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.     

Combining functional weed ecology and crop stable isotope ratios to identify cultivation intensity: a comparison of cereal production regimes in Haute Provence,France and Asturias,Spain

This investigation combines two independent methods of identifying crop growing conditions and husbandry practices—functional weed ecology and crop stable carbon and nitrogen isotope analysis—in order to assess their potential for inferring the intensity of past cereal production systems using archaeobotanical assemblages. Present-day organic cereal farming in Haute Provence, France features crop varieties adapted to low-nutrient soils managed through crop rotation, with little to no manuring. Weed quadrat survey of 60 crop field transects in this region revealed that floristic variation primarily reflects geographical differences. Functional ecological weed data clearly distinguish the Provence fields from those surveyed in a previous study of intensively managed spelt wheat in Asturias, north-western Spain: as expected, weed ecological data reflect higher soil fertility and disturbance in Asturias. Similarly, crop stable nitrogen isotope values distinguish between intensive manuring in Asturias and long-term cultivation with minimal manuring in Haute Provence. The new model of cereal cultivation intensity based on weed ecology and crop isotope values in Haute Provence and Asturias was tested through application to two other present-day regimes, successfully identifying a high-intensity regime in the Sighisoara region, Romania, and low-intensity production in Kastamonu, Turkey. Application of this new model to Neolithic archaeobotanical assemblages in central Europe suggests that early farming tended to be intensive, and likely incorporated manuring, but also exhibited considerable variation, providing a finer grained understanding of cultivation intensity than previously available.     

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     

Taxonomic and functional characteristics of field edge weed communities along a gradient of crop management intensity

The widespread loss of weed diversity and associated ecosystem functions is raising important concerns. Field edges could play a major role in the maintenance of weed functional diversity in arable landscapes as these habitats still harbour high weed diversity, owing to either a reduced farming management intensity and/or to a spillover of species from adjacent perennial field margins. Here, we investigated the taxonomic and functional characteristics of weed species recorded in surveys of field edges and their associated field cores over six consecutive years in 60 arable fields farmed with five crop management strategies. We found that field edges were richer, with species more functionally diverse and composition more stable over years than field core surveys. The distribution of individual functional traits differed between field edges and field cores, with higher values for seed mass and nitrophily (Ellenberg.N), and a wider distribution of specific leaf area values in field edges. The bimodal distribution of plant height and germination period observed in field edges became unimodal in field cores. Field edges harboured species with ecological strategies associated with field cores (ruderal species) plus a conservative strategy which could be explained by a spillover from the adjacent perennial field margins. Crop management strategies impacted field edge flora, though to a lesser extent than the field core flora whereas the functional differences between the field edge and the field core flora were less marked when crop management intensity was lower. These results indicate that field edges harbour a unique assemblage of species and highly contribute to the maintenance of weed diversity in arable landscapes. Future studies should thus focus on the importance of these specific functional traits to the agroecosystem functioning.     

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.     

Early Iron Age agriculture: archaeobotanical evidence from an underground granary at Overbyg»rd in northern Jutland, Denmark

In the Late Pre-Roman Iron Age an underground granary in the village of Overbyg»rd was destroyed by fire and the contents were carbonised. Almost 2000 years later, analyses of the macroremains of the granary, which included a range of processed and unprocessed crops and weed seeds, showed that naked barley (Hordeum vulgare var. nudum) and bread wheat (Triticum aestivum s.l.) were the main crops cultivated, hulled barley (Hordeum vulgare) and flax (Linum usitatissimum) also played a role, whereas emmer (Triticum dicoccum) and gold of pleasure (Camelina sativa) were present as weeds or contaminants. The arable weed flora suggests that crops were sown in spring and that ecological conditions in the arable fields were very variable. The crops were harvested on the straw and may have spent some time drying and maturing in the fields before being transported home to be stored as severed ears in pest-proof granaries. Winnowing or, more probably, casting appears to have been used to clean the crop after threshing. It could not be ascertained if the crops had been sieved. Large collections of weed seeds in the granary were apparently the result of intentional gathering for food, rather than by-products of crop processing. In the light of the investigation it is suggested that future research into Iron Age agrarian practices should include both the analysis of archaeobotanical finds and a programme of practical experiments. This dual approach will give us a much better understanding of arable agriculture, not only in the Iron Age, but in prehistory as a whole.     

The origin of crop weed communities composed of summer annuals

Summary Evidence for the origin and dating of crop weed communities composed of summer annuals is to be found in palaeobotanical and archaeological data. There is archaeological evidence that around the middle of the first millennium BC fields were taken into permanent cultivation, even on the poorer soils.Fertilization with lime and manure and Plaggenwirtschaft were evidently involved in this early agriculture. In this paper the idea is put forward that Plaggenwirtschaft resulted in a sudden increase in the number of weed species brought together in cornfields and that this practise led to the very first beginning of our modern weed communities.Translated into English by Drs. C. van Driel-Murray.     

Functional approaches for assessing plant and invertebrate abundance patterns in arable systems

A functional group approach was developed for plant and invertebrate assemblages from UK arable fields to assess the variation in functional composition of these highly disturbed, managed systems. Data were taken from the Farm-Scale Evaluations (FSE) of genetically modified herbicide-tolerant (GMHT) crops where the impact of management of the GMHT crop has been assessed for winter and spring sown oilseed rape, beet and maize. Twenty plant and 36 invertebrate functional groups were defined according to trophic behaviour and traits that affect resource capture, quality and availability. The functional composition of the plant community was significantly affected by season of sowing, the type of crop sown and, to a lesser extent, herbicide management. The invertebrate community composition was also affected by crop type and sowing season, but not by management. Resource and consumer groups were positively related, and data provide strong evidence for top-down control of herbivore populations. Two main interaction groups were identified within the arable food web: one between omnivores, generalist predators and detritivores, which are positively associated with monocots, and one between omnivores, parasitoids, sap feeders and leaf chewers, which have a stronger association with dicots. Although management has an impact on within-field arable biodiversity, crop type and sowing season have an overriding effect on the functional composition of plant and invertebrate assemblages in arable systems.     

Cereal density and N-fertiliser effects on the flora and biodiversity value of arable headlands

Modern intensive farming caused pronounced changes to the European arable flora. Many species adapted to less intensive traditional farming declined severely, as did the potential of unsown arable vegetation to support higher trophic levels. To reverse these trends, various agri-environment measures were introduced. One such measure is to manage cereal headlands as conservation headlands, involving strict restrictions on pesticide and fertiliser use. An additional modification to management which could reduce crop competition and thus deliver benefits to arable plants is cereal sowing at reduced rates. However, little is known about its benefits to rare and declining arable plants, or to species of value to higher trophic levels, and whether it can be implemented without concomitant increase in undesirable weeds. We set up identical two-factorial experiments in winter wheat and spring barley, combining a nitrogen fertiliser versus no fertiliser treatment with cereal sowing at economic rates versus sowing at rates reduced by 75 %, with added sowing of a mixture of rare arable species. Both experiments also included an uncropped but cultivated control equivalent to another agri-environment measure. Our results show that reduced cereal sowing in conservation headlands can benefit rare and declining species, as well as arable plant diversity, without necessarily resulting in a concomitant increase in undesirable weeds. While such benefits tended to be larger in uncropped cultivated controls, conservation headlands have the advantage of not requiring land being taken out of production. Moreover, as shown in this study, their benefits to arable plants can be maximised by reduced sowing.     

Response of the two rare arable weed species Lithospermum arvense and Scandix pecten-veneris to climate change conditions

Rare weeds are currently under pressure due to intensifying arable management practices, and as a consequence of climate change, these practices will likely become even more intensive, together with a greater uniformity of land use. As a result, ecological stresses will increase for most species of rare weeds, in some cases leading to their further decline or even extinction. Moreover, climate change will alter the suitability of the environment for many plants, since average temperatures are predicted to increase and precipitation extremes to become more common. For most arable weed species it is unclear, whether the anticipated changes in environmental conditions are disadvantageous or beneficial. Little is known about specific biological responses of rare weeds to climate changes, and this study attempts to close some of these knowledge gaps. Here, the rare arable weed Lithospermum arvense and the endangered arable species Scandix pecten-veneris were investigated with regard to the effects of higher temperature and different crop densities on flowering time, shoot development, plant height, dry mass and seed production. Semi-field experiments were conducted with winter wheat crop for 3 years, involving 48 climate cages, in which every second was a variant of warmer temperature and contrasting crop density. We observed that S. pecten-veneris flowered earlier under warmer conditions and had fewer seeds and less biomass in the dense wheat crop compared to control conditions, while L. arvense grew taller, it produced fewer seeds in the high density crop. We suggest that such data concerning the biological responses of weeds can improve the precision of bioclimatic distribution models. Finally, we discuss strategies, such as relocation or non-intrusive management practices, for preventing further disappearances of rare arable weeds. Our results should be of considerable interest for the fields of plant ecology, biodiversity research and conservation.     

Arable weed seeds as indicators of regional cereal provenance: a case study from Iron Age and Roman central-southern Britain

The ability to provenance crop remains from archaeological sites remains an outstanding research question in archaeology. Archaeobotanists have previously identified the movement of cereals on the basis of regional variations in the presence of cereal grain, chaff and weed seeds (the consumer–producer debate), and weed seeds indicative of certain soil types, principally at Danebury hillfort. Whilst the former approach has been heavily criticised over the last decade, the qualitative methods of the latter have not been evaluated. The first interregional trade in cereals in Britain is currently dated to the Iron Age hillfort societies of the mid 1st millennium bc. Several centuries later, the development of urban settlements in the Late Iron Age and Roman period resulted in populations reliant on food which was produced elsewhere. Using the case study of central-southern Britain, centred on the oppidum (large fortified settlement) and civitas capital of Silchester, this paper presents the first regional quantitative analysis of arable weed seeds in order to identify the origin of the cereals consumed there. Analysis of the weed seeds which were present with the fine sieve by-products of the glume wheat Triticum spelta (spelt) shows that the weed floras of samples from diverse geological areas can be separated on the basis of the soil requirements of individual taxa. A preliminary finding is that, rather than being supplied with cereals from the wider landscape of the chalk region of the Hampshire Downs, the crops were grown close to Late Iron Age Silchester. The method presented here requires further high quality samples to evaluate this conclusion and other instances of cereal movement in the past.     

The potential of lentil (Lens culinaris) as a grain legume crop in the UK: an assessment based on a crop growth model

A crop growth model developed in Canterbury, New Zealand was used to assess the potential of lentil (Lens culinaris) as a grain legume crop in the UK. The model was validated using five sowing dates at Durham (54.77°N, 1.58°W) in 1999. Predicted time to flowering was within 7 days of actual time to flowering and predicted seed yields were within 9% of actual yields. Actual yields ranged from 1.40 to 1.65 t ha‐¹. Seed was of high quality. The model was used to predict rate of development and yields of spring and autumn sown lentils at eight sites along a transect from NW Scotland (Stornoway, 58.22°N, 6.32°W) to SE England (East Mailing, 51.28°N, 0.45°E) chosen to encompass important environmental gradients in the UK. In general, for a 1 May sowing with 150 or 250 mm plant available water (PAW) and a 1 October sowing with 150 mm PAW, predicted mean values over the period 1987–95 for maximum crop growth rate, maximum leaf area index, radiation intercepted, total dry matter produced and seed yield were closely positively related to monthly mean values for mean daily air temperature and increased along the transect from NW to SE UK. Time to flowering generally decreased along the transect from NW to SE UK ranging from 28 June to 9 July and from 20 May to 14 June with the May and October sowings respectively. For the 1 May sowing with 250 mm PAW, predicted mean seed yield ranged from 1.00 to 1.90 t ha‐1. For all sites, yield was very stable over the 9 yr period. For the 1 May sowing with 150 mm PAW, predicted mean seed yield ranged from 0.97–1.23 t ha‐1. Yields for the four more southerly sites were more variable at the lower PAW and, in exceptionally dry years, were substantially lower than average. For these sites, autumn sowing increased seed yields in exceptionally dry years and gave similar or greater mean seed yields to spring sowing with 250 mm PAW. For East Mailing, predicted yields for autumn sowing were on average 2.78 t ha‐1. Also, for Stornoway, because of its relatively high overwinter temperatures, the model predicted substantial increases in yield with autumn sowing. It is concluded that lentil has considerable potential as a grain legume crop in the UK.     

The impact of changing the season in which cereals are sown on the diversity of the weed flora in rotational fields in Denmark

1. Surveys have shown that there has been a dramatic decrease in the weed flora of fields under rotational cultivation during the last 30 years. This trend has been particularly noticeable in winter cereals, a crop of increasing importance in the landscape.
2. The weed flora of spring and winter cereals was compared in 19 unsprayed fields during a 5-year study to test the hypothesis that cereal type exerts no effect on the flora or on the absolute and relative abundance of single species.
3. Plant and species densities, and accumulated species richness, were lower in winter than in spring cereals.
4. Floristic similarity was greater among spring cereal fields and between spring and winter cereals within the same fields than among winter cereal fields.
5. Species that occurred with unequal density in spring and winter cereals occurred at higher densities in the spring cereals; these species germinated mainly in the spring. However, for a few species the relative plant abundance was highest in winter cereals; these species were able to germinate both in the spring and autumn.
6. Some species – on the relative scale – occurred indifferently of season of sowing; all but one of these species were able to germinate both in the spring and autumn.
7. Plant species and taxa that are important food resources for arthropod herbivores occurred at greater densities in spring than in winter cereals and, in addition, occurred with the highest relative abundance in spring cereals.
8. Change in land use from spring to winter cereals involves not only an immediate reduction of more than 25% in the density of plants and species, but also a change and increased uncertainty in the composition of the weed flora. These findings may have serious implications for the ecology of wildlife in the agricultural landscape.     

The conservation of arable weeds at crop edges of barley fields in northeast Spain

The effects of weed control practices and fertilisation on weed flora and crop yield were evaluated in crop edges of barley fields in northeastern Spain. The study was carried out in four organic and four conventional barley fields. In each field, four permanent plots were delimited at the crop edge, and fertilisation and weed control treatments in a factorial design were applied over 3 years. Weed composition and the aboveground biomass of weeds and barley were recorded before the crop harvest in the first and the third year. We found relatively low values of species richness per field, as well as low values of weed biomass, especially in the organic crop edges (3.9% of total biomass). Weeds were significantly reduced by herbicide applications on conventional fields and were not affected by weed harrowing on organic fields or fertilisation. These results demonstrate that specific measures are needed to enhance biodiversity at crop edges both in organic and conventional fields. Our results also suggest that under Mediterranean conditions and among impoverished weed communities, limiting the use of herbicides is crucial to enhancing arable diversity and that, contrary to findings found in previous studies in temperate climates, fertilisation and weed harrowing have little effect on weeds.     

Timing of disturbance and vegetation development: how sowing date affects the weed flora in spring‐sown crops

Abstract. The number of annual weeds were recorded in 752 field experiments in spring‐sown cereal crops conducted in Sweden 1972–1993. Two null hypotheses were tested regarding how the sowing date influenced the weed flora. 1. There is no relationship between the weed flora composition and sowing date. A pCCA (with geographic regions, crop species and soil types as covariables) clearly refuted this hypothesis. Hence, the composition of the weed flora varied depending on sowing date. 2. Species classified as summer annuals, winter annuals and germination generalists (that can germinate substantially in both spring and autumn) do not differ in their placement along the first ordination axis in the pCCA, i.e. according to sowing date. An ANOVA was unable to reject this hypothesis. Hence, germination syndrome classification did not explain the observed community differences related to sowing date. These results illustrate the importance of the date of disturbance for any secondary succession involving a seed bank and also the importance of annual dormancy cycles in seed banks.     

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     

Responses of plants and invertebrate trophic groups to contrasting herbicide regimes in the Farm Scale Evaluations of genetically modified herbicide-tolerant crops

Effects of genetically modified herbicide-tolerant (GMHT) and conventional crop management on invertebrate trophic groups (herbivores, detritivores, pollinators, predators and parasitoids) were compared in beet, maize and spring oilseed rape sites throughout the UK. These trophic groups were influenced by season, crop species and GMHT management. Many groups increased twofold to fivefold in abundance between early and late summer, and differed up to 10-fold between crop species. GMHT management superimposed relatively small (less than twofold), but consistent, shifts in plant and insect abundance, the extent and direction of these effects being dependent on the relative efficacies of comparable conventional herbicide regimes. In general, the biomass of weeds was reduced under GMHT management in beet and spring oilseed rape and increased in maize compared with conventional treatments. This change in resource availability had knock-on effects on higher trophic levels except in spring oilseed rape where herbivore resource was greatest. Herbivores, pollinators and natural enemies changed in abundance in the same directions as their resources, and detritivores increased in abundance under GMHT management across all crops. The result of the later herbicide application in GMHT treatments was a shift in resource from the herbivore food web to the detritivore food web. The Farm Scale Evaluations have demonstrated over 3 years and throughout the UK that herbivores, detritivores and many of their predators and parasitoids in arable systems are sensitive to the changes in weed communities that result from the introduction of new herbicide regimes.     

杂草科学管理——理论基础与实施途径

Scientific management of weeds is theoretically based on ecology.The implementation methods incladeIntensifying the weed biology and ecology research, especially those of the heavy weeds;Intensifying the research of developing competition between the crop and the weed;Utilizing a allelopathy the gene engineering and breeding against the weeds;Utilizing allelopathy between the crop and the weed, and utilizing biological and agricultural measurements to control the weeds.