Post-juvenile dispersal of Hazel Grouse Bonasa bonasia in an expanding population of the southeastern French Alps

We studied the post-juvenile dispersal of 18 radiotagged juvenile Hazel Grouse Bonasa bonasia (14 males, four females) in an expanding population in the southeastern French Alps between 1998 and 2001. The mean dispersal distances between the capture sites of juveniles in September–October and the centre of the home range in the following spring was 4 km for males (range 0.1–24.9 km) and 2 km for females (range 0.2–5.6 km). The distances recorded for two long-dispersing males (15 and 24.9 km) are greater than those reported to date for Hazel Grouse. Using our radiotracking data, we interpret the pattern of range expansion that has been occurring since the 1950s around our study area. Barriers to dispersal included rocky ground and other alpine habitats above 2000 m and over 1 km wide, but Hazel Grouse did cross open agricultural land at lower elevation. Two patterns of dispersal movements were recognized in juveniles: erratic movements that led to settlement on or near the natal site, and direct movements to a new range relatively far from the natal area. We discuss the adaptive consequences of these different behaviour patterns.     

Post-fledging behaviour in Golden Eagles Aquila chrysaetos: onset of juvenile dispersal and progressive distancing from the nest

Thirteen juvenile Golden Eagles Aquila chrysaetos were tracked during their first year of life using satellite telemetry. Distances to the nest attained during that period and the age at the onset of juvenile dispersal were explored. The performance of nine different criteria to determine that age was analysed. In general, after a brief period of restricted movements around the nest, the average distance to the nest increased with time. Maximum distances to the nest ranged between 57.7 and 184.3 km, and were considerably greater in females (mean ± sd, 138.5 ± 44.5 km) than in males (70.5 ± 14.0 km). No sex difference was observed in the age at which that distance was attained (males: 329 ± 32 days, females: 312 ± 20 days). The onset of juvenile dispersal took place around the fifth month of life (September in Spain). Eight of the nine criteria provided similar results, suggesting that in Spain dispersal starts when birds are between 140 and 180 days old, and that the post-nestling period lasts between 60 and 120 days. For future studies, to determine the age at which the onset of juvenile dispersal occurs, we recommend the use of either the first day on which individuals were located beyond the mean distance between nests of different pairs (10 km in our study area), or the date of the record midway between the first and the last location recorded during the month in which the maximum variability in the distance to the nest was observed.     

Female-biased natal and breeding dispersal in an alpine lizard, Niveoscincus microlepidotus

We measured two aspects of dispersal in the alpine Australian scincid lizard, Niveoscincus micolepidotus : (1) natal dispersal, i.e. shift in home range over the lizard's first year of life, and (2) breeding dispersal, i.e. shifts of home ranges between breeding attempts as adults. On average, displacements were surprisingly small. Female neonates dispersed about twice as far as did males in the same cohort (means of 12 m vs. 6 m). A female's natal dispersal distance was not correlated with her body size or our estimate of physiological performance (sprint speed). However, larger, faster-running male neonates dispersed further than did smaller, slower males. As was the case for neonates, adult females moved significantly further between breeding seasons than did adult males (14.2 m vs. 9.6 m). Because of a female's long gestation period (more than 1 year), two groups of females occur simultaneously in the population, non-ovulated (i.e. with yolking folicles) and pregnant females (i.e. approaching parturition). Females that were not yet ovulated showed a markedly stronger dispersal in response to high reproductive effort (i.e. clutch size in relation to body condition) than did pregnant females. In adult males, body size was negatively correlated with dispersal distance, suggesting that although males have overlapping territories, they exhibit an increasing level of site tenacity with age and/or size. Thus, selection for the relatively more pronounced site tenacity in adult males may have resulted in the more marked philopatric behaviour compared to females also as neonates.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 277–283.     

Dark-bellied Brent Geese Branta bernicla bernicla, as recorded by satellite telemetry, do not minimize flight distance during spring migration

Nine Dark-bellied Brent Geese Branta bernicla bernicla were equipped with satellite transmitters during spring staging in the Dutch Wadden Sea in 1998 and 1999. The transmitters (in all cases less than 3% of body mass) were attached to the back by a flexible elastic harness. One juvenile female was tracked to the Yamal peninsula in 1998. Eight adult males were selected from a single catch of 75 to span the range of body mass observed on the date of capture (11 May 1999) and all but the lightest individual completed the first lap of the migratory flight to the White Sea, Russia, according to the time schedule normal for the species. Six birds were successfully tracked to Taymyr for a total distance averaging 5004 km (range 45775164) but judging from later movements none bred (although 1999 was a breeding year). Although the routes chosen during spring migration were closely similar, none of the tagged birds migrated together. On average the geese used 16 flights to reach their summer destinations on Taymyr. The longest uninterrupted flights during the first half of the journey (Wadden Sea to Kanin) covered 1056 km (mean of seven adult males, range 7681331), while the corresponding value for the second half of the migration (KaninTaymyr) was only 555 km (mean of six adult males). Only 7% of total time during spring migration was spent in active flight, as contrasted to c.  80% at long-term stopovers. Overall average travelling speed was 118 km/day (range 97148). Including fattening prior to departure the rate of travel falls to 62 km/day (range 4970), in keeping with theoretical predictions. Routes followed deviated from the great circle route, adding at least 700 km (16%) to the journey from Wadden Sea to Taymyr, and we conclude that the coastal route is chosen to facilitate feeding, drinking and resting en route instead of minimizing total flight distance.     

Natal dispersal and its consequences in Black Grouse Tetrao tetrix

Data on dispersal distances of juveniles are important for understanding the genetic structure of populations, population regulation processes and the effects of landscape ecology on metapopulation dynamics. We studied dispersal in juvenile Black Grouse Tetrao tetrix in the French Alps between 1990 and 1998, by radiotracking 39 young grouse captured in August or autumn in a study area of 836 ha. Natal dispersal occurred in two discrete phases, autumn (October) and spring (mid-April to early May), with periods of reduced mobility between. The mean distance travelled by females exceeded that of males in autumn but not in spring. The natal dispersal distance of females was greater than that of males, and resulted in 81% of females leaving the study area to nest 5–29 km from their site of capture. This emigration must have been compensated for by immigrants because the number of hens in the study area increased during the study. Males were more philopatric. By the summer after capture, only 9% had emigrated from the study area. Despite the longer dispersal distances of females, there was no difference in the survival functions of the sexes between the ages of 6 weeks and 13 months.     

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.     

Sex-biased and body condition dependent dispersal capacity in the endangered saproxylic beetle <Emphasis Type="Italic">Osmoderma eremita</Emphasis> (Coleoptera: Cetoniidae)

The dispersal capacity of rare and endangered insect species has rarely been estimated even though it is essential for their management. For these species, laboratory based experiments are considered more appropriate for determining dispersal capacity as well as the factors influencing it. We aimed to characterize dispersal capacity of the endangered saproxylic beetle Osmoderma eremita (Scopoli, 1763) (Coleoptera: Cetoniidae). We studied the influence of sex and body condition on several parameters of dispersal (seven parameters of flight capacity measured in laboratory and pre-flight behaviour observed in the wild). Tethered flight experiments, conducted on 30 individuals collected in several regions of France, revealed: (1) maximal single flight distance of 1,454 m and maximal total flight distance of 2,361 m; (2) higher flight capacity in females than males; (3) flight speed and take-off completion decreasing with increasing body condition only for females. Additionally, 32 individuals displaying pre-flight behaviour in the wild showed similar interacting influences of sex and body condition: females initiating pre-flight behaviour had lower body condition than males. Thus, males and females have different dispersal strategies. We propose that body condition influences on dispersal capacity should be considered for species conservation by, for instance, managing adult food resources at the landscape scale and need to be taken into account in introduction programmes.     

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.     

Development of chicks and predispersal behaviour of young in the Eagle Owl Bubo bubo

Little quantitative information on the development and behaviour of chicks and young is available for many species, despite the crucial importance of such data and the sensitivity of this stage in a bird's life. For Eagle Owls Bubo bubo , despite the large amount of scientific literature on this species, much basic information is lacking. This study provides a photographic and morphometric guide for age estimation of nestlings and fledglings, as well as data on the call behaviour of young, and patterns of movements during the post-fledging dependence period. The most remarkable event in chick development is the rapid increase in mass, and size gain, during the first 30 and 40–45 days, respectively. Because after this time morphometric differences become less evident, young-feather development is more useful for ageing. Patterns of chick call behaviour showed that the time spent calling increased with age and, from 110 days of age, chick vocalizations were usually uniformly distributed through the whole night and most synchronized at sunset and sunrise (the maximum recorded number of vocalizations per chick and per night was 1106 calls). During the post-fledging dependence period, radiotagged Owls moved widely, up to 1500 m from the nest after the age of 80–90 days. During such movements, the mean distance among siblings increased with age, from 168 m on average for juveniles less than 100 days old, to 489 m for those older than 100 days. Definitive dispersal started when young were about 150–160 days old. Information on chick call behaviour and movements is crucial for unbiased census and nest checking, as well as for the definition of young post-fledging areas. Knowledge of the latter is very important in terms of conservation and management (especially for those species that move largely around their nest before dispersal) owing to the high mortality that can occur during this period.     

Significance of spring migration and flexibility in flight-muscle histolysis in waterstriders (Heteroptera, Gerridae)

Abstract. 1. Individuals of long-winged waterstrider (Gerridae) species were found in spring far from their breeding habitats, which indicates that they fly before reproduction.
2. Field samples and laboratory studies show that once they return to their breeding sites, many individuals of three waterstrider species ( Gerris odontogaster (Zett.), Gerris lacustris (L.) and Limnoporus rufoscutellatus (Lat.)) histolyse wing muscles and lose flight ability during their reproductive period.
3. The extent of flight-muscle histolysis varies with environmental factors. Food scarcity affects flight-muscle histolysis in G.odontogaster females. In G.Lacustris , flight-muscle histolysis was more common in the laboratory than in the field samples. Proportionately more females than males lost their flight ability by the end of the reproductive period.
4. Flight ability had direct costs in reproductive potential with (non-flyer) females, which histolysed their flight muscles, laying more eggs than (flyer) females, which maintained flight ability. This was also the case during food scarcity. Non-flyer males of G.odontogaster survived longer than flyer males.
5. Spring migration was distinguished from dispersal during the reproductive period, because these flights serve different functions. Flight-muscle histolysis of females during reproduction is a qualitative reproductive option, with a trade-off between dispersal ability and reproductive potential. Ability to change reproductive behaviour depending on environmental conditions increases an individual's ability to cope with a large variety of habitats.     

Effects of gender and mating status on self-directed dispersal by the whitefly parasitoid Eretmocerus eremicus

1. The impacts of gender and mating on short‐range (< 10 km) dispersal by the whitefly parasitoid Eretmocerus eremicus Rose and Zolnerowich (Hymenoptera: Aphelinidae) were examined. A fluorescent dust‐marking technique was also tested. 2. In a vertical flight chamber, female flight duration was significantly longer than that of males, and unmated parasitoids flew for longer than mated parasitoids. The mean flight durations were: unmated females 34 min, mated females 10 min, unmated males 7 min, mated males < 1 min. 3. The dispersal behaviour of E. eremicus was investigated in the field using fluorescent dust. Before doing so, it was determined in the laboratory that this dust did not affect flight behaviour, was retained over the length of the experiments, and allowed large samples to be processed quickly and inexpensively. 4. In the field, traps were placed along annuli at 3, 5, 7, and 10 m from release points. Eighty‐seven per cent of the 4153 parasitoids captured were males. Sex ratios were near parity on release. 5. The difference in dispersal characteristics between males and females may be resource based, suggesting that certain requirements were met within the field plots for males that were not met for females. 6. Locally, males dispersed in a manner consistent with a simple diffusion model while females engaged in wind‐directed flight soon after leaving release sites. The fact that the genders exhibited dissimilar dispersal characteristics, supports the claim that insect flight, even by small species, can be self‐directed.     

Mutual benefits of associations between breeding and non-breeding White-fronted Geese Anser albifrons

The relationships between yearlings and adult pairs of White-fronted Geese Anser albifrons were studied during pre-nesting, laying and early incubation in the central Canadian Arctic. Prior to nesting, females of lone pairs spent 75–81% of their time feeding, while males spent only 42–47% of time feeding and 46–50% alert. In pairs with one or more associated yearlings, both females and males fed significantly more and spent less time vigilant. Yearlings spent significantly less time (59%) feeding when alone compared with 71–76% when with pairs. Associations between yearlings and paired adults were most frequent before adult females began prospecting for nest sites. No prospecting pairs were associated with yearlings. After the egg-laying period, groups of geese, predominantly yearlings, made distraction flights over humans and terrestrial predators approaching nests, in contrast to the more cryptic behaviour of nesting pairs. The presence of groups of geese associated with some nest sites suggests that continuing parent-offspring relationships may involve assistance with nest defence.     

Breeding dispersal of Northern Flickers Colaptes auratus in relation to natural nest predation and experimentally increased perception of predation risk

After nest predation, breeding dispersal can be an effective strategy to avoid local nest predators. Furthermore, encounters with predators at a nest during the pre-laying stage may be used by parents to judge future risk, such that they may abandon a nest when a nest predator has been encountered. We studied whether the between- and within-year breeding dispersal of Northern Flickers Colaptes auratus was dependent upon the outcome of the previous nesting attempt. We also tested whether pairs presented with a model predator prior to egg-laying were more likely to abandon their nests than were pairs presented with a control model. Between years, males moved significantly further after having their nest depredated than did successful males, and females showed the same trend. However, these movements did not result in greater reproductive success. More pairs switched sites within years after having their nest depredated, but those that remained and those that moved had equal subsequent nest success. Stressful encounters with predators involving nest defence may trigger dispersal both between and within years, although reproductive benefits are unclear. The proportion of pairs abandoning nests did not differ between parents presented with control or predator models, suggesting that a single encounter with a predator is not a sufficient deterrent against continued use of a particular nest.     

Movements of Macquaria ambigua, in the Murray River, south‐eastern Australia

A 2 year (2000–2001) radio‐tagging study was undertaken to investigate the movements of 51 golden perch Macquaria ambigua in the Murray River at Nyah in north‐western Victoria, Australia. During the winter of both years, golden perch did not undertake movements >5 km and displayed strong home range fidelity. In the first year of the study there was an increase in the distance of golden perch movement during late spring which coincided with increasing water temperature and river discharge. Nineteen golden perch were tracked during this period, of which 10 travelled downstream between 11 and 290 km. Seven of these fish moved to an area below the junction of the Murray and Wakool Rivers. Five golden perch travelled upstream between 13 and 35 km, four of which travelled to an area around the junction of the Murray River and Speewa Creek. The remaining four golden perch undertook localized movements of <5 km. Many of the long distance movements undertaken in spring 2000 were rapid and 53% of these golden perch returned to within 3 km of their release locations, indicating homing behaviour. Given that the rapid movements of golden perch in spring coincided with the known spawning season of this species, these long distance movements may be associated with reproductive strategy.     

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.     

Flight metabolic rate has contrasting effects on dispersal in the two sexes of the Glanville fritillary butterfly

Evolution of dispersal is affected by context-specific costs and benefits. One example is sex-biased dispersal in mammals and birds. While many such patterns have been described, the underlying mechanisms are poorly understood. Here, we study genetic and phenotypic traits that affect butterfly flight capacity and examine how these traits are related to dispersal in male and female Glanville fritillary butterflies (Melitaea cinxia). We performed two mark–recapture experiments to examine the associations of individuals’ peak flight metabolic rate (MR_peak) and Pgi genotype with their dispersal in the field. In a third experiment, we studied tethered flight in the laboratory. MR_peak was negatively correlated with dispersal distance in males but the trend was positive in females, and the interaction between MR_peak and sex was significant for long-distance dispersal. A similar but nonsignificant trend was found in relation to molecular variation at Pgi, which encodes a glycolytic enzyme: the genotype associated with high MR_peak tended to be less dispersive in males but more dispersive in females. The same pattern was repeated in the tethered flight experiment: the relationship between MR_peak and flight duration was positive in females but negative in males. These results suggest that females with high flight capacity are superior in among-population dispersal, which facilitates the spatial spreading of their reproductive effort. In contrast, males with high flight capacity may express territorial behaviour, and thereby increase the number of matings, whereas inferior males may be forced to disperse. Thus, flight capacity has opposite associations with dispersal rate in the two sexes.     

On the ecology of the poplar admiral (<Emphasis Type="Italic">Limenitis populi</Emphasis>, Lepidoptera,Nymphalidae) in Eastern Fennoscandia

The ecology of the poplar admiral was studied in the periphery of its range for over 19 years (1990–2008), including seasonal phenomena, spatial distribution, and dynamics of abundance and dispersal of the species. The spatial organization of population of the species was studied in one of the local areas. The flying period of butterflies in Eastern Fennoscandia is divided into three phases: the observed flight of males, latent activity, and observed flight of females. The males emerge more than 10 days earlier than the females. The difference is likely to be one of the reasons for strong reduction in the butterfly abundance. The asynchrony of dynamics in different areas was explained by the influence of local factors. In the years of high abundance, the proportion of occupied areas increased. Analysis of the dispersal and abundance curves showed a trend for colonization of vacant areas. Within the occupied area, the butterflies are irregularly distributed. The presence of areas with constantly moist soils and aspen trees in the forest stands are the main factors responsible for butterfly aggregations. Observations of the marked individuals showed that the maximum distance covered by a butterfly was 4.8 km. The butterfly moved freely across all the study area, and no single accumulation of butterflies isolated from the others was recorded. All the movements are shown to be local. The distance of these movements, according to the exponential model, reached 6–7 km; the grouping itself is classified as a subpopulation, i.e., a structural unit of a large spatially differentiated population. The high mobility of the poplar admiral ensures migration of individuals between the populations and colonization of vacant habitats.     

Alongshore dispersal and site fidelity of juvenile plaice from tagging and transplants

Juvenile plaice Pleuronectes platessa ( n  = 1281) were tagged and released at two locations 300 m apart on a 1 km long sandy beach. Most (>90%) of the fish were recaptured within 100 m of the release site (shown by the colour of the tag), with very few caught >200 m distance after 6 weeks. The changing spatial distribution of marked fish was adequately reproduced by a simple dispersal model with a single parameter: a 78% probability of remaining in a 100 m wide zone from one day to the next, with a 22% chance that fish move into an adjacent zone. In a subsequent similar study at the same beach, fish were either released at the point of capture ( n  = 881) or transplanted to the alternate site ( n  = 910) 100 m distant. After 6 weeks, transplanted fish moved along the shore towards their sites of original capture. Fish replaced at the point of capture showed no such movement along the shore. Further modification of the dispersal model to allow for a distinction between dispersal from home sites and from sites away from the original point of collection, was sufficient to reproduce the behaviour of the populations of both transplanted and control treatment groups. The likelihood of dispersal from home sites was much less than that seen at sites away from home. Juvenile plaice thus have a degree of long‐shore site fidelity not expected of a fish with strong depth‐related migration behaviour in a relatively homogenous habitat.     

Interrelation of mating,flight, and fecundity in navel orangeworm females

The navel orangeworm, Amyelois transitella (Walker) (Lepidoptera: Pyralidae, Phycitini), is an economically important pest of nut crops in California, USA. Improved management will require better understanding of insect dispersal, particularly relative to when mating occurs. A previous study demonstrated a more robust laboratory flight capacity compared to other orchard moth pests, but it was unclear how mating affects dispersal, and how dispersal affects fecundity. In this study, 1‐ and 2‐day‐old females were allowed to fly overnight on a flight mill either before or after mating, respectively, and were then allowed to oviposit. Data on fecundity were compared between treatments to minimally handled or tethered‐only control females. Females that mated before flight flew longer and covered a greater distance than those flying prior to mating. However, timing of flight relative to mating did not affect fecundity, nor did any measure of flight performance. There was no effect on fecundity when females were forced to fly for designated durations from 3 min to 2 h. Together, our data revealed no obvious trade‐off between flight activity and reproductive output. Distances measured on the flight mills (mean ca. 15 km for mated females) may overestimate net displacement in the field where flight tracks are often meandering. The results suggest that most females mate and oviposit in or near their natal habitat, but that some may disperse potentially long distances to oviposit elsewhere.     

OVIPOSITION BEHAVIOUR AND DISPERSAL OF PERTHID A GLYPHOPA COMMON (LEPIDOPTERA: INCURVARIIDAE)

Abstract
Oviposition behaviour of the jarrah leafminer moth Perthida glyphopa was observed during the seasonal flight period in April-May. After alighting on trees, the gregarious males restricted their movements to the branches, whereas the solitary females searched for oviposition sites on the leaves. The females laid most eggs at 14–22 °C in diminishing light or overcast conditions, preferring young leaves in the lower portion of the crown. They dispersed more readily than males.