Orientation Cues for High-Flying Nocturnal Insect Migrants: Do Turbulence-Induced Temperature and Velocity Fluctuations Indicate the Mean Wind Flow?

Migratory insects flying at high altitude at night often show a degree of common alignment, sometimes with quite small angular dispersions around the mean. The observed orientation directions are often close to the downwind direction and this would seemingly be adaptive in that large insects could add their self-propelled speed to the wind speed, thus maximising their displacement in a given time. There are increasing indications that high-altitude orientation may be maintained by some intrinsic property of the wind rather than by visual perception of relative ground movement. Therefore, we first examined whether migrating insects could deduce the mean wind direction from the turbulent fluctuations in temperature. Within the atmospheric boundary-layer, temperature records show characteristic ramp-cliff structures, and insects flying downwind would move through these ramps whilst those flying crosswind would not. However, analysis of vertical-looking radar data on the common orientations of nocturnally migrating insects in the UK produced no evidence that the migrants actually use temperature ramps as orientation cues. This suggests that insects rely on turbulent velocity and acceleration cues, and refocuses attention on how these can be detected, especially as small-scale turbulence is usually held to be directionally invariant (isotropic). In the second part of the paper we present a theoretical analysis and simulations showing that velocity fluctuations and accelerations felt by an insect are predicted to be anisotropic even when the small-scale turbulence (measured at a fixed point or along the trajectory of a fluid-particle) is isotropic. Our results thus provide further evidence that insects do indeed use turbulent velocity and acceleration cues as indicators of the mean wind direction.     

Diurnal flight behavior of Ichneumonoidea (Insecta: Hymenoptera) related to environmental factors in a tropical dry forest

The biology and behavior of insects are strongly influenced by environmental conditions such as temperature and precipitation. Because some of these factors present a within day variation, they may be causing variations on insect diurnal flight activity, but scant information exists on the issue. The aim of this work was to describe the patterns on diurnal variation of the abundance of Ichneumonoidea and their relation with relative humidity, temperature, light intensity, and wind speed. The study site was a tropical dry forest at Ría Lagartos Biosphere Reserve, Mexico; where correlations between environmental factors (relative humidity, temperature, light, and wind speed) and abundance of Ichneumonidae and Braconidae (Hymenoptera: Ichneumonoidea) were estimated. The best regression model for explaining abundance variation was selected using the second order Akaike Information Criterion. The optimum values of temperature, humidity, and light for flight activity of both families were also estimated. Ichneumonid and braconid abundances were significantly correlated to relative humidity, temperature, and light intensity; ichneumonid also showed significant correlations to wind speed. The second order Akaike Information Criterion suggests that in tropical dry conditions, relative humidity is more important that temperature for Ichneumonoidea diurnal activity. Ichneumonid wasps selected toward intermediate values of relative humidity, temperature and the lowest wind speeds; while Braconidae selected for low values of relative humidity. For light intensity, braconids presented a positive selection for moderately high values.     

How to avoid becoming a prey: Predatory encounters between an orb-weaving spider,Araneus pinguis (Karsch) (Araneae: Araneidae) and flying insects

Insects flying into the web of an orb-weaving spiderAraneus pinguis (Karsch) and their avoidance of (pre-hitting process) and escapes from (post-hitting process) the web were examined by direct observation under natural and semi-natural conditions. In the pre-hitting process, mobile insects such as Brachycera, Lepidoptera and Hymenoptera showed a low hitting ratio (number of insects hitting/number of insects flying within 1 m³ space around the web-site) because of active web avoidance and flying activity in layers lower or higher than those in which the webs are usually laid. In contrast, less mobile insects like Heteroptera, Coleoptera and Homoptera showed a high hitting ratio. In the post-hitting process, Brachycera, Lepidoptera and some Nematocera frequently escaped without being detained by the web. Many Orthoptera and Hymenoptera escaped without any sign of detection by the spider. Coleoptera frequently escaped during the spider's attack. Small insects from the Homoptera, Nematocera and Hymenoptera rarely escaped from the web, but were not immediately attacked. Mean escape time of insects was correlated significantly with capture success of the spider. Overall most of the escapes occurred in the early phases of the predation process. This indicates that escapes are unlikely to result in heavy loss of time and energy expenditure due to unsuccessful predation. Escape patterns of insects seem to be related to their mobility.     

To what extent can the tiny parasitoid wasps,Eretmocerus mundus,fly upwind?

Body miniaturization in insects is predicted to result in decreased flight speed and therefore limited ability of these insects to fly upwind. Therefore, tiny insects are often regarded as relying on passive dispersal by winds. We tested this assumption in a wind tunnel by measuring the burst speed of Eretmocerus mundus (Mercet), a beneficial parasitoid wasp with body length <1 mm. Insects were filmed flying upwind towards a UV light source in a range of wind speed 0–0.5 m/s. The Insects flew towards the UV light in the absence and presence of wind but increased their flight speed in the presence of wind. They also changed flight direction to be directly upwind and maintained this body orientation even while drifted backwards relative to the ground by stronger winds. Field measurements showed that the average flight speed observed in the wind tunnel (0.3 m/s) is sufficient to allow flying between plants even when the wind speed above the vegetation was 3–5 folds higher. A simulation of the ability of the insects to control their flight trajectory towards a visual target (sticky traps) in winds show that the insects can manipulate their progress relative to the ground even when the wind speed exceeds their flight speed. The main factors determining the ability of the insects to reach the trap were trap diameter and the difference between insect flight speed and wind speed. The simulation also predicts the direction of arrival to the sticky target showing that many of the insects reach the target from the leeward side (i.e. by flight upwind). In light of these results, the notion that miniature insects passively disperse by winds is misleading because it disregards the ability of the insects to control their drift relative to the ground in winds that are faster than their flight speed.     

The Flight Heights of Chalcid Wasps (Hymenoptera,Chalcidoidea) in a Lowland Bornean Rain Forest: Fig Wasps are the High Fliers1

Tropical rain forests are characterized by their rich plant diversity and highly diverse insect faunas containing mainly rare species. Phytophagous and parasitoid insects utilizing such fragmented resources often must travel considerable distances to find suitable hosts. For small, weak‐flying insects, entry into the fast‐flowing air above the canopy can provide one way by which long‐distance dispersal is achieved. Using sticky traps placed at different heights in a lowland rain forest of Borneo, we compared the diurnal and nocturnal flight heights of chalcids, a group of mainly very small parasitoids and phytophages, to determine if the air above the canopy was used for dispersal. Most families were represented throughout the range of trap heights, including those above the general canopy. A higher proportion of individuals were trapped above the canopy at night than during the day. Fig wasps were exceptional in that they were trapped almost entirely above the canopy. They included species associated with host trees that do not fruit in the canopy, suggesting that these short‐lived, slow‐flying insects actively fly up above the canopy and then use the wind to passively carry them the long distances needed to reach their highly localized and ephemeral hosts. Once the fig wasps detect the species‐specific volatiles released by their host figs, they then may fly down into the canopy, where the lower wind speeds would allow them to fly actively upwind to their hosts.     

Group-Specific Multiplex PCR Detection Systems for the Identification of Flying Insect Prey

The applicability of species-specific primers to study feeding interactions is restricted to those ecosystems where the targeted prey species occur. Therefore, group-specific primer pairs, targeting higher taxonomic levels, are often desired to investigate interactions in a range of habitats that do not share the same species but the same groups of prey. Such primers are also valuable to study the diet of generalist predators when next generation sequencing approaches cannot be applied beneficially. Moreover, due to the large range of prey consumed by generalists, it is impossible to investigate the breadth of their diet with species-specific primers, even if multiplexing them. However, only few group-specific primers are available to date and important groups of prey such as flying insects have rarely been targeted. Our aim was to fill this gap and develop group-specific primers suitable to detect and identify the DNA of common taxa of flying insects. The primers were combined in two multiplex PCR systems, which allow a time- and cost-effective screening of samples for DNA of the dipteran subsection Calyptratae (including Anthomyiidae, Calliphoridae, Muscidae), other common dipteran families (Phoridae, Syrphidae, Bibionidae, Chironomidae, Sciaridae, Tipulidae), three orders of flying insects (Hymenoptera, Lepidoptera, Plecoptera) and coniferous aphids within the genus Cinara. The two PCR assays were highly specific and sensitive and their suitability to detect prey was confirmed by testing field-collected dietary samples from arthropods and vertebrates. The PCR assays presented here allow targeting prey at higher taxonomic levels such as family or order and therefore improve our ability to assess (trophic) interactions with flying insects in terrestrial and aquatic habitats.     

Egomotion estimation with optic flow and air velocity sensors

We develop a method that allows a flyer to estimate its own motion (egomotion), the wind velocity, ground slope, and flight height using only inputs from onboard optic flow and air velocity sensors. Our artificial algorithm demonstrates how it could be possible for flying insects to determine their absolute egomotion using their available sensors, namely their eyes and wind sensitive hairs and antennae. Although many behaviors can be performed by only knowing the direction of travel, behavioral experiments indicate that odor tracking insects are able to estimate the wind direction and control their absolute egomotion (i.e., groundspeed). The egomotion estimation method that we have developed, which we call the opto-aeronautic algorithm, is tested in a variety of wind and ground slope conditions using a video recorded flight of a moth tracking a pheromone plume. Over all test cases that we examined, the algorithm achieved a mean absolute error in height of 7% or less. Furthermore, our algorithm is suitable for the navigation of aerial vehicles in environments where signals from the Global Positioning System are unavailable.     

Spatial and temporal distribution patterns of flying Diptera

The temporal and spatial distributions of flying Diptera have been studied using suction traps hung at three levels and two distances from an emergent hedgerow tree. Most families were far more abundant in 1987 than in 1986, but patterns of spatial and temporal abundance were basically the same. Most families showed a marked concentration close to the tree. Scatopsidae accumulated away from the tree in large numbers. Most families concentrated at the mid-level, except the Anisopodidae and Mycetophilidae (top and bottom levels, respectively). Vertical distributions showed discontinuities for almost all families. It is suggested that the emergent hedgerow tree plays a very important part in the life of flying insects.     

华北地区昆虫秋季迁飞的雷达观测

1999年9－10月利用昆虫雷达对华北地区昆虫秋季回迁的观测表明,秋季迁飞主要发生于19：00－05：00时,飞行高度一般在300－2000m。迁飞方向与风向一致,多为从东北向西南方向迁飞。运行速度和飞行高度成正相关,迁飞个体的平均运行速度由392m高度的4.8m/s增加到2000m处的27.8m/s。空中灯光诱捕表明,甜菜夜蛾和棉铃虫为优势迁飞种类。     

The insect faunas of pear and apple orchards and the effect of windbreaks on their distribution

Two coir netting windbreaks, each 7 m tall and 46 m long, were erected in a pear and an apple orchard, to try to increase the number of insect pollinators present at blossom time. The flying insects were sampled with suction traps and insects visiting trees were sampled by examining marked clusters of blossom. The mean aerial density was calculated for each of the forty-two taxa, mainly families, identified from the aerial population. There were about 50% more insects flying in the pear than in the apple orchard. The distribution of flying insects was greatly affected by the windbreaks, and the sheltered zone contained three times as many of most species, and more than three times as many Chironomidae, Psychodidae and Bibionidae as elsewhere. Small flies were the most abundant insects in the air, in both orchards. In the pear orchard large insects comprised only about 7 % of the total aerial population, of which honeybees constituted 0–7% and wild bees 0–3%. By contrast, large insects accounted for a greater proportion of the population on blossom. The reasons for this are discussed. On apple blossom Syrphidae and honeybees were the insects most often seen, and on pear blossom Bibionidae and Mycetophilidae.     

Visual ground pattern modulates flight speed of male Oriental fruit moth Grapholita molesta

Insects flying in a horizontal pheromone plume must attend to visual cues to ensure that they make upwind progress. Moreover, it is suggested that flying insects will also modulate their flight speed to maintain a constant retinal angular velocity of terrestrial contrast elements. Evidence from flies and honeybees supports such a hypothesis, although tests with male moths and beetles flying in pheromone plumes are not conclusive. These insects typically fly faster at higher elevations above a high‐contrast ground pattern, as predicted by the hypothesis, although the increase in speed is not sufficient to demonstrate quantitatively that they maintain constant visual angular velocity of the ground pattern. To test this hypothesis more rigorously, the flight speed of male oriental fruit moths (OFM) Grapholita molesta Busck (Lepidoptera: Tortricidae) flying in a sex pheromone plume in a laboratory wind tunnel is measured at various heights (5–40 cm) above patterns of different spatial wavelength (1.8–90°) in the direction of flight. The OFM modulate their flight speed three‐fold over different patterns. They fly fastest when there is no pattern in the tunnel or the contrast elements are too narrow to resolve. When the spatial wavelength of the pattern is sufficiently wide to resolve, moths fly at a speed that tends to maintain a visual contrast frequency of 3.5 ± 3.2 Hz rather than a constant angular velocity, which varies from 57 to 611° s^?1. In addition, for the first time, it is also demonstrated that the width of a contrast pattern perpendicular to the flight direction modulates flight speed.     

Bioassay systems for the pyrethrins; water-base sprays against Aëdes aegypti L. and other flying insects

A bioassay system for the pyrethrins is discussed in which the quantity of toxicant that flying insects accumulate is measured, as well as the responses of the insects. The determination of spray pick-up from water-base sprays is described, and the use of primary responses, such as activation and paralysis, in biological assays is advocated. The test system is discussed in detail for Aëdes aegypti L. and the special characteristics of some other species of flying insects noted. It is considered that the location of the site of action of the pyrethrins under these conditions is at the peripheral nervous system of the test insects.     

Seasonal pattern of abundance of honeyeaters and their resources in heathland areas near Sydney

Abundances of honeyeaters, flowers and flying insects, the daily nectar-energy production per flower and the average size of flying insects were estimated every three weeks for circles of radius 20 m located in three heathland areas near Sydney. Seasonal fluctuations in honeyeater density showed no apparent relationship with seasonal fluctuations in nectar-energy productivity or in biomass of flying insects. Variation between circles in honeyeater density was also unrelated to spatial variation in energy productivity and insect biomass. The relatively low incidence of nectar-feeding and high incidence of flying exhibited by birds observed during troughs in nectar-energy production suggest that many of these birds are transient and that their density may consequently be unrelated to local conditions.     

The survival of ingested Serratia marcescens in houseflies (Musca domestica L.) after electrocution with electric fly killers

Electric fly killers (EFKs) are commonly used to control flying insects that enter food establishments. For establishment of the incidence of pathogen-bearing insects in food establishments, insect samples obtained from EFK trays could be used. The principal difficulty with this approach is that the survival time of microorganisms on or within insect corpses after electrocution is unknown. This study determined the survival of Serratia marcescens (as a representative of the enteric bacteria) within houseflies following their electrocution by a commercial EFK. S. marcescens was successfully ingested by houseflies and survived on and within the corpses after electrocution for up to 5 weeks. Maximal levels of bacteria were recovered 24 h postelectrocution. The study also demonstrates the ability of ingested S. marcescens to out-compete resident microbial flora within houseflies. The findings are intended to pave the way for further research to determine the incidence of pathogen-laden flying insects in food establishments.     

Modulation of whitefly take-off and flight orientation by wind speed and visual cues

The effect of different wind speeds on take-off and flight orientation of the sweetpotato whitefly, Bemisia tabaci Gennadius (Homoptera: Aleyrodidae), was studied in the presence of a green visual stimulus which reflected 550 ± 10 nm light, or a white stimulus of the same intensity. When the white light was present, take-off was negatively correlated with wind speed. Analysis of the flight tracks of whiteflies in 0, 15 and 30 cm/s wind with the white light present showed that flight was not directed toward the stimulus in zero wind, and that insects were carried downwind as the wind increased. Net displacement downwind was significantly slower than the wind speed, indicating that B. tabaci can control its rate of displacement relative to its surroundings, and is not always passively transported by the wind. In the presence of the green visual stimulus, take-off and flight behaviour of B. tabaci was markedly different to that observed in the presence of the white light. Taking off was more likely and whiteflies made upwind orientated flights, landing on the illuminated section of the screen when it reflected green light. At all wind speeds tested, the mean ground speeds of B. tabaci were approximately 20 cm/s whether the insects were flying upwind or downwind. This uniformity of ground speed regardless of the changing effects of wind-induced drift in different directions strongly suggests that whiteflies actively control their ground speed using visual flow fields in a manner similar to all other flying insects examined thus far.     

NUMERICAL SIMULATION OF THE PATHWAYS OF MIGRATING INSECTS

Abstract  Based on boundary layer meteorology and behavioural ecology of insect migration, a numerical TRA model was established. The spatial and temporal resolutions of the standard meteorological data are far insufficient for insect migration studies; so, a one dimension TKE closure scheme (E-ε scheme) was adopted to simulate the wind and temperature profiles en route based on the conventional surface and 850 hPa wind and temperature data of Chinese National Meteorological Bureau. Then the migrating behaviour of insects was parameterized as some proper mathematical expressions to determine their timing and their flight height, speed and direction from the wind and temperature profiles simulated by the PBL sub-model, and so their flight pathways could be estimated by a simple algorithm. Finally, the hourly episodes of the airborne migrants were output which consists of the latitude, longitude and flying altitude. The TRA model was verified by mark-release-recapture studies of Mythimna separata, Loxostege sticticalis , and Agrotis ipsilon . The results suggest that the parameterizing scheme of migratory behaviour, the numerical simulation scheme of wind and temperature in PBL and the TRA procedure developed in this paper should be reasonable and feasible. This model provides a useful tool for inter-regional forecast of migratory insect pests.     

Nocturnal insects use optic flow for flight control

To avoid collisions when navigating through cluttered environments, flying insects must control their flight so that their sensory systems have time to detect obstacles and avoid them. To do this, day-active insects rely primarily on the pattern of apparent motion generated on the retina during flight (optic flow). However, many flying insects are active at night, when obtaining reliable visual information for flight control presents much more of a challenge. To assess whether nocturnal flying insects also rely on optic flow cues to control flight in dim light, we recorded flights of the nocturnal neotropical sweat bee, Megalopta genalis, flying along an experimental tunnel when: (i) the visual texture on each wall generated strong horizontal (front-to-back) optic flow cues, (ii) the texture on only one wall generated these cues, and (iii) horizontal optic flow cues were removed from both walls. We find that Megalopta increase their groundspeed when horizontal motion cues in the tunnel are reduced (conditions (ii) and (iii)). However, differences in the amount of horizontal optic flow on each wall of the tunnel (condition (ii)) do not affect the centred position of the bee within the flight tunnel. To better understand the behavioural response of Megalopta, we repeated the experiments on day-active bumble-bees (Bombus terrestris). Overall, our findings demonstrate that despite the limitations imposed by dim light, Megalopta-like their day-active relatives-rely heavily on vision to control flight, but that they use visual cues in a different manner from diurnal insects.     

In situ measurement of calling metabolic rate in an Australian mole cricket, Gryllotalpa monanka

Examination of the energetics of sound production usually requires measurement of species that will produce normal calls under unnatural circumstances. Such measurements are potentially compromised by stress-related changes in calling input (through a reduction in calling effort) or output (through forced use of sub-optimal singing burrows). To determine if such measurements are indeed affected by abstraction from a natural setting, we measured the energetics of song production in undisturbed mole crickets Gryllotalpa monanka and employed a new approach where the animal's singing chamber replaces the respirometry chamber normally used in studies of this type. It was therefore possible to measure metabolic rate (MR) of calling crickets in situ for animals within self-constructed burrows under natural conditions. Calling MR measured under these conditions averaged 13.5-fold higher than standard MR and 2.2-fold higher than MR measured during burrowing in the lab. The calling MR of G. monanka was similar to that measured for other calling insects, and to endothermic insects, but was only 10% of that allometrically predicted for a similarly sized insect (0.89 g) during flight. A male mole cricket is estimated to consume 5.9 ml of oxygen during construction of a calling burrow and a 1-h calling bout; by comparison, a flying female would consume a similar volume in less than 6 min.     

In situ measurement of calling metabolic rate in an Australian mole cricket, Gryllotalpa monanka.

Examination of the energetics of sound production usually requires measurement of species that will produce normal calls under unnatural circumstances. Such measurements are potentially compromised by stress-related changes in calling input (through a reduction in calling effort) or output (through forced use of sub-optimal singing burrows). To determine if such measurements are indeed affected by abstraction from a natural setting, we measured the energetics of song production in undisturbed mole crickets Gryllotalpa monanka and employed a new approach where the animal's singing chamber replaces the respirometry chamber normally used in studies of this type. It was therefore possible to measure metabolic rate (MR) of calling crickets in situ for animals within self-constructed burrows under natural conditions. Calling MR measured under these conditions averaged 13.5-fold higher than standard MR and 2.2-fold higher than MR measured during burrowing in the lab. The calling MR of G. monanka was similar to that measured for other calling insects, and to endothermic insects, but was only 10% of that allometrically predicted for a similarly sized insect (0.89 g) during flight. A male mole cricket is estimated to consume 5.9 ml of oxygen during construction of a calling burrow and a 1-h calling bout; by comparison, a flying female would consume a similar volume in less than 6 min.     

Terrestrial–aquatic transitions: Local abundances and movements of mature female caddisflies are related to oviposition habits but not flight capability

1. Movement behaviours of adult aquatic insects can produce distinct spatial distribution patterns. Studies of adult abundance with distance away from water bodies are common and may invoke flight capability to explain species differences. In contrast, distribution patterns along river channels are poorly described, but are no less important for understanding population dynamics. Longitudinal patterns in adult abundance along short river lengths may differ between sexes and at different life stage transitions between aquatic and terrestrial environments, i.e. at emergence and oviposition. Flight capability is unlikely to influence longitudinal patterns created at emergence, but may influence local abundances of mature females seeking to lay eggs. We tested hypotheses about how local abundances of mature females might differ according to oviposition habits and flight capability.
2. We surveyed abundances of mature female caddisflies at adjacent riffle–pool pairs along short river lengths with homogeneous riparian cover. Our survey included nine species in three families (Hydrobiosidae, Leptoceridae, Hydropsychidae), which encompassed multiple different oviposition habits and a range of wing sizes and shapes. Several of the species oviposit preferentially in riffles. Accordingly, we tested for differences in female abundance between channel units (adjacent riffle–pool pairs). We also tested whether females attained higher abundances in some places along channels than others (i.e. over larger spatial scales and regardless of channel unit) which imply movements along the channel and aggregation in some locations. Wing morphology was used as a proxy measure of flight capability and included measures of wing span, area, aspect ratio and the second moment of wing area.
3. Three distinctly different distribution patterns of mature female caddisflies were identified. The abundance of three species varied over larger scales only (multiple channel units). Six species that oviposit preferentially in riffles had higher female abundances at riffles than pools, but for only one did abundances also vary over larger scales. There was no association between these different patterns and measures of wing morphology, after removing metrics that were correlated and that differed systematically between taxonomic families. However, we could not reject the hypothesis that some aspect of flight behaviour may have contributed to observed patterns.
4. The diverse but distinct distributions of mature female caddisflies we observed along short channel lengths are novel and suggest that species differ in their propensity for movement along streams, which could have consequences for local densities of eggs and juveniles in the aquatic environment. The degree to which population sizes are coupled across the terrestrial-to-aquatic transition is rarely investigated in aquatic insects and may provide fresh insight into sources of spatial variation within populations. Similarly, a more nuanced approach to research on the flight of aquatic insects, including age- and sex-specific phenomena, may provide greater insight into the diverse ecological functions and consequences of movement.