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.     

Environmental conditions for Diaphorina citri Kuwayama (Hemiptera: Liviidae) take‐off

Environmental factors that influence flight activity of Diaphorina citri Kuwayama (Hemiptera: Liviidae) may have implications for Huanglongbing spread and management. In this work, four studies were conducted to evaluate the effect of environmental conditions on D. citri take‐off. In the first, insects were transferred to sweet orange seedlings and confined inside an acrylic cage to verify the take‐off periodicity and the effect of environmental factors on this process. In the second, take‐off temperature threshold was estimated by recording the number of insects that initiated flight from a platform when subjected to gradual temperature increases from 15 to 39°C. In the other studies, we evaluated the effect of different photoperiods and temperature regimes (third study) and of constant temperatures (fourth study) on the propensity for D. citri flight. Insects were confined in clear plastic bottle cages with tubes of 50 ml placed on the cab, to collect emerged adults that initiated flights. Results showed that a small portion of the tested population (maximum 10%) tends to take off from plants and this behaviour is more prevalent in the afternoon (14:00–16:00 h), coinciding with daytimes of lower humidity and higher thermal amplitude. Adults that were submitted to lower temperatures (18°C) and short light periods (10 h) showed less propensity to flight. In contrast, at constant 27°C, the insects were more prone to flight, and this result was confirmed when individuals were submitted to increases in temperature, indicating that 27.14°C is the take‐off temperature threshold of D. citri. Results show that temperature plays an important role in the flight activity of D. citri and suggest that control measures of the insect may be more effective in the morning and in temperatures below 27°C, when the probability to take off from a host is lower.     

Interaction between flight,reproductive development and oviposition in the pine weevil Hylobius abietis

• 1 The development of reproductive and flight capacity of pine weevils Hylobius abietis during the spring and their dispersal to, and subsequent development at, new clearfell oviposition sites comprise key phases in their life cycle in managed forests. At an old clearfell site where autumn‐emerging weevils had overwintered, weevils were trapped as they re‐emerged in the spring and tested for their ability to fly and then dissected to determine the degree of wing muscle and egg development.
• 2 Re‐emerging weevils were most abundant in pine growing at the edge of the clearfell and, over most of the trapping period (April to June), their capacity for flight (proportion flying and wing muscle width) was more advanced than in weevils from the clearfell itself, with a similar trend in the degree of reproductive development (proportion with mature eggs and egg volume).
• 3 In weevils from the clearfell, flight capacity and reproductive development increased concurrently to a peak around mid‐May. In weevils from pine, wing muscles were already well developed at the start of trapping, although few of them flew. Their more advanced development was attributed to the increased opportunities for maturation feeding after emergence in the previous autumn.
• 4 In the spring, weevils reached the canopy of trees for maturation feeding by walking and, to a lesser extent, by flight. Weevils dispersed by flight to oviposition sites in mid‐May when most of them were reproductively mature. After arrival, flight ability and wing muscle size declined rapidly but egg production was maintained until most weevils had stopped flying. When wing muscles reached their minimum size, there was a marked decline in egg size, suggesting that wing muscle breakdown is important in maintaining egg production at oviposition sites. Prospects for further wing muscle and reproductive development are discussed.

     

Spring dispersal ofSitona lineatus: The use of aggregation pheromone traps for monitoring

The spring dispersal ofSitona lineatus L. (Coleoptera; Curculionidae) was investigated on a Danish farm.S. lineatus dispersed by flight in the early spring on sunny, calm days with temperatures above ca. 15°C. Two thirds of the population ofS. lineatus dispersed from perennial leguminous crops (clover and lucerne) in the first period of flight activity. The next dispersal did not occur until one month later despite several intermediate flight activity periods. The first period of dispersal occurred before the germination of the spring sown summer host crop,Vicia faba L. The field bean crop was infested in three later invasions during a period of more than three weeks. The aggregation pheromone, 4-methyl-3,5-heptanedione, had a significant effect on captures of both males and females in cone traps placed on the ground. There was no effect of the pheromone on captures in yellow sticky traps placed 1.5 m above ground. The pheromone effect is discussed in relation to behavioural observations. Both types of traps may be used in a survey system for monitoring spring dispersal ofS. lineatus and optimal timing of insecticide spraying. However, the pheromone cone traps were highly specific whereas all kinds of flying insects were caught in the yellow sticky traps, thus making the latter traps less suitable for monitoring.     

Activity of the forewing stretch receptor in immature and mature adult locusts

Maturation of the flight system of Locusta migratoria occurs during the first two weeks following imaginal ecdysis. One aspect of maturation is an increase in the wingbeat frequency from about 13 Hz to about 23 Hz. We investigated physiological and anatomical mechanisms that may contribute to this process. The difference between the frequencies of the central flight rhythms of immature and mature deafferented preparations was not as great as that between the wingbeat frequencies of immature and mature intact animals. Results from static and dynamic wing elevation showed that the intensity of the forewing stretch receptor response to a given stimulus increased during maturation. The diameter of the main stretch receptor axon was larger and the conduction velocity of signals conveyed along the forewing stretch receptor and the dorsal longitudinal motoneuron was faster in mature than in immature animals. We conclude that during maturation of the flight system the forewing stretch receptor responds to wing elevation with a higher frequency signal that reaches the central circuitry faster. These findings are discussed in the context of a model that describes the influence of stretch receptor input on wingbeat frequency along with other potential mechanisms involved in flight maturation.Abbreviations fDLMn forewing dorsal longitudinal motoneuron - fSR forewing stretch receptor - SR stretch receptor     

Effect of temperature and photoperiod on juvenile hormone biosynthesis and sexual maturation in the cotton bollworm, Helicoverpa armigera: implications for life history traits

The Israeli population of the cotton bollworm, Helicoverpa armigera (Hübner), undergoes a short-day, low-temperature pupal diapause and is also suspected of being a seasonal migrant in the eastern Mediterranean region. H. armigera were reared in the laboratory under several constant temperature and photoperiodic combinations which simulate average conditions encountered in the spring, summer, early-autumn and late-autumn in Israel. Juvenile hormone (JH) biosynthesis, the onset of calling behavior, sex pheromone production and ovarian development were examined in virgin female moths subsequent to eclosion. Allatal maturation, defined as acquisition of competence to synthesize JH, was significantly delayed in moths reared under simulated spring conditions. This was probably the cause for the observed delay in ovarian development and the onset of calling behavior, and to the reduction in sex pheromone biosynthesis. The delay in female sexual maturation, commonly associated with migratory flight, is consistent with presumptive pre-reproductive migration in H. armigera.     

Flight of the bluegreen aphid Acyrthosiphon kondoi Shinji (Homoptera: Aphididae) I. Flight periodicity and phenological relationships

Abstract

Seasonal flight patterns of the bluegreen aphid (BGA), Acyrthosiphon kondoi Shinji (Homoptera: Aphididae) were studied. BGA had flight peaks twice a year in spring and autumn, BGA flight appeared to be diurnal. The main period of activity extended from 5 am to 7.30 pm.

The accumulated spring and autumn flight peaks lasted 422 Degree Days (DD) and 351 DD (2.9 and 2.4 BGA generations) respectively. Accumulated DD between the median of spring and autumn flights, and the autumn and following spring flight, was 1667 and 1676 DD; this corresponds to 11.5 and 11.6 BGA generations respectively. The use of DD for predicting BGA flight is discussed.     

Flight speeds of swifts (Apus apus): seasonal differences smaller than expected

We have studied the nocturnal flight behaviour of the common swift (Apus apus L.), by the use of a tracking radar. Birds were tracked from Lund University in southern Sweden during spring migration, summer roosting flights and autumn migration. Flight speeds were compared with predictions from flight mechanical and optimal migration theories. During spring, flight speeds were predicted to be higher than during both summer and autumn due to time restriction. In such cases, birds fly at a flight speed that maximizes the overall speed of migration. For summer roosting flights, speeds were predicted to be lower than during both spring and autumn since the predicted flight speed is the minimum power speed that involves the lowest energy consumption per unit time. During autumn, we expected flight speeds to be higher than during summer but lower than during spring since the expected flight speed is the maximum range speed, which involves the lowest energy consumption per unit distance. Flight speeds during spring were indeed higher than during both summer and autumn, which indicates time-selected spring migration. Speeds during autumn migration were very similar to those recorded during summer roosting flights. The general result shows that swifts change their flight speed between different flight behaviours to a smaller extent than expected. Furthermore, the difference between flight speeds during migration and roosting among swifts was found to be less pronounced than previously recorded.     

A new experimental approach to investigate migration in Pieris brassicae L.

Abstract.

• 1 It is possible to correlate distinct sequences of flight behaviour in caged females of the migrant butterfly Pieris brassicae with the main flight direction of released individuals of the same stock. The attempts to escape out of the cage point to the same mean direction as the flights of the released butterflies. This provides an experimental tool to investigate some unsolved questions concerning migration behaviour.
• 2 This study examines the constancy of the flight direction of a group of butterflies, the specificity of the direction in two populations of different geographical origin, and the frequently reported change in flight direction of subsequent generations.
• 3 The results indicate that the main flight direction of a tested group is very constant during 2 days of observation. Different directions have been detected in populations originating from Northern Germany and Southern France, respectively. The preferred directions are discussed as an adaptation to the geographical circumstances. Subsequent generations of the population from Northern Germany also show different directions. The spring generation flies to the north, the autumn generation in the opposite direction. This result corresponds to field observations that in P.brassicae a return flight occurs in subsequent generations of a year.

     

Flight thresholds and seasonal variations in flight activity of the light-brown apple moth,Epiphyas postvittana (Walk.) (Tortricidae), in Victoria,Australia

Summary The flight activity of Epiphyas postvittana was studied at two sites near Melbourne with the aid of suction traps, over a period of 4 years. Maximum numbers were found to fly during the period September to March with peak activity coinciding with the emergence of winter, spring and summer generation moths. E. postivittana is predominantly a nocturnal flier with maximum activity around 20.00–24.00 h. The lower temperature threshold of flight was 8–11°C. The upper temperature threshold varied from 20–21°C, 24–25°C and 27–28°C for the winter, spring and summer generation moths respectively. Flight was highly influenced by the prevailing wind. The lower wind speed threshold was 0.5–0.8 m^-s and the upper wind speed threshold was 2.6–2.7 m^-s. The relationship between wind speed and the amount of flight was non-linear, with the frequency of flights decreasing sharply with increasing wind speed. No flights occurred at wind speeds greater than 2.8 m^-s. Variation in relative humidity had no influence on flight, but lack of rain favoured flight. The amount of flight activity and the amount of rainfall were negatively correlated; flights did not occur when the daily precipitation exceeded 32.5 mm, and with a precipitation exceeding 39 mm no flights could be expected. The value of these findings to pest control programmes is discussed.     

Exposure to heat shock affects thermosensitivity of the locust flight system

The natural habitat of the migratory locust, Locusta migratoria, is likely to result in locusts being heat stressed during their normal adult life. It is known that locusts exhibit a heat-shock response: exposure to 45°C for 3 h induces thermotolerance and the expression of heat-shock proteins. We investigated the effects of exposure to heat-shock conditions on the thermosensitivity of flight rhythm generation in tethered, intact animals and in deafferented preparations. Heat shock had no effect on wingbeat frequency measured at the start of flight sequences, nor did it affect the postimaginal maturation of this parameter. During sustained flight, heat shock slowed the characteristic asymptotic reduction of wingbeat frequency. Wingbeat frequency of heat-shocked animals was less sensitive to temperature in the range 24° to 47°C than that of control animals, and the upper temperature limit, above which flight rhythms could not be produced, was 6° to 7°C higher in heat-shocked animals. These results were mirrored in the response of deafferented preparations, indicating that modifications in the properties of the flight neuromuscular system were involved in mediating the response of the intact animal. We propose that exposure to heat shock had the adaptive consequences of reducing thermosensitivity of the neural circuits in the flight system and allowing them to operate at higher temperatures. © 1996 John Wiley & Sons, Inc.     

Interactions between photoperiodism and temperature with respect to the control of dormancy in the adult stage of Pterostichus oblongopunctatus F. (Col., Carabidae)

Summary Pterostichus oblongopunctatus belongs to the type of spring breeding carabids with adult hibernation. Ovarian maturation is under the control of photoperiod: the females need a change of day length to mature. The first step of egg maturation (previtellogenesis) is bound to short day, the second step (vitellogenesis) to long day.The short day effect is achieved almost only at temperatures of 10 to 15°C; it is completely suppressed by 20°.The males mature under very different photoperiods and temperatures, but long day diminishes the percentage of mature males slightly, as well as does the higher temperature of 20°C in short day as compared with 15°C.Simulation of climatic conditions as experienced by the beetles from autumn throughout winter until spring in the laboratory showed that a short span of favourable temperatures at short day as experienced in nature in autumn allows the females to accomplish the short day development. But this maturation effect is still more diminished the longer temperatures below 5°C act on the beetles. Following that, a second period of temperatures between 10–15°C must restore the short day maturation effect if full maturity shall be gained in the following long day.The beetles are able to store reserve materials under long day at 20°C. After such a period they are able to have a normal previtellogenesis in spite of a severe shortage of food during short day. These experiments indicate that the failure in short day development outside the temperature range of 10–15°C is not a matter of an especially well balanced over-all metabolism in this favourable temperature range. It seems rather that hormonal systems involved in the short day phase of oogenesis act only within this span of temperature.Supportet by the Deutsche Forschungsgemeinschaft.     

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.     

Experimental effects of immersion time and water temperature on body condition,burying depth and timing of spawning of the tellinid bivalve <Emphasis Type="Italic">Macoma balthica</Emphasis>

The burying depth of many bivalve molluscs on intertidal mudflats varies throughout the year and differs between places. Many factors are known to influence burying depth on a seasonal or spatial scale, with temperature and tidal regime probably being very important. Burying depth, body condition and gonadal development of Macoma balthica were followed throughout winter and spring in an experiment in which water temperature and immersion time were manipulated. Unexpectedly, relative water temperature, in contrast to the prediction, did not generally affect body condition or burying depth. This was probably a consequence of the exceptionally overall low water temperatures during the experimental winter. Differences in temperature did, however, result in different timing of spawning: M. balthica spawned earlier at higher spring temperatures. Longer immersion times led to higher body condition only late in spring, but led to deeper burying throughout almost the whole period. There was no effect of immersion time on the timing of spawning. We conclude that a longer immersion time leads to deeper burying, independent of body condition. We also conclude that burying behaviour of M. balthica is not determined by the moment of spawning. Electronic Publication     

Successful despite poor flight performance: range expansion is associated with enhanced exploratory behaviour and fast development

Anthropogenic interference forces species to respond to changing environmental conditions. One possible response is dispersal and concomitant range shifts, allowing individuals to escape unfavourable conditions or to track the shifting climate niche. Range expansions depend on both dispersal capacity and the ability to establish populations beyond the former range. We here compare well‐established core populations with recently established edge populations in the currently northward expanding butterfly Lycaena tityrus. Edge populations were characterized by shorter development times and smaller size, a higher sensitivity to high temperature and an enhanced exploratory behaviour. The differences between core and edge populations found suggest adaptation to local climates and an enhanced dispersal ability in edge populations. In particular, enhanced exploratory behaviour may be advantageous in all steps of the dispersal process and may have facilitated the current range expansion. This study describes differences associated with a current range expansion, knowledge which might be useful for a better understanding of species responses to environmental change. We further report on variation between males and females in morphology and flight behaviour, with males showing a longer flight endurance and more pronounced exploratory behaviour than females.     

Spawning behaviour of barred spiny eel,Macrognathus pancalus,in the River Gomti,India

The present study investigated the spawning behaviour of the spiny eel, Macrognathus pancalus, and its relationship with the environment (photoperiod and temperature). The results show that M. pancalus breeds during February to September with peak spawning during February/March and July/August, indicating two main breeding periods. Initiation of breeding occurs during increasing day lengths in early spring (February/March), and regression during decreasing day lengths in autumn (September/October). The breeding status confirmed by gonadosomatic index (GSI) and ovarian histology showed six stages of ovarian maturation (viz. stage I: chromatin nucleaolar; II: perinucleolar; III: cortical alveolar; IV: vitellogenesis; V: ripe; and VI: partially spent), with predominance of one or more stages at a particular time of the year. Males also followed a similar seasonal pattern in gonadal maturation as reflected by their GSI. Results show that M. pancalus is a seasonal breeder with bimodal spawning activity. The breeding pattern suggests an influence of prevailing day length conditions at this latitude.     

Monitoring the dynamics of orchard colonisation by Anthonomus pomorum in spring

Overwintered adult apple blossom weevils, Anthonomus pomorum (L.) (Coleoptera: Curculionidae), colonise apple trees, Malus domestica Borkh. (Rosaceae), in early spring. Information gained from a suitable monitoring technique could serve as a guide to determining the accurate timing for control measures. To assess tree colonisation by flight and crawling, Plexiglas flight traps and commercial screen traps developed for different curculionid species were used. Refuges were mimicked by a newly designed shelter trap based on transparent bubble wrap. The trap catches were contrasted to the limb jarring technique. Microclimate and weather conditions were determined over a trial period of one and a half months. The shelter traps caught a high number of weevils irrespective of weather conditions, and thus performed better than both flight and screen traps. The temperature level within the shelter traps was slightly, but significantly, higher than on the trunk, indicating that weevils exploit microclimatic differences and occupy thermally favourable refuges. The seasonal culmination of colonisation as determined by shelter traps coincided with that as determined by the two methods used to monitor weevil movement towards host trees (flight traps and screen traps). The data indicated that not only flight, but also crawling as quantified by the screen traps contributes substantially to spring colonisation. Captures by all three types of traps over the spring season were significantly positively correlated with each other, but in most respects not with the results of the limb jarring. We conclude that the newly designed shelter trap will be useful for accurately monitoring the seasonal course of colonisation and holds promise for determining the timing of interventions.     

Swarming and mating behaviour ofChironomus flaviplumus (Diptera: Chironomidae), compared with a sympatric congeneric species,C. yoshimatsui

The swarming and mating behaviour ofChironomus flaviplumus was observed and compared with a sympatric congeneric species,C. yoshimatsui. C. flaviplumus males swarmed around sunset near foliage or angles of buildings near the emergence site and copulated with females entering the swarm. Swarming and mating occurred under conditions of higher light intensity in cooler seasons than in warmer ones. Results suggested that temperature had an effect on the timing of flight to the swarming site in both sexes. TheC. flaviplumus swarm marker and swarming behaviour seemed very similar to that toC. yoshimatsui, and their respective daily swarming time zone greatly overlapped. No mixed swarm, however, was observed in the study area. This is probably due to the distance between the species' larval habitats. Possible premating isolation mechanisms between these 2 species are discussed.     

Environmental and morphological factors influencing predatory behaviour by invasive non-indigenous gammaridean species

Predatory behaviour seems to be more frequent in invasive gammaridean species than in native ones. This results in the exclusion of other, mostly native gammaridean species and a change in benthic communities. The present study analysed the influence of environmental factors (water temperature) and morphological factors (sex, body parts involved in catching and holding prey) on the predatory behaviour of Dikerogammarus villosus. A diet study of invasive relatives of D. villosus showed that predation intensity is especially high in spring and summer, that is, at increasing and high temperatures. Experiments with D. villosus in climate rooms at various temperatures, using the native Gammarus fossarum as prey, showed that the average predation rate by both sexes gradually increased over the temperature range from 5 to 30°C. Natural mortality during the experiments was negligible compared to losses due to predation. At each temperature, the predation rate by females was lower than that by males. Males showed a steep allometric growth of body parts involved in the process of catching and holding prey, compared to females at increasing body size in a number of measurements. This may explain the difference in predatory behaviour between males and females, which plays a role in intraguild predation a supposed mechanism for species displacement.     

Seasonal flight activity by the Asian citrus psyllid in east central Florida

The Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Psyllidae), is an important invasive citrus pest in the USA because it vectors a bacterium responsible for huanglongbing, a devastating disease of citrus. Information was lacking on seasonal aspects of flight activity by D. citri, which could have ramifications on psyllid management as well as our understanding of epidemiology of the disease. Of interest from a pest management standpoint would be whether D. citri regularly disperses to or away from citrus on a predictable schedule. In research presented here, seasonal flight activity by D. citri was investigated using yellow sticky traps deployed in citrus trees and in fallow areas adjacent to citrus. Results indicated that flight activity by both male and female D. citri away from citrus can occur at any time of the year with consistent dispersal activity during the spring. The research further indicated citrus is continually subject to infestation by immigrating adults and that there is no time during the year that a citrus grower could be assured immigration would not occur. Growers should be aware that adult dispersal occurs regularly during spring and they should time management tactics accordingly. Adult flight activity 2 m from a citrus tree was more pronounced at 1 m above ground than at 2 or 3 m high. At distances of 8–60 m from trees, numbers of adults on traps were similar among the three heights. Males and females were similar with respect to seasonal flight activity. Numbers of adults captured on traps distant from citrus were not correlated with wind speed, sunlight, or air temperature, but there was some evidence that relative humidity influenced flight activity. Although the D. citri life cycle is dependent on flush, data from these studies did not confirm that psyllid dispersal from citrus consistently increases as citrus flush abundance decreases.