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date: 22 August 2017

Agricultural Dispersals in Mediterranean and Temperate Europe

Summary and Keywords

Along with ceramics production, sedentism, and herding, agriculture is a major component of the Neolithic as it is defined in Europe. Therefore, the agricultural system of the first Neolithic societies and the dispersal of exogenous cultivated plants to Europe are the subject of many scientific studies. To work on these issues, archaeobotanists rely on residual plant remains—crop seeds, weeds, and wild plants—from archaeological structures like detritic pits, and, less often, storage contexts. To date, no plant with an economic value has been identified as domesticated in Western Europe except possibly opium poppy. The earliest seeds identified at archaeological sites dated to about 5500–5200 bc in the Mediterranean and Temperate Europe. The cultivated plants identified were cereals (wheat and barley), oleaginous plant (flax), and pulses (peas, lentils, and chickpeas). This crop package originated in the Fertile Crescent, where it was clearly established around 7500 bc (final Pre-Pottery Neolithic B), after a long, polycentric domestication process. From the middle of the 7th millennium bc, via the Balkan Peninsula, the pioneer Neolithic populations, with their specific economies, rapidly dispersed from east to west, following two main pathways. One was the maritime route over the northwestern basin of the Mediterranean (6200–5300 bc), and the other was the terrestrial and fluvial route in central and northwestern continental Europe (5500–4900 bc). On their trajectory, the agropastoral societies adapted the Neolithic founder crops from the Middle East to new environmental conditions encountered in Western Europe.

The Neolithic pioneers settled in an area that had experienced a long tradition of hunting and gathering. The Neolithization of Europe followed a colonization model. The Mesolithic groups, although exploiting plant resources such as hazelnut more or less intensively, did not significantly change the landscape. The impact of their settlements and their activities are hardly noticeable through palynology, for example. The control of the mode of reproduction of plants has certainly increased the prevalence of Homo sapiens, involving, among others, a demographic increase and the ability to settle down in areas that were not well adapted to year-round occupation up to that point. The characterization of past agricultural systems, such as crop plants, technical processes, and the impact of anthropogenic activities on the landscape, is essential for understanding the interrelation of human societies and the plant environment. This interrelation has undoubtedly changed deeply with the Neolithic Revolution.

Keywords: archaeology, Neolithic, Palaeoethnobotany, ancient cereals, opium poppy, early farming systems


Agriculture is the relationship between an exploiting species and one or more exploited species living in an artificial cultivated ecosystem (Mazoyer & Roudart, 1997). In this sense, humans are not the only ones practicing agriculture. For exemple, Atta and Acromyrmex, two ant genera from the Americas, cultivate fungi that will constitute the essence of their alimentation. However, humans have invented many techniques and artifacts to cultivate a great number of crops. There is currently a wide variety of agricultures in the world, and even now, in the 21st century, agriculture is at the base of most human economies. But what about the first agrarian systems? When did humans become farmers? How did cultivated plants and farming techniques come to Temperate and Mediterranean Europe? Research on the earliest farming systems is highly significant, especially in a globalized context, in which new modes of production have been emerging since the 1980s to handle and patent the gene pool of cultivated plants. In Europe, the privatization of agricultural heritage, as well as experimentation to diffuse agriculture beyond Earth, is coming face to face with a renewal of old varieties and food production on a local scale. It shows that agricultural innovations are complex and not linear, even today (Bonneuil et al., 2006).

In the absence of iconographic and written sources for the ancient period, archaeological materials are the most direct evidence for the early agricultures that appeared in several places across the world from around 11,500 years to 5,000 years ago. Agriculture is one of the main components of Neolithic economy, associated with animal husbandry, potterymaking, and sedentary habitats in Western Europe (Price & Bar-Yosef, 2011). Often described as a revolution (Childe, 1925), the Neolithic period appears today like a transitional phenomenon in the Near East, as well as in the diffusion of its economy to Europe. Agriculture did not arrive suddenly. It took around 3,000 years for domesticates to spread from the Aegean to Great Britain and Ireland. But even if it was not a revolution, the invention of agriculture and its dispersal constituted a crucial change in human economy, as well as its relationship with the natural environment. Before becoming farmers and herders, human beings were predators, living lives of hunting and gathering, for a long time.

This geographical framework includes a territory stretching roughly from the Black Sea in the east to the Atlantic Ocean in the west. In Europe, the Neolithic period is part of the Holocene epoch, which began about 12,000 years ago. The Holocene is characterized by a global warming that followed the frosty and arid climate of the Pleistocene. The epoch is divided into several chronological zones, defined in particular by palynology (i.e., the study of pollen grains conserved in sediment). The beginning of the Neolithic corresponds with the middle of the Atlantic chronozone, characterized by the extension of deciduous oak to the detriment of the boreal forest, composed of birch, pine, and hazelnut, in Temperate Europe, as well as Scots pine and juniper in the Mediterranean. The Mesolithic/Neolithic transition corresponds to the passage from a hunting/gathering to an agropastoral way of life. The transition occurred at different times over three millennia, between 7000 and 4000 bc, according to the regions of Europe and their location on the long route of diffusion of agriculture. The primary Neolithic settlement regions in Europe are, roughly, the Balkans, central Europe, and the central and northwestern areas of the Mediterranean basin. This article aims to present the emergence of agriculture in Europe in the middle of the 7th millennium bc, demonstrating the main routes of cultivated plant diffusion to Temperate and Mediterranean Europe, as well as highlighting assumptions about early Neolithic farming systems (Fig. 1).

Agricultural Dispersals in Mediterranean and Temperate EuropeClick to view larger

Figure 1. Chronological setting of the first agriculture dispersals in Western Europe. The main archaeological cultures, defined mainly by ceramic decoration, are shown in italic.

Studying the Origin of the First Agrarian System and Its Introduction into Europe

How Is the Spread of Early Agriculture Studied?

Proxies Used to Study Early Agriculture

Interdisciplinarity is fundamental to the study of past societies, including early crop diversity and the diffusion of agriculture. Farming systems result from the combination of natural, sociocultural, economic, and technical elements (Jouve, 1988). Several types of residues, plant microfossils (phytoliths, pollen grains and starch, and organic residues), plant macrofossils (charcoal), artifacts (grinding stones and harvesting tools), and archaeozoology (bone pathology) allow the study of most of the components, especially the ecosystem context and technological practices. Synthesizing the results and integrating all the archaeological disciplines and scientific tools used, therefore, is an arduous task. Even then, this synthesis cannot be exhaustive due to the mass of data produced by archaeology (many proxies are used to study the agricultural system), the large territory in question (Europe), and the interest aroused by this huge socioeconomic transition in the scientific community (from which has come an abundant bibliography). This article concentrates on the first moments of Neolithic primary colonization in the Balkans, the Mediterranean, and Temperate Europe, relying on the most direct evidence of past agricultural practices – the plant macroremains recovered from archaeological sites.

Definition and Methodology of Archeobotany

Archaeobotany is the study of seeds, fruits, and inflorescent parts from archaeological sites. Plant macroremains are collected during excavations and mostly come from sedimentary samples. Water sieving allows the extraction of fragile plant remains from their sedimentary matrix. Most often, seeds and fruits are charred in Temperate and Mediterranean Europe. This can result from one or a few specific actions in time and space—as in a burned storage structure—but also disparate events, as in the progressive accumulation of remains resulting from cooking accidents (Bouby, 2000). Others macroremains, such as cereal processing byproducts or weeds, are not related to human food but rather are direct evidence of past farming practices. Furthermore, the study of wild fruits and seeds allows us to consider the importance of gathering practices in the Neolithic productive economy, even when it is assumed that domesticated taxa constitute the main vegetal element of the human diet.

Agricultural Dispersals in Mediterranean and Temperate EuropeClick to view larger

Figure 2. Archaeological sample following water sieving, with a mix of emmer, einkorn, and an important amount of Lapsana communis, a frequent Neolithic weed in Temperate Europe (Remicourt-en-Bia Flo II, around 5000 bc, Belgium).

After careful separation from other macroremains (microfauna, shells, lithics, and mineral residues), seeds, fruits, and parts of the inflorescence are identified using a binocular microscope, with magnifications from about 10 to 50 times (Fig. 2). The archaeobotanical process always begins with the morphological observation of plant remains to identify the taxa. For this purpose, the archaeological specimens, often charred and damaged, are compared with specimens from modern seed comparison collections, as well as morphological botanical descriptions. The cultivated plants and evidence of their mode of production and processing that reach us are fragmentary and most often of accidental origin (as is the case with many archaeological remains). Indeed, certain types of remains, such as leaves, roots, and tubers, are not clearly identified because of their uncharacteristic anatomy and low resistance to carbonization. In addition, different sampling protocols, an unequal analytical corpus, and nonuniform, macro-level counting methods across the discipline make it difficult to integrate data from different researchers (Fig. 3). Thus, limiting filters are to be taken into account when the archaeobotanist seeks to interpret archeobotanical assemblages and create a comparative work between two regions, for example.

Agricultural Dispersals in Mediterranean and Temperate EuropeClick to view larger

Figure 3. Presentation of the main filters that come into archaeobotanical studies, as well as tools used by the archaeobotanist to reconstruct past farming systems.

The first archaeobotanical studies were conducted in Egypt on desiccated seeds from pharaonic tombs, which were studied by Kunth (1826). Results from the Swiss pile-dwelling sites and the Hallstatt salt mines in Austria were published 40 years later (Heer, 1865; Unger, Lesquereux, & Hruschauer, 1851). In 1968, the creation of the International Work Group for Palaeoethnobotany (IWGP) allowed for a united discipline, organizing an international symposium every three years in Europe, in which the study of the first farming system in the world takes an important place. The domestication of plant species and their introduction into new regions are, indeed, among the favorite subjects of archaeobotanists. From the 1980s, an increase in archaeological research has enabled the acquisition of archaeobotanical material. The reference work regarding crop domestication, reprinted several times, discusses the origin of cultivated plants and their distribution in the western and central regions of the Asiatic continent (Zohary, Hopf, & Weiss, 2012). In addition, archeobotanical syntheses and comparative works at the regional and supraregional scale in Europe have appeared in several collective works and articles (e.g., Bogaard & Jones, 2007; Chevalier, Marinova, & Peña-Chocarro, 2014; Colledge & Conolly, 2007a; Coward, Shennan, Colledge, Conolly, & Collard, 2008).

Contribution of Cultivated Plants to Understanding Early Crop Diversity

Cultivated plants (cereals, pulses, and oleaginous/textile materials) are represented by their seeds, as well as, for cereals, their envelopes (lemma and palea) and the rachis that produces the chaff. The chaff corresponds to byproducts from the processing of cereals and allows for the study of postharvesting stages, such as the way that cereals were stored (cleaned seeds, spikelets, or ears). Rachis also provides one of the best morphological traits visible through archaeobotany for discriminating wild and cultivated cereals. In a cultivated cereal, the rachis has robust scars, indicating that the ears are indehiscent (i.e., they cannot disperse by themselves, unlike wild cereals). For the same reason, and because of the cereal processing, the rachis base is often broken. Seeds or chaff imprints, used in building materials or tempered ceramic, can also provide information about the crops and the use of grain products. Seeds and chaff provide information on crop morphological diversity in the earliest stages of Neolithic dispersal.

For cultivated plants, the rank of the species, in the morphological sense of the term, is most often identifiable. Sometimes only the genus (e.g., Triticum/Hordeum) can be identified due to poor preservation of remains. Sometimes species cannot be discriminated because of similar morphological criteria (e.g., Triticum aestivum/durum), preventing the appreciation of the whole diversity of past cultivated plants.

Agricultural Dispersals in Mediterranean and Temperate EuropeClick to view larger

Figure 4. Seeds of Neolithic cultivated plants from Early Neolithic sites in Belgium. (a) T. monococcum (einkorn); (b) T. dicoccum (emmer); (c) P. sativum (pea); (d) L. culinaris (lens); (e) L. usitatissimum (flax).

In addition to the strictly morphological study, genetic research applied to charred plant macrofossils has been in development since the 2000s (Brown et al., 2015; Schlumbaum et al., 2008). This will provide substantial information on the origin of ancient plant remains, the process of domestication, and the genetic diversity within a field, which is not visible through the seeds or cereal rachis morphology.

Contribution of Weeds to Study the Early Farming System

Weed seeds are also conserved, even in charred conditions. Weeds, from the archaeobotanical point of view, correspond to uncultivated plants installed in the fields. They result from the cleaning of cereals or pulses and are direct indicators of the exploited agricultural environments and the agricultural techniques used. In a cultivated plot, the flora is the result of a combination of agronomic factors (e.g., the depth of plowing and the seasonality of seedlings) and the environment (e.g., precipitation, temperature, and soil properties) that interact in complex ways. Archaeological weed assemblages, therefore, are the products of past farming systems.

The identification of weeds within the rank of species is more problematic then for cultivated plants, as it depends on the preservation of the seeds, the experience of the archaeobotanists, and the richness of the reference collections of current plants. Indeed, the diversity of wild plants is much wider than cultivated plants. Moreover, botanical identification depends on several anatomical criteria (e.g., size, color of flowers, number of petals, and arrangement of leaves). However, most often, only the seed is conserved in an archaeological context.

Interpretation of weed assemblages is based on the current flora ecology and on farming experiments (Bogaard, 2002; Jones, 2005; Jones, Charles, Colledge, & Halstead, 1995). When the weed identifications are precise enough (to the specie level), the phytosociology of these plants are indicators of the mode of cultivation. Archaeological weeds can be classified according to their current cooccurrences in fields, in large classes such as Secalinetea (corresponding to winter crops) and Chenopodietea (corresponding to summer crops). However, both classes may cohabit in the same plot, and the weed flora would be influenced by how the land is worked rather than the sowing season (Behre & Jacomet, 1991; Lundström-Baudais, 1986). For example, Chenopodium album, which is regularly identified in Neolithic archaeobotanical assemblages, is found in both summer and winter crops (Bogaard, Jones, Charles, & Hodgson, 2001). It is then much less developed and has a smaller size (Lundström-Baudais, 1986). The main limit to reconstructing the Neolithic farming system in general, including the intensity of crops, the seasonality, and the sustainability of the fields, is due to the current difficulty in finding (by ethnology) or reproducing (by experimentation) farming systems in an ecological and socioeconomical setting comparable to those of past societies.

Table 1. Main Crops and Weeds Mentioned in the Text


Latin Names

English Common Names

Wild and Cultivated Cereals

H. vulgare subsp. spontaneum (K. Koch) Thell.

Wild barley

H. vulgare subsp. vulgare L.

Cultivated hulled barley

H. vulgare subsp. nudum L.

Cultivated naked barley

T. aestivum L.

Bread wheat

T. monococcum L.


T. monococcum subsp. aegilopoides (Link) Thell.

Wild einkorn

T. turgidum L. ssp. dicoccoides (Körn. ex Asch. & Graebn.) Thell.

Wild emmer

T. turgidum L. subsp. dicoccon (Schrank) Thell.

Cultivated emmer

T. turgidum subsp. durum (Desf.) Husn.

Macaroni wheat

Wild and Cultivated Pulses

C. arietinum L.


L. cicera L.

Red pea

L. sativus L.

Grass pea

L. culinaris Medik.


P. sativum L.


V. ervilia (L.) Willd.

Bitter vetch

V. faba L.

Broad bean

V. sativa L.

Common vetch

Oil/Textile/Psychoactive Plants

L. usitatissimum L.


P. somniferum L.

Opium poppy

Papaver somniferum subsp. setigerum (DC.) Arcang.

Wild poppy


C. album L.


B. secalinus L.

Rye brome

B. tectorum L.

Drooping brome

F. convolvulus (L.) Á.Löve


G. aparine L.


L. communis Juss.


S. viridis (L.) P. Beauv.

Wild/green foxtail millet

S. verticillata (L.) P. Beauv.

Bristly foxtail

For the last 15 years, a method called Functional Interpretation of Botanical Survey (FIBS) has been used extensively to interpret weed assemblages. FIBS is based on the autoecology of weed species, including individual attributes such as length of growth period, canopy height, and ability to regenerate after significant soil disturbances (Bogaard, Hodgson, Wilson, & Band, 1998; Jones, Bogaard, Charles, & Hodgson, 2000). It makes it possible to go further in interpreting past agricultural regimes (intensive or extensive) and crops’ seasonality. The growth conditions of the cultivated plants (e.g., water supply and soil fertility) can also be specified thanks to the stable isotopes of carbon (δ‎13C) and nitrogen (δ‎15N) carried out on cereal seeds in particular (Fiorentino, Ferrio, Bogaard, Araus, & Riehl, 2015). These analyses, which will certainly develop exponentially in the future, constitute a promising tool complementary to the interpretations of archaeological assemblages of weeds to understand how plants were cultivated.

At the Origins of Agriculture in the Near East

Geographical Location of Wild Progenitors of Neolithic Crops

There is currently a consensus that crops found in European archaeological sites were not domesticated locally. Most of them come from the Near East. However, the question of domestication attempts of cereals and pulses before the arrival of Neolithic groups is still subject to debate in both Temperate and Mediterranean Europe.

Taxonomic classification, as well as cynogenetic and molecular affinities, make it possible to identify, with more or less precision, the wild ancestors of cultivated plants. However, not all of them have been identified with certainty, especially for pulses. For example, Triticum turgidum ssp. dicoccoideae is the wild progenitor of emmer (Triticum dicoccum), one of the first wheats domesticated in the Near East, whereas the wild ancestor of the faba bean (Vicia faba) is still unknown (Zohary et al., 2012). The presence of wild ancestor populations in one region, therefore, is the main criterion in favor of the local domestication of this plant in the Neolithic period. By the end of the 19th century, botanists such as de Candolle (1883), and later Vavilov (1951), worked on these issues, labeling the Fertile Crescent as one of the world’s centers of plant domestication.

Most putative wild progenitors are, thus, currently absent in Western Europe except for the Balkans. Triticum aegilopoides, the ancestor of einkorn (Triticum monococcum); Lens orientalis, the ancestor of lens (Lens culinaris); and Hordeum spontaneum, the ancestor of barley (Hordeum vulgare) are present in Southeast Europe today (Redden et al., 2015; Valamoti et al., 2007; Zohary et al., 2012). Thus, the hunter-gatherers of the Balkan Peninsula had at their disposal populations of the wild progenitors of cultivated cereals and pulses. In the Franchthi Cave, in the Peloponnese (southern Greece), populations of barley, lentils, and wild oats have been found to be exploited during the Mesolithic period, in archaeological layers dating to around 7900–7500 bc (Valamoti & Kotsakis, 2007). On the following Neolithic archaeological levels, cereals and pulses were identified on the morphological base, which could suggest a domestication process at that site. However, the absence of transitional domestication traits supports the hypothesis of the sudden introduction of cultivars in their fully domesticated morphology during the Initial Neolithic, in the first half of the 7th millennium bc (Perlès, 2001).

Wild pulses and cereals are identified in other Mesolithic layers in Mediterranean Europe. For example, in the L’Abeurador, a cave located in the Massif Central (southern France), numerous charred pulses and seeds of the genera Vicia, Lathyrus, and Pisum have been identified in Mesolithic levels dating to around 9000 bc (Vaquer & Ruas, 2009). These three genera are part of the founder crop package of the Near East. Their identification rank remains imprecise due to poor conservation of macroremains. We do not know whether these Mesolithic wild pulses correspond to wild progenitors. At L’Abeurador, wild pulses have been found along with hazelnut shells, dogwood seeds, and wild grapeseeds, which were probably consumed by inhabitants of the cave. Thus, as in Franchthi Cave, wild pulses also could have been intensively picked up by Mesolithic hunter-gatherers before the introduction of cultivated plants in the Mediterranean region by Neolithic farmers.

These examples demonstrate that exploitation of wild resources, among other wild cereals and pulses that were potentially ancestors of Neolithic crops, are common in Mesolithic sites, mainly the Mediterranean region. However, intensive wild plant gathering does not necessarily involve local domestication processes. Indeed, in Neolithic Europe, the archaeological seeds and elements of cereal chaff are domestic from the morphological point of view. There are no transitory forms, as may be suspected in the Near East (Tanno & Willcox, 2006).

Process of Domestication in the Near East

The domestication of a plant is a long process. First, the wild ancestor is collected, more or less intensively. Humans, in a way, cultivate a wild population before it depends entirely on them for reproduction. The focus of domestication at the origin of the cultivated plants identified on the European Neolithic sites (mainly cereals and pulses), is located in the Fertile Crescent—that is to say, the southeast of Anatolia, the Middle Euphrates, and the Southern Levant.

The process of domestication began in the Epipalaeolithic, about 12,000 years ago. Some authors have estimated that the climatic deterioration of the recent Dryas (12,500–11,500 bp), which corresponds to the end of the last glacial period and which results in a decrease in temperatures, would have played a stimulating role in the choice of cultivation of cereals to compensate for climatic risks. According to Willcox (2005), the models explaining the transition to agriculture place too much importance on this event. Indeed, although the deterioration is undeniable, the climatic conditions had only a moderate effect on vegetation. In addition, agriculture, which depends on a stable climate, was established after the Younger Dryas, at the end of the 9th millennium bc. The availability of wild cereals, ancestors of domestic cultivars, is one of the key factors in understanding the beginning of agriculture. This availability depends on soil types, precipitation, and temperatures. In the 1990s, it was thought that there was only one center of domestication, followed by a dispersal of cultivars. Since the early 2000s, the hypothesis of polycentric domestication in the Fertile Crescent has been accepted on the basis of the distribution of wild populations, archaeobotanical data, and genetic analyses of modern cereal populations (Gebel, 2004; Willcox, 2005). The transition from harvesting to fully established agriculture was gradual, taking several hundred years (Willcox, 2007). The domestication occurred independently in two regions: southeastern Anatolia/Middle Euphrates and the Southern Levant. The presence of a water source in these regions is surely the primary necessity that led to the establishment of villages, even if the wild cereals were sometimes distant (Willcox, 2005).

The Founder Crop Package

At the end of the 8th millennium bc, corresponding to the Pre-Pottery Neolithic B period (PPNB, 7500–7000 bc), domestic plants and animals were at the basis of subsistence. The founding crops, also called the “Neolithic crop package,” consisted of hulled cereals: einkorn (T. monococcum), emmer (T. dicoccum), and barley (H. vulgare subsp. vulgare) (Zohary, 1996). The free-treshing wheat (Triticum aestivum/durum) and naked barley (H. vulgare subsp. nudum) were domesticated in a second phase (Willcox, 2007). Pulses were reprented by lens (L. culinaris), pea (Pisum sativum), chickpea (Cicer arietinum), and vetch (Vicia ervilia). Secondary pulses such as broad bean (Vicia faba) and grass pea (Lathyrus sativus) do not seem to have met the same success, depending on location in the Fertile Crescent, and are thus not included systematically in the founding crop package. Flax (Linum usitatissimum) is the only textile and oilseed plant in the package. Thus, the Neolithic populations that spread into Western Europe have potentially five types of cereals, at least four pulses and one oil and textile plant.

General Chronological Setting of Agriculture Dispersal to Europe

Hypotheses regarding the routes of diffusion and the parameters for the outbreaks of Neolithic migration rely on a large number of proxies, such as radiocarbon dating and the comparison of material productions, genetic, climatic, archaeozoological, and archaeobotanical data. Radiocarbon dating is the strongest tool pointing to the arrival of an agricultural economy from the Near East. Cereal grains, or associated archaeological contexts, are found earlier in the East than in the West. The diffusion westward, from the Near East via Anatolia, started from around the very beginning of the 7th millennium bc. The pioneer front of colonization spread quickly from the core zone, as shown by early evidence of cultivated plants in the late Pre-Pottery Neolithic A (PPNA, 9500–8700 bc) site Klimonas in Cyprus, involving as well an early knowledge of navigation techniques from at least 10,000 years ago (Vigne et al., 2012). Rapid climate deterioration (aridification) or demographic pressures are often suggested as triggers for the diffusion process to Western Europe (Berger & Guilaine, 2009; Bocquet-Appel, 2011; Weninger et al., 2006).

By the mid-1960s, Clark (1965) proposed a model of diffusion from the Near East to Western Europe starting before 5200 bc. Since then, thanks to the increasing number of archaeological excavations and radiocarbon datings, which are also more precise, the diffusion of the Neolithic economy has become better understood. In the early 1970s, a new theory proposed an advancement at an average speed of about 1 km/year (Ammerman & Cavalli-Sforza, 1971). However, the linear progress is questioned because even if it was indeed fast, barriers are recorded in several European regions, as in the Pô plain (northern Italy), the Biscay Bay (north of Spain), and the Hungarian plains. Currently, the arrhythmic diffusion, or the boom-and-bust model, is the most popularly argued theory (Guilaine, 2003; Mazurié de Keroualin, 2003; Shennan, 2013). It should be noted that it is the movements of the components of the Neolithic economy (notably, the seeds of cultivated plants), not strictly those of the people that are considered (Rasse, 2008).

Two main streams of diffusion to Europe are identified: by maritime route, along the north Mediterranean coasts (Impressa/Cardial route), and by land, along the main water courses of Southeast and Central Europe, such as the Danube and the Rhine (Danubian route) (Rowley-Conwy, 2011). Climate (late frosts in temperate zones) and topography (mountainous areas) are probably not the only factors to have influenced the rhythm and the diffusion trajectories of the first farmers in Europe. Internal factors specific to each of the Neolithic societies certainly greatly influenced their way of managing the sustainability of their socioeconomic systems (Manning, Colledge, Crema, Shennan, & Timpson, 2016; Rasse, 2008).

The question of the contribution of local Mesolithic groups to the spread of Neolithic economy is also still widely discussed (e.g., Hadjikoumis, Robinson, & Viner, 2011). Diffusion processes may take slightly different forms (e.g., contacts, exchanges, or acculturation), depending on cultural contexts and geographical locations. For example, despite arguments in favor of the introduction of fully domesticated plants in Franchthi Cave, the artifacts (i.e., stone tools and ornaments) present a strong continuity between the Mesolithic and Neolithic horizons, which could correspond to a brief acculturation episode by local hunter-gatherers during the “Initial Neolithic” (Perlès, 1990). This could be a local phenomenon that cannot be applied generally to all sites with a succession of Mesolithic and Neolithic levels in southern Greece (Perlès, Quiles, & Valladas, 2013). This example illustrates quite well the probably complex process of Neolithic economy diffusion from the Near East to Europe, especially in the Balkans, at the origin of the two main streams of Neolithization to the west and north of the European continent.

Diversity of Crops and Farming Systems in Mediterranean and Temperate Europe in the Neolithic

Diversity of First Cultivated Plants in the Balkans

Chronocultural Background of Agriculture Dispersal in the Balkans

The east of the Balkan Peninsula (Greece, Macedonia, and Bulgaria) is the first point from which the Neolithic economy spread quickly to Europe. Based on the evidence presented in ceramic and lithic artifacts and architecture, this complex has a clear Near Eastern origin, in the cultural continuity of the final PPNB period (Mazurié de Keroualin, 2003). The Neolithic economy arrived in the Peloponnese and Macedonia about 6500 cal bc, before the first evidence in Thessaly and Bulgaria, suggesting that multiple points of contact could have occurred during the second half of the 7th millennium bc in the Aegean and Balkans (Lespez et al., 2013; Perlès et al., 2013). Thus, migration from Anatolia to the Aegean and Thrace regions may not correspond to a single event, but rather to separate waves of diffusion, both by sea and on land (Özdoğan, 2011). The Struma, Maritsa, and Vardar valleys in southwest Bulgaria could be the primary routes of Neolithic diffusion to the interior of the Balkan Peninsula around 6200–6100 bc, with the Karanovo and the Starčevo/Körös/Criş cultures (Lichardus-Itten, Demoule, Perničeva, & Grebska-Kulova, 2006). The first farming groups settled in fertile open lands of the Great Hungarian plain and would give birth to the widespread Linearbandkeramik (LBK) culture in Central Europe (see “Diversity of the First Cultivated Plants in Temperate Europe”). They were probably not greatly constrained by the environment, especially in the eastern Balkan Peninsula. In addition, the hunter-gatherers’ population density was probably very low at the time of their arrival.

The settlement types are varied. There are caves, such as Franchthi, which can be interpreted as a shepherd shelter that is seasonal or complementary to a permanent site that has not been discovered yet (Mazurié de Keroualin, 2003). Perennial sites are also encountered. They present square or rectangular house plans of varying sizes, constructed of brick or cob, as in Neo Nikomedeia in Macedonia or Kovačevo in Bulgaria (Lichardus-Itten, 2012). Both types of sites have provided archaeobotanical material. The fine pottery was generally painted. Sheep and goats seem to predominate at the very beginning of farming diffusion (Mazurié de Keroualin, 2003).

The west of the peninsula is often considered separately in archaeological literature, probably because this region doesn’t correspond, from a cultural point of view, to the Balkano-Anatolian complex. Two routes of diffusion to the Adriatic and south of the Carpathians are identified (Forenbaher et al., 2013; Orton, Gaastra, & Linden, 2016). The maritime route corresponds to the so-called Impressed Ware culture (6100 bc; see “Diversity of First Cultivated Plants in the Mediterranean”).

Crop Diversity in the Balkans

A consequent amount of archaeobotanical studies have been carried out in this pioneer colonization zone [i.e., Bulgaria and Greece (Colledge, Conolly, & Shennan, 2004, 2007b; Marinova, 2007; Marinova, Tonkov, Bozilova, & Vajsov, 2012, Marinova & Valamoti, 2014; Valamoti & Kotsakis, 2007)]. On the other hand, there are not many pieces of data available to the east of the Balkan Peninsula, especially for early Neolithic coastal settlements.

In these regions, the crop package is consistent with what is known in the Near East. It is composed of hulled wheat; hulled and naked barley; free-threshing wheat; a range of pulses such as lentil, pea, chickpea, and bitter vetch; and flax. From a quantitative point of view, hulled wheat varieties (emmer and einkorn) seem to be the most important crops in southeast Europe (Marinova & Valamoti, 2014). However, the quantitative criterion is not very relevant for comparing, as the hulled/naked wheat ratio depends on the chronology of the archaeological sites, the number of sites and samples studied, taphonomy, and past human practices.

Few qualitative variations can be noted in the group of pulses between Greece and southwest Bulgaria. First, chickpea is present but sparse. It seems that this pulse is not present in the north of Greece, although it has been identified in many sites in South Bulgaria, such as Kovačevo. However, in this site, chickpea is associated with the more recent stages (from 5600 bc) of the Neolithic in Southern Bulgaria, while it is not mentioned in the north (Marinova & Popova, 2008; Valamoti & Kotsakis, 2007). This confirms that the Neolithic economy might have penetrated Europe at many different points and times. Thus, not all plants came suddenly, suggesting a potential break in the transmission process. Second, grass pea is continuously present in southeast Europe, regarding the founder crop package, and the importance of this pulse in the Balkans is pointed out (Kislev, 1989; Valamoti & Kotsakis, 2007). Indeed, concentrations of grass pea have regularly been identified in Early Neolithic sites in Bulgaria and Greece (Valamoti, Moniaki, & Karathanou, 2011). Lathyrus cicera, the putative wild ancestor of L. sativus, is distributed in Turkey, as well as Greece and Transcaucasia (Zohary et al., 2012).

Diversity of First Cultivated Plants in the Mediterranean

Chronocultural Background of Agriculture Dispersal in the Mediterranean

A synthesis of radiocarbon dates, many on cereal seeds, from early Neolithic sites in Italy, southern France, and Iberia indicates that the spread of the Neolithic economy took place between 5700 and 5500 bc, by the maritime route (Zilhão, 2014). It therefore took no more than a couple of centuries to spread from the Thyrrenian Sea to the Albora coast. The diffusion of human groups, their economy, and their ideas over more than 3,000 km of coastal territories was then quite rapid, but probably not linear and continuous (Mazurié de Keroualin, 2003). The ceramic style, as well as radiocarbon dating compilation, indicates that different Neolithic groups may have arrived simultaneously around 5700 bc at different points in the northeastern Iberian Peninsula (Morales Hidalgo, Fontanals Torroja, Oms Arias, & Vergès Bosch, 2010). Furthermore, a long period of stasis seems perceptible before the first domesticated plants reached north-central Spain (Zilhão, 2014). Indeed, Biscay Bay, in the Cantabrian region, was not reached until the middle of the 5th millennium, as evidenced by wheat grain emmer dating to 4400 bc (Peña-Chocarro et al., 2005). At first, the pioneering advance seems to have become rather concentrated in the coastal areas before quickly moving inland, and even into higher altitudes via major river systems (Guilaine & Manen, 2007).

These pioneer communities are part of a large cultural entity, defined mainly on ceramic decoration and technology, called the Impressa/Cardial complex. In the central and western Mediterranean Basin, the ceramic paste was decorated with impressions and incisions with fingers, nails, varied tools, or shell edges, such as cardium, which gave its name to the Cardial culture (Mazurié de Keroualin, 2003). Impressa ceramics are more characteristic of the early Neolithic in Italy and southeast France, while the Cardial style is encountered in the western part of the Mediterranean Basin.

Village organization and extent, as well as architectural techniques for housing, are not well known for the Early Neolithic in the Mediterranean. Many settlements are found in the open air with light structures, or in rock shelters, which do not lend themselves to the conservation of archaeological structures or archaeobotanical remains. Rich and fertile soils in the valley floors seem to have been appreciated and utilized by the first farming communities for their settlements, as along the central coast of the Iberian Peninsula (Pérez-Jordà & Peña-Chocarro, 2013). A few sites have been preserved under waterlogged conditions, allowing for an outstanding preservation of organic material. Many wooden posts were discovered at La Marmotta (on Bacciano Lake in central Italy) and at La Draga (on Banyoles Lake in northeast Spain). These huts could correspond to long-term open-air settlements, the building of which would need a substantial labor investment (Fugazzola Delpino & Tinazzi, 2010). These two types of habitats could be interpreted as the coexistence of different mobility patterns (semimobile or permanent) and economic strategies (herders or farmers) from the early Neolithic (Mazurié de Keroualin, 2003). The same variations are visible in the livestock package, in which the proportions of pigs, sheeps/goats, cows, and wild fauna, as well as sheep morphotypes, are different depending on location and whether the sites are attributed to the Impressa or Cardial culture (Rowley-Conwy et al., 2013; Vigne, 2007).

Crop Diversity in the Mediterranean

In Italy, despite the substantial amount of sites studied, the number of remains identified is generally low. However, some Early Neolithic sites, as La Marmotta in Central Italy and Sammardenchia in the Pô Plain, have been subject to systematic archaeobotanical studies, which allow for a substantial overview of the crop package in the coastal Central Mediterranean (Rottoli & Pessina, 2007; Rottoli & Castiglioni, 2009). In France, the amount of data to date is very weak (Gassin et al., 2010). In the Iberian Peninsula, several sites have yielded important and well-preserved data sets, as on the lakeshore site of La Draga in Catalonia and Cova de l’Or and Los Castillejos on the southeastern coast of the peninsula (Antolín & Buxó, 2011, 2012; Rovira, 2007). Recent regional syntheses give substantial data on cultivated plants in these regions and the variation of crop package composition at the supraregional scale (Antolín, Jacomet, & Buxó, 2015; Buxó, 2007; Peña-Chocarro et al., 2013; Pérez Jordà & Peña-Chocarro, 2013; Zapata, Peña-Chocarro, Pérez-Jordá, & Stika, 2004).

From a qualitative point of view, the Neolithic crop package (especially cereals) is equivalent to the Balkans and the Near East. The first farmers in the central and western Mediterranean have grown einkorn; emmer; free-threshing wheat; hulled and naked barley; a wide range of pulses, such as lentils, peas, beans, vetches, and grass peas; and flax. The set of cultivated plants is highly diversified. On Cardial sites from southern France and the Iberian Peninsula, naked cereals predominate, while emmer and einkorn appear to be predominant to the east, in the Central Mediterranean (Antolín & Buxó, 2012; Buxó, 2007; Gassin et al., 2010; Peña-Choccaro et al., 2013). However, the more or less rigorous division between hulled cereals in the Impressa culture and naked cereals in the Cardial culture needs to be confirmed by further studies. Indeed, in the northeast of the Iberian Peninsula, the proportions of naked and hulled cereals vary depending on the site (Antolín & Buxó, 2012). Hulled cereals have been identified in some other sites on the central coast of Catalonia. In other sites, such as La Draga, the naked wheats dominate. There are no taphonomic, geological, or pedological features to explain these differences, indicating that several agricultural traditions in the choice of main cereals may have coexisted as soon as the early Neolithic at the microregional scale (Antolín & Buxó, 2011, 2012). The two types of cereals (naked or hulled) do not have the same requirements (in terms of dehusking, for example) and involve different technical systems. However, the comparison of proportion is based on many cereals identified at the genus level (Triticum or Hordeum), which prevents a clear view of the naked/hulled dominances in the region. Moreover, this example illustrates quite well the limits of the sole morphological identification of the taxa to address early crop diversity. Indeed, genotype differences can be recorded within a field of wheat that looks visibly homogeneous, as it is currently highlighted in the western Mediterranean (Oliveira et al., 2012).

Regarding pulses, chickpea is absent from the Impressa/Cardial route, which indicates that it was not diffused by the very first farmers (Colledge, Conolly, & Shennan, 2005). The reasons for this neglect are still poorly understood. It may or may not have been a conscious choice. Indeed, explanations of an exclusively environmental nature are difficult to substantiate, since all other taxa domesticated in the Near East are identified in the Mediterranean stream. Furthermore, we note the addition of common vetch (Vicia sativa) and broad bean (Vicia faba) as minor components of the crop package in the Mediterranean (Antolín & Buxó, 2012; Rottoli & Pessina, 2007; Rottoli & Castiglioni, 2009). The findings of a large amount of broad bean in early-PPNB occupation levels of Tell el-Kerkh (northwest Syria) and in a middle-PPNB storage structure at Yiftah’el (Israel) could also point toward an early cultivation of this pulse in the area where the Neolithic crop package originated (Kislev, 1989). There is a real difficulty in differentiating wild and domesticated forms because of the overlapping of seed size, especially as these pulses could also be considered as a contaminant in cultivated fields (Zohary et al., 2012). Furthermore, charred archaeological pulses are very fragile, which does note facilitate their identification (Tanno & Willcox, 2006). Thus, the origin and diffusion of pulses remains a largely unknown part of early agriculture across the Mediterranean basin, as well as in the Near East.

An additionnal species, opium poppy (Papaver somniferum), completes the Mediterranean crop package and has not, to date, been identified, further east in the Balkan Peninsula and the Near East (see “Looking for the origin of opium poppy in Western Europe”). At La Marmotta, seeds preserved in carbonized and uncarbonized form, charred capsules, and stigmatic disks were discovered (Rottoli & Pessina, 2007). In Spain, poppy has been identified in sites such as La Draga, Los Murceliagos, and La Lampara (Antolín & Buxó, 2012; Stika, 2005).

Diversity of the First Cultivated Plants in Temperate Europe

Chronocultural Background of Agriculture Dispersal in Temperate Europe

The Neolithic economy spread from south-central Europe, approximately following the Danube watershed, into northwestern continental Europe during the 6th millennium bc. The first farmers of south-central Europe appear in the Starčevo/Körös/Criş complex (6100–5500 bc), which extend from Transdanubia to the Great Hungarian Plain and Transylvania. Around 5600 bc, the LBK culture emerged in southwestern Slovakia and western Hungary, to the north of Lake Balaton (Gronenborn, 1999). In this area, the relationship between LBK culture and the Starčevo/Körös/Criş complex is still subject to debate (Stadler & Kotova, 2010).

Around 5500 bc, the LBK culture began its rapid first phase of expansion westward, reached the Rhine by 5300 bc, and, during a second phase, extended to the Paris Basin, ending around 4900 bc (Bogucki, 2003). To the east, LBK groups settled during the late 6th millennium bc in western Ukraine (Bogucki, 2000). LBK material production shows some variation depending on location and chronology of the site. However, the cultural identity of the pioneer farmers is strong enough to follow the route of early agriculture diffusion through Temperate Europe from Southwest Ukraine to the Paris Basin. This culture is characterized by a broad distribution of similar pottery types and decoration, as well as long houses with a rectangular layout. The decoration of pots—incised linear forms in bands or ribbons—provided the name for this archaeological culture (Linearbandkeramik, Linear Ceramic, or Rubané). Houses and villages are clearly sedentary. The negatives of wood posts, called post holes, make it possible to trace the ground plan of the Danubian houses, which could measure between 10 and 45 meters (Coudart, 1998). The first farmers settled mainly on loessic patches, which are assumed to have a high natural fertility (Catt, 2001), and uninhabited zones separate clusters of villages from each other. LBK emergence is believed by some to be linked to an increase of wetter and colder climatic conditions, and its collapse to warmer and drier conditions (Dubouloz, 2008).

Crop Diversity in Temperate Europe

The Neolithic crop package in Central and Northwest continental Europe is quite well known, thanks to many archaeobotanical studies carried out in LBK sites (e.g., Bakels, 1978; Knörzer, 1997; Kreuz, 2007; Kreuz, Marinova, Schäfer, & Wiethold, 2005; Salavert, 2011).

In the Starčevo/Körös/Criş sphere, at the origin of farming in Central Europe, the diversity of crops may be underestimated. Indeed, the plant economy of the Starčevo/Körös/Criş is mainly known through imprints in daub and ceramics. However, few sites from this period provide charred macroremains, which allow for an overview of the plants cultivated by the first farmers in South-Central Europe (Bogaard, Krause, & Strien, 2011; Colledge & Conolly, 2007b; Gyulai, 2010; Reed, 2015). To date, the list of crops is rather smaller. Both hulled wheats (einkorn and emmer) and, sometimes barley, lens, and peas are identified. The arrival of naked wheat seems to have been delayed to the end of the 6th millennium bc, which corresponds to Late Neolithic (Reed, 2015).

In LBK sites, the range of crops is also relatively narrow (Bakels, 2009; Colledge et al., 2005; Kreuz et al., 2005; Kreuz, 2007; Salavert, 2011). The general pattern shows that einkorn dominated central European sites for the duration of the LBK culture (Kreuz, 2007). Einkorn has a good resistance to lodging that could explain the choice of this cereal, despite its lower yield compared to emmer (Kreuz, 2007). However, west of the Rhine, in the southern Limbourg, Hesbaye, and Hainaut regions, emmer clearly dominates, whether the grains or chaff are taken into account (Salavert, 2011). Thus, a climatic hypothesis cannot be the only motivation behind humans’ choices in their farming strategies. Concerning barley, hulled and naked forms are scarce, which gives an unknown status to this taxon. Barley, as with naked wheat, whose findings are anecdotal, may have been considered a weed in LBK fields rather than a true crop plant (Kreuz, 2007). However, even if the barley form (hulled or naked) is not always specified in archaeobotanical counting tables because of bad conservation of the charred caryopses, naked barley seems more frequent in the western part of the LBK cultural extension, more precisely in the Hainaut and Parisian Basin, and could thus be part of the crop package in these peripheral regions of the primary Neolithic economy dispersal (Bakels, 2009; Salavert, 2011). Indeed, naked barley has been recognized at Wange and Overhespen in the north of Hesbaye (Bakels, 1992) and in four LBK sites in the Aisne valley in France (Bakels, 1999).

The group of pulses is composed of lentils and peas. In the group of fiber/oleaginous plants, in addition to flax, opium poppy (Papaver somniferum) appears only in sites dated from 5300 bc and related to the second phase of LBK extension (Bakels, 1996; Kreuz, 2007; Salavert, 2011).

There is, thus, a clear decrease of crop diversity in Temperate Europe. This reduction corresponds to a moment of rupture in the diffusion of the Neolithic economy. The dynamic slows down, or stops, for several centuries on the level of the Hungarian plain. This pause allowed human societies, as well as the animals and plants that they transported, to adapt to different climatic conditions (more humidity, longer winter) than those encountered previously. However, the cultural argument—that is to say, the choice made by the first farmers of Temperate Europe—cannot be ignored, as hulled wheats also can be successful in temperate environments (Colledge et al., 2005). This break could have enabled the groups to acquire agricultural know-how before being able to spread them into western and eastern Temperate Europe (Bogucki, 2000).

Early Farming Systems in Mediterranean and Temperate Europe

The definition of Neolithic farming systems (more precisely, the seasonality of crops or the maintenance techniques of soil fertility) is at the center of archaeobotanical research, which relies on weeds and stable isotope analyses.

In Central Europe, the specific diversity of weeds is also restricted. The most common species are Bromus secalinus, Chenopodium album, Fallopia convolvulus, and Lapsana communis. These are the main components of the plant association Bromo-Lapsanetum-Praehistoricum, defined by Knörzer (1971) based on weed assemblages. However, this association is no longer observed in cultivated fields in Central Europe. According to several authors, it would show the presence of fields, permanent or not, cultivated every year with the same methods throughout the LBK territory, which would cause a well-defined combination of weeds (Bakels, 1978, 2009; Knörzer, 1971). It is difficult to interpret extinct associations to reconstruct past agricultural practices such as the size of fields, their sustainability, and the seasonality of crops.

For a long time, the practice of slash-and-burn agriculture was suggested to explain the rapid expansion of early farmers in the forest environment of Central Europe. The principle is as follows: A forested area is cultivated over a short period (one to five years) after being cleared and burned. Organic burned material provides nutrients that improve agricultural yield. After this short period, crops are moved to another area that has undergone the same treatment (Bogaard, 2002), allowing the initial site time to recover. However, the presence of long-lived Neolithic LBK sites such as Langweiler 8 and Vaihingen and der Enz (Germany), as well as the spatial proximity of villages, do not offer an economic area wide enough to allow the operation of the system in each village across many generations.

Furthermore, the comparison of weed censuses from current experimental fields, in addition to a large number of archaeological weeds that have been precisely identified and come from deposits resulting from the processing of harvests of emmer and einkorn, exclude the possibility of shifting cultivation in the Early Neolithic in Temperate Europe (Bogaard, 2002). Indeed, with a shifting cultivation system, perennial weeds dominate. However, in archaeological assemblages, perennial weeds account for only 2% of the samples taken into account, and annual weeds greatly dominate.

Thanks to weed autoecology, Bogaard (2004) has suggested that Early Neolithic farmers would have practiced intensive and permanent agriculture on a limited surface, usually near settlements. The sustainability of the fields was permitted, thanks to a strong supply of animal manure and weeding. The main sociotechnological implications are agricultural production at the domestic scale, strong integration between livestock and agriculture, and probably a very significant investment of labour and time (Jones, 2005; Saqalli et al., 2014). For example, manuring and weeding would be six to seven times more labor intensive than shifting cultivation (Pétrequin, Pétrequin, & Schaal, 2015). In the Balkans, the contribution of manure, as evidenced by nitogen isotope values (δ‎15N), was integrated into agricultural systems from the very beginning of the spread of Neolithic economics (Bogaard et al., 2013; Fraser, Bogaard, Schäfer, Arbogast, & Heaton, 2013). Furthermore, at Vaihingen an der Enz, Germany, the integration of archeobotanical analyses, especially weed autoecology (such as soil pH requirements), as well as ceramic typology on a site almost completely excavated, has led to claims of a sort of clanic/group organization, with the distribution of cultivated areas according to group membership and a transfer of plots from generation to generation (Bogaard et al., 2011).

The Early Neolithic farming system in the Western Mediterranean has not been described in detail, mainly because of badly preserved archaeological weed assemblages at current sites. Statistical approaches, with precise identification based on substantial data sets, cannot yet be applied (Antolín & Buxó, 2012). Furthermore, it is far too early to formulate a general trend applicable to the whole Mediterranean region. At Los Castilleros, weed assemblages indicate permanent fields (Rovira, 2007). It seems that “most common taxa are annual plants typical of disturbed areas that could grow as arable weeds in irrigated or dry fields” (Antolín & Buxó, 2012, p. 99). The most common taxa are Chenopodium album and Galium aparine. The very few forest taxa in the assemblages could indicate that early farmers in Spain did not use the shifting cultivation (Antolín, Buxó, Jacomet, Navarrete, & Saña, 2014). The ecological requirements of annual plants and the flowering length indicate relatively intensive perturbations as well as long-term and intensively managed sowing of plots in autumn and possibly spring (Antolín et al., 2015; Peña-Chocarro et al., 2013).

Looking for the Origin of Opium Poppy in Western Europe

Identification of Wild and Cultivated Poppy

Opium poppy is a significant addition in the Neolithic crop package in Meditterranean and Temperate Europe. Opium poppy could be one of the unique plants domesticated in the European territory during the Neolithic. The origin and diffusion of the plant in Western Europe are yet to be understood, especially the question of its wild progenitor.

The family Papaveraceae occurs in temperate and subtropical climates. Today, its main distribution is around the Mediterranean region and the Middle East (Baser & Arslan, 2014). Papaver somniferum subsp. setigerum is often considered to be the wild ancestor of Papaver somniferum subsp. Somniferum, which is the cultivated form (Hammer, 1981; Zohary et al., 2012). However, recent botanical results based on their geographical distribution and the morphological characteristics of poppy identify three subspecies of poppy cultivated at present: P. somniferum subsp. setigerum, P. somniferum subsp. Songaricum, and P. somniferum subsp. somniferum. The putative wild ancestor can therefore also be considered as a cultivated form by botanists (Baser & Arslan, 2014).

The Western and Central Mediterranean is the region of origin for wild poppy that is most often mentioned in archaeobotanical literature (Bakels, 1996; Knörzer, 1971; Schultze-Motel, 1979). However, wild populations of poppy have developed in other geographical areas, including Central Asia, the Caucasus, North Africa, and the Eastern Mediterranean (Salavert, 2010). Given the range of opium poppy, based on the occurrence of its wild ancestor and the evidence of current phytogeography, its origin is difficult to identify. The wild and cultivated forms of P. somniferum are interfertile, and wild populations that are supposed to be “authentic” may be naturalized populations or escaped from cultivated fields (De Candolle, 1883; Ladizinsky, 1998). This is why some authors believe that there are currently no known true wild populations of opium poppy in the world (Merlin, 1984; Chouvy, 2001). The wild ancestor and the natural distribution zone of wild poppy, therefore, are imprecise and may not be confined to the Western Mediterranean.

The morphological criteria to distinguish wild from cultivated seeds in archaeological records are not established. The poppy seed is small (less than 1 mm in diameter) and spherical. It has multiple facets and is pentagonal and hexagonal edged with a slight bulge (Fig. 5). Capsules and stigmatic discs of opium poppy also have been identified at archaeological sites.

Agricultural Dispersals in Mediterranean and Temperate EuropeClick to view larger

Figure 5. Seeds of P. somniferum (opium poppy) from a Linearbandkeramik site in Belgium (around 5000 bc).

Although cultivated seeds are generally larger than wild seeds, there are no valid criteria for differentiating them (Hammer, 1981). The variability of cultivated poppy seeds is very broad and also covers the wild form. It is therefore impossible to distinguish between the seeds, as well as the capsules, of wild and cultivated poppy, especially from the Neolithic period. Furthermore, the number of seeds identified at archaeological sites is generally low. Particular taphonomical conditions must be present to preserve the seeds, and the remains most often originate from a single sample. This rarity can be attributed to the small size of the seeds, which requires a suitable sieving method (0.25 μ‎m mesh). In the same way, the oil content of the seed does not predispose it to preservation. Finally, processes of poppy transformation may not necessitate contact with fire, which weakens the chances that the seeds will be charred and thus preserved at archaeological sites.

Earliest Neolithic Poppy on Archaeological Sites

The largest number of Neolithic archaeological sites that have delivered poppies are located in the Impressa/Cardial and LBK complexes in Western Europe. In the current state of research, about 30 sites dated between 5200 and 5000 bc have delivered seeds of P. somniferum in the LBK. Sites are located in the area delimited by the province of Hesse (Germany) in the east and Hainaut (Belgium) in the west (Salavert, 2010). The seeds are absent from the earliest LBK sites (5500–5300 bc), even in the case of sites located east of the Rhine, and appear only during the second phase of LBK expansion, after 5300 bc. Outside this zone, several determinations are mentioned in France, Austria, and Poland, still at LBK sites dating between 5200 and 5000 bc (Salavert, 2011). Thus, the opium poppy is well established in the LBK context, particularly in the northwestern extension of the culture. In the Impressa/Cardial complex, a large quantity of carbonized and noncharred seeds, capsules, and stigmatic disks was found at the lakeside site of La Marmotta (Italy), dated by radiocarbon and dendrochronology between 5500 and 5400 bc (Kromer, 2009). At the site of La Draga (Spain), charred and watterlogged seeds were identified in one Neolithic structure. The archaeological level is dated between 5400 and 5100 bc, with a majority of dates between 5250 and 5150 bc (Antolín & Buxó, 2011). The opium poppy is mentioned in at least three other Cardial sites dated between 5200 and 4900 bc (Antolín & Buxó, 2012; Salavert, 2011). The importance of the plant in the Neolithic Mediterranean has probably been underestimated because of the small size of the seeds and the mode of conservation (charred), which is not suitable for oleaginous seeds. The Eastern Mediterranean also has delivered poppy. A waterlogged seed was found at the Atlit-Yam Pre-Pottery Neolithic C (PPNC) site in Israel (Kislev, Hartmann, & Galili, 2004). The series of radiometric datings obtained at the site ranges from about 7400 and 5900 bc. This is the single identification of P. somniferum in the Near Eastern Neolithic. The antiquity of the PPNC opium poppy seed can be questioned, on the one hand, by its isolated nature in a region heavily investigated by archaeobotanists, and on the other hand, by the fact that only one (watterlogged) seed was discovered on this submerged site very favorable to the preservation of organic material. No opium poppy has been mentioned in Neolithic sites in Southeast Europe.

The Spanish and Italian arcaheological identifications, in a region supposedly at the origin of the (also supposed) ancestor of the plant, are weakly anterior to LBK Europe. The limitations of the comparisons of dating are real (particularly the inaccuracy of radiocarbon datings), but above all, the fact that it is not the poppy seeds that are directly dated, but other materials (bones, charcoal) sometimes localized in other structures or archaeological layers that date the whole site or the occupation phase.

Scenario of Domestication and Diffusion

Absolute datings from the sites of the Early Neolithic period in Western Europe do not clearly indicate where the opium poppy was cultivated for the first time. One possibility is that the poppy accompanied the dispersal of Neolithic pioneer communities with other cultivated plants originating in the Near East, such as wheats, pulses, and flax. This hypothesis is uncertain because this plant has not been discovered, to date, in the earliest LBK sites (the Starčevo/Körös/Criş or Balkano-Anatolian complex).

If we assume that the origin of the plant is the Western Mediterranean, poppy could have integrated the LBK area under its wild or cultivated form. The contacts between the Neolithic populations of Southern and Northern Europe also have long been observed in archaeological artifacts. They are, for example, established between the Cardial-Epicardial farmers and the LBK populations through ceramic decorations (e.g., Meier-Arendt, 1966; Guilaine & Manen, 1997) and ornaments (Bonnardin, 2009). In addition, wild plants, such as Bromus sterilis/tectorum and perhaps Setaria viridis/verticillata, identified occasionally in LBK sites, originate from the Mediterranean regions (Oberdorfer, 1990). Thus, it is not surprising that seeds of Mediterranean origin are identified in LBK archaeobotanical assemblages.

There have been no discoveries of incised capsules or other elements that would make it possible to understand the use of this plant by early farmers. Poppy can be grown for its oil, or the seeds may have been added to food preparations and consumed. The psychotropic properties may not have been unknown to the Neolithic societies. Furthermore, in Vaihingen an der Enz, it seems that only the houses located in the southeast of the site have adopted this plant. No different taphonomic conditions from the rest of the village could explain this spatial distribution. This could indicate a different social status of certain people as favored contacts with the Southern Cardial (Bogaard et al., 2011) or particular knowledge about poppy cultivation.

Spread of Agriculture to Northern and Western Europe

The presence of cultivated plants is a marker of the spread of the agrarian economy, especially when the seeds are dated directly to the radiocarbon, and are accompanied by weeds. On this basis, there is no convincing evidence of Mesolithic agriculture to date in the north and northwestern margins of Europe which correspond to southern Scandinavia, northern Germany and Poland, as well as Britain and Ireland (Behre, 2007; Sørensen & Karg, 2014; Tresset, 2015). The diffusion of cultivated plants is attested there from 4000 bc, which marks the beginning of the initial Neolithic, a little less than a millennium after the decline of the LBK culture in central and western Temperate Europe (see “Diversity of the First Cultivated Plants in Temperate Europe”). During this millennium, different archaeological cultures developed there, such as Blicquy/Villeneuve-Saint-Germain, Rössen, Epi-Rössen, Michelsberg, or Chasséen. Their social, cultural, and economic structures are significantly different from those of the LBK pioneer farmers who travelled along the Danube from 5500 bc. Concerning cultivated plants, naked cereals have been developed, in particular tetraploid wheat (Triticum durum/turgidum), probably under the influence of neolithic populations from the western Mediterranean (Bakels, 2009).

The modalities for the introduction of cultivars at the beginning of the 4th millennium in the northern and western Europe are still discussed. The particular points are the progressive adoption of agriculture by indigenous mesolithic groups, the sudden arrival of cultivars thanks to small groups of experimented farmers from central Europe, as well as the importance of cereals compared to gathering/fishing/hunting products, in the early Neolithic diet (Rowley-Conwy, 2004; Jones & Rowley-Conwy, 2007; Thomas, 2008; Kreuz et al., 2014; Sørensen & Karg, 2014; Jones & Sibbesson, 2016). Several factors are proposed to explain the relaunch of the crop diffusion to new geographical areas at the beginning of the 4th millennium bc: demographic pressures; a warmer and drier climate around 4000 bc, which would extend the spring growing season for cereals; or the search for good flint sources (Bonsall et al., 2002; Jones et al., 2000; Sørensen & Karg, 2014).

According to radiocarbon dates, only three centuries have been needed to adopt the Neolithic economy, suggesting the arrival of small groups of experienced farmers in southern Scandinavia and northern Germany (Sørensen & Karg, 2014). Similarly, for southern Scandinavia, mitochondrial DNA studies show a close relationship between Neolithic individuals belonging to the northern Funnelbeaker culture (TRB or TBK), developing in northern Germany to the territory of Sweden, with those of Central Europe (Malmström et al., 2015). In Britain and Ireland, pollen records show that the disturbances of initial Neolithic people activities were sudden and rapid in deciduous forest (Woodbridge et al., 2014). However, seeds of cultivated cereals are still often identified in small amounts around 4000 bc (Jones & Rowley-Conwy, 2007). The agrarian economy seems fully established from 3750 bc in Ireland (Whitehouse et al., 2014; McClatchie et al., 2016) and from 3600 bc in northern Germany (Kirleis et al., 2012). However, taphonomic factors may explain the weak data set related to early Neolithic settlements in these areas (Jones & Rowley-Conwy, 2007).

Archaeobotanical syntheses from southern Scandinavia (northern Germany, Denmark, and southern and western Sweden) show that the main cultivated plants identified in the Funnelbeaker context, corresponding to local early and middle Neolithic, are emmer and einkorn, as well as naked barley and bread wheat (Robinson, 2003; Larsson & Broström, 2011; Kirleis & Fischer, 2014). For northern Germany, poppy and pulses are not identified at the very beginning of the period (Kirleis & Fischer, 2014). Recently, discoveries of naked tetraploid wheat in southern Scandinavia support a possible cultural interaction between the Funnelbeaker and contemporaneous neolithic groups in Central Europe, such as the Michelsberg culture, where this cereal is widely cultivated (Kirleis & Fischer, 2014; Kreuz et al., 2014). In the same way, the Michelsberg crop package is characterized by weak evidence of pulses (lentils and pea) and oleaginous/fiber plants (Kreuz et al., 2014). In Ireland, the early Neolithic is characterized by a mixed blend of cereals composed mainly of emmer, some naked wheat (T. astivum/durum/turgidum), and naked barley. Einkorn is rare. Flax is present from the beginning of the Neolithic, while pulses are absent (McClatchie et al., 2014, 2016).

This is a very rapid view of the spread of the Neolithic economy based on cereal cultivation in the northern margins of Europe. The northern and northwestern parts of Europe are therefore characterized by the importance of hulled wheat, but also free-treshing wheat and naked barley, which corresponds to the main cereals cultivated by the Neolithic groups of Central Europe at the beginning of the 4th millennium bc. Pulses are rare (Colledge et al., 2005). The characterization of farming systems is still in progress. In England, preliminary studies indicate a high rate of annual plants compared to perennial weeds characteristic of fields cultivated by a slash-and-burn system (Bogaard et al., 2007). As in Ireland, the intensive system of fixed and perennial fields in the landscape is favored (McClatchie et al., 2014).

Agricultural Dispersals in Mediterranean and Temperate EuropeClick to view larger

Figure 6. Synthetic presentation of the maritime diffusion in Mediterranean (Impressa/Cardial) and continental diffusion in Temperate Europe (Linearbandkeramik), as well as the spread of cultivars in northern and western Europe between 6500 and 4000 bc.


To summarize these findings, the dispersal route of agriculture into Mediterranean and Temperate Europe is rather well known, thanks to archaeology. The diffusion of the Neolithic economy was rapid and spanned about 1,500 years, from 6500 bc in Greece to the beginning of the 4th millennium bc in Ireland (Fig. 6). The maritime route of agriculture dispersal, along the Mediterranean coasts, presents a minimal loss of diversity compared to agricultural origins in the Near East. Only the chickpea did not spread with the very first farming communities in the Balkans. Naked cereals are included in the crop package, but it seems that hulled wheats were preferred at the very beginning of the Neolithic diffusion (Balkans and Impressa). The habitat types (caves, rock shelters, open-air sites) are diversified, as are the crops that the first Mediterranean farmers cultivated from 6500 to 5500 bc. Farming practices need further investigation. Thanks to palaeocological studies, it seems that the first farming groups in the Western Mediterranean may have had a heterogeneous impact, in terms of timing and spatial extent, on the oak forests, which we assume earlier Mesolithic groups modified only sparsely (Revelles, in press; Thiébault, 1988).

In Temperate Europe, the habitat types are more homogeneous and the Neolithic crop package presents a low diversity. Cultivated fields are probably long-lasting and may have been managed on the domestic scale for several family generations. This transfer goes hand in hand with the high investment required for agricultural work and the maintenance of the sustainability of the field (thanks to manure). It also implies a strong territorial settling of the groups of the early Neolithic in the Danubian sphere in the second half of the 6th millennium bc. Studies on daily firewood gathered near the LBK sites of the first farmers in Belgium show the rapid increase in heliophilic taxa of hedges (Maloideae, for example). This dynamic may indicate the rapid opening of forest areas for the establishment of fields and their maintenance near habitats, as well as local anthropic pressure on the landscape (Salavert, Bosquet, & Damblon, 2014a; Salavert & Dufraisse, 2014b). Less than one millennium after the end of the LBK culture, the agriculture saw a second wave of dispersal to northern and western Europe. As during the LBK, the high percentages of Maloideae and Corylus indicate that impact of Funnel beaker farmers on forest may have occurred on a small scale (Jansen et al., 2014).

The two routes, maritime and continental, thus show different dynamics in diffusion processes, cultures, and farming practices. Opium poppy is the main addition in western Europe, and its use concerns both regions. Many points must be highlighted to go further in the research on poppy domestication and uses. First, its wild progenitor is still not recognized with absolute certainty. Second, the status (wild or cultivated) of early Neolithic poppy is not understood, thanks to its seed morphology. Finally, the radiocarbon dating is indirect and not accurate enough to trace the diffusion process with precision. However, thanks to archaeological data, we do know that poppy was probably not integrated into the Neolithic crop package from the Near East or the Balkans. It seems to have been introduced in the Central and Western Mediterranean basin, from about 5400 bc at the earliest, but most of the discoveries are from around 5200 bc, both in Mediterranean and Temperate Europe. The proximity of radiocarbon dating indicates that the diffusion from south to north might have been very rapid. The cultivation or the gathering of opium poppy does not seem to be shared by all farmers within an LBK village. Thus, the plant seems to have had a different status than traditional crops, such as hulled wheat and pulses, in the Neolithic economy.

Cultivated plants are now known to be the basis of human alimentation in the Early Neolithic, although gathering also continued to be practiced, especially for foods such as hazelnuts, wild apples, and wild grapes. However, this part of the Neolithic diet, along with the status of wild trees in the farming system of the Early Neolithic, is not well understood. There are many aspects of the dispersal (notably crop diversity) and components of the farming system that need further investigations. The A-DNA and stable isotope research currently in development will surely enhance our knowledge in this subject without neglecting archaeobotanical studies, which are still at the base of our knowledge of the very first farming communities in Europe.

Further Reading

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Guilaine J. (2000). La diffusion de l’agriculture en Europe: une hypothèse arythmique. La difusión de L’ agricultura en Europa: una hipótesis aritmética. Zephyrus, 53–54, 267–272.Find this resource:


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