A laboratory technique for examining the flight activity of insects under controlled environment conditions

A technique was developed for studying the flight activity of the, black fly,Simulium arcticum, under controlled environment conditions. Wind speed, light, temperature and humidity could be controlled and monitored in the flight chambers. Accurate measurement and recording of individual insect activity were achieved with a high-sensitivity video recording and monitoring system. The controlled-environment monitoring system is adaptable for investigations of the environmental behaviour and physiology of various insect species.     

Activity and distribution of the mushroom phorid fly,Megaselia halterata,in and around commercial mushroom farms

The mushroom phorid fly, Megaselia halterata (Wood) (Diptera: Phoridae), is a key pest in mushroom farming in most parts of the world. Studies on the mushroom phorid fly have focused on its life history within mushroom growing houses, but little is known about the fly's activity outside mushroom growing houses. In this study, daily activity and distribution of adult M. halterata in the areas surrounding mushroom growing houses was studied using yellow sticky traps. Results suggest that M. halterata focuses its flight activity over turf areas rather than windbreaks and spent compost piles, possibly for mating purposes. Our study found no evidence of M. halterata breeding in turf areas surrounding mushroom growing houses. In addition, flight activity is highest in the afternoon until midnight at higher temperatures, yet at lower temperatures activity ceases after sunset. Establishing temperature and daylight thresholds for M. halterata flight activity may be useful in developing integrated pest management (IPM) tactics for this species. The most successful IPM tool that mushroom growers use at present is fly exclusion. Exclusion can be improved by focusing farm operations around temperature and daylight thresholds when fly activity is at its lowest.     

Flight activity ofProstephanus truncatus (Horn) (Coleoptera: Bostrichidae) in relation to population density,resource quality,age, and sex

Higher flight activity has been observed in aged, high-density cultures ofProstephanus truncatus (Horn) (Coleoptera: Bostrichidae), but adults in new, lowdensity culture jars showed less flight activity. In order to understand this change in behavior, the effects of population density, age, resource quality, and sex on the flight ofP. truncatus were studied in a wind tunnel. While an immediate density on the release platform had no significant effect on flight, beetles from high-density cultures were more inclined to fly than those from low-density cultures. Resource quality exerted a major influence on flight; insects in food suitable for boring and oviposition seldomly exhibited flight, however, when food was absent or of inferior quality for boring and oviposition, the dominant behavior was flight. Also, insects maintained for a week in food suitable for boring and oviposition were less ready to fly than those maintained in food unsuitable for boring and oviposition. The optimum age range for flight activity was before the peak of reproduction and insects rarely flew before 4 days or after 32 days of emergence. There were no significant differences between the flight activity of males and that of females. Based on these results, we conclude that age and resource quality are major influences on the flight activity ofP. truncatus and a hypothesis is proposed in which reproductively active male and female beetles disperse from habitats of low resource quality to those that support their reproductive behavior. The practical implications of these results and the possible role of the male-produced aggregation pheromone are discussed.     

黑水虻幼虫蛋白及其酶解产物的抗氧化活性研究

为了探究黑水虻Hermetia illucens幼虫蛋白及酶解产物的抗氧化活性,以鲜活黑水虻幼虫冻虫为原料,采用碱提酸沉法提取黑水虻蛋白,通过碱性蛋白酶、菠萝蛋白酶、风味蛋白酶、木瓜蛋白酶对其蛋白质溶液进行酶解,分别从2,2′-联氮-双-3-乙基苯并噻唑啉-6-磺酸（ABTS）自由基、羟自由基、1,1-二苯基-2-三硝基苯肼（DPPH）自由基3个方面对黑水虻蛋白及其酶解液的抗氧化能力进行测定。结果表明：黑水虻幼虫蛋白及4种酶解后的蛋白肽具有较好的抗氧化活性,其中黑水虻幼虫蛋白对ABTS自由基、羟自由基、DPPH自由基的半抑制浓度（IC50）分别为2.91、0.232、0.764 mg/mL,而酶解过后的蛋白肽具有更强的抗氧化活性,对ABTS自由基、羟自由基、DPPH自由基的最低半抑制浓度（IC50）分别为0.295、0.082、0.354 mg/mL。本研究初步证明了酶解制备抗氧化肽的可行性,为昆虫蛋白资源的利用和无抗饲料的研制提供了新的研究思路。     

Foraging in male bumblebees (Bombus lucorum L.): maximizing energy or minimizing water load?

Summary The O₂, CO₂, and H₂O exchange of single flying male bumblebees (Bombus lucorum and B. terrestris) were measured simultaneously. A respiratory quotient RQ=1 was found for all activities investigated (torpor-flight). The dependence of respiratory CO₂ production in flight on body-weight was measured: for a 220-mg male bumblebee it amounts to 24.5 mg CO₂/h (=56.4 ml O₂/g·h). The corresponding evaporative water loss amounts to 6 mg H₂O/h. Males tranferred to a climatic test chamber and conditioned to artificial flower feeders started to fly, after a few days of acclimatization, in typical scent-marked flight-paths. The daily pattern of flight activity was recorded: the mean total time in flight amounts to 244 min, and the corresponding daily flight length is about 17 km. At 20°C and 50% relative humidity (RH) a daily uptake of 180 l ( 220mg) of 50% sugar solution was measured, equal to the mean body weight of the male bumblebees. Since the body weight remains constant on consecutive days a 24-h energy- and water-budget could be calculated. The energy-budget is balanced; the activities observed can be fuelled with the sugar available. About 70% of the energy is used for the 4 h of flight activity. With the daily nectar volume 110 mg of water is ingested; in the oxidation of 110 mg sugar, 66 mg of metabolic water is produced and 40 mg water is dissipated by the evaporative water-loss. Thus, to have a balanced water-budget, 136 mg of water must be voided in 24 h, which equals the total body-water of the bumblebees. Nectar is a nutrient of high water content which not only provides the sugar necessary for activity but also, in most circumstances, an excess of water. The effect of this high water load in limiting daily activity is discussed and compared with the water- and osmoregulation of hummingbirds. The strategy of maximizing energy for a male bumblebee must be one of minimizing water load.     

BEHAVIORAL EVIDENCE FOR FRUIT ODOR DISCRIMINATION AND SYMPATRIC HOST RACES OF RHAGOLETIS POMONELLA FLIES IN THE WESTERN UNITED STATES

The recent shift of Rhagoletis pomonella (Diptera: Tephritidae) from its native host downy hawthorn, Crataegus mollis, to introduced domesticated apple, Malus domestica, in the eastern United States is a model for sympatric host race formation. However, the fly is also present in the western United States, where it may have been introduced via infested apples within the last 60 years. In addition to apple, R. pomonella also infests two hawthorns in the West, one the native black hawthorn, C. douglasii, and the other the introduced English ornamental hawthorn, C. monogyna. Here, we test for behavioral evidence of host races in the western United States. through flight tunnel assays of western R. pomonella flies to host fruit volatile blends. We report that western apple, black hawthorn, and ornamental hawthorn flies showed significantly increased levels of upwind‐directed flight to their respective natal compared to nonnatal fruit volatile blends, consistent with host race status. We discuss the implications of the behavioral results for the origin(s) of western R. pomonella, including the possibility that western apple flies were not introduced, but may represent a recent shift from local hawthorn fly populations.     

Rapid and repeatable shifts in life‐history timing of Rhagoletis pomonella (Diptera: Tephritidae) following colonization of novel host plants in the Pacific Northwestern United States

Host shifts of phytophagous insect specialists to novel plants can result in divergent ecological adaptation, generating reproductive isolation and potentially new species. Rhagoletis pomonella fruit flies in eastern North America underwent a host shift ~160 ya from native downy hawthorn (Crataegus mollis) to introduced, domesticated apple (Malus domestica). Divergent selection on diapause phenology related to the earlier fruiting time of apples versus downy hawthorns resulted in partial allochronic reproductive isolation between the fly races. Here, we test for how rapid and repeatable shifts in life‐history timing are driving ecological divergence of R. pomonella in the Pacific Northwestern USA. The fly was introduced into the region via larval‐infested apples 40–65 ya and now attacks native black hawthorn (Crataegus douglasii) and introduced ornamental hawthorn (Crataegus monogyna), in addition to early‐ and late‐maturing apple varieties in the region. To investigate the life‐history timing hypothesis, we used a field‐based experiment to characterize the host‐associated eclosion and flight activity patterns of adults, and the feeding times of larvae at a field site in Vancouver, Washington. We also assessed the degree to which differences in host‐fruiting time generate allochronic isolation among apple‐, black hawthorn‐, and ornamental hawthorn‐associated fly populations. We conclude that host‐associated fly populations are temporally offset 24.4% to 92.6% in their seasonal distributions. Our results imply that R. pomonella possesses the capacity for rapid and repeatable shifts in diapause life history to match host‐fruiting phenology, which can generate ecologically based reproductive isolation, and potentially biodiversity in the process.     

Flight Directions in the Migratory Butterfly Pieris brassicae: Results from Semi-natural Experiments

A recently developed experimental set-up has been used to test the constancy of flight direction in the large white butterfly Pieris brassicae, originally collected in Northern Germany. The flight direction, tested with virgin females, depends on the developmental mode. Butterflies from diapause pupae fly north, those from subitan pupae fly south. The constancy of the flight direction has been examined in detail with the spring generation over a period of 14d. The preferred direction does not change with advancing age and is essentially not influenced by temperature or by the sun's azimuth. The flight activity, however, increases with temperature, but this is concealed by an intrinsic response resulting in a diurnal peak of flight activity at 10.00–11.00 hours, long before the maximum temperature of the day is reached. Whether the observed direction can be considered as a parameter of migration behaviour is discussed. It is shown that migration in Pieris brassicae does not end with reproductive maturity but lasts throughout the entire imaginal stage.     

Energy use by the mountain pine beetle (Coleoptera: Curculionidae: Scolytinae) for dispersal by flight

The mountain pine beetle Dendroctonus ponderosae Hopkins is a major native pest of Pinus Linnaeus (Pinaceae) in western North America. Host colonization by the mountain pine beetle is associated with an obligatory dispersal phase, during which beetles fly in search of a suitable host. Mountain pine beetles use stored energy from feeding in the natal habitat to power flight before host colonization and brood production. Lipids fuel mountain pine beetle flight, although it is not known whether other energy sources are also used during flight. In the present study, we compare the level of energy substrates, proteins, carbohydrates and lipids of individual mountain pine beetles flown on flight mills with unflown control beetles. We use a colorimetric method to measure the entire metabolite content of each individual beetle. The present study reveals that mountain pine beetles are composed of more protein and lipid than carbohydrate. Both female and male mountain pine beetles use lipids and carbohydrates as energy sources during flight. There is variation between sexes, however, in the energy substrates used for flight. Male mountain pine beetles use protein, in addition to lipids and carbohydrates, to fuel flight, whereas protein content is not different between flown and control females.     

Factors influencing flight potential of Choristoneura conflictana

Abstract Factors affecting the flight potential of Choristoneura conflictana, an insect that undergoes cyclical changes in population density, are investigated using computer‐linked flight mills. Female and male moths are flown for a 12‐h period and the longest single flight and total flight distance of each moth is recorded. After flight bioassays, moths are subjected to lipid extraction with a soxhlet apparatus to determine the effect of body lipid content on flight. Larger C. conflictana moths fly farther than smaller moths. Female C. conflictana fly farther than male moths but the effects of both mating status and moth age on the distance flown are dependent on sex. Mated females fly farther than virgins and older females fly farther than young females but these relationships do not occur in males. Body lipid content affects the distance flown by both females and males through a significant interaction with sex and age. The factors examined in this study will vary with density in natural populations and are predicted to be important indicators of flight potential and dispersal in this species.     

Dynamic body acceleration increases by 20% during flight ontogeny of greylag geese Anser anser

Despite our knowledge of the biophysical and behavioural changes during flight ontogeny in juvenile birds, little is known about the changes in the mechanical aspects of energy expenditure during early flight development, particularly in migratory species. Here, we investigate in a unique experimental setup how energy expended during flights changes over time beginning with early ontogeny. We calculate overall dynamic body acceleration (ODBA) as a proxy for energy expenditure in a group of hand raised greylag geese Anser anser trained to fly behind a microlight aircraft. We propose two potential hypotheses; energy expenditure either increases with increasing physiological suitability (the ‘physical development hypothesis’), or decreases as a result of behavioural improvements mitigating flight costs (the ‘behavioural development hypothesis’). There was a significant temporal increase of flight duration and ODBA over time, supporting the ‘physical development hypothesis’. This suggests that early on in flight ontogeny behavioural development leading to flight efficiency plays a weaker role in shaping ODBA changes than the increased physical ability to expend energy in flight. We discuss these findings and the implications of flight development on the life history of migratory species.     

Differences in Morphometry and Activity among Tabanid Fly Assemblages in an Andean Tropical Montane Cloud Forest: Indication of Altitudinal Migration?

Evidence suggests that variations along ecological gradients shape organism traits such as behavior or morphometry. We studied the effect of altitude on the flight activity of tropical tabanid fly assemblages of one species of Stypommisa Enderlein along a 1 km altitudinal gradient on the northwestern slopes of the Ecuadorian Andes. Our objectives were as follows: (1) to test the hypothesis that highland individuals present larger flight body structures; and (2) to compare the flight activity patterns of flies’ assemblages among altitudes and correlate it with weather factors. We sampled specimens in Malaise traps at 1180, 1680 and 2180 m of altitude from 0600 to 1830 h for 20 d at each site. Seven weather variables were measured every hour and flight activity was inferred from relative tabanid fly abundances/hour in traps. We measured morphometrical parameters that included tabanid fly body size, thorax volume, wing area and wing loading. Flight activity patterns revealed a bimodal distribution at 1680 m, and two asynchronous unimodal distributions, one at 1180 and one at 2180 m. GLM analyses revealed that temperature, mist and rainfall were the best predictors of fly activity differences among altitudes. Morphometrical analyses showed that body size and thorax volume increased with increasing altitude. Synchronous groups of flies at different altitudes (those between 1180–1680_(pm) m, and 1680_(am)–2180 m) were morphologically similar, suggesting that flies could be capable of migrating from highlands to lowlands at defined hours of the day depending on forest weather conditions.     

Flight activity of three Spodoptera spp., Spodoptera litura, S. exigua and S. depravata, measured by flight actograph

Flight activities of three Spodoptera species were measured by the aid of flight actograph: S. litura and S. exuiga being regarded as long‐distance migratory insects, and S. depravata being non‐migratory and diapause‐inducible species. In all species tested, flight activities were observed only in scotophase, males showed far higher activities than females, being several times higher at the time of maximum flight activity, which was observed within 2 days after adult eclosion. Total flight activity in males was highest in S. litura, some being flyable even 12 days after eclosion, followed by S. exigua being one‐third compared to the former species, while in S. depravata flight activity was nearly half of that of the second species and most ceased to fly within a week after eclosion. There occurred species‐specific daily rhythms in flight activity during respective scotophase. In S. litura, both females and males exhibited a peak of flight activity shortly after light‐off and exhibited the second flight activity in late scotophase, the females slightly but the males more actively compared to early scotophase. In S. exigua, both sexes did not respond to light‐off, did not show a peak of flight activity in early scotophase, whereas males, but not females prominently increased activity toward the end of scotophase. In S. depravata, both sexes exhibited a peak of flight activity in early scotophase, and the males revived flight activity, being maximum shortly before light‐on, but the females did not show a clear rhythm in flight activity. These features observed in flight activity were discussed in relation with migratory capability.     

Vulture flight behavior and implications for aircraft safety

Growing vulture populations represent increasing hazards to civil and military aircraft. To assess vulture flight behavior and activity patterns at the Marine Corps Air Station in Beaufort, South Carolina, we equipped 11 black vultures (Coragyps atratus) and 11 turkey vultures (Cathartes aura) with solar-powered Global Positioning System (GPS) satellite transmitters during a 2-year study (1 Oct 2006–30 Sep 2008). Turkey vultures had larger seasonal home ranges than did black vultures, and 2 turkey vultures made round-trips to Florida. Black vultures consistently spent less time in flight (8.4%) than did turkey vultures (18.9%), and black vultures flew at higher altitudes than did turkey vultures in all seasons except summer when altitudinal distributions (above ground level) did not differ. Although we recorded maximum altitudes of 1,578 m for black vultures and 1,378 for turkey vultures, most flights were low altitude. A matrix of vulture flight altitude versus time of day revealed that >60% of vulture flight activity occurred from 4 hr to 9 hr after sunrise at altitudes below 200 m. Continuation of aggressive harassment coupled with flexible training schedules to avoid times and altitudes of high vulture activity will decrease hazards to aircraft posed by these birds. © 2011 The Wildlife Society.     

Habitat-specific flight period in the cherry fruit fly Rhagoletis cingulata (Loew) (Diptera: Tephritidae)

Flight periods of the cherry fruit fly, Rhagoletis cingulata (Loew), were compared in the major sweet and tart cherry-growing regions of Michigan, among neglected orchards, managed orchards, and natural areas containing the ancestral host, black cherry. Traps were deployed from early June to late September 2005 and 2006. Captures indicated that cherry fruit fly has an early flight (June-July) in neglected orchards, a mid-season flight peaking immediately after harvest (June-August) in managed orchards, and an extended flight covering most of the season (June-September) in natural areas. We found that the period of fruit infestation mirrored the flight period in neglected and managed orchards. In natural areas, we found infestation late in the season only. The relative emergence periods for adults reared from pupae collected from the three habitats and maintained under the same conditions coincided with adult flight periods for each habitat. We also studied factors related to fruit availability that may have a role in shaping the flight periods. Fruit abundance decreased rapidly early in the season in neglected orchards, whereas in managed orchards, fruit left after harvest remained on the trees until late August. Measurements of fruit size and skin firmness revealed that fly activity in neglected and managed orchards began immediately after fruit increased in size and skin firmness decreased, whereas in natural areas, the flight began before fruit matured. In managed orchards, fruit harvest and insecticide sprays likely maintain the late flight period of resident fly populations by preventing the use of fruit earlier in the season. However, a significant proportion of these resident flies may still emerge before harvest and increase the risk of costly fruit infestation.     

Flight activity in the parasitoid waspNasonia vitripennis (Hymenoptera: Pteromalidae)

Flight activity in females of the parasitoid wasp Nasonia vitripennis(Walker) was examined by measuring still-air tethered flight. There was a large amount of variation among females in flight duration. The longest single flight (with no pauses of more than 5 s) was more than 2 h long. Mating status had a significant and large effect on flight: mated females flew twice as long as virgin females. There also was a slight but significant effect of age on flight, with 3-day-old females being less likely to fly than 1-day-old females. Flight duration was not affected by prior exposure to other females, to honey, or to a low or a high host density.     

Investigation on flight and walking activity of Oomyzus sokolowskii Kurdjumov (Hymenoptera: Eulophidae) at different generations under laboratory conditions

The flight and walking activities of second, third, fourth, fifth, sixth and seventh generations of Oomyzus sokolowskii reared on Plutella xylostella L. pupas were compared under laboratory conditions. The parasitoid wasps were obtained by collecting pupa masses of diamondback moth from cabbage fields. The experiments of flight and walking activities were carried out using black PVC tubes and the percentages of inactive, flying and walking wasps to light were calculated. Results showed a reduced flight and walking activities and increased percentage of inactive wasps with the increasing number of generations. In second- third- and fourth-generation average, 52.1 ± 1.27 and 29.6 ± 1.51 of population had flight and walking ability, respectively, and 17.1 ± 1.55 of them were not able to fly and walk. The highest percentage of inactive wasps was observed at seventh generation, whereas wasps showed significantly better flight and walking activities at second, third and fourth generations. Based on the results, flight and walking abilities of O. sokolowskii gradually decreased at continuous laboratory rearing.     

The effect of flight on reproduction in an outbreaking forest lepidopteran

Post‐flight reproductive investment by female insects may be limited as a result of a trade‐off in resource allocation between flight and reproduction. Outbreaking forest pests reduce their habitat quality as a result of severe defoliation when population densities are high. Female relocation to better‐quality habitats can increase offspring survival but reduce their reproductive fitness through flight. In the present study, the effect of flight on the capacity of female Choristoneura conflictana (Walker) (Lepidoptera: Tortricidae) to mate and produce eggs is examined. Females are flown on flight mills, and the subsequent reproductive capacity of each moth is assessed through measures of mating success and egg production. There is no effect of flight on commencement or the duration of mating. Although flight does not affect egg production directly, energy expenditure as a result of flight (as measured by weight loss) shows a negative correlation to potential fecundity, possibly indicating the resorption of eggs in some females. The effect of female size on fecundity is dependent on mating status, suggesting that energy allocated to reproduction is not dependent on flight treatment. Female moth longevity also has a significant effect on egg production but is dependent on flight and mating treatments. There is a relationship between energy expenditure to flight and reproduction in C. conflictana. Females that fly away from dense populations may produce fewer offspring, although this cost may be mitigated by improved offspring survivorship in less defoliated habitats.     

The adaptive significance of alpine melanism in the butterfly Parnassius phoebus F. (Lepidoptera: Papilionidae)

Summary The adaptive significance of alpine melanism, the tendancy for insects to become darker with increased elevation and latitude, was investigated using the butterfly Parnassius phoebus. The effects on temperature dependent activity of five components of overall wing melanism, as well as size, were examined. The components of wing melanism examined were the transparency of the basal hindwing and distal fore-wing areas, the width of the black patch in the basal hind-wing area and the proportion of black to white scales in that area, and the proportion of the distal fore-wing covered by predominantly black scaling.The body temperature of dead specimens was correlated with air temperature, solar radiation, the width of the black patch at the base of the wings, and the proportion of black to white scales at the base of the wings. The minimum air temperatures and solar radiation levels required for initiation of flight did not vary with wing melanism of P. phoebus, in contrast to the results found for Colias butterflies by Roland (1982). However, under environmental conditions suitable for flight initiation, males with a higher proportion of black to white scales in the basal area of the hind-wing, and wider basal black patches, spent a greater proportion of time in flight at low air temperatures and low insolation. Increased basal wing melanism was also associated with increased movement of males within a population. In contrast, melanism in the distal area of the wings has no effect on activities which are dependant on body temperature. The amount of time spent feeding did not vary with differences in wing melanism. I suggest that in dorsal basking, slow-flying butterflies (Parnassius) basal wing color affects body temperature primarily during flight (rather than while basking), such that butterflies with darker wing bases cool down less rapidly because they absorb more solar radiation during flight.     

Lipid economy,flight activity and reproductive behaviour in the speckled wood butterfly: on the energetic cost of territory holding

Insect flight is a highly energy demanding type of locomotion. In butterflies, males may locate females by different behavioural tactics. The tactics correspond to different flight types that, in turn, are assumed to reflect different energetic costs. Costs need to be considered to fully understand the pay‐offs of co‐existing alternative tactics relative to the environmental context and the phenotypes of the individuals. We addressed the issue in the speckled wood Pararge aegeria, in which males either adopt a territorial wait‐and‐fight tactic (i.e. territorial perching) in a sunlit patch on the forest floor, or a fly‐and‐search tactic to locate females in a wider area of the forest (i.e. patrolling). Perching corresponds to high frequency of take‐off flights and aerial combats with high levels of manoeuvrability and is assumed to be energetically more costly than longer, continuous flights at lower speed in patrollers. We tested the effect of different flight activity levels and of the behavioural tactics on lipid reserves and lipid use in males by laboratory and outdoor cage experiments. Low‐activity males that had access to honey water were capable of synthesizing lipids; their lipid reserves increased with age. The effect disappeared in males that actively flew in the outdoor cages. Lipid reserves decreased significantly faster in territorial perching males compared to non‐perching males, but resting metabolic rate did not differ between the alternative behavioural tactics. Territorial perching males had larger flight muscle ratio (i.e. thorax/body mass) than non‐perching males. We discuss the evidence of the physiological costs of perching relative to the co‐existence of perching and patrolling tactics.