Floral volatiles controlling ant behaviour

P>1. Ants show complex interactions with plants, both facultative and mutualistic, ranging from grazers through seed predators and dispersers to herders of some herbivores and guards against others. But ants are rarely pollinators, and their visits to flowers may be detrimental to plant fitness. 2. Plants therefore have various strategies to control ant distributions, and restrict them to foliage rather than flowers. These 'filters' may involve physical barriers on or around flowers, or 'decoys and bribes' sited on the foliage (usually extrafloral nectaries - EFNs). Alternatively, volatile organic compounds (VOCs) are used as signals to control ant behaviour, attracting ants to leaves and/or deterring them from functional flowers. Some of the past evidence that flowers repel ants by VOCs has been equivocal and we describe the shortcomings of some experimental approaches, which involve behavioural tests in artificial conditions. 3. We review our previous study of myrmecophytic acacias, which used in situ experiments to show that volatiles derived from pollen can specifically and transiently deter ants during dehiscence, the effects being stronger in ant-guarded species and more effective on resident ants, both in African and Neotropical species. In these plants, repellence involves at least some volatiles that are known components of ant alarm pheromones, but are not repellent to beneficial bee visitors. 4. We also present new evidence of ant repellence by VOCs in temperate flowers, which is usually pollen-based and active on common European ants. We use these data to indicate that across a wide range of plants there is an apparent trade-off in ant-controlling filter strategies between the use of defensive floral volatiles and the alternatives of decoying EFNs or physical barriers.

Terrestrial carnivores and human food production: impact and management

1. The production of food for human consumption has led to an historical and global conflict with terrestrial carnivores, which in turn has resulted in the extinction or extirpation of many species, although some have benefited. At present, carnivores affect food production by: (i) killing human producers; killing and/or eating (ii) fish/shellfish; (iii) game/wildfowl; (iv) livestock; (v) damaging crops; (vi) transmitting diseases; and (vii) through trophic interactions with other species in agricultural landscapes. Conversely, carnivores can themselves be a source of dietary protein (bushmeat). 2. Globally, the major areas of conflict are predation on livestock and the transmission of rabies. At a broad scale, livestock predation is a customary problem where predators are present and has been quantified for a broad range of carnivore species, although the veracity of these estimates is equivocal. Typically, but not always, losses are small relative to the numbers held, but can be a significant proportion of total livestock mortality. Losses experienced by producers are often highly variable, indicating that factors such as husbandry practices and predator behaviour may significantly affect the relative vulnerability of properties in the wider landscape. Within livestock herds, juvenile animals are particularly vulnerable. 3. Proactive and reactive culling are widely practised as a means to limit predation on livestock and game. Historic changes in species' distributions and abundance illustrate that culling programmes can be very effective at reducing predator density, although such substantive impacts are generally considered undesirable for native predators. However, despite their prevalence, the effectiveness, efficiency and the benefit:cost ratio of culling programmes have been poorly studied. 4. A wide range of non-lethal methods to limit predation has been studied. However, many of these have their practical limitations and are unlikely to be widely applicable. 5. Lethal approaches are likely to dominate the management of terrestrial carnivores for the foreseeable future, but animal welfare considerations are increasingly likely to influence management strategies. The adoption of non-lethal approaches will depend upon proof of their effectiveness and the willingness of stakeholders to implement them, and, in some cases, appropriate licensing and legislation. 6. Overall, it is apparent that we still understand relatively little about the importance of factors affecting predation on livestock and how to manage this conflict effectively. We consider the following avenues of research to be essential: (i) quantified assessments of the loss of viable livestock; (ii) landscape-level studies of contiguous properties to quantify losses associated with variables such as different husbandry practices; (iii) replicated experimental manipulations to identify the relative benefit of particular management practices, incorporating (iv) techniques to identify individual predators killing stock; and (v) economic analyses of different management approaches to quantify optimal production strategies.

Avoidance responses of an aphidophagous ladybird, Adalia bipunctata, to aphid-tending ants

1. Insect predators often aggregrate to patches of high prey density and use prey chemicals as cues for oviposition. If prey have mutualistic guardians such as ants, however, then these patches may be less suitable for predators. 2. Ants often tend aphids and defend them against predators such as ladybirds. Here, we show that ants can reduce ladybird performance by destroying eggs and physically attacking larvae and adults. 3. Unless ladybirds are able to defend against ant attacks they are likely to have adaptations to avoid ants. We show that Adalia bipunctata ladybirds not only move away from patches with Lasius niger ants, but also avoid laying eggs in these patches. Furthermore, ladybirds not only respond to ant presence, but also detect ant semiochemicals and alter oviposition strategy accordingly. 4. Ant semiochemicals may signal the extent of ant territories allowing aphid predators to effectively navigate a mosaic landscape of sub-optimal patches in search of less well-defended prey. Such avoidance probably benefits both ants and ladybirds, and the semiochemicals could be regarded as a means of cooperative communication between enemies. 5. Overall, ladybirds respond to a wide range of positive and negative oviposition cues that may trade-off with each other and internal motivation to determine the overall oviposition strategy.

Owls and rabbits: predation against substandard individuals of an easy prey

The interactions among the multiple factors regulating predator-prey relationships make predation a more complex process than previously thought. The degree to which substandard individuals are captured disproportionately seems to be better a function of the difficulty of prey capture than of the hunting techniques (coursing vs. ambushing predators). That is, when the capture and killing of a prey species is easy, substandard individuals will be predated in proportion to their occurrence in the prey population. In the present study, we made use of eagle owls Bubo bubo and their main prey, the rabbit Oryctolagus cuniculus: (a) the brightness of the white tails of rabbits seems to be correlated with the physical condition of individuals, (b) by using the tails of predated rabbits as an index of individual condition, we found that eagle owls seem to prefer substandard individuals (characterized by duller tails), and (c) by using information from continuous radiotracking of 14 individuals, we suggest that the difficulty of rabbit capture could be low. Although the relative benefits of preying on substandard individuals should considerably decrease when a predator is attacking an easy prey, we hypothesise that the eagle owl preference for substandard individuals could be due to the easy detection of poor individuals by a visual cue, the brightness of the rabbit tail. Several elements allow us to believe that this form of visual communication between a prey and one of its main predators could be more widespread than previously thought. In fact: (a) visual signalling plays a relevant role in intraspecific communication in eagle owls and, consequently, visual signals could also play a role in interspecific interactions, and (b) empirical studies showed that signals may inform the predator that it has been perceived, or that the prey is in a sufficiently healthy state to elude the predator.

Aphidius ervi preferentially attacks the green morph of the pea aphid, Acyrthosiphon pisum

The pea aphid, Acyrthosiphon pisum Harris (Hemiptera: Aphididae) is found in red and green color morphs. Previous work has suggested that the aphidiine parasitoid Aphidius ervi Haliday preferentially attacks green pea aphids in the field. It is not clear whether these results reflect a real preference, or some unknown clonal difference, such as in immunity, between the aphids used in the previous studies. We used three susceptibility-matched pairs of red and green morph pea aphid clones to test for preferences. In a no-choice situation, the parasitoids attacked equal proportions of each color morph. When provided with a choice, A. ervi was significantly more likely to oviposit into colonies formed from green morphs when the neighboring colony was formed from red morph aphids. In contrast, red morphs were less likely to be attacked when their neighboring colony was of the green morph. By preferentially attacking green colonies, A. ervi may reduce the likelihood of intraguild predation, as it is suggested that visually foraging predators preferentially attack red aphid colonies. Furthermore, if this host choice behavior is replicated in the field, we speculate that color morphs of the pea aphid may interact indirectly through their shared natural enemies, leading to intraspecific apparent competition.

Post-dispersal seed predation of non-target weeds in arable crops

Field experiments were conducted to quantify the natural levels of post-dispersal seed predation of arable weed species in spring barley and to identify the main groups of seed predators. Four arable weed species were investigated that were of high biodiversity value, yet of low to moderate competitive ability with the crop. These were Chenopodium album, Sinapis arvensis, Stellaria media and Polygonum aviculare. Exclusion treatments were used to allow selective access to dishes of seeds by different predator groups. Seed predation was highest early in the season, followed by a gradual decline in predation over the summer for all species. All species were taken by invertebrates. The activity of two phytophagous carabid genera showed significant correlations with seed predation levels. However, in general carabid activity was not related to seed predation and this is discussed in terms of the mainly polyphagous nature of many Carabid species that utilized the seed resource early in the season, but then switched to carnivory as prey populations increased. The potential relevance of post-dispersal seed predation to the development of weed management systems that maximize biological control through conservation and optimize herbicide use, is discussed.

Seasonal 'fall out' of sessile macro-fauna from submarine cliffs: quantification, causes and implications

Submarine cliffs are typically crowded with sessile organisms, most of which are ultimately exported downwards. Here we report a 24 month study of benthic fauna dropping from such cliffs at sites of differing cliff angle and flow rates at Lough Hyne Marine Nature Reserve, Co. Cork, Ireland. The magnitude of 'fall out' material collected in capture nets was highly seasonal and composed of sessile and mobile elements. Sponges, ascidians, cnidarians, polychaetes, bryozoans and barnacles dominated the sessile forms. The remainder (mobile fauna) were scavengers and predators such as asteroid echinoderms, gastropod molluscs and malacostracan crustaceans. These were probably migrants targeting fallen sessile organisms. 'Fall out' material (including mobile forms) increased between May and August in both years. This increase in 'fall out' material was correlated with wrasse abundance at the cliffs (with a one month lag period). The activities of the wrasse on the cliffs (feeding, nest building and territory defence) were considered responsible for the majority of 'fall out' material, with natural mortality and the activity of other large mobile organisms (e.g. crustaceans) also being triplicated. Current flow rate and cliff profile were important in amount of 'fall out' material collected. In low current situations export of fallen material was vertical, while both horizontal and vertical export was associated with moderate to high current environments. Higher 'fall out' was associated with overhanging than vertical cliff surfaces. The 'fall out' of marine organisms in low current situations is likely to provide ail important source of nutrition in close proximity to the cliff, in an otherwise impoverished soft sediment habitat. However, in high current areas material will be exported some distance from the source, with final settlement again occurring in soft sediment habitats (as current speed decreases).

Temporal and spatial variability of mobile fauna on a submarine cliff and boulder scree complex: a community in flux

The mobile component of a community inhabiting a submarine boulder scree/cliff was investigated at Lough Hyne, Ireland at dawn, midday, dusk and night over a 1-week period. Line transects (50 m) were placed in the infralittoral (6 m) and circumlittoral (18 m) zones and also the interface between these two zones (12 m). The dominant mobile fauna of this cliff consisted of echinoderms (6 species), crustaceans (10 species) and fish (23 species). A different component community was identified at each time/depth interval using Multi-Dimensional Scaling (MDS) even though both species diversity (Shannon-Wiener indices) and richness (number of species) remained constant. These changes in community composition provided indirect evidence for migration by these mobile organisms. However, little evidence was found for migration between different zones with the exception of the several wrasse species. These species were observed to spend the daytime foraging in the deeper zone, but returned to the upper zone at night presumably for protection from predators. For the majority of species, migration was considered to occur to cryptic habitats such as holes and crevices. The number of organisms declined during the night, although crustacean numbers peaked, while fish and echinoderms were most abundant during day, possibly due to predator-prey interactions. This submarine community is in a state of flux, whereby, community characteristics, including trophic and energetic relationships, varied over small temporal (daily) and spatial (m) scales.

Combining physical, cultural and biological methods: prospects for integrated non-chemical weed management strategies

Physical, cultural and biological methods for weed control have developed largely independently and are often concerned with weed control in different systems: physical and cultural control in annual crops and biocontrol in extensive grasslands. We discuss the strengths and limitations of four physical and cultural methods for weed control: mechanical, thermal, cutting, and intercropping, and the advantages and disadvantages of combining biological control with them. These physical and cultural control methods may increase soil nitrogen levels and alter microclimate at soil level; this may be of benefit to biocontrol agents, although physical disturbance to the soil and plant damage may be detrimental. Some weeds escape control by these methods; we suggest that these weeds may be controlled by biocontrol agents. It will be easiest to combine biological control with. re and cutting in grasslands; within arable systems it would be most promising to combine biological control (especially using seed predators and foliar pathogens) with cover-cropping, and mechanical weeding combined with foliar bacterial and possibly foliar fungal pathogens. We stress the need to consider the timing of application of combined control methods in order to cause least damage to the biocontrol agent, along with maximum damage to the weed and to consider the wider implications of these different weed control methods.

The joint effects of larval density and C-14-cypermethrin on the life history and population growth rate of the midge Chironomus riparius

1. Chemical effects on organisms are typically assessed using individual-level endpoints or sometimes population growth rate (PGR), but such measurements are generally made at low population densities. In contrast most natural populations are subject to density dependence and fluctuate around the environmental carrying capacity as a result of individual competition for resources. As ecotoxicology aims to make reliable population projections of chemical impacts in the field, an understanding of how high-density or resource-limited populations respond to environmental chemicals is essential. 2. Our objective was to determine the joint effects of population density and chemical stress on the life history and PGR of an important ecotoxicological indicator species, Chironomus riparius, under controlled laboratory conditions. Populations were fed the same ration but initiated at different densities and exposed to a solvent control and three concentrations of C-14-cypermethrin in a sediment-water test system for 67 days at 20 +/- 1 degreesC. 3. Density had a negative effect on all the measured life-history traits, and PGR declined with increasing density in the controls. Exposure to C-14-cypermethrin had a direct negative effect on juvenile survival, presumably within the first 24 h because the chemical rapidly dissipated from the water column. Reductions in the initial larval densities resulted in an increase in the available resources for the survivors. Subsequently, exposed populations emerged sooner and started producing offspring earlier than the controls. C-14-cypermethrin had no effect on estimated fecundity and adult body weight but interacted with density to reduce the time to first emergence and first reproduction. As a result, PGR increased with cypermethrin concentration when populations were initiated at high densities. 4. Synthesis and applications. The results showed that the effects of C-14-cypermethrin were buffered at high density, so that the joint effects of density and chemical stress on PGR were less than additive. Low levels of chemical stressors may increase carrying capacity by reducing juvenile competition for resources. More and perhaps fitter adults may be produced, similar to the effects of predators and culling; however, toxicant exposure may result in survivors that are less tolerant to changing conditions. If less than additive effects are typical in the field, standard regulatory tests carried out at low density may overestimate the effects of environmental chemicals. Further studies over a wide range of chemical stressors and organisms with contrasting life histories are needed to make general recommendations.

Climate and biological control in organic crops

Organic farming has increased in popularity in recent years, primarily as a response to the perceived health and conservation benefits. While it is likely that conventional farming will be able to respond rapidly to variations in pest numbers and distribution resulting from climatic change, it is not clear if the same is true for organic farming. Few studies have looked at the responses of biological control organisms to climate change. Here, I review the direct and indirect eects of changes in temperature, atmospheric carbon dioxide and other climatic factors on the predators, parasitoids and pathogens of pest insects in temperate agriculture. Finally, I consider what research is needed to manage the anticipated change in pest insect dynamics and distributions.

Nocturnal ranging behaviour of urban hedgehogs, Erinaceus europaeus, in relation to risk and reward

Urban areas have both positive and negative influences on wildlife. For terrestrial mammals, one of the principle problems is the risk associated with moving through the environment whilst foraging. In this study, we examined nocturnal patterns of movement of urban-dwelling hedgehogs (Erinaceus europaeus) in relation to (i) the risks posed by predators and motor vehicles and (ii) nightly weather patterns. Hedgehogs preferentially utilised the gardens of semi-detached and terraced houses. However, females, but not males, avoided the larger back gardens of detached houses, which contain more of the habitat features selected by badgers. This difference in the avoidance of predation risk is probably associated with sex differences in breeding behaviour. Differences in nightly movement patterns were consistent with strategies associated with mating behaviour and the accumulation of fat reserves for hibernation. Hedgehogs also exhibited differences in behaviour associated with the risks posed by humans; they avoided actively foraging near roads and road verges, but did not avoid crossing roads per se. They were, however, significantly more active after midnight when there was a marked reduction in vehicle and foot traffic. In particular, responses to increased temperature, which is associated with increased abundance of invertebrate prey, were only observed after midnight. This variation in the timing of bouts of activity would reduce the risks associated with human activities. There were also profound differences in both area ranged and activity with chronological year which warrant further investigation.

Accumulation of anticoagulant rodenticides in a non-target insectivore, the European hedgehog (Erinaceus europaeus)

Studies on exposure of non-targets to anticoagulant rodenticides have largely focussed on predatory birds and mammals; insectivores have rarely been studied. We investigated the exposure of 120 European hedgehogs (Erinaceus europaeus) from throughout Britain to first- and second-generation anticoagulant rodenticides (FGARs and SGARs) using high performance liquid chromatography coupled with fluorescence detection (HPLC) and liquid-chromatography mass spectrometry (LCMS). The proportion of hedgehogs with liver SGAR concentrations detected by HPLC was 3-13% per compound, 23% overall. LCMS identified much higher prevalence for difenacoum and bromadiolone, mainly because of greater ability to detect low level contamination. The overall proportion of hedgehogs with LCMS-detected residues was 57.5% (SGARs alone) and 66.7% (FGARs and SGARs combined); 27 (22.5%) hedgehogs contained >1 rodenticide. Exposure of insectivores and predators to anticoagulant rodenticides appears to be similar. The greater sensitivity of LCMS suggests that hitherto exposure of non-targets is likely to have been under-estimated using HPLC techniques.

Local management and landscape drivers of pollination and biological control services in a Kenyan agro-ecosystem

Arthropods that have a direct impact on crop production (i.e. pests, natural enemies and pollinators) can be influenced by both local farm management and the context within which the fields occur in the wider landscape. However, the contributions and spatial scales at which these drivers operate and interact are not fully understood, particularly in the developing world. The impact of both local management and landscape context on insect pollinators and natural enemy communities and on their capacity to deliver related ecosystem services to an economically important tropical crop, pigeonpea was investigated. The study was conducted in nine paired farms across a gradient of increasing distance to semi-native vegetation in Kibwezi, Kenya. Results show that proximity of fields to semi-native habitats negatively affected pollinator and chewing insect abundance. Within fields, pesticide use was a key negative predictor of pollinator, pest and foliar active predator abundance. On the contrary, fertilizer application significantly enhanced pollinator and both chewing and sucking insect pest abundance. At a 1 km spatial scale of fields, there were significant negative effects of the number of semi-native habitat patches within fields dominated by mass flowering pigeonpea on pollinators abundance. For service provision, a significant decline in fruit set when insects were excluded from flowers was recorded. This study reveals the interconnections of pollinators, predators and pests with pigeonpea crop. For sustainable yields and to conserve high densities of both pollinators and predators of pests within pigeonpea landscapes, it is crucial to target the adoption of less disruptive farm management practices such as reducing pesticide and fertilizer inputs.

How predation and landscape fragmentation affect vole population dynamics

Background: Microtine species in Fennoscandia display a distinct north-south gradient from regular cycles to stable populations. The gradient has often been attributed to changes in the interactions between microtines and their predators. Although the spatial structure of the environment is known to influence predator-prey dynamics of a wide range of species, it has scarcely been considered in relation to the Fennoscandian gradient. Furthermore, the length of microtine breeding season also displays a north-south gradient. However, little consideration has been given to its role in shaping or generating population cycles. Because these factors covary along the gradient it is difficult to distinguish their effects experimentally in the field. The distinction is here attempted using realistic agent-based modelling. Methodology/Principal Findings: By using a spatially explicit computer simulation model based on behavioural and ecological data from the field vole (Microtus agrestis), we generated a number of repeated time series of vole densities whose mean population size and amplitude were measured. Subsequently, these time series were subjected to statistical autoregressive modelling, to investigate the effects on vole population dynamics of making predators more specialised, of altering the breeding season, and increasing the level of habitat fragmentation. We found that fragmentation as well as the presence of specialist predators are necessary for the occurrence of population cycles. Habitat fragmentation and predator assembly jointly determined cycle length and amplitude. Length of vole breeding season had little impact on the oscillations. Significance: There is good agreement between our results and the experimental work from Fennoscandia, but our results allow distinction of causation that is hard to unravel in field experiments. We hope our results will help understand the reasons for cycle gradients observed in other areas. Our results clearly demonstrate the importance of landscape fragmentation for population cycling and we recommend that the degree of fragmentation be more fully considered in future analyses of vole dynamics.