Weed vegetation of arable land in Central Europe: Gradients of diversity and species composition

Question: What are the main broad‐scale spatial and temporal gradients in species composition of arable weed communities and what are their underlying environmental variables? Location: Czech Republic and Slovakia. Methods: A selection of 2653 geographically stratified relevés sampled between 1954–2003 was analysed with direct and indirect ordination, regression analysis and analysis of beta diversity. Results: Major changes in weed species composition were associated with a complex gradient of increasing altitude and precipitation and decreasing temperature and base status of the soils. The proportion of hemicryptophytes increased, therophytes and alien species decreased, species richness increased and beta diversity decreased with increasing altitude. The second most important gradient of weed species composition was associated with seasonal changes, resulting in striking differences between weed communities developed in spring and summer. In summer, weed communities tended to have more neophytes, higher species richness and higher beta diversity. The third gradient reflected long‐term changes in weed vegetation over past decades. The proportion of hemicryptophytes and neophytes increased, while therophytes and archaeophytes decreased, as did species richness over time. The fourth gradient was due to crop plants. Cultures whose management involves less disturbances, such as cereals, harboured less geophytes and neophytes, and had higher species richness but lower beta diversity than frequently disturbed cultures, such as root crops. Conclusions: Species composition of Central European weed vegetation is mainly influenced by broad‐scale climatic and edaphic factors, but its variations due to seasonal dynamics and long‐term changes in agricultural management are also striking. Crop plants and crop‐specific management affect it to a lesser, but still significant extent.     

Long‐term compositional and functional changes in alien and native weed communities in annual and perennial irrigated crops

Agricultural intensification in Europe during the past 30 years has led to changes in compositional and functional weed structure in agroecosystems as well as increases in the prominence of alien weeds. Irrigation is a major driver of agricultural intensification, particularly in semi‐arid zones of the Mediterranean. In the past few decades, irrigated land has expanded in semi‐arid agricultural lands in northeastern Spain. The goals of this study were to identify long‐term temporal changes in compositional and functional weed communities in annual (i.e. maize crops) and perennial (i.e. orchards) irrigated crops of this area and determine whether these changes differentially affect native and alien plants. Changes in the diversity, composition and functional groups of the weed communities in fruit‐tree orchards and maize crops were assessed using plant surveys in 1989 and 2009. During the studied period, a decrease was recorded in the diversity of native species in the fruit‐tree orchards; this decrease was accompanied by an increase in alien weed diversity and a general homogenisation of species in the weed community. In the maize crops, the diversity values of native and alien plants changed little during 20 years. The identification of functional groups revealed that most of the species whose cover increased in the fruit‐tree orchards were graminoid alien species that perform C4 photosynthesis and disperse seed via water or a combination of vectors. In the maize crops, the identified functional groups did not differ in the proportion of species whose cover changed between 1989 and 2009. Hence, in irrigated orchards the observed changes in the weed community and the prominence of alien species are mediated by the selection of a set of traits that let species to overcome management filters. Similarly, the stability of functional composition of weed communities in maize fields is the result of the selection of species functionally similar to the crop.     

Plant diversity,species turnover and shifts in functional traits in coastal dune vegetation: Results from permanent plots over a 52‐year period

Question: What is the relationship between plant diversity and species turnover in coastal dune vegetation plots? How is the long‐term change in species composition of vegetation plots related to shifts in functional traits, and what does it tell us about the dominant processes? Location: Coastal dunes, the Netherlands. Methods: Our data set comprised 52 years of vegetation data from 35 permanent plots in grassland/scrub/woodland vegetation. Vegetation dynamics were described in terms of changes in species composition and abundance, and shifts in 13 functional traits related to resources capture and forage quality, regeneration and dispersal. Results: Species turnover in the plots was high, because of local extinction and colonization. Species‐rich plots were more stable in terms of species abundance and composition compared with species‐poor plots. Over time, the plots converged with respect to their abiotic conditions, as reflected by Ellenberg indicator values – indicating that the prevailing process was succession. The high species turnover reflected high invasibility: accordingly, the relative importance of annuals increased. Most newcomer annuals, however, were competitive generalists of little conservation value. The functional trait analysis allowed us to unravel the complexity of effects of disturbances and succession, and yielded information on the processes driving the observed vegetation dynamics. Conclusions: In this study, small‐scale species turnover was negatively related to species diversity, indicating more stability in species‐rich communities. Regarding shifts in trait diversity, unifying filters appeared to be more dominant than diversifying filters. Counteracting this homogenization process poses a challenge for nature management.     

Global to community scale differences in the prevalence of convergent over divergent leaf trait distributions in plant assemblages

Aim The drivers of species assembly, by limiting the possible range of functional trait values, can lead to either convergent or divergent distributions of traits in realized assemblages. Here, to evaluate the strengths of these species assembly drivers, we partition trait variance across global, regional and community scales. We then test the hypothesis that, from global to community scales, the outcome of co‐occurring trait convergence and divergence is highly variable across biomes and communities. Location Global: nine biomes ranging from subarctic highland to tropical rain forest. Methods We analysed functional trait diversity at progressively finer spatial scales using a global, balanced, hierarchically structured dataset from 9 biomes, 58 communities and 652 species. Analyses were based on two key leaf traits (foliar nitrogen content and specific leaf area) that are known to drive biogeochemical cycling. Results While 35% of the global variance in these traits was between biomes, only 15% was between communities within biomes and as much as 50% occurred within communities. Despite this relatively high within‐community variance in trait values, we found that trait convergence dominated over divergence at both global and regional scales through comparisons of functional trait diversity in regional and community assemblages against random (null) models of species assembly. Main conclusions We demonstrate that the convergence of traits occurring from global to regional assemblages can be twice as strong as that from regional to community assemblages, and argue that large differences in the nature and strength of abiotic and biotic drivers of dominant species assembly can, at least partly, explain the variable outcome of simultaneous trait convergence and divergence across sites. Ultimately, these findings stress the urgent need to extend species assembly research to address those scales where trait variance is the highest, i.e. between biomes and within communities.     

Prediction of species composition of plant communities in a rural landscape based on species traits

The species composition of a community is a subset of the regional species pool, and predicting the species composition of a community from ecological traits of organisms is an important objective in ecology. If such a prediction can be made feasible, we could assess the risk of invasion of locally new species (alien species and genetically modified species) into natural communities. We developed and tested statistical models to predict a community’s species composition from ecological traits of the species pool. Various types of communities (forest, meadow, and weed communities) exist in a small area of traditional rural landscape in Japan, and have been maintained by human activities. These communities and the tracheophytes species pool in the 1-km² research area were considered. We used logistic regression and decision-tree analysis to construct predictive models of community species composition based on plant traits, using the presence or absence of species in a community as the dependent variable and ecological traits as independent variables. Plant traits were grouped by cluster analysis, and the average in each trait group was used for model building to avoid multiple collinearity. Statistical prediction models were significant in all communities. About 60–75% of species composition could be predicted from the measured plant traits in forest communities, but 33–56% in the meadow and weed communities. Our results showed the possibility of predicting the species composition of plant communities from the ecological traits of the plant species together with the information on local species pool.     

A functional trait-based approach to understand community assembly and diversity–productivity relationships over 7 years in experimental grasslands

Several multi-year biodiversity experiments have shown positive species richness–productivity relationships which strengthen over time, but the mechanisms which control productivity are not well understood. We used experimental grasslands (Jena Experiment) with mixtures containing different numbers of species (4, 8, 16 and 60) and plant functional groups (1–4; grasses, legumes, small herbs, tall herbs) to explore patterns of variation in functional trait composition as well as climatic variables as predictors for community biomass production across several years (from 2003 to 2009). Over this time span, high community mean trait values shifted from the dominance of trait values associated with fast growth to trait values suggesting a conservation of growth-related resources and successful reproduction. Increasing between-community convergence in means of several productivity-related traits indicated that environmental filtering and exclusion of competitively weaker species played a role during community assembly. A general trend for increasing functional trait diversity within and convergence among communities suggested niche differentiation through limiting similarity in the longer term and that similar mechanisms operated in communities sown with different diversity. Community biomass production was primarily explained by a few key mean traits (tall growth, large seed mass and leaf nitrogen concentration) and to a smaller extent by functional diversity in nitrogen acquisition strategies, functional richness in multiple traits and functional evenness in light-acquisition traits. Increasing species richness, presence of an exceptionally productive legume species (Onobrychis viciifolia) and climatic variables explained an additional proportion of variation in community biomass. In general, community biomass production decreased through time, but communities with higher functional richness in multiple traits had high productivities over several years. Our results suggest that assembly processes within communities with an artificially maintained species composition maximize functional diversity through niche differentiation and exclusion of weaker competitors, thereby maintaining their potential for high productivity.     

How do weeds differ in their response to the timing of tillage? A study of 61 species across the northeastern United States

Previous research has demonstrated that the season in which soil is tilled (spring versus fall) can strongly influence weed community assembly and subsequent species composition and abundance in annual cropping systems. Despite this understanding, it is unknown whether finer‐scale, within‐season variation in the timing of tillage has similar impacts on weed community assembly. We conducted an experiment on four research farms across the northeastern USA to test the effects of tillage timing on weed emergence periodicity. Soil was tilled at 12 different times that were 2 weeks apart from 29 April to 30 September (the entire growing season) and the composition and abundance of the weed seedlings that emerged was measured 6 weeks later. Weed species clustered into three tillage timing groups at the two New York locations and clustered into five tillage timing groups at the New Hampshire and Maine locations. Individual species associated with each window of tillage time varied by location. No single trait or combination of traits were consistently associated with species‐by‐tillage time groupings across locations; however, within each location several traits were associated with particular groups of species, including: (a) seed length, (b) seed weight, (c) cotyledon type, (d) life span, (e) ploidy level and (f) photosynthetic pathway. These results suggest that fine‐scale variation in the timing of tillage can lead to predictable changes in the species composition and trait distribution of weed communities in annually tilled agroecosystems.     

Traits,neighbors, and species performance in prairie restoration

Questions: Are traits related to the performance of plant species in restoration? Are the relationships between traits and performance consistent across the functional groups of annual forbs, perennial forbs and grasses? Do the relationships between traits and performance depend on neighboring functional groups? Location: A former agricultural field, being restored to native upland prairie, in the Willamette Valley of western Oregon, USA. Methods: Twenty‐eight native species, representing three functional groups, were sown in seven different combinations. Eleven functional traits were measured from plants in the laboratory and in the field. Correlations between individual traits and performance variables were measured and regression techniques used to determine which sets of traits were most strongly related to performance. Results: Sets of traits explained up to 56% of variation in cover, and up to 48% of variation in establishment frequency. The relationships between traits and performance were influenced by functional group identity; the functional group identity of neighboring species also influenced species' cover and the relationships between traits and cover. Species' establishment rate in monoculture was the trait most strongly correlated to both establishment and cover in mixtures. In multi‐trait models, annual forb functional group identity was strongly related to establishment in mixtures, and height, leaf weight ratio at 7 d and seed mass were strongly related to cover. Conclusions: Multiple‐trait models should be a useful way of predicting the performance of species prior to sowing in restoration. The functional group identity of each species and the other species being sown may need to be taken into account when making predictions.     

Traits of butterfly communities change from specialist to generalist characteristics with increasing land-use intensity

Land-use intensification leads to species loss and shifts in community composition, but only few studies examine how these dynamics affect ecological and life-history traits. We thus investigated whether ecological and life-history traits differ between butterfly communities of grasslands with different land-use intensity. We conducted butterfly transect surveys in 137 grassland sites in three regions of Germany and compiled 10 species-specific ecological and life-history traits from the literature. These traits are associated with food plant specialisation, dispersal, distribution, reproduction and development. We calculated a land-use intensity gradient based on the amount of fertilise mowing frequency and grazing intensity. We analysed differences of traits characteristics between butterfly communities along the land-use intensity gradient in a fourth-corner analysis, thus considering correlations between traits. Six ecological and life-history traits changed from characteristics associated with specialists to such associated with generalists with increasing land-use intensity. These traits characteristics in intensified grasslands were: high dispersal propensity, large distribution range, low population density, more than one generation per year, hibernation in a more advanced developmental stage and a long flight period. The functional homogenisation of the butterfly communities with changes from specialist to generalist trait characteristics with increasing land-use intensity may have severe consequences for ecosystem functioning and services.     

Species loss and gain in communities under future climate change: consequences for functional diversity

It is anticipated that anthropogenic climate change will lead to substantial reassembly within communities in coming decades as individual species shift their ranges to track optimal conditions for growth and survival. As species are lost and gained in communities, what are the consequences for functional trait diversity? Functional traits are the characteristics of species that affect individual performance and provide the vital link between biodiversity at the species level and ecosystem function. We investigated how projected changes in species richness in plant communities under climate change scenarios for the decade 2050 will affect the distribution and diversity of five functional traits. We aggregated range change projections made in Maxent for the decade 2050 across all species in the regional pool of littoral rainforest vines in eastern Australia (n = 163 species). The effect of richness changes on trait diversity was assessed in nine rainforest reserves along the east coast of Australia. Although richness was predicted to significantly decline across all communities, functional diversity remained stable, indicating a decoupling in response to climate change at these two different levels of biological organization. A high degree of redundancy in trait composition in communities may buffer against the loss of function in these plant communities. Scaling‐up our understanding of the impact of climate change from the species level to communities is a critical step towards developing conservation strategies aimed at preserving ecosystem function.     

Assessing the Importance of Intraspecific Variability in Dung Beetle Functional Traits

Functional diversity indices are used to facilitate a mechanistic understanding of many theoretical and applied questions in current ecological research. The use of mean trait values in functional indices assumes that traits are robust, in that greater variability exists between than within species. While the assertion of robust traits has been explored in plants, there exists little information on the source and extent of variability in the functional traits of higher trophic level organisms. Here we investigated variability in two functionally relevant dung beetle traits, measured from individuals collected from three primary forest sites containing distinct beetle communities: body mass and back leg length. In doing so we too addressed the following questions: (i) what is the contribution of intra vs. interspecific differences in trait values; (ii) what sample size is needed to provide representative species mean trait values; and (iii) what impact does omission of intraspecific trait information have on the calculation of functional diversity (FD) indices from naturally assembled communities? At the population level, interspecific differences explained the majority of variability in measured traits (between 94% and 96%). In accordance with this, the error associated with calculating FD without inclusion of intraspecific variability was low, less than 20% in all cases. This suggests that complete sampling to capture intraspecific variance in traits is not necessary even when investigating the FD of small and/or naturally formed communities. To gain an accurate estimation of species mean trait values we encourage the measurement of 30–60 individuals and, where possible, these should be taken from specimens collected from the site of study.     

Are traits measured on pot grown plants representative of those in natural communities?

Question: The quantification of functional traits in natural communities can be difficult (e.g. root traits, RGR). Can functional traits measured on pot grown plants be reliably applied to natural communities? Alternatively, can below‐ground plant traits be predicted from above‐ground traits? Location: Southeastern Australia. Methods: We compared 17 shoot, root and whole‐plant morphological traits measured on 14 plant species in a native grassland community to those measured under two different pot conditions: unfertilised and fertilised. Results: The majority of trait values for pot grown plants differed to plants in the field, however, species ranking remained consistent for most leaf traits between the field and the two pot growing conditions. In contrast, species ranking was not consistent for most whole plant traits when comparing field plants to fertilised pot grown plants, providing a caution against the tendency to grow plants in controlled conditions at ‘optimal’ (high) resource levels. Moderate to strong correlations were found between below‐ground and above‐ground plant traits, including between root dry matter content and leaf dry matter content, and between specific root area and specific leaf area. Conclusions: The utility of pot grown plants to quantify traits for field plants is highly dependent on the selection of the growing conditions in the controlled environment. The consistency we observed between above‐ground and below‐ground trait strategies suggests that below‐ground traits may be predictable based on above‐ground traits, reducing the need to quantify root traits on cultured plants.     

The influence of fire on phylogenetic and functional structure of woody savannas: Moving from species to individuals

Fire is a key determinant of tropical savanna structure and functioning. High fire frequencies are expected to assemble closely related species with a restricted range of functional trait values. Here we determined the effect of fire on phylogenetic and functional diversity of woody species and individuals in savanna communities under different fire frequencies. We found phylogenetic signals for one third of the functional traits studied. High numbers of fires simultaneously led to phylogenetic overdispersion and functional clustering when communities were represented by mean trait values with all traits that putatively should be affected or respond to fire. This finding is important, because it shows that the relationship between ecological processes and the phylogenetic structure of communities is not straightforward. Thus, we cannot always assume that close relatives are more similar in their ecological features. However, when considering a different set of traits representing different plant strategies (fire resistance/avoidance, physiological traits and regeneration traits), the results were not always congruent. When asking how communities are assembled in terms of individuals (not species) the outcome was different from the species-based approach, suggesting that the realised trait values – rather than mean species trait values – have an important role in driving community assembly. Thus, intraspecific trait variability should be taken into account if we want fully to improve our mechanistic understanding of assembly rules in plant communities.     

Weed species differ in their ability to emerge in no‐till systems that include cover crops

No‐till cropping systems that include cover crops could lead to important changes in weed communities by decreasing some annual weed populations. In this study, we predicted that seed burial depth and the presence of cover crop would affect the emergence and initial growth success of annual weed species. We tested two factors on 14 weed species in a greenhouse: the seed burial depth of weeds (buried versus soil surface) and the presence/absence of a cover crop (ryegrass). We counted the emerged seedlings and measured the height of weeds and cover crops (Hweed, Hcover), the dry matter content of weeds and cover crops (DMCweed, DMCcover) and the number of leaves of weeds (NLweed) on 1433 weed and 390 ryegrass individuals. Emergence of five weed species (AMBEL, ANGAR, BROST, CENCY and EPHHE) was affected by the seed location (?10.3% on average for unburied seeds), five other weed species (ALOMY, CAPBP, SONAS, VERPE and VLPMY) were affected by cover (on average ?9.5% for seeds emerged in the presence of cover crop), and four weed species (GERDI, LAMPU, POAAN and VIOAR) were not affected by either. Weed growth of all weed species also decreased with the presence of a cover crop (on average Hweed: ?49.9%, DMCweed: ?87.2% and NLweed: ?55.4%) and for unburied seeds (on average Hweed: ?33.7%, DMCweed: ?70.6% and NLweed: ?43.3%), with various responses according to species. This study indicates that annual weeds could be disadvantaged by no‐till systems using cover crops.     

Filling the gap in functional trait databases: use of ecological hypotheses to replace missing data

Functional trait databases are powerful tools in ecology, though most of them contain large amounts of missing values. The goal of this study was to test the effect of imputation methods on the evaluation of trait values at species level and on the subsequent calculation of functional diversity indices at community level using functional trait databases. Two simple imputation methods (average and median), two methods based on ecological hypotheses, and one multiple imputation method were tested using a large plant trait database, together with the influence of the percentage of missing data and differences between functional traits. At community level, the complete‐case approach and three functional diversity indices calculated from grassland plant communities were included. At the species level, one of the methods based on ecological hypothesis was for all traits more accurate than imputation with average or median values, but the multiple imputation method was superior for most of the traits. The method based on functional proximity between species was the best method for traits with an unbalanced distribution, while the method based on the existence of relationships between traits was the best for traits with a balanced distribution. The ranking of the grassland communities for their functional diversity indices was not robust with the complete‐case approach, even for low percentages of missing data. With the imputation methods based on ecological hypotheses, functional diversity indices could be computed with a maximum of 30% of missing data, without affecting the ranking between grassland communities. The multiple imputation method performed well, but not better than single imputation based on ecological hypothesis and adapted to the distribution of the trait values for the functional identity and range of the communities. Ecological studies using functional trait databases have to deal with missing data using imputation methods corresponding to their specific needs and making the most out of the information available in the databases. Within this framework, this study indicates the possibilities and limits of single imputation methods based on ecological hypothesis and concludes that they could be useful when studying the ranking of communities for their functional diversity indices.     

Limiting similarity and Darwin’s naturalization hypothesis: understanding the drivers of biotic resistance against invasive plant species

Several hypotheses have been proposed to explain biotic resistance of a recipient plant community based on reduced niche opportunities for invasive alien plant species. The limiting similarity hypothesis predicts that invasive species are less likely to establish in communities of species holding similar functional traits. Likewise, Darwin’s naturalization hypothesis states that invasive species closely related to the native community would be less successful. We tested both using the invasive alien Ambrosia artemisiifolia L. and Solidago gigantea Aiton, and grassland species used for ecological restoration in central Europe. We classified all plant species into groups based on functional traits obtained from trait databases and calculated the phylogenetic distance among them. In a greenhouse experiment, we submitted the two invasive species at two propagule pressures to competition with communities of ten native species from the same functional group. In another experiment, they were submitted to pairwise competition with native species selected from each functional group. At the community level, highest suppression for both invasive species was observed at low propagule pressure and not explained by similarity in functional traits. Moreover, suppression decreased asymptotically with increasing phylogenetic distance to species of the native community. When submitted to pairwise competition, suppression for both invasive species was also better explained by phylogenetic distance. Overall, our results support Darwin’s naturalization hypothesis but not the limiting similarity hypothesis based on the selected traits. Biotic resistance of native communities against invasive species at an early stage of establishment is enhanced by competitive traits and phylogenetic relatedness.     

Evidence of neotropical anuran community disruption on rice crops: a multidimensional evaluation

Agricultural expansion is a major driver of biodiversity loss, especially in the megadiverse tropics. Rice is among the world’s most important food crops, invariably affecting biodiversity worldwide. Although the effects of habitat conversion to rice crops on biodiversity are not completely understood, landscape modification often creates conditions that benefit some species and excludes others. We conducted an integrative evaluation of the effects that habitat conversion to irrigated rice crops has on anuran communities from a Cerrado-Amazon ecotone. We adopted a multidimensional approach to compare anuran communities from agricultural and pristine environments considering (i) taxonomic metrics; (ii) functional and phylogenetic diversity; (iii) selected and excluded traits and (iv) body condition indices. When compared to their pristine counterparts, agricultural waterbodies showed increased functional divergence and decreased species diversity and functional richness. Furthermore, agricultural anuran communities exhibited lower phylogenetic diversity. Nonetheless, taxonomic diversity did not vary significantly, suggesting that it should not be used without complementary metrics. Species with small range, habitat specialization, small clutches and large body size were excluded from rice crops. Furthermore, frogs showed lower body condition in crops than in pristine areas. Understanding how species traits correlate with specific responses to agriculture will allow better predictions of the functional effects of anthropogenic land-use. Maintaining high diversity in anthropogenic environments is important for ecosystem resilience because diverse communities are more likely to hold multiple species capable of contributing to ecological functions. Our results show that converting natural vegetation to irrigated rice crops drives many species to local extinction, and resilient species to exhibit lower body condition.     

Diversity and phenology of weed communities in the Nile Delta region

This study evaluates the species diversity and phe-nological behaviour of the weed communities associated with the common crops in the Nile Delta region (three winter crops: Egyptian clover, broad beans and wheat; and three summer crops: cotton, maize and rice). The weed communities of winter crops are of higher diversity than those of summer crops, presumably due to differences in sowing and weeding processes. The phenological sequences of the weed communities are, in most cases, closely related to those of the crops in which they occur. The correspondence between the crucial periods in the life cycle of the weed and the crop (seedling establishment and fruiting) often leads to severe competition.