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.     

Towards the archaeobotanical identification of intensive cereal cultivation: present-day ecological investigation in the mountains of Asturias, northwest Spain

The region of Asturias, northwest Spain, is highly unusual in that a cereal crop (spelt wheat) is cultivated on a garden scale using horticultural methods. A floristic survey was made of the weeds in 65 spelt plots in this region. The ecological attributes of the weed species were then measured and compared to an earlier study of the functional characteristics of weeds associated with pulse crops on the Greek island of Evvia. In this earlier study, it was possible to distinguish between plots cultivated intensively on a garden-scale and plots cultivated extensively in fields, on the basis of a suite of functional attributes of the weed species. The cereal plots from Asturias were correctly identified as gardens on the basis of the same suite of attributes. The Asturias plots were also compared to weed associations from autumn- and spring-sown crops in Germany, using a different suite of attributes, and were classified either as autumn-sown or ambiguously. This is consistent with the sowing time in Asturias, which is spread over late autumn to winter. These results demonstrate that the suites of functional attributes identified to distinguish intensive and extensive cultivation, and to recognise sowing time, can be applied in another geographical area and to another crop type. This paves the way for the application of these attributes to the identification of past agricultural practices from archaeological weed assemblages. Received September 12, 2001 / Accepted December 12, 2002     

The impact of crop processing on the reconstruction of crop sowing time and cultivation intensity from archaeobotanical weed evidence

Reconstruction of crop sowing time and cultivation intensity, based on arable weed ecology, can resolve archaeological questions surrounding land use and cycles of routine activity, but crop processing may introduce systematic ecological biases in the arable weeds represented in products and by-products. Based on previous ethnoarchaeological work, there is a predicted bias against indicators of spring sowing and intensive cultivation in fine sieve products (and a corresponding over-representation of such species in by-products). Recent work on modern weed floras using functional weed ecology has identified distinctive functional attributes associated with different sowing regimes and cultivation intensity levels. Evaluation of the predicted biases using functional attribute data for modern weed survey studies of different sowing regimes (in Germany) and cultivation intensity levels (in Greece) suggests that there is a likely bias against spring sowing indicators in fine sieve products but not (apparently) against intensive cultivation indicators. An archaeological case study is presented in order to illustrate how bias relating to crop sowing time may be identified and interpreted.     

Questioning the relevance of shifting cultivation to Neolithic farming in the loess belt of Europe: evidence from the Hambach Forest experiment

Despite widespread criticism, the shifting cultivation model continues to inform discussion of Neolithic farming in Europe, beginning with early Neolithic (Linearbandkeramik or LBK) communities concentrated in the loess belt of western-central Europe. Hundreds of LBK and later Neolithic sites have been excavated in this region and many of them sampled for charred plant remains. Archaeobotanical data on the weed floras harvested with crops provide the most direct archaeological evidence of crop husbandry practices, including the permanence of crop fields, but have played a limited role in the debate over shifting cultivation. The Hambach Forest experiment, conducted in the 1970s-80s near Cologne, Germany, provides valuable comparative data on the weed floras growing in newly cleared cultivation plots in an area of longlived mixed oak woodland on loess-based soil. Correspondence analysis of the Hambach weed survey data suggests that weed floras of fields managed under a shifting cultivation regime would be rich in perennial species, including woodland perennials. Comparison of these results with Neolithic weed assemblages from the loess belt of western-central Europe strongly suggests that Neolithic crop fields were not recently cleared of woodland vegetation but were long-established. Received September 5, 2001 / Accepted February 26, 2002     

The distribution, natural habitats and availability of wild cereals in relation to their domestication in the Near East: multiple events, multiple centres

In this article we examine the natural habitats and distribution of the six wild cereals: Triticum urartu (wild urartu wheat), T. boeoticum aegilopoides (single-grained wild einkorn), T. boeoticum thaoudar (two-grained wild einkorn), T. dicoccoides (wild emmer wheat), Secale spp. (wild ryes) and Hordeum spontaneum (wild barley). A comparison of late Pleistocene/early Holocene archaeobotanical assemblages in the Near East with present-day distributions of wild cereals shows a good correlation. The regional variation in the archaeobotanical cereal assemblages and the ensuing domestication provide evidence that different cereal species were domesticated independently in different areas. Some sites were not situated near wild cereal habitats and a few were located outside the limits of distribution, even accounting for moister climatic conditions. I argue here that current models which try to explain the shift to farming have tended to over-emphasize the effect of the Younger Dryas climatic change. First, it would have had only a minor effect on cereal availability. Secondly, agriculture appears to have been established after the Younger Dryas. Thirdly, there is no evidence for a single centre of origin; agriculture arose in widely separated geographic and climatic regions. And fourthly, agriculture depends on stable climatic conditions which were not established until after the Younger Dryas.     

Late Celtic and early Roman plant remains from the oppidum of Bibracte, Mont Beuvray (Burgundy, France)

Two single assemblages of cereals and chaff from the oppidum Bibracte, (Burgundy, France), give evidence of cereal trade, food supply and crop processing in late Celtic and early Roman times. The first deposit consists mainly of Triticum dicoccon chaff which was buried in a wooden box near the sanctuary of La Terrasse. The second assemblage was a store of different wheats and barley excavated in a Roman cellar at the Pature du Couvent. The analyses of these finds are presented, and recent archaeobotanical research at Mont Beuvray is summarized.     

Archaeobotanical analysis of radiocarbon-dated plant remains with special attention to <Emphasis Type="Italic">Secale cereale</Emphasis> (rye) cultivation at the medieval village of Mankby in Espoo (Finland)

Several urban medieval sites in Finland (ad 1000–1500) have been archaeobotanically investigated, but until now only a few rural medieval settlements have been studied. Therefore, many aspects of the crops and agricultural methods of medieval villages in Finland still remain unknown. At the 12th to mid-16th century medieval village of Mankby, Espoo, a systematic sampling for archaeobotanical material was carried out. The analysed samples derived from hearths, building contexts and medieval field layers etc. The archaeobotanical material was carbonised, and it consisted mostly of conifer needles, cereal grains and weed seeds. Secale cereale (rye) and Hordeum vulgare (barley) were the only crop species found, Secale being the most frequent. Seeds of Bromus secalinus (rye brome) and achenes of Centaurea cyanus (cornflower), which are associated with autumn sown winter cereal crops, were also present. The archaeobotanical material shows areas of activity linked to storing crops, as well as cultivation within the village boundaries. Based on radiocarbon dated grains, cultivation of the fields started in the 12th century and continued until the abandonment of the village in 1556. Archaeobotanical material from Mankby has been compared to other contemporary sites in Finland, and differences between the sites can be seen. Results from Mankby have added to our knowledge of medieval agriculture, past areas of activity and economic conditions within the village.     

Informal intercropping of legumes with cereals? A re-assessment of clover abundance in ancient Egyptian cereal processing by-product assemblages: archaeobotanical investigations at Khentkawes town,Giza (2300–2100 <Emphasis Type="SmallCaps">bc</Emphasis>)

The possibility that legumes were specifically cultivated as a separate fodder crop in ancient Egypt has been inferred, usually on the basis of abundance of both legume seeds and or dung in charred macro-botanical samples, combined with a lack of wood charcoal; the implication being that a scarcity of wood led to the use of dung as fuel, and that the legumes in the assemblage derive from livestock which had been fed with cultivated fodder. The archaeobotanical remains from excavations at the Old Kingdom ‘Khentkawes town’ (2300–2100 bc) on the Giza plateau in Egypt contained an abundance of legumes, but also much wood charcoal, and preservation of many fragile and ‘green’ seeds and plant parts. This assemblage has led to questioning of the theory of specific fodder cultivation in Pharaonic Egypt. In this article, alternative interpretations of legume-rich assemblages of cereal processing by-products are investigated. Intercropping of legumes with cereals is one of the most widespread and effective methods of improving crop value and security, and fodder/forage quality. Analysis of this assemblage has led to a hypothesis that Trifolium sp. and other ‘weeds’ may well have been viewed as integral plants within ancient Egyptian cereal fields, due to an awareness of the benefits of intercropping legumes with cereals—as opposed to having been specifically cultivated as a monocrop.     

Evidence of ‘new glume wheat’ from the Late Neolithic (Copper Age) of south-eastern Hungary (4th millennium cal. b.c.)

In 2000, remains of an unknown Triticum species—later named ‘new glume wheat’ (NGW)—were identified in the archaeobotanical material of Neolithic and Bronze Age Greek sites. The presence of NGW was later reported from several other locations across Europe, from the seventh to the first millennium cal. b.c. During the systematic archaeobotanical survey of the multiperiod site of Hódmez?vásárhely–Kopáncs I., Olasz-tanya (5310–2936 cal. b.c.) more than 2,000 cereal remains were recovered. During the morphological analyses, ten spikelet forks showed the distinctive traits of NGW, therefore morphometric analyses were conducted on the remains to reinforce the morphological identification. The results suggest that both approaches—morphological and morphometric—should be applied in parallel to securely separate the NGW remains from Triticum turgidum L. ssp. dicoccum (Schrank) Thell. (emmer) and T. monococcum L. ssp. monococcum (einkorn). All NGW glume bases were recovered from Late Copper Age features (3338–3264 cal. b.c.) of the settlement, which represent the Baden culture of the Great Hungarian Plain. Similarly to other Baden culture sites of the Carpathian Basin einkorn and emmer dominated the crop production of the settlement. The ratio of the NGW remains within the cereal assemblage was measured to be 0.48 %, which suggests that NGW did not have the status of a regular crop; still it may have been part of the accompanying weed flora of the cereal fields during the fourth millennium in the south-eastern Great Hungarian Plain landscape.     

Increasing habitat connectivity in agricultural landscapes as a weed management strategy reconciling ecology and agronomy

While studies have explored how habitat amount drives weed assemblages in agroecosystems, knowledge remains limited of the effects of habitat connectivity. The response-effect trait framework provides insights into the mechanisms underpinning the relationship between landscape structure and the taxonomic diversity and abundance of weed assemblages. This study evaluated how habitat connectivity and habitat amount affect weed diversity and abundance in winter cereal fields, and whether these effects are driven by the functional composition of weed assemblages. We sampled weeds in 27 winter cereal fields. We measured habitat connectivity and habitat amount provided by wooded, grassland and cropland elements. We selected five traits related to the dispersal, establishment, and competitive abilities of weed species likely to respond to landscape structure: seed number per plant, type of reproduction, seed dry mass, plant vegetative height and seed germination rate. The functional composition of weed assemblages was assessed using community weighted mean trait values. Weed diversity and abundance were used as proxies of weed management. The taxonomic approach did not reveal any effect of landscape structure on weed diversity and abundance. Only the grassland elements that contributed to habitat connectivity, and to a lesser extent to habitat amount, drove the functional composition of weed assemblages. High habitat amount favoured species with many seeds, while high habitat connectivity favoured species with fewer seeds, a higher ability to reproduce vegetatively and higher seed germination rates. In turn, higher seed germination rates increased weed evenness and reduced weed abundance. Some of these relationships were influenced by the presence of rare species. Overall, high connectivity provided by grassland elements increases weed evenness and reduces weed abundance by shaping weed functional composition. Our study suggests that land-use planning policies that enhance the connectivity provided by grassland elements could be considered as a weed management strategy reconciling ecology and agronomy.     

From foraging to farming in the southern Levant: the development of Epipalaeolithic and Pre-pottery Neolithic plant management strategies

This paper reviews the archaeobotanical record of the transition from foraging to farming in the southern Levant. The concise presentation of the published botanical evidence follows a critical assessment of: (a) the nature of Epipalaeolithic plant management strategies, (b) the place of the southern Levant in the polycentric development of Near Eastern plant cultivation and domestication, and (c) region-specific pathways for the emergence of domesticated crop “packages”. Some inferences are drawn and suggestions are made concerning the potential contribution of archaeobotanical research to questions of broader archaeological significance about socio-economic change in the southern Levant during the Pre-pottery Neolithic.     

Heterogeneity from increasing crop types: effect on carabid beetles

We investigated the effect of crop type heterogeneity on carabid beetles in intensively managed agricultural landscapes. We compared different crops and habitat type configurations and compositions. In addition we assessed the effect of artificial fragmentation by paved roads and water channels in an agricultural landscape. Crop type heterogeneity negatively affected carabid assemblages. Spatial dissection by artificial structures had an even more negative effect on carabid diversity. The nested structure of the carabid assemblages among crop types was caused by a negative response to heterogeneity in the agricultural landscape. Even though crop types in an intensive agricultural landscape increased, the landscape may tend to fragment habitat. Although artificially fragmented habitats had more unstable carabid assemblages, heterogeneity from different crop types has also been shown to be a characteristic of fragmented landscapes. To produce a positive relationship between biodiversity and heterogeneity in an agricultural landscape, farming intensity should be reduced to allow for the conversion of highly productive lands into more natural habitats.     

Mycorrhiza‐mediated interference between cover crop and weed in organic winter cereal agroecosystems: The mycorrhizal colonization intensity indicator

The mycorrhizal fungi are symbiotic organisms able to provide many benefits to crop production by supplying a set of ecosystem functions. A recent ecological approach based on the ability of the fungi community to influence plant–plant interactions by extraradical mycelium development may be applied to diversified, herbaceous agroecosystems. Our hypothesis is that the introduction of a winter cereal cover crop (CC) as arbuscular mycorrhizal fungi (AMF)–host plant in an organic rotation can boosts the AMF colonization of the other plants, influencing crop–weed interference. In a 4‐years organic rotation, the effect of two winter cereal CC, rye and spelt, on weed density and AMF colonization was evaluated. The AMF extraradical mycelium on CC and weeds roots was observed by scanning electron microscopy analysis. By joining data of plant density and mycorrhization, we built the mycorrhizal colonization intensity of the Agroecosystem indicator (MA%). Both the CC were colonized by soil AMF, being the mycorrhizal colonization intensity (M%) affected by environmental conditions. Under CC, the weed density was reduced, due to the increase of the reciprocal competition in favor of CC, which benefited from mycorrhizal colonization and promoted the development of AMF extraradical mycelium. Even though non‐host plants, some weed species showed an increased mycorrhizal colonization in presence of CC respect to the control. Under intense rainfall, the MA% was less sensitive to the CC introduction. On the opposite, under highly competitive conditions, both the CC boosted significantly the mycorrhization of coexistent plants in the agroecosystem. The proposed indicator measured the agroecological service provided by the considered CCs in promoting or inhibiting the overall AMF colonization of the studied agroecosystems, as affected by weed selection and growth: It informs about agroecosystem resilience and may be profitably applied to indicate the extent of the linkage of specific crop traits to agroecosystem services, contributing to further develop the functional biodiversity theory.     

Cultivation and processing of Linum usitatissimum and Camelina sativa in southern Scandinavia during the Roman Iron Age

Charred seed conglomerates of Linum usitatissimum (flax) and Camelina sativa (gold of pleasure) were found at Uppåkra 2:25, a Roman Iron Age site in Skåne, southern Sweden. The conglomerates showed no mixing with each other, as they were almost pure flax and gold of pleasure respectively. Together with other archaeobotanical data from the site, they provide new evidence on the use, processing and cultivation of these two plants in early Iron Age in Scandinavia. Metric analyses were applied to flax seeds from both conglomerates and other contexts at this site, and compared to seed assemblages from other Roman Iron Age sites in Europe. The comparison showed that the flax cultivation at Uppåkra 2:25 was intended for the production of oil-rich seeds. The contextual relationship indicates that both flax and gold of pleasure seeds were processed in a similar way and used for oil. Furthermore, the pure seed conglomerate of gold of pleasure suggests that this plant was not a weed, but rather an intentionally grown crop which was cultivated separately from flax.     

Diversity in Kodo millet,Paspalum scrobiculatum

Paspalum scrobiculatum is widely distributed in damp habitats across the Old World tropics. It is harvested as a wild cereal in west Africa and in India.The species was domesticated in India some 3,000 yr ago. It is grown in India from Kerala and Tamil Nadu in the south, to Rajasthan, Uttar Pradesh and West Bengal in the north. Kodo millet is variable but lacks racial differentiation. The species was apparently domesticated across its range of present-day cultivation. HybridizationwithwildP. scrobiculatum , which commonly invades fields of kodo millet, obscures racial differentiation. Weedy kodo is harvested with the crop, making it difficult to distinguish wild and cultivated complexes ofP. scrobiculatum. Wild, weed and cultivated kinds merge in all characters studied.     

Long‐term farming systems and last crop sown shape the species and functional composition of the arable weed seed bank

Questions: The assembly of arable weed communities is the result of local filtering by agricultural management and crop competition. Therefore, soil seed banks can reflect the effects of long‐term cumulative field management and crop sequences on weed communities. Moreover, soil seed banks provide strong estimates of future weed problems but also of potential arable plant diversity and associated ecological functions. For this, we evaluated the effects of different long‐term farming systems under the same crop rotation sequence on the abundance, diversity and community assembly of weed seed bank, as well as on the functional diversity and composition. Location: DOK (biodynamic [D], bioorganic [O], conventional [K]) long‐term trial, Therwil, Switzerland. Methods: The effects of long‐term contrasted farming systems (i.e., biodynamic, organic, conventional, mineral and unfertilised systems) and last crop sown (i.e., wheat and maize) were evaluated on different indicators of species and functional diversity and composition of the weed soil seed bank. Results: The results showed significant influences of 40 years of contrasted farming systems on the diversity and composition of the seed bank, with higher diversities being found in unfertilised and organic farming systems, but also higher abundances than those found under conventional systems. Organic farming also allowed higher functional richness, dispersion and redundancy. Different farming systems triggered shifts in species and functional assemblies. Conclusions: The results highlight the importance of organic management for the maintenance of a diverse arable plant community and its functions. However, such results emphasise the need for appropriate yearly management to reduce the abundance of settled weediness and prevent affecting crop production. The farm management filtered community composition based on functional traits. Although the soil seed bank buffers the long‐term farming and crop sequence, the last crop sown and, thus, the yearly management were important determinants of seed bank composition.     

Effects of different cultivation types on native and alien weed species richness and diversity in Moravia (Czech Republic)

Changes in weed species richness and beta-diversity are partly attributable to different types and intensity of disturbance and partly to broad-scale variation in environmental conditions. We compiled a data set of 434 vegetation plots of weed vegetation in root crop and cereal fields in Moravia (eastern Czech Republic) to compare the effects of environmental conditions and different disturbance regimes on species richness and beta-diversity. To detect changes in species richness, we related the variation in species richness to individual environmental conditions. To assess differences in beta-diversity between the vegetation of cereal and root crop fields, we used Whittaker's measure of beta-diversity. The relative importance of each environmental variable for the variation in species composition was evaluated using canonical correspondence analysis. All analyses were done for all vascular plant species and separately for native species, archaeophytes and neophytes. A comparison of weed vegetation of root crops and cereals showed a distinct dichotomy between these two types of weed vegetation. There was no significant difference in total species richness and native species richness; however, cereal fields were richer in archaeophytes and root crop fields were richer in neophytes. The beta-diversity of weed vegetation was higher in root crops. Environmental factors explained a significant part of the variability in richness of both natives and aliens. The richness of native species increased and beta-diversity decreased with increasing precipitation. The opposite relationship was found for archaeophytes, in both cereals and root crops. These results confirmed the importance of climatic factors and management practices for changes in weed species composition. They also showed a distinct pattern of species richness and beta-diversity of native and alien weed species.     

Organic farming expansion drives natural enemy abundance but not diversity in vineyard‐dominated landscapes

Organic farming is seen as a prototype of ecological intensification potentially able to conciliate crop productivity and biodiversity conservation in agricultural landscapes. However, how natural enemies, an important functional group supporting pest control services, respond to organic farming at different scales and in different landscape contexts remain unclear. Using a hierarchical design within a vineyard‐dominated region located in southwestern France, we examine the independent effects of organic farming and semi‐natural habitats at the local and landscape scales on natural enemies. We show that the proportion of organic farming is a stronger driver of species abundance than the proportion of semi‐natural habitats and is an important facet of landscape heterogeneity shaping natural enemy assemblages. Although our study highlights a strong taxonomic group‐dependency about the effect of organic farming, organic farming benefits to dominant species while rare species occur at the same frequency in the two farming systems. Independently of farming systems, enhancing field age, reducing crop productivity, soil tillage intensity, and pesticide use are key management options to increase natural enemy biodiversity. Our study indicates that policies promoting the expansion of organic farming will benefit more to ecological intensification strategies seeking to enhance ecosystem services than to biodiversity conservation.     

Genetic diversity and phylogeography of broomcorn millet (Panicum miliaceum L.) across Eurasia

Broomcorn millet (Panicum miliaceum L.) is one of the world's oldest cultivated cereals, with several lines of recent evidence indicating that it was grown in northern China from at least 10,000 cal bp. Additionally, a cluster of archaeobotanical records of P. miliaceum dated to at least 7000 cal bp exists in eastern Europe. These two centres of early records could either represent independent domestications or cross-continental movement of this cereal that would predate that of any other crop by some 2 millennia. Here, we analysed genetic diversity among 98 landrace accessions from across Eurasia using 16 microsatellite loci, to explore phylogeographic structure in the Old World range of this historically important crop. The major genetic split in the data divided the accessions into an eastern and a western grouping with an approximate boundary in northwestern China. A substantial number of accessions belonging to the 'western' genetic group were also found in northeastern China. Further resolution subdivided the western and eastern genepools into 2 and 4 clusters respectively, each showing clear geographic patterning. The genetic data are consistent with both the single and multiple domestication centre hypotheses and add specific detail to what these hypotheses would entail regarding the spread of broomcorn millet. Discrepancies exist between the predictions from the genetic data and the current archaeobotanical record, highlighting priorities for investigation into early farming in Central Asia.     

Aspects of early medieval farming from sites in Mediterranean France

This paper reviews archaeobotanical remains (seeds and fruits) from some early medieval sites in the Mediterranean area of southern France (Lower Languedoc, Roussillon, Provence and Corsica). Four periods are distinguished, from the 5th-6th to the 11th-12th centuries A.D. The influence upon the results of the reliability of the samples, and the type of features from which they came, is discussed. The few sites with abundant material limit the evidence for the farming system. A comparison with northern France shows differences in the role of some crops in the local economy. The main crop plants were Triticum aestivum/durum, Hordeum vulgare and Vitis vinifera, followed by relatively smaller amounts of Avena sativa. Secale cereale and Panicum miliaceum occur sporadically. Specific identification of hulled wheat (T. dicoccum/spelta and T. cf. monococcum) is equivocal. The occurrence of Vicia faba var. minor, Pisum sativum, Lathyrus cicera and Linum usitatissimum varies between sites. The number of Lens culinaris finds indicates its probable cultivation. The occurrence of Cicer arietinum is limited in assemblages of this area. As in previous centuries, Vitis vinifera is the most common fruit of French medieval sites. Written sources from Catalonia and Languedoc and results from rescue excavations indicate that the nature of viticulture changed after the Gallo-Roman period. Other cultivated fruits, Olea europaea, Prunus dulcis, P. domestica, P. avium, P. persica, Pinus pinea, Morus nigra, Ficus carica, Juglans regia and gathered wild fruit occur rarely. The farming, which was based on mixed farming, includes use of wetland and meadow.