Craniometric data support a mosaic model of demic and cultural Neolithic diffusion to outlying regions of Europe

The extent to which the transition to agriculture in Europe was the result of biological (demic) diffusion from the Near East or the adoption of farming practices by indigenous hunter-gatherers is subject to continuing debate. Thus far, archaeological study and the analysis of modern and ancient European DNA have yielded inconclusive results regarding these hypotheses. Here we test these ideas using an extensive craniometric dataset representing 30 hunter-gatherer and farming populations. Pairwise population craniometric distance was compared with temporally controlled geographical models representing evolutionary hypotheses of biological and cultural transmission. The results show that, following the physical dispersal of Near Eastern/Anatolian farmers into central Europe, two biological lineages were established with limited gene flow between them. Farming communities spread across Europe, while hunter-gatherer communities located in outlying geographical regions adopted some cultural elements from the farmers. Therefore, the transition to farming in Europe did not involve the complete replacement of indigenous hunter-gatherer populations despite significant gene flow from the Southwest Asia. This study suggests that a mosaic process of dispersal of farmers and their ideas was operating in outlying regions of Europe, thereby reconciling previously conflicting results obtained from genetic and archaeological studies.     

Radiocarbon dating the late Middle Paleolithic and the Aurignacian of the Swabian Jura

Many lines of evidence point to the period between roughly 40 and 30 ka BP as the period in which modern humans arrived in Europe and displaced the indigenous Neandertal populations. At the same time, many innovations associated with the Upper Paleolithic--including new stone and organic technologies, use of personal ornaments, figurative art, and musical instruments--are first documented in the European archaeological record. Dating the events of this period is challenging for several reasons. In the period about six to seven radiocarbon half-lives ago, variable preservation, pre-treatment, and sample preparation can easily lead to a lack of reproducibility between samples and laboratories. A range of biological, cultural, and geological processes can lead to mixing of archaeological strata and their contents. Additionally, some data sets point to this period as a time of significant spikes in levels of atmospheric radiocarbon. This paper assesses these questions in the context of the well-excavated and intensively studied caves of Geissenkl?sterle and Hohle Fels in the Swabian Jura of southwestern Germany. We conclude that variable atmospheric radiocarbon production contributes to the problems of dating the late Middle Paleolithic and the early Upper Paleolithic. To help establish a reliable chronology for the Swabian Aurignacian, we are beginning to focus our dating program on short-lived, stratigraphically secure features to see if they yield reproducible results. This approach may help to test competing explanations for the noisy and often non-reproducible results that arise when trying to date the transition from the Middle to the Upper Paleolithic.     

Craniometric Data Supports Demic Diffusion Model for the Spread of Agriculture into Europe

Background

The spread of agriculture into Europe and the ancestry of the first European farmers have been subjects of debate and controversy among geneticists, archaeologists, linguists and anthropologists. Debates have centred on the extent to which the transition was associated with the active migration of people as opposed to the diffusion of cultural practices. Recent studies have shown that patterns of human cranial shape variation can be employed as a reliable proxy for the neutral genetic relationships of human populations.

Methodology/Principal Findings

Here, we employ measurements of Mesolithic (hunter-gatherers) and Neolithic (farmers) crania from Southwest Asia and Europe to test several alternative population dispersal and hunter-farmer gene-flow models. We base our alternative hypothetical models on a null evolutionary model of isolation-by-geographic and temporal distance. Partial Mantel tests were used to assess the congruence between craniometric distance and each of the geographic model matrices, while controlling for temporal distance. Our results demonstrate that the craniometric data fit a model of continuous dispersal of people (and their genes) from Southwest Asia to Europe significantly better than a null model of cultural diffusion.

Conclusions/Significance

Therefore, this study does not support the assertion that farming in Europe solely involved the adoption of technologies and ideas from Southwest Asia by indigenous Mesolithic hunter-gatherers. Moreover, the results highlight the utility of craniometric data for assessing patterns of past population dispersal and gene flow.     

The impossible coincidence. A single‐species model for the origins of modern human behavior in Europe

Few topics in palaeoanthropology have generated more recent debate than the nature and causes of the remarkable transformation in human behavioral patterns that marked the transition from the Middle to the Upper Paleolithic in Europe. 1 - 11 Those of us who have argued for an effective technological and cultural “revolution” at this point in the Paleolithic sequence have emphasized three main dimensions 1 , 2 , 9 , 11 - 14 : the wide range of different aspects of behavior that appear to have been affected (Fig. 1); the relative speed and abruptness with which most of these changes can be documented in the archeological records from the different regions of Europe; and the potentially profound social and cognitive implications of many of the innovations involved. Most striking of all in this context is the abrupt appearance and proliferation of various forms of perforated animal teeth, shells, beads, and other personal ornaments, and the even more dramatic eruption of remarkably varied and sophisticated forms of art, ranging from representations of male and female sex organs, through the highly stylized animal and combined animal‐human figures from southern Germany, to the striking wall paintings of the Chauvet Cave. 8 , 15 - 18 One might add to this the similar proliferation of more enigmatic but potentially equally significant abstract “notation” systems on bone and ivory artifacts. 19 To describe the Upper Paleolithic revolution in Europe as reflecting preeminently an explosion in explicitly symbolic behavior and expression is in no sense an exaggeration, as most prehistorians would now agree. We are probably on safe ground in assuming that symbolic behavior and expression of this level of complexity would be inconceivable in the absence of highly structured language systems and brains closely similar, if not identical to, our own. 5 , 17 , 20 - 28 .     

Tracing the origin and spread of agriculture in Europe

The origins of early farming and its spread to Europe have been the subject of major interest for some time. The main controversy today is over the nature of the Neolithic transition in Europe: the extent to which the spread was, for the most part, indigenous and animated by imitation (cultural diffusion) or else was driven by an influx of dispersing populations (demic diffusion). We analyze the spatiotemporal dynamics of the transition using radiocarbon dates from 735 early Neolithic sites in Europe, the Near East, and Anatolia. We compute great-circle and shortest-path distances from each site to 35 possible agricultural centers of origin—ten are based on early sites in the Middle East and 25 are hypothetical locations set at 5° latitude/longitude intervals. We perform a linear fit of distance versus age (and vice versa) for each center. For certain centers, high correlation coefficients (R > 0.8) are obtained. This implies that a steady rate or speed is a good overall approximation for this historical development. The average rate of the Neolithic spread over Europe is 0.6–1.3 km/y (95% confidence interval). This is consistent with the prediction of demic diffusion (0.6–1.1 km/y). An interpolative map of correlation coefficients, obtained by using shortest-path distances, shows that the origins of agriculture were most likely to have occurred in the northern Levantine/Mesopotamian area.     

Complete mitochondrial genomes reveal neolithic expansion into Europe

The Neolithic transition from hunting and gathering to farming and cattle breeding marks one of the most drastic cultural changes in European prehistory. Short stretches of ancient mitochondrial DNA (mtDNA) from skeletons of pre-Neolithic hunter-gatherers as well as early Neolithic farmers support the demic diffusion model where a migration of early farmers from the Near East and a replacement of pre-Neolithic hunter-gatherers are largely responsible for cultural innovation and changes in subsistence strategies during the Neolithic revolution in Europe. In order to test if a signal of population expansion is still present in modern European mitochondrial DNA, we analyzed a comprehensive dataset of 1,151 complete mtDNAs from present-day Europeans. Relying upon ancient DNA data from previous investigations, we identified mtDNA haplogroups that are typical for early farmers and hunter-gatherers, namely H and U respectively. Bayesian skyline coalescence estimates were then used on subsets of complete mtDNAs from modern populations to look for signals of past population expansions. Our analyses revealed a population expansion between 15,000 and 10,000 years before present (YBP) in mtDNAs typical for hunters and gatherers, with a decline between 10,000 and 5,000 YBP. These corresponded to an analogous population increase approximately 9,000 YBP for mtDNAs typical of early farmers. The observed changes over time suggest that the spread of agriculture in Europe involved the expansion of farming populations into Europe followed by the eventual assimilation of resident hunter-gatherers. Our data show that contemporary mtDNA datasets can be used to study ancient population history if only limited ancient genetic data is available.     

Neandertals and moderns mixed,and it matters

Twenty‐five years ago, the Middle‐to‐Upper Paleolithic transition in Europe could be represented as a straightforward process subsuming both the emergence of symbolic behavior and the replacement of Neandertals by modern humans. The Aurignacian was a proxy for the latter, during which enhanced cognitive capabilities explained ornaments and art. The few instances of Neandertal symbolism were deemed to long postdate contact and dismissed as “imitation without understanding,” if not geological contamination. Such views were strengthened by the recent finding that, in southern Africa, several features of the European Upper Paleolithic, including bone tools, ornaments, and microliths, emerged much earlier. Coupled with genetic suggestions of a recent African origin for extant humans, fossil discoveries bridging the transition between “archaics” and “moderns” in the realm of anatomy (Omo‐Kibish, Herto) seemingly closed the case. Over the last decade, however, taphonomic critiques of the archeology of the transition have made it clear that, in Europe, fully symbolic sapiens behavior predates both the Aurignacian and moderns. And, in line with evidence from the nuclear genome rejecting strict replacement models based on mtDNA alone, the small number of early modern specimens that passed the test of direct dating present archaic features unknown in the African lineage, suggesting admixture at the time of contact. In the realm of culture, the archeological evidence also supports a Neandertal contribution to Europe's earliest modern human societies, which feature personal ornaments completely unknown before immigration and are characteristic of such Neandertal‐associated archeological entities as the Châtelperronian and the Uluzzian. The chronometric data suggest that, north of the Ebro divide, the entire interaction process may have been resolved within the millennium centered around 42,000 calendar years ago. Such a rapid absorption of the Neandertals is consistent with the size imbalance between the two gene reservoirs and further supports significant levels of admixture.     

旧石器时代装饰品研究:现状与意义

装饰品(personal ornament)是旧石器时代中晚期遗址中一类较特殊的遗存,作为早期人类现代行为的重要标志,它的出现及其学术意义一直受到国际学术界的关注和重视。西方考古学界对装饰品的研究开始较早,理论方法成熟,成果丰硕。我国此遗存发现较少,在研究手段和程度方面与西方存在一定的差距。本文从装饰品的定义与分类、发现与研究现状、功能与作用、出现原因及研究意义的探讨等方面入手,对西方学术界对装饰品的研究现状和进展进行梳理,对我国发现的相关材料与研究进展做简要介绍与评述,希望能对我国旧石器时代装饰品的研究提供有价值的参考资料与思路。     

Space Competition and Time Delays in Human Range Expansions. Application to the Neolithic Transition

Space competition effects are well-known in many microbiological and ecological systems. Here we analyze such an effect in human populations. The Neolithic transition (change from foraging to farming) was mainly the outcome of a demographic process that spread gradually throughout Europe from the Near East. In Northern Europe, archaeological data show a slowdown on the Neolithic rate of spread that can be related to a high indigenous (Mesolithic) population density hindering the advance as a result of the space competition between the two populations. We measure this slowdown from a database of 902 Early Neolithic sites and develop a time-delayed reaction-diffusion model with space competition between Neolithic and Mesolithic populations, to predict the observed speeds. The comparison of the predicted speed with the observations and with a previous non-delayed model show that both effects, the time delay effect due to the generation lag and the space competition between populations, are crucial in order to understand the observations.     

The invisible frontier. A multiple species model for the origin of behavioral modernity

Two contradictory theories of human cognitive evolution have been developed to model how, when, and among what hominid groups behavioral modernity emerged. The first model, which has long been the dominant paradigm, links these behavioral innovations to a cultural “revolution” by anatomically modern humans in Europe at around 40,000 years ago, coinciding with the first arrival of our species in this region.^1–4 According to this model, the sudden and explosive character of this change is demonstrated by the appearance in the archeological record of previously unseen carvings, personal ornaments, musical instruments, depictions on cave walls, and new stone and bone technology. A variant of this model sees behavioral modernity resulting from a rapid biological change, a brain mutation producing no apparent change in skull anatomy, which occurred in Europe or, more probably, in Africa at ca. 50,000 years ago.⁵⁶.     

Gravettian ivory ornaments in Central Europe,Moravia (Czech Republic)

In the sense of the gravettian technocomplex of Central Europe in Moravia it is mammoth ivory which was mainly used to manufacture and create personal ornaments. The hunters and gatherers selected and stored mammoth tusks as the raw material for the manufacture of tools, weapons, furniture and ornaments. In Moravia, personal ornaments were found in a great variability of forms and sizes – mostly beads like breast-shaped beads, bilobated beads, bilobated flat beads, tear drop shaped beads and round ornaments – rings, bracelets, discs as well as flat elements – diadems and other pendants (zoomorphs, antropomorphs, geometric forms). Some types of personal ornaments show equivalent forms at other gravettian sites and some are unique in Moravia. This article illustrates the variability of these personal ornaments in the context of the Gravettian in Moravia.     

The oldest anatomically modern humans from far southeast Europe: direct dating, culture and behavior

Background

Anatomically Modern Humans (AMHs) are known to have spread across Europe during the period coinciding with the Middle to Upper Paleolithic transition. Whereas their dispersal into Western Europe is relatively well established, evidence of an early settlement of Eastern Europe by modern humans are comparatively scarce.

Methodology/Principal Finding

Based on a multidisciplinary approach for the study of human and faunal remains, we describe here the oldest AMH remains from the extreme southeast Europe, in conjunction with their associated cultural and paleoecological background. We applied taxonomy, paleoecology, and taphonomy combined with geomorphology, stratigraphy, archeology and radiocarbon dating. More than 160 human bone remains have been discovered. They originate from a well documented Upper Paleolithic archeological layer (Gravettian cultural tradition) from the site of Buran-Kaya III located in Crimea (Ukraine). The combination of non-metric dental traits and the morphology of the occipital bones allow us to attribute the human remains to Anatomically Modern Humans. A set of human and faunal remains from this layer has been radiocarbon dated by Accelerator Mass Spectrometry. The direct-dating results of human bone establish a secure presence of AMHs at 31,900+240/−220 BP in this region. They are the oldest direct evidence of the presence of AMHs in a well documented archeological context. Based on taphonomical observations (cut marks and distribution of skeletal elements), they represent the oldest Upper Paleolithic modern humans from Eastern Europe, showing post-mortem treatment of the dead as well.

Conclusion/Significance

These findings are essential for the debate on the spread of modern humans in Europe during the Upper Paleolithic, as well as their cultural behaviors.     

Skull and limb morphology differentially track population history and environmental factors in the transition to agriculture in Europe

The Neolithic transition in Europe was a complex mosaic spatio-temporal process, involving both demic diffusion from the Near East and the cultural adoption of farming practices by indigenous hunter–gatherers. Previous analyses of Mesolithic hunter–gatherers and Early Neolithic farmers suggest that cranial shape variation preserves the population history signature of the Neolithic transition. However, the extent to which these same demographic processes are discernible in the postcranium is poorly understood. Here, for the first time, crania and postcranial elements from the same 11 prehistoric populations are analysed together in an internally consistent theoretical and methodological framework. Results show that while cranial shape reflects the population history differences between Mesolithic and Neolithic lineages, relative limb dimensions exhibit significant congruence with environmental variables such as latitude and temperature, even after controlling for geography and time. Also, overall limb size is found to be consistently larger in hunter–gatherers than farmers, suggesting a reduction in size related to factors other than thermoregulatory adaptation. Therefore, our results suggest that relative limb dimensions are not tracking the same demographic population history as the cranium, and point to the strong influence of climatic, dietary and behavioural factors in determining limb morphology, irrespective of underlying neutral demographic processes.     

Cultural Diffusion Was the Main Driving Mechanism of the Neolithic Transition in Southern Africa

It is well known that the Neolithic transition spread across Europe at a speed of about 1 km/yr. This result has been previously interpreted as a range expansion of the Neolithic driven mainly by demic diffusion (whereas cultural diffusion played a secondary role). However, a long-standing problem is whether this value (1 km/yr) and its interpretation (mainly demic diffusion) are characteristic only of Europe or universal (i.e. intrinsic features of Neolithic transitions all over the world). So far Neolithic spread rates outside Europe have been barely measured, and Neolithic spread rates substantially faster than 1 km/yr have not been previously reported. Here we show that the transition from hunting and gathering into herding in southern Africa spread at a rate of about 2.4 km/yr, i.e. about twice faster than the European Neolithic transition. Thus the value 1 km/yr is not a universal feature of Neolithic transitions in the world. Resorting to a recent demic-cultural wave-of-advance model, we also find that the main mechanism at work in the southern African Neolithic spread was cultural diffusion (whereas demic diffusion played a secondary role). This is in sharp contrast to the European Neolithic. Our results further suggest that Neolithic spread rates could be mainly driven by cultural diffusion in cases where the final state of this transition is herding/pastoralism (such as in southern Africa) rather than farming and stockbreeding (as in Europe).     

Applying the ecosystem services framework to pasture-based livestock farming systems in Europe

The concept of ‘Ecosystem Services’ (ES) focuses on the linkages between ecosystems, including agroecosystems, and human well-being, referring to all the benefits, direct and indirect, that people obtain from ecosystems. In this paper, we review the application of the ES framework to pasture-based livestock farming systems, which allows (1) regulating, supporting and cultural ES to be integrated at the same level with provisioning ES, and (2) the multiple trade-offs and synergies that exist among ES to be considered. Research on livestock farming has focused mostly on provisioning ES (meat, milk and fibre production), despite the fact that provisioning ES strongly depends on regulating and supporting ES for their existence. We first present an inventory of the non-provisioning ES (regulating, supporting and cultural) provided by pasture-based livestock systems in Europe. Next, we review the trade-offs between provisioning and non-provisioning ES at multiple scales and present an overview of the methodologies for assessing biophysical trade-offs. Third, we present non-biophysical (economical and socio-cultural) methodologies and applications for ES valuation. We conclude with some recommendations for policy design.     

How mankind has shaped European nature

A monographic book volume on the history of the rural landscape of Europe is presented (Emanuelsson, The rural landscape of Europe. How man has shaped European nature, 383, 2009). The focus is on various land-use phases, their history from the Neolithic period to modern time and the survival of traditional farming today in Europe. This is described and illustrated by examples from marginal farming areas, particularly in mountainous regions of southern and northern Europe. Some of these are important as modern analogues when interpreting past cultural landscapes.     

Reconstructing range dynamics and range fragmentation of European bison for the last 8000 years

Aim Understanding what constituted species’ ranges prior to large‐scale human influence, and how past climate and land use change have affected range dynamics, provides conservation planners with important insights into how species may respond to future environmental change. Our aim here was to reconstruct the Holocene range of European bison (Bison bonasus) by combining a time‐calibrated species distribution models (SDM) with a dynamic vegetation model. Location Europe. Method We used European bison occurrences from the Holocene in a maximum entropy model to assess bison range dynamics during the last 8000 years. As predictors, we used bioclimatic variables and vegetation reconstructions from the generalized dynamic vegetation model LPJ‐GUESS. We compared our range maps with maps of farmland and human population expansion to identify the main species range constraints. Results The Holocene distribution of European bison was mainly determined by vegetation patterns, with bison thriving in both broadleaved and coniferous forests, as well as by mean winter temperature. The heartland of European bison was in Central and Eastern Europe, whereas suitable habitat in Western Europe was scarce. While environmentally suitable regions were overall stable, the expansion of settlements and farming severely diminished available habitat. Main conclusions European bison habitat preferences may be wider than previously assumed, and our results suggest that the species had a more eastern and northern distribution than previously reported. Vegetation and climate transformation during the Holocene did not affect the bison’s range substantially. Conversely, human population growth and the spread of farming resulted in drastic bison habitat loss and fragmentation, likely reaching a tipping point during the last 1000 years. Combining SDM and dynamic vegetation models can improve range reconstructions and projections, and thus help to identify resilient conservation strategies for endangered species.     

Plants as human adornment in India

Plants have served for human adornment in India for millennia. Their use as ornaments and cosmetics is not only ancient but survives to the present time. In addition to decoration, adornment is often regarded as having amuletic powers or is used as social diacritical marks. Over 165 plant species used for human adornment in India have been identified from the literature and by personal observations. This study points out the importance of plants as human adornment both to aboriginal groups and modern urban and rural Indians. The widespread employment of vegetal materials for adornment indicates that in India they have traditionally been more important for this use than have mineral or animal substances.     

Environmental Change and Extended Phenotypes: Does Eutrophication Influence Nest Building in Sticklebacks?

Many species use extended phenotypes, such as purpose‐built nests, to increase their reproductive success. These traits have to be adjusted to local environmental conditions to maximize fitness. An important question is whether species are able to adjust their extended phenotypes to human‐induced rapid environmental changes. We investigated whether populations of threespine stickleback, Gasterosteus aculeatus, exposed to different degrees of human‐induced eutrophication during the last decades, have differentiated phenotypically in their nest‐building behaviour. Stickleback males build nests that they use both in mate attraction and for offspring protection and whose characteristics vary with environmental conditions. We allowed males from parallel pairs of mildly and severely eutrophied habitats to build nests under standardized conditions in the laboratory. We recorded the time it took the males to build a nest, the size and neatness of the completed nest and the use of nest ornaments. We found eutrophication at the site of capture to influence nest‐building time – males from eutrophied habitats built faster. However, eutrophication did not alter nest structure or the use of nest ornaments. This is probably because the nests are concealed in vegetation or under stones and females cannot evaluate them before they have followed the male to the nest, and predators cannot detect them before close to the nest. Thus, reduced long‐range visibility does not influence the use of nests as mate choice cues or for offspring protection. This contrasts with a recorded effect of eutrophication on courtship behaviour, whose efficiency depends on long‐range visibility. This suggests that traits are adjusted to eutrophication depending on the influence of eutrophication on the function of the traits.     

Evolution of culture-dependent discriminate sociality: a gene-culture coevolutionary model

Animals behave cooperatively towards certain conspecifics while being indifferent or even hostile to others. The distinction is made primarily according to kinship as predicted by the kin selection theory. With regards to humans, however, this is not always the case; in particular, humans sometimes exhibit a discriminate sociality on the basis of culturally transmitted traits, such as personal ornaments, languages, rituals, etc. This paper explores the possibility that the human faculty of cultural transmission and resultant cultural variation among individuals may have facilitated the evolution of discriminate sociality in humans. To this end, a gene-culture coevolutionary model is developed focusing on competition over control of resource as a context in which discriminate sociality may have evolved. Specifically, two types of culture-dependent discriminate sociality are considered: ingroup favouritism, with ingroup and outgroup being distinguished by the presence or absence of a cultural trait; and prestige hierarchies, with the prestige being conferred on the bearer of a cultural trait. The model specifies the conditions under which emergence and evolutionary stability of the two types of discriminate sociality are promoted by the presence of cultural variation among individuals.