Palaeodemography of the foraging to farming transition: insights from the Danube Gorges Mesolithic-Neolithic transformations
Paléodémographie de la transition chasseurs-cueilleurs agriculteurs : transformations Mésolithique-Néolithique dans les gorges du Danube
p. 113-131
Résumés
The diffusion of the farming way-of-life into environments occupied by Mesolithic hunter-gatherers in Europe has been associated with two major demographic events: the migrations of farmers originating from the Near-East and an unprecedented population increase, the “Neolithic Demographic Transition” (NDT). The Mesolithic-Neolithic transformations in the Danube Gorges provide a context of particular importance for tackling issues of Neolithization, due to its location, temporal depth, and highly contextualized osteo-anthropological record. This chapter compares complementary palaeodemographic proxies and bioarchaeological markers in order to assess the demographic response of local foragers to the Neolithic expansion. Interpreted together, these lines of evidence confirm the predictions of the NDT, and shed light on the relationships between subsistence intensification, sedentism and population growth, between migrations, cultural transmission and adaptations, and between dietary strategies, fertility and morbidity – i.e. on some mechanisms, benefits and costs of the farming transition – in the Central Balkans.
La diffusion du mode de vie agro-pastoral en Europe, notamment dans des milieux occupés par des chasseurs-cueilleurs du Mésolithique, est associée à deux événements démographiques majeurs : les migrations d’agriculteurs originaires du Proche-Orient et un accroissement sans précédent de la population, la « Transition Démographique Néolithique » (TDN). Les sites mésolithiques-néolithiques des gorges du Danube constituent un contexte particulièrement important pour aborder les questions de néolithisation, en raison de leur localisation, de leur profondeur temporelle et de la découverte d’une série ostéo-anthropologique unique. Ce chapitre compare différents marqueurs paléo-démographiques et bioarchéologiques afin d’analyser la réponse démographique des chasseurs-cueilleurs locaux à l’expansion du Néolithique dans la région. Interprétées ensembles, les diverses sources de données examinées confirment les prédictions de la TDN et informent sur les relations entre intensification du mode de subsistance, sédentarisation et croissance démographique, entre migrations, et processus de transmission et d’adaptations culturelles, et entre stratégies alimentaires, fertilité et morbidité au cours du Néolithique ancien dans les Balkans.
Entrées d’index
Mots-clés : transition démographique agricole, Mésolithique - Néolithique, gorges du Danube, Balkans centraux, C distribution des probabilités cumulées, indice de juvénilité, ADN ancien, strontium radiogénique, isotopes stables, état de santé
Keywords : agricultural demographic transition, Mesolithic – Neolithic, Danube Gorges, Central Balkans, C summed probability distribution, juvenility index, ancient DNA, strontium radiogenic, stable isotopes, health status
Remerciements
This chapter illustrates the large influence of Jean-Pierre Bocquet-Appel’s publications, lectures and discussions on the development of the scientific path and thoughts of researchers from the Laboratory for Bioarchaeology of Belgrade University, from Paleodemography theory and methods, the model of the agricultural expansion and bio-cultural adaptations, to the evolution of essential life history events and traits, such as reproduction and birth, motherhood and feeding strategies, or senescence.
Texte intégral
The paradigm of the Neolithic demographic transition (NDT)
1The emergence of the farming system and its sprawling expansion – the Neolithic Transition – induced major changes in the demographic and social structures of human societies and created a new platform for our biocultural evolution during the Holocene (Cohen 2009; Pinhasi and Stock 2011). Population dynamics are central to our understanding of the mechanisms beyond the transition to farming: the appearance of the agro-pastoral lifestyle has been associated with population agglomeration and with a considerable increase in anthropogenic remains (Deevey 1960; Bocquet-Appel and Bar-Yosef 2005), the demographic structure of hunters and farmers has long been observed to differ (Binford and Chasko 1976), and, whether cultural or demic, the expansion of the Neolithic way of life necessarily implied some migrations (Ammerman and Cavalli-Sforza 1971; Bellwood 2005; Fort 2015). As asserted by Bocquet-Appel (2001: 637), “the number of humans is at the heart of this so-called Neolithic Revolution, either as a cause or as an effect of its geographical expansion”.
2Methods to extract demographic information concerning prehistoric populations mainly relied on three types of data: cultural data, such as the density of archaeological sites or phases of occupation (Shennan et al. 2013); cemetery data, such as mortality profiles, growth rates, health status (Bocquet-Appel and Bar Yosef 2005); and genetic or biomolecular data, such as DNA, skeletal proxies for genetic information or isotopic analyses (Ammerman and Cavalli-Sforza 1971; Pinhasi and Pluciennik 2004; Borić and Price 2013). Relying on direct paleodemographic information – growth rates inferred from the proportion of immatures in a skeletal population – Bocquet-Appel has demonstrated that the emergence of agro-pastoral activities was related to a major event of worldwide population growth, caused by an important increase in females’ fertility (Bocquet-Appel 2002; 2011). For many regions, the analyses of the probability distribution of radiocarbon dates have further confirmed the unprecedented population growth (Downey et al. 2014) and have additionally evidenced frequent events of population collapse in the 100-500 years after the appearance of agro-pastoral practices (Shennan et al. 2013).
3Whether the Neolithic population growth has stimulated the subsistence shift or should be considered as a primary consequence of agro-pastoral innovations has been a long-standing debate and remains controversial (Boserup 1965; Cohen 2009). It is possible that subsistence innovations, resulting from the behavioral adaptations of sedentary, growing foraging communities, were selected because they contributed to the reproductive success during periods of climatic variability, and thus participated to increase population growth. To explain the positive effect of the agro-pastoral lifestyle on females’ fertility, Bocquet-Appel (2008) notably suggested, among others, that a set of factors may have collectively contributed to increase birth stacking: changes in mobility and work-load patterns associated with the sedentary lifestyle, an increase in carbohydrate-rich food consumption, a reduction in the duration of breastfeeding and changes in children allo-parenting practices with the village life. These young children in greater number were also the primary victims of the nutritional deficiencies caused by farming subsistence shift and of the appearance of new pathogens (Larsen 2003; Cohen 2008), which may have contributed to “buffer” the farming growth rate (Downey et al. 2014).
4In many regions, the demographic growth should have been one of the driving forces for the agro-pastoral geographic expansion. The farming system has indeed the advantage that it can be exported outside of the original heartland, modified to cope with the new environments, and hence considerably extended, including into regions where hunter-gatherers had their own niche (Zvelebil 2001). Genetic and cultural data demonstrated the role of migrations in the spread of the Neolithic (Fort 2015; Hofmanovà et al. 2016); beyond being a vector of Neolithization, migrations may have also contributed to the demographic transition by emptying areas of Neolithic development and, contrastingly, by stimulating the population growth in secondarily populated areas.
The Neolithic transition in the Central Balkans
5After several millennia of development in the Central Anatolian Plateau, the Neolithic way of life – materialized by a fully developed package (domesticated species, ceramics and ground stone technology, household organization, symbolic practices and aesthetics) – was quite suddenly dispersed into Europe, through Western Anatolia and the Balkan Peninsula from the mid-7th millennium BC (Krauß, 2011; Reingruber 2011). The spatial analyses of substantial sets of radiocarbon dates and cultural material data have indicated that a continental route took the Neolithic package to the Southern Balkans around 6,500-6,200 cal BC and rapidly spread through the Central Balkans and the Southern Pannonian plain around 6,100-6,000 cal BC, where it stabilized for a longer time until 5,500 cal BC (Brami and Zanotti 2015; Fort 2015; Blagojević et al. 2017). This expansion is chronologically associated with the onset and the end of the Rapid Climate Change, cooling conditions which culminate around 6,200 cal BC (the 8.2 cooling event; Weninger et al. 2014). Given the chronological priority of South-Eastern Europe in the spread of the Neolithic into Europe – a receptacle for its adaptation and transmission from Mediterranean to Temperate Europe – the reconstruction of underlying demographic mechanisms in this region has a prominent role in formulating and testing the models of the agro-pastoral expansion.
6The Early Neolithic sites in the Northern and Central Balkan region are related to the Starčevo-Körös-Criş cultural complex. This cultural complex is characterized by cultural similarities with the Southern Balkans and Western Anatolian Neolithic, in various aspects of the Neolithic package and by local specificities in flint industry, settlements and species representations (Garašanin 1982; Trigham 2000; Krauß, 2011). The analyses of the SCPD from substantial dataset from Starčevo sites indicated two significant occupation’s intensity increase ca 6,200-6,000 and 5,700-5,550 cal BC, followed by a significant decline confirming the broad pattern of the NDT (Porčić et al. 2016; Blagojević et al. 2017; Porčić et al. 2021). While there is evidence to suggest exchange of material and knowledge between Neolithic incomers and the Mesolithic groups in Western Anatolia and the Southern Balkans (Reingruber 2011), the Mesolithic presence remained elusive in comparison with the rich documentation available for the Danube Gorges region of the Central Balkans (e.g. Radovanović 1996; Bonsall et al. 2008; Borić 2011).
7Located in the southern fringes of the Carpathian Mountains, the Epipalaeolithic/Mesolithic – Neolithic complex of sites in the Danube Gorges (aka “Iron Gates”) usually refers to two distinct landscape settings: after the Pannonian plain, the Gorges stricto sensu (referred here as “the Inner Gorges”) and the “Downstream Area”, an environment more open on the Wallachian plain (fig. 1). After an Epipalaeolithic occupation in some rock-shelter sites (circa 13,000 – 9,500 cal BC), the human presence is documented on open-air sites in river terraces from the 10th millennium cal BC (Early Mesolithic period, ca. 9,500-7,300 cal BC). During the Late Mesolithic period (ca. 7,300-6,200 cal BC), foragers started to build trapezoidal-shaped buildings and were already probably at-least semi-sedentary (Radovanović 1996; Borić, 2011; Dimitrijević et al. 2016). Numerous palaeodietary studies have indicated that the local population substantially consumed fish, as well as wild games (e.g. Bonsall et al. 1997; Borić and Dimitrijević 2005; Jovanović et al. 2018).
1. Maps of the sites of Danube Gorges and the different regions of South-East Europe mentioned in this chapter. Regions: A. the Danube Gorges (in present day Serbia and Romania); B. Rhodopes-Balkans mountains and South of Wallachian Plain (in present day Bulgaria); C. Central Balkan and South of Pannonian Plain (in present day Serbia); D. North of Pannonian Plain (present day Hungary). Sites of the Danube Gorges: 1. Alibeg; 2. Padina; 3. Lepenski Vir; 4. Vlasac; 5. Cuina Turcului; 6. Climente II; 7. Razvrata; 8. Icoana; 9. Hajdučka Vodenica; 10. Ostrovul Banului; 11. Schela Cladovei; 12. Ostrovul Corbului; 13. Ajmana; 14. Velesnica

8At end of the 7th millennium cal BC (ca. 6,200-5,900 cal BC, Transformation/Early Neolithic period), the Gorges’ inhabitants developed intensive contacts with the Early Neolithic communities recently settled in the neighboring regions. This is archaeologically evidenced by the adoption of some Neolithic technologies, raw materials, and ornaments (Borić 2011). This period saw the complexification of trapezoidal buildings and the creation of unique artistic artifacts (Srejović 1969). Also, some novelties in the funerary practices during this period recall the Anatolian Neolithic sphere, such as burials of neonates under the buildings’ red-plastered floors at the Inner Gorges site of Lepenski Vir (Borić and Stefanović 2004).
9In the Downstream Area around 6,000 cal BC, some Early Neolithic sites display greater cultural affinities with the Starčevo groups than with Inner Gorges’ sites, notably funerary practices such as “multiple burials” with individuals in crouched position (Stalio 1986; Vasić 2008). Further important socio-cultural changes occurred in the Inner Gorges after ca. 6,000 cal BC (Early/Middle Neolithic period, ca. 5,900-5,500 cal BC) in terms of settlements (abandonment of the trapezoidal buildings, new types of domestic structures), material culture (intensified presence of Neolithic technologies), symbolic repertoires (Neolithic inhumations in crouched position; new ornaments and grave goods) and subsistence economy (Neolithic suite of domesticated animals; Borić 2011). Recent archaeobotanical studies also indicate that some micro-plant remains recovered from the dental calculus of individuals dated to the Transformation/Early Neolithic and Early/Middle Neolithic phases could have derived from cereals (Filipović et al. 2017; Jovanović et al. 2021).
A palaeodemographic approach of the Danube Gorges Mesolithic-Neolithic Transformations
10While the chronology of the Neolithic expansion into Europe is now well-documented, we still have little knowledge on how local foragers experienced the transition, and on the mutual influences between the two groups. Considering the temporal depth of the archaeological sequence and the continuity in human occupation, as well as the complex evidence for Mesolithic and Neolithic interactions, the archaeological context of the Danube Gorges plays a central role in the study of Neolithization. These sites are also unique with more than 500 individuals remains, spanning the whole chronological sequence (Roksandić 2000). This skeletal record has provided researchers the opportunity to explore different palaeodemographic issues related to migrations, fertility, mortality, nutritional health and morbidity (Nemeskéri 1978; Grga 1996; Bonsall et al. 1997; Roksandić 2000; Jackes et al. 2008). Today, the constant reassessment of graves’ chronological assignment through the publication of new radiocarbon dates (e.g. Borić 2011; Bonsall et al. 2015b) enables us to reconsider the results of these studies, for instance, concerning health (Radović and Stefanović 2013; Jovanović 2017). Also, the application of isotopic and ancient DNA analyses have recently shed new lights on Mesolithic-Neolithic demographic transformations (Bonsall et al. 1997; Borić and Price 2013; Mathieson et al. 2018; Jovanović et al. 2018). This chapter reviews these complementary bioarchaeological lines of evidence in light of the new available chronological information to assess the local experience and the demographic response of Central Balkans’ foragers to the Neolithic expansion. Particularly, we synthesize the results of:
- New SCPD analyses as a chronological proxy for population dynamics.
- A chronological reassessment of the juvenility index, as indicator of the Neolithic Demographic Transition; Available morphological (skeletal biodistance studies), isotopic (strontium radiogenic), and genetic (ancient DNA) evidence about migrations.
- Available biochemical (stable isotopes) and macroscopic (caries, and non-specific bone and teeth markers of physiological stress) information about dietary adaptations and health status.
1. Radiocarbon dates, intensity of occupation and population dynamics
11During the past 40 years of research on the Danube Gorges, a long-lasting debate focused on the interpretation of the chronological sequence, required condition to understand the origin of farming in the region (e.g. Srejović 1969; Garašanin and Radovanović 2001; Borić et al. 2018). Large radiocarbon dating programs undertaken at the end of the 1990’s have confirmed the very long span of the Epipalaeolithic-Mesolithic-Neolithic human occupation of the Gorges, clarified the chrono-stratigraphy of settlements and refined the chronology of the adoption of different cultural components of the Neolithic package (Cook et al. 2002; Borić 2011; Bonsall et al. 2015b; Borić et al. 2018). Beyond chronological advances, the local radiocarbon record may also provide valuable information related to the long-term demographic patterns. Based on the assumption that one can use “dates as data“, summing the probability distribution of different radiocarbon dates (SCPD) at a given site should inform about the most probable period(s) of occupation. Consequently, summing similar information for numerous phases and sites at the regional level should draw a rough picture of the occupation’s intensity fluctuation, in other words, periods of population growth and/or decline (Shennan et al. 2013). Bonsall et al. (2015a) reconstructed SCPD on the Danube Gorges radiocarbon dataset to specifically explore the local human response to climatic oscillations and local hydrological fluctuation. They found marked discontinuities in the SCPD curves circa 7,500-7,000, 6,700-6,000 and after 5,800 cal BC, corresponding to major climatic anomalies: the 9.3 and 8.2 ky cold events, and possibly colder temperatures circa 5,800-5,500 cal BC. Thus, they suggested that dips in the local SCPD curves should “reflect periods of higher annual river discharge and an increase in flood magnitude during Holocene “neoglacial” events, associated with generally cooler, wetter conditions in the Danube catchment upstream of the Iron Gates, and that the increased flood risk led to a reduction in the intensity with which people used certain sites or the lower parts of sites bordering the river” (Bonsall et al. 2015a: 6).
12However, the version of the SCPD method used by Bonsall et al. (2015a) did not account for sampling bias – an important issue in the Danube Gorges – or taphonomy, and did not provide a statistical test for the significance of the observed patterns. To address these issues, we used the SCPD approach formulated by Shennan et al. (2013) and Timpson et al. (2014)1. In addition to applying a more rigorous statistical analysis, we also include 31 new radiocarbon dates (Penezić et al. 2020; Porčić et al. 2021; Jovanović et al. 2021; Jovanović et al. in prep.). A total of 312 audited dates from 12 Danube Gorges sites is included in the analysis. A correction for the freshwater reservoir effect was performed for all the samples with δ15N values above the threshold following Cook et al. (2002).
2. Statistical tests of the SCPD of the Danube Gorges radiocarbon dataset.

13In general, the SCPD curve shows significant deviations from the null model (global p<0.001; fig. 2). After the onset of the Holocene, the SCPD curve begins to increase slowly and irregularly, suggesting the presence of small and fluctuating population size in the Gorges, which is broadly consistent with archaeological information (Radovanović 1996; Borić 2011). The first significant drop can be observed between 9,200 and 8,800 cal BC and might coincide with a slight decrease in the NGRIP δ18O palaeoclimate proxy (Bronk Ramsey 2009). The drop associated with the 9.3 ky cold event is less pronounced than in Bonsall et al. (2015a) and it is not statistically significant (p= 0.276) according to the specific SCPD test (Edinborough et al. 2017). The intensity of occupation as reflected by radiocarbon dates then statistically increases again after 7,000 cal BC, reaching its maximum ~6,500 BC (6,700-6,400 cal BC). Changes in the patterns of occupation of the sites during this period are also supported by the discovery of the earliest forms of trapezoidal buildings at sites such as Vlasac and Schela Cladovei, and of the first forms of red-plastered floors at Vlasac (Borić 2011). This episode of intense occupation is followed by a rapid and significant decline after 6,500 BC (ca 6,400-6,200 cal BC; specific SCPD test p=0.0218), roughly coinciding with the Hudson Bay Rapid Climate Change (6,600-6,000 cal BC) and particularly with the beginning of the sharp cooling episode of the 8.2 ka event (Weninger et al. 2014). Although the possibility that this drop in the SCPD curve could have been caused by a dramatic demographic crisis cannot be rejected (pathogen?), this scenario appears quite unlikely in the light of the age structure of the buried population and of available information on health status (cf infra part 2 and 4). A simpler interpretation might imply the possible departure of part of the local (semi-)sedentary population to settlements located outside, in the immediate vicinity of the Danube banks (or temporary return to a more nomadic lifestyle?). Such depopulating could have been caused by climatic oscillations and increased risks of floods, by temporary changes in the availability of some fish species, and/or by particular social tension or structural/territorial reorganization, perhaps in relation with the increased demographic expansion of the first Early Neolithic groups settled in the south of the Balkan Peninsula.
3. Comparison of the SCPD of the Danube Gorges with the SCPD of neighboring regions presented in figure 1 (results of Porčić et al. 2016 and Blagojević et al. 2017).

14The main increase in the curve occurred around 6,200-5,900 cal BC, during the Transformation-Early Neolithic phase in the Gorges, concomitantly with the arrival of Early agro-pastoralist communities in the surrounding areas of the Central Balkans (figs. 2 and 3; Porčić et al. 2016; Blagojević et al. 2017; Porčić et al. 2021). This phase represents the apex of Lepenski Vir culture which reflects an original cultural identity featuring at some sites unique architectural elaborations, artistic creations and symbolic ways of expressions, and Mesolithic-Neolithic cultural hybritization (Borić 2011; Borić et al. 2018). The contacts between these two worlds should have thus contributed to intensify the occupation at localities such as Lepenski Vir. The foragers’ knowledge of local environmental conditions may have represented an important advantage for agro-pastoralists’ adaptations to the new biomes of Temperate Europe, which may have contributed to the fast rate of the Neolithic spread into the Central Balkans.
15However, while this “contact phase” may have stimulated the growth of population grouped at some localities, the subsequent Early Neolithic demographic expansion in the surrounding regions of the Balkans ca 5,900-5,500 cal BC coincides with a gradual decline in the intensity of human occupation in the Danube Gorges (fig. 3). Although further socio-cultural changes occurred in the region after 6,000 cal BC, including the adoption of animal husbandry, the Gorges environment may not have been suitable (rugged topography) or attractive (new Neolithic social ethos) enough for the demographically expanding Starčevo groups and their agro-pastoral practices. After 5,500 cal BC, the entire Early Neolithic population of the Central Balkans also experienced a rapid demographic collapse, either caused by important emigrations or by some catastrophic events (fig. 3; Porčić et al. 2016; Blagojević et al. 2017; Porčić et al. 2021).
2. Mortality profiles, growth rates and female fertility
16The evidence for an intense occupation of the Gorges during the 7th millennium BC calls into question the possibility of an important local population growth prior to the adoption of agricultural practices. Several authors applied diverse palaeodemographic indices on the grouped osteoanthropological sample and inferred a rather stable and stationary population with both low fertility and low mortality, comparable to other European Mesolithic population (Meiklejhon et al. 1997; Bocquet-Appel 2002). Jackes et al. (2008) further examined these data per sites and chronological periods, adding samples from sites located in the Downstream Area (Ajmana and Velesnica). Their results confirmed previous observations for the Mesolithic period, and indicated a fall in fertility at Lepenski Vir during the time of the Mesolithic-Neolithic contact, interpreted either in relation to an over-representation of adults due to the influx of adults from outside or as an actual effect of changing and unstable conditions. In contrast, the grouped Neolithic rather indicate an increase in fertility (Lepenski Vir Early Middle Neolithic and Downstream Early Neolithic sites), consistent with the assumptions of the NDT. The new radiocarbon dates and the reassessment of chrono-cultural sequences has allowed us to re-assign individuals to different periods and to re-interpret these results. To limit the “squeezing effect” which may result from averaging data for numerous generations or for sites with different mortuary practices, we first inferred growth and birth rates from the juvenility index (the proportion of children aged 5-19 to the whole population; Bocquet-Appel 2002) calculated per sites’ chronological phases and then per broader chrono-geographical groups (Table 1 and fig. 4)2.
4. Growth rates and 95% confidence intervals inferred from the juvenility index (15p5) for the Mesolithic-Neolithic sites of the Gorges and comparison with Mesolithic (n=9) and Neolithic (n=91) European populations (data from Bocquet Appel 2001 and Downey et al. 2014). The cross refers to the growth rate value inferred for the Tranformational period of Lepenski Vir from house-floor-based Bayesian demographic methods (Porčić and Nikolić 2014).

Table 1. Growth and birth rates inferred from the juvenility index (15P5), computed using the proportions of individuals aged 5-19 years old and more than 20 years old, per sites and archaeological phases.
Phase - Sites | n 5-19 | n 20+ | 15P5 | Growth rate (95% confidence interval) | Birth rate (95% confidence interval) |
Early Meso. Lep. Vir | 3 | 14 | 0.17 | 0.001 (-0.009 – 0.01) | 0.036 (0.03 – 0.04) |
Early Meso. Padina | 2 | 41 | 0.0046 | -0.025 (-0.035 – -0.014) | 0.014 (0.08 – 0.020) |
Late Meso. - Trans. Vlasac | 14 | 127 | 0.099 | -0.01 (-0.02 – -0.001) | 0.023 (-0.023 – -0.002) |
Late Meso. - Trans. Haj. Vod. | 5 | 40 | 0.1 | -0.01 (-0.02 – 0.007) | 0.025 (0.019 – 0.031) |
Trans. Lep. Vir | 12 | 62 | 0.16 | -0.001 (-0.01 – 0.094) | 0.034 (0.028 – 0.040) |
Early-mid. Neo. Lep. Vir | 11 | 36 | 0.23 | 0.009 (-0.0018 – 0.002) | 0.046 (0.040 – 0.051) |
Early Neo. Ajmana (Downstr.) | 7 | 7 | 0.5 | 0.036 (0.025 – 0.047) | 0.086 (0.080 – 0.092) |
Early Neo. Velesn. (Downstr.) | 3 | 5 | 0.375 | 0.025 (0.014 – 0.035) | 0.068 (0.061 – 0.073) |
17Reassessed growth and birth rates confirm that the Mesolithic population was rather stable and stationary (Table 1 and fig. 4). The particularly low growth rate inferred for the Early Mesolithic period could result from the habitual mobility patterns of foragers communities using these locations as burials and/or residential sites, and to a biased representation of different age classes possibly caused by higher level of daily or seasonal mobility. Differential models of mobility and of territories occupations could indeed influence the representativeness of skeletal data either in relation to the place where the death occurred or because of associated mortuary practices, and one should keep in mind that the “squeezing” generational effect should be higher for long-term Mesolithic phases of occupation. These results also suggest that the apparent Late Mesolithic population increase reflected by the SCPD should rather be related to a change in the occupation patterns than to a demographic process internal to the population. Possible explanations include an increase in the number of localities used simultaneously for fishing activities, for instance, changes in the organization of subsistence and daily activities, changes in the territorial structuration and social organization perhaps related to an increased sedentism. Although excluded from these palaeodemographic indicators for methodological concerns, remains of numerous neonates have been recovered associated with adults in Late Mesolithic contexts3 (Borić and Stefanović 2004; Stefanović and Borić 2008). An even higher number of neonates have been uncovered in the Transformational phase of Lepenski Vir below the house-floors4, a cultural practice which points of the Anatolian and Southern Balkans Neolithic sphere (ibidem). Therefore, the deposition of these babies below the house-floors could reflect some modification of reproductive behavior and/or young children mortality rates which took place at the time of the first contacts with the Early Neolithic communities.
18We did not find the drop in fertility observed by Jackes et al. (2008) for the Transformational period of occupation of Lepenski Vir: the growth and birth rates appear slightly higher than inferred for other Late Mesolithic-Transformational contexts, but still range in the lower part of European Mesolithic population variability (Table 1 and fig. 4). Thus, the important occurrence of neonates at Lepenski Vir might be related to their better preservation due to new mortuary rituals, and perhaps also to the site-specific symbolic nature. An increase in females’ fertility and thus neonatal mortality could also produce such a pattern. Porčić and Nikolić (2015) recently estimated growth rate for the period of Transformations/Early Neolithic at the site of Lepenski Vir by combining the archaeological settlement evidence (data on house floor areas and stratigraphic information) and ethnoarchaeological information (floor area-to-household size, house-use life, hunter gatherer group size) with mathematical models of house accumulation and population growth within the Approximate Bayesian Computation framework. According to the authors, the initial population at Lepenski Vir must have been low, within the range of 5-20 people, while the final population size corresponds to ethnographically documented village size of people dependent on aquatic resources (30-40 people; estimated growth rate: ca 0.01-0.02; fig. 4). The growth rates estimated between skeletal data (-0.007-0.014) consistently overlap with house-floor inference (0.004–0.02). The slight difference between the estimated ranges might simply be related to the differential resolution and nature of these indicators: the growth rate calculated from skeletal data groupings should be tightly correlated to birth rates, and could be affected by mortuary practices (differential eligibility of some parts of the population to different burial places), whereas the growth rates based on archaeological stratigraphic information may be affected by alternate demographic or cultural factors, such as migrations or changes in the function or use of the houses.
19While Jackes et al. (2008) pooled individuals from the sites of the Downstream Area (samples of Ajmana and Velesnica) with the Early-Middle Neolithic assemblage of Lepenski Vir, recent radiocarbon dates suggest that they might be earlier (Velesnica: ca 6,215-5,845 cal BC; Ajmana: 6075-5715 cal BC; Bonsall et al. 2015b). Palaeodemographic indicators reconstructed for these Early Neolithic sites located downstream suggest higher fertility and childhood mortality rates (Table 1 and fig. 4). Although these two samples are smaller in size (total number of individuals = 26), these different demographic parameters coincide with the notable cultural differences with the sites of the Inner Gorges, including funerary patterns (Stalio 1986; Vasić 2008). One cannot disregard the possibility that the inferred growth rate might be affected by specific mortuary practices for subadults. However, these results coincide with the broad differences observed between European Mesolithic and Neolithic demographic patterns (fig. 4) and fits with the results of the SCPD analyses of Mesolithic and Early Neolithic occupation in the Central Balkans (fig. 3). Because demographic models and mortuary practices may not be entirely unrelated, the differential birth and childhood mortality rates, as observed amongst European Mesolithic and Neolithic societies, may also influence the community idea about household and parenting, the social roles of the youngest amongst the community, and possibly their places amongst the deaths of the group. Thus, the peak in the Danube Gorges SCPD curve identified around 6,200-5,900 cal BC could be explained by the attractiveness of Lepenski Vir in the Inner Gorges and by the local presence of Early Neolithic communities with higher growth rates and probably different social organization. However, the growth rate inferred for the Transformational and Neolithic phases of Lepenski Vir (respective average ca -0.001 and 0.009) still range in the lower part of European Neolithic farmers’ variability and coincides with the average value for European Mesolithic hunter-gatherers. As also suggested by the gradual drop noticed in the SCPD curve between 5,900-5,500 cal BC, the demography of local foragers may not have been sustained with the expansion of agro-pastoral populations of Pannonia and of the Central Balkans valleys, and the consequent marginalization of the Gorges.
3. Early Neolithic migrations, interactions, and admixture
20Besides the changes in the population age structure, the osteoarcheological sample provided a direct opportunity to explore migrations and admixture issues and test Neolithic diffusion models. In the first studies conducted on the osteoanthropological assemblages from the Gorges, some authors observed a temporal trend towards gracilization interpreted as the effect of new dietary adaptations and/or of migrations (Mikić 1981; Menk and Nemeskéri 1989). Particularly, individuals buried at Downstream Area sites (Velesnica and Ajmana) display more gracility than most Mesolithic individuals buried at Inner Gorges’ sites but are comparable to some individuals discovered in the Neolithic layers of Lepenski Vir (Živanović et al. 1986; Radosavljević-Krunić 1986). Roksandić formalized these observations further through a biodistance study of individuals buried at different sites of the Inner Gorges (Roksandić 2000). Her results indicate a local biological continuity in the region with a gradual temporal ordering and a more pronounced difference at the time of contact (Roksandić 2000). The chronological re-assessment of these results, as well as Principal Component Analyses additionally performed on a set of cranial measurements, have also shown that the Early Neolithic individuals buried in the Downstream Area range on the margin of the local Mesolithic-Neolithic distribution (de Becdelièvre et al. 2015).
5. A: frequency of Danube Gorges prehistoric individuals from different mitochondrial haplotypes (from Hofmanová 2017; González-Fortes et al. 2017; Mathieson et al. 2018). B: frequency of Danube Gorges prehistoric individuals with a local or non-local strontium radiogenic signal (Borić and Price 2013); C: number of locals and non-locals assigned to U or non-U maternal lineages.

21The results of strontium radiogenic (87Sr/86Sr) and palaeogenetic (aDNA) studies have provided further insight into the causes for this morphological temporal variation pattern (Borić and Price 2013; Hofmanová 2017; Mathieson et al. 2018). Strontium radiogenic analyses have directly documented a dramatic increase in the number of individuals of non-local origin (first generation migrants), mostly females, buried in the Danube Gorges at two sites – Lepenski Vir (Inner Gorges) and Ajmana (Downstream Area) – during the period of Transformation-Early Neolithic, and during the Early/Middle Neolithic period (fig. 5B and C; Borić and Price 2013). Palaeogenomic studies have recently evidenced that an important proportion of individuals buried at Lepenski Vir during the Transformational and Neolithic phase belong to maternal clades which were found amongst Near Eastern and European individuals, but which were mostly absent amongst analyzed genetic sequences of European Mesolithic populations (fig. 5A; Hofmanová 2017; González-Fortes et al. 2017; Mathieson et al. 2018). Most individuals of non-local origins belong to Near-Eastern Neolithic-like maternal haplogroups. These observations have been confirmed on formal genetic distance analyses and additional Bayesian biostatistical analyses have also confirmed that a few individuals buried in the Gorges had an admixed European Mesolithic-like and Near-Eastern Neolithic-like ancestry (Hofmanová 2017; Mathieson et al. 2018). Foragers’ interactions with Early Neolithic communities settled the downstream Wallachian plain and the southern Pannonian plain, and the presence of migrants descended from Anatolian Neolithic groups may thus explain the increased density of occupation observed in the Gorges ca 6,200-5,900 cal BC at the time of the flourishment of the Lepenski Vir culture.
22Finally, the possibility that these interactions may have led to an increased pattern of social tension and inter-group conflicts has been explored through the examination of traumatic lesions on skeletal remains (fractures and projectiles injuries; Roksandić et al. 2006). The individuals who bear traces of wounds were mostly dated to the Late Mesolithic period (14 Late Mesolithic out of 16 individuals with traces of wounds), indicating that these traumas cannot have resulted from violent foragers/farmers oppositions but rather from sporadic episodes of interpersonal violence, maybe in relation to the increased density of occupation of the region during the first half of the 7th millennium BC.
4. Behavioural interactions, adaptations and the mechanisms of the NDT
23The information provided by numerous paleodietary studies and by the analyses of nutritional and physiological health markers conducted on the Mesolithic-Neolithic sample from the Danube Gorges provides the opportunity to consider the relationships between population dynamics and subsistence adaptations, and to examine locally some behavioral mechanisms beyond the NDT.
24Stable isotopes studies have confirmed that fishing played a central role in the local ways of subsistence during the whole Mesolithic period (Jovanović et al. 2018). Various oral health studies have also suggested the greater consumption of protein-rich food, over carbohydrate-rich resources (low incidence of caries; high rates of dental calculus and high degree of teeth abrasion; e.g. Grga 1996; Radović and Stefanović 2013; Jovanović 2017). Some scholars have related the elevated rates of some skeletal markers of essential nutrients deficiencies to the possible presence of local aquatic food-borne parasitic infections (high frequency of porotic hyperostosis and presence of cribra orbitalia ; Meiklejohn and Zvelebil 1991; Jovanović 2017). More specifically, the observation of a shift up in δ34S values suggests that anadromous fish should have become particularly important in the local subsistence after ca 7,000 cal BC, as a possible result of change in the availability of local species and/or in fishing strategies (fig. 6A; Jovanović et al. 2018; de Becdelièvre et al. 2020a). Therefore, we can suggest a relationship between the adaptation of subsistence strategies towards an intensified-specialized fishing and the intensified occupation reflected by the SCPD ca 7,000-6,500 cal BC, at the time when the earliest forms of trapezoidal buildings appear along the Danube. During warmer and better climate periods, a change in patterns of residential mobility amongst these semi-sedentary communities could have been associated with an intensified exploitation of local resources, the development of specialized fishing strategies, population agglomeration and increased density of occupation at some localities along the Gorges.
6. A: sulfur and nitrogen stable isotope ratios (δ34S, δ15N) for Mesolithic-Neolithic individuals from the Central Balkans (aged >10 years old) and prehistoric animals discovered in the Danube Gorges (data from Nehlich et al. 2009; Borić 2011; Jovanović et al. 2018; de Becdelièvre et al. 2020a). B: carbon stable isotope “weaning trendlines” for Transformational-Early Neolithic children from the Inner Gorges and the Downstream Area (smoothed δ13C values obtained on several dental tissues using a Loess statistical procedure; adapted from de Becdelièvre et al. 2020a).

25Zooarchaeological studies have documented a continuity in the local ways of subsistence during the period of Transformations and the Neolithic; animal husbandry has indeed only been introduced after 6,000/5,900 cal BC in the Inner Gorges, where it likely remained subsidiary to fishing and hunting and played a minor role than at most surrounding Starčevo settlements (Bartosiewicz et al. 2001; Borić and Dimitrijević 2006). While crop remains have been found at numerous Early Neolithic sites of the Central Balkans, the documentation remains scarce concerning sites of the Gorges (Filipović et al. 2017; Jovanović et al. 2021). In accordance with the evidence for Early Neolithic migrations, the stable isotope analyses of Transformational – Neolithic individuals have revealed a broadening of the local variability towards lower values, indicating that some individuals consumed substantially fewer aquatic resources (fig. 6A; Bonsall et al. 1997; Jovanović et al. 2018). While individuals buried at most sites of the Inner Gorges exhibit high δ15N ratios in continuity with the previous period of occupations (high aquatic resources consumption), some buried at the central site of Lepenski Vir in the Inner Gorges display lower values, similar to all individuals buried in the Downstream Area (Ajmana and Velesnica) and to those discovered in other Starčevo contexts from the broader Central Balkan regions (fig. 6B; Jovanović et al. 2018; de Becdelièvre et al. 2020a). The individuals with lower δ15N and δ34S values whose strontium radiogenic signal was analyzed have been identified as first-generation migrants: they likely originated from communities with a mixed farming-foraging subsistence economy (de Becdelièvre et al. 2020b). In contrast, all their putative descendants – the individuals with a local strontium signature born in the region of the Gorges but belonging to a Near-Eastern Neolithic-like lineages – exhibit elevated stable isotope ratios, which confirm their adaptation to the local ways of subsistence (ibidem).
26Intra-individual stable isotope analyses (the longitudinal analyses of different tissues formed between birth and teenage) have also shown significant differences in feeding strategies between the Mesolithic and Transformational children buried in the Inner Gorges and the Early Neolithic children buried downstream (fig. 6B). The later experienced a more abrupt transition to solid food and were supplemented with preparations made of lower trophic-level resources than the former whose weaning preparations likely contained aquatic resources (de Becdelièvre 2020a). The children buried at the downstream site of Ajmana are also the only one to display evidence of dental caries, only observed in the Gorges on a few Transformational-Early-Middle Neolithic adults discovered at Lepenski Vir and Ajmana (Jovanović et al. 2017; de Becdelièvre 2020a). With cereal-based weaning preparations (carbohydrate-rich food resources), deciduous teeth should have been particularly at risks for dental caries. Besides, although the occurrence of enamel hypoplasia, caused by events of physiological stress experienced during childhood, remains low in the Gorges throughout the Mesolithic-Neolithic sequence, a greater frequency of affected children has been observed at the Early Neolithic sample of Ajmana, downstream (ibidem). The skeletal examination of Early Neolithic discovered in the Broader Central Balkans and Southern Pannonian Plain has also revealed higher number of dental caries and of skeletal markers of nutritional deficiencies, confirming the pattern reported for several other Near-Eastern and European early farming communities (Jovanović 2017).
27These results are in the line with some of the assumptions for the causes of the NDT, which posit that differences in terms of children feeding practices may contribute to the reported demographic differences between foragers and farmers (Bocquet-Appel 2008): the presence of potential new weaning strategies and technologies may have facilitated an earlier decrease in the intensity and the frequency of suckles and released the Neolithic mothers from the energetic burden of lactation, increasing fertility rates; in turn, these new feeding practices may not have been optimal for children oral and physiological health.
28The analyses of anomalies on the annual deposits of cementum of Mesolithic and Neolithic individuals from the Danube Gorges and the broader Central Balkans region have also reported that Neolithic individuals experienced more frequent episode of stress during adulthood (Penezić et al. 2020). This tendency was statistically significant only for females which, combined with the fact that pregnancies are one of the major causes of stress layer formation in tooth cementum, might also be related to the Early Neolithic increased fertility in the region. In the trade-off with individual fitness (life-expectancy and well-being), it seems that the expansion of the farming niche dramatically favored populational fitness (reproductive success; Lambert 2009).
Conclusions: Neolithized foragers and Mesolithized farmers, an original case-study for palaeodemographic research
29Demography plays a central role in all aspects of Anthropology, from the study of populating patterns and cultural transmission processes to the mechanisms of evolution and social adaptations. Research have shown that the onset of the farming system induced an unprecedented demographic growth, both triggering and stimulated by a sprawling geographic expansion, migrations, cultural transmission, and ecological adaptations. The archaeological complex of the Danube Gorges has provided a unique opportunity to reconstruct long-term trends in demographic fluctuations in the Central Balkans, a key region for our understanding of the Neolithization of Europe. Analyses of the radiocarbon dataset indicate an increase in the intensity of human occupation in the Gorges during the mid-7th millennium BC, which might be associated with a change in settlements patterns of occupation and a reduction of residential mobility. Interpreted in the light of stable isotope values and archaeozoological data, this intensified occupation of some localities may be related to an intensified exploitation of aquatic resources and the development of more specialized fishing strategies. This period of intense occupation was followed by an important decline (ca 6,400-6,200 cal BC), which coincides with the harsher conditions of the Hudson Bay Rapid Climate change, and perhaps with the departure of part of the local (semi-)sedentary population to more distant settlements.
30The main increase in the intensity of occupation dates to the end of the 7th millennium BC and is concomitant with the arrival and demographic increase of the first farming communities in the broader Central Balkans, and the adoption of ceramics in the Gorges. Genetic analyses revealed that individuals genetically closer to Near-Eastern Neolithic communities were buried at Lepenski Vir near individuals genetically closer to European Mesolithic individuals. Also, there is genetic evidence for admixture. During this period Lepenski Vir may have been an attractive site for a broader population, and a central place for contacts, exchanges, and interactions between the foragers of the Gorges and the first agro-pastoralists settled in surrounding regions.
31At the same time, bioanthropological and cultural material data suggest that the settlements located in the Downstream Area may have represented some Starčevo Neolithic cultural enclaves which may have played a role in the diffusion of Neolithic practices in the Inner Gorges. The growth rates inferred from the Transformational and Neolithic skeletal assemblages of Lepenski Vir are rather similar as data available for European Mesolithic population; in contrast, the demographic parameters inferred from the Early Neolithic osteo-archaeological assemblages of the Downstream Area are at the higher end of the prehistoric agricultural societies’ variability. All individuals discovered in this area exhibit lower stable isotopes values than individual buried in the Inner Gorges. Children dental isotope analyses also demonstrated that their feeding strategies significantly differed, with higher rates of dental caries and greater frequency of enamel hypoplasia. It is possible that such different dietary behaviors may have differently affected some demographic parameters – females’ fertility and children health – of the Danube Gorges foragers and of Central Balkans Early Neolithic communities.
32After 5,900 cal BC, the intensity of human occupation in the Gorges gradually declined over the 6th millennium BC, at the time when the Early Neolithic demographic expansion in neighboring regions was likely fueled by the high fertility rates of the first agro-pastoralists. Further larger generations of migrants were discovered at Lepenski Vir and probably brought different socio-cultural Neolithic novelties. Yet, agro-pastoral activities remained rather subsidiary in the local ways of subsistence in comparison with fishing and hunting. The general pattern of health decline usually associated with the onset of the farming system is not documented in the region of the Danube Gorges: the important consumption of micro-nutrient and protein-rich food resources probably contributed to buffer the adverse effect of the Neolithic Transition, which contrasts with the situation observed at other Early Neolithic contexts of the Central Balkans where the greater occurrence of physiological and nutritional markers of stress could be a by-product of niche colonization, early farming practices, diet, and elevated fertility rates. At that time, it is possible that the environment of the Gorges was less suitable for the agropastoral practices of Starčevo people, and that the former foragers’ way of living became increasingly forsaken by the appearing Neolithic social ethos.
33The resolution of (bio-)archaeological information now available from the region of the Danube Gorges has shed lights on some local mechanisms of the NDT. The same relationships which have been observed between reduced residential mobility and intensified exploitation of local food resources amongst Near-Eastern Epipalaeolithic hunter-gatherers may have also sporadically caused similar increases in the size of some groups of European Mesolithic foragers, in the Gorges, the Dniepr, along the Atlantic, North, and Baltic seas coasts. Yet, as exemplified by the Late Mesolithic inhabitants of the Danube Gorges, it is possible that the extractive way-of-subsistence may have not be enough to sustain larger-size population of foragers for long period of time, and the comparison of the SCPD with paleoclimatic proxies suggests that the demography of these larger groups of sedentary foragers should have been quite dependent on climatic oscillations.
34The population dynamics inferred at the onset of the agro-pastoral way of life in the Central Balkans is consistent with the predictions of the NDT, as there is evidence for Neolithic migrations followed by population growth which preceded a demographic decline. While the first contacts between foragers and first agro-pastoral communities should have contributed to stimulating population growth in areas populated by foragers, the presence of foragers may have also played a role in the rapid geographic expansion and adaptation of the Neolithic package in the region. Although there is evidence for greater proportions of admixture with foragers in the Central Balkans than in the regions previously populated by Early farmers, their biological contribution to future generations of Neolithic farmers remained limited, as observed in other regions of South-Eastern, Central and Western Europe. This situation likely contrasts with Northern and Eastern European regions where interactions may have lasted longer or where different aspects of the Neolithic were gradually adopted without significant demographic contribution from migrants.
35Notable differences in terms of children feeding strategies between the Mesolithic – Neolithic foragers of the Inner Gorges and the Early Neolithic communities settled in the Downstream Area could have differently influenced some aspects of females’ metabolism or reproductive development, as well as childhood health. Such differences in parenting strategies may have also contributed to the rapid Neolithic demographic expansion in the region. The analyses of children feeding practices amongst other transitional populations could allow further testing. In the Central Balkans, it is possible that local foragers’ demographic behavior could not sustain high population density facing the explosive demography of the first agro-pastoral communities.
36Conducting further research on the demographic dynamics during periods of subsistence transition may contribute to deepen our understanding of the mechanisms of human adaptability in relation with key events of niche construction and colonization. Far from saying “Farewell to palaeodemography” (Bocquet-Appel and Masset 1982), welcoming diverse bioarcheological and biomolecular approaches in paleodemographic studies has provided a wealth of comparable data that allows not only for reconstructing population dynamics, but also for exploring the causes and the consequences of historical demographic cycles, an essential knowledge to contemplate better paths towards sustainability in a connected and globalized world with billions of people.
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10.4312/dp.28.1 :Notes de bas de page
1 This version of the SCPD method accounts for the research (sampling) bias by applying a binning procedure (we used 100 radiocarbon years as a threshold, resulting in 102 bins). The effects of taphonomy are accounted for by setting the taphonomic model as the null model for the Monte Carlo simulation. We used the exponential decay model published in Surovell et al. (2009) as the taphonomic model – the null model that assumes stationary population size with exponential effects of taphonomic loss on the SCPD curve (see Porčić et al. 2016 for details). The method also provides a test for the statistical significance of the deviations of the empirical SCPD from the null model by means of Monte Carlo simulation (10000 iterations). The method is implemented in the R (version 3.6.3), using the Rcarbon package (Crema and Bevan 2020).
2 The age of children was reassessed based on dental development when possible or the length of long bones (de Becdelièvre et al. 2020a). We followed the procedure of MNI calculation developed by Jackes et al. (2008), just re-assessing the final accounts by-assigning individuals to different periods (using chronological information from Borić and Price 2013; Borić et al. 2014; Bonsall et al. 2015b; Stefanović 2016; Borić et al. 2018; data in de Becdelièvre et al. 2020a).
3 A proportion of 28 foetus and neonates for 127 Late Mesolithic – Transformational adults (mostly dated to the Late Mesolithic period – Borić and Stefanović 2004; Borić et al. 2014).
4 A proportion of 41 neonates for 62 adults (Borić and Stefanović 2004; Stefanović 2016)
Auteurs
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Camille de Becdelièvre
Laboratory for Bioarchaeology, Faculty of Philosophy, University of Belgrade, Serbia
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Tamara Blagojević
Biosense Institute, University of Novi Sad, Serbia
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Jelena Jovanović
Laboratory for Bioarchaeology, Faculty of Philosophy, University of Belgrade, Serbia – Biosense Institute, University of Novi Sad, Serbia
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Sofia Stefanović
Biosense Institute, University of Novi Sad, Serbia – Laboratory for Bioarchaeology, Faculty of Philosophy, University of Belgrade, Serbia
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Zuzana Hofmanová
Department of Archaeogenetics, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany – Department of Archaeology and Museology, Masaryk University, Brno, Czechia Republic
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Marko Porčić
Laboratory for Bioarchaeology, Faculty of Philosophy, University of Belgrade, Serbia – Biosense Institute, University of Novi Sad, Serbia
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