Agroecological weed control using a functional approach: a review of cropping systems diversity

Agriculture since the 1950s has shown pronounced trends toward specialisation and intensification. Intensive measures have been taken for crop protection against pests through the widespread use of chemical pesticides in order to reduce the loss of agriculture yield. Although crop protection practices have reduced the overall potential losses of 50 % to actual losses of about 30 %, crop losses due to pests still vary from 14 to 35 % according to the country. Moreover, consequences of this intensive agriculture are now well known with an important increase of atmospheric CO2 concentrations, water pollution and biodiversity loss. Current challenge is thus to design alternative sustainable cropping systems which maintain food production while reducing externalities. Application of ecological principles to agroecosystems has been proposed for that purpose. Nonetheless, it is difficult since crop systems are characterised by frequent and high disturbances, high nutrient input and high pressure of human activities. Here, we review the current knowledge in weed science and functional ecology and propose a conceptual framework to analyse weed community assembly in arable fields. Cropping systems are highly variable in their combination of agricultural techniques. We designed a trait-based approach of functional diversity (1) to establish a comparative description of the environmental gradients created by cropping systems and (2) to characterise the response of weeds to environmental gradients. We categorise the effects of cropping systems on the environment into disturbance and resource gradients. Disturbances induced by actual and previous agricultural practices are split into physical and chemical components, whose regime are defined by disturbance timing and frequency. Resource availability in arable fields is described by the value of effect traits of crops, such as plant height, that are related to their use of resources. Finally, we provide a list of relevant response traits of weeds to each component of the two gradients.

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title: Identifying functional groups for response to disturbance in an abandoned pasture.
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title: The role of weeds in supporting biological diversity within crop fields
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title: Plant biodiversity enhances bees and other insect pollinators in agroecosystems. A review
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title: TILLAGE EFFECTS ON WEED SEED RETURN AND SEEDBANK COMPOSITION
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title: An on-farm approach to investigate the impact of diversified crop rotations on weed species richness and composition in winter wheat
abstract: Weed species diversity may benefit from organic farming due to enhanced temporal diversification of crop species in a rotation and omission of herbicide applications. However, in intensively managed conventional systems, little evidence exists as to what extent diversified crop rotations contribute to higher weed species richness. Using an on-farm approach, the effect of crop rotation (organic, conventional diverse (CD) and conventional simple (CS) crop rotations) and weed control (with vs. without) on weed species richness, cover, community composition and crop biomass, was analysed in 24 winter wheat fields. Weed species with beneficial functions for invertebrates and birds were analysed separately. Weed species richness was higher in the organic crop rotation, but did not differ between CD and CS crop rotations. Weed control treatment reduced species richness in both conventional rotations, but not in the organic one. Redundancy analyses revealed that crop rotation intensity accounted for the largest part of the explained variation in weed species composition. Results from the study indicate that the maintenance of weed species richness and conservation of species with important ecological functions requires not only temporal diversification of crop species in the rotation, but also an adjustment of weed control strategies.

title: Consequences of organic farming and landscape heterogeneity for species richness and abundance of farmland birds
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abstract: Recent studies have shown that relatively undisturbed plant communities, such as woodland and pasture, have generally low seed persistence, while seed longevity in frequently disturbed habitats, such as arable fields, is high. In addition, seed mass and shape were found to be closely linked to the living conditions of plants. The objective of the present study was to show how farming practice modifies these seed traits in the arable weed seed bank. On 67.4 ha of arable land at the Scheyern Research Station in Germany, conventional arable use was converted to organic farming, to a reduced-tillage system and to set-aside. During the six subsequent years, seed bank data were collected at 283 sampling points to analyse the effects of (1) the farming systems (long-term effects), (2) individual crops (short-term effects) and (3) vegetation cover. Set-aside arable land favoured a disk- or needle-like seed shape, greater mass and reduced seed longevity. Similarly, organic farming significantly increased seed mass and decreased longevity. Therefore, both types of land use reduced the selection for small and persistent seeds with a spherical shape. By contrast, an increasing persistence under reduced tillage suggested a higher selection pressure. The most consistent effect was that seed longevity increased with tillage frequency, independent from the farming system. Both high seed masses and a compact seed shape were frequently associated with a high crop cover. The results prove that beyond the properties of living plants the arable farming practice also significantly impacts the seed traits in the soil seed bank.

title: Management Filters and Species Traits: Weed Community Assembly in Long-Term Organic and Conventional Systems
abstract: Community assembly theory provides a useful framework to assess the response of weed communities to agricultural management systems and to improve the predictive power of weed science. Under this framework, weed community assembly is constrained by abiotic and biotic “filters” that act on species traits to determine community composition. We used an assembly approach to investigate the response of weed seed banks to 25 yr of management-related filtering in three different row-crop management systems in southeastern Pennsylvania: organic manure-based, organic legume-based, and conventional. Weed seed banks were sampled in April of 2005 and 2006 and quantified by direct germination in a greenhouse. We also assessed the filtering effects of weed management practices and relationships between assembled seed bank and emergent weed communities by allowing or excluding weed control practices within each management system and measuring emergent weed community response. Germinable weed seed bank densities and species richness in the final year of the study were over 40% and 15% higher, respectively, in the organic systems relative to the conventional system. Seed bank community structure in the organic systems was different from the conventional system, and the relationships between assembled seed banks and the emergent flora varied. Primary tillage, weed control, timing of planting, and fertility management appeared to be the main filters that differentiated weed seed banks in the three systems. Weed life history, emergence periodicity, seed size, and responsiveness to soil fertility and hydrology appeared to be the most important functional traits determining how weed species responded to management-related filters. Our results suggest that management systems can exert strong filtering effects that can persist over relatively long (greater than one growing season) time scales. Legacy effects of community-level filtering might be more important than previously assumed, and should be incorporated into predictive models of weed community assembly.

title: Environmental and management factors determining weed species composition and diversity in France
abstract: Multivariate analysis of data from approximately 700 arable fields from France was carried out to partition the respective importance of environmental factors versus management practices on weed species richness and composition. Overall, canonical correspondence analysis indicated that the major variations in species composition between fields were associated with human management factors; (1) the current crop type and (2) the preceding crop type. Three main weed communities were identified according to sowing season: winter, spring and summer-sown crops. The third most important gradient was associated with soil pH and soil texture to a lesser degree, resulting in highly contrasting weed communities on basic clay soils against those on acidic sandy soils. The influence of climate and geographical region was less pronounced and identified mainly through relationships with precipitation and longitude. Within individual crop types, the effect of other management practices became more prominent. Species richness is dependant on factors other than, or in addition to those influencing species composition, like those describing landscape organisation and/or tillage depth. Species richness (α-diversity) and community composition (β-diversity) had, for example, contrasting relationship to altitude: 300–450 m altitude giving high species richness but low species turnover. The variations observed in this large scale data set help to identify the agricultural practices which have had the most significant impact on the loss of species diversity in arable fields in recent decades.

title: Advances, challenges and a developing synthesis of ecological community assembly theory
abstract: Ecological approaches to community assembly have emphasized the interplay between neutral processes, niche-based environmental filtering and niche-based species sorting in an interactive milieu. Recently, progress has been made in terms of aligning our vocabulary with conceptual advances, assessing how trait-based community functional parameters differ from neutral expectation and assessing how traits vary along environmental gradients. Experiments have confirmed the influence of these processes on assembly and have addressed the role of dispersal in shaping local assemblages. Community phylogenetics has forged common ground between ecologists and biogeographers, but it is not a proxy for trait-based approaches. Community assembly theory is in need of a comparative synthesis that addresses how the relative importance of niche and neutral processes varies among taxa, along environmental gradients, and across scales. Towards that goal, we suggest a set of traits that probably confer increasing community neutrality and regionality and review the influences of stress, disturbance and scale on the importance of niche assembly. We advocate increasing the complexity of experiments in order to assess the relative importance of multiple processes. As an example, we provide evidence that dispersal, niche processes and trait interdependencies have about equal influence on trait-based assembly in an experimental grassland.

title: Trait-based approaches to unravelling the assembly of weed communities and their impact on agro-ecosystem functioning
abstract: Navas M-L (2012). Trait-based approaches to unravelling the assembly of weed communities and their impact on agro-ecosystem functioning. Weed Research52, 479–488. 
 
 
The trait-based approach to plant functional ecology has gained considerable attention over the last two decades, allowing ecologists to address questions relating to species distribution, community assembly and ecosystem functioning. We show here how this approach can be used to address these issues for weed ecology in a new way, allowing research to shift from purely weed control issues to a more global understanding of the impact of weed communities on the agro-ecosystem. We review how weed species are sorted by environmental factors and management according to the value of traits and the role thereof in the assembly of weed communities. How weed trait values and their distribution within communities affect agro-ecosystem processes is discussed in relation to loss of crop production. We also introduce the question of the impact of weed functional structure on ecosystem services and suggest some directions for research at species, community and agro-ecosystem levels.

title: A General Hypothesis of Species Diversity
abstract: A new hypothesis, based on differences in the rates at which populations of competing species approach competitive equilibrium (reduction or exclusion of some species), is proposed to explain patterns of species diversity. The hypothesis assumes that most communities exist in a state of nonequilibrium where competitive equilibrium is prevented by periodic population reductions and environmental fluctuations. When competitive equilibrium is prevented, a dynamic balance may be established between the rate of competitive displacement and the frequency of population reduction, which results in a stable level of diversity. Under conditions of infrequent reductions, an increase in the population growth rates of competitors generally results in decreased diversity. This model clarifies an underlying pattern of variation in diversity and points out the common elements of previous hypotheses. Rather than arguing that either competition, predation, or productivity control diversity, it demonstrates that all of these may contribute to the same basic mechanism. In doing so, it not only explains the correlations of the other hypotheses with patterns of diversity, but also explains the exceptions that these hypotheses could not explain. This hypothesis may be applied to variations of diversity both on a latitudinal gradient and within specific regions.

title: From Plant Traits to Plant Communities: A Statistical Mechanistic Approach to Biodiversity
abstract: We developed a quantitative method, analogous to those used in statistical mechanics, to predict how biodiversity will vary across environments, which plant species from a species pool will be found in which relative abundances in a given environment, and which plant traits determine community assembly. This provides a scaling from plant traits to ecological communities while bypassing the complications of population dynamics. Our method treats community development as a sorting process involving species that are ecologically equivalent except with respect to particular functional traits, which leads to a constrained random assembly of species; the relative abundance of each species adheres to a general exponential distribution as a function of its traits. Using data for eight functional traits of 30 herbaceous species and community-aggregated values of these traits in 12 sites along a 42-year chronosequence of secondary succession, we predicted 94% of the variance in the relative abundances.

title: Predicting changes in community composition and ecosystem functioning from plant traits: revisiting the Holy Grail
abstract: The concept of plant functional type proposes that species can be grouped according to common responses to the environment and/or common effects on ecosystem processes. However, the knowledge of relationships between traits associated with the response of plants to environmental factors such as resources and disturbances (response traits), and traits that determine effects of plants on ecosystem functions (effect traits), such as biogeochemical cycling or propensity to disturbance, remains rudimentary. 2. We present a framework using concepts and results from community ecology, ecosystem ecology and evolutionary biology to provide this linkage. Ecosystem functioning is the end result of the operation of multiple environmental filters in a hierarchy of scales which, by selecting individuals with appropriate responses, result in assemblages with varying trait composition. Functional linkages and trade-offs among traits, each of which relates to one or several processes, determine whether or not filtering by different factors gives a match, and whether ecosystem effects can be easily deduced

title: Scaling environmental change through the community-level: a trait-based response-and-effect framework for plants
abstract: Predicting ecosystem responses to global change is a major challenge in ecology. A critical step in that challenge is to understand how changing environmental conditions influence processes across levels of ecological organization. While direct scaling from individual to ecosystem dynamics can lead to robust and mechanistic predictions, new approaches are needed to appropriately translate questions through the community level. Species invasion, loss, and turnover all necessitate this scaling through community processes, but predicting how such changes may influence ecosystem function is notoriously difficult. We suggest that community-level dynamics can be incorporated into scaling predictions using a trait-based response-effect framework that differentiates the community response to environmental change (predicted by response traits) and the effect of that change on ecosystem processes (predicted by effect traits). We develop a response-and-effect functional framework, concentrating on how the relationships among species' response, effect, and abundance can lead to general predictions concerning the magnitude and direction of the influence of environmental change on function. We then detail several key research directions needed to better scale the effects of environmental change through the community level. These include (1) effect and response trait characterization, (2) linkages between response-and-effect traits, (3) the importance of species interactions on trait expression, and (4) incorporation of feedbacks across multiple temporal scales. Increasing rates of extinction and invasion that are modifying communities worldwide make such a research agenda imperative.

title: Patterns of plant traits in annual vegetation of man-made habitats in central Europe
abstract: Man-made habitats in central Europe can be broadly divided into arable land with weed vegetation, and settlements and their surroundings, harbouring ruderal vegetation. The former is a predictable environment with frequent, regular and large-scale disturbances, while the latter is an unpredictable environment with irregular disturbances of varying spatial extent producing heterogeneous mosaics of different successional stages. We hypothesize that these differences in disturbance regimes select for different sets of biological and ecological plant traits in these two habitats. A data set of 2715 vegetation plots sampled in man-made habitats dominated by annual plants in the Czech Republic was combined with data on biological and ecological traits of vascular plants, mostly taken from the BiolFlor database. Differences due to temporal variation and location of plots in different climatic zones were partialled out using partial canonical correspondence analysis. Then the differences in traits of the plants growing on arable fields and in settlements were analysed using logistic and least-square regression models, both with and without phylogenetic correction. Plants growing on arable land were more often annuals, R-strategists, with overwintering green leaves, insect or self-pollinated, reproducing by seeds, with persistent seed banks and archaeophytes (i.e. those aliens that arrived prior to 1500). Plants growing in human settlements were more often biennials or perennials, C-strategists, wind-pollinated, flowering in mid summer, reproducing both by seeds and vegetatively, dispersed by wind or humans, neophytes (i.e. those aliens that arrived after 1500), species with high demands for light and nutrients and with more continental distribution ranges. Most associations between plant traits and habitats did not change after taking phylogenetic relationships into account. Traits strongly linked to phylogeny were especially modes of pollination and dispersal. By contrast, traits weakly linked to phylogeny included life strategy and alien status.

title: A functional group approach to the management of UK arable weeds to support biological diversity
abstract: Weeds have an important role in maintaining farmland biodiversity. This needs to be balanced with their potential negative impact on crop yield and quality. Mechanistic models of crop-weed competition are an important tool in striking this balance. A range of common UK annual weeds were screened for the eco-physiological traits required by the models. Using multivariate techniques, a number of functional groups with a similar pattern of productivity and competition were identified, based on trade-offs between traits. A scheme was developed to assign species outside of the data set to one of the groups, based on life cycle, seed mass, maximum height and time of first flowering. As well as having a similar competitive ability, species within a group also appeared to have a similar ecosystem function, in terms of supporting higher trophic groups. Two beneficial groups of species were identified that combined a relatively low competitive ability with a high importance for invertebrates and birds. The identification of functional groups in the UK arable flora is a useful tool for assessing a weed community in the context of reconciling biodiversity provision with crop production. Preserving beneficial plant functional types within the crop would complement non-cropped wildlife refuges, such as field margins.

title: A functional analysis of large‐scale temporal shifts from 1970 to 2000 in weed assemblages of sunflower crops in France
abstract: What are the relationships between weed species traits and their change in distribution over a 30-year period? What does it tell us about factors that have driven shifts in the composition of weed communities? 
 
Location: France. 
 
 We analysed the links between change in the status of weed species in sunflower crops (decreasing or increasing) and a set of 17 traits using data sets collected in the 1970s and the 2000s, respectively. We analysed the contribution of traits to explain changes in the status of species both individually and in a multivariate way by mean of a clustering of species into functional groups. 
 
 69% of the most widespread species had significantly changed their frequency rank status over the last 30 years. Nearly two thirds of the increasing species belonged to a single functional group, out of the five groups identified in this analysis. Overall, the weed flora occurring in sunflower crops has specialised since the 1970s in favour of ‘sunflower mimicking’ functional groups: increasing species were more nitrophilous, more heliophilous, less sensitive to sunflower herbicides and shared a rapid summer life cycle. 
 
 The individual trait approach gave some indication as to the environmental factors likely to have caused the shift in sunflower weed communities. The functional group approach seemed to outperform direct trait comparisons as it accounted for major traits combinations i.e. cases where a species has a number of favourable traits, but is severely disadvantaged by the possession of one or a few deleterious traits.

title: Functional traits relating arable weed communities to crop characteristics
abstract: Which weed traits respond to the community assembly filters imposed by cropping regimes and how do they respond? Which crop characteristics (traits or aspects of field management) represent the strongest filters on weed traits? 
 
Location: France. 
 
 In 561 arable fields, we measured associations between crop characteristics and (i) mean values of quantitative weed traits at each site and (ii) composition of optimally defined weed functional types, based on both qualitative and quantitative weed traits and typologies. The crop characteristics included crop height, propagule size, family and sowing date; we also used basic types (winter wheat, maize, etc.) for comparison. 
 
 Crop sowing date was strongly related to many weed traits, whereas crop type was not the strongest predictor of any trait. With later sowing of crops, weeds started flowering later, germinated later and had shorter flowering periods. Sowing date was also associated with the distribution of Raunkiaer life forms, while crop architecture was associated with the distribution of weed heights and a C-S-R classification. 
 
 Cropping regimes can usefully be summarized by the crop sowing date. Phenological traits of weeds as well as classifications based on life form and C-S-R are important descriptors of weed community composition. Such findings may help predict the effects of particular crop rotations, novel crop varieties and invasive weed species.

title: Trajectories of weed communities explained by traits associated with species’ response to management practices
abstract: Two large-scale weed surveys conducted on winter wheat crops in France in the 1970s and the 2000s were used to determine the influence of management on weed communities. A trait-based approach was used to identify the mechanisms associated with the changing status of arable weeds over a 30 year period. A three-table ordination method (RLQ analysis) of the data set was performed to relate the environmental table to the species trait table using a species composition table to extract the joint structure (synchronic analysis). We then conducted a diachronic analysis to investigate the relationship between traits, alone or in combination, and the changing status of weeds. The synchronic analysis showed that tillage intensity filtered weeds according to height, seed weight, life forms and dispersal. Conversely, herbicides selected for species with delayed germination, which allows them to escape herbicide treatments. The diachronic analysis showed that successful weeds that have expanded were small plants, with rather light seeds, that can germinate over a long time frame during the vegetative period. This trait syndrome was probably favoured by profound changes in crop rotation and by increasing herbicide pressure. Our approach provides an excellent example of how future shifts in weed communities can be predicted and hence how weed management can be adapted so as to avoid promoting selection of problematic weeds.

title: A new model for the continuum concept
abstract: A reformulation of the continuum concept is presented after considering the implications of the community/continuum controversy and current niche theory. Community is a spatial concept dependent on landscape pattern while the continuum is an environmental concept referring to an abstract space. When applying niche theory to plants, the mechanisms of competition are ill-defined and the assumption of bell-shaped response curves for species unrealistic.

title: Remarkable changes of weed species in Spanish cereal fields from 1976 to 2007
abstract: Management practices, geographical gradients and climatic factors are factors explaining weed species composition and richness in cereal fields from Northern and Central Europe. In the Mediterranean area, the precise factors responsible for weed distribution are less known due to the lack of data and surveys. The existence of weed survey data of year 1976 in the Zaragoza province of the Aragón region, Spain, offered us the opportunity to compare present weed species with weed species growing 30 years ago. No detailed comparison of changes in weed species composition in cereal fields in that period of time has been conducted in the Mediterranean area. Here a survey was conducted in the Aragón region from 2005 to 2007. Weeds were surveyed in 138 winter cereal fields in ten survey areas where winter cereals are the main crops, using the same methodology applied 30 years ago. In the Zaragoza province, 36 fields were chosen in the same municipalities than in the previous survey. Several management, geographic and climatic variables of each field were recorded and related to weed species with multivariate analysis. Diversity index were calculated and related to survey area and altitude. Our results show that out of the 175 species only 26 species were found in more than 10% of the surveyed fields. The main species were Papaver rhoeas, Lolium rigidum, Avena sterilis and Convolvulus arvensis found in more than half of the surveyed fields. L. rigidum was related to dryland, while the other species were found overall. Furthermore, we found that management, geographical and climatic factors were significantly related to weed species distribution. In particular altitude, survey areas, irrigation and herbicide use in post-emergence were the most driving factors explaining weed species distribution. Species richness was higher in survey areas with extensive management practices and increased with altitude excepting a very productive area with intensive management practices at high altitude where richness was as low as in the irrigated lowlands. The main differences found between the 1976 and the 2005–2007 surveys were (1) the striking increase of grass weeds, (2) the high decrease of mean weed species number found in each field declining from 9 to 3 and (3) the frequency decrease of many weed species probably caused by agriculture intensification in that period of time. The growing importance of other weed species is probably related to their adaptation to minimum tillage, which is a widespread technique nowadays.

title: Floristic and ecological differences between recent and ancient forests growing on non-acidic soils
abstract: The aim of this work was to investigate differences in soil chemistry and understory composition between recent forests (sites afforested in the last 170 years) and ancient forests growing on non-acidic soils. The study was carried out on hardwood forests at moderate elevation (400–600 m asl) in the Jura Mountains (N.E. France) on four main pedological substrates with different characteristics. The floristic composition of 127 stands from recent forests (n = 65) or ancient forests (n = 62) was surveyed. Some functional traits and the Ellenberg indicator values of the surveyed species were recorded. In addition, the topsoil from 30 stands was analysed. The composition of the flora was analysed by Detrended Correspondence Analysis and the species which were typical of one class of forest age were identified using a chi-square (χ2) test. The difference between forest classes for plant traits, their indicator values, or soil chemistry was tested using the generalized linear model and Bonferroni t-tests (or Kruskall–Wallis tests). The floristic composition of the ancient forests was significantly different from that of the recent forests and was characterized by a high occurrence of shrub species in recent forests. These differences were associated with higher specific leaf area, low-range seeds dispersal, and some life forms like geophytes. There was no clear difference in soil chemistry between the two classes of forests, except for δ15N values. The weakness of the difference in the soil between ancient and recent forests suggested that changes in soil chemistry caused by a former agricultural land use were not responsible for the differences in understory composition recorded. The differences in functional traits between the two forest classes supported this conclusion. We finally concluded that (i) past land use modifies the vegetation composition of current forests, even on neutral soils and that (ii) in our context, biological filters were probably responsible for these changes.

title: Relative climatic, edaphic and management controls of plant functional trait signatures
abstract: To identify the relative roles of climatic, edaphic and management factors in controlling the weighted mean traits of vegetation. Eleven sites in Europe and one in Israel undergoing transitions in land use. Standardised methods were used to collect information on species traits and attributes from plots covering a range of land uses at each site. This was combined with abundance data to create a plot x trait matrix. Variance partitioning was used to identify the relative roles of climate, soil and management on the weighted and unweighted mean traits of the vegetation in the full data set, and the data set divided into vegetative traits (including life-form, clonality, defence and a range of leaf traits) and traits linked to regeneration via seeds (including seed mass, dispersal and pollination mechanism). Variance partitioning of the full data set showed that climate (18.7%), explained more variance in the weighted mean traits of the vegetation than climate and soil together (9.2), soil (6.9) and management (6.1). There was a similar distribution of variance explained for both vegetative and regeneration via seed traits, although more variance was explained for the latter. This restricted set of climatic, edaphic and management variables could explain 45-50% of the variance in the weighted mean traits of the vegetation between plots. There were only small differences between analyses of the weighted and unweighted data. Despite large variations in climate and soils between sites, there was still a separate and recognisable impact of management on the mean weighted traits of the vegetation. There was also a degree of shared variation between the three groups of factors, indicating that the response of plant traits to one group of factors may not be predictable because they may be modulated by their response to other groups.

title: Dynamics of weed populations
abstract: Dynamics of weed populations , Dynamics of weed populations , مرکز فناوری اطلاعات و اطلاع رسانی کشاورزی

title: Seed supply, drought, and grazing determine spatio-temporal patterns of recruitment for native and introduced invasive pines in grasslands
abstract: The rate of grassland invasion by trees depends on the ability of the species to invade, i.e. their invasiveness, and on the susceptibility of the environments to invasion, i.e. their invasibility. Knowledge of the invasiveness of native and introduced tree species and of the environmental factors that contribute to invasibility is necessary to understand landscape evolution and assess required management measures. Our main aim was to explore this by estimating the separate effects of propagule pressure and environmental factors on the spatio-temporal patterns of sapling recruitment, a key stage in the tree life cycle. Location Causse Mejean calcareous plateau (southern France).  The effects of seed supply and environmental variables (grazing, geological substrate, and duration or intensity of drought) on the spatio-temporal patterns of sapling recruitment were assessed for the native Scots pine ( Pinus sylvestris L.) and the introduced black pine ( Pinus nigra Arn. ssp. nigra ). Estimates were derived by inverse modelling with data of locations and ages of 4- to 20-year-old saplings and seed-bearing trees in 32 sites. Yearly indices of drought were derived from a soil‐water content model.  For both species, seed supply was as important as the whole set of environmental factors in explaining sapling recruitment rates. Grazing and the duration of drought from July to August decreased sapling recruitment rates, which were also lower on hard limestone than on dolomite. Dispersal distances and effective fecundities were higher for the introduced P. nigra , which was less susceptible to drought but more affected by grazing than the native P. sylvestris . In grazed grasslands, shrubs facilitated sapling establishment of both species.  This study shows how seed supply and environmental factors shape spatio-temporal patterns of sapling recruitment for trees invading grasslands. Implications for landscape evolution and management, of the difference in invasiveness of the two pine species and of the hierarchy of environmental factors in determining invasibility, are discussed.

title: Plant response traits mediate the effects of subalpine grasslands on soil moisture.
abstract: * In subalpine grasslands, changes in abiotic conditions with decreased management intensity alter the functional composition of plant communities, leading to modifications of ecosystem properties. Here, it is hypothesized that the nature of plant feedbacks on soil moisture is determined by the values of key traits at the community level.  As community functional parameters of grasslands change along a gradient of land uses, those traits that respond most to differences in abiotic conditions produced by land use changes were identified. A vegetation removal experiment was then conducted to determine how each plant community affected soil moisture.  Soil moisture was negatively correlated with community root length and positively correlated with canopy height, whereas average leaf area was associated with productivity. These traits were successfully used to predict the effects on soil moisture of each plant community in the removal experiment. This result was validated using data from an additional set of fields.  These findings demonstrate that the modification of soil moisture following land use change in subalpine grasslands can be mediated through those plant functional traits that respond to water availability.

title: Physiological Plant Ecology: Ecophysiology and Stress Physiology of Functional Groups
abstract: Plant ecophysiology is the science of the vital processes of plants in interaction with the environment. This advanced text offers insights into the laws governing the carbon, nitrogen, mineral and water balance, development, vital capacity and adaptability of plants. An extensive chapter deals with the effects of natural and anthropogenic stress factors. The book focuses on the ways the different plant species and functional types react in various locations and in all climatic zones.

title: Plant traits related to competition: how do they shape the functional diversity of communities?
abstract: The identification of functional traits critical to plant responses to the environment promotes our understanding of assembly of communities which relies on environmental filtering. However, the recent trait-community approaches mostly ignore the influence of plant-plant interactions by mainly focusing on traits related to abiotic filtering processes. The conceptual framework we propose aims to clarify how the functional diversity of communities depends on the filtering effect of competition on relevant traits. We define two types of competition-related traits: competitive effect traits reflect the changes in local resource levels due to plant activity while competitive response traits are related to plant response to these resource depletions. We then suggest that the contribution of both types of competition-related traits to functional diversity depends on the importance of competition, previously defined as the effect of competition on plant fitness relative to that of other environmental factors. The...

title: Intraspecific seed trait variations and competition: passive or adaptive response?
abstract: The phenotype of offspring depends on the abiotic and biotic environment in which the parents developed. However, the direct effects of competition experienced by parent plants on single-seed traits are poorly documented despite their impact on plant fitness. 2. We hypothesize that single-seed traits can differentially respond to the resource deficiencies of parent plants due to competition: seed quality may decrease as seed number does, magnifying the negative effects of competition for offspring ('passive response' hypothesis), or increase and then enhance offspring fitness to offset the reduction in offspring number ('adaptive response' hypothesis). Here we tested these hypotheses for four single-seed traits. We assessed the sensibility of their responses to changes in competition intensity due to species with different competitive effects and to contrasting soil nitrogen conditions. 3. In a common-garden experiment, four single-seed traits related to fitness - seed mass, seed nitrogen concentration (SNC), germinability and the timing of germination - were measured on a phytometer species transplanted in 14 different neighbours grown in monoculture with and without soil nitrogen limitation. 4. Under nitrogen-limiting conditions, the responses of SNC and of the timing of germination were passive and mainly related to the effects of neighbours on soil nitrogen availability, as shown by the increase in SNC with N-fixing neighbours. Within-individual seed mass variability decreased with increasing competition intensity, as an adaptive response to counterbalance the reduction in seed production. With nitrogen supplementation, competitors had no detectable effect on single-seed traits despite an overall increase in SNC and germination rate, confirming their nitrogen-dependent passive responses to competition. Germinability did not change among treatments. 5. The impact of competition on single-seed traits depends on both phytometer trait identity and resource modulation by neighbours. The passive response of seed chemical composition to competitors may magnify the competitive effects on offspring. By contrast, the adaptive response of seed size variability may offset these competitive effects. As a consequence, experiments looking at the fitness consequences of competition should not only consider the effects on fitness parameters of a target plant but also on the offspring.

title: Asymmetric competition in plant populations.
abstract: Recently there has been much interest in the hypothesis that competition between individual plants is asymmetric or onesided: larger individuals obtain a disproportionate share of the resources (for their relative size) and suppress the growth of smaller individuals. This has important implications for population structure, for the analysis of competition between plants at the individual, population and community levels, and for our understanding of competition as a selective force in the evolution of plant populations.

title: Evolutionary Agroecology: the potential for cooperative, high density, weed-suppressing cereals
abstract: Evolutionary theory can be applied to improve agricultural yields and/or sustainability, an approach we call Evolutionary Agroecology. The basic idea is that plant breeding is unlikely to improve attributes already favored by millions of years of natural selection, whereas there may be unutilized potential in selecting for attributes that increase total crop yield but reduce plants’ individual fitness. In other words, plant breeding should be based on group selection. We explore this approach in relation to crop‐weed competition, and argue that it should be possible to develop high density cereals that can utilize their initial size advantage over weeds to suppress them much better than under current practices, thus reducing or eliminating the need for chemical or mechanical weed control. We emphasize the role of density in applying group selection to crops: it is competition among individuals that generates the ‘Tragedy of the Commons’, providing opportunities to improve plant production by selecting for attributes that natural selection would not favor. When there is competition for light, natural selection of individuals favors a defensive strategy of ‘shade avoidance’, but a collective, offensive ‘shading’ strategy could increase weed suppression and yield in the high density, high uniformity cropping systems we envision.

title: Decomposition Analysis of Competitive Symmetry and Size Structure Dynamics
abstract: An analysis is introduced, based on the decomposition of relative growth rates, to examine the mode of competition (i.e. whether competition is symmetric or asymmetric), the size-dependence of growth, and their interdependence. In particular, the basis for two commonly held views is examined: (1) that the type of resource limitation determines the mode of competition, and (2) that asymmetric competition always leads to size-divergence between unequal competitors. It is shown that in field-grown millet plants, competition for light was symmetric at low density and asymmetric at high density. However, size variation at low density decreased during growth, because small plants had greater relative growth rates than larger plants. Size variation stayed constant at high density, since plants of all sizes had equal average relative growth rates. Based on these results and a general discussion, it is proposed that the type of resource limitation does not determine the mode of competition. Competition for light can be symmetric, and foraging for heterogeneously distributed soil resources can produce asymmetric competition below-ground. Furthermore, the mode of competition alone does not determine size structure dynamics. Size-dependence of resource conversion efficiency and allocation can modify the effects of resource uptake on growth.

title: Plant Ecological Strategies: Some Leading Dimensions of Variation Between Species
abstract: ▪ Abstract An important aim of plant ecology is to identify leading dimensions of ecological variation among species and to understand the basis for them. Dimensions that can readily be measured would be especially useful, because they might offer a path towards improved worldwide synthesis across the thousands of field experiments and ecophysiological studies that use just a few species each. Four dimensions are reviewed here. The leaf mass per area–leaf lifespan (LMA-LL) dimension expresses slow turnover of plant parts (at high LMA and long LL), long nutrient residence times, and slow response to favorable growth conditions. The seed mass–seed output (SM-SO) dimension is an important predictor of dispersal to establishment opportunities (seed output) and of establishment success in the face of hazards (seed mass). The LMA-LL and SM-SO dimensions are each underpinned by a single, comprehensible tradeoff, and their consequences are fairly well understood. The leaf size–twig size (LS-TS) spectrum has obvio...

title: A handbook of protocols for standardised and easy measurement of plant functional traits worldwide
abstract: There is growing recognition that classifying terrestrial plant species on the basis of their function (into 'functional types') rather than their higher taxonomic identity, is a promising way forward for tackling important ecological questions at the scale of ecosystems, landscapes or biomes. These questions include those on vegetation responses to and vegetation effects on, environmental changes (e.g. changes in climate, atmospheric chemistry, land use or other disturbances). There is also growing consensus about a shortlist of plant traits that should underlie such functional plant classifications, because they have strong predictive power of important ecosystem responses to environmental change and/or they themselves have strong impacts on ecosystem processes. The most favoured traits are those that are also relatively easy and inexpensive to measure for large numbers of plant species. Large international research efforts, promoted by the IGBP–GCTE Programme, are underway to screen predominant plant species in various ecosystems and biomes worldwide for such traits. This paper provides an international methodological protocol aimed at standardising this research effort, based on consensus among a broad group of scientists in this field. It features a practical handbook with step-by-step recipes, with relatively brief information about the ecological context, for 28 functional traits recognised as critical for tackling large-scale ecological questions.

title: Can traits predict the competitive response of herbaceous Mediterranean species
abstract: This study tested whether (i) functional traits, measured on individually grown plants, can be used to predict species response to competition; (ii) species competitive response depends on the standing biomass of neighbouring vegetation. Nine herbaceous Mediterranean species (targets) were grown as isolated plants or in mixture with one of three grasses (neighbours) differing in RGR, in pots during 4 months. Ten traits of targets, related to plant size, resource acquisition and conservation ability, and the above-ground biomass of neighbours were measured. Species with large basal area when individually grown were least suppressed by competition. No relationship was found between species competitive response and other traits measured on plants grown in isolation, including height and RGR. This result suggests that plant basal area could be used to predict the competitive ability of species in herbaceous communities. The competitive response of targets varied with the standing biomass of neighbours: it decreased with increasing neighbour biomass for low-productive mixtures, then reached a minimal value as neighbour biomass became larger, whatever the identity of neighbours. The implications of this relationship are discussed.

title: Leaf dry matter content and lateral spread predict response to land use change for six subalpine grassland species
abstract: Land-use change has a major impact on terrestrial plant communities by affecting fertility and disturbance. We test how particular combinations of plant functional traits can predict species responses to these factors and their abundance in the field by examining whether trade-offs at the trait level (fundamental trade-offs) are linked to trade-offs at the response level (secondary trade-offs).  Central French Alps.  We conducted a pot experiment in which we characterized plant trait syndromes by measuring whole plant and leaf traits for six dominant species, originating from contrasting subalpine grassland types. We characterized their response to nutrient availability, shading and clipping. We quantified factors linked with different land usage in the field to test the relevance of our experimental treatments.  We showed that land management affected nutrient concentration in soil, light availability and disturbance intensity. We identified particular suites of traits linked to plant stature and leaf structure which were associated with species responses to these environmental factors. Leaf dry matter content separates fast and slow growing species. Height and lateral spread separated tolerant and intolerant species to shade and clipping.  Two fundamental trade-offs based on stature traits and leaf traits were linked to two secondary trade-offs based on response to fertilization shade and mowing. Based on these trade-offs, we discuss four different species strategies which could explain and predict species distributions and traits syndrome at community scale under different land-uses in subalpine grasslands.  Tutin et al. (1964–1980).

title: Let the concept of trait be functional
abstract: In its simplest definition, a trait is a surrogate of organismal performance, and this meaning of the term has been used by evolutionists for a long time. Over the last three decades, developments in community and ecosystem ecology have forced the concept of trait beyond these original boundaries, and trait-based approaches are now widely used in studies ranging from the level of organisms to that of ecosystems. Despite some attempts to fix the terminology, especially in plant ecology, there is currently a high degree of confusion in the use, not only of the term "trait" itself, but also in the underlying concepts it refers to. We therefore give an unambiguous definition of plant trait, with a particular emphasis on functional trait. A hierarchical perspective is proposed, extending the "performance paradigm" to plant ecology. "Functional traits" are defined as morpho-physiophenological traits which impact fitness indirectly via their effects on growth, reproduction and survival, the three components of individual performance. We finally present an integrative framework explaining how changes in trait values due to environmental variations are translated into organismal performance, and how these changes may influence processes at higher organizational levels. We argue that this can be achieved by developing "integration functions" which can be grouped into functional response (community level) and effect (ecosystem level) algorithms.

title: Flowering phenology and height growth pattern are associated with maximum plant height, relative growth rate and stem tissue mass density in herbaceous grassland species
abstract: We hypothesize that flowering phenology correlates with plant height growth pattern and that the pattern is associated with functional traits including maximum plant height (Hmax), RGR, stem tissue mass density (SD), hollow ratio (proportion of central hollow of stem cross-sectional area) and leaf mass per area (LMA) in grassland herbaceous species. 2. We investigated plant height growth trajectories and flower phenology, and measured LMA, SD and hollow ratio for 25 herbaceous species including 20 dicot forb species and five monocot species in an old-field grassland of New England, USA. Hmax, RGR, T10 and T90 (Julian day when plant height was 10% and 90% Hmax respectively) were derived from a logistic function for each species and were analysed in relation to LMA and SD. 3. Hmax was positively correlated with T10, T90 and flowering onset time (Julian day when the first 10% of flowers were blossoming) across species and across evolutionary-correlated divergences. Early growing and flowering species were shorter than late ones, and species reaching Hmax earlier flowered earlier than their counterparts. 4. There was a positive relationship between T90 and RGR, in which early growing species were usually at mid-to-high levels of RGR, while late-growing ones had widely varied RGR. A similar relationship was found between flowering onset time and RGR. RGR was significantly negatively correlated with SD and LMA but positively with hollow ratio, as indicated by correlation analysis and phylogenetically independent comparative analysis. 5. Based on the above results, we propose that herbaceous species have two major dimensions of height growth strategies (early vs. late and fast vs. slow growth), collectively resulting in three extreme cases (early and fast, late and slow, and late and fast). Different height growth trajectories resulting from these strategies may reduce asymmetric competition among co-existing species in dense grasslands. 6. Synthesis. Flowering phenology and height growth patterns are significantly associated with functional traits such as RGR, LMA and hollow ratio in herbaceous grassland species. The difference in height growth trajectories and associated functional traits may allow species coexistence possibly at both plant and consumer trophic levels.

title: The influence of sowing rate and row spacing on the plant density and yield of red beet
abstract: Three experiments examined the effects of sowing rate and between-row spacing on the plant density and yield of red beet. The proportion of seeds which produced mature plants decreased when the mean distance to the nearest neighbour was less than 5 cm. In these experiments, this distance was governed by within-row spacing. Thus, plots with narrow-spaced rows achieved a higher plant density than those with wide-spaced rows, when sown with the same weight of seed. Total yield of beet per unit area decreased with increasing plant density. Maximum yields per unit area of small beet were achieved at high plant densities, whereas maximum yields of large beet were achieved at low plant densities. The effect of between-row spacing on yield was much smaller than that of density, and was important only for crops harvested early. Shoot yield per unit area was measured in two experiments and was not affected by row spacing in either. Shoot yield was not affected by plant density in one experiment, but, in the other, tended to a maximum value with increasing plant density.

title: A parsimonious combination of functional traits predicting plant response to disturbance and soil fertility
abstract: 1. How many traits associated with persistence and regeneration are necessary to predict the response of plants to soil fertility and disturbance? 2. Are correlated changes in trait expressions linked to the response of functional groups to fertility and disturbance?  Lower Frankonia, Germany.  On 120 plots located in managed and abandoned grasslands, fields, and vineyards, we recorded species composition, disturbance intensity, soil water and nutrients, and ten candidate traits for 75 species. We used a novel method which is based on three steps: (1) logistic regression to separate responsive from non-responsive species; (2) iterative clustering of all possible combinations of the candidate traits including regression of each cluster in response to the environmental variables; (3) selection of the trait combination that performed best with respect to goodness of fit of all clusters from this combination. Bivariate trait relationships across functional groups were analysed with reduced major axis regression (RMA).  The parsimonious trait combination consisted of life span, specific leaf area (SLA), canopy height, and seed number. The ‘acquisitive’ functional groups in terms of SLA and height were linked to higher fertility and earlier disturbance, while the ‘retentive’ groups related to lower fertility and later disturbance. Investment in reproduction, however, displayed a reverse relation. SLA and canopy height showed correlated shifts in two pairs of co-occurring functional groups.  A small number of traits is sufficient to predict the response of species. Plants need a higher increment in SLA to reach the same height, if start of disturbance is earlier. Linkages between traits shift from generative to vegetative with increasing fertility and earlier start of disturbance. Functional groups enable shifts in scaling relationships between traits to be analysed, in contrast to the analysis of single traits.  Rothmaler (1994).

title: Weed seedbank response to tillage, herbicides, and crop rotation sequence.
abstract: Changes in the weed seedbank due to crop production practices are an important determinant of subsequent weed problems. Research was conducted to evaluate effects of primary tillage (moldboard plowing and chisel plowing), secondary tillage (row cultivation), and herbicides on weed species changes in the soil seedbank in three irrigated row crop rotational sequences over a 3-yr period. The cropping sequences consisted of continuous corn for 3 yr, continuous pinto beans for 3 yr, or sugarbeets for 2 yr followed by corn in the third year. Cropping sequence was the most dominant factor in- fluencing species composition in the seedbank. This was partly due to herbicide use in each cropping sequence producing a shift in the weed seedbank in favor of species less susceptible to applied herbicides. A comparison between moldboard and chisel plowing indicated that weed seed of predominant species were more prevalent near the soil surface after chisel plowing. The number of predominant annual weed seed over the 3-yr period increased more rapidly in the seedbank after chisel plowing compared to moldboard plowing unless effective weed control could be maintained to produce a decline in seedbank number. In this case, seedbank decline was generally more rapid after moldboard plowing. Row cultivation generally reduced seedbanks of most species compared to uncultivated plots in the pinto bean and sugarbeet sequences. A simple model was developed to validate the observation that rate of change in the weed seedbank is influenced by type of tillage and weed control effectiveness.  Corn, Zea mays L.; pinto beans, Phaseolus vulgaris L.; sugarbeets, Beta vulgaris L. Additional index words: Weed shifts, reduced tillage, conservation tillage, population dynamics, modeling, KCHSC, AMARE, CHEAL, SOLSA, SETVI, ERACN.

title: Integrated Weed Management systems allow reduced reliance on herbicides and long-term weed control
abstract: Current concerns about the environmental impacts of pesticide use in agriculture require the investigation of novel cropping systems that would reduce their reliance on pesticides. In this paper, we report on an experiment carried out over 6 years to assess the performance of four cropping systems based on Integrated Weed Management (IWM) as compared to a reference standard system. Systems differed in crop rotations, soil tillage, mechanical and chemical weeding and crop management. Fewer herbicides were applied in IWM-based systems as compared to the reference, resulting in a lower environmental impact. Over the 6 years, we detected no significant increase in the density of winter or summer annual broad-leaved, and of grassy weeds in IWM systems, with one exception that is discussed. These results indicate that the combination of various IWM techniques allow both the long-term control of arable weeds and a significantly reduced reliance on herbicides.

title: Modelling rotations: can crop sequences explain arable weed seedbank abundance?
abstract: We investigated the effects of crop sequences on monocotyledon, dicotyledon and total weed seedbank abundance. Using seedbank data sampled from the conventionally cropped part of the GB farm-scale evaluations of genetically modified, herbicide-tolerant (GMHT) crops, we asked whether it is possible to identify crop sequence effects, to identify their duration and to simplify crop sequences into crop management classes with similar effects on weed seedbanks. This work showed that it is possible to detect historical effects of past crops, sown in sequence, on weed seedbanks for up to 3 years and that crop sequences may be simplified to crop management classes describing the season of sowing, crop type and weed target for herbicide application. Model estimates for the seedbanks were validated against an independent, follow-up seedbank data set. The analysis provided abundance estimates that ranged over 3 and 1.7 orders of magnitude for the monocotyledon and dicotyledon weed seedbanks for different crop sequences. This work yields a methodology for estimating seedbank abundance in current crop sequences, potentially allowing sequences to be identified that better reconcile the competing needs for weed control to maintain crop productivity and the demand for increased farmland biodiversity.

title: Weed seed predation increases with vegetation cover in perennial forage crops
abstract: Vegetation cover may affect weed seed predation by modifying the habitat quality for predatory organisms. Post-dispersal weed seed predation was measured by placing ‘seed cards’ in two perennial crops (alfalfa, cocksfoot) with and without crop cutting and in plots with bare soil. Each treatment was repeated four times in a randomized complete block design. Vegetation cover was measured by canopy light interception. Predation trials lasted two weeks and were repeated three times. Seed predation rates varied among three weed species (highest for Viola arvensis, intermediate for Alopecurus myosuroides, lowest for Sinapis arvensis). Vertebrate exclusion cages (12 mm × 12 mm openings) strongly reduced seed predation rates. Positive relationships were observed between vegetation cover and seed predation rates by both vertebrates and invertebrates for all weed species and trials, except when overall predation rates were very low. Predation rates were highest in uncut alfalfa, lowest on bare soil, but 16–64% of this variation could equally be explained by vegetation cover. The factorial design indicated that cutting had a stronger impact than crop species (legume or grass). Results suggest that weed seed predation may be enhanced by maintaining a high and temporally extended vegetation cover.

title: Timing of tillage is an important filter on the assembly of weed communities
abstract: A trait-based community assembly approach to weed management may enhance our understanding of how weed communities respond to specific management practices and increase the utility of weed management based on ecological principles. Therefore, identifying management practices that operate as assembly filters and the species traits upon which they act is an important first step in developing a more predictive weed science. Here, I report results from a 3-yr investigation of the effects of timing of annual tillage (spring vs. fall) on the annual assembly of arable weed communities. The timing of tillage had consistent and dramatic effects on the composition of weed communities; spring tillage led to weed communities dominated by early emerging spring annual forbs and C4 grasses, and fall tillage led to communities dominated by later-emerging forbs and C3 grasses. Traits determining a species' susceptibility to tillage time likely include germination syndrome and life cycle, both of which influence how species respond to changes in soil resource levels and light availability driven by seasonal disturbance regime. Manipulating the timing of tillage and other major soil disturbances may therefore be an important tool for managers interested in influencing community composition or targeting species with similar germination and life-history traits.

title: 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 regard- ing how the sowing date influenced the weed flora. 1. There is no relationship between the weed flora composition and sow- ing 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 so wing 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, germina- tion syndrome classification did not explain the observed com- munity 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.

title: The Search for Generality in Studies of Disturbance and Ecosystem Dynamics
abstract: Studies of disturbance have a long tradition in vegetation ecology (Cooper 1926; Raup 1941; White 1979) and have increased lly during the last 30 years (Dayton 1971; Heinselman 1973; Levin and Paine 1974; Borman and Likens 1979; Sousa 1979a,b, 1984; Pickett 1980; Pickett and White 1985; Van der Maarel 1993; Bornette and Amoros 1996; Paine et al. 1998; Freiich and Reich 1999; White et al. 1999). We have learned a tremendous amount about the significance of disturbance as an ecological factor in various habitats and communities (Knapp 1974; Grubb 1977; Miles 1979; Oliver 1981; Pickett and White 1985; Goldberg 1988; Frelich and Lorimer 1991; Milton et al. 1997), about disturbance regimes (Romme 1982; Turner et al. 1993; White et al. 1999), about functional adaptations of plants (Garcia-Mora et al. 1999; Walker et al. 1999), about responses of ecosystems (Bornette and Amoros 1996; Johnson et al. 1998; Engelmark et al. 1999) and about restoring disturbance as an ecosystem process (White and Walker 1997; Covington et al. 1999). During this period, a few theories and synthetic concepts have been proposed, but we do not yet have an inclusive general paradigm for this important body of work.

title: Characteristics of seeds and seedlings from weeds treated with sublethal herbicide doses
abstract: Size, germination and viability of seeds as well as growth of seedlings derived from three weed species were studied in a pot trial. Fallopin convolvulus (L.) A. Loeve, Galium. spurium L. and Thlaspi arvense L. were treated with MCPA or tribenuron-methyl at four doses and at five growth stages, from seedling stage to flowering, In G. spurium subnormal doses of tribenuron-methyl, applied at intermediate growth stages, greatly reduced seed weight, gennination, viability, seedling shoot biomass and root biomass. Germination and viability, as well as the shoot biomass and root biomass of seedlings, were highly correlated with seed weight. In addition, but to a smaller extent, seed weight was reduced in F. convolvidus and T. arvense by tribenuron-methyl and in G. spurium by MCPA. Germination was reduced in F. convolvulus by MCPA and in T. arvense by tribenuron-methyl. However, the effects varied greatly depending on the growth stage at application.

title: Morphological characterisation of soil structure in tilled fields: from a diagnosis method to the modelling of structural changes over time
abstract: Characterisation of soil structure within the tilled layer of cultivated fields is crucial because the importance of this soil characteristic on the biological, chemical and physical properties of the soil and its repercussions on water cycle, root growth and functioning. We present in this paper a method for field characterisation of soil structure. This method, practised since the 1970s, was designed for field diagnosis of the effects of cropping systems on soil structure. It is based on a stratification of the observation face of a pit dug perpendicular to the direction of tillage and traffic: spatial compartments are distinguished, according to the nature of the mechanical stresses they have been submitted to during tillage and crop management. Characterisation of soil structure is performed on a morphological basis, using two criteria, each of them addressing a specific organisation level of the soil: firstly, clods size distribution, proportion of fine soil and the way the clods are brought together are considered; then, secondly, the clods are classified in three types, on the basis of the importance and the origin of their internal structural porosity. Physical measurements (bulk density, compaction test, and water retention) are presented, which demonstrate that physical behaviour is different between clod types. These results justify the use of the method to model changes with time in soil structure, under the effects of the main factors affecting soil structure dynamics in tilled fields: compaction, fragmentation, climate and biological activity. A model which simulates at the field scale the changes over time of the proportion of compacted clods within the tilled layer is presented. In this model, the tilled layer is represented as a set of 1 cm 2 pixels, regularly located on a square grid. Each pixel is defined by its co-ordinates and a specific structure, compacted or non compacted. The pixel co-ordinates are modified during ploughing, for which the model calculates the lateral and vertical displacement of the soil. The structure of any individual pixel can be changed, depending on the soil condition and the operation type. This model was evaluated by comparing its outcomes to measurements obtained from a long term field experiment designed to study soil structure dynamics.

title: Integration of soil structure variations with time and space into models for crop management. A review
abstract: Soil structure plays a major role in the design of new crop management systems. For instance, the transition from conventional to no-tillage changes soil structure, which, in turn, has implications on crop yield greenhouse gas emissions, and pesticide and nitrate leaching. Modelling soil structure at field scale faces two main issues: (1) the spatial variability and (2) the temporal variability. Here, we review how spatial variability of soil structure is taken into account in water transfer models at field scale. We discuss the effects of soil structure on hydraulic properties. We present options to model soil structure effects using pedotransfer functions or calculations based on pore network geometry. Then we review studies on water transfer. Here, we show the utility of one-dimensional (1-D) and 2-D models, and the range of soil profile partitions. In the second part, we study a mean to model the temporal variation of soil structure. We propose an indicator of soil structure dynamics based on the proportion of compacted clods in the tilled layer. This indicator was measured from the observation face of soil pits. We studied this indicator in a long-term field experiment involving various risks of compaction. The results showed that this indicator gave a more precise description of the time course changes in soil structure than the mean soil bulk density measured on the same experimental plots. Lastly, we discuss the principles of a model that predicts the evolution of this indicator under different soil tillage and climatic conditions. This model can be used to evaluate the effects of different crop management systems on soil structure and soil water transfer.

title: Weed seedling emergence and seed survival: separating the effects of seed position and soil modification by tillage
abstract: Tillage causes both vertical redistribution of weed seeds and changes in soil physical properties. These two factors are rarely distinguished in studies of the impact of tillage on seedling emergence or seed survival. In this study, seeds of Chenopodium album L., Amaranthus retroflexus L., and Abutilon theophrasti Medik, were planted at particular depths in pots of undisturbed or stirred soil to separate these effects. Emergence and survival data were analysed by non-linear regression to determine the nature of significant differences between treatments. Emergence increased with slight burial and then decreased exponentially at greater depths. Average emergence over all depths was generally greater in tilled sou than in unfilled soil, particularly for C. album and A. retroflexus. Seed survival approached a maximum with increasing depth. Average survival of seeds that did not produce emerged seedlings was greater in tilled soil than in untilled soil for C. album and A. retroflexus. Thus, tillage affects emergence and seed survival of weeds through changes in soil conditions independently of effects resulting from redistribution of seeds in the soil profile.

title: Effects of soil depth and aggregate size on weed seed distribution and viability in a silt loam soil
abstract: The position of weed seeds within the soil matrix plays an important role in seedling emergence and seed survival. The relationship of weed seeds with soil aggregates and soil depth was evaluated in a Waukegon silt loam soil that had been under a long-term, conventional tillage, annual crop management system. Soil aggregates were separated and classified into eight size classes from ≤5 to >12 mm and weed seeds were extracted from the aggregates. Amaranthus spp., Chenopodium album L. (common lambsquarters), Polygonum pensylvanicum L. (Pennsylvania smartweed), Setaria faberi Herrm. (giant foxtail), and Solanum ptycanthum Dun. (eastern black nightshade) accounted for the majority of seeds recovered. In general, seed viability declined from April to June, but increased in October following seed deposition. Seeds of individual species were most abundant in the aggregate size class most closely matching its seed size. However, seeds were commonly found associated with aggregates larger than 9 mm. Highest seed viability was found in the aggregate fraction closest to the seed size, however, S. faberi viability was also high in the >12 mm aggregate size class. Regardless of aggregate size, seed numbers were generally greatest in the upper 5 cm of soil. The results of this research were species-dependent and variable and demonstrated the complexity of weed seed/soil aggregate associations. However, they did show that seed placement within the soil matrix may play an important role in weed population dynamics.

title: Effects of seed depth and soil aggregates on the emergence of weeds with contrasting seed traits
abstract: Gardarin A, Durr C & Colbach N (2010). Effects of seed depth and soil aggregates on the emergence of weeds with contrasting seed traits. Weed Research50, 91–101. 
 
 
 
Summary 
 
Tillage has a strong impact on weed emergence by burying seeds in the soil and modifying soil structure. Its influence can vary according to seed and seedling characteristics. This article focuses on shoot and radicle elongation in the soil and on seedling mortality caused by soil obstacles. Germinated seeds of nine contrasting weeds were planted in pots and grown in the dark to measure shoot and radicle elongation. Second, the proportion of seedlings blocked under aggregates (20–50 mm) was measured. Shoot growth rate, maximal shoot and radicle lengths were positively correlated with seed mass. Large-seeded species (e.g. Avena fatua) were more likely to emerge from greater depths (exceeding 20 cm deep). Seedling mortality increased with increasing obstacle size for all species; it was greater for monocotyledonous than for dicotyledonous species and decreased with the shoot diameter. Weed seed depth and soil structure influence emergence of weed species differently, depending on seed mass, shoot diameter and taxa. Including the results of this study in a cropping system-weed dynamics model would help to predict responses of weeds species to tillage.

title: Seasonal restrictions of bud growth on roots of Cirsium arvense and Sonchus arvensis and rhizomes of Elymus repens
abstract: Brandsaeter LO, Fogelfors H, Fykse H, Graglia E, Jensen RK, Melander B, Salonen J & Vanhala P (2010). Seasonal restrictions of bud growth on roots of Cirsium arvense and Sonchus arvensis and rhizomes of Elymus repens. Weed Research50, 102–109. 
 
 
 
Summary 
 
The success of weed management aimed at depleting the regenerative structures of perennial weeds depends largely on the sprouting activity of rhizome and root buds. Seasonal variation in sprouting of these buds on Cirsium arvense, Sonchus arvensis and Elymus repens was studied for plants collected from Denmark, Finland, Norway and Sweden. At 2-week intervals from July to October, 5-cm fragments of roots or rhizomes were cut from plants grown in buckets and planted into soil in pots, half of which were placed immediately into growth chambers at 18°C for 4 weeks. The other half of the pots were initially placed in a dark room at 2°C for 4 weeks before being transferred to the same growth chamber, also for 4 weeks. During the growth chamber period, the numbers of emerged shoots in each pot were counted weekly. The sprouting activity of C. arvense and E. repens was relatively uniform during this period and bud dormancy was not apparent. In all ecotypes of S. arvensis, innate bud dormancy developed during the latter part of the growing season. For all three species, differences in sprouting readiness were found among ecotypes. The results imply that C. arvense and E. repens are more likely to be controlled by mechanical measures in autumn than S. arvensis.

title: Cumulative effect of annually repeated passes of heavy agricultural machinery on soil structural properties and sugar beet yield under two tillage systems
abstract: The aim of this study was to determine potential cumulative effects of repeated passes with current heavy agricultural machinery on topsoil (0–0.3 m) and subsoil (below 0.3 m) physical properties of a Luvisol as affected by long-term tillage (annual mouldboard ploughing to 0.3 m depth (MP), shallow-mixing conservation tillage to 0.1 m depth (SM) with a wing-bladed rigid tine cultivator). Moreover, sugar beet yield was determined. Wheeling was conducted with a six-row self-propelled sugar beet harvester representing contemporary heavy agricultural machinery (wheel load 7.8–11.7 Mg, average ground contact pressure 100–145 kPa). Wheeling was applied once per year over three consecutive years after harvest of sugar beet, cereal and cereal, and moreover, independent from regular plot management with light experimental machinery. Soil moisture at wheeling (0–0.6 m depth) was around 100% field capacity in most years, which was secured by irrigation before wheeling if necessary. Repeated wheeling negatively affected penetration resistance, macropore volume (equivalent diameter >50 μm) and air permeability of topsoil (0.05–0.1 m, 0.18–0.23 m) and subsoil (0.4–0.45 m) layers, while biopore number and surface water infiltration remained unaffected. SM compared to MP tillage increased penetration resistance while decreasing macropore volume and air permeability in the 0.18–0.23 m layer, whereas reverse effects occurred in 0.4–0.45 m depth. Sugar beet yield was decreased by wheeling and SM tillage compared to the control treatments. No significant interactions between wheeling and tillage occurred in any parameter investigated. Conclusively, SM tillage did not provide better subsoil resistance against compaction compared to MP treatment under wheeling and soil conditions prevalent in our experiment. Repeated wheeling with heavy agricultural harvest machinery is obviously at risk to exceed the bearing capacity of susceptible soils. Although (i) under regular harvest conditions just small parts of arable fields (except headlands) are wheeled with high loads, (ii) harvest is by far not every year conducted under high soil moisture, and (iii) effects in the subsoil were small, such risks have to be taken into account. Reduction of tillage depth to

title: Weeds and Weed Management on Arable Land: An Ecological Approach
abstract: Classification of plants based on traits of ecological significance annual and perennial crops weed commodities looked upon as early stages in secondary vegetation succession weeds with diverse life forms in various types of crop germination, emergence and establishment of crop and weed plants competition in plant stands of short duration weed flora and weed plant adaption to environment and competitive conditions measurements of competition and competitiveness in plant stands of short duration soil tillage effects on weeds chemical weed control as an element in the cropping system special management measures knowledge of importance for understanding the occurrence and rational management of weeds. (Part contents)

title: Challenging Theophrastus: A common core list of plant traits for functional ecology
abstract: Ecologists need a common language of plant traits in order to make comparisons across regions and scales, pool data, and maximize the utility of the data. To develop such a set of traits we began with the primary challenges faced by plants: dispersal, establishment, and persistence in order to identify fundamental traits. Most of these traits are hard to measure, but advances in comparative ecology have suggested a number of easy to measure analogs. Unfortunately, some of the fundamental traits have no simple analog. The common core list includes: seed mass, seed shape, dispersal mode, clonality, specific leaf area, leaf water content, height, above-ground biomass, life history, onset of flowering, stem density, and resprouting ability. Most of the traits can be measured quantitatively, but several traits on the list must still be measured qualitatively due to logistical problems or lack of an easy analog. Key problem areas include: dispersal ability, capacity for vegetative spread, germination, palatability, plasticity, and all the various below-ground traits. Comparative studies need to address these problem areas. The common core list is suggested as a common starting point for studies of functional ecology. The idiosyncrasies of regional floras and specific research agendas will dictate which traits can be ignored and those that need to be added.

title: A leaf-height-seed (LHS) plant ecology strategy scheme
abstract: A leaf-height-seed (LHS) plant ecology strategy scheme is proposed. The axes would be specific leaf area SLA (light-capturing area deployed per dry mass allocated), height of the plant's canopy at maturity, and seed mass. All axes would be log-scaled. The strategy of a species would be described by its position in the volume formed by the three axes.The advantages of the LHS scheme can be understood by comparing it to Grime's CSR scheme, which has Competitors, Stress-tolerators and Ruderals at the corners of a triangle. The CSR triangle is widely cited as expressing important strategic variation between species. The C–S axis reflects variation in responsiveness to opportunities for rapid growth; in the LHS scheme, SLA reflects the same type of variation. The R axis reflects coping with disturbance; in the LHS scheme, height and seed mass reflect separate aspects of coping with disturbance.A plant ecology strategy scheme that permitted any species worldwide to be readily positioned within the scheme could bring substantial benefits for improved meta-analysis of experimental results, for placing detailed ecophysiology in context, and for coping with questions posed by global change. In the CSR triangle the axes are defined by reference to concepts, there is no simple protocol for positioning species beyond the reference datasets within the scheme, and consequently benefits of worldwide comparison have not materialized. LHS does permit any vascular land plant species to be positioned within the scheme, without time-consuming measurement of metabolic rates or of field performance relative to other species. The merits of the LHS scheme reside (it is argued) in this potential for worldwide comparison, more than in superior explanatory power within any particular vegetation region.The LHS scheme avoids also two other difficulties with the CSR scheme: (a) It does not prejudge that there are no viable strategies under high stress and high disturbance (the missing quadrant in the CSR triangle compared to a two-axis rectangle); (b) It separates out two distinct aspects of the response to disturbance, height at maturity expressing the amount of growth attempted between disturbances, and seed mass (inverse of seed output per unit reproductive effort) expressing the capacity to colonize growth opportunities at a distance.The advantage of LHS axes defined through a single readily-measured variable needs to be weighed against the disadvantage that single plant traits may not capture as much strategy variation as CSR's multi-trait axes. It is argued that the benefits of potential worldwide comparison do actually outweigh any decrease in the proportion of meaningful variation between species that is captured. Further, the LHS scheme opens the path to quantifying what proportion of variation in any other ecologically-relevant trait is correlated with the LHS axes. This quantification could help us to move forward from unprofitable debates of the past 30 years, where CSR opponents have emphasized patterns that were not accommodated within the scheme, while CSR proponents have emphasized patterns that the scheme did account for.

title: Evidence for the Existence of Three Primary Strategies in Plants and Its Relevance to Ecological and Evolutionary Theory
abstract: It is suggested that evolution in plants may be associated with the emergence of three primary strategies, each of which may be identified by reference to a number of characteristics including morphological features, resource allocation, phenology, and response to stress. The competitive strategy prevails in productive, relatively undisturbed vegetation, the stress-tolerant strategy is associated with continuously unproductive conditions, and the ruderal strategy is characteristic of severely disturbed but potentially productive habitats. A triangular model based upon the three strategies may be reconciled with the theory of r- and K-selection, provides an insight into the processes of vegetation succession and dominance, and appears to be capable of extension to fungi and to animals.

title: Distribution and Survival of Quackgrass (Elytrigia repens) Rhizome Buds1
abstract: An experiment was established at three sites to determine the longevity and distribution of rhizome buds of two quackgrass biotypes in the soil profile under two tillage systems (no-till or fall-plow). The experiments were established in May 1987 at sites initially free of quackgrass. Rhizome segments with three nodes were planted 2 to 5 cm deep to give an initial density of 21 buds m-2. Production of new buds was prevented in subsequent years. Rhizome bud populations were sampled every 6 mo between October 1987 and October 1989. Buds were produced until snow cover, indicating the need to control quackgrass until late fall to prevent a bud population increase. Under no-till conditions, 94% of quackgrass buds occurred in the top 10 cm of soil while 68 and 19% were found below 10 and 20 cm, respectively, in the fallplow treatments. A control program against quackgrass should aim at keeping the majority of rhizomes in the top 10 cm in order to facilitate control by having most shoots emerging at the same time. There was no statistically significant difference in bud longevity between biotypes, sites, and tillage systems. Viability of the rhizome buds declined rapidly during the first year. In most cases it took 2 yr for the bud populations to reach extinction although 0.6% of buds of one biotype survived as long as 30 mo in one of the three sites. Therefore, a minimum of 2 yr of total control would be required to eradicate quackgrass from most locations.  Bromoxynil, 3,5-dibromo-4-hydroxybenzonitrile; dicamba, 3,6-dichloro-2-methoxybenzoic acid; paraquat, 1,1'dimethyl-4,4'-bipyridinium ion; quackgrass, Elytrigia repens (L.) Nevski #3 AGRRE.  Agropyron repens, Elymus repens, rhizome bud longevity, AGRRE.

title: Ecological interpretation of weed flora dynamics under different tillage systems
abstract: In northern Italy, on soil managed with three different tillage systems (conventional tillage, ridge tillage, and no-tillage) and submitted to standard cultural practices (crop rotation, and chemical weed control), the weed vegetation was assessed at the beginning of the trial (1987) and after six, and eight years. The aims were to evaluate (1) the effect of tillage systems on the weeds; and (2) the possibility of linking the floristic changes under reduced disturbance to the theory of ecological succession. The weeds were categorised according to life-forms (biological groups), periodicity types (ecophysiological groups), dispersal types and seed longevity. Data were analysed using Sorenson's Indices of Similarity, the Independence test, and Principal Components Analysis. The tillage systems profoundly altered the weed community: in undisturbed soils the importance of the geophyte and hemicryptophyte species, and among the annuals, Digitaria sanguinalis, Conyza canadensis and Kickxia elatine increased, as well as that of the wind-dispersed weeds. The species linked to disturbance were annuals and in particular Amaranthus spp., Chenopodium album and Echinochloa crus-galli. After eight years the floristic evolution in the reduced tillage system can be interpreted on the basis of ecological succession. The community that has formed assumes, from the quantitative point of view, characteristics of a pioneer community of secondary succession with a predominance of annual species and a large number of wind-dispersed plants. Qualitatively there is a movement towards a more mature community that could become similar to that of the woodland edge, with more perennial species, shrubs, and bird-dispersed plants. The implications of these conclusions are discussed in terms of weed management.

title: Management in a modified no-tillage corn–soybean–wheat rotation influences weed population and community dynamics
abstract: Conservation tillage systems, such as no-tillage, are ecologically advantageous because they reduce soil erosion; however, they rely heavily on herbicide use. Our goal was to determine how weed communities of no-tillage systems are affected when the system is modified to reduce herbicide use through a combination of banded herbicides and interrow cultivation. To this end, we conducted a 9-yr study in a no-tillage corn–soybean–winter wheat rotation. All management systems had a preplant application of glyphosate, followed by either broadcast PRE herbicides (conventional no-tillage), interrow cultivation with banded PRE herbicides, or interrow cultivation alone. Aboveground weed densities were assessed each year and data were grouped into early (1991 to 1993) and late (1996 to 1998) time periods. Over time, weed communities became more distinct, showing a strong response to management and crop. In the early years, weed communities separated more in response to management than crop. In the late years, this was reversed. Weed communities in systems with interrow cultivation were more diverse than those in conventional no-tillage. The response to weed management system and crop was species specific. For example, the abundance of yellow foxtail was higher when interrow cultivation was employed, but abundance was equal in all crops. Dandelion was more abundant in conventional no-tillage of corn and soybean; however, it was equally abundant in all management systems in wheat. Seed bank species richness increased over time and was highest in systems with interrow cultivation. Herbicide use can be reduced in a modified no-tillage corn–soybean–wheat rotation by incorporating interrow cultivation, with or without banded herbicides, into the management plan. The weed community trajectory changes, and the weed community becomes more diverse. A more diverse weed community will not necessarily alter how we manage weeds.  Glyphosate; yellow foxtail, Setaria glauca (L.) Beauv.; dandelion, Taraxacum officinale Weber; corn, Zea mays L.; soybean, Glycine max (L.) Merr.; wheat, Triticum aestivum L.

title: The Selective Memory of Weed Seedbanks after 18 Years of Conservation Tillage
abstract: A conservation tillage study provided the opportunity to test whether tillage effects on the germinable weed seedbank would be consistent across different crop rotations and to investigate the potential residual effects of herbicide treatments terminated 12 yr earlier. Our objective was to measure the effects of tillage (moldboard plow [MP] vs. chisel plow [CP] vs. no-till [NT]), crop rotation (2-yr barley–red clover followed by 4-yr barley–canola–wheat–soybean rotation, compared to a cereal monoculture), and of a prior weed management factor (three intensity levels of herbicide use) on the density, diversity, and community structure of weed seedbanks. Species richness, evenness (Shannon's E), and diversity (Shannon's H′) of spring seedbanks varied little across treatments and over time. Total seedbank density generally increased as tillage was reduced, with some variations due to weed management in 1993 and crop rotation in 2006. Crop rotations generally had smaller seedbanks with fewer species than the monoculture. In 1993, seedbanks with minimum weed management were twice as dense as those with intensive or moderate weed management (approximately 6,000 vs. 3,000 seed m−2). By 2006, seed density averaged 6,838 seed m−2 across intensive and moderate weed management regardless of tillage, but was nearly twice as large in NT (12,188 seed m−2) compared to MP (4,770 seed m−2) and CP (7,117 seed m−2) with minimum weed management (LSD0.005  =  4488). Species with abundant seedbanks responded differently to treatments. Barnyardgrass and green foxtail had larger seedbanks in the monoculture than in the rotation. Common lambsquarters and pigweed species had large seedbanks in tilled treatments in the rotation, whereas yellow foxtail and field pennycress contributed to the large seedbanks observed in NT treatments. The latter two species were also associated with residual effects of weed management treatments (terminated 12 yr earlier) in NT. The differential seedbank response of weed species, attributed in part to contrasting weed emergence patterns and agronomic practice effects on seed rain, explained some of the weak treatment effects observed for total seedbank density and diversity. The large weed seedbanks observed in NT plots after 18 yr confirms the importance of seed rain and seedbank management for the sustainability of NT systems.  Barnyardgrass, Echinochloa crus-galli (L.) Beauv.; common lambsquarters, Chenopodium album L.; green foxtail, Setaria viridis (L.) Beauv.; field pennycress, Thlaspi arvense L.; pigweed species, Amaranthus sp.; yellow foxtail, Setaria pumila (Poir.) Roem & Schult.

title: Weed surveys in different tillage systems in southwestern Ontario field crops
abstract: A weed survey of 593 corn, soybean and winter wheat fields in southwestern Ontario was conducted during 1988 and 1989 to determine the abundance and distribution of weeds under a variety of tillage systems. The survey was conducted after all weed-control measures had been carried out. A total of 82 weed species and groups of species were recorded. Many weeds were found infrequently. The most abundant weeds were green foxtail (Setaria viridis (L.) Beauv.), lamb’s-quarters (Chenopodium album L.), quack grass (Agropyron repens (L.) Beauv.), redroot pigweed (Amaranthus retroflexus L.), common ragweed (Ambrosia artemisiifolia L.) and dandelion (Taraxacum officinale Weber). These weeds accounted for 54% of the total relative abundance. Weed communities in individual fields were highly variable. Most fields had fewer than 10 species, and nearly half of the fields had fewer than 6 weeds m−2. Six weeds, yellow foxtail (Setaria glauca (L.) Beauv.), quack grass, crab grass spp.

title: Using phenology prediction in weed management: a review
abstract: The success of weed management based on ecological principles and weed biology will depend on a better understanding of the effect of environment on lift history strategies, growth, and competition of weeds; and crops, and particularly upon the ability to predict weed and crop phenology, This paper reviews the importance of phenotypic plasticity to weed and crop competition and other biological interactions. We also discuss the utility of phenological predictions in weed management and review current weed phenology models that are based on thermal time. By understanding the variables that drive plant phenotypic responses, new approaches and more long-term solutions for weed problems can be developed.

title: The worldwide leaf economics spectrum
abstract: Bringing together leaf trait data spanning 2,548 species and 175 sites we describe, for the first time at global scale, a universal spectrum of leaf economics consisting of key chemical, structural and physiological properties. The spectrum runs from quick to slow return on investments of nutrients and dry mass in leaves, and operates largely independently of growth form, plant functional type or biome. Categories along the spectrum would, in general, describe leaf economic variation at the global scale better than plant functional types, because functional types overlap substantially in their leaf traits. Overall, modulation of leaf traits and trait relationships by climate is surprisingly modest, although some striking and significant patterns can be seen. Reliable quantification of the leaf economics spectrum and its interaction with climate will prove valuable for modelling nutrient fluxes and vegetation boundaries under changing land-use and climate.

title: Modelling assimilation rates of 14 temperate arable weed species as a function of the environment and leaf traits
abstract: Information on the response of assimilation rate to environmental factors is lacking for many less competitive weed species that need to be considered in the context of increasing farm biodiversity. A pot experiment was sown to parameterize gross assimilation rate at light saturation and initial light use efficiency for 14 common UK annual weeds and winter wheat at four leaf temperatures. Field experiments were also sown to measure inter-specific differences in specific leaf area (SLA), leaf nitrogen content and assimilation rates in the field at near-optimum temperatures. A generic relationship describing the response of assimilation rate to temperature and light using SLA and leaf nitrogen content as conversion factors successfully predicted inter-specific differences in assimilation rates in the field. This relationship could be incorporated into weed-crop competition models to predict the productivity and competitive impact of weed mixtures, including species outside the current data set. Assimilation rates at light saturation in the field were determined largely by SLA. This trait was variable between species and within a species across the growing season and needs to be well described in mechanistic competition models to accurately calculate instantaneous assimilation rates.

title: Competition, traits and resource depletion in plant communities
abstract: Although of primary importance to explain plant community structure, general relationships between plant traits, resource depletion and competitive outcomes remain to be quantified across species. Here, we used a comparative approach to test whether instantaneous measurements of plant traits can capture both the amount of resources depleted under plant cover over time (competitive effect) and the way competitors perceived this resource depletion (competitive response). We performed a large competition experiment in which phytometers from a single grass species were transplanted within 18 different monocultures grown in a common-garden experiment, with a time-integrative quantification of light and water depletion over the phytometers’ growing season. Resource-capturing traits were measured on both phytometers (competitive response traits) and monocultures (competitive effect traits). The total amounts of depleted light and water availabilities over the season strongly differed among monocultures; they were best estimated by instantaneous measurements of height and rooting depth, respectively, performed when either light or water became limiting. Specific leaf area and leaf water potential, two competitive response traits measured at the leaf level, were good predictors of changes in phytometer performance under competition, and reflected the amount of light and water, respectively, perceived by plants throughout their lifespan. Our results demonstrated the relevance of instantaneous measures of plant traits as indicators of resource depletion over time, validating the trait-based approach for competition ecology.

title: Seed mass and the competition/colonization trade-off: Competitive interactions and spatial patterns in a guild of annual plants
abstract: We used neighbourhood modelling to estimate individual-level competition coefficients for seven annuals growing in limestone grassland over 2 years. We calculated the relative strength of intra- and interspecific competition and related this to differences in seed size and plant size between targets and neighbours. 2 Significant differences in the impact of neighbours on each target species were observed in half the models fitted, allowing us to reject a null hypothesis of competitive equivalence. 3 In one year we found that as the seed size or plant size of neighbours increased relative to targets, so did their competitive effect. Although this is consistent with the competition/colonization trade-off model the competitive interactions were not sufficiently asymmetric to allow coexistence. In a second year we found only weak interspecific competition and no relationship with plant or seed size. 4 We found no overall relationship between competition coefficients and the degree of segregation, contradicting the spatial segregation hypothesis for coexistence. However, segregation was linked to differences in plant traits: when targets were smaller than neighbours the degree of segregation increased with relative neighbour size. 5 Most species were positively associated with each other due to a shared preference for otherwise unvegetated patches. The degree of association was negatively correlated with differences in plant and seed size, particularly when interspecific competition was weak. This might reflect (i) decreasing overlap in microhabitat use with increasing trait divergence or (ii) density-dependent mortality. 6 Seed size is a key trait within this group of species, determining both competitive and colonizing ability. The presence of such a competition/colonization trade-off undoubtedly stabilizes community dynamics although other mechanisms may also be at work.

title: Early season height differences as robust predictors of weed growth potential in maize: new avenues for adaptive management?
abstract: Weed interference in annual cropping systems can be highly variable from year-to-year, as well as spatially heterogeneous. These factors have confounded efforts to develop simple methods for predicting competitive outcomes early in the growing season. A 2-year study was conducted with maize in competition with four annual weed species (Setaria faberi, Abutilon theophrasti, Chenopodium album and Amaranthus retroflexus) to: (i) characterise the relationship between early crop–weed height differences and weed growth potential, and (ii) assess threshold-type decision rules based on these relationships. Analysis of 742 weed individuals suggests that early season height disparities are robust predictors of season-long growth. For example, weeds £8 cm tall when maize canopy was 60 cm tall had a very low probability (c. 2%) of growing larger than 150 cm and this applied across weed species, relative emergence dates and spatial proximity to the maize row. In our dataset, weeds less than 150 cm tall were associated with low crop yield losses and weed fecundity. Precision management of weed individuals based on early season height differences may provide a reliable and practical basis for selective post-emergence weed control that is relevant to both yield loss and longer-term seedbank

title: Differential response to competition in weedy biotypes of proso millet
abstract: Five weedy biotypes of proso millet (Panicum miliaceum L.) were grown under conditions of both intra- and inter-biotype competition. These biotypes represent the range of weediness from crop-like, large, white-seeded types with dense, drooping panicles to the most weed-like strain, which produces small, dormant, black seeds and has open, shattering panicles. Under noncompetitive conditions, the five biotypes of proso millet (WHITE, GOLDEN, ORANGE, CROWN and BLACK) differ strikingly in plant size, fresh weight distribution patterns, flowering time, and seed size. All biotypes showed decreased biomass per plant and delayed flowering time with increasing density when grown in monoculture. Variation in total biomass among individuals in each biotype was reduced at increasing densities resulting in positively skewed frequency distributions. The crop-like biotypes (WHITE and GOLDEN) demonstrated the greatest intrabiotype competitive effects, i.e., were the least tolerant of increased density in monoculture show...

title: Life-history variation in the annual arable weed Diplotaxis erucoides (Cruciferae)
abstract: Variation in life-history traits such as emergence, survival, time of flowering, and fecundity were studied in Diplotaxis erucoides, a mediterranean winter annual weed, by analyzing cohorts that emerged in autumn, early spring, and spring. The response of the plants to the environment, as reflected by plant architecture and pattern of biomass allocation, was also studied. Seedlings that germinate in autumn produced from 3 to 10 times more seeds than those that germinated in spring. The main factor affecting the number of seeds produced appears to be the life-span. Reduction of the growing period led to a decrease in both number and length of modular units, which resulted in decreased numbers of leaves, flowers, and fruits of each module. In semelparous D. erucoides plants, differences in the pattern of biomass allocation to reproduction are related to plant size. Our field data indicate that an increase of reproductive effort with size occurs in small individuals; however, a decrease occurs for vegetative...

title: Morphological analysis of herbaceous communities under different grazing regimes
abstract: A methodology for the morphological analysis of herbaceous communities is presented, together with an exam- ple of its application in montane grasslands in the province of C6rdoba (Argentina) subject to grazing and burning. The method, based on multivariate ordination and classifi- cation techniques, enabled the detection of morphological changes at three levels in response to disturbance: (a) charac- terization of the spatial structure of the vegetation; (b) identi- fication of morphological plant groups; and (c) quantification of morphological shifts among different individuals of a single species. The architecture of the vegetation changed toward a pro- gressive miniaturization of photosynthetic structures and con- centration of biomass close to the ground, as disturbance intensity increased. Six morphological plant groups (modes of response) showing different behaviour in relation to competi- tion for light and pressure from large herbivores were identi- fied. Some species, highly preferred by ungulate grazers, showed high morphological variability among morphs grow- ing in different grazing situations, whereas some others were morphologically uniform.

title: Quantifying trait selection driving community assembly: a test in herbaceous plant communities under contrasted land use regimes
abstract: Plant traits are particularly important in determining plant community structure. However, how can one identify which traits are the most important in driving community assembly? Here we propose a method 1) to quantify the direction and strength of trait selection during community assembly and 2) to obtain parsimonious lists of traits that can predict species relative abundances in plant communities. We tested our method using floristic data from 32 plots experiencing different treatments (fertilisation and grazing) in southern France. Twelve functional traits were measured on 68 species. We determined the direction and strength of selection on these 12 traits using a metric derived from a maximum entropy model (i.e. lambda). We then determined our parsimonious list of traits using a backward selection of traits based on these lambda values (for all treatments and in each treatment separately). We finally compared our method to two other methods: one based on iterative RLQ and the other based on an entropy-based forward selection of traits. We found major differences in the direction and strength of selection across the 12 traits and treatments. From the 12 traits, plant vegetative and reproductive heights, leaf dry matter content leaf nitrogen content, specific leaf area, and leaf phosphorus content were particularly important for predicting species relative abundances when considering all treatments together. Our method yielded results similar to those produced by the entropy-based approach but differed from those produced by the iterative RLQ, whose selected traits could not significantly predict species relative abundances. Together these results suggest that the assembly of these communities is primarily driven by a small number of key functional traits. We argue that our method provides an objective way of determining a parsimonious list of traits that together accurately predict community structure and which, despite its complementarities with entropy-based method, offers significant advantages.

title: Seed mass and the competition/colonization trade-off: a sowing experiment
abstract: 1  A seed-addition experiment using seven co-occurring annual plant species with a range of seed masses was carried out in a limestone grassland in South Wales. 
 
 
 
2  If seedlings compete for establishment sites, then large seed size may confer enhanced competitive ability. However, the simple reciprocal relationship found between seed mass and per capita seed output showed that species producing larger seeds suffer reduced fecundity. Seed size may therefore act as a surrogate in a competition/colonization trade-off. 
 
 
 
3  Equal numbers of seeds of all species were sown in a mixture over a range of densities. As sowing density increases, all species should reach a higher proportion of the available microsites. If large-seeded species are the best competitors they are expected to win all the sites they reach, and hence to occupy an increasing proportion of sites as sowing density increases. 
 
 
 
4  The three species with the largest seeds made up 49% of individuals at low-density sown plots but 83% of individuals in high-density sown plots. In addition, seed mass and plant density were not correlated in unsown plots, but were strongly correlated in high-density sown plots. However, all small-seeded species maintained a presence in sown plots. 
 
 
 
5  Although species were sown at random with respect to one another, individuals were up to five times more likely than expected to have a conspecific as a nearest neighbour. This could be caused by interspecific competition and/or by environmental heterogeneity that favours different species in different patches. 
 
 
 
6  The results suggest that seedlings do compete for establishment sites and that large-seeded species generally win when in direct competition. In unsown areas small-seeded species win many sites by forfeit (because large-seeded species are strongly recruitment limited) but there may be a restricted subset of potential sites for which they are the best competitors and which they can win outright.

title: Seedling survival and seed size: a synthesis of the literature
abstract: 1Large-seeded species have long been known to have higher survivorship during establishment than small-seeded species. Here, we assessed the size of this advantage by compiling published data on survival through seedling emergence, seedling establishment and sapling establishment. 
2We found no relationship between seed mass and survival through the transition from viable seed in or on the soil to newly emerged seedlings (P = 0.47, n = 33 species). 
3Synthesis of data from experimental studies on the advantages of large-seeded species establishing under particular hazards (such as shade, drought or herbivory) confirmed that seedlings of large-seeded species perform better than those of small-seeded species in most situations. However, the magnitude of this advantage was not sufficient to counterbalance the greater number of seeds produced by small-seeded species m−2 of canopy outline year−1. 
4Synthesis of data from field studies of populations under natural conditions also showed that large-seeded species have higher survival through early seedling establishment than small-seeded species (P = 0.006, n = 112 species). However, the magnitude of this advantage would only be sufficient to counterbalance the greater number of seeds produced by small-seeded species m−2 of canopy outline year−1 if mortality continued at the same rate for some time. 
5The time required for a species with 10-fold larger seeds to recoup the advantage gained by a smaller-seeded species during seed production ranged from 8.8 weeks for the smallest seeded species in the data set, up to an implausible 4.2 years for the largest-seeded species. Thus, while large-seeded species do have a survival advantage over small-seeded species during seedling establishment, the available evidence suggests that advantages must also accrue during other stages in the life cycle. One possibility is that the greater seed production of small-seeded species (m−2 of canopy outline year−1) is partly offset by larger canopies and longer reproductive life spans in large-seeded species.

title: Which model species for weed seedbank and emergence studies? A review
abstract: Gardarin A, Durr C & Colbach N (2009). Which model species for weed seedbank and emergence studies? A review. Weed Research49, 117–130. 
 
 
 
Summary 
 
Models predicting the effects of cropping systems on weed demography are important tools for testing new rules for integrated weed management that may reduce the use of herbicides and preserve the biodiversity of agro-ecosystems. Such models already exist for a few species and should now be extended to a larger flora, in order to predict and understand the effects of agricultural practices on the evolution of weed communities. This review analysed the literature from 1973 to 2006, focusing on 45 species, to identify past reasons for choosing particular species when modelling the effects of cropping systems on the processes leading to seedling emergence. The frequency or harmfulness of the species were the main reason for studying them. It appears that the studied species were mainly autumn-emerging in north-western Europe cropping systems and summer-emerging in North America; the effects of deep soil tillage were studied mainly in Europe, as simplified sowing techniques are more often practised in North America. A voluminous literature exists on seed persistence in the soil, dormancy, germination and emergence, but rarely with the attempt of establishing generic relationships between species characteristics and model parameters. Until now, such an approach has been mostly developed in ecological studies. Taxa, as well as ecological preferences, seed size and the relationships of these characteristics with weed emergence model parameters should be considered when selecting a range of species for multi-specific modelling purposes.

title: Game-Theoretical Evolution of Seed Mass in Multi-Species Ecological Models
abstract: Within plant communities seed mass often varies over 3 to 5 orders of magnitude, yet simple evolutionary models predict a single optimum seed mass. Here we explore a class of models where seed mass determines 1) the number of seeds produced via a size-number trade-off and 2) competitive ability - plants arising from large seeds are assumed to have a competitive advantage over those derived from small seeds. In this setting the existence of a single-species global ESS seed mass requires the competitive advantage of large seeds over small ones to be unbounded. If there is a limit on the competitive advantage that large seeds obtain then it is always possible to find a smaller seed mass that will successfully invade. In such circumstances there might be a multi-species coevolutionarily stable coalition of several species each with a different seed mass. In this way a wide range of seed masses could be promoted by evolution. In general the adaptive landscape generated by these models is extremely flat leading to slow evolutionary dynamics. The implications of these results for the interpretation of observational, comparative and experimental studies are discussed.

title: Seed size, shape and vertical distribution in the soil: indicators of seed longevity
abstract: 1. We investigated the vertical distribution of seeds in the soil, using data from nine studies in five European countries. We discovered significant correlations between seed shape and distributio ...

title: Seed mortality in the soil is related to seed coat thickness
abstract: Models that quantify the effects of cropping systems on weed dynamics are useful tools for testing innovative cropping systems. In these models, seed mortality in the soil is a key parameter to account for the cumulated effect of cropping systems over time via the soil seed-bank. Since seed mortality is difficult to measure, our objective was to develop a method to estimate it from easily accessible information. Seeds of 13 weed species were buried 30 cm deep in fields and were recovered regularly for 2 years to measure their viability. Seed mass, dimensions, shape, and protein and lipid contents as well as coat thickness were measured. To estimate seed mortality of species not included in the study, we searched for relationships between mortality rates and seed traits. Seed viability mainly decreased during the second year of burial, with mortality rates ranging from 0.01 to 0.63 seeds·seeds− 1·year− 1, depending on the species. Seed mortality decreased with increasing seed coat thickness. No correlation was found with other measured traits or with seed persistence data in the literature. These results were confirmed when the effects of phylogenetic relatedness with phylogenetically independent contrasts were included. The thickness of the seed coat, which varied between 17 and 231 μm over the range of species studied, can protect the seed from external attacks in the soil and slow down seed decay. This trait can be easily measured via X-ray images and could be used to estimate the seed mortality rate for a wider range of species.

title: Seeds Ecology Biogeography And Evolution Of Dormancy And Germination
abstract: Thank you very much for downloading seeds ecology biogeography and evolution of dormancy and germination. As you may know, people have look hundreds times for their favorite novels like this seeds ecology biogeography and evolution of dormancy and germination, but end up in malicious downloads. Rather than reading a good book with a cup of tea in the afternoon, instead they cope with some harmful bugs inside their desktop computer.

title: Hydrophobic trichome layers and epicuticular wax powders in Bromeliaceae.
abstract: The distinctive foliar trichome of Bromeliaceae has promoted the evolution of an epiphytic habit in certain taxa by allowing the shoot to assume a significant role in the uptake of water and mineral nutrients. Despite the profound ecophysiological and taxonomic importance of this epidermal structure, the functions of nonabsorbent trichomes in remaining Bromeliaceae are not fully understood. The hypothesis that light reflection from these trichome layers provides photoprotection was not supported by spectroradiometry and fluorimetry in the present study; the mean reflectance of visible light from trichome layers did not exceed 6.4% on the adaxial surfaces of species representing a range of ecophysiological types nor was significant photoprotection provided by their presence. Several reports suggesting water repellency in some terrestrial Bromeliaceae were investigated. Scanning electron microscopy (SEM) and a new technique-fluorographic dimensional imaging (FDI)-were used to assess the interaction between aqueous droplets and the leaf surfaces of 86 species from 25 genera. In the majority of cases a dense layer of overlapping, stellate or peltate trichomes held water off the leaf epidermis proper. In the case of hydrophobic tank-forming tillandsioideae, a powdery epicuticular wax layer provided water repellency. The irregular architecture of these indumenta resulted in relatively little contact with water droplets. Most mesic terrestrial Pitcairnioideae examined either possessed glabrous leaf blades or hydrophobic layers of confluent trichomes on the abaxial surface. Thus, the present study indicates that an important ancestral function of the foliar trichome in Bromeliaceae was water repellency. The ecophysiological consequences of hydrophobia are discussed.

title: Herbicide deposition on leaf surfaces.
abstract: Leaf surface morphology and physical characteristics of herbicide deposits on leaf surfaces can influence herbicide performance. Leaf surface topography, the degree and type of epicuticular wax formation, and the presence, type, and distribution of trichomes all influence the distribution of a given herbicide formulation sprayed onto a leaf surface. Depressions above anticlinal cell walls accumulate herbicide, thus lessening uniform distribution. As the amount of particulate wax increases, the size of individual spray drop deposits on the leaf decreases, thus resulting in reduced coverage. In many instances the presence of trichomes reduces optimal epidermal coverage by intercepting spray drops before they reach the epidermal surface. Adjuvants reduce the adverse influence of leaf topography, epicuticular wax, and trichomes on herbicide distribution, but their use usually does not yield an even coating over the entire leaf surface. Many herbicides, in pure form, are solids (i.e., crystals) rather than liquids. For most applications, herbicides are dissolved, dispersed, or emulsified in a water-based spray solution. After spraying, water and any solvents evaporate from the leaf surface and herbicides often return to their solid crystalline form. In the few cases that have been studied, less herbicide is absorbed when present on the leaf surface as a solid rather than as a liquid. In many instances, greater effectiveness of a postemergence herbicide may be obtained if attention is given to optimizing the distribution and physical form on sprayed leaf surfaces.

title: Managing arable weeds for biodiversity.
abstract: As a result of the recent intensification of crop production, the abundance and diversity of UK arable weeds adapted to cultivated land have declined, with an associated reduction in farmland birds. A number of questions need to be addressed when considering how these declines can be reversed. Firstly, can the delivery of crop production and biodiversity be reconciled by spatially separating cropping from designated wildlife areas? A number of subsidised environmental schemes in the UK take this approach and are focused on establishing vegetation cover on uncropped land. However, because of the lack of regular disturbance in these habitats, they are dominated by perennials and they therefore have limited potential for promoting the recovery of annual weed populations. A number of farmland bird species also rely on the provision of resources in field centres, and it is therefore likely that the recovery of their populations will rely on weed management options targeted at the cropped areas of the field. This raises two further questions. Firstly, is it possible to identify beneficial weed species that are relatively poor competitors with the crop and also have biodiversity value? Secondly, are the tools available to manage these species at acceptable levels while controlling pernicious weeds? A number of approaches are being employed to answer these questions, including predicting yield loss from weed competition models and exploiting herbicide selectivity. The further development of these tools is crucial if farmer opposition to managing weeds in crops is to be overcome.

title: Towards an assessment of multiple ecosystem processes and services via functional traits
abstract: Managing ecosystems to ensure the provision of multiple ecosystem services is a key challenge for applied ecology. Functional traits are receiving increasing attention as the main ecological attributes by which different organisms and biological communities influence ecosystem services through their effects on underlying ecosystem processes. Here we synthesize concepts and empirical evidence on linkages between functional traits and ecosystem services across different trophic levels. Most of the 247 studies reviewed considered plants and soil invertebrates, but quantitative trait–service associations have been documented for a range of organisms and ecosystems, illustrating the wide applicability of the trait approach. Within each trophic level, specific processes are affected by a combination of traits while particular key traits are simultaneously involved in the control of multiple processes. These multiple associations between traits and ecosystem processes can help to identify predictable trait–service clusters that depend on several trophic levels, such as clusters of traits of plants and soil organisms that underlie nutrient cycling, herbivory, and fodder and fibre production. We propose that the assessment of trait–service clusters will represent a crucial step in ecosystem service monitoring and in balancing the delivery of multiple, and sometimes conflicting, services in ecosystem management.

title: The strength of seeds and their destruction by granivorous insects
abstract: The influence of seed structure and strength on their destruction by granivores is central to understanding the dynamics of granivore-plant interactions. For up to nine seed species, the effects of seed size (cm3), mass (mg), density (mg/cm3) and coat strength (MPa) on the damage inflicted by three post-dispersal granivores (Harpalus pensylvanicus, Anisodactylus sanctaecrucis, and Gryllus pennsylvanicus) were evaluated. Seed destruction rates by G. pennsylvanicus were statistically unrelated to the size and toughness of the seeds. Seed densities significantly affected their destruction by A. sanctaecrucis and H. pensylvanicus, as did seed size, mass, and strength in H. pensylvanicus under choice conditions. The carabid beetles destroyed more of the small, denser seeds with stronger seed coats. The results show that different granivores are able to distinguish the structural strength and physical density of seeds as well as seed size. The relative ability of granivores to detect these seed characteristics offers a way in which diverse communities of post-dispersal insect granivores can persist within a single habitat. The authors redefine how the strength of biological structures should be evaluated in ecological studies, using guidelines commonplace in the field of engineering.