Talkin' About a Revolution. Changes and Continuities in Fruit Use in Southern France From Neolithic to Roman Times Using Archaeobotanical Data (ca. 5,800 BCE - 500 CE).

The use and socio-environmental importance of fruits dramatically changed after the emergence of arboriculture and fruit domestication in the eastern Mediterranean, between the 5th and the 3rd millennia BCE. Domesticated fruits together with cultivation techniques apparently reached the western Mediterranean via colonial activities during the 1st millennium BCE - early 1st millennium CE. However, the pace and chronology of this diffusion as well as the recompositions in diversity, to adapt to new socio-environmental conditions, remain poorly known. In this study we investigate archaeobotanical records in Southern France from the Neolithic to the end of the Roman empire (ca. 5,800 BCE - 500 CE) to assess changes in fruit use as well as the emergence, spread and evolution of fruit cultivation. We explore changes in native traditions faced with innovations brought by Mediterranean colonization and how domesticated fruit cultivation spread from the Mediterranean to more temperate areas. Archaeobotanical data from 577 assemblages were systematically analyzed distinguishing two datasets according to preservation of plant remains (charred vs. uncharred), as this impacts on the quantity and diversity of taxa. The 47 fruit taxa identified were organized in broad categories according to their status and origin: exotic, allochtonous cultivated, indigenous cultivated, wild native. We also analyzed diversity, quantity of fruits compared to the total of economic plants and spatio-temporal variations in the composition of fruit assemblages using correspondence factor analyses. Archaeobotanical data reflect variations and continuities in the diversity of species used through time and space. In the Mediterranean area, significant changes related to the arrival of new plants and development of fruit cultivation occurred mainly, first during the Iron Age (6th-5th c. BCE), then in the beginning of the Roman period. Large cities played a major role in this process. In agreement with archeological information, archaeobotanical data reveal the predominance of viticulture in both periods. However, arboriculture also included other fruit species that have been subject to less intensive and specialized cultivation practices. Most significantly, this study pinpoints the continuous contribution of native, supposedly wild fruits throughout the chronology. Despite the homogenizing Roman influence, results reveal clear differences between the Mediterranean and temperate regions.

Pip shape echoes grapevine domestication history.

The pip, as the most common grapevine archaeological remain, is extensively used to document past viticulture dynamics. This paper uses state of the art morphological analyses to analyse the largest reference collection of modern pips to date, representative of the present-day diversity of the domesticated grapevine from Western Eurasia. We tested for a costructure between the form of the modern pips and the: destination use (table/wine), geographical origins, and populational labels obtained through two molecular approaches. Significant structuring is demonstrated for each of these cofactors and for the first time it is possible to infer properties of varieties without going through the parallel with modern varieties. These results provide a unique tool that can be applied to archaeological pips in order to reconstruct the spatio-temporal dynamics of grape diversity on a large scale and to better understand viticulture history. The models obtained were then used to infer the affiliations with archaeobotanical remains recovered in Mas de Vignoles XIV (Nîmes, France). The results show a twofold shift between the Late Iron Age and the Middle Ages, from table to wine grape varieties and from eastern to western origins which correlates with previous palaeogenomic results.

Evolution and history of grapevine (Vitis vinifera) under domestication: new morphometric perspectives to understand seed domestication syndrome and reveal origins of ancient European cultivars.

BACKGROUND AND AIMS: In spite of the abundance of archaeological, bio-archaeological, historical and genetic data, the origins, historical biogeography, identity of ancient grapevine cultivars and mechanisms of domestication are still largely unknown. Here, analysis of variation in seed morphology aims to provide accurate criteria for the discrimination between wild grapes and modern cultivars and to understand changes in functional traits in relation to the domestication process. This approach is also used to quantify the phenotypic diversity in the wild and cultivated compartments and to provide a starting point for comparing well-preserved archaeological material, in order to elucidate the history of grapevine varieties. METHODS: Geometrical analysis (elliptic Fourier transform method) was applied to grapevine seed outlines from modern wild individuals, cultivars and well-preserved archaeological material from southern France, dating back to the first to second centuries. KEY RESULTS AND CONCLUSIONS: Significant relationships between seed shape and taxonomic status, geographical origin (country or region) of accessions and parentage of varieties are highlighted, as previously noted based on genetic approaches. The combination of the analysis of modern reference material and well-preserved archaeological seeds provides original data about the history of ancient cultivated forms, some of them morphologically close to the current 'Clairette' and 'Mondeuse blanche' cultivars. Archaeobiological records seem to confirm the complexity of human contact, exchanges and migrations which spread grapevine cultivation in Europe and in Mediterranean areas, and argue in favour of the existence of local domestication in the Languedoc (southern France) region during Antiquity.

Eco-evo-devo implications and archaeobiological perspectives of trait covariance in fruits of wild and domesticated grapevines.

The phenotypic changes that occurred during the domestication and diversification of grapevine are well known, particularly changes in seed morphology, but the functional causes and consequences behind these variations are poorly understood. Wild and domesticate grapes differ, among others, in the form of their pips: wild grapes produce roundish pips with short stalks and cultivated varieties have more elongated pips with longer stalks. Such variations of form are of first importance for archaeobotany since the pip form is, most often, the only remaining information in archaeological settings. This study aims to enlighten archaeobotanical record and grapevine pip development by better understanding how size and shape (co)variates between pip and berry in both wild and domesticated Vitis vinifera. The covariation of berry size, number of seeds per berry ("piposity"), pip size and pip shape were explored on 49 grapevine accessions sampled among Euro-Mediterranean traditional cultivars and wild grapevines. We show that for wild grapevine, the higher the piposity, the bigger the berry and the more elongated the pip. For both wild and domesticated grapevine, the longer is the pip, the more it has a "domesticated" shape. Consequences for archaeobotanical studies are tested and discussed, and these covariations allowed the inference of berry dimensions from archaeological pips from a Southern France Roman site. This systematic exploration sheds light on new aspects of pip-berry relationship, in both size and shape, on grapevine eco-evo-devo changes during domestication, and invites to explore further the functional ecology of grapevine pip and berry and notably the impact of cultivation practices and human selection on grapevine morphology.

Early Neolithic (ca. 5850-4500 cal BC) agricultural diffusion in the Western Mediterranean: An update of archaeobotanical data in SW France.

Farming economy was first introduced to the coastal areas of Southern France by Impressa groups (ca. 5850-5650 cal BC), originating from Italy, and subsequently spread to the hinterland by Cardial/Epicardial communities (ca. 5400-4500 cal BC). Fruit and seed remains preserved in archaeological sites provide direct evidence of the botanical resources cultivated and collected by these ancient social groups. But the transition from hunter-gathering to agricultural subsistence strategies is still poorly known in the area, due to insufficient and sometimes outdated archaeobotanical studies. Here we present new results and a critical review of all the available archaeobotanical data, in order to characterize food plant resources, cultivation practices and their variations in time and space. The archaeological dataset is composed of 19 sites (20 site/phases) mostly located in the Mediterranean lowlands. Our results demonstrate that farming economy of the Impressa groups was focused on the cultivation of hulled wheats, with only slight differences compared to their South Italian origins. The contribution of naked cereals increased in the Cardial/Epicardial agriculture, in agreement with the situation in other areas of the Western Mediterranean. The subsistence economy of hinterland sites seems to include a wider contribution of wild fruits and more limited contribution of crops. However, the poor evidence of cultivation activities in the hinterland is likely due first to the difficulties to find and excavate the sites and perform large-scale archaeobotanical sampling. It is likely that agriculture played a significant but variable role between sites and territories.

Palaeogenomic insights into the origins of French grapevine diversity.

The Eurasian grapevine (Vitis vinifera) has long been important for wine production as well as being a food source. Despite being clonally propagated, modern cultivars exhibit great morphological and genetic diversity, with thousands of varieties described in historic and contemporaneous records. Through historical accounts, some varieties can be traced to the Middle Ages, but the genetic relationships between ancient and modern vines remain unknown. We present target-enriched genome-wide sequencing data from 28 archaeological grape seeds dating to the Iron Age, Roman era and medieval period. When compared with domesticated and wild accessions, we found that the archaeological samples were closely related to western European cultivars used for winemaking today. We identified seeds with identical genetic signatures present at different Roman sites, as well as seeds sharing parent-offspring relationships with varieties grown today. Furthermore, we discovered that one seed dated to ~1100 CE was a genetic match to 'Savagnin Blanc', providing evidence for 900 years of uninterrupted vegetative propagation.

Bioarchaeological insights into the process of domestication of grapevine (Vitis vinifera L.) during Roman times in Southern France.

Grapevine (Vitis vinifera), one of the most important fruit species in the Classical Mediterranean world, is thought to have been domesticated first in South-Western Asia, during the Neolithic. However, the domestication process remains largely unknown. Crucial unanswered questions concern the duration of the process (rapid or slow?) and the related geographical area (single or multiple-origins?). Seeds from domesticated grapevine and from its wild ancestor are reported to differ according to shape. Our work aims, first, to confirm this difference and secondly to identify the extent of domestication in the grapes cultivated by Romans in Southern France during the period 50 BCE-500 CE. We had the opportunity to analyze uncharred waterlogged grape pips from 17 archaeological sites. Based on an extended reference sample of modern wild grapevines and cultivars our work shows that both subspecies can be discriminated using simple measurements. The elongation gradient of the pip's body and stalk may be regarded as an indicator of the strength of the selection pressures undergone by domesticated grapes. Grapevines cultivated during the Roman period included a mix of morphotypes comprising wild, intermediate and moderately selected domesticated forms. Our data point to a relative shift towards more selected types during the Roman period. Domestication of the grapevine appears to have been a slow process. This could result from the recurrent incorporation into cultivation of plants originating from sexual reproduction, when grape cultivation essentially relies on vegetative propagation.