The Effect of Abiotic Factors on the Male Mate Searching Behavior and the Mating Success of Aphidius ervi (Hymenoptera: Aphidiidae)

The effect of wind speed and distance from the source on the male response of the aphid parasitoid, Aphidius ervi (Hymenoptera: Aphidiidae), to a pheromone source was studied in a wind tunnel. The number of males taking flight, entering the plume and successfully reaching the source, decreased at wind speeds >50 cm/s. Furthermore, the proportion of those attempting upwind flight that fell to the ground increased with increasing wind speed. In contrast, distance from the source had no significant effect on any of the parameters examined. While male flight behavior was significantly reduced at 70 cm/s, some males walked to the source when there was a bridge connecting the pheromone source and the release platform. This suggests that ambulatory behavior could be a significant component of male mate searching in A. ervi when wind conditions are too strong for upwind flight. The possible effects of variation in atmospheric pressure on male flight behavior to the long distance pheromone, as well as to the short distance one, were also investigated. No significant effects of atmospheric pressure were observed. These findings differ significantly from those previously reported for another aphid parasitoid, A. nigripes, and the reasons for such differences are discussed.     

Emergence, Mating, and Postmating Behaviors of the Oriental Beetle (Coleoptera: Scarabaeidae)

In a previous field-trapping study of the oriental beetle, Exomala orientalis (Waterhouse), by using synthetic sex pheromone on golf course fairways, numerous males were observed and trapped during the hours of peak mating activity. However, very few beetles were observed in the same areas when synthetic pheromone was absent. To investigate the hypothesis that mating in nature occurs cryptically within vegetation at the soil surface, laboratory studies on female emergence and pheromone release, male emergence and mate-locating, and female and male mating behaviors were conducted. Mate acquisition and copulation occurred on the soil surface near the female emergence site, with both sexes engaging in pheromone-mediated behaviors after having emerged from the soil. A highly stereotyped female pheromone release, or calling, behavior was observed, consisting of insertion of the female's head into the soil and elevation of the tip of her abdomen into the air. Bioassays conducted in a wind tunnel that simulated a turf fairway environment showed that walking and flying were both important in the upwind response of males to females. Mating and copulation occurred without an obvious complex courtship, but observations of postmating behaviors suggested that mate guarding occurs.     

Effects of methoxyfenozide-treated surfaces on the attractiveness and responsiveness of adult leafrollers

Abstract The attractiveness and responsiveness of the adult redbanded leafroller, Argyrotaenia velutinana (Walker), and obliquebanded leafroller, Choristoneura rosaceana (Harris), exposed to surfaces treated with the ecdysteroid agonist methoxyfenozide were investigated in wind tunnel and orientation tube assays. For both species, when males were exposed to water‐treated surfaces for 48 h, and regardless of what treatment surfaces the females had been exposed to, the mean percentage recaptures of such treated males in the wind tunnel assays were always significantly greater than the recaptures of males that had been exposed to methoxyfenozide. However, exposure to surfaces treated with surfactant (Latron B‐1956) also negatively affected the responsiveness (or attractiveness) of both moth species. In the orientation tube assays, males exposed to methoxyfenozide‐treated surfaces always had significantly lower mean levels of individuals exhibiting sexual excitability, and the mean distances traveled upwind, regardless of females’ exposure. In addition, exposure to surfactant‐treated surfaces did not affect mean levels of sexual excitibility and distances traveled upwind. The two bioassays demonstrated that male A. velutinana and C. rosaceana exposed to methoxyfenozide‐treated surfaces were not as responsive to calling females (treated and non‐treated) as were the non‐treated males. Conversely, females exposed to methoxyfenozide‐treated surfaces were just as attractive to non‐treated males as water‐treated females. It appears that a male's ability to respond to a calling female is more negatively affected by the ecdysone agonist than a female's ability to call and attract males.     

Oriented flight responses of female Mediterranean fruit flies to calling males, odor of calling males, and a synthetic pheromone blend

Male Mediterranean fruit flies,Ceratitis capitata (Wiedemann), produce volatile chemicals thought to be attractive sex pheromones. We demonstrated for the first time that male odor elicits upwind flight and zigzagging upwind flight patterns in mature unmated females. Such flight patterns indicate the mechanisms involved in female location of the pheromone source and arrival at that source. Similar female oriented upwind flight responses occurred with a three component blend comprised of ethyl-(E)-octenoate, geranyl acetate, andE,E-alpha farnesene. These findings clarify the role of male sex pheromone in mate-location strategies in this species and provide new bioassay criteria for evaluating attraction responses to male pheromone and synthetic blends.     

Examining disruption of pheromone communication in Choristoneura rosaceana and Pandemis limitata using microencapsulated (Z)-11-tetradecenyl acetate applied in a laboratory flight tunnel

Pheromone‐based mating disruption of lepidopteran pests (Tortricidae) of pome fruits using hand‐applied dispensing systems has become standard management practice for many producers in western North America. Sprayable microencapsulated (MEC) pheromone formulations that enable the application of pheromone controls with other orchard sprays and assist in the development of multispecies mating‐disruption systems are currently under development. Responses of male Choristoneura rosaceana (Harris) and Pandemis limitata (Robinson) (Lepidoptera: Tortricidae) to calling females in clean air, and air treated with their major pheromone component (Z)‐11‐tetradecenyl acetate (Z11‐14:OAc), released from a 3M sprayable pheromone formulation containing proprietary 3M Phase I microcapsules, applied at doses of 1, 10, and 100 mg of active ingredient (ai) m^?2 to the upwind end of a flight tunnel (equivalent to field rates of 10, 100, and 1000 g ai ha^?1) were compared in laboratory flight tunnels. In both species, disorientation was found to be dose‐dependent, because relative to male orientation to calling females in clean air, the orientation of male P. limitata was disrupted 23.3, 46.3, and 71.3%, and orientation by male C. rosaceana was disrupted 31.6, 37.7, and 45.8% by treatment doses of 1, 10, and 100 mg m^?2, respectively. Latency of male responses to calling females in a background of Z11‐14:OAc relative to responses in clean air was also dose‐dependent. Albeit short, the disruption lasted 26, 74, and 218 h in P. limitata and 30, 54, and 174 h in C. rosaceana at each application rate, respectively. Disruption by pheromone treatment was greater in P. limitata than in C. rosaceana. This difference may be correlated with species’ differences in the pheromone release rates of females. Mechanisms of disruption invoked by this 3M MEC pheromone formulation are discussed in relation to issues of its longevity and observed differences in the effects against the two species. It appears possible to evaluate relative activity of MEC pheromones in a laboratory setting which may aid in development of new formulations for mating disruption.     

The sublethal effects of tebufenozide on the precopulatory and copulatory activities of Choristoneura fumiferana and C. rosaceana

The sublethal effects of tebufenozide, an ecdysone agonist, on the reproductive biology of Choristoneura fumiferana (Clem) and of Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae), treated during the larval stage, were evaluated using two treatment methods: the force‐feeding method and the diet method. The percentage of mortality and the developmental time of survivors increased linearly with the concentration of tebufenozide used. This ecdysone analogue proved to be more toxic to C. fumiferana than to C. rosaceana. In C. rosaceana, the weight of males and females decreased proportionally with the dose ingested, but females were affected to a greater extent. This difference might be due to a greater consumption of the treated diet, or to a differential vulnerability to tebufenozide. Tebufenozide did not modify the pre‐copulatory activities associated with chemical communication in the females. However, the consumption of tebufenozide delayed ovarian maturation, causing a reduction in the fecundity of females. Treated males had smaller spermatophores and fewer eupyrene sperms in their bursa copulatrix and spermatheca, along with lower mating success. In C. fumiferana, tebufenozide delayed the females’ onset time of calling the first night after emergence, but did not affect the mean time spent calling or the production of the main component of the sex pheromone. The males showed significantly greater difficulty in executing oriented flight in a wind tunnel, although their mating success was not affected. We concluded that tebufenozide interferes with various aspects of the reproductive biology of males and females of C. fumiferana and C. rosaceana, including some pre‐copulatory behaviors associated with sex pheromone communication.     

Behaviors of Naïve vs. Pheromone-Exposed Leafroller Moths in Plumes from High-Dosage Pheromone Dispensers in a Sustained-Flight Wind Tunnel: Implications for Mating Disruption of These Species

Brief exposures of male Choristoneura rosaceana and Argyrotaenia velutinana to the plumes generated by lures releasing 3-component pheromone blends specifically tuned for each species or by commercially distributed Isomate OBLR/PLR Plus pheromone rope dispensers induced markedly different subsequent behavioral responses to pheromone. A greater proportion of C. rosaceana males took flight and successfully oriented toward lures 24 h after preexposure to a lure, a rope, or the lure–rope combination in a sustained-flight wind tunnel compared to naïve moths. Flights were also longer for preexposed than naïve moths. Preexposed male C. rosaceana were not more likely to fly toward ropes 24 h after preexposure. By contrast, fewer male A. velutinana oriented to lures 24 h after preexposure than did naïve moths. Those preexposed A. velutinana successfully locking onto plumes from lures flew for significantly shorter intervals than did unexposed moths. Electroantennograms revealed no changes at the periphery 15 min and 24 h after preexposure. For A. velutinana, the long-lasting effect was decreased attraction to a lure and increased attraction to a rope. For C. rosaceana, pheromone preexposure increased responsiveness to its authentic blend. This behavioral evidence is sufficient to explain why sexual communication of C. rosaceana is more difficult to disrupt than that of A. velutinana. Furthermore, it suggests a more complete blend of pheromone may be necessary to disrupt the former species but not the latter when using rope dispensers.     

Effects of Wind Speed and Atmospheric Pressure on Mate Searching Behavior in the Aphid Parasitoid Aphidius nigripes (Hymenoptera: Aphidiidae)

The effects of wind speed and atmospheric pressure on male mate searching behavior, modulated by a female sex pheromone, were investigated in the aphid parasitoid Aphidius nigripes (Hymenoptera: Aphidiidae). Male A. nigripe generally did not reach females at wind speeds of 100 cm/sec, as the majority of individuals taking flight in the pheromone plume (81.8%) were unable to sustain upwind flight. At lower wind velocities, male responsiveness to females generally decreased with distance from the source. However, wind speeds approaching the upper threshold (100 cm/sec) tended to eliminate this distance effect. Therefore, there appears to be a trade-off between the need for higher wind speeds to detect the pheromone source from long distances, and a reduction in male flight capacity as wind velocity increases. Our results also indicate that chemical communication in A. nigripes could be affected by variations in atmospheric pressure, as we observed a relationship between pressure fluctuations in the 24 hr prior to testing and male responsiveness to females. The importance of these abiotic factors on mate searching behavior is discussed within the context of the reproductive biology of A. nigripes.     

Mate-location flight of the red-necked longicorn beetle, <Emphasis Type="Italic">Aromia bungii</Emphasis> (Coleoptera: Cerambycidae): an invasive pest lethal to Rosaceae trees

The red-necked longicorn beetle, Aromia bungii (Faldermann) (Coleoptera: Cerambycidae), which has recently invaded Japan, is a serious pest species. The larvae kill cherry and orchard Rosaceae trees, such as peach, plum, and apricot. To clarify their features for mate location, male and female flight behaviors were observed in wind tunnels with caged males and females as lures. In a small wind tunnel (50 cm in diameter, 2 m in length), both sexes showed takeoff behavior according to increasing airflow. The rate of female takeoff against the male lure, a live male in a wire-netting cage, tended to be higher than for other combinations. In a large wind tunnel (ca. 1.6 m in diameter, 4 m in length), both sexes ascended and showed momentary hovering and astatic flight. When male lure cages were placed windward in the wind tunnel, females showed “upwind flight,” while males did not. It is assumed that females were induced to fly upwind by a factor derived from live males, likely to be an airborne pheromone component.     

Disruption of pheromone communication of Choristoneura rosaceana (Lepidoptera: Tortricidae) using microencapsulated sex pheromones formulated with horticultural oil

Sprayable, microencapsulated (MEC) sex pheromone formulations represent a promising tool for achieving mating disruption, yet often lack sustained effectiveness in the field, making repeated applications necessary. This study evaluated the impact of adding Purespray Green horticultural oil as an adjuvant to 3M MEC-LR, an MEC formulation of (Z)-11-tetradecenyl acetate, on disruption of mate-finding behavior in Choristoneura rosaceana (Harris) in small-plot trials in experimental apple orchards. Treatments consisting of MEC-LR in water, MEC-LR in water + 2% (vol:vol) Purespray Green, and a water control were applied to plots of apple using an airblast sprayer at a rate of 100 g of pheromone/ha. Disruption caused by foliar treatments was evaluated over a 7-wk period using mark-release recapture experiments in the field and concurrent bioassays in a flight tunnel. Disruption of orientation to 2-d-old, calling, virgin females was used as a measure of treatment effect in all experiments. Both pheromone alone and pheromone + oil treatments significantly disrupted male mate-finding behavior for a period of > or =21 d in flight tunnel assays and > or =42 d in mark-recapture field trials. The addition of oil did not significantly enhance the disruption activity nor increase the longevity of the MEC pheromone formulation. Our results show the compatibility of spraying MEC pheromone with a horticultural oil, and techniques for applying an oil-pheromone formulation to maximize the control impact of this combination are discussed.     

旋花蛾性信息素成分的鉴定

提取和鉴定了杂草田旋花Convolvulus arvensis的天敌旋花蛾Tyta luctuosa的性信息素主要成分,并应用风洞实验对雄蛾进行了测试。结果表明,腺体提取物和雌蛾求偶时所释放的主要性信息素成分均为2种化合物：顺-9-十四碳烯醛（Z9-14∶Ald）和顺-11-十六碳烯醛（Z11-16∶Ald）。腺体提取物中2种化合物的总量在个体间差异很大（22～167 ng）,但二者的比率相对稳定（Z9-14∶Ald/Z11-16∶Ald=0.3±0.15）。雌蛾所释放的性信息素量及比率在个体间也存在差异。每只雌蛾求偶时平均每小时释放94 ng Z9-14∶Ald和45 ng Z11-16∶Ald,平均比率为2.2。风洞实验结果显示,48%的雄蛾可被合成的性信息素混合物（0.4 μg Z9-14∶Ald, 2 μg Z11-16∶Ald）引诱完成逆风飞行并最终触及刺激源,而2种对照组（以求偶期雌蛾或性信息素腺体提取物为刺激源）的引诱率则分别为62%和44%。     

Attractiveness of year-old polyethylene Isonet sex pheromone dispensers for Lobesia botrana

In this study, we have compared the release of sex pheromone from mating disruption dispensers exposed in the field for 12 months and from calling females. The main pheromone component of the grapevine moth, Lobesia botrana (D. and S.) (Lepidoptera: Tortricidae), is (E)‐7,(Z)‐9‐dodecadienyl acetate, and a minor component is (Z)‐9‐dodecenyl acetate. Aged dispensers from two different years emitted a much higher amount of both pheromone components than calling females. However, the summer temperature during field exposure influenced the release from mating disruption dispensers the following year. In the wind tunnel, male L. botrana were equally attracted to 12‐month, field‐exposed dispensers, a standard monitoring pheromone lure, and to synthetic (E)‐7,(Z)‐9‐dodecadienyl acetate sprayed at the rate of 0.6–60 ng h^?1. Field trapping tests confirmed that aged dispensers from both years were at least as attractive to L. botrana males as a standard monitoring pheromone lure. The possible contribution of previously applied dispensers to the mating disruption efficacy during following applications is discussed.     

Sublethal exposure to methoxyfenozide-treated surfaces reduces the attractiveness and responsiveness in adult oriental fruit moth (Lepidoptera: Tortricidae)

The chemical communication (female attractiveness and male responsiveness) of adult oriental fruit moth, Grapholita molesta (Busck) (Lepidoptera: Tortricidae), exposed to surfaces treated with the ecdysteroid agonist methoxyfenozide for 48 h were investigated in two laboratory wind tunnel assays. The recapture assay examined the ability of treated males to orient to a single cage of treated females, and the data gathered were mean percentage of males recaptured per treatment. The male sexual behavior assay examined some specific orientation behaviors (associated with sexual excitability) of treated males when they were given a choice of two competing pheromone sources (cages of treated females), and the data gathered were mean time males spent in upwind plume orientations and at source contact (female cage) per treatment. Data from the recapture assay suggests that exposure to methoxyfenozide impacts male responsiveness more than female attractiveness. In contrast, data from the sexual behavior assay strongly revealed that exposure to methoxyfenozide-treated surfaces does negatively impact both the ability of calling females to attract males and of aroused males to display sustained upwind flight behavior and time spent at the female cages.     

Pheromone pre-exposure and mating modulate codling moth (Lepidoptera: Tortricidae) response to host plant volatiles

Abstract 1 Two codling moth Cydia pomonella kairomonal attractants, ethyl (E,Z)‐2,4‐decadienoate (pear ester) and (E)‐β‐farnesene, were tested in an insecticide‐sprayed apple orchard and an orchard treated for mating disruption with synthetic pheromone (E,E)‐8,10‐dodecadienol (codlemone). Male captures with pear ester were higher in the pheromone‐treated than in the insecticide‐treated orchard, whereas captures with (E)‐β‐farnesene were not different. Subsequent wind tunnel experiments confirmed that pre‐exposure to sex pheromone codlemone increased the behavioural response of codling moth males to pear ester. This supports the idea that male attraction to the plant volatile pear ester and sex pheromone codlemone is mediated through the same sensory channels. 2 Pear ester is a bisexual codling moth attractant and even captures of female moths were significantly increased in the pheromone‐treated orchard. In the laboratory wind tunnel, pheromone pre‐exposure had no effect on female response to pear ester, but significantly more mated than unmated codling moth females flew upwind towards a pear ester source. Differences in mating status in insecticide‐treated vs. pheromone‐treated orchards may thus account for the differences in female trap captures with pear ester. 3 These findings are important with respect to monitoring of codling moth with pear ester in mating disruption orchards. They also emphasize the importance of host plant volatiles in pheromone‐mediated mating disruption, which has been neglected to date.     

Factors influencing the landing of maleEpiphyas postvittana (Walker) exhibiting pheromone-mediated flight (Lepidoptera: Tortricidae)

The effects of changes in various visual and olfactory properties of a white card surface on the landing position of male Epiphyas postvittanaexhibiting pheromone-mediated flight were studied in a wind tunnel. Males landed predominantly at the most downwind position of a surface in line with the pheromone source, regardless of the strength of the source. The position on the surface that males landed was strongly influenced by visual factors. The landing position of males appeared to be influenced by visual cues along all three axes of the surface. Decreases in either the dimension horizontally perpendicular to the wind direction or the vertical dimension resulted in greater numbers of males landing farther upwind on the surface than the downwind edge. Visual changes in the axis along the wind direction also affected the position at which males landed. For example, when presented with two white card surfaces with a 4- cm gap between them, males tended to land on the downwind edge of the upwind surface (on which the source was located). When the gap was bridged with clear Mylar, the landing pattern was significantly different, with the greater proportion of males landing on the downwind surface. However, when Mylar was placed on the plexiglass floor of the tunnel (in addition to bridging the gap), the landing pattern on the surface was not significantly different from that on the two surfaces without the Mylar bridge. It is suggested that during the prelanding and landing phases of pheromone-mediated flight, male moths orient to visual features of the surface containing the pheromone source rather than to visual features of the source (conspecific female moth) itself.     

Behaviour of male oriental fruit moth, Grapholita molesta, in overlapping sex pheromone plumes in a wind tunnel

The behaviour of Grapholita molesta (Busck) (Lepidoptera: Tortricidae) males was studied in overlapping sex pheromone plumes in a wind tunnel. The ultimate aim of the study was to assess the effect of different treatments on male behaviour and consider the observed changes within the context of the suggested mechanisms underlying mating disruption. Two baits were placed either in series or parallel using both synthetic pheromone blends and female extracts. One bait, the reference containing (Z)-8-dodecenyl acetate/(E)-8-dodecenyl acetate/(Z)-8-dodecenol in a ratio of 100/6/10 was kept constant at a dose of 100 g of the main component, giving a composition and a release rate close to that of a female. The dose of the other bait varied between 0.1 and 100 times the concentration of the reference and was a mixture of one, two or three pheromone components. Males clearly discriminated between different blends and doses in the overlapping plumes, for regardless if the lures were presented in series or in parallel they followed the complete plume. Complete suppression of the response to the reference was only achieved with 300 g of the optimal three-component blend on the other lure. When tested singly, a bait consisting of Z8-12:OAc/E8-12:OAc/Z8-12:O Hin a 100/0.2/0.4 ratio, attracted a high proportion of the males when placed 75 cm upwind of the male release site, but no males from 150 cm. Our data suggest that complete pheromone blends should be more effective than any incomplete blends in mating disruption formulations for G. molesta.     

Effects of pheromone plume structure and visual stimuli on the pheromone-modulated upwind flight of male gypsy moths (Lymantria dispar) in a Forest (Lepidoptera: Lymantriidae)

The pheromone-modulated upwind flight ofLymantria dispar males responding to different pheromone plume structures and visual stimuli designed to mimic trees was video recorded in a forest. Males flying upwind along pheromone plumes of similar structure generated tracks that were similar in appearance and quantitatively similar in almost all parameters measured, regardless of the experimentally manipulated visual stimuli associated with the pheromone source. Net velocities, ground speeds, and airspeeds of males flying in point-source plumes were slower than those of males flying in the wider, more diffuse plumes issuing from a cylindrical baffle. The mean track angle of males flying in plumes issuing from a point source was greater (oriented more across the wind) than that of males flying in plumes issuing from a transparent cylindrical baffle. Males flying in point-source plumes also turned more frequently and had narrower tracks overall than males responding to plumes from a cylindrical baffle. These data suggest thatL. dispar males orienting to pheromone sources (i.e., calling females) associated with visible vertical cylinders (i.e., trees) use predominantly olfactory cues to locate the source and that the structure of the pheromone plume markedly affects the flight orientation and the resultant track.     

Relationship between sex pheromone elicited behaviour and response of single olfactory receptor neurones in a wind tunnel

Responses from pheromone‐specific receptor neurones in male Agrotis segetum (Denis & Schiffermüller) (Lepidoptera: Noctuidae) were recorded in a laboratory wind tunnel. Stimuli were: (1) rubber septum dispensers loaded with single components or a four‐component pheromone blend, (2) excised glands from female A. segetum, (3) constrained A. segetum females with extruded glands. Dose–response curves for three neurone‐types with different specificity were established. The neurones were specifically tuned to respond to either one of the two pheromone components (Z)‐5‐decenyl acetate and (Z)‐7‐dodecenyl acetate, or to the behavioural antagonist (Z)‐5‐decenol. In parallel, a behavioural dose–response curve with males flying upwind to a four‐component pheromone blend was established. There was a clear correlation between behavioural arrestment of upwind flight and maximum spiking activity in Z5–10:OAc‐specific neurones. The pheromone release rates of individual females and synthetic dispensers were compared. A load of 50–200 ng of Z5–10:OAc on a rubber septum elicited approximately the same neural response as one female gland.     

Flight manoeuvers used by a parasitic wasp to locate host-infested plant

Cotesia rubecula Marshall (Hymenoptera: Braconidae) is a specialist larval parasitoid of the butterfly Pieris rapae L. which itself feeds almost exclusively upon cruciferous plants. Female wasps are attracted to the odour of host-infested plant (plant-host complex: PHC) and the probability of flights in a wind tunnel depends on females' prior oviposition experience with the PHC and on the concentration of the PHC odour. This study considers the effect of both factors on characteristics of oriented flight upwind towards the PHC. The flight track parameters that we measured and calculated were not significantly affected by these factors. C. rubecula females exhibited high average flight velocity and relatively straight flight tracks. There was a considerable variability between individuals, however, in their odour-modulated upwind flight tracks. Some females generated a zigzagging upwind flight track similar to those commonly observed from male moths responding to female sex pheromone. Other females flew along a straight track directly upwind. The flight tracks of most female wasps were intermediate between these extremes. The full range of these flight performances was observed to all experimental treatments.     

Detection of short pure-tone stimuli in the noctuid ear: what are temporal integration and integration time all about?

The interception of a pheromone filament induces flying moths to surge briefly nearly straight upwind; in the absence of pheromone moths cease upwind progress and zigzag crosswind. We tested males of the almond moth, Cadra cautella (Lepidoptera, Pyralidae), in a low-turbulence wind tunnel in wind velocities of 20, 40 and 80 cm s⁻¹. A mechanical pulse generator was set to produce plumes either with same pheromone pulse frequency (pulse generation frequency of 2.9 Hz, interpulse distances from 7 cm to 28 cm) or plumes with same interpulse distance across the three wind velocities (interpulse distance of 14 ± 2 cm, pulse generation frequency of 1.7–5.0 Hz). In plumes of similar pulse frequency, the faster the speed of the wind the slower the ground speed of flight. However, in plumes of similar interpulse distance, ground speed remained relatively constant independent of the wind speed. A `realized' frequency of pulse interception for males flying along the various combinations of pulse frequencies and wind velocities was calculated using the males' average airspeed and the spatial distribution of pheromone pulses in the plume. Realized frequency of pulse interception ranged from 1.3- to 3.0-fold higher than the frequency of pulse generation. The flight tracks of males reflected the regime of realized pulse interception. These results suggest that upwind flight orientation of male C. cautella to pheromone in different wind velocities is determined by the flux of filament encounter. Accepted: 3 September 1997