Cloud-radar observations of insects in the UK convective boundary layer

Radar has been applied to the study of insect migration for almost 40 years, but most entomological radars operate at X-band (9.4 GHz, 3.2 cm wavelength), and can only detect individuals of relatively large species, such as migratory grasshoppers and noctuid moths, over all of their flight altitudes. Many insects (including economically important species) are much smaller than this, but development of the requisite higher power and/or higher frequency radar systems to detect these species is often prohibitively expensive. In this paper, attention is focussed upon the uses of some recently-deployed meteorological sensing devices to investigate insect migratory flight behaviour, and especially its interactions with boundary layer processes. Records were examined from the vertically-pointing 35 GHz ‘Copernicus’ and 94 GHz ‘Galileo’ cloud radars at Chilbolton (Hampshire, England) for 12 cloudless and convective occasions in summer 2003, and one of these occasions (13 July) is presented in detail. Insects were frequently found at heights above aerosol particles, which represent passive tracers, indicating active insect movement. It was found that insect flight above the convective boundary layer occurs most often during the morning. The maximum radar reflectivity (an indicator of aerial insect biomass) was found to be positively correlated with maximum screen temperature.

Insects, weather and climate

Applications of atmospheric science are relevant to a range of themes within science and society; application to entomology was the main focus of this meeting organised by Dr Curtis Wood (University of Reading). This meeting was held jointly with the Royal Entomological Society. The talks were designed to appeal to the broader scientific community by showcasing topics near the join of the two disciplines. The audience heard about exciting topics within weather and climate change, how they are applied to entomological science and how insects can be used to advance atmospheric science. The meeting included the 2009 Margary Lecture given by Prof. Philip Mellor from the Institute for Animal Health (IAH) at Pirbright.

Radar studies of the vertical distribution of insects migrating over southern Britain: the influence of temperature inversions on nocturnal layer concentrations

Insects migrating over two sites in southern UK (Malvern in Worcestershire, and Harpenden in Hertfordshire) have been monitored continuously with nutating vertical-looking radars (VLRs) equipped with powerful control and analysis software. These observations make possible, for the first time, a systematic investigation of the vertical distribution of insect aerial density in the atmosphere, over temporal scales ranging from the short (instantaneous vertical profiles updated every 15 min) to the very long (profiles aggregated over whole seasons or even years). In the present paper, an outline is given of some general features of insect stratification as revealed by the radars, followed by a description of occasions during warm nights in the summer months when intense insect layers developed. Some of these nocturnal layers were due to the insects flying preferentially at the top of strong surface temperature inversions, and in other cases, layering was associated with higher-altitude temperature maxima, such as those due to subsidence inversions. The layers were formed from insects of a great variety of sizes, but peaks in the mass distributions pointed to a preponderance of medium-sized noctuid moths on certain occasions.

The accuracy of Doppler radar wind retrievals using insects as targets

Insect returns from the UK's Doppler weather radars were collected in the summers of 2007 and 2008, to ascertain their usefulness in providing information about boundary layer winds. Such observations could be assimilated into numerical weather prediction models to improve forecasts of convective showers before precipitation begins. Significant numbers of insect returns were observed during daylight hours on a number of days through this period, when they were detected at up to 30 km range from the radars, and up to 2 km above sea level. The range of detectable insect returns was found to vary with time of year and temperature. There was also a very weak correlation with wind speed and direction. Use of a dual-polarized radar revealed that the insects did not orient themselves at random, but showed distinct evidence of common orientation on several days, sometimes at an angle to their direction of travel. Observation minus model background residuals of wind profiles showed greater bias and standard deviation than that of other wind measurement types, which may be due to the insects' headings/airspeeds and to imperfect data extraction. The method used here, similar to the Met Office's procedure for extracting precipitation returns, requires further development as clutter contamination remained one of the largest error contributors. Wind observations derived from the insect returns would then be useful for data assimilation applications.

Organic farming in isolated landscapes does not benefit flower-visiting insects and pollination

Organic farming has often been found to provide benefits for biodiversity, but the benefits can depend on the species considered and characteristics of the surrounding landscape. In an intensively farmed area of Northeast Italy we investigated whether isolated organic farms, in a conventionally farmed landscape, provided local benefits for insect pollinators and pollination services. We quantified the relative effects of local management (i.e. the farm system), landscape management (proportion of surrounding uncultivated land) and interactions between them. We compared six organic and six conventional vine fields. The proportion of surrounding uncultivated land was calculated for each site at radii of 200, 500, 1000 and 2000 m. The organic fields did not differ from the conventional in their floral resources or proportion of surrounding uncultivated land. Data were collected on pollinator abundance and species richness, visitation rates to, and pollination of experimental potted plants. None of these factors were significantly affected by the farming system. The abundance of visits to the potted plants in the conventional fields tended to be negatively affected by the proportion of surrounding uncultivated land. The proportion fruit set, weight of seeds per plant and seed weight in conventional and organic fields were all negatively affected by the proportion of surrounding uncultivated land. In vine fields the impact of the surrounding landscape was stronger than the local management. Enhancement of biodiversity through organic farming should not be assumed to be ubiquitous, as potential benefits may be offset by the crop type, organicmanagement practices and the specific habitat requirements in the surrounding landscape.

Influence of agricultural management, sward structure and food resources on grassland field use by birds in lowland England.

Agricultural management of grassland in lowland Britain has changed fundamentally in the last 50 years, resulting in spatial and structural uniformity within the pastoral landscape. The full extent to which these changes may have reduced the suitability of grassland as foraging habitat for birds is unknown. This study investigated the mechanisms by which these changes have impacted on birds and their food supplies. We quantified field use by birds in summer and winter in two grassland areas of lowland England (Devon and Buckinghamshire) over 3 years, relating bird occurrence to the management, sward structure and seed and invertebrate food resources of individual fields. Management intensity was defined in terms of annual nitrogen input. There was no consistent effect of management intensity on total seed head production, although those of grasses generally increased with inputs while forbs were rare throughout. Relationships between management intensity and abundance of soil and epigeal invertebrates were complex. Soil beetle larvae were consistently lower in abundance, and surface-active beetle larvae counts consistently higher, in intensively managed fields. Foliar invertebrates showed more consistent negatively relationships with management intensity. Most bird species occurred at low densities. There were consistent relationships across regions and years between the occurrence of birds and measures of field management. In winter, there was a tendency towards higher occupancy of intensively managed fields by species feeding on soil invertebrates. In summer, there were few such relationships, although many species avoided fields with tall swards. Use of fields by birds was generally not related to measures of seed or invertebrate food abundance. While granivorous species were perhaps too rare to detect a relationship, in insectivores the strong negative relationships (in summer) with sward height suggested that access to food may be the critical factor. While it appears that intensification of grassland management has been deleterious to the summer food resources of insectivorous birds that use insects living within the grass sward, intensification may have been beneficial to several species in winter through the enhancement of soil invertebrates. Synthesis and applications. We suggest that attempts to restore habitat quality for birds in grassland landscapes need to create a range of management intensities and sward structures at the field and farm scales. A greater understanding of methods to enhance prey accessibility, as well as abundance, for insectivorous birds is required.

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.

Summer drought alters plant-mediated competition between foliar- and root-feeding insects

Summer droughts are predicted to increase in severity and frequency in the United Kingdom, due to climate change. Few studies have addressed the impacts of drought on interactions between species, and the majority have focussed on increases in CO2 concentration and changes in temperature. Here, the effect of experimental summer drought on the strength of the plant-mediated interaction between leaf-mining Stephensia brunnichella larvae and root-chewing Agriotes larvae was investigated. Agriotes larvae reduced the abundance and performance of S. brunnichella feeding on a mutual host plant, Clinopodium vulgare, as well as the rate of parasitism of the leaf-miner. The interaction did not, however, occur on plants subjected to a severe drought treatment, which were reduced in size. Changes to summer rainfall, due to climate change, may therefore reduce the occurrence of plant-mediated interactions between insect herbivores.

Farm level tree planting in Pakistan: The role of farmers' perceptions and attitudes

The low proportion of forested land and continuing degradation of existing forest cover are serious threats to the sustainability of forestry in Pakistan. Farm forestry has been identified as a feasible solution, particularly in the plain areas. Applying the Theory of Planned Behaviour in a survey of 124 farmers in Dera Ismail Khan district of Pakistan's North West Frontier Province showed that farmers' willingness to grow trees on their farms is a function of their attitudes towards the advantages and disadvantages of growing trees, their perception of the opinions of salient referents and factors that encourage and discourage farm level tree planting. Farmers viewed farm forestry as economically beneficial and environmentally friendly. Tree planting was perceived as increasing income, providing wood for fuel and furniture, controlling erosion and pollution and providing shade for humans and animals. Farmers saw hindrance in agricultural operations and the harbouring of insects, pests and diseases as negative impacts of tree planting; however, these were outweighed by their perceptions of positive impacts. Tree growing decisions of farmers were influenced by the opinions of family members, owners/tenants, fellow farmers and village elders. The factors that significantly predicted farm level tree planting were availability of barren land, lack of markets, lack of nurseries and damage caused by animals and humans. Farm forestry programmes are more likely to be successful if they acknowledge and address the factors which underlie farmers' reasons for planting or not planting trees.

The effect of acetylsalicylic acid and oxalic acid on Myzus persicae and Aphidius colemani

Plants can respond to damage by pests with both induced direct defences and indirect defences by the attraction of their natural enemies. Foliar application of several plant-derived chemicals, such as salicylic acid and oxalic acid, can induce these defence mechanisms. The effect of acetylsalicylic acid and oxalic acid on the aphid Myzus persicae Sulzer (Homoptera: Aphididae) and its parasitoid Aphidius colemani Viereck (Hymenoptera: Aphidiidae) was investigated. Experiments were carried out with direct application of acetylsalicylic and oxalic acids on these insects, as well as choice and no-choice tests using foliar application of both chemicals on Brussels sprouts plants, Brassica oleracea var. gemmifera L. (Brassicaceae). Parasitoids were given a choice between treated and untreated plants for oviposition, and the effects of the chemicals on aphid and parasitoid development were determined. Although direct application of both chemicals increased aphid mortality, their foliar application did not induce resistance against aphids. The foliar application of such compounds, even in low concentration as shown in the choice tests, has the potential to induce indirect plant defences against aphids by encouraging aphid parasitisation. Although the direct application of both chemicals reduced parasitoid emergence from their hosts, the foliar application of acetylsalicylic acid and low concentrations of oxalic acid did not have a negative effect on parasitoid emergence ability. However, 10 mm oxalic acid reduced the number of emerged parasitoids in no-choice experiments. This study shows that foliar application of acetylsalicylic and oxalic acids has the potential to encourage aphid parasitisation, but care is needed as high concentrations of oxalic acid can have a negative effect on these beneficial organisms.

Was the removal of rabbits and house mice from Selvagem Grande beneficial to the breeding of Cory's shearwaters Calonectris diomedea borealis?

The breeding success of Cory's shearwaters Calonectris diomedea borealis at its important Atlantic colony on Selvagern Grande has been monitored periodically at two study plots since 1982. A successful eradication programme was implemented to remove two alien invasive mammals, rabbits Oryctolagus cuniculus and house mice Mus musculus, from the island during 2002. The availability of long-term breeding data for Cory's shearwaters on Selvagern Grande provided a unique opportunity to study the effects of the removal of rabbits and mice on seabird breeding. Annual observation of approximately 400 Cory's nests showed that significantly more birds fledged from both study sites in the five breeding seasons after the eradication than in the 13 seasons prior to it for which reliable breeding data were available. The numbers of young birds present at the time of fledging were an average of 47 and 23% greater than pre-eradication numbers at the two study sites. The eradication of rabbits and mice was simultaneous and, therefore, it was impossible to attribute the increased breeding success of Cory's shearwaters to the removal of one or other species. However, both are known to have adverse impacts on the breeding of nesting seabirds. These observations provide important justification for the implementation of further programmes for the removal of alien invasive mammals from oceanic islands.

An evaluation of the costs of making specific secondary metabolites: does the yield penalty incurred by host plant resistance to insects due to competition for resources?

The presumption that the synthesis of 'defence' compounds in plants must incur some 'trade-off' or penalty in terms of annual crop yields has been used to explain observed inverse correlations between resistance to herbivores and rates of growth or photosynthesis. An analysis of the cost of making secondary compounds suggests that this accounts for only a small part of the overall carbon budget of annual crop plants. Even the highest reported amounts of secondary metabolites found in different crop species (flavonoids, allylisothiocyanates, hydroxamic acids, 2-tridecanone) represent a carbon demand that can be satisfied by less than an hour's photosynthesis. Similar considerations apply to secondary compounds containing nitrogen or sulphur, which are unlikely to represent a major investment compared to the cost of making proteins, the major demand for these elements. Decreases in growth and photosynthesis in response to stress are more likely the result of programmed down-regulation. Observed correlations between yield and low contents of unpalatable or toxic compounds may be the result of parallel selection during the refinement of crop species by humans.

On the stability of populations of mammals, birds, fish and insects

A key concern for conservation biologists is whether populations of plants and animals are likely to fluctuate widely in number or remain relatively stable around some steady-state value. In our study of 634 populations of mammals, birds, fish and insects, we find that most can be expected to remain stable despite year to year fluctuations caused by environmental factors. Mean return rates were generally around one but were higher in insects (1.09 +/- 0.02 SE) and declined with body size in mammals. In general, this is good news for conservation, as stable populations are less likely to go extinct. However, the lower return rates of the large mammals may make them more vulnerable to extinction. Our estimates of return rates were generally well below the threshold for chaos, which makes it unlikely that chaotic dynamics occur in natural populations - one of ecology's key unanswered questions.