Wild cereal grain consumption among Early Holocene foragers of the Balkans predates the arrival of agriculture
Figures
![](https://iiif.elifesciences.org/lax/72976%2Felife-72976-fig1-v2.tif/full/617,/0/default.jpg)
Sites in the central Balkans investigated in the article, which provided dental calculus and ground stone tools.
EM = Early Mesolithic; LM = Late Mesolithic; M/N = Mesolithic-Neolithic; EN = Early Neolithic; BA = Bronze Age; MA = Medieval.
![](https://iiif.elifesciences.org/lax/72976%2Felife-72976-fig3-v2.tif/full/617,/0/default.jpg)
Late Mesolithic ground stone tools from the site of Vlasac featuring use-wear traces and residues related to plant food processing.
![](https://iiif.elifesciences.org/lax/72976%2Felife-72976-fig4-v2.tif/full/617,/0/default.jpg)
Starch granules from Mesolithic and Neolithic dental calculus.
Early Mesolithic: (a) Type Ib (PAD11); (b) Type Ib (PAD9); (c) Type Ib (PAD11); (d) Type V (PAD12); (e) Type III (PAD11); (f) Type Ib (PAD15); Late Mesolithic: (g) Type II (VL82c); (h) Type IV (VL31); (i) Type VI (VL70); (j, k) multicellular structures of long cells embedded in dental calculus (HV25/26, VL70); (l) Type Ia (HV11); Mesolithic–Neolithic: (m) Type Ia (HV16). Neolithic: (n) Type III (LV32a); (o–v) damaged Type I granules (A-Type granules) (VEL-2D); (w) Type I (A-Type granule) (VEL-2D); (x) single dendritic cell (Gârleşti); (y) Type I (A-Type granule) (VEL-2D; VEL-2A).
![](https://iiif.elifesciences.org/lax/72976%2Felife-72976-fig5-v2.tif/full/617,/0/default.jpg)
Other dietary and nondietary debris found in Mesolithic dental calculus from the Danube Gorges.
Early Mesolithic: (a) Type V (PAD11); (b, c) Type III (PAD12); (d) Type I (A-Type granule) (VEL-2D; PAD12); (e) Type I (A-Type granule) (PAD12); (f) smoke particle (LV50); (g, h) plant fiber embedded in calculus (PAD16); (i) Type Ib (PAD9); (j) feather barbule embedded in calculus (PAD9); Late Mesolithic: (k) Type II (PAD2); (l) Polylobate phytolith (US64 x.11); (m) Phytoliths (VL79); (n–p) Type VI (VL70,VL83); (q) feather barbules embedded in calculus (HV25/26); (r) feather barbule embedded in calculus (VL80a); (s) Echinate pollen grain in calculus (VL83); (t) plant tissue (LV79a); (u) Type II (VL43); (v) Type Ia (HV11); Mesolithic-Neolithic: (w) Type I (HV16); (x) wood particle (PAD4); (y) phytoliths (LV28); (z) feather barbule (PAD4).
![](https://iiif.elifesciences.org/lax/72976%2Felife-72976-fig6-v2.tif/full/617,/0/default.jpg)
Starch granule morphological variability within the species of the genus Aegilops and domestic species of the Triticeae tribe.
(a) Aegilops cylindrica; (b) A. neglecta; (c) A. speltoides tauschii; (d) A. caudata; (e) A. triuncialis; (f) A. comosa; (g) A. uniaristata; (h) A. ventricosa; (i) A. geniculata; (j) A. crassa; (k) A. peregrina; (l) Elymus caninus; (m) Bromus tectorum; (n) Agropyron pungens; (o) A. farctus; (p) Dasypyron villosum; (q) Triticum monococcum; (r) Hordeum vulgare; (s) T. dicoccum; (t) T. aestivum.
![](https://iiif.elifesciences.org/lax/72976%2Felife-72976-fig7-v2.tif/full/617,/0/default.jpg)
Experimental reference for starch granules identified in the dental calculus and ground stone tools.
(a) Aegilops triuncialis; (b) A. crassa; (c, d) Avena strigosa; (e, f) Setaria italica; (g) Vicia cracca; (h) V. sylvatica; (i) Quercus pubescens; (j) Q. robur; (k) Q. colurna; (l) Cornus mas.
![](https://iiif.elifesciences.org/lax/72976%2Felife-72976-fig8-v2.tif/full/617,/0/default.jpg)
Experimental macroresidues and micropolish associated with grass grains processing compared to macroresidue and micropolish identified on archaeological ground stone tools from the site of Vlasac.
(a–e) Yellowish organic film covering the crystal grains on experimental GSTs used to process oat (a), downy brome (b), wild grass grains (c), and millet (d); smooth domed and flat micropolish developing over the high and low microtopographies associated with oat (Avena barbata) grinding; (f) smooth flat and domed micropolish developing over the surface high and low microtopographies and characterized by narrow microstriations associated with grinding downy brome (Bromus tectorum L.); (g) smooth flat micropolish developed over the high and low microtopographies characterized by sporadic narrow striations associated with grinding wild grass grains (Aegilops ventricosa Tausch); (h) smooth domed polish developed over the high and low microtopographies associated with the grinding of foxtail millet (Setaria italica (L.) P. Beauvois); (i–l) spots of organic film, yellowish in color covering the crystal grains across the surface of archaeological GSTs; smooth domed micropolish identified on archaeological GSTs developing over the high and low surface microtopographies and associated with microstriations (m-o). Starch granules identified on archaeological GSTs. (q) Type I (GST no. INV.80); (r) Type I (GST no. INV.146); (s) Type III (GST no. INV.28); (t) Type VI (GST no. INV.67); (u) Type VI (GST no. INV.10); (v) Type VI (GST no. INV.146); (w) Type I (GST no. INV.71).
![](https://iiif.elifesciences.org/lax/72976%2Felife-72976-fig9-v2.tif/full/617,/0/default.jpg)
Starch granule length in modern wild and domestic cereal species.
(a) Distribution of starch granule length in wild species; (b) distribution of domesticated species; (c) violin plot of comparing the length of starch granules in wild and domesticated species; (d) interquartile ranges (IQRs) of wild and domestic species. IQR corresponds to the difference in the medians of the lower and upper half of the data.
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Figure 9—source data 1
Starch granule length in modern wild and domestic cereal species.
- https://cdn.elifesciences.org/articles/72976/elife-72976-fig9-data1-v2.xlsx
![](https://iiif.elifesciences.org/lax/72976%2Felife-72976-fig10-v2.tif/full/617,/0/default.jpg)
Controls for contamination.
(a–w) Evidence of pollutants retrieved from clean working surfaces and dust traps located in different areas of the DANTE Laboratory at Sapienza University of Rome; (x–Ll) dust recorded in storage boxes where groundstone tools were stored at the Faculty of Philosophy, University of Belgrade.
Tables
Details of dental calculus sampled for the study (n = 60).
*No stable isotope values are currently available for this individual in order to correct the obtained radiocarbon date for the reservoir effect, and the calibrated range should probably be considered too old for its actual age, likely being 200–500 years younger. All calibrated ranges have end points rounded outwards to 5 years. The dates were individually calibrated using OxCal 4.4 and IntCal 20 (Reimer et al., 2020).
Site | Burial no. | Period attribution | AMS dates | Calculus location | |||||
---|---|---|---|---|---|---|---|---|---|
Lab code and source | 14C age (BP) | Reservoir effect corrected age (BP) | 95.4 % confidence, cal BC | Tooth | Surface | Weight (mg) | |||
Padina | PAD20 | Early Meso | 17 | Buccal | 9.6 | ||||
Padina | PAD25 | Early Meso | 38 | Buccal | 9.59 | ||||
Padina | PAD15 | Early Meso | OxA-17145 (Borić, 2011) | 9310 ± 44 | 8870 ± 63 | 8240–7770 | 38 | Lingual | 9.58 |
Padina | PAD16a | Early Meso | PSU-2407 (Mathieson et al., 2018) | 9340 ± 35 | 8907 ± 66 | 8275–7815 | 34 | Buccal | 9.62 |
Padina | PAD18b | Early Meso | PSU-2376 (Mathieson et al., 2018) | 9715 ± 40 | 9424 ± 55 | 9115–8550 | 48 | Lingual | 9.67 |
Padina | PAD9 | Early Meso | AA-57771 (Borić, 2011) | 9920 ± 100 | 9480 ± 110 | 9225–8495 | 42, 46 | Lingual | 9.59 |
Padina | PAD11 | Early Meso | OxA-16938 (Borić, 2011) | 9665 ± 54 | 9225 ± 70 | 8620–8290 | 27 | Lingual | 9.57 |
Padina | PAD12 | Early Meso | BM-1146 (Borić, 2011) | 9331 ± 58 | – | 8750–8350 | 27 | Lingual | 9.76 |
Padina | PAD17 | Early Meso | PSU-2375 (Mathieson et al., 2018) | 9505 ± 35 | 9105 ± 62 | 8540–8230 | 25 | Buccal | 9.64 |
Lepenski Vir | LV50 | Early Meso | BA-10651 (Borić et al., 2018) | 9455 ± 38 | 9082 ± 62 | 8540–8020 | 35 | Buccal | 9.99 |
Lepenski Vir | LV20 | Early Meso | OxA-39629(this paper) | 10,268 ± 38 | 9928 ± 58 | 9740–9270 | 48 | Lingual | 9.73 |
Padina | PAD26 | Early Meso | 14 | Buccal | 9.58 | ||||
Padina | PAD6 | Early Meso | 47 | Lingual | 9.65 | ||||
Padina | PAD2 | Late Meso | BM-1143 (Borić, 2011) | 7738 ± 51 | – | 6650–6465 | 36 | Lingual | 9.68 |
Hajdučka Vodenica | HV25/26 | Late Meso | 44 | Buccal | 9.60 | ||||
Hajdučka Vodenica | HV29 | Late Meso | AA-57774 (Borić, 2011) | 8151 ± 60 | 7711 ± 75 | 6690–6425 | 48 | Lingual | 10.72 |
Hajdučka Vodenica | HV8 | Late Meso | OxA-13613 (Borić, 2011) | 8456 ± 37 | 8016 ± 58 | 7075–6695 | 48 | Buccal | 9.61 |
Hajdučka Vodenica | HV11 | Late Meso | 48 | Buccal | 9.71 | ||||
Hajdučka Vodenica | HV profil A | Late Meso | 27 | Buccal | 9.70 | ||||
Hajdučka Vodenica | HV30 | Late Meso | 27 | Buccal | 9.56 | ||||
Vlasac | VL82c | Late Meso | BRAMS-2588 (Jovanović et al., 2021a) | 8035 ± 28 | 7595 ± 53 | 6590–6270 | 42 | Buccal | 9.68 |
Vlasac | VL2 | Late Meso | 14 | Buccal | 9.54 | ||||
Vlasac | VL80a | Late Meso | 26 | Lingual | 9.84 | ||||
Vlasac | VL55 | Late Meso | BRAMS-2583 (Jovanović et al., 2021b) | 8377 ± 29 | 7837 ± 63 | 7035–6500 | 33 | Lingual | 9.64 |
Vlasac | VL74 | Late Meso | BRAMS-2587 (Jovanović et al., 2021b) | 8149 ± 28 | * | 7315–7055* | 28 | Lingual | 9.70 |
Vlasac | VL83 | Late Meso | OxA-5826 (Borić, 2011) | 8200 ± 90 | 7760 ± 100 | 7030–6420 | 24 | Lingual | 9.62 |
Vlasac | VL43 | Late Meso | 27 | Lingual | |||||
Vlasac | VL31 | Late Meso | AA-57777 (Borić, 2011) | 8196 ± 69 | 7756 ± 82 | 6900–6430 | 26 | Buccal | 9.58 |
Vlasac | VL45 | Late Meso | AA-57778 (Borić et al., 2004) | 8117 ± 62 | 7677 ± 77 | 6655–6400 | 38 | Buccal | 9.50 |
Vlasac | VL70 | Late Meso | 17 | Buccal | 10.42 | ||||
Vlasac | VL79 | Late Meso | BRAMS-2448 (Jovanović et al., 2021b) | 8005 ± 29 | 7565 ± 54 | 6565–6250 | 16 | Buccal | 9.60 |
Vlasac | U44 | Late Meso | 27 | Buccal | 9.96 | ||||
Vlasac | H232 | Late Meso | OxA-20702 (Borić, 2011) | 7725 ± 40 | 6640–6470 | 28 | Lingual | 9.92 | |
Vlasac | H317 | Late Meso | PSU-2381 (Mathieson et al., 2018) | 8110 ± 35 | 7625 ± 71 | 6645–6270 | 26, 36 | Lingual | 9.73 |
Vlasac | U115 | Late Meso | 28 | Buccal | 9.95 | ||||
Vlasac | U326 | Late Meso | PSU-2382 (Mathieson et al., 2018) | 8045 ± 30 | 7728 ± 51 | 6645–6465 | 17 | Buccal | 9.94 |
Vlasac | U326 | Late Meso | PSU-2382 (Mathieson et al., 2018) | 8045 ± 30 | 7728 ± 51 | 6650–6460 | 1, 2 | Buccal | 9.1 |
Vlasac | U64 x.11/H81 | Late Meso | OxA-20762 (Borić, 2011) | 8125 ± 45 | 7685 ± 64 | 6645–6430 | 20, 26, 27, 29, 30, 31 | Lingual | 9.92 |
Vlasac | H341 | Late Meso | 1 | Buccal | 10.12 | ||||
Vlasac | VL48 | Late Meso | 34 | Lingual | 10.06 | ||||
Vlasac | U222 x.18 | Late Meso | 2 | Buccal | 9.54 | ||||
Lepenski Vir | LV28 | Meso-Neo | – | 43 | Buccal | 9.58 | |||
Lepenski Vir | LV79a | Meso-Neo | OxA-25091 (Bonsall, 2008) | 7605 ± 38 | 7119 ± 74 | 6220–5805 | 33 | Buccal | 9.69 |
Hajdučka Vodenica | HV16 | Meso-Neo | 36 | Lingual | 9.54 | ||||
Hajdučka Vodenica | HV19 | Meso-Neo | 37 | Buccal | 9.58 | ||||
Hajdučka Vodenica | HV13 | Meso-Neo | AA-57773 (Borić, 2011) | 7435 ± 70 | 6995 ± 83 | 6020–5720 | 17 | Lingual | 9.56 |
Padina | PAD4 | Meso-Neo | AA-57769 (Ottoni et al., 2021) | 7518 ± 72 | 7078 ± 85 | 6080–5745 | 48 | Buccal | 9.73 |
Padina | PAD5 | Meso-Neo | AA-57770 (Borić, 2011) | 7598 ± 72 | 7158 ± 85 | 6230–5845 | 15 | Buccal | 8.10 |
Vlasac | U24 x.30 | Meso-Neo | 32 | Lingual | 9.97 | ||||
Vlasac | H53 | Meso-Neo | OxA-16544 (Borić et al., 2014) | 7035 ± 40 | – | 6015–5805 | 3, 28, 29 | Lingual | 10.04 |
Lepenski Vir | LV4 | Early Neo | 33 | Buccal | 9.64 | ||||
Lepenski Vir | LV73 | Early Neo | BA-10652 (Borić et al., 2018) | 7265 ± 30 | 6973 ± 48 | 5980–5735 | 34 | Buccal | 9.69 |
Lepenski Vir | LV8 | Early Neo | AA-58319OxA-25207 (Borić et al., 2018) | 6825 ± 517097 ± 36 | 6690 ± 546984 ± 39 | 5715–55205985–5750 | 44 | Lingual | 9.69 |
Lepenski Vir | LV32A | Early Neo | OxA-5828 (Bonsall et al., 2013) | 7270 ± 90 | 7032 ± 95 | 6065–5730 | 42, 43, 36 | Buccal | 9.77 |
Lepenski Vir | LV17 | Early Neo | AA-58320 (Borić et al., 2018) | 7007 ± 48 | 6787 ± 53 | 5775–5565 | 15 | Lingual | 9.10 |
Padina | PAD30 | Bronze Age | PSU-2379 | 2140–1765 | 47 | Buccal | 9.69 | ||
Velesnica | 2A | Early Neo | OxA-19191 (Bonsall, 2008) | 7409 ± 38 | 7196 ± 47 | 6220–5930 | 8 | Lingual | 9.74 |
Velesnica | 2D | Early Neo | OxA-19210 (Bonsall, 2008) | 7327 ± 38 | 7183 ± 42 | 6215–5925 | 9 | Lingual | 7.2 |
Gârleşti | Early Neo | 2 | Lingual | 8.22 | |||||
Lepenski Vir | LV30 | Medieval | OxA-25218 (Bonsall, 2008) | 427 ± 23 | AD1440–1490 | 16 | Lingual | 9.67 |
Details of the microdebris (starch granules and other microremains) found in the archaeological dental calculus samples (PO = pollen; W = wood; Ch = microcharcoal/burnt debris; Gr = grit; P = phytoliths; FE = feathers; FI = fibers; FU = fungi; S = smoke) (n = 51).
Site | Burial label | Chronocultural attribution | Type I Triticeae | Type IIAveneae | Type III Paniceae | Type IVFabeae | Type V Fagaceae | Type VI Cornaceae | Indet. | Other | |
---|---|---|---|---|---|---|---|---|---|---|---|
1 | Padina | PAD20 | Early Meso | 7 | 4 | 3 | 1P/10FI/2FE/2W/1Ch/Gr | ||||
2 | Padina | PAD25 | Early Meso | 1 | 1P/1Ch/Gr | ||||||
3 | Padina | PAD15 | Early Meso | >100 | 4 | 6 | 1P/4PO/1W/1Ch/Gr | ||||
4 | Padina | PAD16a | Early Meso | 20 | 3 | 1P/10FI/8Ch/Gr | |||||
5 | Padina | PAD9 | Early Meso | >200 | 2FI/2FE | ||||||
6 | Padina | PAD11 | Early Meso | >100 | 8 | 1 | |||||
7 | Padina | PAD12 | Early Meso | 36 | 12 | 1 | 1 | ||||
8 | Lepenski Vir | LV50 | Early Meso | 1 | 1 | 1FI;S | |||||
9 | Lepenski Vir | LV20 | Early Meso | 4 | 1 | ||||||
10 | Padina | PAD2 | Late Meso | 13 | 1 | 7 | 1 | 1PO/1FI/2FE/2W/1Ch/Gr | |||
11 | Hajdučka Vodenica | HV25/26 | Late Meso | 5 | 15 | 1 | 2P/1PO/2FI/3FE/1Ch/3FU/Gr | ||||
12 | Hajdučka Vodenica | HV29 | Late Meso | 5 | 3 | 5 | 3P/1PO/1FI/1FE/13W/1FU/Gr | ||||
13 | Hajdučka Vodenica | HV8 | Late Meso | 1 | |||||||
14 | Hajdučka Vodenica | HV11 | Late Meso | >100 | 8 | 1 | |||||
15 | Hajdučka Vodenica | HV profil A | Late Meso | 14 | 1 | 3PO/1FI/2FE/3W/1Ch/1FU/Gr | |||||
16 | Hajdučka Vodenica | HV30 | Late Meso | 1 | |||||||
17 | Vlasac | U222 x.18 | Late Meso | 2 | 1P | ||||||
18 | Vlasac | U326 | Late Meso | >60 | 1 | 1P | |||||
19 | Vlasac | VL82c | Late Meso | 4 | 23 | 7 | 2 | 5 | 1P/3PO/1FE/1W/1Ch/Gr | ||
20 | Vlasac | VL2 | Late Meso | 2 | 12 | 2P/1PO/2FI/1FE/2FU/Gr | |||||
21 | Vlasac | VL80a | Late Meso | 3 | 15 | 5 | 5 | 1 | 1P/4FE/1W/1Ch/1FU/Gr | ||
22 | Vlasac | VL55 | Late Meso | 6 | 1 | 1P/1PO/1FI/1W/1FU/Gr | |||||
23 | Vlasac | VL74 | Late Meso | 1 | 1 | 1P/3PO/17FI/1Ch/Gr | |||||
24 | Vlasac | VL83 | Late Meso | 6 | 8 | 1 | 1P/1PO/1FE/2W/5Ch/Gr | ||||
25 | Vlasac | VL43 | Late Meso | >200 | 12 | 1 | 1 | 2 | 2FE/1W/2Ch/1FU/Gr | ||
26 | Vlasac | VL31 | Late Meso | 18 | 3 | 3 | 4PO/5FE/1Ch/2FU/Gr | ||||
27 | Vlasac | VL45 | Late Meso | 23 | 8 | 20 | 1 | 14 | 1PO/2FE/2Ch | ||
28 | Vlasac | VL70 | Late Meso | 3 | 3 | 1 | 14 | 4P/1FI/7Ch | |||
29 | Vlasac | VL79 | Late Meso | 1P/2FI | |||||||
30 | Vlasac | U44 | Late Meso | 3 | 2 | ||||||
31 | Vlasac | H232 | Late Meso | <100 | 4 | 1 | 1PO/1FE | ||||
32 | Vlasac | U115 | Late Meso | 1 | |||||||
33 | Vlasac | U64 x.11 | Late Meso | >200 | 10 | 32 | 4 | 2P/4FI/2FE/3Ch/1FU | |||
34 | Vlasac | H341 | Late Meso | 1 | |||||||
35 | Lepenski Vir | LV28 | Meso-Neo | 4 | 3 | 1 | 6 | 2P/2PO/1FE/2W/4Ch/Gr | |||
36 | Hajdučka Vodenica | HV16 | Meso-Neo | >200 | 1FU | ||||||
37 | Hajdučka Vodenica | HV19 | Meso-Neo | 1 | 1FE | ||||||
38 | Hajdučka Vodenica | HV13 | Meso-Neo | 1 | |||||||
39 | Padina | PAD4 | Meso-Neo | 1 | 7 | 2PO/1FI/3FE/4W/2Ch/1FU/Gr | |||||
40 | Padina | PAD5 | Meso-Neo | 6 | |||||||
41 | Vlasac | U24 x.30 | Meso-Neo | 10 | 1P/2Ch | ||||||
42 | Vlasac | H53 | Meso-Neo | 22 | >200 | 5 | 1FE/1W | ||||
43 | Lepenski Vir | LV4 | Early Neo | 1 | 3 | 1 | |||||
44 | Lepenski Vir | LV73 | Early Neo | 12 | 9 | 1 | 7P/2PO/17FI/1FE/3FU/Gr | ||||
45 | Lepenski Vir | LV8 | Early Neo | 11 | 4 | 1W | |||||
46 | Lepenski Vir | LV32A | Early Neo | 8 | >200 | 1P/2FE | |||||
47 | Lepenski Vir | LV17 | Early Neo | 14 | 5 | ||||||
48 | Velesnica | 2A | Early Neo | 4FU | |||||||
49 | Velesnica | 2D | Early Neo | 12 | 2P/2FU | ||||||
50 | Gârleşti | Early Neo | 1 | 1P/1Ch | |||||||
51 | Lepenski Vir | LV30 | Medieval | 12 | 4 | 1P/1PO/4Ch/1FU/Gr | |||||
Total | >1446 | 324 | >409 | 43 | 8 | 24 | 284 |
Late Mesolithic ground stone tools from the site of Vlasac.
Inv. no. | Archaeological context | Shape | Tool type | Length (cm) | Width (cm) | Thickness (cm) | Weight (g) | Volume (cm3) | State of preservation | PDM | Micropolish description | Micropolish location | Microstriation description | Microstriation orientation | Cristal grain modification | Gesture |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
10 | a1-III | Subangular | Handstone/grinder | 12.7 | 10.5 | 7.88 | 1542 | 645 | Preserved | Light soil concretion | Smooth and domed | High microtopographies | Short narrow with a matt bottom | Unidirectional | Y | Mixed |
13 | a1-VIII | Round | Handstone/grinder | 11 | 8.16 | 5.68 | 680 | 287 | Preserved | None | Smooth and domed with sporadic pits | High and low microtopographies | NA | NA | N | Longitudinal |
23 | BV/C/IV-X | Subangular | Indeterminable | 11.7 | 8.79 | 8.76 | 823 | 367 | Preserved | Light soil concretion | Rough to smooth with domed and flat spots | High and low microtopographies | NA | NA | N | Longitudinal |
28 | BIII-C/V | Oval | Passive base | 13.3 | 11.8 | 7.18 | 1283 | 499 | Fractured | None | Smooth | High microtopographies | NA | NA | Y | Longitudinal |
56 | A/II–XIII | Round | Passive base | 15.9 | 14.8 | 7.7 | 1038 | 368 | Preserved | Heavy surface concretion on one surface | Smooth domed and flat | High microtopographies | NA | NA | Y | Mixed |
63 | b/17-XV | Round | Handstone/grinder | 8.4 | 6.6 | 4.47 | 403 | 141 | Preserved | Light surface abrasion | Rough to smooth with reticulated and flat spots | High and low microtopographies | NA | NA | N | Longitudinal |
65 | C/I–VI | Round | Passsive base | 100 | 84.3 | 55.5 | 680 | 298 | Broken | Fractures | Smooth domed and reticulated | High microtopographies | Narrow with a matt bottom | Mixed | N | Longitudinal |
66 | C/I II/V | Round | Indeterminable | 9.5 | 8.5 | 7.6 | 1170 | 437 | Preserved | None | Smooth domed and cratered | High and low topographies | NA | NA | Y | Mixed |
67 | C/I-C/II–III | Subangular | Passive base | 10.8 | 8.4 | 5.91 | 633 | 253 | Broken | Light soil concretion and surface abrasion | Smooth and reticulated | High microtopographies | Short and deep with a matt bottom | Unidirectional | N | Longitudinal |
71 | C/I–V | Round | Handstone/grinder | 72.1 | 57.9 | 45.9 | 309 | 119 | Preserved | None | Smooth domed to flat | High microtopographies | Long and shallow with a polished bottom | Mixed | Y | Longitudinal |
75 | b/18V | Subangular | Handstone/grander | 5.53 | 5.29 | 3.45 | 137 | 57 | Broken | Fractures | Smooth and domed | High and low microtopographies | Short narrow with a polished bottom | Unidirectional | N | Longitudinal |
80 | C/II-II/6 | Ovate | Passive base | 9.85 | 8.35 | 3.72 | 547 | 225 | Broken | None | Smooth domed | High microtopographies | Short and narrow with a matt bottom | Unidirectional | Y | Longitudinal |
134 | b/V3-XII | Subangular | Indeterminable | 12.2 | 9.57 | 8.13 | 1433 | 565 | Preserved | None | Smooth domed | High and low microtopographies | Short deep with a matt bottom | Unidirectional | Y | Longitudinal |
141 | B/I 0–8.9 | Round | Handstone/grinder | 10.8 | 9.92 | 9.22 | 370 | NA | Preserved | Soil concretion | Smooth domed and flat | High microtopographies | NA | NA | Y | Longitudinal |
146 | B/I-below hearth 9 | Round | Handstone/grinder | 6.65 | 5.7 | 4.66 | 275 | 106 | Preserved | Light soil concretion | Smooth domed and reticulated | High microtopographies | Short narrow with a matt bottom | Unidirectional | N | Longitudinal |
162 | A/16X | Round | Handstione/grinder | 10.6 | 9.3 | 7.52 | 1143 | 424 | Preserved | None | Smooth domed | High and low microtopographies | Shirt narrow with a matt bottom | Unidirectional | Y | Longitudinal |
167 | a/15-VII | Round | Indeterminable | 8.94 | 8.82 | 5.65 | 611 | 241 | Broken | Light surface abrasion | Rough granular and domed | High and low topographies | NA | NA | Y | Orthogonal |
Summary statistics of the length (μm) of wild grass grains and domestic cereal starch granules.
Species | Min. | Max. | Mean | Median | St. Dev. | Range | IQR |
---|---|---|---|---|---|---|---|
A. caudata | 5.29 | 59.3 | 21.6 | 16.7 | 15.17 | 5.29–59.33 | 26.55 |
A. comosa | 7.95 | 34.5 | 21.5 | 21.7 | 9.78 | 7.95–34.54 | 20.09 |
A. crassa | 13.38 | 53.7 | 35.3 | 33.7 | 11.09 | 13.38–53.69 | 19.08 |
A. cylindrica | 8.52 | 54.0 | 24.2 | 23.7 | 13.07 | 8.52–54.05 | 21.6 |
A. geniculata | 11.61 | 47.0 | 26.3 | 26.0 | 8.39 | 11.61–47.03 | 12.87 |
A. neglecta recta | 10.54 | 62.7 | 35.0 | 36.2 | 14.46 | 10.54–62.71 | 26.5 |
A. peregrina | 9.84 | 53.6 | 27.8 | 25.9 | 9.89 | 9.84–53.62 | 11.34 |
A. speltoides tauschii | 13.25 | 40.0 | 23.5 | 22.2 | 5.93 | 13.25–39.97 | 8.39 |
A. triuncialis | 5.60 | 50.1 | 28.2 | 28.2 | 11.24 | 5.60–50.06 | 15.18 |
A. uniaristata | 14.35 | 62.4 | 38.2 | 39.3 | 12.87 | 14.35–62.38 | 22.83 |
A. ventricosa | 14.10 | 40.0 | 26.3 | 25.7 | 7.44 | 14.10–40.04 | 12.77 |
H. vulgare distichon | 5.19 | 29.6 | 19.7 | 22.2 | 8.12 | 5.19–29.59 | 8.32 |
T. dicoccum | 6.17 | 41.5 | 16.5 | 12.8 | 8.66 | 6.17–41.55 | 14.07 |
T. monococcum | 6.68 | 36.6 | 20.1 | 19.1 | 7.11 | 6.68–36.61 | 10.44 |
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Table 4—source data 1
Summary statistics of the length of wild grass grains and domestic cereal starch granules.
- https://cdn.elifesciences.org/articles/72976/elife-72976-table4-data1-v2.docx
Pairwise Wilcoxon test performed on the length distribution of modern starches from Aegilops, Hordeum, and Triticum species (p value: not significant/ns >0.05; *<0.05; **<0.01; ***<0.001).
A. caudata | A. comosa | A. crassa | A. cylindrica | A. geniculata | A. neglecta recta | A. peregrina | A. speltoides tauschii | A. triuncialis | A. uniaristata | A. ventricosa | H. vulgare distichon | T. dicoccum | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
A. comosa | ns | – | – | – | – | – | – | – | – | – | – | – | – |
A. crassa | *** | *** | – | – | – | – | – | – | – | – | – | – | – |
A. cylindrica | ns | ns | *** | – | – | – | – | – | – | – | – | – | – |
A. geniculata | ns | * | *** | ns | – | – | – | – | – | – | – | – | – |
A. neglecta recta | *** | *** | ns | *** | *** | – | – | – | – | – | – | – | – |
A. peregrina | * | ** | ** | ns | ns | ** | – | – | – | – | – | – | – |
A. speltoides tauschii | ns | ns | *** | ns | ns | *** | * | – | – | – | – | – | – |
A. triuncialis | * | ** | ** | ns | ns | * | ns | * | – | – | – | – | – |
A. uniaristata | *** | *** | ns | *** | *** | ns | *** | *** | *** | – | – | – | – |
A. ventricosa | ns | * | *** | ns | ns | *** | ns | ns | ns | *** | – | – | – |
H. vulgare distichon | ns | ns | *** | ns | *** | *** | *** | * | *** | *** | *** | – | – |
T. dicoccum | ns | * | *** | ** | *** | *** | *** | *** | *** | *** | *** | ns | – |
T. monococcum | ns | ns | *** | ns | *** | *** | *** | * | *** | *** | *** | ns | * |
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Table 5—source data 1
Length of modern starch granules of Aegilops, Hordeum, and Triticum species.
- https://cdn.elifesciences.org/articles/72976/elife-72976-table5-data1-v2.xlsx