How does agricultural intensification modulate changes in plant community composition?

Agricultural intensification, at local and landscape scales, has caused a decrease in plant diversity and changes in species composition in cereal fields. To better understand the role of landscape complexity and farming systems in shaping plant assemblages, it is of interest to focus on functional traits rather than on floristic composition, which may help to highlight trends in vegetation patterns. We analysed the relative abundance of various functional attributes (different life forms, growth forms, wind-pollinated species and wind-dispersed species) at three contrasted field positions (boundary, edge and centre) of 29 organic and 29 conventional cereal fields distributed in 15 agrarian localities of NE Spain. Agricultural intensification affected the biological attributes of the vegetation in dryland Mediterranean cereal fields; local factors (farming system and position) had a more prominent role in affecting plant functional composition than the surrounding landscape. Local factors were important for life form distribution, growth form and pollination type, whereas landscape complexity mainly affected the proportion of wind-dispersed species. Therefore, depending on the objective of the study, it is important to select functional attributes sensitive to the different scales of agricultural intensification, especially because landscape complexity and land-use intensity are commonly related.     

Organic versus conventional arable farming systems: Functional grouping helps understand staphylinid response

The response of different feeding groups of staphylinid beetles to organic management, distance to perennial boundary and landscape context was studied in 42 paired organic and conventional winter wheat fields. Management effects were found to strongly depend on feeding group. While the activity-density of predators was higher in the conventional fields, both activity-density and species richness of detritivores were higher in the organic fields. Activity-density and species richness of detritivores were positively correlated to crop yield in the conventional but not in the organic fields. Unexpectedly, species richness as high as in the less productive organic fields was thus found in intensified conventional systems. No significant effects of landscape context could be found on activity-density or species richness of the different feeding groups. More species were caught near the field edge than in the middle, showing the importance of spill-over from field boundaries into arable crops for diversity. In conclusion, separation of species into feeding groups revealed patterns that have not been shown before. Decomposer diversity (but not that of predators and fungivores) was higher in organic fields, but reached similar levels in high-yielding conventional fields. Thus resource quality (purely organic or organic/synthetic fertiliser) and quantity appear to play a major role for this functional group.     

Plant species in arable field margins and road verges of central Norway

Plant species composition and species number were studied in two types of field margins: 31 arable field margins and 33 road verges. Both field margin types were adjacent to intensively managed grass or cereal fields. Effects of eight variables on field margin vegetation were studied. Despite having many plant species in common, composition and species number differed between the two field margin types, due to different ecological conditions and margin management. Arable field margins were composed of tall and/or nutrient demanding ubiquitous species and were characterised by species of later successional stages than those of road verges. The vegetation of road verges was lower and species numbers were higher than in arable field margins. The road verges contained several semi-natural meadow species that are declining in abundance. A CCA ordination of the field margins and the recorded variables showed that plant species composition was significantly affected by thickness of litter, slope, width, moisture level and type of crop in the adjacent field. Number of plant species was significantly higher in mown than unmown margins and generally higher in margins adjacent to fields with mainly grass production than in margins adjacent to fields with mainly cereal crops. In order to preserve botanical diversity in this agricultural landscape, the maintenance of regularly mown road verges should be prioritized.     

Weed and arthropod populations in conventional and genetically modified herbicide tolerant fodder beet fields

The introduction of genetically modified herbicide tolerant (GMHT) crops has raised concerns from both scientists and non-governmental organisations about possible effects on arable flora and fauna due to the changes in herbicide application and management that such crops involve. Three consecutive studies were performed, covering flora and fauna in fields of GMHT and conventional fodder beets over the season, at different locations and under different spraying regimes. At all locations and in the 3 years, a denser and more diverse weed flora and arthropod fauna were found in GMHT beets in early and mid-summer than in conventional beets when glyphosate-treatment occurred at or after label recommendation. Following the herbicide applications the GMHT fields had fewer weed species and seeds and lower weed densities and biomass than conventional fields. However, application of glyphosate earlier than recommended resulted in an extremely low weed diversity, density and biomass during the entire season. Timing of the first glyphosate applications, i.e. the duration of the herbicide free period, was essential in terms of biodiversity improvements. In the long term reduced production of weed seeds in GMHT fields may deplete the weed flora if the GMHT strategy becomes widely adopted.     

Farming systems and insect predators

Predatory carabids, staphylinids and spiders were monitored from 1981 to 1987 in a Dutch project concerning the development of arable farming systems. During this period epigeic predators in the conventional, integrated and organic systems were sampled using pitfall traps. The effects of crop type and farming system on species diversity, abundance and guild structure was analysed using trap data from wheat, pea, sugar beet, potato, onion and carrot fields. Predator abundance and species composition were found to be clearly affected by the farming system. However, in most cases the type of crop appeared to be of greater importance. Crops with a greater cover early in the season, like winter wheat and peas appeared to be more favourable than late and open crops like onions or carrots. Both system and crop effects were more apparent in carabids than in spiders or staphylinids. Only minor effects of farm management were found on species richness. Again, the crop itself seemed to be the main structuring factor. It is concluded that the presence and quality of predator populations is mainly determined by crop structure and crop-related factors. The role of favourable crops and field size in predator enhancement is discussed in relation to the agroecological infrastructure of the landscape.     

Epigeic Collembola in winter wheat under organic,integrated and conventional farm management regimes

Community characteristics of Collembola assemblages in conventional, integrated and organic fields of winter wheat were compared among three randomly chosen areas in England using analysis of similarities, cluster analysis, multi-dimensional scaling and several measures of diversity and evenness. Indicator values were used to identify indicator species. Significant differences were found in the abundance of most species and in community structure among the three geographical regions but few differences between the farming regimes were significant. Despite a lack of significant differences among regimes, Entomobrya multifasciata and Isotomurus spp. were consistently, although not significantly more common in conventional than organic fields whereas the opposite was true for Isotoma viridis and Isotoma notabilis. Farming regime significantly affected the abundance of Sminthurinus elegans and Sminthurus viridis but the effect differed between geographical regions. Community composition and species dominance were influenced by farming regime, but no species were indicative of the different farming systems, as most occurred ubiquitously in all fields. Organically and conventionally farmed fields were found not to differ significantly from each other in community composition, but both differed from integrated fields. These findings are compared with the results from other recent European studies of the effects of farming systems on arthropods and their wider ecological implications are discussed.     

Predominance of self-compatibility in hummingbird-pollinated plants in the Neotropics

Both plant traits and plant–pollinator interactions are thought to influence plant mating systems. For hummingbird-pollinated plants, foraging strategy (territorial or traplining) is also expected to influence plant mating. We hypothesize that the traplining behavior of hermits promotes outcrossing, whereas the behavior of non-hermits favours self-incompatibility. Thus, selection is expected to maintain self-incompatibility in plants pollinated by non-hermits. We explore the incidence of self-incompatibility in Neotropical hummingbird-pollinated plants and its association with hummingbird behavior and plant traits. We conducted a literature review (56 species) and performed hand-pollination experiments in 27 hummingbird-pollinated plants in an Atlantic rainforest. We found that self-incompatibility (measured as <0.3 for the Index of Self-incompatibility [ISI]) occurred in only 33 % of the Neotropical hummingbird-pollinated plants. The interaction of hummingbird and habit type affected ISI, as did phylogenetic relationships. Specifically, herbs pollinated by non-hermits had higher ISI than woody plants pollinated by non-hermits, and herbs pollinated by both hermits and non-hermits. For the Atlantic rainforest plant guild, 30 % of the species were self-incompatible. ISI was higher in herbs than in woody species and increased with plant aggregation but was not dependent on foraging behavior, plant density, or floral display. Although hummingbirds differ in their foraging strategies, these behavioral differences seem to have only a minor influence on the incidence of self-incompatibility. Phylogenetic relatedness seems to be the strongest determinant of mating system in Neotropical hummingbird-pollinated plants.     

Bird pollination of Canary Island endemic plants

The Canary Islands are home to a guild of endemic, threatened bird-pollinated plants. Previous work has suggested that these plants evolved floral traits as adaptations to pollination by flower specialist sunbirds, but subsequently, they appear to have co-opted generalist passerine birds as sub-optimal pollinators. To test this idea, we carried out a quantitative study of the pollination biology of three of the bird-pollinated plants, Canarina canariensis (Campanulaceae), Isoplexis canariensis (Veronicaceae) and Lotus berthelotii (Fabaceae), on the island of Tenerife. Using colour vision models, we predicted the detectability of flowers to bird and bee pollinators. We measured pollinator visitation rates, nectar standing crops as well as seed-set and pollen removal and deposition. These data showed that the plants are effectively pollinated by non-flower specialist passerine birds that only occasionally visit flowers. The large nectar standing crops and extended flower longevities (>10 days) of Canarina and Isoplexis suggests that they have evolved a bird pollination system that effectively exploits these low frequency non-specialist pollen vectors and is in no way sub-optimal. Seed set in two of the three species was high and was significantly reduced or zero in flowers where pollinator access was restricted. In L. berthelotii, however, no fruit set was observed, probably because the plants were self-incompatible horticultural clones of a single genet. We also show that, while all three species are easily detectable for birds, the orange Canarina and the red Lotus (but less so the yellow-orange Isoplexis) should be difficult to detect for insect pollinators without specialised red receptors, such as bumblebees. Contrary to expectations if we accept that the flowers are primarily adapted to sunbird pollination, the chiffchaff (Phylloscopus canariensis) was an effective pollinator of these species.     

Influence of hedgerow and grassy field borders on ground beetle (Coleoptera: Carabidae) activity in fields of corn

Agricultural landscapes may be manipulated in ways that benefit predatory invertebrates by providing alternate food sources, overwintering sites, and refuge from farming activities. Ecological theory predicts that complex plant communities should support a richer community of natural enemies of pest insects than a simple plant community. A study was conducted in Iowa, USA to investigate the influence of the vegetative diversity of field boarders on the activity, species richness, and community similarities of predatory beetles occurring in corn fields.Ground beetle (Coleoptera: Carabidae) populations were compared among corn fields bounded by either complex hedges or simple grass edges. Directional pitfall traps were used to investigate activity patterns of beetles between border types and their adjacent corn fields. Beetles were trapped during four seasonal periods, based on the growth stage of corn. During corn emergence (May–June) when fields are barren, carabids were more active and species richness was higher in the corn fields bordered by woody hedges. The carabid species Scarites quadriceps, Scarites subterraneus, and Harpalus pensylvanicus, were more dominant in hedge sites as compared to grass sites at this time. Following corn–canopy closure, carabids were now more active in fields bordered by grassy edges, but beetle activity also remained high in the fields adjacent to woody hedges. Further analysis of the carabid communities by Bray–Curtis Similarity Index showed no difference among field edge types at any time of the season. Results indicate that both complex and simple field border habitats support abundant and diverse populations of carabids during most of the growing season. However, during the early growing season hedges appear to be more important than grass edges in supplying carabid beetles to corn fields. Woody hedges may serve as very important overwintering sites and as an early season refuge for predatory beetles in corn.     

Set-aside management: How do succession, sowing patterns and landscape context affect biodiversity?

European Union (EU) member states set aside between 5 and 15% of arable land during the last two decades, but abolition of the set-aside scheme in 2008 caused a sudden loss in habitat availability and biodiversity in agricultural landscapes. Management of set-aside has many facets and in this perspective paper we focus on the biodiversity effects of successional age, sowing strategies and landscape context. Young, 1-2-year-old set-asides have been initially considered to be too ephemeral to have any conservation value. However, when a rich seed and bud bank is available, a species-rich natural (secondary) succession can be observed. Arable (annual) weed communities in the first two years of succession can even include endangered plant species with associated rare insect consumers. Furthermore, many bird species benefit from early-successional habitats, whereas small mammal communities are richer in older habitats. If the local plant species pool is poor, sowings of diverse mixtures from regional seed collections can be recommended. Set-aside managers using species-rich sowings often experience that dominant weeds suppress the less competitive annual species. This trend to species-poor communities can be avoided by intraspecific aggregation of competitively weak species. Broadening the spatial scale from the plot to the landscape, efficiency of set-aside is highest in simple landscapes, where set-aside exhibits greatest effect in enhancement of biodiversity and associated services such as pollination and biological control. In complex landscapes, however, additional set-aside does not add much to the high level of biodiversity and ecological processes already present. Twenty percent of semi-natural, non-crop habitat appears to be a rough threshold for enhancing biodiversity and sustaining services such as pollination and biological control, but improved set-aside management should have the potential to reduce the percentage of semi-natural non-crop habitat needed. EU policy should tailor set-aside schemes for the maintenance of biodiversity and also consider that management efficiency is higher in simple than complex landscapes.     

Host pollination mode and mutualist pollinator presence: net effect of internally ovipositing parasite in the fig–wasp mutualism

The Ficus–their specific pollinating fig wasps (Chalcidoidea, Agaonidae) interaction presents a striking example of mutualism. Figs also shelter numerous non-pollinating fig wasps (NPFW) that exploit the fig–pollinator mutualism. Only a few NPFW species can enter figs to oviposit, they do not belong to the pollinating lineage Agaonidae. The internally ovipositing non-agaonid fig wasps can efficiently pollinate the Ficus species that were passively pollinated. However, there is no study to focus on the net effect of these internally ovipositing non-agaonid wasps in actively pollinated Ficus species. By collecting the data of fig wasp community and conducting controlled experiments, our results showed that internally ovipositing Diaziella bizarrea cannot effectively pollinate Ficus glaberrima, an actively pollinated monoecious fig tree. Furthermore, D. bizarrea failed to reproduce if they were introduced into figs without Eupristina sp., the regular pollinator, as all the figs aborted. Furthermore, although D. bizarrea had no effect on seed production in shared figs, it significantly reduced the number of Eupristina sp. progeny emerging from them. Thus, our experimental evidence shows that reproduction in Diaziella depends on the presence of agaonid pollinators, and whether internally ovipositing parasites can act as pollinators depends on the host fig’s pollination mode (active or passive). Overall, this study and others suggest a relatively limited mutualistic role for internally ovipositing fig wasps from non-pollinator (non-Agaonidae) lineages.     

On the success of a swindle: pollination by deception in orchids

A standing enigma in pollination ecology is the evolution of pollinator attraction without offering reward in about one third of all orchid species. Here I review concepts of pollination by deception, and in particular recent findings in the pollination syndromes of food deception and sexual deception in orchids. Deceptive orchids mimic floral signals of rewarding plants (food deception) or mating signals of receptive females (sexual deception) to attract pollen vectors. In some food deceptive orchids, similarities in the spectral reflectance visible to the pollinator in a model plant and its mimic, and increased reproductive success of the mimic in the presence of the model have been demonstrated. Other species do not mimic specific model plants but attract pollinators with general attractive floral signals. In sexually deceptive orchids, floral odor is the key trait for pollinator attraction, and behaviorally active compounds in the orchids are identical to the sex pheromone of the pollinator species. Deceptive orchids often show high variability in floral signals, which may be maintained by negative frequency-dependent selection, since pollinators can learn and subsequently avoid common deceptive morphs more quickly than rare ones. The evolution of obligate deception in orchids seems paradoxical in the light of the typically lower fruit set than in rewarding species. Pollination by deception, however, can reduce self-pollination and encourage pollen flow over longer distances, thus promoting outbreeding. Although some food deceptive orchids are isolated through postzygotic reproductive barriers, sexually deceptive orchids lack post-mating barriers and species isolation is achieved via specific pollinator attraction. Recent population genetic and phylogenetic investigations suggest gene-flow within subgeneric clades, but pollinator-mediated selection may maintain species-specific floral traits.     

Breeding skylarks (Alauda arvensis) on organic and conventional arable farms in The Netherlands

The aim of this study was to analyse the effects of differences in cropping pattern between organic and conventional arable farms on the breeding activity of skylarks and to assess the effects of arable crop management on skylark nest survival. Skylark nest density was seven times higher on organic farms than on conventional farms (0.63 vs. 0.09 nest per 10 ha). Skylarks showed a strong preference for spring cereals, lucerne and grass leys, all of which were mainly or exclusively grown on organic farms. On organic farms nests were initiated during the entire breeding season, but on conventional farms no nesting activity was found during the peak of the season (early May to early June). On organic farms 27% of all nests was successful. Increasing the availability of suitable breeding habitat during the peak of the breeding season on conventional farms might provide one means of enhancing breeding skylark populations. On organic farms, crop management should focus on reducing nest loss due to farming operations.     

Indirect interactions between invasive and native plants via pollinators

In generalised pollination systems, the presence of alien plant species may change the foraging behaviour of pollinators on native plant species, which could result in reduced reproductive success of native plant species. We tested this idea of indirect interactions on a small spatial and temporal scale in a field study in Mauritius, where the invasive strawberry guava, Psidium cattleianum, provides additional floral resources for insect pollinators. We predicted that the presence of flowering guava would indirectly and negatively affect the reproductive success of the endemic plant Bertiera zaluzania, which has similar flowers, by diverting shared pollinators. We removed P. cattleianum flowers within a 5-m radius from around half the B. zaluzania target plants (treatment) and left P. cattleianum flowers intact around the other half (control). By far, the most abundant and shared pollinator was the introduced honey bee, Apis mellifera, but its visitation rates to treatment and control plants were similar. Likewise, fruit and seed set and fruit size and weight of B. zaluzania were not influenced by the presence of P. cattleianum flowers. Although other studies have shown small-scale effects of alien plant species on neighbouring natives, we found no evidence for such negative indirect interactions in our system. The dominance of introduced, established A. mellifera indicates their replacement of native insect flower visitors and their function as pollinators of native plant species. However, the pollination effectiveness of A. mellifera in comparison to native pollinators is unknown. Christine B. Müller, deceased 7 March 2008.     

Influence of habitat type and surrounding landscape on spider diversity in Swedish agroecosystems

Lycosid and linyphiid spiders were collected over a full cropping season around Uppsala, Sweden, in eight organic spring sown cereal fields in three different habitat types: field margin, field, and the edge between the two. The sites were located in landscapes with different proportions of non-crops, forest, perennial crops, annual crops, and number and sizes of arable fields. The field margin, compared with the field habitat, was found to be important for the activity density of lycosids, and for the species richness and composition of linyphiids. Landscape parameters were central for the activity density of linyphiids and for the species richness and composition of Lycosidae. A diverse landscape with easy access to perennial crops and forest in addition to field margins will augment both the number of species and individuals of the two spider families.     

Conspecific flowers of <Emphasis Type="Italic">Sinapis arvensis</Emphasis> are stronger competitors for pollinators than those of the invasive weed <Emphasis Type="Italic">Bunias orientalis</Emphasis>

Biological invasions can affect the structure and function of ecosystems and threaten native plant species. Since most weeds rely on mutualistic relationships in their new environment, they may act as new competitors for pollinators. Pollinator competition is likely to be density dependent, but it is often difficult to disentangle competition caused by flower quality from effects caused by flower quantity. In order to test the effects of the presence and number of flowers of the invasive weed Bunias orientalis on the insect visitation rates in a native species (Sinapis arvensis), we performed two replacement experiments using plants with standardised flower numbers. The visitation rates in S. arvensis were significantly higher than in B. orientalis and the number of insect visits dropped significantly with increasing density of S. arvensis flowers. These results suggest that intraspecific competition among flowers of S. arvensis is stronger than the competitive effect of alien flowers. As flowers of B. orientalis do not seem to distract visitors from S. arvensis, it is unlikely that pollinator competition between these two plant species plays a crucial role. However, it cannot be excluded that mass blossom stands of B. orientalis may distract flower visitors from native species.     

Long-term changes in soil fertility in organic arable farming systems in England,with particular reference to phosphorus and potassium

Organic farming has undergone significant expansion in Europe over the last decade and it is often seen as a sustainable alternative to intensive agricultural systems. If it is to be truly sustainable, it must maintain levels of soil fertility sufficient for economic crop production in the long-term, whilst also protecting the environment. This paper presents results comparing soils managed organically for at least 15 years, with soils under conventional management, on four arable farms in England. There were no significant differences in total soil organic matter, total nitrogen or C:N ratio between the conventionally and organically managed soils. However, concentrations of extractable potassium and phosphorus were significantly lower in soils managed organically. The largest difference between the conventional and organic fields in potassium concentration was on the oldest organic farm. These results support the argument that organic mixed arable systems are mining reserves of potassium and phosphorus, built up during conventional management, and that changes to organic management practices to increase inputs of potassium and phosphorus are required, if long-term declines in soil fertility, and thus, yields are to be avoided.     

Yield loss in conventional and natural rice farming systems

Compared to modern, conventional agriculture, alternative agricultural production systems may rely on biologically different mechanisms (syndromes) to attain similar production goals. Yield loss to rice in conventional and natural farming rice paddies in Japan was evaluated by simulated injury (leaf-clipping) and monitoring plants damaged by insect herbivores. Rice grown under natural farming practices was more tolerant of simulated injury and injury from Oulema oryzae than rice grown under conventional practices. Natural farming rice retained proportionately more tillers and had a higher proportion of mature seeds than conventionally grown rice. In conventional paddies, the simulated injury may have made the rice plants more susceptible to plant pathogens than their non-injured counterparts, resulting in higher disease attack and proportionately greater yield loss. These results suggest that, pests may affect yield loss independently in natural farming, but in conventional paddies, multiple pest injury may interact synergistically, compounding yield loss.     

Effects of organic and soil conservation management on specialist bird species

Bird abundance was assessed on a total of 58 farms across the Seine-et-Marne department, France (12 organic, 19 conservation-tillage and 27 conventional farms). Local abundance variations among the three farming systems were related to two species traits, i.e. habitat specialisation and diet, considering both farmland and non-farmland species. It was found that organic farming favoured specialist birds, either considering the whole community or non-farmland birds only. On the opposite, specialist farmland species were found to be less abundant in conservation-tillage farms than in conventional ones. Invertebrate-feeders were found to benefit from conservation-tillage practices compared to omnivorous species but not compared to granivorous ones; an interaction between species diet and the species specialisation level was also found. Granivorous species tended to increase with the conservation-tillage duration and in particular specialist birds.