Early life conditions play an important role in determining adult body size. In particular, childhood malnutrition and disease can elicit growth delays and affect adult body size if severe or prolonged enough. In the earliest stages of farming, skeletal growth impairment and small adult body size are often documented relative to hunter-gatherer groups, though this pattern is regionally variable. In Central/Southeast Europe, it is unclear how early life stress, growth history, and adult body size were impacted by the introduction of agriculture and ensuing long-term demographic, social, and behavioral change. The current study assesses this impact through the reconstruction and analysis of mean stature, body mass, limb proportion indices, and sexual dimorphism among 407 skeletally mature men and women from foraging and farming populations spanning the Late Mesolithic through Early Medieval periods in Central/Southeast Europe (~7100 calBC to 850 AD). Results document significantly reduced mean stature, body mass, and crural index in Neolithic agriculturalists relative both to Late Mesolithic hunter-gatherer-fishers and to later farming populations. This indication of relative growth impairment in the Neolithic, particularly among women, is supported by existing evidence of high developmental stress, intensive physical activity, and variable access to animal protein in these early agricultural populations. Among subsequent agriculturalists, temporal increases in mean stature, body mass, and crural index were more pronounced among Central European women, driving declines in the magnitude of sexual dimorphism through time. Overall, results suggest that the transition to agriculture in Central/Southeast Europe was challenging for early farming populations, but was followed by gradual amelioration across thousands of years, particularly among Central European women. This sex difference may be indicative, in part, of greater temporal variation in the social status afforded to young girls, in their access to resources during growth, and/or in their health status than was experienced by men.
Citation: Macintosh AA, Pinhasi R, Stock JT (2016) Early Life Conditions and Physiological Stress following the Transition to Farming in Central/Southeast Europe: Skeletal Growth Impairment and 6000 Years of Gradual Recovery. PLoS ONE 11(2): e0148468. https://doi.org/10.1371/journal.pone.0148468
Editor: Luca Bondioli, Museo Nazionale Preistorico Etnografico 'L. Pigorini', ITALY
Received: August 6, 2015; Accepted: January 17, 2016; Published: February 4, 2016
Copyright: © 2016 Macintosh et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Data Availability: All relevant data are within the paper and its Supporting Information files.
Funding: This study was supported by funding from the Cambridge Commonwealth, European and International Trust (AAPM), the Social Sciences and Humanities Research Council of Canada (AAPM), the European Research Council (RP; ERC Starting Grant, ERC-2010-StG263441), and the Natural Environment Research Council (JTS; NERC Grant Number NE/M/S/2003/00069). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
Competing interests: The authors have declared that no competing interests exist.
Human skeletal growth and development are complex biological processes progressing within a genetic/hormonal, environmental, and cultural framework [1–6]. The intimate relationship between human biology and culture is particularly evident in the Holocene, with the development of agriculture as the primary mode of subsistence. Though not a universal pattern, subsistence shifts at the transition to farming have been associated with reduced adult body size and/or skeletal and dental evidence of poor diet and health when compared to the corresponding profiles of related hunter-gatherer populations [7–17]. These studies and others have provided insight into the difficult early life conditions often experienced by the first farming populations as they adjusted to major subsistence, socioeconomic, and demographic change.
There are few inter-population assessments of the trends in adult body size and sexual dimorphism across the transition to farming in Central/Southeast Europe, and Mesolithic remains from Central Europe are scarce. Piontek and Vančata  reported a reduction in body size among some Early Neolithic Central European farming groups relative to preceding Mesolithic individuals from Germany, Britain, France, Spain, and Crimea. Similarly, among a larger sample of European Mesolithic and Neolithic populations, Berner and colleagues  identified a reduction in sexual dimorphism in stature and body mass across the transition to farming. However, the relationship between changes in body size, genetic, and environmental conditions during early life across the shift to agriculture is not well understood.
In the Iron Gates region, Serbia (Southeast Europe), Mesolithic and Neolithic lifeways coexisted for at least a thousand years, much longer than elsewhere in the Balkans . Here, the Mesolithic period was followed by a Transitional phase (~6200–5900 calBC) during which contact and exchange between Mesolithic and Neolithic populations occurred [20–21]. This period of Neolithic influence, combined with the stable environment and rich and predictable subsistence of the Iron Gates region [19, 22], likely facilitated the development of the relatively large and permanent settlements of Mesolithic foragers found here. In addition to their semi-sedentary lifestyle, Iron Gates Mesolithic communities were also unusual for their advanced architecture, artistic expression, burial customs, and specialized reliance on fishing . Though Late Mesolithic Iron Gates groups are not necessarily representative of hunter-gatherer communities elsewhere in Europe, this transitional Mesolithic-Neolithic semi-settled lifestyle and cultural complexity prior to intensive reliance on agriculture and domesticated animals [19, 22, 24] presents an interesting opportunity to explore the impact of food production on early life conditions experienced through the transition to full-scale agriculture in Central and Southeast Europe.
Subsistence in the Late Mesolithic-Transitional phase of the Iron Gates involved the seasonal exploitation of resources and cultivation of wild crops , the gathering of floral resources, hunting of red deer, roe deer, wild boar, and elk [24–26] and a heavy reliance on freshwater fish [19, 27–31]. Analyses of growth trajectories among the Late Mesolithic inhabitants of the Iron Gates suggest that early life stress was only moderate, and was short enough in duration to allow for catch-up growth prior to growth plate closure . However, with the movement of intensive agriculture through the Balkans into Central Europe, the diet and health of the human populations here appear to have changed considerably.
The intensification of small-scale cultivation in Central Europe began with the appearance of the Neolithic Linearbandkeramik (LBK) cultural group, which emerged in Transdanubia around 5600–5500 calBC . Within 500 years, LBK communities had spread through to France in the west and the Ukraine in the east [25,34–36]. The LBK practiced intensive agriculture and were heavily reliant on C3 plants such as wheat [37–41], supplemented by meat and milk from domesticated animals, particularly cattle, and the continued exploitation of wild game [25, 37, 39, 42–44]. Unlike the Iron Gates Late Mesolithic communities, skeletal and dental evidence from the LBK groups included in this study (Vedrovice, Nitra, Schwetzingen, Stuttgart-Mühlhausen) documents high childhood physiological stress and pathogen load [39, 45–47], particularly among women. These same LBK populations also performed intensive physical activity [48–49] that likely began, at least for some young boys, in early childhood . Thus, existing evidence suggests that early life conditions may have been difficult for these Early Neolithic LBK populations, and it is expected that adult body size and/or patterns of sexual dimorphism will differ from those documented among the Late Mesolithic inhabitants of the Iron Gates. This expectation is supported by the findings of Piontek and Vančata : the German LBK group in their study did exhibit drastically reduced body size relative to Late Upper Paleolithic and Mesolithic European groups as well as to contemporaneous Early Neolithic Corded Ware cultures. Sexual dimorphism in body size was also exceptionally high in this LBK group, reflecting higher values among males.
After this initial period of change with the introduction of farming, cereal agriculture and stock rearing became widespread, providing the main subsistence base for millennia [51–58]. Einkorn, emmer, and bread wheat, hulled six-row barley, millet, spelt, oats, rye, lentils, and peas remained staple cultivars in Central/Southeast European agricultural populations through to the Medieval period [51, 55, 59]. However, the introduction of the plough in the Mid-Late Neolithic increased agricultural productivity, supporting greater population densities and larger communities . The plough also enabled the clearance of larger tracts of land for livestock grazing, and by the Middle Neolithic and Early Bronze Age, archaeological evidence for the intensification of dairying and cheese production becomes visible [52–53]. These demographic and dietary changes drove genetic selection among agricultural populations, particularly in alleles conferring disease resistance [60–61] and enhanced nutrient processing capabilities [62–66]. For instance, evidence of selection for lactase persistence in response to intensive dairying among Central European agriculturalists appears at least by the Late Bronze Age , if not slightly earlier [68–69].
Phenotypic variation has also been influenced by dietary change and the development of social complexity after the introduction of farming. Social inequality according to status and sex is often visible in the archaeological record among the farming communities in the current study, regardless of time period, not just in grave goods and burial rites [45, 70–73], but also in health [46, 74–76], and access to plant-based dietary components, milk, milk products, and animal protein [37–41, 77–82]. In particular, differences in the degree to which men and women were afforded social status, and related variation in their overall growth conditions, were common [75, 77, 83–86], and these differences can impact sexual dimorphism in adult body size. Typically, systemic genetic, hormonal, and physiological controls on pubertal growth, skeletal maturation, and final bone length result in the standard levels of body size dimorphism between men and women [87–89]. However, these genetic and hormonal growth trajectories can be impacted substantially by cultural factors, such as differential access to resources by sex  or the preferential treatment of offspring, particularly under stressful conditions [90–91]. For instance, at the Iron Age site of Tápiószele in Hungary, and another nearby site, men's stature increased relative to a Bronze Age site from the same region, while women's stature declined . Women also exhibited fewer temporal improvements in health conditions than contemporaneous men and experienced higher childhood morbidity and dental caries than men . Thus, factors such as increasing socioeconomic complexity, population density, and greater reliance on agriculture may have impacted the diet and health of men and women quite differently. Though common among many of the cultures included in the current study, high gender and social inequality was not ubiquitous [92–93]. When present, inequalities in access to resources among the members of the increasingly more socioeconomically complex populations in this study are expected to contribute to trends in body size and/or sexual dimorphism long after the initial shift to farming in Central/Southeast Europe.
Several millennia after the initial onset of farming, better technologies and more efficient methods of crop production, as well as improvements in sanitation and healthcare, all likely increased the availability and reliability of adequate nutrition and reduced the likelihood of infection among agricultural populations [15–16, 94–96]. For example, though sex differences in health and stress were evident at Tápiószele, overall increases in life expectancy at birth and in the first year of life, and reductions in developmental stress among subadults relative to Bronze Age groups, suggest that at least some aspects of health did improve over time [75–76]. Among living humans, similar improvements in diet, health, and socioeconomic opportunity can elicit rapid recovery in growth and adult body size, leading to significant height and body mass differences that can emerge within a single generation following migration from developing to industrialized nations [97–100]. Further, industrialization since the early 19th century has been associated with rapid secular increases in height and body mass with changes in nutrition, hygiene, social class, and stress level [101–112].
It is this impact of early life conditions on adult body size and shape that allows for the interpretation of overall growth history and relative physiological stress from adult skeletal remains. There is undoubtedly an underlying genetic basis to the growth trajectory [1–3], long bone length and stature [3, 5, 113–120], body and limb proportions [4, 91, 121–122], and body mass . However, among living humans, an individual's genetically-determined growth trajectory can be modified substantially by the redirection of energy from growth to more essential functions, eliciting major delays [100, 103–104, 117, 119–120, 123–124]. One of the most powerful sources of growth impairment is the simultaneous effect of childhood malnutrition and infection , while the combination of high-impact loading and inadequate energy intake can also have lasting influences on growth and body size [126–130]. The developmental canalization of growth is such that catch-up growth may occur following less severe or prolonged stress, either through a markedly accelerated rate of growth or by the extension of the growth period [2, 103, 119, 131–133]. However, catch-up growth is costly [124, 133] and cannot always make up for previous delays [119, 134]. Thus, proximate and long-term trends in adult body size, limb proportions, and sexual dimorphism among foragers and agriculturalists have the potential to inform on a variety of factors during life, including the combined effects of dietary and health status during growth, relative physiological stress, biological and life history differences between men and women, cultural differences in access to resources, and the sex-specific assignment of tasks.
Tracking the impact of early life conditions on body size and limb proportions
Archaeological evidence from foraging and agricultural groups in Central/Southeast Europe documents cultural, social, and sex differences in the distribution of grave goods, burial rituals, life expectancy, diet, disease load, dental health, developmental stress, and habitual behaviors [24–26, 48–49, 75–77, 79–82, 84–86, 92–93, 135–139]. These differences suggest that human populations living before and following the introduction of intensive food production in these regions would have experienced variation in the severity of stress experienced during growth, its timing relative to local growth velocities, the available opportunities for catch-up growth prior to adulthood, and ultimately in adult body size/shape. The current study tracks diachronic change and regional differences in mean adult stature, body mass, limb proportions, and sexual dimorphism in these parameters through ~8000 years of subsistence and cultural change in Central/Southeast Europe (~7100 to ~850 AD). Body size and limb proportion variables are interpreted in combination with existing skeletal and dental evidence of developmental stress, health status, access to resources, and habitual behavior among the cultures and cemeteries included. This approach allows for insight to be gained into the impact of agricultural intensification, increasing sedentism, demographic and social change over thousands of years on the early life conditions and overall growth history of young men and women in Central/Southeast Europe.
Materials and Methods
The Late Mesolithic-Transitional phase hunter-gatherer-fisher population (~7131–5838 calBC) comprised skeletal remains from the cemeteries of Vlasac and Lepenski Vir in the Iron Gates region. These sites are located three kilometers apart along the Danube River in the Gospodjin Vir gorge in Vojvodina, northern Serbia. Agricultural populations included in this study represent portions of four archaeological time periods following the transition to agriculture: the Early/Middle Neolithic (~5300–4600 calBC), Early/Middle Bronze Age (~2300–1450 BC), Early through Late Iron Age (~850 BC-100 AD), and Early Medieval (~800–850 AD). These cemeteries are primarily located in Central Europe, in modern-day Baden-Württemberg (Germany), Moravia (Czech Republic), Lower Austria, western Slovakia, and the northern Carpathian Basin in Hungary. A further two agricultural cemeteries are located in the southern Carpathian Basin of Southeast Europe, in modern-day Vojvodina (northern Serbia). The Moravian Eneolithic Bell Beaker cemetery of Hoštice 1 za Hanou (~2600–2000 BC) was included in regional analyses for temporal trends within Moravia. The geographical locations of all cemeteries from which the skeletal remains originated are presented in Fig 1.
1. Vlasac 2. Lepenski Vir 3. Vedrovice 4. Nitra Horné Krškany 5. Schwetzingen 6. Stuttgart-Mühlhausen 7. Polgár-Ferenci-hát 8. Hrtkovci-Gomolava 9. Brno-Tuřany 10. Ostojićevo 11. Polgár Kenderföld 12. Brno-Maloměřice 13. Tápiószele 14. Pottenbrunn 15. Hoštice. *: this site was used only for regional analyses.
Details of the cemetery collections utilized in this study are presented in Table 1, and all individuals included in analyses are listed in S1 Table, in which specimen numbers and all relevant individual-level data are also provided. All collections were pre-excavated and are housed in museum or university collections (see Table 1). Please see S1 Appendix for further details on the cemeteries included in analyses. Collections were accessed with permission from the relevant curators. Sex assessment and age estimation was performed according to the methods outlined in Buikstra and Ubelaker . Sex assessments incorporated as many morphological criteria as possible from the cranium and pelvis. In the event of ambiguity, priority was given to pelvic criteria. Neither limb bone dimensions nor any of the variables in which sexual dimorphism was explored were utilized in sex assessment of the skeletons. Individuals sexed as 'Possible Male' or 'Possible Female' were pooled with male and female groups, respectively, for analyses. Only skeletally mature adults with all long bone epiphyses fused were included in this study, with priority given to individuals aged approximately 20–40 years at death.
To reduce variation attributable solely to genetic differences in body size and build, latitudinal variation among populations selected for inclusion was minimized, as much of the population specificity of body size and shape variables ultimately stems from long-term adaptation to climatic conditions [4, 141–149]. Further, where possible, temporal sequences were obtained from the same local regions (e.g., Moravia, Vojvodina) or the same cemetery (e.g., Gomolava).
Bone measurements and estimated body size variables
An osteometric board was used to record maximum lengths of the humerus, radius, femur, and tibia parallel to the long axis of the diaphysis, as well as bicondylar length of the femur, and biomechanical length of the tibia [150–153]. All length measurements were recorded to the nearest 0.5 millimeter. Left and right supero-inferior (S-I) femoral head diameters were measured to the nearest 0.1 millimeter using digital sliding calipers. For Pottenbrunn, some S-I femoral head diameters were obtained directly from three-dimensional laser scans of femora obtained by A. Macintosh using the measuring tool in Rapidform XOR (see Macintosh and colleagues  for laser scan methodology).
Variation in the relative lengths of the proximal and distal segments were evaluated using brachial and crural indices, which are the ratio of distal to proximal segment lengths in the upper and lower limb, respectively . Limb proportion indices were calculated using maximum lengths of the humerus, radius, and tibia, and bicondylar length of the femur [145, 147, 154–155). Poor preservation of both left and right elements, most often involving the medial malleoli, required estimation of average maximum element length in eight of 219 brachial indices and 23 of 262 crural indices included in analyses. Individuals for whom approximate average element length was used to calculate limb proportions are identified in S1 Table. Stature and body mass were estimated utilizing the equations for European Holocene populations derived by Ruff and colleagues . These equations were derived for a large sample (stature: N = 501; body mass: N = 1145) spanning the Mesolithic to modern day (~7000 BC to ≥ 1900 AD) across most of Europe, including samples from the Balkans, Czech Republic, Germany, and Eastern Austria. Stature was estimated using maximum femoral length. If the femora were missing or too poorly preserved to provide a length measurement, tibial maximum length was used. However, ecogeographic variation in distal limb lengths means that region-specific equations are required when estimating stature from the tibia . In the current study, the 'northern' equation of Ruff and colleagues  was used to estimate stature in 7% of German, Czech, Slovakian, and Austrian individuals (14 of 187 individuals), and the 'southern' equation was used in 5% of Serbian and Hungarian individuals (10 of 193 individuals). These individuals are indicated in S1 Table.
Body mass was estimated from femoral head SI breadth utilizing the equations for European Holocene populations derived by Ruff and colleagues . Among Medieval individuals for whom S-I breadths were measured directly from 3D laser scans, measurements were taken on two separate occasions one year apart (AAM) and the average value was used to estimate body mass. Body mass estimates quantified from both scan- and caliper-derived S-I breadths did not differ significantly in a test sample of femora for which both measurements were available, so all Medieval body mass estimates were pooled together for analyses. Individuals for whom body mass was estimated from a laser scan model are indicated in S1 Table. No age adjustment factor was applied to stature or body mass estimates, following Ruff and colleagues . Where possible, all estimated body size variables were calculated from an average of left and right side elements, in order to reduce laterality biases and minimize the effects of asymmetry [152–153, 155–156].
Mean stature and body mass were expected to be larger in men than women simply due to genetic and hormonal differences between the sexes [87, 89, 157–158]. Thus, the current study investigated change in a ratio of male to female size that expresses the male mean as a percentage of the female mean for a given variable: (male mean / female mean)*100 [159–160]. However, this index does not take sample size or standard deviation into account. Thus, in addition, a measure of sexual dimorphism quantified using the regions of non-overlap of male and female distributions was employed (D) , whereby values range from 0 (no dimorphism, complete overlap) to 1 (completely dimorphic, no overlap). Further, effect sizes for sexual dimorphism were calculated using Pearson's correlation coefficient r, according to the following method outlined by Mascie-Taylor : where df = n-2 and
An r-value greater than 0.8 was considered a large effect size, indicating a high magnitude of difference between males and females, and a value of 0.5 was considered a medium effect . These effect sizes are more conservative than the widely followed suggestions of Cohen [163–164], whereby any r-value greater than 0.5 is considered a large effect, so all values >0.5 are highlighted in the results tables.
Sample data distributions were assessed for normality using z-scores for skewness and kurtosis, Q-Q plots, and histograms. All outliers more than three standard deviations from the mean were removed and all data included in analyses were normally distributed. One-way analysis of variance (ANOVA) was used to test for temporal change in body size variables and limb proportions among male and female Central/Southeast Europeans. Post-hoc comparisons were performed with either the Hochberg's GT2 or Games-Howell test. Independent samples t-tests were used to explore sexual dimorphism in body size variables and limb proportions. Statistical analyses were conducted in SPSS v23, and statistical significance was considered as a p-value less than 0.05.
Trends by time period
Summary statistics for body size and limb proportion variables by sex and time period are presented in Table 2 and results of one-way ANOVAs by sex and time period are presented in Table 3. Boxplots for temporal change by sex and time period in stature, body mass, brachial index, and crural index are presented in Fig 2. When agricultural populations are compared to the Late Mesolithic hunting/gathering/fishing group, the relatively large body size of the latter is apparent, particularly among men. Late Mesolithic Iron Gates men are significantly taller (p<0.001) and heavier (p<0.006) than Neolithic men, significantly taller (p<0.001) than Bronze Age men, and significantly taller (p<0.018) and heavier (p<0.021) than Iron Age men. Similarly, Late Mesolithic women are significantly taller than both Early Neolithic (p<0.001) and Iron Age (p<0.013) women. Though mean brachial and crural indices do not change significantly, these mean values are also higher in Late Mesolithic men and women than their Neolithic counterparts.
A) Stature, B) Body mass, C) Brachial index, D) Crural index. Brackets indicate significant differences (* = p<0.05; ** = p<0.01).
Through the ~6150 years spanned by the agricultural populations examined in this study (Early Neolithic through Early Medieval periods), average male body size does not change significantly. However, Neolithic men consistently have the lowest mean estimated stature and body mass of all agriculturalists. Among Neolithic women, deficits in body size variables and crural index are particularly pronounced relative to subsequent agricultural women. Neolithic women have significantly lower mean estimated stature than Early Medieval women (p<0.001), significantly lower mean body mass than Bronze Age women (p<0.012), and significantly lower mean crural indices (shorter tibiae relative to femora) than Bronze Age (p<0.012) and Early Medieval (p<0.001) women. Early Medieval women are also significantly taller on average than their counterparts in the Iron Age (p<0.007).
Results of independent samples t-tests for sexual dimorphism in body size and limb proportions by time period are presented in Table 4. Males at all time periods examined have significantly higher stature and body mass than females (p<0.001 for all). Effect sizes for the magnitude of sexual dimorphism in stature and body mass are very large in the Mesolithic period (r-values of 0.72 and 0.86, respectively), and decline consistently through to the Early Medieval period (r-values of 0.55 and 0.42, respectively). Temporal declines in D values for stature and body mass also indicate that male and female distributions for these variables overlap to a progressively greater extent through time, becoming more similar. Indices of sexual dimorphism parallel this trend to a certain extent, declining consistently from the Mesolithic through Bronze Age for both variables. By contrast, significant sexual dimorphism in limb proportions is rare, though brachial index is significantly higher among Neolithic (p<0.001) and Bronze Age (p<0.017) males than females. Effect sizes are also small (most r<0.2, maximum of 0.4) in almost all instances for limb proportions, as are D values, and neither indicate any consistent trend through time in the sexual dimorphism of limb proportions.
Trends by region
Summary statistics for body size and limb proportion variables are presented for men and women by geographic region and cemetery in S2 Table. Results for regional differences within each time period by sex are presented in Table 5. Among Iron Gates foragers, women at Lepenski Vir are of significantly lower stature (p<0.001) and body mass (p<0.011) than women at Vlasac, while men have significantly reduced crural indices (p<0.032). Among Neolithic farming populations, there are almost no significant differences in body size or limb proportions among the LBK, ALP, and Vinča men and women, despite these groups spanning Germany to Vojvodina. Hungarian ALP men at Polgár-Ferenci-hát have significantly reduced brachial indices compared to LBK men in Moravia (Vedrovice; p<0.034) and Baden-Württemberg (Schwetzingen; p<0.033), but this difference is not replicated in any other body size or proportion variable. In the Bronze and Iron Ages however, Central and Southeast Europe diverge; in these time periods, men and women from Vojvodina exhibit significantly smaller body size and/or relatively shortened distal lower limb segments relative to broadly contemporaneous groups in Central Europe (Moravia and Hungary; see Table 5). There are never any significant differences between Hungarian and Moravian groups for any variable in either sex. Within Moravia/Lower Austria, the Eneolithic population from Hoštice does not differ significantly in body size or limb proportions from preceding or subsequent agriculturalists in the region. However, regional analyses must be interpreted with caution, as the division of time periods into constituent cemeteries greatly reduces sample size, particularly among the female group and at certain cemeteries. Sampling biases may be affecting results among these small cemetery samples, such as at Polgár-Ferenci-hát, Hoštice, and Gomolava.
This study explores the impact of the transition to farming and ~6150 years of agricultural intensification and cultural change on adult body size and limb proportions among men and women in Central/Southeast Europe. Mean stature and body mass were significantly lower among Neolithic agriculturalists than Late Mesolithic hunter-gatherer-fishers, with particularly large body size among Late Mesolithic men and particularly small body size among Neolithic women. Following the Neolithic period, mean stature, body mass, and crural index among agricultural populations increased very gradually across thousands of years in both sexes, but significantly so almost exclusively among women. Though men at all time periods were significantly taller and heavier than women, the extent to which this was the case was highest among Late Mesolithic foragers and Neolithic farmers, and declined through time among agriculturalists alongside temporal increases in female body size.
Interestingly, among women, mean stature and body mass was significantly higher at Vlasac than Lepenski Vir in the current study, as were male crural indices, despite the contemporaneity of the sites and their close proximity to each other. Existing reconstructions of stature among men and women at Vlasac using the equations of Trotter and Gleser  provided mean statures slightly taller than those in the current study: 176.6 cms for males and 163.2 cms for females . These are significantly taller than stature estimates from Western European Mesolithic populations, and would not be expected in a population experiencing high nutritional stress and/or lacking opportunities for catch-up growth . At Vlasac, most human remains have thus far dated to the Late Mesolithic, with its most intensive occupation occurring from ~7131 to 5838 calBC, while Neolithic Starčevo pottery does not appear at Vlasac until after ~6000–5800 calBC [26, 166]. In contrast, human occupation at Lepenski Vir was most intensive during the Transitional phase (Lepenski Vir I-II), between ~6240–5845 calBC . The appearance of the 'Neolithic package' at the site dates to only slightly later: the remains of domesticated pigs, goats, and cattle found in association with Early Neolithic features, such as domed ovens, at Lepenski Vir date to ~6005 to 5798 calBC . The Transitional phase of habitation at Lepenski Vir also contains both Early Neolithic pottery and polished stone axes typical of Neolithic groups [20, 167–168]. This clear documentation of contact between Mesolithic and Neolithic cultures at Lepenski Vir suggests the possibility that hunting/fishing populations living at the site may have experienced some of the same factors that contributed to reduced body size and limb proportions among Early Neolithic groups.
Individuals with non-local trace element signatures have also been identified during the Transitional phase at Lepenski Vir , indicative of individuals being buried at the site that originated outside of the Iron Gates. Also, the Iron Gates at this time experienced wetter conditions and considerable flooding [29, 170], likely contributing to evidence for the exploitation of a wider array of dietary resources among post-contact individuals at Lepenski Vir [28, 30–31, 171], with a significantly higher proportion of terrestrial foods in the diet at this time . In addition to any of the above factors, results could also be reflecting the small sample sizes created by the division of time periods into constituent cemeteries. There are also a range of biases known to typify archaeological assemblages, including differential preservation of human remains from different age groups, partial samples of a larger cemetery, variations in the demographic profiles of the cemetery samples, and the overall selectivity affecting the formation of any mortuary sample . It is possible that such biases, which cannot be assessed at the present study, contributed to some extant to the observed temporal patterns, both temporally and regionally. However, none of the above-mentioned factors varies in its direction and intensity between the time periods analysed here.
Despite differences in body size and crural indices between Vlasac and Lepenski Vir, the majority of the Mesolithic group overall was typified by large body size and high limb proportion indices relative to most of the agricultural populations analyzed. The semi-settled and socially complex lifestyle of the Late Mesolithic Iron Gates foragers is not representative of typical European hunter-gatherer lifeways, and their body size appears to be larger than their Western European Mesolithic counterparts [27, 173]. However, the reduction in mean stature and body mass between Late Mesolithic Iron Gates foragers (Southeast Europe) and Neolithic Central/Southeast European agriculturalists (predominantly LBK) identified in this study was similar to that identified by Piontek and Vančata between Mesolithic European groups and Early Neolithic Central European LBK . Further, the large changes in body size identified in the current study between Late Mesolithic and Neolithic populations do parallel existing evidence of substantially different early life conditions for these populations. For instance, the moderate early life stress and/or opportunities for catch-up growth in the Late Mesolithic Iron Gates foragers  contrast sharply with high childhood developmental stress, shifting of the childhood growth spurt, and high pathogen load documented among the Early Neolithic LBK at Vedrovice and Stuttgart-Mühlhausen [45–47]. The Early Neolithic LBK populations in this study also show evidence of high lower limb bone loading among both men and women relative to the subsequent agricultural groups, likely associated with higher terrestrial mobility . There is also evidence that high mobility was a component of the early childhoods of at least some young LBK boys . This intensive physical activity, undertaken at a young age in at least some individuals, may have affected already limited energetic resources during growth among LBK populations. It is probable that the combination of a homogeneous carbohydrate-rich diet, high developmental stress, intensive physical activity at a young age, and sex differences within these all contributed to the impairment of adult body size and crural index among Neolithic populations relative to both hunter-gatherers and subsequent farming populations.
Despite large body size in most Iron Gates men and women, particularly at Vlasac, compared to their agricultural counterparts, results suggest that disparity in growth conditions between the sexes may have been present that negatively affected women. The magnitude of sexual dimorphism in stature and body mass in the Iron Gates Late Mesolithic group was the largest of all time periods examined: men were, on average, 128% the body mass of women, a very large effect (r = 0.86). Significant differences have been documented in male and female diet at Vlasac and Lepenski Vir  that may be playing a role in this result. Similar to the trends documented by Berner and colleagues , the overall magnitude of sexual dimorphism in stature and body mass in Central/Southeast Europe decreased slightly across the transition to farming. However, effect sizes for dimorphism in stature and body mass among Neolithic communities overall was still high (r>0.6 for body mass; r>0.7 for stature), and this was the case across both the LBK and ALP and across a geographic region spanning from southwest Germany to Hungary (e.g., r = 0.94 for stature at Schwetzingen, Nitra, and Polgár-Ferenci-hát).
The substantially smaller body size of Early Neolithic agricultural women relative to men parallels evidence for sex differences in diet and health among these populations, with negative effects for women. Relative to men, LBK women consumed less animal protein, had a higher frequency and severity of dental caries, and had a reduced life expectancy [36–41, 45–46, 74, 78]. Thus, despite the different subsistence systems of Mesolithic and Neolithic populations in the current study, gender inequality that negatively affecting the growth and development of young girls may have been a shared factor of both of these early and socially complex ways of life. Sex differences in the adequacy of resources supplied to young boys and girls, in levels of early life stress, health status, divisions of labor, age at initiation of work, and/or life history could all be playing a role in this particularly high sexual dimorphism in adult body size. Sex differences in sensitivity to poor conditions during childhood and in the likelihood of subsequent recovery could also be playing a role, though the relative impairment of female body size does not appear to support the typical suggestion of higher sensitivity among males [90, 105, 107–108, 123, 134, 174–175].
Following the initial introduction of agriculture, the magnitude of sexual dimorphism declined progressively through time, alongside gradual increases in mean stature, body mass, and crural index that were significant almost exclusively among women. From the Neolithic to the Bronze Age, women's mean body mass increased significantly, and their tibiae became significantly longer relative to femora, a trend which continued into the Iron Age as well. These temporal increases in mean body size/proportions were consistent across Moravia/Lower Austria, inclusive of the Eneolithic sample, and the Carpathian Basin of Hungary and western Slovakia, but were less pronounced within Vojvodina, where Bronze and Iron Age men and women were of relatively small body size. The regional similarity of Neolithic groups throughout Central and Southeast Europe may suggest that the adoption of farming initially had widespread negative impacts on growth and adult body size and shape. Subsequent regional variation in the Bronze Age corresponds to a time of greater socioeconomic complexity and technological innovation [43, 58, 176–181], changing behaviors and divisions of labor for women [76, 139, 182–183] and major migration and admixture events [67, 184–187], that all likely played a role in driving temporal and regional change in body size variables.
The large average body size and high limb proportions of Bronze Age women in Moravia and Hungary, and the low magnitude of sexual dimorphism at this time, do correspond to existing evidence from these regions of relatively high gender equality , similar dental health/diet between the sexes, and low physiological stress during growth at this time . For example, by the Bronze Age, the small Polgár region of Hungary was at the intersection of important east-west and north-south trade links, capable of sustaining several Füzesabony communities and their richly furnished cemeteries within just a few kilometers of each other . Socioeconomic conditions were likely good for the inhabitants of these communities, including at Polgár Kenderföld, and women here could clearly attain high social status; the richest grave at the site was that of a 35–40 year old woman, adorned with amber and bronze artifacts . Good conditions in the Polgár region during the Bronze Age, particularly for women, correspond to an increase of ~ nine cms in average female stature across the ~3500 years separating the Füzesabony women here from their ALP counterparts in the Early Neolithic.
Recent genetic evidence has also begun to shed light on major population migrations and admixture between the Neolithic/Eneolithic and Bronze Age periods in Europe, mainly from the Eurasian Steppe regions such as Western Russia and Ukraine [67, 184–187]. An influx of genetically different populations could clearly be contributing to divergence in the body size of Central and Southeast European populations in the Bronze Age. In addition, the gateway position of Vojvodina between influences from both central and southeastern Europe encouraged cultural mixing [188–189], and the genetic diversity of its inhabitants may have been impacted in different ways than the inhabitants of Central Europe.
The Iron Age also brought an influx of eastern cultural influences and genes into the Carpathian Basin [67, 188], coinciding with the movement of Scythian groups from the Eurasian steppes , and Celtic migration was also high [188, 191] The small mean body size of Iron Age Bosut men and women from Gomolava relative to their Scythian and Celtic counterparts in Central Europe is thus notable here. In particular, mean stature among Bosut men from Gomolava was very low, so much so that they differed little from Bosut women, and sexual dimorphism in stature at the site was consequently minimal (r = 0.11). There were no significant differences in any body size variables between 'Males' and 'Possible Males' at the site that would suggest the erroneous presence of females in the male sample. However, despite the unusually low body size of the Bosut men in this study, mean male crural index rose significantly in the ~3750 years separating the Neolithic Vinča and Iron Age Bosut men at Gomolava (p<0.047). Because childhood malnutrition impacts the growth of the distal limb segments in particular , gradual increases in mean crural index among agricultural men in Vojvodina through time could be indicative of declining early life stress. It is likely that sampling biases are affecting the male sample of Bosut from Gomolava, and more research is needed to characterize body size among this Early Iron Age population in Vojvodina, as few Bosut skeletal remains are known from this time in the region .
Despite the high degree of migration among Scythians and Celts, the Iron Age population overall did not differ significantly in body size from that of any other agricultural group, though crural indices continued to increase relative to the Neolithic among women. Among the Celts and Scythians, it is rather the differential treatment of men and women that is interesting, as the Iron Age group overall exhibited a relatively high degree of sexual dimorphism in estimated stature. Warrior status was an important social differentiator among Celtic men at the necropolis of Brno-Maloměřice , and Celtic warriors were afforded better access to animal protein than non-warriors [81–82]. High social status was not restricted to men, but it does appear to have taken more time for women to attain [92–93]. It is unknown if any of the Celtic men included in analyses from Brno-Maloměřice were warriors, but the greater likelihood of attaining high social status among young men than women at this time could be contributing to the high sexual dimorphism in stature at Brno-Maloměřice (r = 0.75) relative to the other Bronze Age, Iron Age, and Medieval sites examined (r values ranging from 0.11 to 0.73), with the exception of Tápiószele. At the Iron Age Scythian site of Tápiószele in Hungary, conditions for women relative to men were very poor, and the effect size for stature at the site was correspondingly very large (r = 0.84). As described above, Scythian women here showed evidence of greater childhood morbidity, an earlier average age of developmental stress, and a higher reliance on carbohydrates than men , and experienced fewer temporal improvements than did men in body size, levels of developmental stress and dental caries, and health status relative to a Bronze Age sample from the same region . These poor conditions for women relative to men could be contributing, at least in part, to the high levels of sexual dimorphism in body size at Tápiószele.
Though there is still evidence of high dietary and environmental stress during growth at the Early Medieval Slavic site of Pottenbrunn [84, 135], women here were significantly taller than those of the preceding Iron Age group overall. Among men, it was also the Medieval period in which was found the highest mean stature and body mass of all agricultural time periods examined. This large body size, particularly among women, could again reflect genetic differences, as the Slavs were an immigrant population into eastern Austria, arriving sometime around the late 6th or 7th centuries AD . However, in stark contrast to the earliest farming populations in this study, women at Pottenbrunn exhibited fewer indicators of developmental stress than men, and there is no isotopic indication of any dietary differences between the sexes at the site . Correspondingly, effect sizes of sexual dimorphism in the Medieval group were the lowest of all time periods, and mean estimated stature among women at Pottenbrunn was second only to pre-farming Late Mesolithic foraging women from Vlasac. In fact, the average Medieval individual at Pottenbrunn could expect to live a decade longer than the average individual at the Early Neolithic cemetery of Vedrovice from ~6150 years earlier (average life expectancy of 37.7 years relative to 27.6 years, respectively) [46, 84]. Thus, developmental insults at Pottenbrunn may have been less severe than those experienced by earlier agricultural groups, or recovery may have been possible prior to growth plate closure, reducing the impact of these insults on adult body size.
However, the relationship between human growth and development and culturally driven environmental change since the transition to agriculture is complex, and it is interesting to note the gradual nature with which body size and proportions changed across the ~8000 years examined by this study. For instance, following impairment in the earliest stages of farming, average body size and limb proportions among men changed very little across thousands of years, despite advances in agricultural efficiency, technology, and sanitation. Even the more pronounced positive temporal change among agricultural women was very gradual, and young girls in even some of the more recent agricultural populations examined (e.g., Tápiószele) still appeared to have had profoundly more severe or prolonged episodes of early life stress than contemporaneous men. However, ultimately, following a pronounced reduction in body size among early farmers relative to foragers, gradual temporal increases in mean stature, body mass, and crural index and reductions in sexual dimorphism among most Central/Southeast Europe agriculturalists do suggest that the overall impact of millennia of cultural, socioeconomic, and technological change was positive, particularly for women.
This study examined both overall and regional trends in adult body size, limb proportions, and sexual dimorphism among Central/Southeast European foragers and farmers in the context of existing evidence of diet and health from the same populations. This study documented significant reductions in mean stature and body mass among the earliest Neolithic agriculturalists in Central/Southeast Europe relative to preceding Late Mesolithic Iron Gates hunter-gatherer-fishers. The relatively large body size of Late Mesolithic Iron Gates hunter-gatherer-fishers, particularly at Vlasac, contrasted with the relatively small body size and low crural indices of Neolithic agricultural men and women. These deficits in body size and limb proportions among the earliest farmers were followed by gradual increases in mean values across thousands of years, though these were significant almost exclusively among women and were not documented in the Bronze and Iron Ages of Southeast Europe. Particularly high magnitudes of sexual dimorphism in mean stature and body mass in the Late Mesolithic and Neolithic periods suggested that young women at these times were more likely than young men to experience more severe and/or prolonged developmental stress and poor health. These deficits among women relative to men declined through time alongside significant increases in the body size of agricultural women. The very gradual nature by which body size and limb proportions changed through time among agriculturalists, and the regional differences identified in the Bronze and Iron Ages, suggests that temporal improvements to technology and agricultural efficiency, as well as to factors like infrastructure and sanitation, did not necessarily equate to universal, rapid, or population-wide increases in body size. This may reflect the influence of increasing socioeconomic complexity and urbanization on the unequal distribution of resources among individuals and on disease load, and of major admixture and migration events in the Bronze and Iron Ages. However, by the Early Medieval period, mean body size and crural index among both men and women overall had increased markedly from the introduction of farming in Central/Southeast Europe, significantly so among the women. These results parallel dramatic improvements in average life expectancy already documented across the ~6150 years of agriculture covered by this study, and track alongside existing evidence for improved health and access to dietary resources through time among at least some members of Central/Southeast European farming communities.
S1 Appendix. Supplementary Information on Cemeteries.
S1 Table. Body size and limb proportion variables by cemetery and individual.
The authors would like to thank Dr. Luca Bondioli and two anonymous reviewers for their insightful comments that greatly improved this manuscript. The authors also wish to thank the following individuals for their hospitality and for access to the skeletal collections utilized in this research: Michael Francken at the University of Tübingen (Tübingen, Germany), Joachim Wahl at the University of Tübingen (Tübingen, Germany) and the Stuttgart Regional Council State Conservation Office- Osteology (Konstanz, Germany), Lidija Milašinović at the National Museum of Kikinda (Kikinda, Serbia), Marija Jovanović and Darko Radmanović at the Museum of Vojvodina (Novi Sad, Serbia), Sofija Stefanović at the University of Belgrade (Belgrade, Serbia), Eva Drozdová at Masaryk University (Brno, Czech Republic), Zdeněk Tvrdý at the Moravian Museum (Brno, Czech Republic), Ildiko Pap at the Hungarian Natural History Museum (Budapest, Hungary), and Maria Teschler-Nicola at the Vienna Natural History Museum (Vienna, Austria). The authors also wish to thank Dr. Emma Pomeroy (University of Cambridge) for her assistance with production of the map (Fig 1).
Conceived and designed the experiments: AAM RP JTS. Performed the experiments: AAM JTS. Analyzed the data: AAM. Contributed reagents/materials/analysis tools: AAM JTS. Wrote the paper: AAM RP JTS.
- 1. Ruff CB. 2002. Variation in human body size and shape. Annu Rev Anthropol 31:211–232.
- 2. Zelditch ML. 2005. Developmental regulation of variability. In: Hallgrímsson B, Hall BK, editors. Variation: A central concept in biology. London: Elsevier. pp 249–276.
- 3. Dubois L, Girard M, Girard A, Tremblay R, Boivin M, Pérusse D. 2007. Genetic and environmental influences on body size in early childhood: A twin birth-cohort study. Twin Res Hum Genet 10:479–485. pmid:17564506
- 4. Frelat MA, Mittereocker P. 2011. Postnatal ontogeny of tibia and femur form in two human populations: A multivariate morphometric analysis. Am J Hum Biol 23:796–804. pmid:21957036
- 5. Morris DH, Jones ME, Schoemaker MJ, Ashworth A, Swerdlow AJ. 2012. Familial concordance for height and its components: Analyses from the breakthrough generations study. Am J Hum Biol 24:22–27. pmid:22121080
- 6. Vercellotti G, Piperata BA. 2012. The use of biocultural data in interpreting sex differences in body proportions among rural Amazonians. Am J Phys Anthropol 147:113–127. pmid:22120650
- 7. Nickens PR. 1976. Stature reduction as an adaptive response to food production in Mesoamerica. J Archaeol Sci 3:31–41.
- 8. Angel 1984. Health as a crucial factor in the changes from hunting to developed farming in the eastern Mediterannean. In: Cohen MN, Armelagos GJ, editors. Paleopathology at the origins of agriculture. Florida: Academic Press, Inc. pp 51–73.
- 9. Rathbun TA. 1984. Skeletal pathology from the Paleolithic through the Metal Ages in Iran and Iraq. In: Cohen MN, Armelagos GJ, editors. Paleopathology at the origins of agriculture. Orlando: Academic Press, Inc. pp 137–167.
- 10. Ruff CB, Larsen CS, Hayes W. 1984. Structural changes in the femur with the transition to agriculture on the Georgia coast. Am J Phys Anthropol 64:125–136. pmid:6465303
- 11. Larsen CS. 1995. Biological changes in human populations with agriculture. Ann Rev Anthropol 24:185–213.
- 12. Larsen CS. 2006. The agricultural revolution as environmental catastrophe: Implications for health and lifestyle in the Holocene. Quat Int 150:12–20.
- 13. Alfonso MP, Standen VG, Castro MV. 2007. The adoption of agriculture among Northern Chile Populations in the Azapa Valley, 9000-1000BP. In: Cohen MN, Crane-Kramer GMM, editors. Ancient health: Skeletal indicators of agricultural and economic intensification. Florida: University Press of Florida. pp 113–129.
- 14. Douglas MT, Pietrusewsky M. 2007. Biological consequences of sedentism: Agricultural intensification in Northeastern Thailand. In: Cohen MN, Crane-Kramer GMM, editors. Ancient health: Skeletal indicators of agricultural and economic intensification. Florida: University Press of Florida. pp 300–319.
- 15. Starling AP, Stock JT. 2007. Dental indicators of health and stress in early Egyptian and Nubian agriculturalists: A difficult transition and gradual recovery. Am J Phys Anthropol 134:520–528. pmid:17786997
- 16. Mummert A, Esche E, Robinson J, Armelagos GJ. 2011. Stature and robusticity during the archaeological transition: Evidence from the bioarchaeological record. Econ Hum Biol 9:284–301
- 17. Piontek J, Vančata V. 2012. Transition to agriculture in Central Europe: Body size and body shape amongst the first farmers. Interdisciplinaria Archaeologica: Natural Sciences in Archaeology 3:23–42.
- 18. Berner M, Sládek V, Ruff C, Holt BM, Niskanen M, Galeta P, et al., 2012. Variation in sexual dimorphism of postcranial robusticity and body proportions in European Holocene populations. Am J Phys Anthropol 147:98.
- 19. Radovanović, I. 1996. The Iron Gates Mesolithic. Ann Arbor: International Monographs in Prehistory, Archaeological Series 11.
- 20. Borić D. 2002. The Lepenski Vir conundrum: reinterpretation of the Mesolithic and Neolithic sequences in the Danube Gorges. Antiquity 76:1026–1039
- 21. Borić D, Dimitrijević V. 2007. Absolute chronology and stratigraphy of Lepenski Vir. Starinar 9–55.
- 22. Srejović D, Letica Z. 1978. Vlasac: A Mesolithic Settlement in the Iron Gates Region: Volume 1- Archaeology. Belgrade: Serbian Academy of Sciences and Arts Monographies Vol DXII, Department of Historical Sciences Vol. 5.
- 23. Wallduck RJ. 2013. Post-mortem body manipulation in the Danube Gorges' Mesolithic-Neolithic: A taphonomic perspective. PhD dissertation, University of Cambridge.
- 24. Jochim M. 2002. The Mesolithic. In: Milisauskas S, editor. European prehistory: A survey. New York: Kluwer Academic. pp 115–141.
- 25. Whittle A. 1996. Europe in the Neolithic: The creation of new worlds. Cambridge: Cambridge University Press.
- 26. Borić D, French CAI, Stefanović S, Dimitrijević V, Cristiani E, Gurova M, et al., 2014. Late Mesolithic lifeways and deathways at Vlasac (Serbia). J Field Archaeol 39:4–31.
- 27. Meiklejohn C, Zvelebil M. 1991. Health status of European populations at the agricultural transition and the implications for the adoption of farming. In: Bush H, Zvelebil M, editors. Health in past societies. Oxford: British Archaeological Reports, International Series 567. pp 129–145.
- 28. Bonsall C, Lennon R, McSweeney K, Stewart C, Harkness D, Boroneant V et al., 1997. Mesolithic and Early Neolithic in the Iron Gates: A paleodietary persective. J Eur Archaeol 5:50–92.
- 29. Bonsall C, Cook G, Lennon R, Harkness D, Scott M, Bartosiewicz L et al., 2000. Stable isotopes, radiocarbon and the Mesolithic-Neolithic transition in the Iron Gates. Documenta Praehistorica 27:119–132.
- 30. Bonsall C, Cook GT, Hedges REM, Higham TFG, Pickard C, Radovanović I. 2004. Radiocarbon and stable isotope evidence of dietary change from the Mesolithic to the Middle Ages in the Iron Gates: New results from Lepenski Vir. Radiocarbon 46:293–300.
- 31. Roksandic M. 2012. Mobile and terrestrial but firmly rooted on the river banks: Biological anthropology of Lepenski Vir and the Iron Gates Gorge Mesolithic. Advances in Anthropol 2:117–124.
- 32. Pinhasi R, Stefanovic S, Papathanasiou A, Stock J. 2011. Variability in long bone growth patterns and limb proportions within and amongst Mesolithic and Neolithic populations from Southeast Europe. In: Pinhasi R, Stock J, editors. Human bioarchaeology of the transition to agriculture. Chichester: John Wiley & Sons, Ltd. pp 177–202.
- 33. Bánffy E, Oross K. 2010. The earliest and earlier phase of the LBK in Transdanubia. In: Gronenborn D, Petrasch J, editors. Die Neolithisierung Mitteleuropas: international Tagung, Mainz 24. bis 26. Junis 2005 (The Spread of the Neolithic to central Europe: proceedings of the international symposium, Mainz 24th-26th June 2005). Verlag des Römisch-Germanischen Zentralmuseums. pp 255–272.
- 34. Jeunesse C. 1997. Pratiques funéraires au Néolithique ancien: sépultures et nécropoles Danubiennes 5500–4900 av. J. C. Paris: Éditions Errance.
- 35. Bocquet-Appel JP, Naji S, Linden MV, Kozlowski JK. 2009. Detection of diffusion and contact zones of early farming in Europe from the space-time distribution of 14C dates. J Archaeol Sci 36:807–820.
- 36. Whittle A, Bentley RA, Bickle P, Dočkalová M, Fibiger L, Hamilton J et al., 2013. Moravia and western Slovakia. In: Bickle P, Whittle A, editors. The first farmers of Central Europe: Diversity in LBK lifeways. Oxbow Books: Oxford. pp 101–158.
- 37. Smrčka V, Bůzek F, Erban V, Berkovec T, Dočkalová M, Neumanová K et al., 2005. Carbon, nitrogen and strontium isotopes in the set of skeletons from the Neolithic settlement at Vedrovice (Czech Republic). Anthropologie 43:315–323.
- 38. Smrčka V, Mihaljevic M, Zobcová J, Humpolová A, Berkovec T. 2006. Multielementární chemická analýza z kosterních pozůstatků lidí a zvířat neolitického sídliště ve Vedrovicích (Česka Republika). Ve Službách Archeologie 7:329–340.
- 39. Richards MP, Montgomery J, Nehlich O, Grimes V. 2008. Isotopic analysis of humans and animals from Vedrovice. Anthropologie 46:185–194.
- 40. Jarošová I. 2008. Dietary inferences using buccal microwear analysis on the LBK population from Vedrovice, Czech Republic. Anthropologie 46:175–184.
- 41. Zvelebil M, Pettitt P. 2008. Human condition, life, and death at an early Neolithic settlement: Bioarchaeological analyses of the Vedrovice cemetery and their biosocial implications for the spread of agriculture in Central Europe. Anthropologie 46:195–218.
- 42. Milisauskas S. 2002. Early Neolithic: The first farmers in Europe, 7000-5500/5000 BC. In: Milisauskas S, editor. European prehistory: A survey. New York: Kluwer Academic. pp 143–192.
- 43. Milisauskas S, Kruk J. 2002. Middle Neolithic continuity, diversity, innovations, and greater complexity, 5500/5000-3500/3000 BC. In: Milisauskas S, editor. European prehistory: A survey. New York: Kluwer Academic. pp 193–246.
- 44. Müller HH. 1964. Die Haustiere der Mitteldeutschen Bandkeramik. Berlin: Deutsche Akademie der Wissenschaften.
- 45. Price TD, Wahl J, Knipper C, Burger-Heinrich E, Kurz G, Bentley RA. 2003. Das bandkeramische Gräberfeld von Stuttgart-Mühlhausen: Neue Untersuchungsergebnisse zum Migrationsverhalten im frühen Neolithikum. In: Funda DT, editor. Fundberichte aus Baden-Württemberg. Stuttgart: Kommissionsverlag Konrad Theiss Verlag. pp 23–58.
- 46. Lillie M. 2008. Vedrovice: Demography and paleopathology in an early farming population. Anthropologie 46:135–152.
- 47. Welte B, Wahl J. 2010. Auxologische Studien an Skelettresten fruehneolithischer Kinder und Jugendlicher aus Suedwestdeutschland. FuBer BadWürt 31:7–28.
- 48. Macintosh A, Pinhasi R, Stock JT. 2014. Lower limb biomechanics track long-term decline in mobility across ~6150 years of agriculture in Central Europe. J Archaeol Sci 52:376–390.
- 49. Macintosh A, Davies TG, Pinhasi R, Stock JT. 2015. Declining tibial curvature parallels ~6150 years of decreasing mobility in Central European agriculturalists. Am J Phys Anthropol 157:260–275. pmid:25677783
- 50. Knipper C. 2009. Mobility in a sedentary society: Insights from isotope analysis of LBK human and animal teeth. In: Hofmann D, Bickle P, editors. Creating communities: New advances in Central European Neolithic research. Oxford and Oakville: Oxbow Books. pp 142–158.
- 51. van Zeist W. 1975. Preliminary report on the botany of Gomolava. J Archaeol Sci 2:315–325.
- 52. Coles JM, Harding AF. 1979. The Bronze Age in Europe: An introduction to the prehistory of Europe c. 2000-700BC. London: Methuen.
- 53. Harding A. 2002. The Bronze Age. In: Milisauskas S, editor. European prehistory: A survey. New York: Kluwer Academic. pp 271–334.
- 54. Küster H. 1992. Kulturpflanzenanbau in Südbayern seit der Jungsteinzeit. In: Hahn M, Prammer J, editors. Bauern in Bayern: Von den Anfängen bis zur Römerzeit. Straubing: Gäubodenmuseum. pp 137–155.
- 55. Bottema S, Ottaway BS. 1982. Botanical, malacological and archaeological zonation of settlement depots at Gomolava. J Archaeol Sci 9:221–246.
- 56. Wells PS. 2002. The Iron Age. In: Milisauskas S, editor. European prehistory: A survey. New York: Kluwer Academic. pp 335–383.
- 57. Rösch M. 2008. New aspects of agriculture and diet of the early Medieval period in Central Europe: Waterlogged plant material from sites in south-western Germany. Veget Hist Archaeobot 17:S225–S238.
- 58. Bartelheim M. 2009. Elites and metals in the Central European Early Bronze Age. In: Kienlin TL, Roberts BW, editors. Metals and Societies: Studies in honour of Barbara S. Ottoway. Bonn: Dr. Rudolf Habelt GmbH. pp 34–46.
- 59. Smrčka V, Drozdová E, Bůzek F, Zocová J, Matějíčková A. 2007. Stabilní izotopy C, N, v souboru skeletů lidu kultury zvoncových pohárů z Hoštic 1 za Hanou (Morava). Ve Službách Archeologie Brno: Moravské zemské muzeum 2:110–114.
- 60. Tishkoff SA, Varkonyi R, Cahinhinan N, Abbes S, Argyropoulos G, Destro-Bisol G et al. 2001. Haplotype diversity and linkage disequilibrium at human G6PD: Recent origin of alleles that confer malarial resistance. Science 293:455–462. pmid:11423617
- 61. Galvani AP, Novembre J. 2005. The evolutionary history of the CCR5- Δ32 HIV-resistance mutation. Microbes Infect 7:302–309. pmid:15715976
- 62. Tishkoff SA, Reed FA, Ranciaro A, Voight BF, Babbitt CC, Silverman JS et al. 2006. Convergent adaptation of human lactase persistence in Africa and Europe. Nat Genet 39:31–40. pmid:17159977
- 63. Enattah NS, Jensen TGK, Nielsen M, Lewinski R, Kuokkanen M, Rasinpera H et al. 2008. Independent introduction of two lactase-persistence alleles into human populations reflects different history of adaptation to milk culture. Am J Hum Genet 82:57–72. pmid:18179885
- 64. Holden C, Mace R. 2009. Phylogenetic analysis of the evolution of lactose digestion in adults. Hum Biol 81:597–619. pmid:20504185
- 65. Hancock AM, Witonsky DB, Ehler E, Alkorta-Aranburu G, Beall C, Gebremedhin A et al. 2010. Human adaptations to diet, subsistence, and ecoregion are due to subtle shifts in allele frequency. Proc Natl Acad Sci 107:8924–8930. pmid:20445095
- 66. Peng Y, Shi H, Qi XB, Xiao CJ, Zhong H, Ma RZ et al. 2010. The ADH1B Arg47His polymorphism in East Asian populations and expansion of rice domestication in history. BMC Evol Biol 10:15. pmid:20089146
- 67. Gamba C, Jones ER, Teasdale MD, McLaughlin RL, Gonzalez-Fortes G, Mattiangeli V et al. 2014. Genome flux and stasis in a five millennium transect of European prehistory. Nat Commun 5:5257. pmid:25334030
- 68. Itan Y, Powell A, Beaumont MA, Burger J, Thomas MG. 2009. The origins of lactase persistence in Europe. PLoS Comput Biol 5(8):e1000491. pmid:19714206
- 69. Mathieson I, Lazaridis I, Rohland N, Mallick S, Llamas B, Pickrell J et al. 2015. Eight thousand years of natural selection in Europe. bioRxiv:016477
- 70. Milašinović L. 2009. Review on possibility of social structure reconstruction on the Maros culture finds from the necropolis at Ostojićevo. English summary. Rad Muzeja Voyvodina pp 65–70.
- 71. Bentley RA, Bickle P, Francken M, Gerling C, Hamilton J, Hedges R, Stephan E, Wahl J, Whittle A. 2013. Baden-Württemberg. In: Bickle P and Whittle A, editors. The first farmers of Central Europe: Diversity in LBK lifeways. pp 251–288.
- 72. Pankowská A. 2009. Comparison of health status in human skeletal remains disposal in settlements and necropolises in the early Bronze Age (in Central Moravia, Czech Republic). Anthropologie XLVII: 215–228.
- 73. Kala J, Konášová K, Smrčka V. 2008. Pohřby ze zásobních jam na okraji únětického pohřebiště v Brně-Tuřanech. [The Burials in Settlement Features on The Edge of The Early Bronze Age Burial a Settlement site in Brno Tuřany]. In: Acta Archaeologica Opaviensia. Opava: Ústav historických věd FPF Slezské Univerzity v Opavě, pp. 61–78.
- 74. Frayer DW. 2004. The dental remains from Krškany (Slovakia) and Vedrovice (Czech Republic). Anthropologie 42:71–103.
- 75. Ubelaker DH, Pap I. 1998. Skeletal evidence for health and disease in the Iron Age of Northeastern Hungary. Int J Osteoarchaeol 8:231–251.
- 76. Ubelaker DH, Pap I. 1996. Health profiles of a Bronze Age population from northeastern Hungary. Annls Hist Nat Mus Nat Hung 88:271–296.
- 77. Smrčka V, Velemínský P, Bůzek F, Zocová J. 2008. Stable C, N isotopes in human skeletal material from the Great Moravian burial site at Mikulčice-Kostelisko. In: Velemínský P, Poláček L, editors. Studien zum Burgwall von Mikulčice VIII. Brno: Archeologický ústav AV ČR. pp 169–175.
- 78. Zvelebil M, Pettitt P. 2013. Biosocial archaeology of the Early Neolithic: Synthetic analyses of a human skeletal population from the LBK cemetery of Vedrovice, Czech Republic. J Anthropol Archaeol 32:313–329.
- 79. Schutkowski H. 1995. What you are makes you eat different things—Interrelations of diet, status, and sex in the early Medieval population of Kirchheim unter Teck, FGR. Hum Evol 10:119–130.
- 80. Schutkowski H, Herrmann B, Wiedemann F, Bocherens H, Grupe G. 1999. Diet, status and decomposition at Weingarten: trace element and isotope analyses on Early Mediaeval skeletal material. J Archaeol Sci 26:675–685.
- 81. Le Huray JD, Schutkowski H. 2005. Diet and social status during the La Tène period in Bohemia: carbon and nitrogen stable isotope analysis of bone collagen from Kutná Hora-Karlov and Radovesice. J Anthropol Archaeol 24:135–147.
- 82. Le Huray JD, Schutkowski H, Richards M. 2009. Stable isotope analysis as an indicator of diet and status in La Tène Bohemia. In: Smrčka V, Walker PL, editors. Proceedings of the symposium "Social history and anthropology". Prague: Karolinum Press. pp 145–152.
- 83. Ameen S, Staub L, Ulrich S, Vock P, Ballmer F, Anderson SE. 2005. Harris lines of the tibia across centuries: A comparison of two populations, medieval and contemporary in Central Europe. Skeletal Radiol 34:279–284. pmid:15586281
- 84. Herold M. 2008. Sex differences in mortality in Lower Austria and Vienna in the Early Medieval Period: An investigation and evaluation of possible contributing factors. Universität Wien: PhD Dissertation.
- 85. Stránská P, Velemínský P, Velemínská J. 2008. The state of dentition in the Great Moravian population—A comparison of the Mikulčice centre and its hinterland. In: Velemínský P, Poláček L, editors. Studien zum Burgwall von Mikulčice VIII. Brno: Archeologický ústav AV ČR. pp 121–139.
- 86. Stránská P, Velemínský P, Poláček L. 2015. The prevalence and distribution of dental caries in four early medieval non-adult populations of different socioeconomic status from Central Europe. Arch Oral Biol 60:62–76. pmid:25255473
- 87. Tupman GS. 1962. A study of bone growth in normal children and its relationship to skeletal maturation. J Bone Joint Surg 44B: 42–67.
- 88. Ruff CB, Jones HH. 1981. Bilateral asymmetry in cortical bone of the humerus and tibia: Sex and age factors. Hum Biol 53:69–86. pmid:7239493
- 89. Cutler GB. 1997. The role of estrogen in bone growth and maturation during childhood and adolescence. J Steroid Biochem Mol Biol 61:3–6.
- 90. Stinson S. 1985. Sex differences in environmental sensitivity during growth and development. Am J Phys Anthropol 28:123–147.
- 91. Stinson S. 2012. Growth variation: Biological and cultural factors. In: Stinson S, Bogin B, O'Rourke D, editors. Human biology: An evolutionary and biocultural perspective, 2nd edition. Hoboken: John Wiley & Sons. p 587–635.
- 92. Scheeres M, Knipper C, Hauschild M, Schönfelder M, Siebel W, Pare C et al. 2014. "Celtic migrations": Fact or fiction? Strontium and oxygen isotope analysis of the Czech cemeteries of Radovesice and Kutná Hora. Am J Phys Anthropol 155:496–512.
- 93. Čižmářová J. 2006. Les nécropoles celtiques de la Moravie. In: Dossiers Archeologie et sciences des origines No. 313:26–33.
- 94. Cook DC. 1984. Subsistence and health in the lower Illinois Valley: Osteological evidence. In: Cohen MN, Armelagos GJ, editors. Paleopathology at the origins of agriculture. Orlando: Academic Press. pp 235–269.
- 95. Zakrzewski SR. 2007. Population continuity or population change: Formation of the ancient Egyptian state. Am J Phys Anthropol 132:501–509. pmid:17295300
- 96. Stock JT, O'Neill MC, Ruff CB, Zabecki M, Shackelford L, Rose J. 2011. Body size, skeletal biomechanics, mobility, and habitual activity from the Late Paleolithic to the Mid-Dynastic Nile Valley. In: Pinhasi R, Stock JT, editors. Human bioarchaeology of the transition to agriculture. London: John Wiley & Sons, Ltd. pp 347–370.
- 97. Lasker GW. 1946. Migration and physical differentiation: A comparison of immigrant with American-born Chinese. Am J Phys Anthropol 4:273–300. pmid:20276031
- 98. Greulich WW. 1957. A comparison of the physical growth and development of American-born and native Japanese children. Am J Phys Anthropol 15:489–515. pmid:13533526
- 99. Bogin BA, Smith P, Orden AB, Varela Silva MI, Loucky J. 2002. Rapid change in height and body proportions of Maya America children. Am J Hum Biol 14:753–761 pmid:12400036
- 100. Bogin BA, Rios L. 2003. Rapid morphological change in living humans: Implications for modern human origins. Comp Biochem Physiol A Mol Integr Physiol 136:71–84. pmid:14527631
- 101. Trotter M, Gleser GC. 1951. Trends in stature of American Whites and Negroes born between 1840 and 1924. Am J Phys Anthropol 9:427–440. pmid:14903067
- 102. Tanner JM, Hayashi T, Preece MA, Cameron N. 1982. Increase in length of leg relative to trunk in Japanese children and adults from 1957 to 1977: Comparison with British and with Japanese Americans. Ann Hum Biol 9:411–423. pmid:7137939
- 103. Tanner JM. 1989. Foetus into man: Physical growth from conception to maturity. 2nd ed. Ware: Castlemead Publications.
- 104. Eveleth PB. Tanner JM. 1990. Worldwide variation in human growth. 2nd ed. Cambridge: Cambridge University Press.
- 105. Kuh DL, Power C, Rodgers B. 1991. Secular trends in social class and sex differences in adult height. Int J Epidemiol 20:1001–1009. pmid:1800396
- 106. Meadows L, Jantz RL. 1995. Allometric secular change in the long bones from the 1800s to the present. J Forensic Sci 40:762–767. pmid:7595319
- 107. Jantz LM, Jantz RL. 1999. Secular change in long bone length and proportion in the United States, 1800–1970. Am J Phys Anthropol 110:57–67. pmid:10490468
- 108. Cavelaars A, Kunst AE, Geurts JJM, Crialesi R, Grötvedt L, Helmert U et al. 2000. Persistent variations in average height between countries and between socioeconomic groups: An overview of 10 European countries. Ann Hum Biol 27:407–421. pmid:10942348
- 109. Freedman DS, Kettel Khan L, Serdula MK, Srinivasan SR, Berenson GS. 2000. Secular trends in height among children during 2 decades: The Bogalusa Heart Study. Arch Pediatr Adolesc Med 154:155–161. pmid:10665602
- 110. Frisancho AR, Gilding N, Tanner S. 2001. Growth of leg length is reflected in socio-economic differences. Acta Med Auxol 33:47–50.
- 111. Gronkiewicz L. 2001. Social-class differences in Poland pronounced in stature, but absent in gene frequencies. Przeglad Antropologiczny—Anthropol Review 64:73–79.
- 112. Shin DH, Oh CS, Kim YS, Hwang YI. 2012. Ancient-to-modern secular changes in Korean stature. Am J Phys Anthropol 147:433–442. pmid:22270697
- 113. Alberman E, Filakti H, Williams S, Evans SJW, Emanuel I. 1991. Early influences on the secular change in adult height between the parents and children of the 1958 birth cohort. Ann Hum Biol 18:127–136. pmid:2024947
- 114. Silventoinen K, Kaprio J, Lahelma E, Koskenvuo M. 2000. Relative effect of genetic and environmental factors on body height: Differences across birth cohorts among Finnish men and women. Am J Pub Health 90: 627–630.
- 115. Silventoinen K, Bartels M, Posthuma D, Estourgie-van Burk GF, Willemsen G, van Beijsterveldt TC et al. 2001. Genetic regulation of growth in height and weight from 3 to 12 years of age: A longitudinal study of Dutch twin children. Twin Res Hum Genet 10:354–363.
- 116. Silventoinen K, Sammalisto S, Perola M, Boomsma DI, Cornes BK, Davis C et al. 2003. Heritability of adult body height: A comparative study of twin cohorts in eight countries. Twin Res 6:399–408. pmid:14624724
- 117. Silventoinen K. 2003. Determinants of variation in adult body height. J Biosoc Sci 35:263–285. pmid:12664962
- 118. Wadsworth MEJ, Hardy RJ, Paul AA, Marshall SF, Cole TJ. 2002. Leg and trunk length at 43 years in relation to childhood health, diet, and family circumstances: Evidence from the 1946 national birth cohort. Int J Epidemiol 31:383–390. pmid:11980800
- 119. Li L, Manor O, Power C. 2004. Early environment and child-to-adult growth trajectories in the 1958 British birth cohort. Am J Clin Nutr 80:184–192.
- 120. Jelenkovic A, Ortega-Alonso A, Rose RJ, Kaprio J, Rebato E, Silventoinen K. 2011. Genetic and environmental influences on growth from late childhood to adulthood: A longitudinal study of two Finnish twin cohorts. Am J Hum Biol 23:764–773. pmid:21957002
- 121. Golden MH. 1994. Is complete catch-up possible for stunted malnourished children? E J Clin Nutr 48:S58.
- 122. Livshits G, Roset A, Yakovenko K, Trofimov S, Kobyliansky E. 2002. Genetics of human body size and shape: Body proportions and indices. Ann Hum Biol 29:371–289.
- 123. Jenkins CL. 1981. Patterns of growth and malnutrition among preschoolers in Belize. Am J Phys Anthropol 56:169–178. pmid:6798879
- 124. Leonard WR, DeWalt KM, Stansbury JP, McCaston MK. 2000. Influence of dietary quality on the growth of highland and coastal Ecuadorian children. Am J Hum Biol 12:825–837. pmid:11534073
- 125. Katona P, Katona-Apte J. 2008. The interaction between nutrition and infection. Clin Infect Dis 46:1582–1588. pmid:18419494
- 126. Tveit-Milligan P, Spindler AA, Nichols JF. 1993. Genes and gymnastics: A case study of triplets. Sports Med, Train, Rehab 4:47–52.
- 127. Lindholm C, Hagenfeldt K, Ringertz BM. 1994. Pubertal development in elite juvenile gymnasts: Effects of physical training. Acta Obstet Gynecol Scand 73:269–273. pmid:8122511
- 128. Haapasalo H, Sievänen H, Kannus P, Heinonen A, Oja P, Vuori I. 1996. Dimensions and estimated mechanical characteristics of the humerus after long-term tennis loading. J Bone Miner Res 11:864–872. pmid:8725185
- 129. Constantini NW, Brautber C, Manny N, Ish-Shalom S. 1997. Differences in growth and maturation in twin athletes. Med Sci Sports Exerc 29:S150.
- 130. Bass SL, Bradney M, Pearce G, Hendrich E, Inge K, Stuckey S et al. 2000. Short stature and delayed puberty in gymnasts: Influence of selection bias on leg length and the duration of training on trunk length. J Pediatr 136:149–155. pmid:10657818
- 131. Prader A, Tanner JM, Von Harnack GA. 1963. Catch-up growth following illness or starvation: An example of developmental canalization in man. J Pediatr 62:646–659. pmid:13985887
- 132. Adair LS. 1999. Filipino children exhibit catch-up growth from age 2 to 12 years. J Nutr 129:1140–1148. pmid:10356078
- 133. Metcalfe NB, Monaghan P. 2001. Compensation for a bad start: Grow now, pay later? Trends Ecol Evol 16:254–260. pmid:11301155
- 134. Greulich WW. 1976. Some secular changes in the growth of American-born and native Japanese children. Am J Phys Anthropol 45:553–568. pmid:187065
- 135. Pinhasi R, Teschler-Nicola M, Knaus A, Shaw P. 2005. Cross-population analysis of the growth of long bones and the os coxae of three Early Medieval Austrian populations. Am J Hum Biol 17:470–488. pmid:15981184
- 136. Kujanová M, Bigoni L, Velemínská J, Velemínský P. 2008. Limb bones asymmetry and stress in Medieval and Recent Populations of Central Europe. Int J Osteoarch 18:476–491.
- 137. Havelková P, Villotte S, Velemínský P, Poláček L, Dobisíková M. 2011. Enthesopathies and activity patterns in the Early Medieval Great Moravian population: Evidence of division of labour. Int J Osteoarchaeol 21:487–504.
- 138. Bigoni L, Krajiček V, Sládek V, Velemínsky P, Velemínská J. 2012. Skull shape asymmetry and the socioeconomic structure of an Early Medieval Central European society. Am J Phys Anthropol 150:349–364.
- 139. Macintosh A, Pinhasi R, Stock JT. 2014. Divergence in male and female manipulative behaviors with the intensification of metallurgy in Central Europe. PLoS ONE 9(11): e112116. pmid:25389972
- 140. Buikstra JE, Ubelaker DH. 1994. Standards for data collection from human skeletal remains. Fayetteville: Arkansas Archeological Survey Research Series No. 44.
- 141. Katzmarzyk PT, Leonard WR. 1998. Climatic influences on human body size and proportions: Ecological adaptations and secular trends. Am J Phys Anthropol 106:483–503. pmid:9712477
- 142. Martorell R, Malina RM, Castillo RO, Mendoza FS, Pawson IG. 1988. Body proportions in three ethnic groups: Children and youths 2–17 years in NHANES II and HHANES. Hum Biol 60:205–222. pmid:3371962
- 143. Ruff CB. 1991. Climate and body shape in hominid evolution. J Hum Evol 21:81–105.
- 144. Ruff CB. 1994. Morphological adaptation to climate in modern and fossil hominids. Am J Phys Anthropol 37:65–107.
- 145. Holliday TW. 1997. Body proportions in Late Pleistocene Europe and modern human origins. J Hum Evol 32:423–447. pmid:9169992
- 146. Holliday TW. 1997. Postcranial evidence of cold adaptation in European Neandertals. Am J Phys Anthropol 104:245–258. pmid:9386830
- 147. Holliday TW. 1999. Brachial and crural indices of European Late Upper Paleolithic and Mesolithic humans. J Hum Evol 36:549–566. pmid:10222169
- 148. Holliday TW, Hilton CE. 2010. Body proportions of circumpolar peoples as evidenced from skeletal data: Ipiutak and Tigara (Point Hope) versus Kodiak Island Inuit. Am J Phys Anthropol 142:287–302. pmid:19927367
- 149. Temple DH, Okazaki K, Cowgill LW. 2011. Ontogeny of limb proportions in late through final Jomon period foragers. Am J Phys Anthropol 145:415–425. pmid:21541923
- 150. Martin R. 1928. Lehrbuch der anthropologie. Jena: Fischer.
- 151. Martin R. 1957. Lehrbuch der Anthropologie in Systematischer Darstellung mit Besonderer Berücksichtigung der Anthropologischen Methoden. Zweiter Band: Kraniologie Osteologie, third edition. Verlag von Gustav Fischer, Stuttgart.
- 152. Auerbach BM, Ruff CB. 2006. Limb bone bilateral asymmetry: Variability and commonality among modern humans. J Hum Evol 50:203–218. pmid:16310833
- 153. Ruff CB, Holt BM, Niskanen M, Sládek V, Berner M, Garofalo E et al. 2012. Stature and body mass estimation from skeletal remains in the European Holocene. Am J Phys Anthropol 148:601–617. pmid:22639191
- 154. Holliday TW, Ruff CB. 2001. Relative variation in human proximal and distal limb segment lengths. Am J Phys Anthropol 116:26–33. pmid:11536114
- 155. Auerbach BM, Sylvester AD. 2011. Allometry and apparent paradoxes in human limb proportions: Implications for scaling factors. Am J Phys Anthropol 144:382–391. pmid:21302265
- 156. Auerbach BM, Ruff CB. 2004. Human body mass estimation: A comparison of “morphometric” and “mechanical” methods. Am J Phys Anthropol 125:331–342. pmid:15472894
- 157. Jansson JO, Eden S, Isaksson O. 1983. Sites of action of testosterone and estradiol on longitudinal bone growth. Am J Physiol Endocrinol Metab 244:E135–E140.
- 158. Mauras N, Rogol AD, Haymond MW, Veldhuis JD. 1996. Sex steroids, growth hormone, insulin-like growth factor-I: Neuroendocrine and metabolic regulation in puberty. Horm Res Paediatr 45:74–80.
- 159. Relethford JH, Hodges DC. 1985. A statistical test for differences in sexual dimorphism between populations. Am J Phys Anthropol 66:55–61. pmid:3976870
- 160. Nikitovic D, Bogin B. 2014. Ontogeny of sexual size dimorphism and environmental quality in Guatemalan children. Am J Hum Biol 26:117–123. pmid:24302534
- 161. Chakraborty R, Majumder PP. 1982. On Bennett's measure of sex dimorphism. Am J Phys Anthropol 59:295–298. pmid:7158663
- 162. Mascie-Taylor CGN. 2010. Research designs and statistical methods in both labouratory and field settings. In: Mascie-Taylor CGN, Yasukouchi A, Ulijaszek S, editors. Human variation: From the labouratory to the field. Boca Raton FL: CRC Press Taylor and Francis Group. pp 239–268.
- 163. Cohen J. 1988. Statistical power analysis for the behavioural sciences, 2nd edition. New York: Academic Press.
- 164. Cohen J. 1992. A power primer. Psychol Bull 112:155–159. pmid:19565683
- 165. Trotter M, Goldine G. 1952. Estimation of stature from long bones of American Whites and Negroes. Am J Phys Anthropol 10:463–514. pmid:13007782
- 166. Borić D, Dimitrijević V. 2007. When did the 'Neolithic package' reach Lepenski Vir? Radiometric and faunal evidence. Documenta Praehistorica 34:53–72.
- 167. Borić D. 2011. Adaptations and Transformations of the Danube Gorges Foragers (c. 13.000–5500 BC): An overview. In: Krauß R, editor. Archaeolical and Anthropological Sciences Beginnings—New research in the appearance of the Neolithic between Northwest Anatolia and the Carpathian Basin; Papers of the International Workshop 8th–9th April 2009, Istanbul. Leidorf, Rahden/Westf. pp 157–203.
- 168. Garašanin M, Radovanović I. 2001. A pot in house 54 at Lepenski Vir I. Antiquity 75:118–125.
- 169. Borić D. 2006. Cultural continuities and ruptures: Tradition, social memory and representation in Balkan prehistory I. Lisbon: Union of Pre- and Protohistoric Sciences.
- 170. Bonsall C, Macklin MG, Payton RW, Boroneanţ A. 2002. Climate, floods and river gods: Environmental change and the Meso- Neolithic transition in Southeast Europe. Before Farming 3–4:1–15.
- 171. Borić D, Grupe G, Peters J, Mikić Ž. 2004. Is the Mesolithic-Neolithic subsistence dichotomy real? New stable isotope evidence from the Danube gorges. Eur J Archaeol 7:221–248.
- 172. Pinhasi R., Bourbou C. 2008. How representative are human skeletal assemblages for population analysis and interpretation? Implications for palaeopathological and palaeoepidemiological investigations. In: Pinhasi R, Mays S, editors. Advances in Palaeopathology: Methodological and Biocultural Perspectives. New York: Wiley-Liss. pp. 31–44.
- 173. Meiklejohn C, Shentag CT, Venema A, Key P. 1984. Socioeconomic change and patterns of pathology and variation in the Mesolithic and Neolithic of Western Europe: Some suggestions. In: Cohen MN, Armelagos G, editors. Palaeopathology at the Origins of Agriculture. London: Academic Press. pp 75–100.
- 174. Stini WA. 1969. Nutritional stress and growth: Sex difference in adaptive response. Am J Phys Anthropol 31:417–426. pmid:4313260
- 175. Vercellotti G, Stout SD, Boano R, Sciulli PW. 2011. Intrapopulation variation in stature and body proportions: Social status and sex differences in an Italian medieval population (Trino Vercellese, VC). Am J Phys Anthropol 145:203–214. pmid:21312185
- 176. Banner J. 1956. Die Péceler Kultur. Budapest: Archaeologia Hungarica.
- 177. Němejcová-Pavúkova V. 1973. Zu Ursprung und Chronologie der Boleraz-Gruppe. In: Chropovský B, editor. Symposium über die Entstehung und Chronologie der Badener Kultur. Bratislava: Verlag der slowakischen Akademie der Wissenschaften. pp 297–316.
- 178. Kalicz N. 1976. Novoya nakhodka modeli povozki epokhi eneolita iz okresnostej Budapeshta. Sov Archeol 2:106–117.
- 179. Bakker JA, Kruk J, Lanting AE, Milisauskas S. 1999. Bronocice, Flintbek, Uruk and Jebel Aruda: The earliest evidence of wheeled vehicles in Europe and the Near East. Antiquity 73:778–790.
- 180. Greenfield HJ. 2001. European Early Bronze Age. In: Peregrine P, Ember M, editors. Encyclopaedia of Prehistory: Vol. 4, Europe, for the Human Area Relations Files. New York: Kluwer Academic/Plenum Publishers. pp 124–137.
- 181. Sherratt A. 2006. La traction animale et la transformation de l’Europe néolithique: In: Pétrequin P, Arbogast R-M, Pétrequin A-M, Van Willigen S, Bailly M, editors. Premiers chariots, premiers araires. La traction animale en Europe pendant les ive et les millénaires avant notre ére. Paris: CNRS Éditions. pp 329–360.
- 182. Dani J, Máthé M, Szabö G. 2001. Ausgrabungen in der bronzezeitlichen Tell-Siedlung und im Gräberfeld von Polgár-Kenderföld (Vorbericht über die Freilegung des mittelbronzezeitlichen Gräberfeldes von Polgár-Kenderföld, Majoros-Tanya). In: Kacsó C, editor. Bronzezeitliche Kulturerscheinungen im Karpatischen Raum. Die Beziehungen zu den benachbarten Gebieten. Ehrensymphosium für Alexandru Vulpe zum 70. Geburtstag, Baia Mare Oktober 10–13, 2001. pp 93–118.
- 183. Sosna D. 2007. Social differentiation in the Late Copper Age and the Early Bronze Age in South Moravia (Czech Republic). Electronic Theses, Treatises and Dissertations. Paper 1621. The Florida State University.
- 184. Allentoft ME, Sikora M, Sjögren KG, Rasmussen S, Rasmussen M, Stenderup J et al. 2015. Population genomics of Bronze Age Eurasia. Nature 522:167–174. pmid:26062507
- 185. Haak W, Forster P, Bramanti B, Matsumura S, Brandt G, Tänzer M et al. 2005. Ancient DNA from the first European farmers in 7500-year-old Neolithic sites. Science 310:1016–1018. pmid:16284177
- 186. Olalde I, Lalueza-Fox C. 2015. Modern humans' paleogenomics and the new evidences on the European prehistory. STAR 1: STAR20152054892315Y.0000000002
- 187. Metzner-Nebelsick C. 2010. Aspects of mobility and migration in the Eastern Carpathian Basin and adjacent areas in the Early Iron Age (10th-7th centuries BC). In: Dzięgielewski K, Przybyła MS, Gawlik A, editors. Migration in Bronze and Early Iron Age Europe. Kraków: Jagiellonian University Institute of Archaeology. pp 121–152.
- 188. Tasić N. 2004. Istorijska slika razvoja ranog gvozdenog doba u srpskom Podunavlju. [Historical picture of development of Early Iron Age in the Serbian Danube Basin]. Balcanica XXXV:5–23.
- 189. Tasić N. 2003–2004. Historical picture of development of Bronze Age cultures in Vojvodina. Starinar 53–54:25–34.
- 190. Rolle R. 1989. The world of the Scythians. Berkeley and Los Angeles: University of California Press.
- 191. Dezső G. 1966. A population of the Scythian period between the Danube and the Tisza. Anthropol Hung VII:36–57
- 192. Zoffmann ZK. 2011. Antropološka analiza skeleta iz ranog gvozdenog sa arheološkog lokaliteta Vranj-Hrtkovci. Rad Muzeja Vojvodine 53:93–96.
- 193. Friesinger H. 1972. Frühmittelalterliche Körpergräber aus Pottenbrunn, Stadtgemeinde St. Pölten, NÖ. Archaeol Austriaca 51:113–189.
- 194. Tasić N. 1972. An Early Iron Age collective tomb at Gomolava. Archaeol Iugosl 13:27–37.
- 195. Podborský V. 1993. Pravěké dějiny Moravy. Brno: Muzejní a Vlastivĕdná Společnost v Brně.
- 196. Frey OH. 2004. A new approach to Early Celtic art. Proceedings of the Royal Irish Academy, Section C: Archaeology, Celtic Studies, History, Linguistics, Literature 104C:107–129.
- 197. Pettitt P, Hedges R. 2008. The age of the Vedrovice cemetery: The AMS radiocarbon dating programme. Anthropologie 46:125–134.
- 198. Borić D. 2009. Absolute dating of metallurgical innovations in the Vinča culture of the Balkans. In: Kienlin TL, Roberts BW, editors. Metals and Societies: Studies in honour of Barbara S. Ottaway. Bonn: Habelt. pp 191–245.
- 199. Bentley RA, Bickle P, Fibiger L, Nowell GM, Dale CW, Hedges REM et al. 2012. Community differentiation and kinship among Europe’s first farmers. P Natl Acad Sci 109:9326–9330.
- 200. Fischl KP, Kiss V, Kulcsár G, Szeverényi V. 2013. Transformations in the Carpathian Basin around 1600 B. C. In: Meller H, editor. 1600 BC—Cultural change in the shadow of the Thera-Eruption? Halle: Landesamt für Denkmalpflege und Archäologie Sachsen-Anhalt—Landesmuseum für Vorgeschichte. pp 355–372.
- 201. Whittle A, Anders A, Bentley RA, Bickle P, Cramp L, Domboróczki L et al. 2013. Hungary. In: Bickle P, Whittle A, editors. The first farmers of Central Europe: Diversity in LBK lifeways. Oxford: Oxbow Books. pp 49–100.
- 202. Bonsall C, Vasić R, Boroneant A, Roksandic M, Soficaru A, McSweeney K, Evatt A, Aguraiuja Ü, Pickard C, Dimitrijević V, Higham T, Hamilton D, Cook G. 2015. New AMS 14C dates for human remains from Stone Age sites in the Iron Gates reach of the Danube, Southeast Europe. Radiocarbon 57:33–46.
- 203. Drozdová E, Balueva TS, Veselovskaya E, Smrčka V, Benešová J, Bůzek F, Erban V, Kanický V, Ovesná P, Nejezchlebová H, Vaňharová M, Zocová J, Matějíčková A. 2011. Hoštice I za Hanou. Výsledky antropologické analýzy pohřebiště lidu kultury zvoncovitých pohárů. [Hoštice I za Hanou. Results of anthropological survey of human skeletal remains dated to Bell Beaker culture. 1. vyd. Brno: Masarykova univerzita.
- 204. Raczky P, Anders A. 2009. Settlement history of the Middle Neolithic in the Polgár micro-region (The development of the Alföld Linearband Pottery in the Upper Tisza Region, Hungary). In: Kozłowski JK, editor. Interactions between different models of Neolithization north of the Central European agro-ecological barrier. Papers presented on the symposium organized by the EU project FEPRE. Kraków: Polska Akademia Umiejętności. pp 31–50.