
All of the charred food remains were further examined under a Meiji MT6500 darkeld/
brighteld incident light microscope (magnication ×50500) and subsequently mounted
on SEM aluminium stubs and gold sputter coated (to a thickness of 20nμ) to allow for
more detailed observation (following established analytical protocols: Carretero et al.
2017; Heiss et al.2017; Valamoti et al.2021). Identication of plant constituents, including
specic plant cell types and patterns, made use of the published literature on plant cell iden-
tication, including comprehensive guides on specic genera and families, and comparisons
to modern specimens held in the University of Liverpool Archaeobotany Laboratory plant
reference collection.
Results
Franchthi Cave
Four charred food fragments, recovered from four otation samples from Franchthi Cave,
were analysed. Three of these contain fused pulse seed, seed coat and other tissue fragments
such as macrosclereids, set in a fully or partially gelatinised matrix (Figures 13). There is evi-
dence for starch and protein cell deformation, alongside areas of vitrication most commonly
associated with the effects of soaking and heating. The smooth edges of the seed fragments
embedded in the charred matrix indicate fragmentation before carbonisation. The seed frag-
ment sizes are generally variable and might indicate coarse grinding and/or pounding (as
opposed to nely ground our-like mixtures). The formation of the gelatinised matrix
around the fragments of the seeds with adhering seed coats further suggests that preparation
of the food item might have started with soaking whole dry seeds, or with the use of fresh
seeds that had a high moisture content. The interpretation of pounding prior to charring,
as indicated by the smooth edges of the seed fragments and their variable sizes, is based
on recent experimental and archaeobotanical research on cereal food preparations, which
has established criteria for detecting this sequence of events (Valamoti et al.2021). Further
experimental work focusing on pulse processing, including grinding, mashing and soaking, is
needed to provide additional reference data for the interpretation of pulse-rich archaeological
food remains. The food fragments (Figures 13) also contain abundant remains of the
papillose seed coat pattern and macrosclereid cells characteristic of the tribe Fabeae (limited
to lentil, vetch and grass pea; Butler 1990; see also Figures S1 & S2 in the online
supplementary materials (OSM)). The food fragment in Figure 1D, for example, is suf-
ciently well preserved to permit the identication of bitter vetch (Vicia ervilia) (Butler
1990: 49395, pls 57; see also Figure S2). Our observations regarding the presence of
pulse species are further supported by previous studies, which report the abundance of
charred lentil, vetch and pea seeds in the archaeobotanical assemblage from Franchthi
Cave (Hansen 1991; see also Asouti et al.2018).
Unlike the food fragments described above, the fourth fragment of food remains is close-
textured and lacks seed fragments, observable seed coat or epidermal cells (Figure 4). Its
matrix contains voids and cracks of varying sizes, indicating a heat-affected, expanded
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starch-rich matrix. This structure strongly resembles experimental preparations and archaeo-
botanical examples of charred bread-like foods or nely ground cereal meals, such as those
reported from various Neolithic and later prehistoric settlements (Valamoti et al.2008; Car-
retero et al.2017). Probablyas a result of the advanced state of vitrication and gelatinisation,
the food fragment does not preserve identiable plant cells or other characteristic components
that would permit identication of specic plant species. The presence of starch-rich plant
food sources at Franchthi Cave, however, is well established, including grasses (oats and bar-
ley) and nuts (almonds and Pistacia) documented in the archaeobotanical assemblage (Han-
sen 1991; Asouti et al.2018).
Shanidar Cave
Five charred food fragments from the Upper Palaeolithic (Baradostian) layers at Shanidar
Cave were analysed in detail (Figures 59). All contain crushed and fused remains of
pulses, including of the genera Lathyrus and Pisum. The mounded-papillose seed coat pattern
(Figure 3CD) is frequently observed in medium-sized Lathyrus species (Butler 1990). The
height of the macrosclereids suggests that they probably belong to L. cassius,L. hirsutus or
L. nissolia (Butler 1990: 55051, pls 6263; Günes2013; see also Figure S3).
Figure 1. Pulse-rich charred plant food fragment from Franchthi Cave (context no. H1A 167, Final Palaeolithic,
Epigravettian): A) overview; B) close-up of pulse seed coat and seed fragment; C) close-up of pulse seed coat surface;
D) view of Vicia ervilia seed coat papillose cells; E) close-up of pulse seed coat surface (SEM micrographs taken by
C. Kabukcu).
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Based on the height of the macrosclereid layer and the mounded-papillose pattern, the
food fragment shown in Figure 6BC closely resembles pea (Pisum fulvum and P. sativum
subsp. elatius) (Werker et al.1979; Zablatzká et al.2021; see also Figure S4). Additionally,
Figure 3. Pulse-rich charred plant food fragment from Franchthi Cave (context no. H1A 177, Upper Palaeolithic,
Mediterranean Gravettian): A) overview; BC) close-up of pulse seed fragments and seed coat (SEM micrographs
taken by C. Kabukcu).
Figure 2. Pulse-rich charred plant food fragment from Franchthi Cave (context no. H1A 168, Final Palaeolithic,
Epigravettian): A) overview; BD) close-ups of pulse seed fragments and seed coat remains (SEM micrographs taken
by C. Kabukcu).
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Figure 5. Pulse-rich charred plant food remains from Shanidar Cave (context no. 1812, Upper Palaeolithic,
Baradostian): A) overview; BD) close-up of pulse seed coat surface and mounded-papillose seed coat pattern of
Lathyrus sp. (likely Lathyrus cassius, L. hirsutus or L. nissolia) (SEM micrographs taken by C. Kabukcu).
Figure 4. Charred plant food remains from Franchthi Cave, with a homogenised matrix (context no. H1A 172, Upper
Palaeolithic, Mediterranean Gravettian): A) overview; B) close-up showing variable sizes of voids (SEM micrographs
taken by C. Kabukcu).
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the fragments with well-preserved reticulate seed coats and globular-shaped cotyledons (e.g.
Tantawy et al.2004; Gabr 2018) are probably wild mustards (Brassicaceae). Two of the
charred food fragments also contain plant tissues resembling Pistacia nutshell and pericarp
fragments; these appear heavily deformed, possibly due to the effects of food preparation
and/or post-depositional taphonomic processes on the morphology of the plant tissues
(see Figures 8B,8D and 9CD). SEM images of carbonised modern reference Pistacia speci-
mens are included in the OSM (Figure S5).
The single fragment of charred food remain from a Mousterian layer at Shanidar Cave
(Figure 10) contains pulse seed and seed coat fragments. Unlike the Baradostian food frag-
ments, however, it also includes the long cells characteristic of grasses (Poaceae) (Figure 10D).
Discussion
The food remains from Upper/Final Palaeolithic strata at Franchthi Cave and Middle/Upper
Palaeolithic strata at Shanidar Cave reported here currently represent the earliest direct macro-
botanical evidence of Palaeolithic plant food processing in the Eastern Mediterranean and
South-west Asia. They represent a signicant addition to an accumulating body of archaeo-
botanical data from these regions that points to selective plant foraging by Palaeolithic
Figure 7. Charred plant food remains from Shanidar Cave (context no. 1823, Upper Palaeolithic, Baradostian)
containing wild mustard: A) overview; BC) close-up of mustard seed fragment and seed coat pattern (SEM
micrographs taken by C. Kabukcu).
Figure 6. Charred plant food fragment from Shanidar Cave (context no. 1866, Upper Palaeolithic, Initial
Baradostian): A) overview; BC) close-up of wild pea (Pisum fulvum or P. sativum subsp. elatius) seed coat (SEM
micrographs taken by C. Kabukcu).
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hunter-gatherers. At Middle Palaeolithic Kebara Cave (Mount Carmel, Israel), for example,
pulse seeds (vetches, grass pea and lentils) constitute the majorityof the charred plant macro-
fossils (Lev et al.2005). The Epipalaeolithic occupation at El Wad (also in Mount Carmel)
similarly contains an archaeobotanical assemblage dominated by pulses, with a signicant
proportion of vetches (Caracuta et al.2016). Epipalaeolithic Palegawra Cave (Iraqi Kurdi-
stan) provides evidence for a more diverse range of foraged plants, including wild pulses,
grasses, nuts, tubers and mustards (Asouti et al.2020). In the Levantine Epipalaeolithic,
there is also increasing evidence for the use of tubers (e.g. Shubayqa I, Jordan;
Arranz-Otaegui et al.2018) and mustards (e.g. Kharaneh IV, Jordan; Bode et al.2022).
The remarkably well-preserved assemblage from Ohalo II (Israel) also provides evidence
for the use of wild grasses during this period (Weiss et al.2004).
Other notable Epipalaeolithic sites in South-west Asia from which archaeobotanical data
are available, including the Karain and Öküzini Caves (Martinoli 2004) and the Pınarbası
Figure 8. Charred plant food remains from Shanidar Cave (context no. 1866, Upper Palaeolithic, Initial Baradostian):
A) overview; B) & D) close-up of cf. Pistacia nutshell/pericarp remain; C) close-up of pulse seed coat pattern (Fabeae)
(SEM micrographs taken by C. Kabukcu).
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rockshelter (Baird et al.2013) in Anatolia, demonstrate an emphasis on nuts (almonds and
terebinth), pulses and various wild fruits. The broadly contemporaneous occupation at Haua
Fteah in north-east Libya similarly provides evidence for the use of pine nuts and wild vetches
(Barker et al.2010). Several European Upper Palaeolithic sites attest to the use of wild grasses
and tubers. At Klisoura 1 Cave in Peloponnese (Greece), phytolith and micromorphological
data from Upper Palaeolithic clay-lined hearths indicate grass seed roasting (Karkanas et al.
2004). Starch and ground stone use-wear data from the Gravettian occupation at Grotta
Paglicci in southern Italy suggest the cooking and processing (grinding/crushing) of wild
oats and the processing of tannin-rich oak acorns (Lippi et al.2015). Similarly, starch and
use-wear evidence on grinding stones from the Late Stone Age occupation at Haua Fteah,
dated to c. 31 000 years ago, point to the regular processing of goat grass (Aegilops sp.) (Barton
et al.2018).
Most of the carbonised food remains reported here contain variously sized fragments of
pulse seeds, probably representing processing by coarse grinding, cracking and/or pounding.
This type of preparation differs from the ner grinding required for our and does not neces-
sitate the use of at grinding stones. It could have been undertaken using only percussive tools
Figure 9. Charred plant food fragment from Shanidar Cave (context no. 636, Upper Palaeolithic, Baradostian): A)
overview; BD) close-up of cf. Pistacia nutshell/pericarp fragment (SEM micrographs taken by C. Kabukcu).
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and/or perishable implements. Use-wear data from the Acheulian site of Gesher Benot Yaa-
qov (Israel) point to the early use of stone tools for nut cracking and plant food preparation
(Goren-Inbar et al.2015). In the Upper Palaeolithic starch-rich charred food fragment from
Franchthi Cave (Figure 4), the absence of large particles and the high degree of homogenisa-
tion of the matrix suggest that the plants were processed via ne grinding and/or boiling and
mashing. Similar charred plant aggregates comprising ne-ground starch plant tissue and
occasionally no identiable plant cells have also been reported from Epipalaeolithic, Meso-
lithic and Neolithic contexts in South-west Asia and Europe, and have been interpreted as
breadsor porridges(Kubiak-Martens et al.2015; Carretero et al.2017; Arranz-Otaegui
et al. 2018).
Pulse seeds, especially bitter vetch (Vicia ervilia) and grass pea (Lathyrus cassius,L. hirsutus
and L. nissolia), contain notable quantities of alkaloids and tannins, resulting in a bitter and
astringent taste. These compounds are concentrated in the seed coats. While cooking tech-
niques, such as soaking and boiling, can remove a large portion of tannins and other bitter,
astringent and toxic compounds (Ressler et al.1997), hulling (the removal of the seed coat)
Figure 10. Charred plant food remains from Shanidar Cave containing pulses and grasses (context no. 1924, Middle
Palaeolithic, Mousterian): A) overview; BC) close-up of pulse seed coat and seed fragments; D) close-up of long cells of
Poaceae seed (SEM micrographs taken by C. Kabukcu).
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would have been a far more efcient method. This technique is commonly practised today
(Ressler et al.1997), as well as in the past (Melamed et al.2008; Valamoti et al.2011),
for processing vetches and grass pea. Thesoaking of wild pulses, as indicated by the Franchthi
and Shanidar charred food fragments, would have enabled their safe consumption and
improved their palatability by removing most of the bitter-tasting compounds. The presence
of seed coat fragments, however, suggests that a low level of plant chemicals, including some
tannins and alkaloids, may have been intentionally retained in plant food preparations. This
evidence adds to an increasing body of archaeobotanical studies suggesting a persistent reli-
ance on, and tolerance of, bitter- and astringent-tasting plant foods such as pulses, mustards,
almonds and terebinths, from as early as the Middle Palaeolithic through to the later prehis-
toric periods. Beyond the Eastern Mediterranean and South-west Asia, archaeobotanical
studies at sites such as Niah Cave (Sarawak, Borneo) have revealed evidence for the processing
of the highly toxic Dioscorea (yam) and Pangium edule nuts from as early as 50 000 years ago,
underscoring the complexity and deep ancestry of such food preparation practices (Barker
et al.2007; Barton et al.2016).
Apart from detoxication, food preparation practices such as soaking and pounding would
also have improved the bioavailability of bulk nutrients. Multi-proxy data from Early Upper
Palaeolithic sites in the Pontic steppe, derived from starch residue extraction, spectroscopic
and spectrometric techniques, point to the processing by pounding of a diverse group of
tubers, possibly with the aim of tenderising them for consumption (including some less com-
monly observed C
4
carbon-xing plant species; see Longo et al.2021). Other aspects of plant
resource choice and use, including raw materials and medicinal uses, have also been high-
lighted with regard to Lower and Middle Palaeolithic foraging (Hardy et al.2012,2022;
Hardy 2018).
Conclusion
The evidence presented here supports previous hypotheses regarding the diversity and com-
plexity of Palaeolithic plant use. It provides direct evidence for previously undocumented
food preparation practices and brings into focus the diversity of specialised cooking practices
developed by Middle and Upper Palaeolithic hunter-gatherers, which involved multiple
preparation steps and different plant components (sensu Jones 2009). Our results reinforce
current understanding that the use of plants in the Palaeolithic regularly relied on starch-rich
tubers and grasses (Henry et al.2011; Hardy et al.2022) and further demonstrate that the
labour-intensive processing of a broad spectrum of plant foods, including bitter, astringent
and potentially toxic plants for human consumption, was an integral part of hunter-gatherer
resource management strategies. The use of plant food preparation techniques was prevalent
across the Eastern Mediterranean and South-west Asia from as early as the Middle Palaeolithic
and appears to be independent of uctuations in forage and prey ceilings due to climatic con-
ditions (Hardy 2018; Power & Williams 2018). Crucially, our results demonstrate that food
choices and preparation practices traditionally associated with the intensication of plant
resource use that is linked to climatic amelioration at the PleistoceneHolocene boundary
and the origin of farming (Smith & Zeder 2013) clearly have a deep history that precedes
the earliest evidence for plant cultivation by several tens of thousands of years.
Ceren Kabukcu et al.
© The Author(s), 2022. Published by Cambridge University Press on behalf of Antiquity Publications Ltd
24
https://doi.org/10.15184/aqy.2022.143 Published online by Cambridge University Press
Acknowledgements
C.K. designed the study, carried out the analysis and produced the original draft. All authors
contributed to writing, review and editing of the nal draft. E.A. provided access to the
Franchthi Cave samples; E.H., C.H., E.P., T.R. and G.B. designed and carried out sampling
for archaeobotanical remains at Shanidar Cave and provided access to the samples. We thank
the Kurdistan Regional Government for inviting G.B. to plan and direct new excavations at
Shanidar Cave, and the Kurdistan General Directorate of Antiquities for granting excavation
permits and permission to analyse the nds. Additional modern plant reference materials
were made available to C.K. by the USDA Agricultural Research Service, Plant Germplasm
Introduction and Testing Unit.
Supplementary materials
To view supplementary material for this article, please visit https://doi.org/10.15184/aqy.
2022.143.
Funding statement
C.K. acknowledges funding from the Leverhulme Trust (Early Career Fellowship, ECF
284); G.B., E.P., C.H. and T.R. acknowledge funding from the Leverhulme Trust (Research
Grant RPG2013105), Rust Family Foundation, British Academy, Wenner-Gren Founda-
tion, Society of Antiquaries, McDonald Institute of Archaeological Research at the University
of Cambridge and Natural Environment Research Councils Oxford Radiocarbon Dating
Facility (grant NF/2016/2/14).
Data statement
Archaeological and modern plant reference specimens are archived at the University of
Liverpool Archaeobotany Laboratory, Department of Archaeology, Classics and Egyptology.
All data are available in the main text and the OSM.
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