Effect of a vertical object on the sex pheromone-mediated flight and landing behaviors of the male lightbrown apple moth,Epiphyas postvittana (walker) (Lepidoptera: Tortricidae)

A small (2.5-cm-wide) vertical (10-cm-high) white object influenced the sex pheromone-mediated flight and landing behaviors of maleEpiphyas postvittana. When the vertical object was positioned on a horizontal surface to the side (3–5 cm) and upwind of a pheromone source (in the middle of the surface), the distribution of landing positions of males on the surface was different from that when the object was not present; males tended to land in positions skewed toward the side of the source that the object was on. The closer the object was positioned to the source, the greater the number of males that landed on the object (rather than on the horizontal surface). This difference in landing positions (when the object was present) corresponded with changes in the flight tracks; the tracks of males flying to the surface with an object were skewed toward the object and had higher amplitude intertrack reversal distances than the tracks of males flying to a surface without a vertical object. Positioning of a vertical object progressively upwind of the source resulted, apparently, in decreased effects on the landing (and presumably flight) behavior of males. The effect of the vertical object on the flight and landing behaviors of males corresponded largely with changes in pheromone plume structure (visualized with smoke) induced by the extra turbulence in the airflow over the source. Thus it appears that the vertical object influences the behavior of maleE. postvittana largely through the olfactory sensory modality. However, when a clear, Mylar object, in place of the white object, was placed on the surface, more males landed on the Mylar object (than did on the white object), suggesting that the vertical object may also influence the behavior of males through the visual modality.     

Pheromone-mediated optomotor anemotaxis and altitude control exhibited by male oriental fruit moths in the field

Abstract. In the field over short grass, pheromone-stimulated oriental fruit moth males, Grapholita molesta (Busck), flying under high windspeeds tended to steer courses more into the wind and to increase their airspeeds compared with those flying in low windspeeds.Thus, optomotor anemotaxis enabled the males to steer relatively consistent upwind track angles and to maintain an upwind progress of between c. 50–100 cm/s despite variable wind velocities.Zigzagging flight tracks were observed at both 10 m and 3 m from the source, as were tracks with no apparent zigzags.Transitions from casting to upwind flight or vice-versa were observed.The durations of the intervals between reversals during both upwind zigzagging flight and casting were consistent with those observed in previous wind-tunnel experiments.The control of altitude was more precise during upwind zigzagging flight than during casting.In general, the side-to-side deviations in the tracks were greater than the up-and-down deviations, with both the side-to-side and vertical distances and their ratios being consistent with previous wind-tunnel studies of pheromone-mediated flight.One difference between the field and laboratory flight tracks was that males in the field exhibited much higher airspeeds than in the wind tunnel.Males occasionally were observed to progress downwind faster than the wind itself, and further analysis showed that they were steering a downwind course in pheromone-free air following exposure to pheromone, which is the first time this has been recorded in moths.We propose that such downwind flight may aid in the relocation of a pheromone plume that has been lost due to a wind-shift, by enabling the moth to catch up to the pheromone as it recedes straight downwind away from the source.     

Impact of (Z)-7-dodecenol and turbulence on pheromone-mediated flight manoeuvres of male Trichoplusia ni

Abstract. Turbulence and chemical noise are two factors which may influence pheromone-mediated flight manoeuvres of a moth in natural habitats. In this study, the effects of turbulence and the behavioural antagonist (Z)-7-dodecenol on flight manoeuvres of male Trichoplusia ni (Hübner) were evaluated in a wind tunnel. Male moths increase airspeed and course angles when turbulence is increased. This leads to significant increases in the length of flight tracks, but significant reductions in the time taken to reach a pheromone source. In less disturbed pheromone plumes, distributions of course angles and track angles of male T.ni show a prominent peak centred about 0° relative to the upwind direction, indicating that moths can temporarily steer directly upwind toward a pheromone source.
When (Z)-7-dodecenol is released 10 cm upwind of a pheromone source to form an overlapping plume downwind, course angles, airspeeds and ground-speeds of male T.ni are reduced significantly compared with those in uncon-taminated pheromone plumes. This results in a longer flight time to reach a pheromone source. The decrease in flight speed would decrease the rate of contact with filaments, and thereby perhaps allow the moth to detect uncon-taminated pheromone filaments independently from filaments containing the behavioural antagonist.     

Three-dimensional analysis of aphid landing behaviour in the laboratory and field

The final second of the landing approach of black bean aphids, Aphis fabae, was analysed in three dimensions using video techniques. A yellow landing platform was placed upwind or downwind from aphids aggregating under a ceiling light in a laboratory wind tunnel with 10, 20, 30, 40 or 50 cm s^–1 wind speeds, and up-tunnel or down-tunnel in still air. As individual aphids flew to the platform, body orientation (assessed by direct observation) was predominantly into-wind whether the initial flight direction to the landing platform was upwind or downwind. A greater proportion showed into-wind body orientation as wind speed increased. Flight track parameters which differed significantly between wind speeds were the track length, linear start to finish distance, linearity index, horizontal ground speed, speed vertical to the ground, vertical turning rate, and horizontal turning rate. The position of the landing platform was important for track length, linear start to finish distance, horizontal ground speed, three-dimensional turning rate, horizontal turning rate, vertical turning rate, and sinuosity. As wind speed increased above 30 cm s^–1 the ground speed became more consistent and indicated considerable variation in air speed to adjust for ground speed. For the majority of aphids there was a strong preference (88%) for into-wind landings with initial upwind directed flight, while for downwind flights a significant number (55%) of insects reversed initial flight direction and landed into-wind. Field recorded landings showed that 66% of aphids landed into-wind and there was a mean bearing to the wind of 71 ± 42°, a similar finding to wind-tunnel studies.     

Olfactory orientation responses of the eucalyptus woodborer, Phoracantha semipunctata, to host plant in a wind tunnel

The eucalyptus woodborer, Phoracantha semipunctata Fabricius (Coleoptera: Cerambycidae), attacks mainly species of Eucalyptus (Myrtaceae). This study investigated walking and flight behaviour of P. semipunctata males and females exposed to an odour plume originating from a log of E. globulus placed vertically in the upwind end of a wind tunnel. In control experiments, beetles were exposed to a PVC drainpipe in the same position as the log, providing a visual stimulus without host‐tree odour. No statistical differences were found between behavioural responses of either sex when exposed to the log or PVC pipe. No beetles landed on the PVC pipe, whereas 49% of the beetles exposed to host‐tree odour plume landed on the log. Beetles aged over 24 days after emergence from the host tree were more responsive than beetles aged 20–24 days, and accounted vor 86% of the beetles that landed on the log. While walking, host‐tree odour affected the behaviour of the beetles that landed on the log as follows: upwind movement and path linearity increased, whereas turning rate, stopping frequency, mean stopping time and time to take‐off flight decreased. During flight, host‐tree odour affected the behaviour of the beetles that landed on the log as follows: increased upwind flight, turning rate, flight time, flight distance, and decreased flight speed. For beetles that never lost contact with the odour plume, flight progressed upwind with narrow zigzags, and showed higher directedness upwind, path linearity, faster flight speed and lower turning rate than for beetles that lost contact with the odour plume. After loosing contact with the plume, beetles tended to decrease their upwind progression, exhibiting a sharp turn or quick counterturns followed by crosswind or downwind excursions. This led to regaining contact with the odour plume and resumed upwind progression at higher speed provided they flew within the boundaries of the plume. The results showed that host‐tree odour affects both walking and flight behaviour of P. semipunctata beetles, inducing a more directed upwind movement and landing on the visual stimulus of a tree trunk.     

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.     

Investigations of Mechanisms of Pheromone Communication Disruption of Choristoneura rosaceana (Harris) in a Wind Tunnel

Although atmospheric treatment with synthetic pheromone is used commercially to control several lepidopteran pests, little is known about how mate-finding behavior is altered by this procedure. Mechanisms of disruption of the mate-finding behavior of Choristoneura rosaceana (Lepidoptera: Tortricidae) were examined in wind tunnel bioassays. Male moths were exposed to synthetic sex pheromone or pheromone components prior to or during upwind flight to a calling virgin female. The proportion of males successfully contacting a calling female was reduced only when the synthetic pheromone treatment was presented simultaneously. A synthetic source of the attractive four-component pheromone placed upwind of a calling female was more effective than a less attractive two-component blend in achieving disorientation. Habituation of the central nervous system does not appear to be a significant factor in disruption of mate-finding behavior of male C. rosaceana, as exposure to pheromone prior to the bioassay did not alter the proportion of males that achieved subsequent upwind flight. Disruption of mate-finding behavior of C. rosaceana in a wind tunnel is probably the result of a combination of mechanisms including adaptation of antennal receptors, camouflage of the female-produced plume, and false-trail following which contributed to the additive disruption effect observed with the most attractive four-component pheromone.     

Influence of plume structure and pheromone concentration on upwind flight of Cadra cautella males

Abstract. The effects of pheromone plume structure and its concentration on the pheromone-mediated flight of male Cadra cautella (Lepidoptera: Phycitinae) were investigated in a laminar-flow wind tunnel. When two C. caurella males flew simultaneously along a ribbon plume of mixed smoke and pheromone, their inflight behaviour was dependent on the instantaneous structure of the plume they encountered. When a male intercepted an intact ribbon filament, he sustained a crosswind course, whereas when he intercepted a turbulent filament (created by an upwind male fragmenting the ribbon plume), he adopted a flight course more due upwind. These results indicate that C. cautella males altered their in-flight manoeuvres in response to instantaneous changes in the fine structure of the pheromone plume. We also demonstrated that differences in the fine structure of the plume had more influence on the flight pattern of C. cautella males than a 1000-fold range in pheromone dose. The size of the plume was increased by adding wind deflectors upwind of the pheromone source, independent of source dosage, males following ribbon plumes flew slow zigzag tracks, whereas males following large, turbulent plumes flew directly to the source in fast, straight tracks with less counterturning.     

Attraction range of a sex pheromone trap for the damson‐hop aphid Phorodon humuli (Hemiptera: Aphididae)

The attraction range of olfactory response by winged female gynoparae (autumn migrants that give birth to oviparae, the sexual females) and male damson–hop aphids Phorodon humuli (Schrank) is investigated in field experiments over 2 years by analyzing the spatial patterns of catches in concentric circles of yellow‐painted traps (60 in total) around a central trap releasing the species' sex pheromone, (1RS,7S,7aS)‐nepetalactol. Males are more likely than females to be found in the central trap, with 65.6% of the 1824 males caught there compared with 11.2% of 1346 females. Both morphs are more numerous in traps axial with the mean wind direction and centred on the pheromone‐release trap than at other angles. Males are approximately five‐fold more numerous in traps downwind than at similar distances upwind of the pheromone, showing that its presence stimulates landing. For males, the estimated active space of the lure extends 6 m downwind. Catches of females are equally numerous up and downwind of the pheromone lure because females orienting on the axis of the pheromone source continue to respond to visual cues in their flight path if they overshoot the olfactory one. For females, the active space of a pheromone lure is less than 2 m downwind. It is unimportant for either morph whether the pheromone‐release trap is yellow or transparent. In these experiments, both morphs orient with, track and probably arrive in the pheromone source trap from at least 26 m, the distance to the nearest aphid‐infested hops.     

Light versus pheromone-bearing wind in the control of flight direction by bark beetles, Scolytus multistriatus

ABSTRACT. A wind tunnel lit from above was used to measure the relative effectiveness of light as opposed to pheromone-bearing wind in directing the flight of male S.multistriatus. Upward flight in the tunnel was overridingly phototactic not geotactic, while horizontally upwind flight toward the pheromone source was anemotactic not chemotactic, since the pheromone concentration was uniform in the tunnel. The flight tracks of newly-emerged unfed and previously unflown beetles were preponderantly upwards and downwind; the longer the beetles had spent in flight activity beforehand, the more of them flew upwind.     

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.     

Trapping and sex pheromone-mediated flight and landing behaviour of maleCtenopseustis obliquana

A sex pheromone-baited delta trap was found to be inefficient at eliciting landing and entering of the trap by maleCtenopseustis obliquana. The inefficiency of the delta trap related to turbulence altering the pheromone plume and the concomitant effect on the flight manoeuvres of male moths. In the wind tunnel, high proportions of males flew upwind and landed on the sides, outside, of the trap, but only a relatively small proportion of these males entered the trap and contacted the sticky surface. When males approached the delta trap, they tended to fly in wide zigzags (i.e., large inter-track reversal distances) and at an altitude near the top of the trap, where the trap was relatively narrow in width (compared to the bottom). Thus, these flight manoeuvres largely precluded males from entering the trap. Greater numbers of male moths entered the trap when: (i) the front barriers of the delta trap were removed, (ii) pheromone dosage was increased to 300 μg, and (iii) the trap design was changed to a rectangular one. The first two changes appeared to influence the flight manoeuvres of males (who appeared to fly with narrower inter-track reversal distances), while the third change apparently did not affect the glight manoeuvres of males, but rather allowed more males to enter the trap because of the greater area of the entrance. The low trap catches of maleC. obliquana in the field are also a consequence of the glue which is currently used. After contact with this glue most males are able to escape, flying off the sticky surface and losing contact with the pheromone. A field trial found that delta traps with another glue caught more than three times the number of males of the related tortricid moth,Epiphyas postvittana, than delta traps with the currently used glue.     

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.     

Orientation of flying male Anobium punctatum (Coleoptera: Anobiidae) to sex pheromone: separating effects of visual stimuli and physical barriers to wind

Abstract. Flying male woodworm, Anobium punctatum , are able to locate a source of female pheromone offered as a point source but land more quickly if the same odour source is presented on a model 'tree' (cylinder). We tested the response to a two-dimensional black 'tree' marked as a silhouette on the upwind screen of the wind tunnel, with or without a hidden upwind barrier to the wind. The beetles responded equally to pheromone presented on a visual silhouette with wind barrier and to the hidden upwind wind-barrier alone, suggesting that the wind-barrier itself has a significant effect on insect orientation and landing: visual stimuli do not add to the landing response if the barrier is present. However, without the barrier upwind, landing was faster when the visual stimulus was present than without. This is the third forest-living species to be shown to respond in this way, which suggests this phenomenon may be more widespread. The possibility that this phenomenon is present in other, non-forest, phytophagous insects, their predators and parasitoids is discussed.     

Landing Surface Color Preferences of Spathius agrili (Hymenoptera: Braconidae), a Parasitoid of Emerald Ash Borer, Agrilus planipennis (Coleoptera: Buprestidae)

The color preferences for landing surfaces were examined for Spathius agrili (Hymenoptera: Braconidae), a parasitic wasp introduced for biocontrol of emerald ash borer, Agrilus planipennis (Coleoptera: Buprestidae). Lures with the 3-component pheromone blend of male S. agrili were used to activate upwind flight by virgin female S. agrili in a laminar flow wind tunnel. Paper discs with halves of two different colors (combination pairs of black, white, red, yellow, green, or purple), with the pheromone lure in the center, were tested to quantify preferences for landing on one color over another. Females landed preferentially on green, yellow, and white surfaces, and landed the least frequently on red, black, and purple surfaces. Changes in color preferences due to adjacent colors were observed and discussed.     

Elevation of pheromone response threshold in almond moth males pre-exposed to pheromone spray

Abstract. High percentages of naive Cadra cautella (Walker) (Lepidoptera: Pyralidae) males not pre-exposed to pheromone flew upwind to sources containing 50 ng (83%) and 500 ng (97%) of pheromone, but not to sources containing 5 μg (23%) and 50 μg (4%).Of the naive males that flew upwind in response to 50 ng sources, 67% located and landed on the source, whereas fewer than 19% of the naive males that flew upwind in response to higher doses located and landed on the sources.A 2-minute pre-exposure of C.cautella males to a spray cloud containing 50 ng, 500 ng, 5 μg or 50 μg of pheromone, induced shifts in response levels such that in wind-tunnel bioassays performed 1 h later, there was an increase in the doses that optimally elicited upwind flight and landing on the source that was proportional to the pre-exposure dose.Few of the pre-exposed males flew upwind to (10–43%) and landed on (0–33%) 50 ng sources, whereas they now perferentially flew upwind to(58–81% and 52–73%) and landed on (33–68% and 55–60%) pheromone sources of doses of 500 ng and 5 μg, respectively.Therefore pre-exposure to pheromone promoted a shift of threshold for response, and not an overall reduction in responsiveness to pheromone.     

Behaviour of flying oriental fruit moth males during approach to sex pheromone sources

Abstract. The pheromone-modulated upwind flight tracks of Grapholita molesta (Busck) males were video recorded as they approached a point-source of pheromone in a wind tunnel. The field of view of the video recording was divided longitudinally into 33 cm sections and the flight behaviour of the males in these sections was measured and compared as they approached from 233 cm to 50 cm downwind of the pheromone source. As the males approached the source, their mean ground speeds decreased. The mean values of their track angles increased with respect to due upwind (0), indicating movement more across the wind. These changes resulted mainly from the males decreasing their air speeds as they progressed up the plume toward the source. They did not change the average direction of their steering (course angle). Thus, the increase in track angles resulted from the males allowing themselves to drift more in the wind as they approached the odour source. The males also increased their average rate of counterturning as they approached the source. The net result of all these behavioural changes was a track that slowed and grew narrower, giving the impression that the males were 'homing-in' on the pheromone source as they approached. Causes of these systematic changes in behaviour are considered with respect to the known systematic changes in pheromone plume structure as the distance to the source decreases.     

Field and laboratory electroantennographic measurements of pheromone plume structure correlated with oriental fruit moth behaviour

Abstract. Peak-to-trough electroantennogram amplitudes (bursts), caused by the individual filaments of a plume of female pheromone, diminish as high-emission-rate sources are approached by male Grapholita molesta , and this reduction is correlated with in-flight arrestment (ceasing to advance upwind). These findings are consistent with the hypothesis that one cause of in-flight arrestment in response to high-concentration point sources is the attenuation of the peak-to- trough amplitudes close to the source. High burst frequency, high pheromone flux, or low levels of continuous neuronal activity all are less well correlated with arrestment. Rather, arrestment appears due to a reduction of chemosensory input to the CNS during flight up the plume, even though the actual molecular concentration continues to increase. In a laboratory wind tunnel, upwind flight initiation by more than 20% of males was elicited only by pheromone source concentrations evoking significant fluctuations in EAG amplitudes at downwind release points. The burst frequencies that evoked high levels of upwind flight initiation ranged from a mean of 0.4-2.2 bursts/s. Because a previous study revealed that flying male G. molesta change their course angle within 0.15 s of losing or contacting pheromone, these EAG burst frequencies indicate that during flight in a pheromone plume, many manoeuvres are probably made in response to contact with individual plume filaments. Thus, upwind flight tracks may be shaped by hundreds of steering reactions in response to encounters with individual pheromone filaments and pockets of clean air. Field-recorded EAGs reveal that burst amplitudes diminish from 3 to 30 m downwind of the source, whereas burst frequencies do not, averaging c. 1/s at 3, 10 and 30 m downwind.     

Visual feedback in the control of pheromone-mediated flight ofHeliothis virescens males (Lepidoptera: Noctuidae)

MaleHeliothis virescens (F.) (Lepidoptera: Noctuidae) were made to fly into a uniformly white and translucent tube within a large wind tunnel while responding to sex pheromone. Different visual patterns placed within the tube greatly affected the ability of the male moths to maintain upwind progress or remain oriented to the wind while in contact with the plume. Over 89% of males attempting to fly through a blank tube, lacking visual patterns, became disoriented, the males gaining or losing altitude and repeatedly hitting the sides of the tube. Patterns of 20–40 dots placed on the sides of the tube at or slightly above plume level resulted in high levels of sustained upwind flight (47–74%) relative to patterns placed directly below (30–40%), directly above (35%), or slightly below the level of the flight path (26–44%). Optimal upwind progression in pheromone-responding males occurred when image motion could be resolved both transversely (T), orthogonally to the longitudinal axis of the body relative to the horizontal plane of the environment, and longitudinally (L), along the body axis. Even very sparse patterns (single rows of dots) could elicit high levels of sustained upwind flight (53–63%) when positioned within the tube such that the males' movements would create both L and T image motion. However, successful negotiation of the tube was also unexpectedly facilitated by patterns apparently providing no horizontal transverse component for flying males but providing longitudinal flow while centering the moth in the plume through a symmetrical left-right input (4–40%).     

Orientation of male gypsy moths,Lymantria Dispar (L.), to pheromone sources: The role of olfactory and visual cues

The role of olfaction and vision in the close-ranging flying and walking orientation of male gypsy moths, Lymantria dispar(L.), to females was studied in the forest and in the laboratory. In the forest, feral males found an isolated pheromone source as readily as one supplemented with female visual cues; dead, acetonerinsed females deployed without pheromone received virtually no visitations. In flight tunnel choice experiments using cylinders as surrogate trees and pheromone in different spatial configurations, visual attributes of the female did not influence either the males' choice of landing site or the efficiency with which they located the female. Rather, the presence of pheromone on the cylinder was necessary to elicit orientation as well as landing and walking on the cylinder. When a female visual model was placed in various positions around a pheromone source, walking males oriented primarily to the chemical stimulus. There were, however, indications that males would alter their walking paths in response to female visual cues over short distances (<5 cm), but only if they continued to receive pheromone stimulation. When visual and chemical cues were abruptly uncoupled by altering the trajectory of the pheromone plume, most males responded to the loss of the odor cue rather than to visual cues from the female. Temporal pheromone stimulation patterns affected male walking orientation. When stimulated by pheromone, males oriented toward the source; loss of the odor cue prompted an arearestricted local search characterized by primarily vertical and oblique movements with frequent reversals in direction. Presumably these maneuvers enhance the likelihood of recontacting the plume or serendipitously encountering the female. The apparent lack of visual response to the female is discussed in light of morphological and behavioral evidence suggesting that gypsy moths were formerly nocturnal.