Long flights in Cydia pomonella L. (Lepidoptera: Tortricidae) measured by a flight mill: influence of sex, mated status and age

Abstract. The flight capacity of Cydia pomonella L. (Lepidoptera: Tortricidae) was measured in the laboratory by using computer-linked flight mills. Codling moths showed a large variation in flight capacity between individuals. We defined arbitrarily a longest single flight (LSF) of more than 5 km as an index for long-flyers. About 16.7% of virgin and 10.0% of mated males and 20.0% of virgin and 7.4% of mated females undertook such flights. Based on the LSF and the total distance flown (TDF^we concluded that males and females have little or no difference in flight capacity and that both the within-and between-habitat flights are similar in number and magnitude for both sexes. In the field, females are therefore potentially able to undertake flights of up to 11 km, as reported for males by other authors. This ability was highest at ages of 2–7 days after emergence, i.e. the first third of their lifetime, for virgin and mated male moths and for virgin female moths. Mated females showed peak flight capacity between 1 and 3 days after eclosion, which corresponded with the major egg-laying period. Few long flights were undertaken before oviposition. These findings do not agree with the oogenesis flight syndrome described by other authors, and this theory is believed not to apply to C.pomonella. Our laboratory results are discussed in relation to field experiments in general and hypotheses are developed about the significance of long-flyers for this species.     

Sexual differences in the flight performance of the oriental fruit moth, Cydia molesta

Cydia molesta Busck (Lepidoptera: Tortricidae), a major pest of stone fruits and an increasingly important late season pest of apple, is predominately monitored by pheromone trapping of male moths. We investigated flight performance in relation to sex, age, and mating status using computer-linked flight mills, and also examined the relationship between female flight and reproduction. The crepuscular flight pattern of the experimental moths in relation to photointensity was very similar to that reported from field studies. Female moths significantly outperformed males in all measured flight parameters including total distance flown, distance of longest single flight, and velocity. The proportion of long-flying females (categorised as those completing an unbroken flight of greater than 1 km) was three to six times greater than that of males. Female flight performance was not related to mating status, but mated males displayed significantly greater flight than unmated males. The maximal flight period of mated females commenced on the third day after eclosion, following 30% egg deposition. Male flight was not significantly related to age. The data suggest that a limited proportion of the population, in particular females, may have the capacity to make inter-orchard flights. The limitations of monitoring C. molesta populations in apple orchards solely by pheromone trapping of males are discussed.     

Diel flight pattern and flight performance of Cactoblastis cactorum (Lepidoptera: Pyralidae) measured on a flight mill: influence of age, gender, mating status, and body size

Cactoblastis cactorum (Berg) (Lepidoptera: Pyralidae) is an invasive herbivore that poses a serious risk to Opuntia cacti in North America. Knowledge of the flight behavior of the cactus moth is crucial for a better understanding of natural dispersal, and for both monitoring and control. We used computer-linked flight mills to investigate diel flight activity and flight performance in relation to gender, age, mating status, and body size. Maximal flight activity for both mated and unmated moths occurred during twilight, whereas flight activity was low during photophase. The total distance flown and the number of initiated flights within a diel cycle were higher in both unmated and mated females than in males, but the longest single flight was similar in both genders. These findings suggest that pheromone trap captures of males likely indicate the simultaneous presence of females and that mated females might even be in areas where males are not detected yet. Flight performance heterogeneity was large, with a small portion of the population (both males and females) performing long unbroken flights, whereas the majority made short flights. Females had higher pupal and adult body size and shorter longevity than males. A few individuals, particularly young mated females, flying long distances may be important for active spread of a population and the colonization of new habitats. Implications of this study in the control of the cactus moth by using the sterile insect technique are discussed.     

Laboratory evaluation of flight activity of the common cutworm, Spodoptera litura (Lepidoptera: Noctuidae)

Abstract  The flight activity of Spodoptera litura in tethered conditions is evaluated using a computer-mediated flight-mill in the laboratory. The 3–4-day-old moths fly longer and farther than those of other ages. Male and female moths exhibit similar flight activity. Mating status does not influence the flight duration and distance of 2-day-old females. However, these two flight parameters with a 6-day-old mated female is significantly lower than that of unmated ones. The optimum temperature for flight ranged from 16–24°C, whereas the optimum RH ranged from 60%–100%. During 72-h period, the total flight duration and distance of 1-day-old male and female moths were 19.6 h (± 5.8) and 83.3 km (± 28.4), and 24.0 h (± 7.0) and 105.4 km (± 37.4), respectively. These results indicate that S. litura has a great potential to undertake long-distance migratory flights.     

Simultaneous monitoring of flight and oviposition of individual velvetbean caterpillar moths

ABSTRACT. A 'pivot' flight actograph was combined with a rolling oviposition surface to characterize the flight and oviposition behaviour of velvetbean caterpillar moths, Anticarsia gemmatalis Hubner. Tethered, caged control females laid significantly more eggs that those flown on the actograph. Mated females laid more eggs than unmated ones. However, mating did not affect longevity nor fight frequency and duration. Laboratory-reared and wild-type females also did not differ signficantly in longevity and flight. Both laboratory and wild mated females laid most of their eggs during the first 7 days whereas unmated ones delayed their oviposition. None of the female types produced a definite hourly, daily or lifetime pattern in flight frequency or duration. Of the 7672 recorded flights, about 3% were greater than 0.5 h. 'Long' flights (>0.5h) were made by some females before oviposition, as in colonization migration, but most interspersed flight with oviposition, as in extended search migration.     

粘虫飞行对生殖及寿命的影响

该文报道了粘虫Mythimna separata（Walker）成虫飞行后产卵、交配及寿命的研究结果。1日龄成虫飞行6 h、12 h、18 h、24 h后的产卵前期均显著短于对照的,其中飞行6 h、12 h的比对照的短2天以上,产卵量均比对照的高。对1～5日龄成虫分别飞行23.5 h后的研究结果表明,1日龄飞行的产卵前期和上述结果相一致。2～4日龄飞行的与对照的没有显著差异,但产卵量则随飞行日龄的延迟而逐渐减少。5日龄飞行的产卵前期显著延长,产卵量已不到对照的一半。所有经过飞行的成虫产卵高峰日比对照的早1天。不同日龄成虫飞行时间、距离与成虫产卵量的关系为：1～3日龄飞行时间、距离长的个体产卵量也高;但4～5日龄的成虫飞行时间与距离越长,其产卵量越少,表现出明显的卵子发生飞行拮抗症(oogenesis-flight syndrome)。除了5日龄飞行的成虫交配率有所下降以外,所有经过飞行的成虫产卵历期、交配率及寿命与对照的没有显著差异。最后,根据这些结果,对粘虫迁飞的起飞时期,迁飞在粘虫生殖、种群动态及成灾规律中的作用进行了讨论。     

Flight Studies on Photic Communication by the Firefly Photinus pyralis

Flash communication by the firefly Photinus pyralis was studiedin a stationary, simulated flight apparatus in which an individualof either sex could be "flown" and its flashing behavior andflight orientation recorded in response to photic stimulation.Males made long "flights" showing many of the characteristicsof their natural, female-seeking patrol flights. Males orientedtheir flight vectors towards light emitting diode (LED) flashesthat mimicked the responses of females to their patrol flashes.Females flew and responded to male-emulating LED flashes, makinga previously unknown early response followed by the typical2 sec delayed response characteristic of the dialoging perchedfemale, including abdominal aiming of the flash. Pairs consistingof males, in tethered flight, and females, perched, were runin an integrating sphere photometer, permitting the first determinationsof flash intensities of both sexes during courtship dialog.The implications of this work on thought about evolution ofphotic behavior in fireflies are considered.     

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.     

Monitoring codling moth (Lepidoptera: Tortricidae) with passive interception traps in sex pheromone-treated apple orchards

Male and female codling moth, Cydia pomonella (L.), were monitored with passive interception traps (PI-traps) in apple orchards treated with sex pheromone dispensers. The proportion of mated females recaptured by PI-traps was significantly higher than the proportion released after the release of both sexes into a codling moth-infested orchard. However, no significant difference occurred between the proportion of mated females recaptured and released when only females were released into uninfested orchards. Replicated nine-tree apple plots situated either on the edge or in the center ofpheromone-treated apple orchards were monitored with PI-traps during first moth flight in 1995 and during both flights in 1996. Moths caught on PI-traps were predominately males. The first male moths were captured 7-10 d before females during the first flight in both years. Initial capture of virgin and mated females on PI-traps coincided in 1995. Mated females were captured 14 d after the first virgin females in 1996. The mean proportion of females that were mated ranged from 32 to 55% during the first flight and 85 to 92% during the second flight. Moth catch and fruit injury were significantly higher in the edge versus the center plots. The numbers of total and female moths caught with PI-traps were significantly correlated with fruit injury for each generation. The percentage of female moths caught on PI-traps that were mated was 32% lower and the mean oocyte load of all females was 42% higher in a pheromone-treated apple orchard than in the untreated crabapple grove monitored during May and June 1997.     

Flight Performance of <Emphasis Type="Italic">Agrilus planipennis</Emphasis> (Coleoptera: Buprestidae) on a Flight Mill and in Free Flight

Computer-monitored flight mills were used to record the flight performance of the emerald ash borer, Agrilus planipennis Fairmaire. Flight performance of fed and unfed, and mated and unmated beetles of both sexes were recorded and compared. Mated females flew further per day and longer than unmated females or males. Mated females that were allowed to feed between flight periods flew an average of 1.3 km/day for four days; 10% flew more than 7 km/day. Measurements of free-flight speed using a mirror and a high-speed camera were used to calibrate the flight mill results, permitting absolute estimates of flight performance to be made. Free-flight speeds were approximately three times the speeds recorded by the flight mills. The median corrected distance flown by mated females was >3 km with 20% flying >10 km and 1% flying >20 km. The flight performance of mated females suggests a considerable capacity for range expansion by this invasive species. The implications for quarantine and control are discussed.     

The migration syndrome in the African armyworm moth, Spodoptera exempts: allocation of resources to flight and reproduction

Abstract. Selection for the capacity for prolonged tethered flight in Spodoptera exempta resulted in heavier moths, with significantly larger abdominal glyceride glycerol contents in females of two flight-selected strains (87% and 49% higher than in the comparable non-selected strain) and in males of one of them (80% higher). All flight-selected strains contained individuals of both sexes with very high abdominal glyceride levels. There was no significant relationship between abdominal total glyceride glycerol after flight and flight duration for non-selected moths, but an inverse linear relationship was evident in both sexes from a flight-selected strain. Oxygen consumption during tethered flights by flight-selected moths ranged from 28.2 to 56.6 ml O₂g^-1h^-1.
Using these and previous data, notional energy budgets were calculated to account for flight, reproduction and resting metabolism for non-selected and flight-selected S. exempta flown on the flight balances and provided thereafter with distilled water. The results confirm the trade-off between flight and reproduction reported by Gunn et al. (1989). A similar approach using data for two non-selected strains from the field in Kenya indicated genetic variation in migratory potential, reflected both in pre-reproductive period and resources available for flight. We conclude that elevated glyceride levels are a component of the migratory syndrome in S. exempta and that this is the major factor underlying the curvilinear relationship between flight duration and fecundity obtained by Gunn et al. (1989).     

Relative flight responses of Rhagoletis indifferens as influenced by crowding, sex, and resources

Although flight is believed to be the primary mechanism for dispersal in the Western cherry fruit fly, Rhagoletis indifferens Curran (Diptera: Tephritidae), an orchard pest of both sweet (Prunus avium L.) and sour (Prunus cerasus L.) (Rosaceae) cherry crops, the movement of these flies between host patches is difficult to quantify in the field. A tethered flight mill system was used in the laboratory to examine the flight behaviour of sexually mature flies exposed to different levels of conspecific contact and resource availability. A complete 2 × 2 × 3 factorial design compared the relative influence of the factors ‘context’ (crowded, isolated), ‘sex’ (female, male), and ‘resources’ (low = food only; medium = food + leaf; high = food + leaf + cherries) on flight performance measures including distance flown, net trial time, and stopping patterns. Rather than using a minimum time or distance to determine trial length, flight observations were continued for each fly until a behavioural protocol based on stopping time was met. In this protocol each successful trial was composed of three consecutive flight intervals and included a minimum of three stops lasting a combined total of 5 min. Of the 160 flies tested, 86.9% flew <500 m on the flight mill. Individuals from both sexes were capable of maximum flights in the same order of magnitude, ca. 3 km on the flight mill. Distance flown was significantly influenced by ‘context’ such that crowded individuals flew >1.5-fold farther than isolated individuals. Sex influenced the frequency and duration of stops made, with females stopping more often and longer than males. Although females and males in high resource treatments had the shortest net trial times, the factor ‘resources’ did not produce any highly significant main effects, but did generate significant interaction terms with the factors ‘context’ and ‘sex’, suggesting that past experience with ‘resources’ modifies individual flight behaviour. We have shown for the first time using a tethered flight mill system that R. indifferens flight behaviour is context dependent and sensitive to adult crowding. The implications of this study for improved field experiments on dispersal are discussed.     

Investigations of the oogenesis-flight syndrome inAnthonomus grandis (Coleoptera: Curculionidae) using tethered flight tests

This paper investigates the relationship between oogenesis and flight duration and the use of tethered flight as an indicator of tendency to migrate inAnthonomus grandis grandis Boheman (Coleoptera: Curculionidae), the boll weevil. When boll weevils were flown to exhaustion in tethered flight tests, many flew between 2 and 3 h, with several flying more than 4 h. To test the validity of the tethered flight test as an indicator of tendency to migrate, comparisons of mean flight duration were made between boll weevils trapped in pheromone traps far from any cultivated cotton and those trapped at the edge of heavily infested, flowering cotton fields. There was a significant difference in mean flight time between the two groups, supporting the assumption that long-duration tethered flight in the laboratory reflects the tendency to make long-distance flights in the field. Groups of weevils of different ages were killed after flight testing, and the degree of ovarian development and fat body status were determined and related to duration of tethered flight. Insects with undeveloped or partially developed ovaries were the most likely to make long flights. Weevils with ovaries bearing chorionated eggs made very few long flights. Flight duration was positively correlated with degree of fat body development. In contrast, there were no significant differences in the degree of ovarian development with fat body status. We conclude from these experiments thatA. grandis grandis is capable of long-distance flight, that this species displays some behavioral and physiological characteristics typical of many insect migrants including an oogenesis-flight syndrome, and that a tethered flight test is an appropriate means of measuring migratory tendency in this species.     

甘蓝夜蛾Mamestra brassicae Linnaeus飞行能力研究

【目的】测定日龄、性别、交配等因素对农业迁飞性害虫甘蓝夜蛾Mamestra brassicae Linnaeus飞行能力的影响,为其迁飞行为研究和异地测报提供理论依据。【方法】在气温为(24±1)℃、相对湿度为75%±5%的实验室环境下,利用飞行磨系统对甘蓝夜蛾实验室种群1~5日龄未交配个体、已交配的3日龄个体以及野外迁飞种群分别进行连续24 h吊飞测试,记录其累计飞行时间、累计飞行距离、平均飞行速率等参数。【结果】甘蓝夜蛾实验室种群的飞行能力随日龄增长而变化,初羽化时即具有较强的飞行能力,2~3日龄达到峰值,4~5日龄的飞行能力显著下降;雌性甘蓝夜蛾的飞行能力强于雄性甘蓝夜蛾;交配的甘蓝夜蛾平均飞行速率显著小于处女甘蓝夜蛾,但二者的平均飞行时间和飞行距离均无显著差异。此外,渤海湾迁飞种群中晚季节南迁种群的飞行能力显著强于早季节北迁种群和过渡种群。【结论】日龄和交配状态是影响甘蓝夜蛾飞行能力的关键因素。     

Flight behavior of the bean bug,Riptortus clavatus (Thunberg) (Hemiptera: Alydidae), relative to adult age,sex and season

Studies related to the flight behavior of the bean bug, Riptortus clavatus, an insect pest of soybean, provide information, which can aid the development of management tactics. R. clavatus flight activities were determined using the tethered flight technique in the laboratory and a field study. We compared the flight parameters (flight distance, duration, speed and frequency) of laboratory-reared mated or unmated females and males of different adult age groups, and field-collected females and males from different seasons in a year. Mating effect was only significant for flight frequency, which was higher for unmated adults. Only the flight frequency was different between females and males. Among age groups, 25 d old individuals were shown to have higher flight parameters while the 35 or 45 d old groups showed the lower flight. Among the season, flight activities decreased as the season progressed. The flight distance of R. clavatus in a day was estimated to be 1.6–5.1 km with an average speed of 0.8 m/s from the laboratory data. However, from the field study, it was shown that bean bugs flew a 45–54 m distance with a flight speed of 3.0–3.6 m/s for the first single flight. The second flight was much shorter and slower. We discuss the possible difference of flight parameters between the laboratory and field studies with features of flight mill, variable field conditions and host plant finding behaviors. Further study on flight behavior may provide a better understanding of R. clavatus biology which may guide proper management.     

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.     

Effect of sex,age and morphological traits on tethered flight of Bactrocera dorsalis (Hendel) (Diptera: Tephritidae) at different temperatures

The oriental fruit fly, Bactrocera dorsalis (Hendel) (Diptera: Tephritidae), is a pest of fruit and vegetable production that has become established in 42 countries in Africa after its first detection in 2003 in Kenya. It is likely that this rapid expansion is partly due to the reported strong capacity for flight by the pest. This study investigated the tethered flight performance of B. dorsalis over a range of constant temperatures in relation to sex and age. Tethered flight of unmated B. dorsalis aged 3, 10 and 21 days was recorded for 1 h using a computerized flight mill at temperatures of 12, 16, 20, 24, 28, 32 and 36 °C. Variations in fly morphology were observed as they aged. Body mass and wing loading increased with age, whereas wing length and wing area reduced as flies aged. Females had slightly larger wings than males but were not significantly heavier. The longest total distance flown by B. dorsalis in 1 h was 1559.58 m. Frequent short, fast flights were recorded at 12 and 36 °C, but long-distance flight was optimal between 20 and 24 °C. Young flies tended to have shorter flight bouts than older flies, which was associated with them flying shorter distances. Heavier flies with greater wing loading flew further than lighter flies. Flight distances recorded on flight mills approximated those recorded in the field, and tethered flight patterns suggest a need to factor temperature into the interpretation of trap captures.     

Reproduction-flight relationship in the beet armyworm, Spodoptera exigua (Lepidoptera: Noctuidae)

To evaluate the relationship between reproduction and migratory flight, we examined ovaries, tethered flight, and egg laying in the beet armyworm, Spodoptera exigua (Hübner). Ovarian development in female S. exigua was classified into five stages in relation to the postemergence age of the moths. Oviposition peaked at ovarian stages III and IV. Mating accelerated ovarian development in this species, and thus the mated females developed faster than the virgin moths in ovaries. In virgin adults, flight capacity increased from the first day to the third day after emergence and remained at a high level until the moths were 7 d old. In contrast, the mated moths reached the maximum flight capacity in 2 d after emergence and maintained a high level until the age of 7 d. However, mating status could not influence the flight capacity of the beet armyworm, although there were general differences in flight capacity between virgin and mated adults at the same age. Moreover, significant correlations between the number of eggs laid and the flight capacity were not performed by the mated moths. These results showed that mating status and ovarian development did not seem to influence flight capacity in the beet armyworm, which was not in agreement with the definitions of the oogenesis-flight syndrome observed in many other migratory insects.     

Mating frequency in dispersing potato tuber moth, Phthorimaea operculella, and its influence on the design of refugia to manage resistance in Bt transgenic crops

Mating of potato tuber moth, Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae), was investigated in relation to the dispersal of males in laboratory and field trials. The effect of stimulating the flight of males to light sources in a large cage on their mating ability was estimated for three age groups, and compared with similar estimates for confined moths. Although the mating of males declined with ages of up to 15 days, simulated dispersal had no effect on subsequent mating when the males were paired with virgin females. The dispersal of male moths was also categorised by the initial flight activity of untethered moths to a light source. Scores for poor, moderate, and good flight provided a repeatable measure of initial male flight activity, but the degree of activity was not related to their subsequent mating ability. In the field, virgin female potato tuber moths were tethered at various distances from the edge of isolated potato crops and then dissected to determine their mating status. Female mating frequency averaged 75% at the crop margin, remained above 50% up to 200 m, and then declined to 19% at 360 m from the margin. Derivation of the mating probability for an individual male potato tuber moth confirmed earlier work by other researchers that has indicated a tendency for dispersal prior to mating, and that males retain their ability to mate as they disperse from a crop. The influence of dispersal and mating on gene flow between crops, and its potential effects on refuge size required to minimise the development of resistance to Bt transgenic potato crops was examined.     

Longevity,flight ability and reproductive performance of the diamondback moth,Plutella xylostella (L.) (Lepidoptera: Yponomeutidae), related to adult body size

Plutella xylostella in the temperate zone shows a clear seasonal change in adult body size. In the laboratory, large and small moths were produced during immature stages at 15°C and 25°C, respectively. These moths were then used to evaluate longevity, age-specific flight ability, flight ability of mated and unmated females, and the influence of flight experience on the subsequent reproductive success. The large moths lived longer and displayed a greater flight ability over 3 weeks. Irrespective of body size, unmated females flew for a longer time than mated females, and flight experience affected their subsequent reproductive success. Females of both sizes mated and laid eggs soon after emergence, without any obvious pre-reproductive period. More flight experience did not delay oviposition, but did reduce egg production. It is likely that large moths with a longer adult life span and greater flight ability are better fitted for long-distance flight and more fecund than small ones. These experimental results may explain why long-distance migration ofP. xylostella is mostly seen during cool seasons, when relatively large moths with long forewing appear in the field.