Testing solutions in grass-dominated landscapes: a review of current research

In this paper we review the experimental development of agri-environment measures for use on grasslands. Sward structure has been shown to have a strong influence on birds' ability to forage in grasslands, but the effects of food abundance on foraging behaviour are poorly understood and this hinders development of grassland conservation measures. The experiments described have a dual purpose: to investigate the foraging ecology of birds on grasslands and to test candidate management measures. Most of the work featured focuses on increasing invertebrate food resources during the summer by increasing habitat heterogeneity. We also identify important gaps in the habitats provided by existing or experimental measures, where similar dual-purpose experiments are required.

The influence of livestock management on habitat quality for farmland birds

This review covers research linking foraging habitat quality for birds to livestock management in lowland farmland. Based on this research we propose a framework for predicting the value of grazing systems to birds. This predictive framework is needed to guide the development of agri-environment measures to address farmland bird declines in pastoral areas. We show that the exacting requirements of declining granivorous birds pose the greatest challenges, while the needs of soil invertebrate feeding species are more easily met.

Leaving final-cut grass silage in situ overwinter as a seed resource for declining farmland birds

The loss of seed-rich wintering habitats has been a major contributory cause of farmland bird population declines in western Europe. Agricultural grasslands are particularly poor winter foraging habitats for granivorous birds, which have declined most in the pastoral farming regions of western Britain. We describe an experiment to test the utility of fertile ryegrass (Lolium) swards as a potentially rich source of winter seed for declining farmland birds. Four patches of final-cut grass silage were allowed to set seed and were left in situ overwinter. Half of each patch was lightly aftermath grazed in an attempt to increase the accessibility of the seed to foraging birds and reduce the perceived predation risk. Large numbers of yellowhammers (Emberiza citrinella) and reed buntings (E. schoeniclus) foraged on the seeded plots throughout the winter. They preferred to forage on ungrazed seeded plots, where the accumulation of senescent foliage resulted in a 14% average loss in silage yield in the following season. However, seed produced on the plots also led to sward regeneration, increasing subsequent yields on some plots. The technique offers clear benefits as a potential future agri-environment measure for declining granivorous birds, with wide applicability, but requires further development to minimise sward damage and costs to the farmer. Autumn grazing should reduce sward damage, but at the cost of reduced usage by buntings. Using the technique just prior to reseeding would be one way of avoiding any costs of sward damage.

The roles of olfaction and vision in host-plant finding by the diamondback moth, Plutella xylostella

The relative roles of olfaction and vision in the crepuscular host-finding process of a major lepidopteran pest of cruciferous crops, the diamondback moth Plutella xylostella are investigated in a series of laboratory and semi-field experiments. Flying female moths use volatile plant chemical cues to locate and to promote landing on their host, even in complex mixed-crop environments in large cages. Multiple regression analysis shows that both the plant position (front, middle or back rows) and the type of plant (host plant, nonhost plant) are needed to explain the distribution of insects in such a mixed-crop situation. This strong plant position effect indicates that, when host plants are present in a mixture, foraging P. xylostella are more likely to alight on the first row of the plants. The findings are discussed with regard to current theories of host-plant location by phytophagous insects and the possible implications for integrated pest management.

Very short delays prior to escape from potential predators may function efficiently as adaptive risk-assessment periods

Periods between predator detection and an escape response (escape delays) by prey upon attack by a predator often arise because animals trade-off the benefits such a delay gives for assessing risk accurately with the costs of not escaping as quickly as possible. We tested whether freezing behaviour (complete immobility in a previously foraging bird) observed in chaffinches before escaping from an approaching potential threat functions as a period of risk-assessment, and whether information on predator identity is gained even when time available is very short. We flew either a model of a sparrowhawk (predator) or a woodpigeon (no threat) at single chaffinches. Escape delays were significantly shorter with the hawk, except when a model first appeared close to the chaffinch. Chaffinches were significantly more vigilant when they resumed feeding after exposure to the sparrowhawk compared to the woodpigeon showing that they were able to distinguish between threats, and this applied even when time available for assessment was short (an average of 0.29 s). Our results show freezing in chaffinches functions as an effective economic risk assessment period, and that threat information is gained even when very short periods of time are available during an attack.

Ecological specificity of arbuscular mycorrhizae: evidence from foliar- and seed-feeding insects

Arbuscular mycorrhizal (AM) fungi have a variety of effects on foliar-feeding insects, with the majority of these being positive, although reports of negative and null effects also exist. Virtually all previous experiments have used mobile insects confined in cages and have studied the effects of one, or at most two, species of mycorrhizae on one species of insect. The purpose of this study was to introduce a greater level of realism into insect-mycorrhizal experiments, by studying the responses of different insect feeding guilds to a variety of AM fungi. We conducted two experiments involving three species of relatively immobile insects (a leaf-mining and two seed-feeding flies) reared in natural conditions on a host (Leucanthemum vulgare). In a field study, natural levels of AM colonization were reduced, while in a phytometer trial, we experimentally colonized host plants with all possible combinations of three known mycorrhizal associates of L. vulgare. In general, AM fungi increased the stature (height and leaf number) and nitrogen content of plants. However, these effects changed through the season and were,dependent on the identity of the fungi in the root system. AM fungi increased host acceptance of all three insects and larval performance of the leaf miner, but these effects were also season- and AM species-dependent. We suggest that the mycorrhizal effect on the performance of the leaf miner is due to fungal-induced changes in host-plant nitrogen content, detected by the adult fly. However, variability in the effect was apparent, because not all AM species increased plant N content. Meanwhile, positive effects of mycorrhizae were found on flower number and flower size, and these appeared to result in enhanced infestation levels by the seed-feeding insects. The results show that AM fungi exhibit ecological specificity, in that different. species have different effects on host-plant growth and chemistry and the performance of foliar-feeding insects. Future studies need to conduct experiments that use ecologically realistic combinations of plants and fungi and allow insects to be reared in natural conditions.

Pollination and other ecosystem services produced by mobile organisms: a conceptual framework for the effects of land-use change

Many ecosystem services are delivered by organisms that depend on habitats that are segregated spatially or temporally from the location where services are provided. Management of mobile organisms contributing to ecosystem services requires consideration not only of the local scale where services are delivered, but also the distribution of resources at the landscape scale, and the foraging ranges and dispersal movements of the mobile agents. We develop a conceptual model for exploring how one such mobile-agent-based ecosystem service (MABES), pollination, is affected by land-use change, and then generalize the model to other MABES. The model includes interactions and feedbacks among policies affecting land use, market forces and the biology of the organisms involved. Animal-mediated pollination contributes to the production of goods of value to humans such as crops; it also bolsters reproduction of wild plants on which other services or service-providing organisms depend. About one-third of crop production depends on animal pollinators, while 60-90% of plant species require an animal pollinator. The sensitivity of mobile organisms to ecological factors that operate across spatial scales makes the services provided by a given community of mobile agents highly contextual. Services vary, depending on the spatial and temporal distribution of resources surrounding the site, and on biotic interactions occurring locally, such as competition among pollinators for resources, and among plants for pollinators. The value of the resulting goods or services may feed back via market-based forces to influence land-use policies, which in turn influence land management practices that alter local habitat conditions and landscape structure. Developing conceptual models for MABES aids in identifying knowledge gaps, determining research priorities, and targeting interventions that can be applied in an adaptive management context.

Searching efficiency and the functional response of a pause-travel forager

1. The feeding rates of many predators and parasitoids exhibit type II functional responses, with a decelerating rate of increase to reach an asymptotic value as the density of their prey or hosts increases. Holling's disc equation describes such relationships and predicts that the asymptotic feeding rate at high prey densities is set by handling time, while the rate at which feeding rate increases with increased prey density is determined by searching efficiency. Searching efficiency and handling time are also parameters in other models which describe the functional response. Models which incorporate functional responses in order to make predictions of the effects of food shortage thus rely upon a clear understanding and accurate quantification of searching efficiency and handling time. 2. Blackbird Turdus merula exhibit a type II functional response and use pause-travel foraging, a foraging technique in which animals search for prey while stationary and then move to capture prey. Pause-travel foraging allows accurate direct measurement of feeding rate and both searching efficiency and handling time. We use Blackbirds as a model species to: (i) compare observed measures of both searching efficiency and handling time with those estimated by statistically fitting the disc equation to the observed functional response; and (ii) investigate alternative measures of searching efficiency derived by the established method where search area is assumed to be circular and a new method that we propose where it is not. 3. We find that the disc equation can adequately explain the functional response of blackbirds feeding on artificial prey. However, this depends critically upon how searching efficiency is measured. Two variations on the previous method of measuring search area (a component of searching efficiency) overestimated searching efficiency, and hence predicted feeding rates higher than those observed. Two variations of our alternative approach produced lower estimates of searching efficiency, closer to that estimated by fitting the disc equation, and hence more accurately predicted feeding rate. Our study shows the limitations of the previous method of measuring searching efficiency, and describes a new method for measuring searching efficiency more accurately.

Response of plant-pollinator communities to fire: changes in diversity, abundance and floral reward structure

Globally, plant-pollinator communities are subject to a diverse array of perturbations and in many temperate and semi-arid systems fire is a dominant structuring force. We present a novel and highly integrated approach, which quantifies, in parallel, the response to fire of pollinator communities, floral communities and floral reward structure. Mt Carmel, Israel is a recognised bee-flower biodiversity hotspot, and using a chronosequence of habitats with differing post-fire ages, we follow the changes in plant-pollinator community organisation from immediately following a burn until full regeneration of vegetation. Initially, fire has a catastrophic effect on these communities, however, recovery is rapid with a peak in diversity of both flowers and bees in the first 2 years post-fire, followed by a steady decline over the next 50 years. The regeneration of floral communities is closely matched by that of their principal pollinators. At the community level we quantify, per unit area of habitat, key parameters of nectar and pollen forage known to be of importance in structuring pollinator communities. Nectar Volume, nectar water content, nectar concentration and the diversity of nectar foraging niches are all greatest immediately following fire with a steady decrease as regeneration proceeds. Temporal changes in energy availability for nectar, pollen, total energy (nectar + pollen) and relative importance of pollen to nectar energy show a similar general decline with site age, however, the pattern is less clear owing to the highly patchy distribution of floral resources. Changes in floral reward structure reflect the general shift from annuals (generally low-reward open access flowers) to perennials (mostly high-reward and restricted access flowers) as post-fire regeneration ensues. The impact of fire on floral communities and their associated rewards have clear implications for pollinator community structure and we discuss this and the role of other disturbance factors on these systems.

Noise, predation risk compensation and vigilance in the chaffinch Fringilla coelebs

Background noise should in theory hinder detection of auditory cues associated with approaching danger. We tested whether foraging chaffinches Fringilla coelebs responded to background noise by increasing vigilance, and examined whether this was explained by predation risk compensation or by a novel stimulus hypothesis. The former predicts that only inter-scan interval should be modified in the presence of background noise, not vigilance levels generally. This is because noise hampers auditory cue detection and increases perceived predation risk primarily when in the head-down position, and also because previous tests have shown that only interscan interval is correlated with predator detection ability in this system. Chaffinches only modified interscan interval supporting this hypothesis. At the same time they made significantly fewer pecks when feeding during the background noise treatment and so the increased vigilance led to a reduction in intake rate, suggesting that compensating for the increased predation risk could indirectly lead to a fitness cost. Finally, the novel stimulus hypothesis predicts that chaffinches should habituate to the noise, which did not occur within a trial or over 5 subsequent trials. We conclude that auditory cues may be an important component of the trade-off between vigilance and feeding, and discuss possible implications for anti-predation theory and ecological processes

Scavenging or infection? Possible host choosing by entomopathogenic nematodes

Entomopathogenic nematodes are able to survive by scavenging. We tested Steinernema feltiae, S. affine and Heterorhabditis megidis alone or in different combinations to evaluate the responses of these nematodes when dead or live Galleria mellonella larvae were offered. Steinernema feltiae and S. affine scavenged upon dead G. mellonella larvae and about 30% more dead larvae were penetrated than live ones. By contrast, H. megidis penetrated more live larvae than dead ones. When the nematode species were combined, the results varied among the combinations, but the dead larvae were always used as a host. The behaviour of natural field populations of S. feltiae and S. affine was also compared. Steinernema feltiae showed no difference between scavenging and performing 'normal infections', whereas S. affine scavenged to a reduced amount (around 60% less); this difference could be related to the particular foraging strategy of these nematodes.

Measuring the functional responses of farmland birds: an example for a declining seed-feeding bunting

1. Many farmland bird species have undergone significant declines. It is important to predict the effect of agricultural change on these birds and their response to conservation measures. This requirement could be met by mechanistic models that predict population size from the optimal foraging behaviour and fates of individuals within populations. A key component of these models is the functional response, the relationship between food and competitor density and feeding rate. 2. This paper describes a method for measuring functional responses of farmland birds, and applies this method to a declining farmland bird, the corn bunting Miliaria calandra L. We derive five alternative models to predict the functional responses of farmland birds and parameterize these for corn bunting. We also assess the minimum sample sizes required to predict accurately the functional response. 3. We show that the functional response of corn bunting can be predicted accurately from a few behavioural parameters (searching rate, handling time, vigilance time) that are straightforward to measure in the field. These parameters can be measured more quickly than the alternative of measuring the functional response directly. 4. While corn bunting violated some of the assumptions of Holling's disk equation (model 1 in our study), it still provided the most accurate fit to the observed feeding rates while remaining the most statistically simple model tested. Our other models may be more applicable to other species, or corn bunting feeding in other locations. 5. Although further tests are required, our study shows how functional responses can be predicted, simplifying the development of mechanistic models of farmland bird populations.

Arthropod diversity and community structure in relation to land use in the mekong delta, vietnam

Declining biodiversity in agro-ecosystems, caused by intensification of production or expansion of monocultures, is associated with the emergence of agricultural pests. Understanding how land-use and management control crop-associated biodiversity is, therefore, one of the key steps towards the prediction and maintenance of natural pest-control. Here we report on relationships between land-use variables and arthropod community attributes (for example, species diversity, abundance and guild structure) across a diversification gradient in a rice-dominated landscape in the Mekong delta, Vietnam. We show that rice habitats contained the most diverse arthropod communities, compared with other uncultivated and cultivated land-use types. In addition, arthropod species density and Simpson's diversity in flower, vegetable and fruit habitats was positively related to rice cover in the local landscape. However, across the landscape as a whole, reduction in heterogeneity and the amount of uncultivated cover was associated, generally, with a loss of diversity. Furthermore, arthropod species density in tillering and flowering stages of rice was positively related to crop and vegetation richness, respectively, in the local landscape. Differential effects on feeding guilds were also observed in rice-associated communities with the proportional abundance of predators increasing and the proportional abundance of detritivores decreasing with increased landscape rice cover. Thus, we identify a range of rather complex, sometimes contradictory patterns concerning the impact of rice cover and landscape heterogeneity on arthropod community attributes. Importantly, we conclude that that land-use change associated with expansion of monoculture rice need not automatically impact diversity and functioning of the arthropod community.

The potential of grass field margin management for enhancing beetle diversity in intensive livestock farms

1. Declining populations of UK grassland flora and fauna have been attributed to intensification of agricultural management practices, including changes in cutting, fertilizer, grazing and drainage regimes. We aimed to develop field margin management practices that could reverse declines in intensively managed grassland biodiversity that would have application in the UK and Europe. Here we focus on one aspect of grassland biodiversity, the beetles. 2. In four intensively managed livestock farms in south-west England, 10-m wide field margins in existing grasslands were managed to create seven treatments of increasing sward architectural complexity. This was achieved through combinations of inorganic (NPK) fertilizer, cattle grazing, and timing and height of cutting. To examine the potential influence of complexity on faunal diversity, beetles were identified to species level from suction samples taken between 2003 and 2005, and their assemblage structure was related to margin management, floral assemblages and sward architecture. 3. Beetle abundance, and species richness and evenness were influenced by margin management treatment and its interaction with year. Correlations with sward architecture and the percentage cover of dominant forbs and grasses were also found. Functional groups of the beetles showed different responses to the management treatments. In particular, higher proportional abundances of seed/flower-feeding guilds were found in treatments not receiving NPK fertilizer. 4. The assemblage structure was shown to respond to margin management treatments, sward architecture and the percentage cover of dominant forbs and grasses. The most extensively managed treatments were characterized by distinct successional trajectories from the control treatment. 5. Synthesis and applications. This study provides management options suitable for use within agri-environment schemes intended to improve faunal diversity associated with intensively managed lowland grasslands. Field margins receiving either no management or a single July silage cut were shown to support greater abundances and species richness of beetles, although subtler modifications of conventional management may also be beneficial, for example the absence of NPK fertilizer while maintaining grazing and silage cutting systems.

Grazing management of calcareous grasslands and its implications for the conservation of beetle communities

Calcareous grasslands are an important habitat for floral and faunal communities in the UK and Europe. Declines due to changes in management, scrub invasion and agricultural improvement have left much of the remnants of this habitat in a degraded and fragmented state. Grazing, by cattle or sheep, is one of the main management practices used to maintain and improve the floral and faunal quality of calcareous grassland. The long-term impacts of different grazing regimes, however, are poorly understood, particularly in terms of the invertebrate communities. This study contrasted the impacts of recently introduced and long-term sheep or cattle grazing on beetle communities present on one of the largest areas of calcareous grassland in Europe, the Salisbury Plain military training Area, UK. No effects of grazing management on beetle abundance, species. richness or evenness were found, but plant diversity and overall percentage cover of grasses did influence beetle diversity. Proportions of the total number of individuals and overall species richness within beetle guilds (predatory, phytophagous, flower/seed feeders, root feeders and foliage feeders) were strongly influenced by both the duration and type of grazing animal. At the species level, beetle community structure showed significant differences between ungrazed, long-term cattle and long-term sheep grazing treatments. Changes in plant community structure were found to influence beetle community structure. The significance of these results is discussed in terms of the long-term impacts of grazing on beetle community structure, and the benefits of different grazing regimes for the conservation management of calcareous grasslands. (c) 2005 Elsevier Ltd. All rights reserved.