Is naked barley an eastern or a western crop? The combined evidence of archaeobotany and genetics

Forms of Hordeum vulgare ssp. vulgare (barley) that possess a naked caryopsis are an important human staple and are mainly found today in eastern Asia. However, naked barley has not always been an eastern crop: archaeobotanical data show that it was prevalent in Europe and the Near East during various periods in prehistory. In this review we have collated data on the incidence of hulled and naked barley at archaeological sites in Europe and the Near East from two sources: archaeobotanical literature reviews and an archaeobotanical database, both assembled by Helmut Kroll. We have also examined the incidence of hulled and naked barleys in extant germplasm collections. Our compilation of this archaeobotanical data has enabled us to elucidate long-term changes in the ratio of hulled to naked barley under cultivation in these regions; specifically, these records show that naked barley begins to disappear from the archaeobotanical record from the Chalcolithic/Bronze Age onwards in the Near East, and from the Iron Age/Roman periods onwards in Europe. We discuss the possible causes of this decline in naked barley cultivation in these regions, along with the present-day prevalence of naked barley landraces in eastern Asia, particularly in relation to genetic evidence, which shows that naked barley has a single origin.     

Seed bank diversity in mesic grasslands in relation to vegetation type,management and site conditions

Questions: 1. Do different management types (i.e. hay meadow, silage meadow, meadow‐pasture, pasture) have different impact on the size and composition of the seed bank of mesic grassland (Arrhenatheretalia)? 2. How strong is the effect of management on the seed bank in relation to above‐ground vegetation, edaphic factors and land‐use history? 3. Are there differences in C‐S‐R plant strategy types and seed longevity under different management regimes? Location: Lahn‐Dill Highlands in central‐western Germany. Methods: Above‐ground vegetation and the soil seed bank of 63 plots (at 21 sites) in mesic grasslands were studied. Differences between management types in quantitative seed bank traits and functional characteristics were tested by ANOVA. The impact of management, above‐ground vegetation, site conditions and land‐use history on seed bank composition were analysed by partial CCA. Results: Management had no significant impact on species richness and density of the seed bank but significantly influenced their floristic composition and functional characteristics. CCA revealed that even after adjustment for soil chemical parameters and above‐ground vegetation management still had significant impact on seed bank composition. ANOVA revealed that silage meadows contained higher proportions of R‐strategy compared to hay meadows. In contrast, in hay meadows and meadow‐pastures proportions of S‐strategy were higher than in silage meadows. Conclusions: The type of grassland management has little impact on quantitative seed bank traits. Management types with a high degree of disturbance lead to an increase of species following a ruderal strategy in the seed bank. Irrespective of management type only a limited proportion of characteristic grassland species is likely to re‐establish from the seed bank after disappearance from above‐ground vegetation.     

Mitochondrial DNA and Y-chromosomal diversity in ancient populations of domestic sheep (Ovis aries) in Finland: comparison with contemporary sheep breeds

Background

Several molecular and population genetic studies have focused on the native sheep breeds of Finland. In this work, we investigated their ancestral sheep populations from Iron Age, Medieval and Post-Medieval periods by sequencing a partial mitochondrial DNA D-loop and the 5’-promoter region of the SRY gene. We compared the maternal (mitochondrial DNA haplotypes) and paternal (SNP oY1) genetic diversity of ancient sheep in Finland with modern domestic sheep populations in Europe and Asia to study temporal changes in genetic variation and affinities between ancient and modern populations.

Results

A 523-bp mitochondrial DNA sequence was successfully amplified for 26 of 36 sheep ancient samples i.e. five, seven and 14 samples representative of Iron Age, Medieval and Post-Medieval sheep, respectively. Genetic diversity was analyzed within the cohorts. This ancient dataset was compared with present-day data consisting of 94 animals from 10 contemporary European breeds and with GenBank DNA sequence data to carry out a haplotype sharing analysis. Among the 18 ancient mitochondrial DNA haplotypes identified, 14 were present in the modern breeds. Ancient haplotypes were assigned to the highly divergent ovine haplogroups A and B, haplogroup B being the major lineage within the cohorts. Only two haplotypes were detected in the Iron Age samples, while the genetic diversity of the Medieval and Post-Medieval cohorts was higher. For three of the ancient DNA samples, Y-chromosome SRY gene sequences were amplified indicating that they originated from rams. The SRY gene of these three ancient ram samples contained SNP G-oY1, which is frequent in modern north-European sheep breeds.

Conclusions

Our study did not reveal any sign of major population replacement of native sheep in Finland since the Iron Age. Variations in the availability of archaeological remains may explain differences in genetic diversity estimates and patterns within the cohorts rather than demographic events that occurred in the past. Our ancient DNA results fit well with the genetic context of domestic sheep as determined by analyses of modern north-European sheep breeds.     

Agricultural practices in Arctic Norway during the first millennium b.c.

Palaeoecological investigations of mires suggest that agriculture was established north of the Arctic Circle in Norway during the late Bronze Age (1100?C500 b.c.) and Pre-Roman Iron Age (500?C1 b.c.). The lack of archaeobotanical and archaeological investigations has made it difficult to assess the nature of this early agricultural expansion into the Arctic in any detail. Here we present the first well documented archaeobotanical investigation from north Norway that covers this agricultural pioneer phase. Remains of charred seeds show that barley (Hordeum) was already being cultivated in the late Bronze Age, and that wheat (Triticum) was introduced in the Pre-Roman Iron Age. Large amounts of crowberry (Empetrum) seeds are also typical of the Pre-Roman Iron Age and were obviously an important food plant at the time, at least locally. Charcoal rich layers dated to the late Bronze Age suggest that the local birch forest was initially cleared away with the help of fire, possibly related to a slash-and-burn cultivation practice. Lithostratigraphic and pollen-analytical results indicate that the cultivation practice of the Pre-Roman Iron Age was a form of bush-fallow system with intensive soil re-working alternated with long periods of fallow.     

Pollen analysis of Iron Age cow dung in southern Africa

Thick accumulations of consolidated cow dung occur in ancientkraals (byres or corrals) in the bushveld and highveld areas of Zimbabwe, Botswana, and South Africa dating from the last 2000 years. They originated from long-term cattle herding by Iron Age people. The vitrified or baked dung deposits are thought to be a product of the burning of cow dung as fuel, either for domestic purposes or for iron smelting. In order to establish the palaeoecological potential of this material, 36 samples of cow dung from archaeological sites within the present-day savanna and grassland biomes were analyzed for pollen and other microfossils. Of the samples, 29 contained pollen together with other microfossils that support a faecal origin of the material such as sordariaceous ascospores,Thecaphora, Gelasinospora, andChaetomium, and eggs of the intestinal parasiteTrichuris. Similar microfossils were also found in recent fresh cow dung from the same study areas. The presence of pollen grains and spores in most of the Iron Age samples lead to the assumption that they survived the burning because fire temperatures were not high enough to destroy them. Pollen in these cow dung pieces is apparently sealed and can be preserved under open-air conditions at sites under which pollen in other deposits like soils, will decay away. Good pollen preservation and palynomorph diversity were found with mainly Poaceae, and secondly Chenopodiaceae and Cyperaceae as the most important pollen types, while trees and shrubs indicating savanna are rare. In the case of the samples that came from the subtropical savanna biome the latter result is unexpected and suggests that the cattle were kept in more open vegetation than the woody environments of today. Recent cow dung samples reflect the composition of present-day vegetation by showing considerably higher proportions of tree pollen than the fossil assemblages.     

Holocene vegetation and land-use history in the environs of Lake Kahala, northern Estonia

Pollen profiles, based cores taken in Lake Kahala and from the adjoining mire, were used to establish general vegetation history and to reconstruct the extent and types of land-use over most of the Holocene. Modern pollen deposition was studied using moss polsters and the results were used in the interpretation of the fossil pollen data in terms of former land-use practices. The modern-day samples are from settlements, hay meadow and pasture, and overgrown pasture. Indications of human activity can be traced back to the Stone Age. At ca. 6400 cal. B.C., the first indications of possible woodland utilisation by humans are recorded. This may have involved grazing within the forests. From 4200 cal. B.C. onwards, animal husbandry with changing intensity was practised. Arable farming, involving cereals, was introduced to the area at ca. 1800 cal. B.C., but it was only at ca. 500 cal. B.C. that it assumed an important role in the farming economy.Secale cereale (rye) was introduced during the Roman Iron Age, intensive rye cultivation started at the end of the Iron Age, at ca. cal. A.D. 800. Ever increasing farming pressures triggered the formation of openalvars. Open landscape similar to that of today has persisted, with minor forest regeneration phases, since at least 500 cal. B.C.     

The functional ecology of present-day arable weed floras and its applicability for the identification of past crop husbandry

Five studies of present-day weed floras are brought together in this paper to address the issue of ancient crop husbandry and how different agricultural practices may be identified from the archaeobotanical record. These studies have provided valuable insights into the complexity of the relationship between the functional attributes of species and their ecological significance in relation to habitat conditions and husbandry regimes. Using a suite of characteristics, practices such as irrigation, fallowing, crop rotation, time of sowing and intensity of cultivation (hoeing, weeding, manuring etc.) can be recognised on the basis of the functional ecology of the weed species accompanying the crops. The geographic variation in functional attributes is limited and permits the identification of husbandry practices in different climatic zones. Recommendations are made for the application of weed ecological methods to archaeobotanical assemblages.     

Condiments before Claudius: new plant foods at the Late Iron Age oppidum at Silchester,UK

Our understanding of the introduction and adoption of new plant foods in Roman Britain is currently limited by a lack of data from a group of Late Iron Age settlements commonly referred to as oppida (large pre-Roman towns). This paper presents the first evidence of several imported plant foods from Late Iron Age Britain in the form of waterlogged plant remains from the oppidum at Silchester. These were recovered from the basal contexts of two wells, dated to the early first century a.d. One olive stone and several seeds of celery, coriander and dill were identified. The results are compared to archaeobotanical data from elsewhere in Britain and northwestern Europe, demonstrating that Silchester is part of the wider phenomenon of the adoption of new flavourings and fruits in Late Iron Age Europe.     

Approaches to reconstruction of early Celtic land-use in the cent ral Neckar region in southwestern Germany

In four early Celtic settlements in the central Neckar region of Baden-Württemberg (Germany), 250 archaeobotanical samples were collected and studied for their plant macrofossils.Hordeum vulgare L. (hulled barley) andTrilicum spella L. (spelt wheat) could be recognized as the main crops, as well asPanicum miliaceum L. (millet) in some places. Legumes andLinum usitatissimum L. (flax) were important crops. Typical field weeds dominated the wild plant remains, but some present-day grassland taxa were abundantly and frequently found. The analysis of the data set of finds of the cereal remains, field weeds and grassland plants was done by Correspondence Analysis, but no significant differences between the various groups were found. Samples dominated by remains of grassland taxa, which could indicate that there were hay meadows, were absent. Consideration of plant macrofossil results as well as archaeological, osteological and pollen analytical data shows that there is likely to have been a land management system of arable fields and grassland. The densely settled fertile central Neckar region could therefore have been intensively utilised, using all available resources. Permanent grasslands in the form of pasture on river floodplains and on dry soils were certainly present. Hay meadows, on the other hand, were not identified. Cultivated plants such asFicus carica L. (fig) andVicia ervilia (L.) Willd. (bitter vetch) as well as some field weeds suggest connections with the Mediterranean lands.     

Ancient DNA Reveals Matrilineal Continuity in Present-Day Poland over the Last Two Millennia

While numerous ancient human DNA datasets from across Europe have been published till date, modern-day Poland in particular, remains uninvestigated. Besides application in the reconstruction of continent-wide human history, data from this region would also contribute towards our understanding of the history of the Slavs, whose origin is hypothesized to be in East or Central Europe. Here, we present the first population-scale ancient human DNA study from the region of modern-day Poland by establishing mitochondrial DNA profiles for 23 samples dated to 200 BC – 500 AD (Roman Iron Age) and for 20 samples dated to 1000–1400 AD (Medieval Age). Our results show that mitochondrial DNA sequences from both periods belong to haplogroups that are characteristic of contemporary West Eurasia. Haplotype sharing analysis indicates that majority of the ancient haplotypes are widespread in some modern Europeans, including Poles. Notably, the Roman Iron Age samples share more rare haplotypes with Central and Northeast Europeans, whereas the Medieval Age samples share more rare haplotypes with East-Central and South-East Europeans, primarily Slavic populations. Our data demonstrates genetic continuity of certain matrilineages (H5a1 and N1a1a2) in the area of present-day Poland from at least the Roman Iron Age until present. As such, the maternal gene pool of present-day Poles, Czechs and Slovaks, categorized as Western Slavs, is likely to have descended from inhabitants of East-Central Europe during the Roman Iron Age.     

Nitrogen dynamics in alpine ecosystems of the northern Caucasus

Net N mineralization, nitrification, microbial biomass N and ¹⁵N natural abundance were studied in a toposequence of representative soils and plant communities in the alpine zone of the northern Caucasus. The toposequence was represented by (1) low-productive alpine lichen heath (ALH) of wind-exposed ridge and upper slope; (2) more productive Festuca varia grassland (FG) of middle slope; (3) most productive Geranium gymnocaulon/Hedusarum caucasicummeadow (GHM) of lower slope; (4) low-productive snowbed community (SBC) of the slope bottom. N availability, net N mineralization and nitrification were higher in soils of alpine grassland and meadow of the middle part of the toposequence compared with soils of lichen heath and snowbed community of extreme habitats in the alpine zone. There was no correlation between intensities of N transformation processes and favorable (low soil acidity, low C/N ratio, long vegetation period, relatively high temperature, absence of hydromorphic features) and unfavorable (opposite) factors, indicating that the intensity of N mineralization and nitrification in the alpine soils is controlled by a complex combination of these factors. Potential net N mineralization and nitrification in alpine soils determined in the short-term laboratory incubation were considerably higher than those determined in the long-term field incubation. The differences of potential nitrification between soils of various plant communities did not correspond to the field determined pattern indicating the importance of on-site climatic conditions for control of nitrification in high mountains. The result of comparison of N transformation potentials in incubated and native soils indicated that nitrification potential was significantly increased after long-term soil incubation. It means that net nitrification determined in the field was probably overestimated, especially in the meadow soils. A soil translocation experiment indicated that low temperature was an important factor limiting net N mineralization and nitrification in alpine soils: net N mineralization and especially nitrification increased when alpine soils were translocated into the subalpine zone and mean annual temperature increased by about 3°C. Additional N input increased N availability (NH₄ ⁺-N) and potential nitrification in soils of the lower part of the toposequense (GHM and SBC), and potential net N mineralization in two soils of extreme habitats (ALH and SBC). A positive correlation was found between soil ¹⁵N and net N mineralization and nitrification; the relative ¹⁵N enrichment was characteristic of grassland and meadow ecosystems. ¹⁵N of total soil N pool increased during the field mineralization experiment; there was a positive tendency between the change in ¹⁵N and net N mineralization and nitrification, however the relationship was not significant. Foliar ¹⁵N of dominant plant species varied widely within community, however, a tendency of higher foliar ¹⁵N for species growing on the soils with higher net N mineralization, nitrification and ¹⁵N was observed.     

Spontaneous regeneration of dry grasslands on set-aside fields

Dry grasslands are among the most threatened plant communities of Central Europe. We explore the time scale of spontaneous regeneration of dry grasslands on abandoned fields in an area of Central Europe, where also ancient grassland communities occur (Saxony-Anhalt, Germany). On three permanent plots with shallow soils we monitored during 10 years change of species composition and analysed whether spontaneous succession leads to assemblages similar to the ancient dry grassland communities in the direct surroundings. We found that dry grassland species are able to invade the permanent plots and during the 10 years of succession the number of dry grassland species increased. But even after 10 years there was a clear difference between ancient dry grassland communities and the assemblages on the permanent plots. Our findings suggest two important conclusions: First, spontaneous succession on abandoned fields is a cheap possibility for the conservation of some dry grassland species, at least on shallow soils. Second, the time scale of the regeneration process, however, is rather long. Hence, conservation of remnants of ancient grassland communities needs special attention.     

Growth,photosynthesis and storage of carbohydrates and nitrogen in Phaseolus lunatus in relation to resource availability

External hyphae of vesicular-arbuscular mycorrhizal (VAM) fungi were quantified over a growing season in a reconstructed tallgrass prairie and an ungrazed cool-season pasture. In both sites, hyphal lengths increased throughout the growing season. Peak external hyphal lengths were 111 m cm^–3 of soil in the prairie and 81 m cm^–3 of soil in the pasture. These hyphal lengths calculate to external hyphal dry weights of 457 g cm^–3 and 339 g cm^–3 of soil for prairie and pasture communities, respectively. The relationships among external hyphal length, root characteristics, soil P and soil moisture were also determined. Measures of gross root morphology [e.g., specific root length (SRL) and root mass] have a strong association with external hyphal length. Over the course of the study, both grassland communities experienced a major drought event in late spring. During this period a reduction in SRL occurred in both the pasture and prairie without a measured reduction in external hyphal length. Recovery for both the pasture and prairie occurred not by increasing SRL, but rather by increasing external hyphal length. This study suggests that growth is coordinated between VAM hyphae and root morphology, which in turn, are constrained by plant community composition and soil nutrient and moisture conditions.     

内蒙古典型草原区3种不同草地利用模式下植物功能群及其多样性的变化

以呼伦贝尔草原辉河国家级自然保护区草原群落为研究对象, 选取3块生境条件基本相同的样地(250 m × 250 m) ——放牧场、割草场及公共草场, 每块样地设置3条样带, 每条样带上设置8个样方(1 m × 1 m), 研究了不同草地利用模式下草原群落的物种组成、生物多样性及群落的功能群特征。结果表明: 在放牧场、割草场和公共草场3种不同利用模式下, 植物功能群结构及其多样性发生了变化。灌木、半灌木的累积优势度在各利用模式间差异显著( p< 0.05), 在割草场中最高; 多年生禾草的累积优势度在各利用模式间差异显著(p< 0.05); 一、二年生草本的累积优势度在各利用模式间差异显著(p< 0.05)。旱生植物功能群在公共草场中所占比例最高, 为50%; 中旱生植物在割草场中的百分比最低, 为19%, 且在各利用模式间差异显著(p< 0.05); 各利用模式下, 灌木、半灌木功能群生物量均较低; 随着割草场、公共草场及放牧场3种草地利用模式下利用强度的增加, 多年生禾草和多年生杂类草两大功能群的生物量递减, 而一、二年生草本功能群生物量递增。草地利用强度的增加, 使得草地植物群落结构简单、脆弱。丰富度指数和多样性指数在放牧场最低, 在割草场最高; 均匀度变化并不明显。割草场利用模式有助于改善草原的健康状况, 而放牧场及公共草场利用模式均存在不同程度的过度放牧现象, 尤其是公共草场利用模式由于所有权不明确而缺乏有效的管理, 导致草原退化明显。     

Grassland Butterfly Fauna under Traditional Animal Husbandry: Contrasts in Diversity in Mown Meadows and Grazed Pastures

Semi-natural grasslands resulting from traditional land use practices (mowing and grazing) are severely endangered throughout Europe due to the intensification of agriculture. The ecological impact of mowing and grazing on grassland butterflies was studied in eight mown meadows and eight grazed pastures under traditional animal husbandry in NW Russia and adjacent Finland. Transect count data over 3 years (1997–1999) covered a total of 48 species and 5742 individuals. The butterfly fauna was rather similar under both management forms; species richness, diversity and total abundance did not differ significantly between meadows and pastures, yet meadows were preferred by more species. In both groups, the most abundant species were Aphantopus hyperantus, Pieris napi and Thymelicus lineola. Of 37 species observed as a minimum of five individuals, Polyommatus amandus, Ochlodes sylvanus and A. hyperantus showed a significant preference for mowing management. According to the ordination, butterfly communities were affected more by the origin and age of the grassland than the present management method. Landscape factors (meadow or pasture surrounded by forests or open environments), the abundance of nectar plants and the intensity of tilling were the most important factors differentiating older grasslands from the younger ones evolved from old Finnish hay fields cultivated prior to the 1940s.     

干旱区不同草地土壤植物系统水盐的特征

 土壤水盐状况对干旱区草地植物群落的种类组成有直接的影响。本文通过3年对土壤水盐动态(1m土壤含盐量、潜水埋深和矿化度)的定位定点监测,系统地划分出水盐等级指标,并结合草地调查,阐述了不同草地类型与水盐状况的关系,将新疆呼图壁草地生态试验站天然草地划分为沼泽化低地草甸、盐化低地草甸、重盐化低地草甸、盐土荒漠和重盐土荒漠五个亚类。     

京津风沙源治理工程对草地土壤有机碳库的影响——以内蒙古锡林郭勒盟为例

研究草地生态建设工程对土壤有机碳库的影响,对于草地生态建设工程成效评估及草地碳循环研究具有重要意义.本文以内蒙古锡林郭勒盟京津风沙源治理工程区为例,采用IPCC推荐的碳收支清单法,分析了2000-2006年京津风沙源治理工程对草地土壤有机碳库的影响,并对3种管理措施下（人工种草、飞播牧草和围栏封育）草地土壤达到最大有机碳密度的时间进行估算.结果表明： 2000-2006年,京津风沙源治理工程对草地土壤碳汇具有极大的促进作用,工程区整体表现为碳汇,碳汇量为59.26×10⁴ t C;不同草地管理方式下草地土壤固碳速率和效益差异显著,其中,人工种草的土壤固碳速率较快,为0.25 t C·hm^-2·a^-1,而围栏封育导致土壤固碳效益较高（0.63亿元）;与其他草地类型相比,低地草甸和温性草甸草原的土壤固碳速率较快,均为0.14 t C·hm^-2·a^-1.通过管理措施使草地达到潜在最大土壤有机碳密度是一个长期的过程,相对而言,人工种草所需的时间较短（57.75年）.     

Weed finds as indicators for the cultivation regime of the early Neolithic Bandkeramik culture?

Ethnographic data combined with the characteristics of the weed species from Bandkeramik settlement sites give hints for the reconstruction of Early Neolithic agricultural practises in Central Europe. In contrast to the Balkan situation with a high diversity in cultivated crops, Bandkeramik field management can be reconstructed as a simple agricultural system with emphasis on summer crop growing. Permanent fields were treated with hoes, digging sticks or similar tools, sown in spring and grazed in autumn and winter. The intensity of field management seems to increase through time as shown by diachrone comparison of archaeobotanical data from Neolithic, Iron Age and Roman times. The absence of winter-cereals such as naked wheat, grown in the Balkan Peninsula, gives a hint of a certain emphasis on stock breeding. Summer crop growing would have had the advantage that the Bandkeramik fields could be grazed after harvest until next spring and would therefore be manured at the same time.     

Grassland management for meadow birds in the Netherlands is unfavourable to pollinators

Agricultural intensification and loss of semi-natural grassland have contributed to biodiversity decline, including pollinator species, in pastures around the world. To reverse the decline, agri-environmental schemes have been implemented, varying widely in effectiveness. In addition, many countries, including the Netherlands, have established nature reserves in which semi-natural grasslands are restored and are often managed for specific groups of species, e.g. meadow birds or plants. The effects of such measures on insect biodiversity are not well known but recent reports on the dramatic decline of insect biomass in nature reserves have put even more attention to the impact of land use and management on biodiversity. This study compares pollinator abundance and species richness in three common semi-natural grassland management types in the Netherlands: (1) hay meadows, (2) herb-rich grasslands and (3) meadow bird grasslands. Pollinator abundance and species richness were assessed in eleven study areas, each with all three management types present. Standardized transects, insect sampling within a standard 20 min time frame and plot-based flower surveys were used in spring and summer to assess the relationships between management regime, floral abundance and diversity and pollinator communities. The results show that meadow bird grasslands have lower pollinator abundance and diversity and a less unique pollinator assemblage than both other types. Moreover, flower abundance has a positive effect on pollinator abundance and flower diversity has a positive effect on pollinator species richness. These results indicate that meadow-bird grasslands are a comparatively unfavourable habitat for bees, hoverflies and butterflies, which may be explained by a lack of flowers as well as unsuitable mowing practices. Measures benefitting both insectivorous birds and flower-visiting insects, such as rotational mowing, could remediate this imbalance.     

The agricultural consequences of colonial contacts on the Iberian Peninsula in the first millennium <Emphasis Type="SmallCaps">b</Emphasis>.<Emphasis Type="SmallCaps">c</Emphasis>

The Iron Age archaeobotanical record on the Iberian Peninsula shows how the Phoenician and Greek colonisers caused the indigenous Iberians to change the management of the agricultural resources and the crops which they grew. These colonisers also brought about the development of viticulture and olive cultivation. The importance of agricultural products in the trade network which was stimulated by the colonisers may have encouraged new farming systems, as well as surplus capacity in the native agriculture in the region.