Culture Matters in the Neolithic Transition and Emergence of Hierarchy in Thy, Denmark: Distinguished Lecture

How did the emergence of hierarchical social structure that followed the domestication of plants and animals in the Neolithic actually come about? I suggest that material media were instrumental in this transformation, as culture was changed by incorporating such newmedia as landscape constructions and elaborate prestige objects. During the Neolithic transition in Thy, Denmark, local corporate groups formed, and, subsequently, Bronze Age chieftains came to power. Shifts in material culture suggest possible connections to these institutional changes, namely the materialization of property rights by burial monuments and permanent domestic architecture and the centralization of power through the controlled production of wealth objects. I conclude that, as part of social process, the nature of culture has been transformed by incorporating material culture with specific characteristics of scale, permanency, and control that were vital to institutional change.     

A comparison of early Neolithic crop and weed assemblages from the Linearbandkeramik and the Bulgarian Neolithic cultures: differences and similarities

The spread of early agriculture from the Mediterranean to central Europe is still poorly understood. The new subsistence reached western central Europe during the second half of the 6th millennium cal b.c. This paper presents a comparison of crop and weed species from 33 Bandkeramik sites from Austria and Germany and six Bulgarian Neolithic sites. The aim is to investigate whether the early cultivation system brought in from the eastern Mediterranean was adapted to European conditions in Bulgaria or further West. Some characteristics of the potential weeds are interpreted with respect to the cultivation systems and the origin of the species.     

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.     

Evidence for plant exploitation and vegetation history from three Early Neolithic pre-pottery sites on the Euphrates (Syria)

Archaeobotanical results based on a limited number of samples from three aceramic sites dating from 9800 to 7800 B.P., which are under excavation in the valley of the Middle Euphrates, are discussed. The finds are presented simply by presence, and are compared to the contemporary vegetation and finds from similar sites. Carbonised plant remains recovered by flotation from levels dated to between 9800 and 9200 B.P. (Dja'de and Jerf al Ahmar) indicate that wild cereals (einkorn wheat, rye and barley) and pulses (lentils, pea and bitter vetch) were exploited. Other plants such as wild grasses, Pistacia, wild almond and oak, suggest that the local vegetation provided a rich diversity of resources. A study of possible weed taxa is being carried out in order to see whether this assemblage could be used to identify the cultivation of morphologically wild cereals for this period. Ninth millennium B.P. levels at Halula see the appearance of domestic crops such as emmer, naked wheat and barley, but wild-type cereals persist. The cultivars appear to have been introduced from elsewhere and later ninth millennium B.P. species include olive and flax. Ash, vine, maple, plane, alder and elm from the gallery forest, wild rye, wild einkorn, deciduous oak, wild almond, Pistacia, and Pyrus, from the hinterland, indicate cooler conditions.     

The history of crops and crop weeds in south-western Germany from the Neolithic period to modern times, as shown by archaeobotanical evidence

The results of more than 100 archaeobotanical investigations from south-western Germany have been arranged into chronological groups and analyzed with a spreadsheet to obtain presence data and the numbers of taxa present in the various time periods. They show an increase in the number of plant taxa cultivated in gardens since the Roman period, a decrease in the crops that were the earliest to be cultivated, especially emmer, and an increase of spelt, rye and oats in the course of time. New crop weeds appear, including an increasing number of southern plants originating from open vegetation. This is more likely to be a result of the intensification of agriculture than from increased human mobility. The crop weeds, grouped together according to their requirements for soil nutrients, show increasing soil acidification through the course of time.     

Anthracological analysis from Kovacevo, southwest Bulgaria: woodland vegetation and its use during the earliest stages of the European Neolithic

Wood charcoal analysis from Kovacevo in southwest Bulgaria, one of the earliest Neolithic sites in southeastern Europe, provided information about the first stages of anthropogenic impact on vegetation during the Early Neolithic (6159–5630 cal b.c.). Deciduous oak was the most abundant and frequently used taxon in the wood charcoal assemblages. Cornus charcoal was also abundant, probably connected with the use of its twigs as building material in wattle and daub structures. The dominant deciduous oak forest was opened during the Kovacevo I period, as shown by evidence from the Kovacevo Ia and Kovacevo Ib occupation phases. Other types of vegetation, like Black pine (Pinus nigra) woodland, riverine forests and some sub-Mediterranean elements, were used only sporadically, indicating high and sustained availability of wood resources in the oak forests. Anthropogenic impacts were gradual, a pattern that matches contemporary studies elsewhere in the region.     

Morphological studies of free-threshing wheat ears from a Neolithic site in southwest Germany, and the history of the naked wheats

Two hundred and seventy carbonized naked wheat ears from the Neolithic pile dwelling at Hornstaad-Hörnle IA/Lake Constance (Germany) were analysed. The characteristics of internodes and glumes show that only tetraploid naked wheat types were cultivated. Following the history of free-threshing wheats it seems conceivable that the tetraploid naked wheats spread from their point of origin in the Middle East (7th/8th millennium B.C.) via a Mediterranean route to southwest Europe, reaching the northern foothills of the Alps about 4000 B.C.     

Dispersal and the transition to sympatry in vertebrates

Under allopatric speciation models, a key step in the build-up of species richness is population dispersal leading to the co-occurrence of previously geographically isolated forms. Despite its central importance for community assembly, the extent to which the transition from spatial segregation (allopatry or parapatry) to coexistence (sympatry) is a predictable process, or alternatively one governed by chance and the vagaries of biogeographic history, remains poorly understood. Here, we use estimated divergence times and current patterns of geographical range overlap among sister species to explore the evolution of sympatry in vertebrates. We show that rates of transition to sympatry vary predictably according to ecology, being faster in marine or strongly dispersive terrestrial clades. This association with organism vagility is robust to the relative frequency of geographical speciation modes and consistent across taxonomic scales and metrics of dispersal ability. These findings reject neutral models of dispersal assembly based simply on evolutionary age and are not predicted by the main alternative view that range overlap is primarily constrained by biotic interactions. We conclude that species differences in dispersal limitation are fundamental in organizing the assembly of ecological communities and shaping broad-scale patterns of biodiversity over space and time.     

Ancient mitochondrial DNA from the northern fringe of the Neolithic farming expansion in Europe sheds light on the dispersion process

The European Neolithization process started around 12 000 years ago in the Near East. The introduction of agriculture spread north and west throughout Europe and a key question has been if this was brought about by migrating individuals, by an exchange of ideas or a by a mixture of these. The earliest farming evidence in Scandinavia is found within the Funnel Beaker Culture complex (Trichterbecherkultur, TRB) which represents the northernmost extension of Neolithic farmers in Europe. The TRB coexisted for almost a millennium with hunter–gatherers of the Pitted Ware Cultural complex (PWC). If migration was a substantial part of the Neolithization, even the northerly TRB community would display a closer genetic affinity to other farmer populations than to hunter–gatherer populations. We deep-sequenced the mitochondrial hypervariable region 1 from seven farmers (six TRB and one Battle Axe complex, BAC) and 13 hunter–gatherers (PWC) and authenticated the sequences using postmortem DNA damage patterns. A comparison with 124 previously published sequences from prehistoric Europe shows that the TRB individuals share a close affinity to Central European farmer populations, and that they are distinct from hunter–gatherer groups, including the geographically close and partially contemporary PWC that show a close affinity to the European Mesolithic hunter–gatherers.     

Pollen and phytolith evidence for rice cultivation during the Neolithic at Longqiuzhuang, eastern Jianghuai, China

Phytolith and pollen analyses were carried out at the archaeological site at Longqiuzhuang in Gaoyou, Jiangsu, southern China. The results indicate that the key morphological phytolith types associated with cultivated rice (Oryza) are common in the Neolithic cultural layers at this site. The evidence strongly suggests that cultivated rice (mainlyO. japonica) was grown locally during the Neolithic. The archaeopalynological record provides information about the impact of human activity and, in particular, farming on the natural vegetation. The evergreen and deciduous broad-leaved forest was substantially altered, and herbaceous taxa, including ruderals, expanded. Based on the results from the phytolith and pollen analyses, two distinct phases of human activity have been recognized, namely (1) phase A (7000-6300 B.P., i.e. early Neolithic) a warm and humid period when arable farming, including rice cultivation, was pursued but the variation in the size of the carbonized rice grains was low, and (2) phase B (6300-5500 B.P., late Neolithic age), a period of relatively cold and/or arid climate when cultivated rice was of major importance and was morphologically similar to present-day rice. Environment, and in particular climate change in the late Neolithic, were important factors affecting the development of rice as a cultivated crop. It was mainly during this period that artificial selection favoured the emergence of forms similar to those of today.     

Asymmetric larval mobility and the evolutionary transition from siblicide to nonsiblicidal behavior in parasitoid wasps

The widespread evolution of gregarious development in parasitoidwasps presents a theoretical challenge because the conditionsunder which larval tolerance can spread in an intolerant populationare very stringent (the individual fitness of larvae developingtogether must increase with clutch size). Recent empirical workhas suggested that gregarious development can arise throughthe loss of larval mobility rather than through the gain oftolerant behavior. Using analytical genetic models, we exploredwhether decreased mobility presents a less stringent route togregariousness than the gain of tolerance. Reduced mobilitycan spread under a wide range of conditions. The critical conditionfor the spread of immobility is much less stringent than thatfor larval tolerance. In contrast with previous models of tolerance,the criterion for the spread of a rare immobility allele isindependent of any bias in the sex ratio and the likelihoodof single sex broods. Superparasitism increases the stringencyof the criterion for the spread of immobility, whereas doublekilling relaxes the criterion. Tolerance can subsequently replaceimmobility if there is any cost to the retention of fightingability. Our results suggest that asymmetric larval mobilitymay explain many instances of the evolution of gregarious development.     

Cultivated plant remains of the late Neolithic Michelsberg Culture at Heilbronn-Klingenberg (southwest Germany) - a comparison of different features, find assemblages and preservation conditions relating to the representation of archaeobotanical remains

A large area investigation was undertaken of the plant remains from a Michelsberg Culture (late Neolithic) settlement. The charred macroscopic remains and imprints in pieces of daub were expected to show both the spectrum of the cultivated plants there, and also the degree of their cultivation and use. The loess covered hill-top lies in a landscape with favourable climatic and soil conditions and is blocked off by two parallel ditches running in an arc. Ditches and pits filled with different sediments were investigated. The daub, unearthed in several pits, had been deliberately mixed with chaff of the glume wheats einkorn and emmer to temper it when it was originally made. The investigated imprints and charred plant remains give hints of spatial distribution of crop processing activities. To determine the amounts of the crops that were cultivated and used, it is necessary to study the charred remains. The degree of ubiquity (frequency of occurrence) of grains in the pit sediments seems to be the best indicator of the representation of cereals. Four main cereals were found: Triticum monococcum, T. dicoccum, T. aestivum/T. durum and Hordeum vulgare var. nudum. Pisum sativum also was an important cultivated plant, much more than Lens culinaris. The role of Linum usitatissimum and Papaver somniferum is less clear.     

C4 photosynthetic modifications in the evolutionary transition from land to water in aquatic grasses

Cladistic analysis supports the conclusion that the Orcuttieae tribe of C₄ grasses reflect evolution from a terrestrial ancestry into seasonal pools. All nine species in the tribe exhibit adaptations to the aquatic environment, evident in the structural characteristics of the juvenile foliage, which persist submerged for 1–3 months prior to metamorphosis to the terrestrial foliage. Aquatic leaves of the least derived or basal genus Neostapfia have few morphological and anatomical characteristics specialized to the aquatic environment and have retained full expression of the C₄ pathway, including Kranz anatomy. Orcuttia species have many derived characteristics and are more specialized to the aquatic environment. These latter species germinate earlier in the season and persist in the submerged stage longer than Neostapfia and evidence from the literature indicates length of submergence is positively correlated with fitness components. Aquatic leaves of Orcuttia species lack Kranz or PCR bundle sheath anatomy, yet ¹⁴C-pulse chase studies indicate >95% malate + aspartate as the initial products of photosynthesis and these products turn over rapidly to phosphorylated sugars, indicating a tight coupling of the C₄ and C₃ cycles. Presence of the C₄ pathway is further supported by enzymological data. Contemporary dogma that Kranz anatomy is a sine qua non for operation of the C₄ pathway is contradicted by the patterns in Orcuttia; however, it is unknown whether the pathway acts as a CO₂ concentrating mechanism in these aquatic plants. Received: 12 June 1997 / Accepted: 10 February 1997     

The Early Neolithic site at Hoge Vaart, Almere, the Netherlands, with particular reference to non-diffusion of crop plants, and the significance of site function and sample location

The construction of a new motorway necessitated the excavation of an Early Neolithic site in the Dutch Flevopolder near the Hoge Vaart canal. In 1994-1996, an area of ca. 1700 m² was excavated in 0.5x0.5-m squares. The excavation revealed 150 surface hearths and more than a hundred deep hearth pits. The finds, mainly flint and pottery, as well as¹⁴C dates from charcoal demonstrated that the area had been used mainly during the Early Neolithic (ca. 4900–4500 cal. B.C.). The investigation of the Early Neolithic remains included detailed botanical analyses of 182 samples and hand-sorting by the excavation team, which included archaeobotanists, of tens of thousands of samples sieved over 2-mm-mesh sieves. No remains of cultivated plants were recognised among the 87 taxa (40 identified to species level) that were preserved, mainly in carbonised form. These results are compared with other Early Neolithic sites studied in the Netherlands, such as Swifterbant S3 and Schokland P14. The possible significance of site function and sample location as important factors influencing presence/absence of cultivated plant remains at a particular site is discussed.     

Linking recruitment to trophic factors: revisiting the Beverton--Holt recruitment model from a life history and multispecies perspective

The Beverton--Holt recruitment model can be derived from arguments about evolution of life history traits related to foraging and predation risk, along with spatially localized and temporarily competitive relationships in the habitats where juvenile fish forage and face predation risk while foraging. This derivation explicitly represents two key biotic factors, food supply (I) and predator abundance (R), which appear as a risk ratio (R/I) that facilitates modelling of changes in trophic circumstances and analysis of historical data. The same general recruitment relationship is expected whether the juvenile life history is simple or involves a complex sequence of stanzas; in the complex case, the Beverton--Holt parameters represent weighted averages or integrals of risk ratios over the stanzas. The relationship should also apply in settings where there is complex, mesoscale variation in habitat and predation risk, provided that animals sense this variation and move about so as to achieve similar survival at all mesoscale rearing sites. The model predicts that changes in food and predation risk can be amplified violently in settings where juvenile survival rate is low, producing large changes in recruitment rates over time.     

Sex,long life and the evolutionary transition to cooperative breeding in birds

Long life is a typical feature of individuals living in cooperative societies. One explanation is that group living lowers mortality, which selects for longer life. Alternatively, long life may make the evolution of cooperation more likely by ensuring a long breeding tenure, making helping behaviour and queuing for breeding positions worthwhile. The benefit of queuing will, however, depend on whether individuals gain indirect fitness benefits while helping, which is determined by female promiscuity. Where promiscuity is high and therefore the indirect fitness benefits of helping are low, cooperation can still be favoured by an even longer life span. We present the results of comparative analyses designed to test the likelihood of a causal relationship between longevity and cooperative breeding by reconstructing ancestral states of cooperative breeding across birds, and by examining the effect of female promiscuity on the relationship between these two traits. We found that long life makes the evolution of cooperation more likely and that promiscuous cooperative species are exceptionally long lived. These results make sense of promiscuity in cooperative breeders and clarify the importance of life-history traits in the evolution of cooperative breeding, illustrating that cooperation can evolve via the combination of indirect and direct fitness benefits.     

Fish remains from the Neolithic site of El Mirador cave (Atapuerca,Spain): Seasonality and resource management

Ichthyoarchaeological analyses of the freshwater fish remains from levels 19 and 20 in El Mirador cave (Atapuerca, Spain) have been conducted. Fish were always present as a source of animal protein, although their importance in the human diet was not fully exploited by people during the Neolithic on the Iberian Peninsula. Two principal goals are treated here: a taxonomic study of the fish remains and a characterization of the exploitation of this resource. The results show that the human community of El Mirador cave practiced fishing, and that fish was part of their diet and social life.     

Fossil biogeography: a new model to infer dispersal,extinction and sampling from palaeontological data

Methods in historical biogeography have revolutionized our ability to infer the evolution of ancestral geographical ranges from phylogenies of extant taxa, the rates of dispersals, and biotic connectivity among areas. However, extant taxa are likely to provide limited and potentially biased information about past biogeographic processes, due to extinction, asymmetrical dispersals and variable connectivity among areas. Fossil data hold considerable information about past distribution of lineages, but suffer from largely incomplete sampling. Here we present a new dispersal–extinction–sampling (DES) model, which estimates biogeographic parameters using fossil occurrences instead of phylogenetic trees. The model estimates dispersal and extinction rates while explicitly accounting for the incompleteness of the fossil record. Rates can vary between areas and through time, thus providing the opportunity to assess complex scenarios of biogeographic evolution. We implement the DES model in a Bayesian framework and demonstrate through simulations that it can accurately infer all the relevant parameters. We demonstrate the use of our model by analysing the Cenozoic fossil record of land plants and inferring dispersal and extinction rates across Eurasia and North America. Our results show that biogeographic range evolution is not a time-homogeneous process, as assumed in most phylogenetic analyses, but varies through time and between areas. In our empirical assessment, this is shown by the striking predominance of plant dispersals from Eurasia into North America during the Eocene climatic cooling, followed by a shift in the opposite direction, and finally, a balance in biotic interchange since the middle Miocene. We conclude by discussing the potential of fossil-based analyses to test biogeographic hypotheses and improve phylogenetic methods in historical biogeography.     

A model explaining the size distribution of gene and protein families

This article deals with the theoretical size distribution of gene and protein families in complete genomes. A simple evolutionary model for the development of such families in which genes in a family are formed or selected against independently and at random, and in which new families are formed by the random splitting of existing families, is used to derive the resulting size distribution. Mathematically this turns out to be the distribution of the state of a homogeneous birth-and-death process after an exponentially distributed time, which it is shown will under certain conditions exhibit the power-law behaviour observed for gene and protein family sizes.