Skip to content
BY 4.0 license Open Access Published by De Gruyter Open Access September 14, 2021

Developing a Reference Collection for Starch Grain Analysis in Early Neolithic Western Temperate Europe

  • Clarissa Cagnato EMAIL logo , Caroline Hamon , Aurélie Salavert and Michelle Elliott
From the journal Open Archaeology


While we know that cereals played an important role in the diet of Linearbandkeramik (LBK) and Blicquy/Villeneuve-Saint-Germain (BVSG) populations in the Paris Basin, many questions remain to be answered as to the real contribution of other plants. To assess this topic, the recovery of other lines of data beyond macrobotanicals is crucial: starch grains have the potential to reveal additional information regarding past plant use. However, in Western Europe, in particular, for the Neolithic period, there is a significant lag in the development of the discipline. We, therefore, present how our current reference collection (composed of nearly 100 taxa spread across 35 families) was established, the reasoning behind our plant selections, and where the material comes from. Overall, our work shows that even though not all the selected plant organs produce diagnostic starch grains, it may be possible to broaden the spectrum of plants likely consumed by Early Neolithic (and beyond) populations in the Paris Basin, in particular concerning the use of wild plants and specific plant parts, especially underground storage organs (tubers, rhizomes, roots, bulbs, etc.). We believe our research will help guide future scholars in the creation of their own starch grain reference collection and to carry out such analyses on archaeological material from this region by consulting our image database. We conclude by providing a brief summary of what the starch grain record in the Paris Basin tells us to date on ancient plant use.

1 Introduction

Plant macroremains (those visible to the naked eye) are some of the most direct evidence of the Neolithic diet, especially when they are found charred in a domestic context. Nevertheless, charred plant remains alone permit only a limited knowledge of the diversity of plants and plant parts that were consumed, as fire also damages the most fragile plant parts (Fritz & Nesbitt, 2014). This statement is applicable to the Early Neolithic period in the Paris Basin where our knowledge of plant use is primarily based on the archaeobotanical record of sites ranging from the Aisne Valley in France to Hesbaye in central Belgium (Figure 1) (Bakels, 1984, 1999, 2009; Berrio, 2011; Dietsch-Sellami, 2004; Hamon, Salavert, Dietsch-Sellami, & Monchablon, 2019; Salavert, 2010, 2011). The Linearbandkeramik (LBK) culture, which originated in central Europe and expanded rapidly westwards reaching the Paris Basin around 5100 BC (Salavert, 2017), was subsequently replaced around 4900 BC by another group known as the Blicquy/Villeneuve-Saint-Germain (BVSG).

Figure 1 
               Map showing the geographical extent of the Paris Basin covering northern France and parts of Belgium (in orange), and sites mentioned in the text. Note: sites shown are not necessarily contemporaneous. Map modified from CC-BY-SA-3.0 by Alexrk2.
Figure 1

Map showing the geographical extent of the Paris Basin covering northern France and parts of Belgium (in orange), and sites mentioned in the text. Note: sites shown are not necessarily contemporaneous. Map modified from CC-BY-SA-3.0 by Alexrk2.

While we know that cereals played an important role in the diet of these LBK and BVSG populations (Bakels, 2014; Hamon, 2008; Salavert, 2011), many questions remain to be answered regarding the real contribution of other plants, as no single dataset can provide the entire spectrum of plants used in the past (Colledge & Conolly, 2014). For example, what other plants were processed and consumed in western LBK regions, and more specifically the Paris Basin? Did Neolithic populations consume fewer wild resources than those of the Mesolithic (or none, see Bogucki, 2000, p. 204)? What was the function of different tools such as grinding stones and ceramics? To answer these questions, the recovery of other lines of data is crucial, and this is where starch grains have the potential to reveal additional information regarding past plant use.

To create our reference collection, we began by consulting the existing macrobotanical record of our region, but as mentioned, it is limited by preservation issues. Thus, to amplify our reference collection, we turned to archaeobotanical data from neighboring countries in temperate Western Europe, notably the Netherlands, Switzerland, Germany, Denmark, and Poland for both the preceding period (Mesolithic, c. 8500–3500 BC) but also for slightly later periods, notably the Bronze Age (c. 2200 BC). We also consider ethnobotanical reviews of plants used today across Europe to provide a wider perspective of possible plant use. By doing so, we were able to consider the potential and limits of the methodology: what starch grains can we expect to find and identify in the archaeobotanical record? Which domesticated plants, recovered in macrobotanical form, can we identify to species? Which ones are truly diagnostic? Which plants will we be unable to document through this methodology? We thus share our reference collection in the form of an image database[1] in the hope that similar analyses will multiply in the Early Neolithic contexts of temperate Europe, broadening our perception, and understanding of the exploitation of plants for this period and region.

2 Starch Grain Analysis

Starch grains are energy storage units of plants, ranging from 1 to 100 μm (1 μm = 0.001 mm) and composed of two different glucose chains, amylose and amylopectin (for an in-depth review of their complex structure see Buléon, Colonna, Planchot, & Ball, 1998). The grains are synthesized in plastids found in leaves as a result of photosynthesis and then stored in various organs in a plant; however, dense amounts of starch are often concentrated in seeds, tubers, and fruits (Haslam, 2004). Like other macro- and microbotanical remains, starch grains display a range of characteristics, which include size and shape, and are the result of their genetic makeup (Copeland & Hardy, 2018). Although morphological similarities exist between and within species, in some cases, the starch grains can be highly diagnostic to a particular plant taxon. Other important features that often allow them to be identified to taxon include the location of the hilum (the point from which the grain starts to grow), presence of lamellae (the growth rings), fissures, and the extinction cross, also known as the Maltese cross, a feature visible only when viewed under cross-polarized light (Gott, Barton, Samuel, & Torrence, 2006).

While starch grains are susceptible to digestive enzymes (Hardy et al., 2009), they are resistant to many types of processes including grinding and drying (Cortella & Pochettino, 1994), and surprisingly, can survive processes such as cooking and carbonization (Babot, 2003; Chantran & Cagnato, 2021; Crowther, 2009; Pagán Jiménez, Guachamin-Tello, Romero-Bastidas, & Vásquez-Ponce, 2017). While starch is insoluble in cold water, it is permanently affected by extreme changes in temperature and moisture (Haslam, 2004). During cooking, gelatinization occurs, which results in the loss of native structure and morphology (Crowther, 2012). Although the long-term survival[2] of starch grains in the archaeological record has yet to be fully explained (see Copeland & Hardy, 2018; Mercader et al., 2018a for an excellent review on the topic), the presence of these remains makes it especially useful for understanding plants that are often under-represented in the archaeological record, including roots, tubers, corms, and rhizomes. We will use the widely employed term of underground storage organ (USO) to refer to these organs, which are poorly understood since they do not preserve well in the archaeological record, partially because of their preparation styles (Hather & Hammond, 1994; Pearsall, 2000).

Besides indicating the presence of certain plants in the past, when recovered on artifacts, their presence can indicate the function of these objects, from for example studying the function of grinding stones (Cagnato & Ponce, 2017; Hamon, Cagnato, Barbier-Emery, & Salavert, 2021; Hayes, Cnuts, Lepers, & Rots, 2017; Liu et al., 2010) to what plants were contained in vessels (Duncan, Pearsall, & Benfer, 2009; Wang et al., 2016). Finally, parts of the diet of an individual (human or animal) can be reconstructed through the study of dental calculus (Henry, Brooks, & Piperno, 2011), intestinal remains (Cagnato et al., 2021), and coprolites (Horrocks, Irwin, Jones, & Sutton, 2004). It should be noted, however, that microremains recovered in the dental calculus may not always reflect diet but also the ingestion of medicines and/or craft activities (Copeland & Hardy, 2018; Hardy, Buckley, & Copeland, 2018).

On a broad regional scale, notable works making use of starch grain analysis include those for Southwest Asia (Hart, 2014), Eastern North America (Messner, 2011), Central America (Piperno & Holst, 1998), the Canadian Plains (Zarrillo & Kooyman, 2006), Indonesia (Lentfer, 2009), and Sub-Saharan Africa (Mercader et al., 2018b). Many studies focusing on the Neolithic, and in turn implying the beginnings of sedentary lifestyles and agriculture, have made use of the potential of starch grains. In particular, they have made it possible to establish the importance of certain resources, such as tubers, on Chinese grindstones and ceramics (Liu, Duncan, Chen, Liu, & Zhao, 2015; Liu, Kealhofer, Chen, & Ji, 2014; Wan et al., 2012; Yang et al., 2013, 2015, Yao et al., 2016). Other work on Neolithic contexts has been carried out in North Africa (Lucarini, Radini, Barton, & Barker, 2016), Israel (Nadel, Piperno, Holst, Snir, & Weiss, 2012), South East Asia (Barker & Richards, 2013, Barker et al., 2007, but on sediments), Oceania (Denham, Haberle, & Lentfer, 2004; Horrocks, Bulmer, & Gardner, 2008), and Japan (Shibutani, 2017), but also on the American continent (Dickau, Ranere, & Cooke, 2007; Inomata et al., 2020; Iriarte et al., 2004; Pearsall, Chandler-Ezell, & Zeidler, 2004; Piperno, Ranere, Holst, & Hansell, 2000). Beyond analyses on grinding stones and ceramics, dental calculus trapping starch grains and other microremains has made it possible to carry out analyses of the Early Neolithic in Sudan (Madella, García-Granero, Out, Ryan, & Usai, 2014; Out et al., 2016) and Iraq (Scott Cummings, Yost, & Sołtysiak, 2018).

Western Europe is therefore a real gap in the development of the discipline, in particular for the Neolithic period. Few works exist in our geographical area, one example is the work by Chevalier and Bosquet (2013, 2017) who studied grinding stones from Remicourt “En Bia Flo II” in Belgium, an LBK site. Beyond our immediate area, starch grain analysis has provided insights on the Neolithic period in other parts of Europe. At Tiszasziget (Hungary), starch grains have been extracted from ceramics dating to the Late Neolithic (5000–4500 BC, Pető, Gyulai, Pópity, & Kenéz, 2013). In Northern Germany at the site of Neustadt, Saul et al. (2012) extracted starch grains from charred residues, also collected in ceramic vessels dating between 4600 and 3700 cal BC. The presence of tubers was identified based on the collection of parenchyma fragments from inside ceramics from Late Mesolithic and Early Neolithic sites in the Netherlands[3] (Kubiak-Martens, Brinkkemper, & Oudemans, 2015; Raemaekers, Kubiak-Martens, & Oudemans, 2013), but also from Neolithic domestic sites, causewayed enclosures, and megalithic graves in Germany and Denmark (4000–1800 BC) (Klooss, Fischer, Out, & Kirleis, 2016).

We sought to address this gap by beginning a preliminary starch grain study on a range of Early Neolithic tools and ceramics from across the Paris Basin[4] (Hamon et al., 2021). To carry out a successful study, however, a solid reference collection is necessary. Unlike other parts of the world, where detailed plant lists and in some cases images of starch grains have been produced, this does not exist for Western Europe. In this paper, we present the characteristics of the reference collection we assembled (how and why the plants were included), to facilitate future analyses in this area in other Early Neolithic contexts in Western Temperate Europe.

3 Materials and Methods

3.1 Selecting Species for the Reference Collection

In this section, we consider how our current reference collection was established, the reasoning behind our plant selections, and where the material comes from.

We began by considering the lists of species identified in the macrobotanical record in archaeological assemblages of the Early Neolithic from northwestern Temperate Europe (Bakels, 1984, 1999; Berrio, 2011; Dietsch, 1997; Jadin & Heim, 2003; Salavert, 2011; Saqalli et al., 2014). These data indicate the presence of a range of both domesticated and pulses, as well as wild/weedy plants including Poaceae (i.e., wild cereals). A large majority of these plants were available in the reference collection (“carpothèque”) of the UMR 7209 at the Muséum National d’Histoire Naturelle (MNHN) in Paris, France. We thus included hulled wheat (Triticum monococcum, T. turgidum dicoccon) as these are the most important crops for the LBK. Naked barley (Hordeum vulgare subsp. nudum L.) and hulled barley (Hordeum vulgare subsp. vulgare L.), recovered in minor quantities in archaeobotanical assemblages, are also included. Two oat species (Avena sativa L., A. strigosa Schreb.) were also selected. Domesticated oats are clearly attested in Europe starting in the Bronze Age (Zohary, Hopf, & Weiss, 2012), while wild oat (e.g., A. strigosa) is documented in Late Neolithic contexts in France (Marinval, 1988 in Fairweather & Ralston, 1993). Domesticated pulses included peas (Pisum sativum L.) and lentils (Lens culinaris Medik.), the most frequent species, as well as vetches (Vicia sativa L./V. ervilia (L.) Willd.), and grass pea (Lathyrus sativus/L. cicera). Plants used for their oil, fibers, or for their psychoactive properties include flax (Linum usitatissimum L.) and opium poppy (Papaver somniferum L.).

Fruits are widespread recovered in the archaeobotanical record of the Neolithic period, whether in LBK contexts in the Paris Basin (Berrio, 2011; Salavert, 2011) or at sites in western-Central Europe spanning between 4400 and 2400 cal BC (Colledge & Conolly, 2014). While it is known that ripened fruits in general tend to be relatively poor in starch grains (with for example the exception of tropical fruits such as the banana or avocado), we still tested some fruits (Crataegus monogyna Jacq., Prunus spinosa L., Malus sylvestris (L.) Mill., Sambucus sp.). The presence of hazelnuts (Corylus avellana L.) and acorns[5] from deciduous oak (Quercus sp.) are reported from LBK sites in our region of interest and were therefore included (Berrio, 2011). We also included the starch-rich fruits of water caltrop (Trapa natans L.), which have been reported from Mesolithic sites in the Netherlands and at Schwarzenberg Lake in the Czech Republic (Divišová & Šída, 2015), and from Neolithic sites (3500–3000 cal. BC) in Slovenia (Tolar, Jacomet, Velušček, & Čufar, 2011).

Wild/weedy plants were also considered. For the most part, these were collected at the MNHN and completed with seeds from the collection at the Archaeobotany lab of ArScAn (UMR 7041, Arch. Env) in Nanterre, France. Seeds found commonly in Early Neolithic Paris Basin contexts include goosefoot (Chenopodium album L.) various species of a dock (Rumex ssp.), nipplewort (Lapsana communis L.), cleaver (Gallium ssp.), knotgrass (Polygonum ssp.), bromes (Bromus ssp.), green foxtail millets (S. viridis (L.) P. Beauv., S. verticillata (L.) P. Beauv.), black bindweed (Fallopia convolvulus (L.) Á. Löve), vetches (Vicia hirsuta (L.) Gray/V. tetrasperma (L.) Schreb.), hemp-nettle (Galeopsis sp.), and orache (Atriplex ssp.) (Bakels, 1999; Berrio, 2011; Dietsch, 1996; Salavert, 2011). The wild carrot (Daucus carota L.), in seed form, has been recovered from the region, albeit in small quantities (Berrio, 2011; Dietsch, 1996). It is unclear which part(s) would have been consumed, but roots (raw) and young stewed leaves are today consumed in some parts of Spain (Tardío, Pardo-de-Santayana, & Morales, 2006). We were able to test both the wild carrot's root and seed. The common reed (Phragmites australis (Cav.) Trin. ex Steud.), a completely edible plant – in particular the young stems and rhizomes – was also included in our collection as it has been reported from Neolithic sites in central Europe (Colledge & Conolly, 2014) but also from Mesolithic contexts (Kubiak-Martens, 1999). Also, in the Poaceae family, and found in small quantities in the Paris Basin archaeobotanical record, are cockspur grass (Echinochloa crus-galli (L.) Beauv.) and annual meadow grass (Poa annua L.), which we included (Bakels, 1999; Berrio, 2011). Hairy crab-grass (Digitaria sanguinalis (L.) Scop.) does not seem to be mentioned in the archaeobotanical record of the Paris Basin, yet the seeds can be ground into flour (Simkova & Polesny, 2015), and it was present along the Mediterranean for potential use by the Neolithic populations between the early sixth millennium to the late third millennium BC (Delhon, Binder, Verdin, & Mazuy, 2020).

We have also considered plants that are reported to have only appeared or become important in the region both earlier and later. For example, the earliest gold of pleasure (Camelina sativa (L.) Crantz) has been reported in Switzerland around 4000 BC, becoming more common only between 1800 and 1200 BC in south-eastern and central Europe (Zohary et al., 2012, cited by Larsson, 2013). Although there is some limited evidence of Camelina in France in the Middle Neolithic, it is argued that cultivation in Western France likely began in the Late Bronze Age (Toulemonde, 2010). The same can be said about the sweet chestnut (Castanea sativa Mill.), whose introduction to southern France dates to probably the Roman period (Buonincontri, Saracino, & Di Pasquale, 2015).

One of the major problems in trying to establish the actual spectrum of plant use is to a certain extent due to the lack of USOs, recovered in macrobotanical form, in part due to the way they are consumed or prepared (fresh or boiled), making it unlikely that they will end up carbonized (Scheel-Ybert, 2001). While exceptions exist regarding the presence and probable use of USOs for the entire Neolithic period from contexts across Northern, Central, and Western Europe, there is a real gap of information on the presence of USOs at Early Neolithic sites in the Paris Basin. The little data that we have beyond the Paris Basin come in the form of tubers of lesser celandine (Ficaria verna Huds.), which have been widely reported from Early Neolithic (4000–3400 BC) contexts in northern Germany and Denmark (Klooss et al., 2016), while wild garlic (Allium ursinum L.) is reported as having been consumed by the Neolithic populations living near the Chalain Lake in the Jura, France (Dommelier, Bentrad, Paicheler, Pétrequin, & Bouchet, 1998 citing Pétrequin & Pétrequin, 1988). Turnip[6] (Brassica rapa var. rapa) has been reported from waterlogged contexts (4400–2400 cal BC) in Central Europe (Colledge & Conolly, 2014) and in seed form from Stare gmajne (Slovenia) dated between 3500 and 3000 cal BC (Tolar et al., 2011). Other remains include a Liliaceae bulb from Early Neolithic (5240–4990 cal BC) contexts at the site of Taï near the Mediterranean (Bouby, Durand, Rousselet, & Manen, 2019), and tuber oat-grass bulbs (Arrhenatherum elatius subsp. bulbosum) from Late Neolithic Germany (3500–2800 cal BC) (Kirleis, Klooß, Kroll, & Müller, 2012). In fact, more information concerning the use of USOs comes from European Mesolithic sites in Poland, the Netherlands, and Scotland[7] (see full references in Kubiak-Martens, 2016), where Scanning Electron Microscope (SEM) techniques have made it possible to identify parenchyma. Archaeological research at these hunter-gatherer sites has yielded a rich collection of starchy foods in the form of knotgrass rhizomes (Polygonum sp.), tubers of arrowhead (Sagittaria cf. sagittifolia), and horsetail (Equisetum sp.). Moreover, a potential sedge family (Cyperaceae) corm/stem base along with Schoenoplectus lacustris (L.) Palla was also recovered. In Neolithic contexts, a few mentions are made for the recovery of Scirpus, Carex, Cyperus, and Bolboschoenus (Kirleis et al., 2012), although not all species will necessarily be related to food consumption (e.g., Scirpus lacustris, Dietsch, 1996). Bolboschoenus maritimus (syn. Scirpus bolboschoenus) charred tubers were recovered from Late Neolithic contexts in the Netherlands (Kubiak-Martens et al., 2015); the stem bases, nutlets (achenes) and tubers can all be consumed (Kubiak-Martens, 1999). It was interpreted that tubers of a related taxon (Bolboschoenus glaucus) were used to produce flat bread-like products at Shubayqa 1, a Natufian hunter-gatherer site in northeastern Jordan (Arranz-Otaegui, Carretero, Ramsey, Fuller, & Richter, 2018).

Late Mesolithic datasets (c. fifth mill. BC), reported from Tybrind Vig and Halsskov, both lacustrine areas in Denmark, testify to the presence of sea beet (Beta vulgaris ssp. maritima), whose roots are rich in starch and sugar, and pignut (Conopodium majus (Gouan) Loret.) tubers (Kubiak-Martens, 1999, 2002); we were unfortunately unable to get a hold of these latter two plants. Seeds of waterlilies (Nuphar and Nymphaea) have been reported from Mesolithic and Neolithic contexts, indicating that they were probably consumed (see Bouby, Dietsch-Sellami, Martin, Marinval, & Wiethold, 2018; Dietsch, 1996; Kirleis et al., 2012;[8] Kubiak-Martens, 2002, 2016, pp. 128–129; Raemaekers et al., 2013). However, it is known that waterlily rhizomes are also edible, and their consumption is widely attested in the ethnographic record (Kubiak-Martens, 2016). The Mesolithic archaeobotanical record is also a good source of information on other plants besides USOs. For example, a charred caryopsis of floating sweetgrass (Glyceria fluitans (L.) R. Br.) was recovered at Tybrind Vig (Kubiak-Martens, 1999). Ethnobotanical studies in the Czech Republic indicate that these seeds can be ground into flour (Simkova & Polesny, 2015). We were unable to get a hold of this species but tested a relative-G. maxima-instead. Cattail reeds (Typha sp.) have a long history of use, as far back as the Upper Palaeolithic, where hunter-gatherers at the Bilancino site in Italy prepared flour from the starchy rhizomes (Aranguren, Becattini, Lippi, & Revedin, 2007). They are also reported in Mesolithic contexts from northern Netherlands and Poland (Kubiak-Martens, 1999; Perry, 1999). Two species are noted as likely being present at the time, T. angustifolia and T. latifolia, and we were able to include the latter in our reference collection.

The presence of a rare celery (Apium graveolens L.) schizocarp[9] from Parkhaus Opera, a Neolithic site on the shore of Lake Zurich, Switzerland (3176–3153 BC), indicates that this plant may have been used as a bread condiment (Heiss et al., 2017). We, therefore, included this plant, along with wild celery (Apium graveolens var. graveolens) in our reference collection. Other members of the Apiaceae family, although not reported in the literature to date, were considered: bulbous chervil (Chaerophyllum bulbosum L.) and celeriac (Apium graveolens var. rapaceum), both of which were purchased from local markets in France.

Raemaekers et al. (2013) cogently noted that species recovered in seed form from Early Neolithic sites in the Netherlands produce edible stems or fleshy shoots. These include glasswort (Salicornia europaea L.), sea aster (Aster tripolium L.), stinging nettle (Urtica dioica L.), greater burdock (Arctium lappa L.), and chickweed (Stellaria media (L.) Vill.), to mention but a few. Remains of Arctium minor and stinging nettle are reported from central European Neolithic sites, dated between 4400 and 2400 cal BC (Colledge & Conolly, 2014), although it is unclear which parts of the plants were recovered. The authors do note that for Arctium minor, the entire plant is edible, while the stinging nettle is usually used for its leaves and oil. We were unable to find any archaeobotanical evidence of sea aster or of glasswort. However, the leaves of the former are well known for their edibility, while the young stems of the latter are consumed. Seeds of chickweed are reported from the Paris Basin in small quantities, but not necessarily from the Early Neolithic (see Bakels, 1999).

The recovery of ferns is rare in the archaeobotanical record, yet some examples do exist and document their use as food or medicines (see for example Fiorin et al., 2018 who studied dental calculus from people in Medieval Majorca). In the case of bracken fern (Pteridium aquilinum (L.) Kuhn), this plant has been widely used around the world, especially for its rhizomes and young fronds (Divišová & Šída, 2015). Mesolithic period charred parenchyma studied by Kubiak-Martens (2008) was shown to belong to bracken and likely male fern (Dryopteris filix-mas (L.) Schott). Bracken fern was also recovered from Late Bronze (905–869 BC) settlements in the French Alps (Bouby & Billaud, 2005).

As there is a lack of data on USOs in the archaeobotanical record, we turned to ethnobotanical reviews that mentioned roots, tubers, and corms as being consumed to expand our reference collection. These studies for example mentioned the use of consuming raw rampion bellflower (Campanula rapunculus L.) roots,[10] as well as their shoots and leaves (Mattalia, Quave, & Pieroni, 2013; Simkova & Polesny, 2015). Other USOs consumed include rhizomes of Lords-and-Ladies (Arum maculatum L.), used as a flour or boiled, and tubers of bulbous chervil, which are noted as being used in boiled dishes (Simkova & Polesny, 2015). The rhizomes of common comfrey (Symphytum officinale L.) are also noted as being eaten. The consumption of the tubers of the tuberous pea (Lathyrus tuberosus L.) seems to be common in parts of Italy, France, and the Netherlands, but more rarely in Spain (Mattalia et al., 2013; Tardío et al., 2006). We also included the tubers of great pignut (Bunium bulbocastanum L.) as these are reported as being consumed in the Western Italian Alps (Mattalia et al., 2013). The bitter roots of the great yellow gentian (Gentiana lutea L.) are used to prepare a digestive liquor (Abbet et al., 2014). Dandelion (Taraxacum officinale (L.) Weber ex F.H. Wigg) leaves and flowers are used in multiple ways (Abbet et al., 2014), and so are the roots, which can be consumed either raw or cooked. Horseradish (Armoracia rusticana G.Gaertn., B.Mey. & Scherb.) is another plant that could have been gathered for its roots (Saul et al., 2012), even though to date no archaeobotanical remains have been recovered suggesting its use in ancient times. Parsnip (Pastinaca sativa L.) is a commonly consumed root but not well-known in the archaeological record: according to Zohary et al. (2012) it has been recovered from Roman sites in Europe.

To date, we have collected and tested 99 species that cover 35 families (Table 1). Out of those, we tested fruits (n = 11), seeds (n = 69), underground storage organs (n = 22), and stems (n = 2). For three taxa (Daucus carota, Rumex acetosa, and Schoenoplectus lacustris), we tested both the seeds and the USOs, while we also tested acorns at different stages of maturity. Thus, we have a total of 103 samples. Besides those obtained from the herbaria, most of the plants were either purchased from local markets (turnip, chestnut, bulbous chervil) or nurseries (e.g., lesser celandine, common comfrey). A small percentage of the taxa were directly gathered from the wild (e.g., dandelion, wild carrot, wild garlic).

Table 1

Plant taxa included in our reference collection with provenience information and whether starch grains are present

Binomial name Common name Family Source Plant part tested Starch present
Avena sativa L. Oats Poaceae MNHN UMR 7209 Seed Y
Avena strigosa Schreb. Bristle oats Poaceae MNHN UMR 7209 Seed Y
Hordeum vulgare subsp. nudum L. Cultivated naked barley Poaceae Caroline Hamon Seed Y
Hordeum vulgare subsp. vulgare L. Cultivated hulled barley Poaceae MNHN UMR 7209 Seed Y
Triticum aestivum L. Bread wheat Poaceae MNHN UMR 7209 Seed Y
Triticum durum Desf. Macaroni wheat Poaceae MNHN UMR 7209 Seed Y
Triticum turgidum Desf. Durum wheat Poaceae MNHN UMR 7209 Seed Y
Triticum monococcum L. Einkorn Poaceae MNHN UMR 7209 Seed Y
Triticum cf. timopheevi New glume wheat Poaceae MSH Mondes Seed Y
Triticum dicoccon Schrank. Cultivated emmer Poaceae MNHN UMR 7209 Seed Y
Lathyrus sativus L. Grass pea Fabaceae MNHN UMR 7209 Seed Y
Lens culinaris Medik. Lentil Fabaceae MNHN UMR 7209 Seed Y
Pisum sativum L. Pea Fabaceae MNHN UMR 7209 Seed Y
Vicia ervilia (L.) Willd. Bitter vetch Fabaceae MNHN UMR 7209 Seed Y
Vicia sativa L. Common vetch Fabaceae MSH Mondes Seed Y
Oily/Fiber Plants
Linum usitatissimum L. Flax Linaceae MNHN UMR 7209 Seed Y
Papaver somniferum L. Opium poppy Papaveraceae MNHN UMR 7209 Seed N
Pinus sp. Pine kernel/nut Pinaceae Market Seed Y
Wild Plants
Aesculus hippocastanum L. Horse chestnut Sapindaceae Laura Longo Fruit Y
Aethusa cynapium subsp. cynapium Fool’s parsley Apiaceae MSH Mondes Seed N
Allium ursinum L. Wild garlic Amaryllidaceae Charlène Bouchaud USO N
Armoracia rusticana Horseradish Brassicaceae Nursery USO Y
Aster tripolium (Jacq.) Dobrocz Sea aster Asteraceae MNHN UMR 7209 Seed N
Apium graveolens L. Celery Apiaceae Market Stem N
Apium graveolens var. graveolens Wild celery Apiaceae MSH Mondes Seed N
Apium graveolens var. rapaceum Celeriac Apiaceae Market USO N
Atriplex hortensis L. Garden orache Amaranthaceae MSH Mondes Seed N
Beta vulgaris ssp. maritima Sea beet Amaranthaceae MSH Mondes Seed Y
Bolboschoenus maritimus (L.) Palla Sea club-rush Cyperaceae MSH Mondes Seed Y
Brassica rapa var. rapa Turnip Brassicaceae Market Seed Y
Bromus secalinus L. Rye brome Poaceae MSH Mondes Seed Y
Bromus sterilis L. Barren brome Poaceae MSH Mondes Seed Y
Bromus tectorum L. Drooping brome Poaceae MSH Mondes Seed Y
Bunium bulbocastanum L. Great pignut Apiaceae Nursery USO Y
Camelina sativa (L.) Crantz Gold of pleasure Brassicaceae Francoise Toulemonde Seed N
Campanula rapunculus L. Rampion bellflower Campanulaceae Nursery USO N
Capparis spinosa subsp. rupestris Capparaceae MSH Mondes Seed N
Carex hirta L. Sedge Cyperaceae MSH Mondes Seed Y
Castanea sativa Mill. Chestnut Fagaceae Market Fruit Y
Chaerophyllum bulbosum L. Bulbous chervil Apiaceae Market USO Y
Chenopodium album L. Goosefoot Amaranthaceae MSH Mondes Seed Y
Corylus avellana L. Hazelnut Betulaceae Market Fruit N
Crataegus monogyna Jacq. Common hawthorn Rosaceae MNHN UMR 7209 Fruit N
Cyclamen sp. Cyclamen Primulaceae Laura Longo USO Y
Cyperus esculentus L. Yellow nutsedge Cyperaceae MSH Mondes USO Y
Cyperus rotundus L. Purple nutsedge Cyperaceae Laura Longo USO Y
Daucus carota L. Wild carrot Apiaceae Aurélie Salavert USO N
Daucus carota L. Wild carrot Apiaceae Aurélie Salavert Seed N
Digitaria sanguinalis (L.) Scop. Hairy crab-grass Poaceae MSH Mondes Seed Y
Echinochloa crus-galli (L.) Beauv. Cockspur grass Poaceae MNHN UMR 7209 Seed Y
Elymus caninus L. Bearded wheatgrass Poaceae MSH Mondes Seed Y
Equisetum spp. Horsetail Equisetaceae Nursery Stem Y
Erythronium dens-canis L. Dogs’ tooth-violet Liliaceae Nursery USO Y
Fagus sylvatica L. Common beech Fagaceae MSH Mondes Seed Y
Fallopia convolvulus (L.) Á.Löve Black-bindweed Polygonaceae MSH Mondes Seed Y
Festuca arundinacea Schreb. Tall fescue Poaceae MSH Mondes Seed Y
Ficaria verna Huds. Lesser celandine Ranunculaceae Nursery USO Y
Galeopsis segetum Neck. Hemp-nettle Lamiaceae MSH Mondes Seed N
Galium aparine L. Cleaver Rubiaceae MSH Mondes Seed N
Gentiana lutea L. Great yellow gentian Gentianaceae Nursery USO Y
Glyceria maxima (Hartm.) Holmb. Greater sweetgrass Poaceae MSH Mondes Seed N
Iris sibirica L. Siberian iris Iridaceae Laura Longo USO Y
Lapsana communis Juss. Nipplewort Asteraceae MSH Mondes Seed N
Lotus corniculatus L. Common bird’s-foot trefoil Fabaceae MSH Mondes Seed N
Lupinus albus L. White lupin Fabaceae MSH Mondes Seed N
Malus sylvestris (L.) Mill. Crabapple Rosaceae Market Fruit N
Medicago lupulina L. Black medick Fabaceae MSH Mondes Seed N
Pastinaca sativa L. Parsnip Apiaceae Market USO Y
Phleum pratense L. Timothy-grass Poaceae MNHN UMR 7209 Seed Y
Phragmites australis (Cav.) Trin. ex Steud. Reed Poaceae Nursery USO Y
Plantago major L. Broadleaf plantain Plantaginaceae MSH Mondes Seed Y
Poa annua L. Annual meadow grass Poaceae MSH Mondes Seed Y
Polygonum lapathifolium L. Pale persicaria Polygonaceae MSH Mondes Seed Y
Polygonum bistorta Delarbre Common bistort Polygonaceae MSH Mondes Seed Y
Polygonum persicaria Gray Lady’s thumb Polygonaceae MSH Mondes Seed Y
Prunus spinosa L. Blackthorn Rosaceae Wild Fruit N
Quercus ilex L. Acorn (immature) Fagaceae Laura Longo Fruit Y
Quercus ilex L. Acorn (ripe) Fagaceae Laura Longo Fruit Y
Rumex acetosa L. Common sorrel Polygonaceae MSH Mondes Seed Y
Rumex acetosa L. Common sorrel Polygonaceae Nursery USO Y
Rumex crispus L. Curly dock Polygonaceae MNHN UMR 7209 Seed Y
Rumex obtusifolius L. Bitter dock Polygonaceae MSH Mondes Seed Y
Sagittaria sagittifolia L. Arrowhead Alismataceae Nursery USO Y
Sambucus nigra L. Black elder Adoxaceae MSH Mondes Fruit N
Schoenoplectus lacustris (L.) Palla Common club-rush Cyperaceae Nursery USO Y
Schoenoplectus lacustris (L.) Palla Common club-rush Cyperaceae MSH Mondes Seed Y
Setaria verticillata (L.) P. Beauv. Bristly foxtail Poaceae MSH Mondes Seed Y
Setaria viridis (L.) P. Beauv. Green foxtail millet Poaceae MNHN UMR 7209 Seed Y
Sinapsis arvensis L. Wild mustard Brassicaceae MNHN UMR 7209 Seed N
Solanum nigrum L. Black nightshade Solanaceae MNHN UMR 7209 Seed N
Stellaria media (L.) Vill. Chickweed Caryophyllaceae MSH Mondes Seed Y
Stipa capillata L. Very slender feather grass Poaceae MSH Mondes Seed Y
Symphytum officinale L. Common comfrey Boraginaceae Nursery USO Y
Taraxacum sp. Dandelion Asteraceae Market USO N
Tetragonolobus purpureus Moench Winged pea Fabaceae MSH Mondes Seed N
Trapa natans L. Water caltrop Lythraceae Laura Longo Fruit Y
Trifolium dubium Sibth. Field clover Fabaceae MSH Mondes Seed N
Typha domingensis Pers. Cattail reed Typhaceae MNHN UMR 7209 Seed N
Typha latifolia L. Broadleaf cattail Typhaceae Nursery USO Y
Urtica dioica L. Stinging nettle Urticaceae MNHN UMR 7209 Seed N
Vicia hirsuta (L.) Gray Hairy vetch Fabaceae MSH Mondes Seed Y
Vicia tetrasperma agg. Four-seeded vetch Fabaceae MSH Mondes Seed Y
Veronica hederifolia agg. Ivy-leaved speedwell Plantaginaceae MSH Mondes Seed N

3.2 Processing the Modern Plant Samples and Recording the Starch Grains

Fresh plant material was first cleaned and peeled (when dealing with tubers, roots, and fruits). Otherwise, material from herbaria or already dried was directly processed. The sample (seed, fruit, USO) was then gently crushed using a mortar and pestle. To avoid damaging the starch grains in an extremely hard seed (e.g., cereals) they were placed for 1–2 h in distilled water, before cutting off a small piece with a clean scalpel and then gently rubbing it against a clean microscope slide (using a clean toothpick when necessary). A drop of 1:1 glycerine:water solution was added before covering with a coverslip. The glycerine solution allows for the starch grains to be more easily rotated and viewed in both cross-polarized and transmitted light. Although the sample will eventually dry, it can be re-hydrated before observation using this afore-mentioned solution. The reference slides were then examined at 600× magnification using a Nikon E600 POL microscope and starch grains were measured using NIS Elements software, which provided statistical data including the mean, standard deviation, and range of sizes. The number of grains counted by different specialists when establishing their reference collection varies greatly, from at least 50 grains (Hart, 2014; Li, Pagán‐Jiménez, Tsoraki, Yao, & Van Gijn, 2020; Musaubach, Plos, & Babot, 2013; Piperno, Weiss, Holst, & Nadel, 2004), to at least 300 (Mercader et al., 2018b). We measured 50 simple starch grains whenever possible, and these were randomly chosen on the slide. Photographs were taken under transmitted and cross-polarized light, and attributes such as the shape of the grain, the type and position of the hilum as well as the presence or absence of facets and fissures, were described (see Supplementary materials for photos and descriptions). Whenever possible, we kept materials (stored separately) in case new slides need to be prepared. Some species in our reference collection (e.g., Trapa natans, Iris sibirica) were obtained from Dr Laura Longo (Università Ca’ Foscari), who prepared ultrapure starch pellets extracted using a sequential water/ethanol protocol.[11] In this case, a small amount of the resulting powder was placed on slides, where a drop of 1:1 glycerine:water solution was then added, before adding a coverslip.

4 Results and Discussion

We tested 103 different plant parts, but since we tested immature and mature acorns, our calculations are based on a total of 102 different taxa. Out of these, we found that 69 produced starch grains (68%), with a large majority of USOs producing starch grains (77%), followed by seeds (68%). There is a 50/50 chance of recovering starch from fruits or stems, but this probability may well vary if we tested a wider number of stems. It is immediately clear that compared to Hart (2014), who found that only 10 out of 64 species present across Southwest Asia produced starch grains, we found a much greater number of species that do produce these microbotanical remains. This is of course a positive outcome, but we must also keep in mind that even when starch is produced, not all starch grains are diagnostic to genus and even less to species. Moreover, some species will produce exceedingly small grains that will be hard to effectively see using a stereomicroscope. When the grains produced are below 5 μm, these are difficult to characterize with a light microscope, even at 600×. This is the case for species included in our collection such as sea beet, goosefoot, chickweed, as well as some taxa in the grass family (Festuca, Poa). In this case, additional higher resolution microscopy such as that obtained by SEM may help to further identify these starch grains (Jane, Kasemsuwan, Leas, Zobel, & Robyt, 1994).

The fruits we investigated for most part did not contain starch, except for acorns, water caltrop, chestnut, and horse chestnut. It has been reported that unripe fruits will contain more starch than ripe fruits, as these will convert the starch to sugar as the fruit ripens (Gott et al., 2006). We tested both green and mature acorns, and while our results do not suggest that there is a significant difference, more tests involving other taxa are necessary.

We also found that the presence of starch in seeds is variable. Cereals, notably those in the Triticeae tribe (e.g., wheat and barley), have a bimodal distribution (grains come in two sizes), and produce abundant quantities of starch grains, with the larger ones being lenticular in shape. The morphology of some wild grasses has also been explored in depth (Hart, 2014; Piperno et al., 2004; see Yang & Perry, 2013 for a specific focus on species found in China). We found starch grains in all the grass seeds we collected, except for greater sweetgrass. The smallest grains were produced by cockspur, stipa, timothy-grass, and Poa, followed by Digitaria and Setaria; with barren brome seeds producing the largest starch grains among the wild grasses sampled.

The Fabaceae family produces very recognizable starch grains with a distinct longitudinal cleft. We extracted starch from all the domesticated taxa tested (peas, lentils, beans grass pea, and bitter vetch), and from both wild Vicia species. We discovered that not all members of this family produce starch grains, this was the case for species of Lotus, Medicago, and Lupinus.

As expected, seeds with high oil/fat content, and used for extracting oil such as opium poppy and hazelnuts, did not contain starch grains. As noted by Gott et al. (2006), in these plants, the main storage is lipid, and therefore, little to no starch will be produced or stored. We did, however, find starch grains in the seeds of flax, but only in the ultrapure pellets prepared by Laura Longo and Elena Badetti. Even then, these starch grains remained very tightly packed within the rest of the seed matrix. For this reason, it may be harder to detect them in the archaeological record if they are simply gently pounded or ground.

For this initial reference collection, we focused on seeds and USOs, and set aside the study of the leaves. In the latter, starch grains, known as transient or transitory starch, have been determined to be small (less than 7 μm; Gott et al., 2006; Haslam, 2004), and therefore probably not diagnostic. However, leaves will have to be included in the future as, more recently, the presence of larger (non-transient) starch grains in the leaves and stems of domesticated and wild plants have been noted (Liu, Wang, & Levin, 2017; Yang et al., 2014). Neolithic populations likely had access to a large range of plants they could have used for their greens (leaves, stems): for example, stinging nettle, wild carrot, dandelion, and dock/sorrel (Mattalia et al., 2013; Simkova & Polesny, 2015). It will be of interest to test these in the future to determine whether the leaves of such species produce diagnostic starch grains. After testing an important number of seeds from wild plants, we found that starch grains were present in a majority of these. However, based on their tendency to be on the smaller side (between 2 and 11 μm), the probability of correctly identifying the ones we recover in the archaeobotanical record is rather slim, unless an SEM is used.

One issue that is clearly at the forefront of starch grain analysis is its utility in identifying underground storage organs in the archaeobotanical record. However, knowing which taxa to include in a reference collection has been problematic when we rely on solely macrobotanical remains. For the Neolithic, few taxa have been reported (but see Klooss et al., 2016; Kubiak-Martens, 2016), yet it is evident that multiple plants could have been gathered by these populations. Hart (2014) already made a similar observation for studies of USOs in Southwest Asia. We believe that our biggest contribution here is to provide a list of putative plants, besides those already found at other sites in the Neolithic, whose USOs may have been consumed by LBK/BVSG populations in Paris Basin. It should be noted that not all USOs contain starch grains, in fact, other reserve carbohydrates can be present (either in place of or in addition). Both monocotyledons and dicotyledons have fructan-containing species, and their families include Poaceae, Campanulaceae, Amaryllidaceae, Iridaceae, Asteraceae, and Liliaceae (Ranwala & Miller, 2008). In our case, we were unable to find starch in the roots of wild garlic (Amaryllidaceae) or dandelion (Asteraceae); however, the Siberian iris (Iridaceae), dogs’ tooth-violet (Liliaceae), and the common reed (Poaceae) were all rich in starch. We, therefore, believe that it is important to also test species in these families. Overall, we found that a vast majority of the USOs we collected contained starch grains, for the most part, the grains are typical of USOs: elongated with eccentric hila, but other forms of starch grains exist. Taxa that did not produce starch grains in their USOs include members of the Apiaceae family (wild carrot, wild celery, celeriac), but this is not a fast rule as other members in the same family (bulbous chervil and parsnip) have starch grains (although neither produces particularly diagnostic grains).

Our current work makes it clear that not all taxa will produce diagnostic starch grains and in turn be useful for archaeobotanical studies. If we disregard those that are too small to be clearly visible with a microscope at 600× (species which have already been discussed earlier in the text), we can propose the following observations regarding those taxa that are more likely to be diagnostic or identifiable. Cereals such as wheat and barley are easily categorized due to the presence of lenticular starch grains. However, to differentiate between these taxa, is not as straightforward, even when a large reference collection is available (see for example Bocanegra & Sáez, 2012). Moreover, when age and taphonomic factors are added to this mixture, differentiating wheat and barley may be even more problematic. One taxon that could potentially be confused with wheat or barley is Bromus sterilis; however, the grains are wider when viewed from the side and the extinction cross is not often bilateral. Oats will likely be identifiable if they are still packed into clusters, but we were unable to distinguish between the two species we tested. When the grains are loosened, these could be mistaken for Digitaria sanguinalis. The latter produces slightly more angular grains than those produced by both A. sativa and A. strigosa. Within the Cyperaceae family, the starch grains in the USOs of both Cyperus esculentus and C. rotundus are diagnostic. The same cannot be said for the starch grains in the seeds of Carex hirta and Bolboschoenus maritimus. The grains produced by taxa in the Fabaceae family are extremely diagnostic. Distinguishing between the various species may be possible, with peas, lentils, and Lathyrus sativus deemed especially diagnostic. Within the vetches, we found that V. hirsuta and V. tetrasperma lacked distinct lamellae, which are more visible on V. sativa and V. ervilia. We did find that several taxa produced small (5–8 μm on average), polyhedral or angular starch grains with a centric hilum. These include seeds of Echinochloa crus-galli, Phleum pratense, Bolboschoenus maritimus, and species of Setaria. However, the latter, in particular S. verticillata, will be likely distinguished by the presence of a distinct continuous double border. USOs of various species, as noted earlier, are more likely to be identified as such given their shape and relatively larger sizes. We propose that Brassica rapa var. rapa, Gentiana lutea, Erythronium dens-canis, and Ficaria verna could thus be identified in the archaeobotanical record. Finally, fruits of Aesculus hippocastanum and Quercus are also good candidates for identification.

While we were unable to gather all the plants that we had hoped to sample, for example, sea beet, pignut, tuber oat-grass, and tuberous pea[12] (see Table 2), the creation of a reference collection is often a work in progress, and we plan to continue expanding it. We believe our work thus far provides a starting base for starch grain analysis in the region. Besides continuing our search for new plant species that could have been used and present in the region during the Early Neolithic, some additional types of samples need to be considered. For example, we need to collect plants at different stages of maturation but also from different environments, as these factors may affect the size of the starch grains (see references in Gott et al., 2006). Other plant parts could have been utilized, for example, the inner bark tissue of pine and birch, which are starch-rich resources (Gott et al., 2006; Kubiak-Martens, 2016; Sandgathe & Hayden, 2003) and will also need to be considered. Experimental work that comprises mechanical processing (grinding, pounding), as well as thermal exposure (cooking) and fermentation, is essential to gather a broader picture of how starch grains are modified (see for example Cagnato, 2019; Chantran & Cagnato, 2021; Henry, Hudson, & Piperno, 2009; Li et al., 2020; Wang et al., 2017). Preliminary work (Cagnato, Hamon, & Salavert, in press) on domesticated species (cereals and pulses) has already been carried out, but we plan to expand this collection with the processing of wild plants, especially tubers and rhizomes, due to their presence in archaeological samples from the Paris Basin (Hamon et al., 2021).

Table 2

Plant taxa to be considered in the future to test for the presence of starch grains (list not exhaustive)

Binomial name Common name Family Plant part(s) Comments
Arctium lappa L. Greater burdock Asteraceae Root
Arrhenatherum elatius subsp. bulbosum Tuber oat-grass Poaceae Root
Arum maculatum L. Lords-and Ladies Araceae Root
Beta vulgaris ssp. maritima Sea beet Amaranthaceae Root
Cichorium intybus var. sativum Root chicory Asteraceae Root
Conopodium majus (Gouan) Loret. Pignut Apiaceae Root
Elymus repens (L.) Gould Coach grass Poaceae Rhizome Average measurement of grains given in Juhola, Etu-Sihvola, Näreoja, and Ruohonen (2014)
Glyceria fluitans (L.) R.Br. Flotating sweetgrass Poaceae Seed Average measurement of grains given in Juhola et al. (2014)
Heracleum sphondylium L. Hogweed Apiaceae Root
Lathyrus cicera L. Red pea Fabaceae Seed
Lathyrus tuberosus L. Tuberous pea Fabaceae Root
Lilium sp. Lily Liliaceae Bulb Photos of American species in Messner (2011)
Nuphar lutea (L.) Sm. Waterlily Nymphaeaceae Seed/Rhizome Tuber starch grain photos in Henry et al. (2011) and Messner (2011)
Nymphaea sp. Waterlily Nymphaeaceae Seed/Rhizome N. alba Rhizome starch grain photos in Henry et al. (2011)
Polygonum aviculare L. Common knotgrass Polygonaceae Seed Starch grain photos in Juhola et al. (2014)
Pteridium aquilinum (L.) Kuhn Bracken fern Dennstaedtiaceae Rhizome Starch grain photos in Horrocks et al. (2004)
Ruscus aculeatus L. Butcher’s-broom Asparagaceae Root
Salicornia europaea L. Glasswort Amaranthaceae Root
Typha angustifolia L. Reed Typhaceae Tuber Starch grain photos in Revedin et al. (2010)

Overall, our work here shows that through starch grain analysis, it may be possible to broaden the spectrum of plants likely consumed by Early Neolithic (and beyond) populations in the Paris Basin, in particular concerning the use of wild plants and specific plant parts, especially underground storage organs. Now that a large selection of starch-rich species has been identified, future research can focus on determining whether it will be possible to clearly differentiate between them. We hope that our research helps guide future scholars in the creation of their own starch grain reference collection as there is the necessity for a solid database, and this across disciplines.

4.1 What Does Starch Grain Analysis Tell Us Thus Far About LBK Food Processing?

To conclude, we provide a brief synthesis of what we know about food transformation to date in the Paris Basin. The application of starch grains has revealed that cereals were not the only foods processed on grinding stones. Notably, 9 LBK grinding stones from Remicourt “En Bia Flo II” in Belgium revealed the presence of wheat, barley, oats, peas, and acorns (Chevalier & Bosquet, 2013, 2017). Our research (Hamon et al., 2021), carried out on a large corpus of grinding stones (n = 32) from across LBK and BVSG sites in the Paris Basin, supports the notion that grinding stones were multipurpose as we also found starch grains of wheat, barley, and peas. Cooked (or at least heated) plants were also processed with these stone tools, based on our data. Finally, we also found evidence for the processing of different types of USOs (Hamon et al., 2021; Cagnato, Hamon, Salavert, & Elliott, in prep.), thereby proving that starch grain research can be fruitful and provide a new vision of past plant use.

Special Issue: THE EARLY NEOLITHIC OF EUROPE, edited by F. Borrell, I. Clemente, M. Cubas, J. J. Ibáñez, N. Mazzucco, A. Nieto-Espinet, M. Portillo, S. Valenzuela-Lamas, & X. Terradas


We wish to thank Laura Longo for kindly providing pellets of starch extracted from various plant species, as well as Françoise Toulemonde and Charlène Bouchaud for providing some of the species included in this research. Finally, we are grateful to Julien Wiethold for his help with proposing which plants to include in this study.

  1. Funding information: The work was financed by a DIM MAP Ile de France postdoctoral fellowship and the ANR Homes grant (ANR-18-CE27-0011).

  2. Author contributions: All authors have accepted responsibility for the entire content of this manuscript and approved its submission. CH, CC, AS, and ME designed the project, and CC prepared the reference collection. CC prepared the manuscript with contributions from all co-authors.

  3. Conflict of interest: Authors state no conflict of interest.

  4. Data availability statement: The datasets generated and analyzed during the current study are available in the collections of UMR 7041 ArScAn, Archéologies environnementales (MSH Mondes, Nanterre), UMR 8215 Trajectoires (9 Rue Malher, Paris), and UMR 7209 Archéozoologie, Archéobotanique: Sociétés, Pratiques et Environnements (AASPE) (55 Rue Buffon, Paris). Many of the dataset generated is also included in this article in its supplementary information file. Additional photos generated during the current study are available from the corresponding author on reasonable request.


Abbet, C. , Mayor, R. , Roguet, D. , Spichiger, R. , Hamburger, M. , & Potterat, O. (2014). Ethnobotanical survey on wild alpine food plants in Lower and Central Valais (Switzerland). Journal of Ethnopharmacology, 151(1), 624–634.10.1016/j.jep.2013.11.022Search in Google Scholar

Aranguren, B. , Becattini, R. , Lippi, M. M. , & Revedin, A. (2007). Grinding flour in Upper Palaeolithic Europe (25,000 years bp). Antiquity, 81(314), 845.10.1017/S0003598X00095946Search in Google Scholar

Arranz-Otaegui, A. , Carretero, L. G. , Ramsey, M. N. , Fuller, D. Q. , & Richter, T. (2018). Archaeobotanical evidence reveals the origins of bread 14,400 years ago in northeastern Jordan. Proceedings of the National Academy of Sciences, 115(31), 7925–7930.10.1073/pnas.1801071115Search in Google Scholar

Babot, M. D. P. (2003). Starch grain damage as an indicator of food processing. In D. M. Hart & L. A. Wallis (Eds.), Phytolith and starch research in the Australian-Pacific-Asian regions: The state of the art, Terra Australis 19 (pp. 69–82). Canberra, Australia: Pandanus Books.Search in Google Scholar

Bakels, C. (1999). Archaeobotanical investigations in the Aisne valley, northern France, from the Neolithic up to the early Middle Ages. Vegetation History and Archaeobotany, 8(1–2), 71–77.10.1007/BF02042844Search in Google Scholar

Bakels, C. (2014). The first farmers of the Northwest European Plain: Some remarks on their crops, crop cultivation and impact on the environment. Journal of Archaeological Science, 51, 94–97.10.1016/j.jas.2012.08.046Search in Google Scholar

Bakels, C. C. (1984). Carbonized seeds from northern France. Analecta Praehistorica Leidensia, 17, 1–27.Search in Google Scholar

Bakels, C. C. (2009). The Western European loess belt: Agrarian history, 5300 BC–AD 1000. New York: Springer Science & Business Media.10.1007/978-1-4020-9840-6Search in Google Scholar

Barker, G. , & Richards, M. B. (2013). Foraging–farming transitions in island Southeast Asia. Journal of Archaeological Method and Theory, 20(2), 256–280.10.1007/s10816-012-9150-7Search in Google Scholar

Barker, G. , Barton, H. , Bird, M. , Daly, P. , Datan, I. , Dykes, A. , & Higham, T. (2007). The ‘human revolution’ in lowland tropical Southeast Asia: The antiquity and behavior of anatomically modern humans at Niah Cave (Sarawak, Borneo). Journal of Human Evolution, 52(3), 243–261.10.1016/j.jhevol.2006.08.011Search in Google Scholar

Berrio, L. (2011). L'économie végétale au Néolithique ancient. Comparaison des données carpologiques des sites rubanés et Blicquy/Villeneuve-Saint-Germain dans le Bassin parisien et en Moyenne-Belgique. Mémoire Master 1 Archéologie Et Environnement Université De Paris I – Panthéon-Sorbonne. Paris, France: UFR D’histoire de L’art et Archéologie.Search in Google Scholar

Bocanegra, F. J. A. , & Sáez, J. A. L. (2012). Caracterización morfológica de almidones de los géneros Triticum y Hordeum en la Península Ibérica. Trabajos de prehistoria, 69(2), 332–348.10.3989/tp.2012.12095Search in Google Scholar

Bogucki, P. (2000). How agriculture came to north-central Europe. In T. D. Price (Ed.), Europe’s first farmers (pp. 197–218). Cambridge, UK: Cambridge University Press.10.1017/CBO9780511607851.009Search in Google Scholar

Bouby, L. , & Billaud, Y. (2005). Identifying prehistoric collected wild plants: A case study from Late Bronze Age settlements in the French Alps (Grésine, Bourget Lake, Savoie). Economic Botany, 59(3), 255–267.10.1663/0013-0001(2005)059[0255:IPCWPA]2.0.CO;2Search in Google Scholar

Bouby, L. , Dietsch-Sellami, M. F. , Martin, L. , Marinval, P. , & Wiethold, J. (2018). Ressources végétales et économie de subsistance au Néolithique en France (6000–2000 av. J.-C.). In D. Garcia & J. Guilaine (Eds), La Protohistoire de la France (pp. 141–152). Paris, France: Hermann.10.3917/herm.garci.2018.01.0142Search in Google Scholar

Bouby, L. , Durand, F. , Rousselet, O. , & Manen, C. (2019). Early farming economy in Mediterranean France: Fruit and seed remains from the Early to Late Neolithic levels of the site of Taï (ca 5300–3500 cal bc). Vegetation History and Archaeobotany, 28(1), 17–34.10.1007/s00334-018-0683-xSearch in Google Scholar

Buléon, A. , Colonna, P. , Planchot, V. , & Ball, S. (1998). Starch granules: Structure and biosynthesis. International Journal of Biological Macromolecules, 23(2), 85–112.10.1016/S0141-8130(98)00040-3Search in Google Scholar

Buonincontri, M. P. , Saracino, A. , & Di Pasquale, G. (2015). The transition of chestnut (Castanea sativa Miller) from timber to fruit tree: Cultural and economic inferences in the Italian peninsula. The Holocene, 25(7), 1111–1123.10.1177/0959683615580198Search in Google Scholar

Cagnato, C. (2019). Hervir y moler: Descifrando técnicas de elaboración de alimentos a través del análisis microscópico de los granos de almidón recolectados en contextos arqueológicos mesoamericanos. Itinerarios, 29, 9–33.10.7311/ITINERARIOS.29.2019.01Search in Google Scholar

Cagnato, C. , Goepfert, N. , Elliott, M. , Verano, J. , Prieto, G. , & Dufour, E. (2021). Eat and die: The last meal of sacrificed Chimú Camelids at Huanchaquito–Las Llamas, Peru, as revealed by starch grain analysis. Latin American Antiquity, 32(3), 595–611. 10.1017/laq.2021.19.Search in Google Scholar

Cagnato, C. , Hamon, C. , & Salavert A. (in press). Starch grain analysis of Early Neolithic (Linearbandkeramik and Blicquy/Villeneuve-Saint-Germain) contexts: Experimental grinding tests of cereals and legumes. Submitted to Access Archaeology. Search in Google Scholar

Cagnato, C. , Hamon, C. , Salavert, A. , & Elliott, M. (in prep). The use of underground storage organs in the Early Neolithic (Linearbandkeramik and Blicquy/Villeneuve-Saint-Germain) in the Paris Basin: the contribution of starch grain analyses. To be submitted to Revue d’ethnoécologie.Search in Google Scholar

Cagnato, C. , & Ponce, J. M. (2017). Ancient Maya manioc (Manihot esculenta Crantz) consumption: Starch grain evidence from late to terminal classic (8th–9th century CE) occupation at La Corona, northwestern Petén, Guatemala. Journal of Archaeological Science: Reports, 16, 276–286.10.1016/j.jasrep.2017.09.035Search in Google Scholar

Chantran, A. , & Cagnato, C. (2021). Boiled, fried, or roasted? Determining culinary practices in Medieval France through multidisciplinary experimental approaches. Journal of Archaeological Science: Reports, 35. in Google Scholar

Chevalier, A. , & Bosquet, D. (2013). Culture matérielle, exploitation du territoire et identités socio-culturelle rubanées en Belgique: Analyses de microfossiles sur des instruments de mouture. In P. Anderson , C. Cheval , & A. Durand (Eds.), An interdisciplinary focus on plant-working tools (pp. 189–204). Antibes, France: ADPCA.Search in Google Scholar

Chevalier, A. , & Bosquet, D. (2017). Integrating archaeological data toward a better understanding of food plants choices and territory exploitation in the Northwestern European Early Neolithic: The Case of Remicourt “En Bia Flo II”. In M. P. Sayre & M. C. Bruno (Eds.), Social perspectives on ancient lives from paleoethnobotanical data (pp. 15–54). Switzerland: Springer.10.1007/978-3-319-52849-6_2Search in Google Scholar

Colledge, S. , & Conolly, J. (2014). Wild plant use in European Neolithic subsistence economies: A formal assessment of preservation bias in archaeobotanical assemblages and the implications for understanding changes in plant diet breadth. Quaternary Science Reviews, 101, 193–206.10.1016/j.quascirev.2014.07.013Search in Google Scholar

Copeland, L. , & Hardy, K. (2018). Archaeological starch. Agronomy, 8(1), 4.10.3390/agronomy8010004Search in Google Scholar

Cortella, A. R. , & Pochettino, M. L. (1994). Starch grain analysis as a microscopic diagnostic feature in the identification of plant material. Economic Botany, 48(2), 171.10.1007/BF02908212Search in Google Scholar

Crowther, A. (2009). Morphometric analysis of calcium oxalate raphides and assessment of their taxonomic value for archaeological microfossil studies. In M. Haslam , G. Robertson , A. Crowther , S. Nugent , & L. Kirkwood (Eds.), Archaeological science under a microscope: Studies in residue and ancient DNA analysis in Honour of Thomas H. Loy (pp. 102–128). Canberra, Australia: ANU E Press.10.22459/TA30.07.2009.08Search in Google Scholar

Crowther, A. (2012). The differential survival of native starch during cooking and implications for archaeological analyses: A review. Archaeological and Anthropological Sciences, 4(3), 221–235.10.1007/s12520-012-0097-0Search in Google Scholar

Delhon, C. , Binder, D. , Verdin, P. , & Mazuy, A. (2020). Phytoliths as a seasonality indicator? The example of the Neolithic site of Pendimoun, south-eastern France. Vegetation History and Archaeobotany, 29(2), 229–240.10.1007/s00334-019-00739-0Search in Google Scholar

Denham, T. , Haberle, S. , & Lentfer, C. J. (2004). New evidence and revised interpretations of early agriculture in Highland New Guinea. Antiquity, 78(302), 839. 10.1017/S0003598X00113481Search in Google Scholar

Dickau, R. , Ranere, A. J. , & Cooke, R. G. (2007). Starch grain evidence for the preceramic dispersals of maize and root crops into tropical dry and humid forests of Panama. Proceedings of the National Academy of Sciences, 104(9), 3651–3656.10.1073/pnas.0611605104Search in Google Scholar

Dietsch, M. F. (1996). Gathered fruits and cultivated plants at Bercy (Paris), a Neolithic village in a fluvial context. Vegetation History and Archaeobotany, 5(1–2), 89–97.10.1007/BF00189438Search in Google Scholar

Dietsch, M. F. (1997). Milieux humides pré-et protohistoriques dans le bassin parisien: L'étude des diaspores. (Doctoral dissertation). Université Paris-X, France.Search in Google Scholar

Dietsch-Sellami, M. F. (2004). L’alternance céréales à grains vêtus, céréales à grains nus au Néolithique: Nouvelle données, premières hypothèses. In INTERNEO 5 Actes de la Journée d’information du 20 novembre 2004 (pp. 125–135). Paris, France: Société Préhistorique Française.Search in Google Scholar

Divišová, M. , & Šída, P. (2015). Plant use in the Mesolithic period. Archaeobotanical data from the Czech Republic in a European context–A review. Interdisciplinaria Archaeologica: Natural Sciences in Archaeology, 6(1), 95–106.10.24916/iansa.2015.1.7Search in Google Scholar

Dommelier, S. , Bentrad, S. , Paicheler, J. C. , Pétrequin, P. , & Bouchet, F. (1998). Parasitoses liées à l’alimentation chez les populations néolithiques du lac de Chalain (Jura, France). Anthropozoologica, 27, 41–49.Search in Google Scholar

Duncan, N. A. , Pearsall, D. M. , & Benfer, R. A. (2009). Gourd and squash artifacts yield starch grains of feasting foods from preceramic Peru. Proceedings of the National Academy of Sciences, 106(32), 13202–13206.10.1073/pnas.0903322106Search in Google Scholar

Fairweather, A. D. , & Ralston, I. B. (1993). The Neolithic timber hall at Balbridie, Grampian Region, Scotland: The building, the date, the plant macrofossils. Antiquity, 67(255), 313.10.1017/S0003598X00045373Search in Google Scholar

Fiorin, E. , Sáez, L. , & Malgosa, A. (2018). Ferns as healing plants in medieval Mallorca, Spain? Evidence from human dental calculus. International Journal of Osteoarchaeology, 29(1), 82–90.10.1002/oa.2718Search in Google Scholar

Fritz, G. , & Nesbitt, M. (2014). Laboratory analysis and identification of plant macroremains. In J. M. Marston , J. D’Alpoim Guedes , & C. Warinner , Method and theory in paleoethnobotany (pp. 115–146). Colorado: University Press.10.5876/9781607323167.c007Search in Google Scholar

Gott, B. , Barton, H. , Samuel, D. , & Torrence, R. (2006). Biology of starch. In R. Torrence & H. Barton (Eds.), Ancient starch research (pp. 35–45). Walnut Creek, California: Left Coast Press.Search in Google Scholar

Hamon, C. (2008). Functional analysis of stone grinding and polishing tools from the earliest Neolithic of north-western Europe. Journal of Archaeological Science, 35(6), 1502–1520.10.1016/j.jas.2007.10.017Search in Google Scholar

Hamon, C. , Cagnato C. , Barbier-Emery, A. , & Salavert, A. (2021). European first farmers food practices: Combined use-wear and microbotanical approaches of LBK and BVSG grinding tools from the Paris Basin. Journal of Archaeological Science: Reports, 36. in Google Scholar

Hamon, C. , Salavert, A. , Dietsch-Sellami, M. F. , & Monchablon, C. (2019). Cultiver et consommer les plantes au Néolithique entre Seine et Meuse: Technologie des meules et analyses carpologiques. In F. Bostyn , C. Hamon , A. Salavert , & F. Giligny (Eds.), L’exploitation du milieu au Néolithique dans le quart nord-ouest de l’Europe: Contraintes environnementales, identités techniques et choix culturels, Session 4 des Actes du XXVIIIe Congrès préhistorique de France (pp. 119–137). Paris, France: Société Préhistorique française.Search in Google Scholar

Hardy, K. , Blakeney, T. , Copeland, L. , Kirkham, J. , Wrangham, R. , & Collins, M. (2009). Starch granules, dental calculus and new perspectives on ancient diet. Journal of Archaeological Science, 36(2), 248–255.10.1016/j.jas.2008.09.015Search in Google Scholar

Hardy, K. , Buckley, S. , & Copeland, L. (2018). Pleistocene dental calculus: Recovering information on Paleolithic food items, medicines, paleoenvironment and microbes. Evolutionary Anthropology: Issues, News, and Reviews, 27(5), 234–246.10.1002/evan.21718Search in Google Scholar

Hart, T. C. (2014). Analysis of starch grains produced in select taxa encountered in Southwest Asia. Ethnobiology Letters, 5, 135–145.10.14237/ebl.5.2014.251Search in Google Scholar

Haslam, M. (2004). The decomposition of starch grains in soils: Implications for archaeological residue analyses. Journal of Archaeological Science, 31(12), 1715–1734.10.1016/j.jas.2004.05.006Search in Google Scholar

Hather, J. G. , & Hammond, N. (1994). Ancient Maya subsistence diversity: Root and tuber remains from Cuello, Belize. Antiquity, 68(259), 330–335.10.1017/S0003598X00046639Search in Google Scholar

Hayes, E. H. , Cnuts, D. , Lepers, C. , & Rots, V. (2017). Learning from blind tests: Determining the function of experimental grinding stones through use-wear and residue analysis. Journal of Archaeological Science: Reports, 11, 245–260.10.1016/j.jasrep.2016.12.001Search in Google Scholar

Heiss, A. G. , Antolín, F. , Bleicher, N. , Harb, C. , Jacomet, S. , Kühn, M. , & Valamoti, S. M. (2017). State of the (t) art. Analytical approaches in the investigation of components and production traits of archaeological bread-like objects, applied to two finds from the Neolithic lakeshore settlement Parkhaus Opéra (Zürich, Switzerland). PLoS One, 12(8), e0182401.10.1371/journal.pone.0182401Search in Google Scholar

Henry, A. G. , Brooks, A. S. , & Piperno, D. R. (2011). Microfossils in calculus demonstrate consumption of plants and cooked foods in Neanderthal diets (Shanidar III, Iraq; Spy I and II, Belgium). Proceedings of the National Academy of Sciences, 108(2), 486–491.10.1073/pnas.1016868108Search in Google Scholar

Henry, A. G. , Hudson, H. F. , & Piperno, D. R. (2009). Changes in starch grain morphologies from cooking. Journal of Archaeological Science, 36(3), 915–922.10.1016/j.jas.2008.11.008Search in Google Scholar

Horrocks, M. , Bulmer, S. , & Gardner, R. O. (2008). Plant microfossils in prehistoric archaeological deposits from Yuku rock shelter, Western Highlands, Papua New Guinea. Journal of Archaeological Science, 35(2), 290–301.10.1016/j.jas.2007.03.005Search in Google Scholar

Horrocks, M. , Irwin, G. , Jones, M. , & Sutton, D. (2004). Starch grains and xylem cells of sweet potato (Ipomoea batatas) and bracken (Pteridium esculentum) in archaeological deposits from northern New Zealand. Journal of Archaeological Science, 31(3), 251–258.10.1016/S0305-4403(03)00018-9Search in Google Scholar

Inomata, T. , Triadan, D. , López, V. A. V. , Fernandez-Diaz, J. C. , Omori, T. , Bauer, M. B. M. , … & Nasu, H. (2020). Monumental architecture at Aguada Fénix and the rise of Maya civilization. Nature, 582(7813), 530–533.10.1038/s41586-020-2343-4Search in Google Scholar

Iriarte, J. , Holst, I. , Marozzi, O. , Listopad, C. , Alonso, E. , Rinderknecht, A. , & Montaña, J. (2004). Evidence for cultivar adoption and emerging complexity during the mid-Holocene in the La Plata basin. Nature, 432(7017), 614–617.10.1038/nature02983Search in Google Scholar

Jadin, I. , & Heim, J. (2003). Sur la voie de l’orge et du pavot: Macrorestes végétaux et agriculture rubanée du Haut Geer dans un cadre européen. Trois petits tours et puis s’ en vont… La fin de la présence danubienne en Moyenne-Belgique. ERAUL, 109, 345–392.Search in Google Scholar

Jane, J. , Kasemsuwan, T. , Leas, S. , Zobel, H. , & Robyt, J. F. (1994). Anthology of starch granule morphology by scanning electron microscopy. Starch‐Stärke, 46(4), 121–129 . 10.1002/star.19940460402Search in Google Scholar

Juhola, T. , Etu-Sihvola, H. S. K. , Näreoja, T. , & Ruohonen, J. (2014). Starch analysis reveals starchy foods and food processing from Finnish archaeological artefacts. Fennoscandia Archaeologica, XXXI, 79–100.Search in Google Scholar

Kirleis, W. , Klooß, S. , Kroll, H. , & Müller, J. (2012). Crop growing and gathering in the northern German Neolithic: A review supplemented by new results. Vegetation History and Archaeobotany, 21(3), 221–242.10.1007/s00334-011-0328-9Search in Google Scholar

Klooss, S. , Fischer, E. , Out, W. , & Kirleis, W. (2016). Charred root tubers of lesser celandine (Ficaria verna HUDS.) in plant macro remain assemblages from Northern, Central and Western Europe. Quaternary International, 404, 25–42.10.1016/j.quaint.2015.10.014Search in Google Scholar

Kubiak-Martens, L. (1999). The plant food component of the diet at the late Mesolithic (Ertebolle) settlement at Tybrind Vig, Denmark. Vegetation History and Archaeobotany, 8(1–2), 117–127.10.1007/BF02042850Search in Google Scholar

Kubiak-Martens, L. (2002). New evidence for the use of root foods in pre-agrarian subsistence recovered from the late Mesolithic site at Halsskov, Denmark. Vegetation History and Archaeobotany, 11(1–2), 23–32.10.1007/s003340200003Search in Google Scholar

Kubiak-Martens, L. (2008). Voedseleconomie: Parenchym en andere plantaardige macroresten (Hanzelijn Oude Land – Knooppunt Hattemerbroek). BIAXiaal, 386, 1–23.Search in Google Scholar

Kubiak-Martens, L. (2016). Scanning electron microscopy and starchy food in Mesolithic Europe: The importance of roots and tubers in Mesolithic diet. In K. Hardy & L. Kubiak-Martens (Eds), Wild Harvest. Plants in the Hominin and pre-Agrarian Human Worlds (pp. 113–134). Oxford and Philadelphia: Oxbow Books.10.2307/j.ctvh1dmjj.12Search in Google Scholar

Kubiak-Martens, L. , Brinkkemper, O. , & Oudemans, T. F. (2015). What’s for dinner? Processed food in the coastal area of the northern Netherlands in the Late Neolithic. Vegetation History and Archaeobotany, 24(1), 47–62.10.1007/s00334-014-0485-8Search in Google Scholar

Larsson, M. (2013). Cultivation and processing of Linum usitatissimum and Camelina sativa in southern Scandinavia during the Roman Iron Age. Vegetation History and Archaeobotany, 22(6), 509–520.10.1007/s00334-013-0413-3Search in Google Scholar

Lentfer, C. J. (2009). Building a comparative starch reference collection for Indonesia and its application to palaeoenvironmental and archaeological research. In M. Haslam , G. Robertson , A. Crowther , S. Nugent , & L. Kirkwood (Eds.), Archaeological science under a microscope: Studies in residue and ancient DNA analysis in Honour of Thomas H. Loy (pp. 80–101). Canberra, Australia: ANU E Press.10.22459/TA30.07.2009.07Search in Google Scholar

Li, W. , Pagán‐Jiménez, J. R. , Tsoraki, C. , Yao, L. , & Van Gijn, A. (2020). Influence of grinding on the preservation of starch grains from rice. Archaeometry, 62(1), 157–171.10.1111/arcm.12510Search in Google Scholar

Liu, L. , Duncan, N. A. , Chen, X. , Liu, G. , & Zhao, H. (2015). Plant domestication, cultivation, and foraging by the first farmers in early Neolithic Northeast China: Evidence from microbotanical remains. The Holocene, 25(12), 1965–1978.10.1177/0959683615596830Search in Google Scholar

Liu, L. , Field, J. , Fullagar, R. , Bestel, S. , Chen, X. , & Ma, X. (2010). What did grinding stones grind? New light on early Neolithic subsistence economy in the Middle Yellow River valley, China. Antiquity, 84(325), 816–833. 10.1017/S0003598X00100249.Search in Google Scholar

Liu, L. , Kealhofer, L. , Chen, X. , & Ji, P. (2014). A broad-spectrum subsistence economy in Neolithic Inner Mongolia, China: Evidence from grinding stones. The Holocene, 24(6), 726–742.10.1177/0959683614526938Search in Google Scholar

Liu, L. , Wang, J. , & Levin, M. J. (2017). Usewear and residue analyses of experimental harvesting stone tools for archaeological research. Journal of Archaeological Science: Reports, 14, 439–453.10.1016/j.jasrep.2017.06.018Search in Google Scholar

Lucarini, G. , Radini, A. , Barton, H. , & Barker, G. (2016). The exploitation of wild plants in Neolithic North Africa. Use-wear and residue analysis on non-knapped stone tools from the Haua Fteah cave, Cyrenaica, Libya. Quaternary International, 410, 77–92.10.1016/j.quaint.2015.11.109Search in Google Scholar

Madella, M. , García-Granero, J. J. , Out, W. A. , Ryan, P. , & Usai, D. (2014). Microbotanical evidence of domestic cereals in Africa 7000 years ago. PLoS One, 9(10), e110177.10.1371/journal.pone.0110177Search in Google Scholar

Marinval, P. (1988). L’alimentation végétale en France du Mésolithique jusqu’à l’Age du Fer. Toulouse, France: CNRS.Search in Google Scholar

Mattalia, G. , Quave, C. L. , & Pieroni, A. (2013). Traditional uses of wild food and medicinal plants among Brigasc, Kyé, and Provençal communities on the Western Italian Alps. Genetic Resources and Crop Evolution, 60(2), 587–603.10.1007/s10722-012-9859-xSearch in Google Scholar

Mercader, J. (2009). Mozambican grass seed consumption during the Middle Stone Age. Science, 326(5960), 1680–1683.10.1126/science.1173966Search in Google Scholar

Mercader, J. , Abtosway, M. , Bird, R. , Bundala, M. , Clarke, S. , Favreau, J. , … Walde, D. (2018b). Morphometrics of starch granules from sub-Saharan plants and the taxonomic identification of ancient starch. Frontiers in Earth Science, 6, 146.10.3389/feart.2018.00146Search in Google Scholar

Mercader, J. , Akeju, T. , Brown, M. , Bundala, M. , Collins, M. J. , Copeland, L. , … Xhauflair, H. (2018a). Exaggerated expectations in ancient starch research and the need for new taphonomic and authenticity criteria. Facets, 3(1), 777–798.10.1139/facets-2017-0126Search in Google Scholar

Messner, T. C. (2011). Acorns and Bitter roots: Starch grain research in the prehistoric Eastern woodlands. Tuscaloosa: University of Alabama Press.Search in Google Scholar

Musaubach, M. G. , Plos, A. , & Babot, M. D. P. (2013). Differentiation of archaeological maize (Zea mays L.) from native wild grasses based on starch grain morphology. Cases from the Central Pampas of Argentina. Journal of Archaeological Science, 40(2), 1186–1193.10.1016/j.jas.2012.09.026Search in Google Scholar

Nadel, D. , Piperno, D. R. , Holst, I. , Snir, A. , & Weiss, E. (2012). New evidence for the processing of wild cereal grains at Ohalo II, a 23,000-year-old campsite on the shore of the Sea of Galilee, Israel. Antiquity, 86(334), 990–1003.10.1017/S0003598X00048201Search in Google Scholar

Out, W. A. , Ryan, P. , García-Granero, J. J. , Barastegui, J. , Maritan, L. , Madella, M. , & Usai, D. (2016). Plant exploitation in Neolithic Sudan: A review in the light of new data from the cemeteries R12 and Ghaba. Quaternary International, 412, 36–53.10.1016/j.quaint.2015.12.066Search in Google Scholar

Pagán Jiménez, J. R. , Guachamin-Tello, A. M. , Romero-Bastidas, M. E. , & Vásquez-Ponce, P. X. (2017). Cocción experimental de tortillas de casabe (Manihot esculenta Crantz) y de camote (Ipomoea batatas [L.] Lam.) en planchas de barro: Evaluando sus efectos en la morfometría de los almidones desde una perspectiva paleoetnobotánica. Americae, 2, 27–43.Search in Google Scholar

Pearsall, D. M. (2000). Paleoethnobotany: A handbook of procedures. Walnut Creek: Left Coast Press.Search in Google Scholar

Pearsall, D. M. , Chandler-Ezell, K. , & Zeidler, J. A. (2004). Maize in ancient Ecuador: Results of residue analysis of stone tools from the Real Alto site. Journal of Archaeological Science, 31(4), 423–442.10.1016/j.jas.2003.09.010Search in Google Scholar

Perry, D. (1999). Vegetative tissues from Mesolithic sites in the northern Netherlands. Current Anthropology, 40(2), 231–237.10.1086/200008Search in Google Scholar

Pető, Á. , Gyulai, F. , Pópity, D. , & Kenéz, Á. (2013). Macro-and micro-archaeobotanical study of a vessel content from a Late Neolithic structured deposition from southeastern Hungary. Journal of Archaeological Science, 40(1), 58–71.10.1016/j.jas.2012.08.027Search in Google Scholar

Pétrequin, A. M. , & Pétrequin, P. (1988). Le Néolithique des lacs: Préhistoire des lacs de Chalain et de Clairvaux (4000–2000 av. J.-C.). France: Éd. Errance.Search in Google Scholar

Piperno, D. R. , & Holst, I. (1998). The presence of starch grains on prehistoric stone tools from the humid neotropics: Indications of early tuber use and agriculture in Panama. Journal of Archaeological Science, 25(8), 765–776.10.1006/jasc.1997.0258Search in Google Scholar

Piperno, D. R. , Ranere, A. J. , Holst, I. , & Hansell, P. (2000). Starch grains reveal early root crop horticulture in the Panamanian tropical forest. Nature, 407(6806), 894–897.10.1038/35038055Search in Google Scholar

Piperno, D. R. , Weiss, E. , Holst, I. , & Nadel, D. (2004). Processing of wild cereal grains in the Upper Palaeolithic revealed by starch grain analysis. Nature, 430(7000), 670–673.10.1038/nature02734Search in Google Scholar

Raemaekers, D. C. , Kubiak-Martens, L. , & Oudemans , T. F. (2013). New food in old pots–charred organic residues in Early Neolithic ceramic vessels from Swifterbant, the Netherlands (4300–4000 cal BC). Archäologisches Korrespondenzblatt, 43(3), 315–334.Search in Google Scholar

Ranwala, A. P. , & Miller, W. B. (2008). Analysis of nonstructural carbohydrates in storage organs of 30 ornamental geophytes by high‐performance anion‐exchange chromatography with pulsed amperometric detection. New Phytologist, 180(2), 421–433.10.1111/j.1469-8137.2008.02585.xSearch in Google Scholar

Revedin, A. , Aranguren, B. , Becattini, R. , Longo, L. , Marconi, E. , Lippi, M. M. , & Svoboda, J. (2010). Thirty thousand-year-old evidence of plant food processing. Proceedings of the National Academy of Sciences, 107(44), 18815–18819.10.1073/pnas.1006993107Search in Google Scholar

Salavert, A. (2010). Le pavot (Papaver somniferum) à la fin du 6e millénaire av. J.-C. en Europe occidentale. Anthropobotanica, 1(3), 3–16.Search in Google Scholar

Salavert, A. (2011). Plant economy of the first farmers of central Belgium (Linearbandkeramik, 5200–5000 BC). Vegetation History and Archaeobotany, 20(5), 321–332.10.1007/s00334-011-0297-zSearch in Google Scholar

Salavert, A. (2017). Agricultural dispersals in Mediterranean and temperate Europe. In Oxford research encyclopedia of environmental science. in Google Scholar

Sandgathe, D. M. , & Hayden, B. (2003). Did Neanderthals eat inner bark? Antiquity, 77(298), 709–718.10.1017/S0003598X00061652Search in Google Scholar

Saqalli, M. , Salavert, A. , Bréhard, S. , Bendrey, R. , Vigne, J. D. , & Tresset, A. (2014). Revisiting and modelling the woodland farming system of the early Neolithic Linear Pottery Culture (LBK), 5600–4900 BC. Vegetation History and Archaeobotany, 23(1), 37–50.10.1007/s00334-014-0436-4Search in Google Scholar

Saul, H. , Wilson, J. , Heron, C. P. , Glykou, A. , Hartz, S. , & Craig, O. E. (2012). A systematic approach to the recovery and identification of starches from carbonised deposits on ceramic vessels. Journal of Archaeological Science, 39(12), 3483–3492.10.1016/j.jas.2012.05.033Search in Google Scholar

Scheel-Ybert, R. (2001). Man and vegetation in southeastern Brazil during the late Holocene. Journal of Archaeological Science, 28(5), 471–480.10.1006/jasc.2000.0577Search in Google Scholar

Scott Cummings, L. , Yost, C. , & Sołtysiak, A. (2018). Plant microfossils in human dental calculus from Nemrik 9, a Pre-Pottery Neolithic site in Northern Iraq. Archaeological and Anthropological Sciences, 10(4), 883–891.10.1007/s12520-016-0411-3Search in Google Scholar

Shibutani, A. (2017), What did Jomon people consume for starchy food? A review of the current studies on archaeological starch grains in Japan. Japanese Journal of Archaeology, 5, 3–25.Search in Google Scholar

Simkova, K. , & Polesny, Z. (2015). Ethnobotanical review of wild edible plants used in the Czech Republic. Journal of Applied Botany and Food Quality, 88(1), 49–67.Search in Google Scholar

Tardío, J. , Pardo-de-Santayana, M. , & Morales, R. (2006). Ethnobotanical review of wild edible plants in Spain. Botanical Journal of the Linnean Society, 152(1), 27–71.10.1111/j.1095-8339.2006.00549.xSearch in Google Scholar

Tolar, T. , Jacomet, S. , Velušček, A. , & Čufar, K. (2011). Plant economy at a late Neolithic lake dwelling site in Slovenia at the time of the Alpine Iceman. Vegetation History and Archaeobotany, 20(3), 207–222.10.1007/s00334-010-0280-0Search in Google Scholar

Toulemonde, F. (2010). Camelina Sativa: L’or végétal du Bronze et du Fer. Anthropobotanica, 1, 3–14.Search in Google Scholar

Wan, Z. , Yang, X. , Ge, Q. , Fan, C. , Zhou, G. , & Jiang, M. (2012). Starch grain analysis reveals Late Neolithic plant utilization in the middle reaches of the Ganjiang River. Science China Earth Sciences, 55(12), 2084–2090.10.1007/s11430-012-4512-2Search in Google Scholar

Wang, J. , Liu, L. , Ball, T. , Yu, L. , Li, Y. , & Xing, F. (2016). Revealing a 5,000-y-old beer recipe in China. Proceedings of the National Academy of Sciences, 113(23), 6444–6448.10.1073/pnas.1601465113Search in Google Scholar

Wang, J. , Liu, L. , Georgescu, A. , Le, V. V. , Ota, M. H. , Tang, S. , & Vanderbilt, M. (2017). Identifying ancient beer brewing through starch analysis: A methodology. Journal of Archaeological Science: Reports, 15, 150–160.10.1016/j.jasrep.2017.07.016Search in Google Scholar

Yang, X. , & Perry, L. (2013). Identification of ancient starch grains from the tribe Triticeae in the North China Plain. Journal of Archaeological Science, 40(8), 3170–3177.10.1016/j.jas.2013.04.004Search in Google Scholar

Yang, X. , Barton, H. J. , Wan, Z. , Li, Q. , Ma, Z. , Li, M. , … Wei, J. (2013). Sago-type palms were an important plant food prior to rice in southern subtropical China. PLoS One, 8(5), e63148.10.1371/journal.pone.0063148Search in Google Scholar

Yang, X. , Fuller, D. Q. , Huan, X. , Perry, L. , Li, Q. , Li, Z. , … Lu, H. (2015). Barnyard grasses were processed with rice around 10000 years ago. Scientific Reports, 5, 16251.10.1038/srep16251Search in Google Scholar

Yang, X. , Ma, Z. , Li, Q. , Perry, L. , Huan, X. , Wan, Z. , … Zheng, J. (2014). Experiments with lithic tools: Understanding starch residues from crop harvesting. Archaeometry, 56(5), 828–840.10.1111/arcm.12034Search in Google Scholar

Yao, L. , Yang, Y. , Sun, Y. , Cui, Q. , Zhang, J. , & Wang, H. (2016). Early Neolithic human exploitation and processing of plant foods in the Lower Yangtze River, China. Quaternary International, 426, 56–64.10.1016/j.quaint.2016.03.009Search in Google Scholar

Zarrillo, S. , & Kooyman, B. (2006). Evidence for berry and maize processing on the Canadian plains from starch grain analysis. American Antiquity, 71, 473–499.10.1017/S0002731600039779Search in Google Scholar

Zohary, D. , Hopf, M. , & Weiss, E. (2012). Domestication of Plants in the Old World: The origin and spread of domesticated plants in Southwest Asia, Europe, and the Mediterranean Basin. Oxford, UK: Oxford University Press.10.1093/acprof:osobl/9780199549061.001.0001Search in Google Scholar

Received: 2020-10-31
Revised: 2021-05-24
Accepted: 2021-06-20
Published Online: 2021-09-14

© 2021 Clarissa Cagnato et al., published by De Gruyter

This work is licensed under the Creative Commons Attribution 4.0 International License.

Downloaded on 22.2.2024 from
Scroll to top button