Response of both sexes of Trichoplusia ni (Lepidoptera: Noctuidae) to virgin females and to synthetic pheromone

Abstract. 1. Male and female Trichoplusia ni were caught at three locations in California in 1974 and 1975, in traps baited with virgin female T. ni or with synthetic female pheromone.
2. 71% of the females were captured between 20.00 and 21.00 hours; only 22% of the males.
3. Both mated and unmated females were trapped.
4. Comparisons of live trap vs. dead trap catches suggest that females are responding to the virgin females (or synthetic pheromone) and not to a combination of stimuli from the first trapped males plus female pheromone.     

Vertical distribution of codling moth adults in pheromone-treated and untreated plots

The vertical distribution of codling moth,Cydia pomonella (L.) within pheromone-treated and untreated apple and pear orchard canopies was determined using tethered virgin females, unbaited sticky traps, and blacklight observation of released moths. Mating of virgin females tethered at various heights in untreated orchard canopies increased with placement height from 1–4 m. Application of pheromone dispensers for mating disruption at 2 and 4 m above the ground greatly decreased mating. Greatest capture of males and females on unbaited sticky traps occurred at mid- and upper-canopy heights. Total capture of males and females in pheromone-treated plots was not statistically different than in untreated plots. The percentage of mated females captured on sticky traps did not vary with trap height or pheromone treatment. Released moths marked with flourescent powder and observed at dark with a blacklight indicated that moths are primarily distributed high in the canopy. However, males shifted to a position lower in the canopy when pheromone dispensers were placed 2.1 m above the ground. Results suggest that pheromone dispensers be placed in the upper canopy for optimal disruption of codling moth mating.     

Pheromone‐trapping the nun moth,Lymantria monacha (Lepidoptera: Lymantriidae) in Inner Mongolia,China

The nun moth, Lymantria monacha L., is one of the most important defoliators of Eurasian coniferous forests. Outbreaks during 2011–2015 in the natural/planted larch, and larch‐birch mixed forests of the Greater Khingan Range in Inner Mongolia, China, caused tremendous timber losses from severe defoliation and tree mortality. A series of trapping experiments were conducted in these outbreak areas to evaluate the efficacy of a synthetic species‐specific pheromone lure based on the female pheromone blend of European nun moth populations. Our results clearly show that the nun moth in Inner Mongolia is highly and specifically attracted to this synthetic pheromone, with few gypsy moths (Lymantria dispar) captured. Flight activity monitoring of L. monacha male moths using pheromone‐baited Unitraps at 2 locations during the summer of 2015 indicated that the flight period started in mid‐July, peaking in early August at both locations. Based on male moth captures, there was a strong diurnal rhythm of flight activity throughout the entire scotophase, peaking between 22:00 and 24:00. Unitraps and wing traps had significantly and surprisingly higher catches than the gypsy moth traps. Unitraps fastened to tree trunks 2 m above ground caught significantly more male moths than those at the ground level or at 5 m height. Male L. monacha moths can be attracted to pheromone‐baited traps in open areas 150–200 m distant from the infested forest edge. Our data should allow improvement on the performance of pheromone‐baited traps for monitoring or mass‐trapping to combat outbreaks of this pest in northeastern China.     

Mating attraction by Stenotus rubrovittatus (Heteroptera: Miridae) females and its relationship to ovarian development

Stenotus rubrovittatus (Matsumura) (Heteroptera: Miridae) causes brown or black marks on rice, Oryza sativa L., grains (pecky rice), and it is becoming of increasing importance in Japan. Attractiveness of adult females or males of S. rubrovittatus to conspecific individuals was examined in the field in 2003 and 2004. Unmated female-baited traps captured significantly more males than did the unmated male-baited traps. However, the numbers of females captured by female- or male-baited traps were low, and they were not significantly different from the numbers caught by the control traps. No nymphs were captured by any traps. In 2004, we examined the effects of age and mating experience on female attraction ability with the goal of understanding the role of reproductive development in the observed behavior. The daily number of males captured by young unmated female (3-d-old)-baited traps increased from the first day until the fourth day of experiments, and then capture started to decrease. The peak in the number of captured males corresponded to the preoviposition period. When we observed ovarian development of S. rubrovittatus females under 25 degrees C and a photoperiod of 16:8 (L:D) h, we found that vitellogenesis had already started in approximately 30-50% of 1-d-old individuals. By the fifth day after emergence, 50-70% of individuals had mature eggs. These results indicated that the attractiveness of females is the strongest when egg laying becomes possible. Therefore, S. rubrovittatus females attract males selectively for mating, and it is probable that females use a sex pheromone for the attraction.     

Effect of rearing strategy and gamma radiation on field competitiveness of mass-reared codling moths (Lepidoptera: Tortricidae)

We compared the field competitiveness of sterile codling moth, Cydia pomonella (L.), males mass-reared through diapause or standard production protocols and treated with either 150 or 250 Gy of gamma radiation. Evaluations were performed during spring and summer 2003 by using field release-recapture tests. Released males were recaptured using traps baited with synthetic pheromone or with virgin females. In addition, mating tables baited with virgin females were used in the summer to assess the mating competitiveness of the released moths. Field performance of released males was significantly improved by rearing through diapause and by lowering the dose of radiation used to treat the insects. These effects were observed during spring when evening temperatures were relatively cool and in summer when evening temperatures were high. These effects were observed regardless of the sampling method (i.e., capture in pheromone-baited traps, virgin female-baited traps, or in mating tables). There were significant interactions between larval rearing strategy and radiation dose with respect to day of recapture. The effect of rearing strategy on male performance was observed immediately after release, whereas the effect of dose of radiation was usually delayed by 2-3 d. In general, the best treatment for improving codling moth male field performance was a combination of rearing through diapause and using a low dose of radiation (150 Gy). The difference in performance when insects were treated with 150 or 250 Gy was greater when males had been reared using standard (nondiapause) rearing protocols, suggesting that diapause rearing may attenuate some of the negative effects of the higher doses of radiation.     

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

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

Monitoring grape berry moth (Paralobesia viteana: Lepidoptera) in commercial vineyards using a host plant based synthetic lure

For some Lepidopteran pests, such as the grape berry moth Paralobesia viteana (Clemens), poor correlation between males captured in traps baited with sex pheromone and oviposition activities of female moths has called into question the value of pheromone-based monitoring for these species. As an alternative, we compared the capture of female and male grape berry moth in panel traps baited with synthetic host volatiles with captures of males in pheromone-baited wing traps over two growing seasons in two blocks of grapes in a commercial vineyard in central New York. Lures formulated in hexane to release either 7-component or 13-component host volatile blends captured significantly more male and female grape berry moth on panel traps compared with the numbers captured on panel traps with hexane-only lures. For both sexes over both years, the same or more moths were captured in panel traps along the forest edge compared with the vineyard edge early in the season but this pattern was reversed by mid-season. Male moths captured in pheromone-baited wing traps also displayed this temporal shift in location. There was a significant positive correlation between captured males and females on panel traps although not between females captured on panel traps and males captured in pheromone-baited traps for both years suggesting pheromone traps do not accurately reflect either female or male activity. Male moths captured in pheromone traps indicated a large peak early in each season corresponding to first flight followed by lower and variable numbers that did not clearly indicate second and third flights. Panel trap data, combining males and females, indicated three distinct flights, with some overlap between the second and third flights. Peak numbers of moths captured on panel traps matched well with predictions of a temperature-based phenology model, especially in 2008. Although effective, panel traps baited with synthetic host lures were time consuming to deploy and maintain and captured relatively few moths making them impractical, in the current design, for commercial purposes.     

Factors influencing the effectiveness of an attracticide formulation against the Oriental fruit moth, Grapholita molesta

An attracticide formulation, LastCall?OFM, was tested against the Oriental fruit moth Grapholita molesta (Busck) (Lepidoptera: Tortricidae) in replicated small plot field trials in apple, Malus domestica (Borkhausen), orchards in South‐eastern Pennsylvania, USA. Attracticide treatments were applied using a calibrated hand pump, and treated plots were compared to similar untreated plots. Male moth activity was monitored using virgin female‐baited traps, and the potential for reduction in mating activity was assessed using sentinel virgin females. A comparison of application rates showed that 1500 droplets per ha of the attracticide formulation was as effective as 3000 droplets per ha, and both application rates reduced captures in synthetic pheromone‐baited traps for prolonged periods. Droplets placed either at high or low positions within the canopy significantly reduced trap capture and mating with sentinel females. In addition, the only sentinel females that mated in the treated plots were located in the untreated portion of the tree canopy. Mate finding behaviour was equally disrupted by formulations with and without insecticide. Therefore, under the test conditions, the mechanism by which the attracticide formulation worked was by disruption of male orientation, and not by the removal of males due to insecticide poisoning. Two field cage experiments tested the impact of population density on the competitiveness of the attracticide formulation compared to virgin females. A significant proportion of males were captured in female‐baited traps at the highest female‐to‐droplet ratio tested. Equal proportions of males were captured in attracticide‐baited traps at male moth densities of 10, 20, 40, and 80 males per cage. These results clarify some of the factors influencing the effectiveness and possible mechanisms of an attracticide management tactic against the Oriental fruit moth.     

Mating behavior of the coffee leaf-miner Leucoptera coffeella (Guérin-Mèneville) (Lepidoptera: Lyonetiidae)

Despite the importance of Leucoptera coffeella (Guérin-Mèneville) in coffee production worldwide, there is a lack of information on its reproductive biology. This knowledge will help in mass rearing, as well as support the development of behavioral control techniques for this insect. The purpose of the study was to determine the periodicity of mating and male capture and describe the mating behavior L. coffeella. In laboratory, we observed the periodicity of mating with virgin couples of different ages, zero to five days after emergence. Male activity was studied in a 0.7 ha coffee plantation, cv. Catuaí, where Delta traps were installed at 0.5 m above ground, using either virgin females or rubber septa lured with the synthetic sex pheromone. The sequence of mating behavior was studied by making visual observations and recorded of pairs placed on individual plastic tubes. Mating occurred between 4h and 6h of photophase, when the highest frequencies involved pairs with ages of one and three days after emergence, with peak of mating occurring in 5th hour of photophase. The young or old pairs showed significantly copulation frequency and the peak of matings advance in 1h. The highest male capture occurred at 12p.m. and 13 p.m. by traps with virgin females or traps with synthetic sex pheromone lures, respectively. L. coffeella is one insect with diurnal mating and the mating behavior was not different from what is know for other Lepidoptera species.     

Factors affecting captures of male citrus leafminer, Phyllocnistis citrella Stainton, in pheromone-baited traps

The citrus leafminer, Phyllocnistis citrella Stainton (Lepidoptera: Gracillariidae), is an important world‐wide pest of citrus. Larval mining within leaf flush impacts yield and predisposes trees to infection by citrus canker, Xanthomonas axonopodis pv. citri. The present series of studies sought to identify factors affecting male P. citrella catch in pheromone‐baited traps with the intent of developing effective monitoring. A commercially available pheromone lure (Citralure, ISCA Technologies, Riverside, CA, USA) was highly effective in attracting male P. citrella to traps. Pherocon VI Delta (Trécé Inc., Adair, OK, USA) traps baited with a Citralure captured more male P. citrella than identically baited Pherocon IC Wing traps (Trécé Inc.). The superiority of the Delta‐style trap was found to be due to a 3 cm long closing latch that likely prevents males from flying directly through the trap without capture. Within canopies of mature citrus trees (approximately 3.5 m high), traps at mid‐canopy height (2.0 m) captured more males than traps placed higher (3.5 m) or lower (0.6 m). On the canopy perimeter and in between canopies, traps near ground level (0.6 m height) captured more males than traps at 2.0 and 3.5 m heights. Male catch was greater within the tree canopy or on the canopy perimeter than 2.0 away from the canopy. Traps deployed in trees on the edge of groves captured more males than traps placed 120 and 240 m away from the grove edge and within the grove interior. In non‐pheromone‐treated grove plots, the optimal dosage for catching males was between 0.1 and 1.0 mg of the 3 : 1 blend of (Z,Z,E)‐7,11,13‐hexadecatrienal and (Z,Z)‐7,11‐hexadecadienal; however, in pheromone‐treated plots a higher 10.0 mg dosage lure was most effective. Male catch in pheromone‐baited traps exhibited a diel rhythm with most males captured during scotophase (22:00–23:00 h) and no males captured during photophase.     

Sex ratio and female sexual status of the coconut pest, Oryctes monoceros (Coleoptera: Dynastidae), differ in feeding galleries and pheromone-baited traps

Oryctes monoceros is a serious coconut pest, causing up to 40% damage in tropical Africa. Synthetic aggregation pheromone, ethyl 4-methyloctanoate, has been used to lure adults to traps. Traps with pheromone plus decaying palm material captured a high proportion of males. This raises the question whether individuals, which damage palms are receptive to the pheromone. We studied the sex ratio of the insects feeding on coconuts and those attracted to pheromone traps. Sixty two percent of adults from feeding galleries on living coconut palms were females. Pheromone with rotting palm material lured 43% females. To investigate the reasons for this difference, we compared the reproductive system of females lured to the odour traps or feeding in coconut galleries, or present in old rotting stems. Ninety six percent of the females trapped by pheromone had mated, and were sexually mature. In the galleries on living palms, 46% of females were immature, and 24% had not mated. In old rotting stems where eggs are laid and larvae develop, a mixture of 52% mated and 48% virgin females was found. Therefore, the pheromone together with the odour of rotting coconut stems signals a reproduction site to beetles, particularly mature females. In practice, the pheromone-baited traps will help in reducing the dissemination of gravid females, but will not affect directly the numbers of immature ones attacking palms. Our results show that when using pheromones for monitoring or controlling insects, the physiological status of the insects may have unexpected effects on the outcome.     

Development and optimization of methods for using sex pheromone for monitoring the mealybug Planococcus ficus (Homoptera: Pseudococcidae) in California vineyards

The sex pheromone of the vine mealybug Planococcus ficus Signoret has been identified as a single component, lavandulyl senecioate. Racemic lavandulyl senecioate was as attractive to male mealybugs as the insect-produced (S)-enantiomer, indicating that the unnatural enantiomer is not inhibitory. Lavandulol, which also was found in extracts from virgin females, antagonized attraction of males at higher doses. Rubber septum lures loaded with 10- to 1,000-microg doses of the pheromone were equally attractive, and lures loaded with 100 microg of racemic pheromone remained attractive for at least 12 wk under field conditions. Delta traps were more effective than double-sided sticky cards and minimized captures of nontarget insects. Pheromone-baited traps had an effective range of at least 50 m. Comparison of visual sampling methods and sampling of males with pheromone-baited traps revealed that trap catches were significantly correlated with the results from visual sampling methods, and with economic damage.     

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

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

The relative performance of pheromone and light traps in monitoring the seasonal activity of both sexes of the eastern hemlock looper, Lambdina fiscellaria fiscellaria

The seasonal flight activity of both sexes of the eastern hemlock looper, Lambdina fiscellaria fiscellaria Guenée (Lepidoptera: Geometridae) was studied during two consecutive years in Quebec and Newfoundland, using light (L), pheromone (P) and combined light and pheromone (LP) traps. Moth density significantly affected the performance of the different traps, with P traps being more effective at low than high density. However, P trap catches decreased just prior to the onset of female captures, probably as a result of competition between traps and virgin calling females. Nearly all females caught in L and LP traps were already mated and even the first females caught had laid at least half of their egg complement. In Quebec under warm nights, the pattern of male activity occurred at different times, with peak P catches being later in the scotophase than those of L traps, but overall similar numbers of males were caught in both traps. In contrast, under cool nights, males were caught early in the night in both P and L traps, suggesting a strong competition effect between traps, although more males were caught in P than L traps overall. In Newfoundland, the pattern of male captures in L and P traps was similar at both high and low temperatures, so competition between trap types would always be high. Under these conditions P traps were more effective than L traps. Irrespective of the region, year or temperature, significantly more males were captured in LP, with the effect of L and P being additive. In both regions, females responded similarly to L and LP traps with peak activity occurring early in the night. Captures of females were lower than those of males under cool temperatures, suggesting that the temperature threshold for flight is higher for females. The use of L and P traps simultaneously and/or in combination is discussed in relation to integrated pest management programs and ecological considerations.     

Should I stay or should I go? Modelling dispersal strategies in saproxylic insects based on pheromone capture and radio telemetry: a case study on the threatened hermit beetle <Emphasis Type="Italic">Osmoderma eremita</Emphasis>

To predict how organisms cope with habitat fragmentation we must understand their dispersal biology, which can be notoriously difficult. We used a novel, multi-pronged approach to study dispersal strategies in the endangered saproxylic hermit beetle Osmoderma eremita, exploiting its pheromone system to intercept high numbers of dispersing individuals, which is not possible with other methods. Mark-release-recapture, using unbaited pitfall traps inside oak hollows and pheromone-baited funnel traps suspended from tree branches, was combined with radio telemetry (in females only) to record displacements. Dispersal, modelled as a probability distribution of net displacement, did not differ significantly between sexes (males versus females recaptured), observation methods (females recaptured versus radio-tracked), or sites of first capture (pitfall trap in tree versus pheromone trap – distance from original dispersal point unknown). A model including all observed individuals yielded a mean displacement of 82 m with 1% dispersing > 1 km. Differences in body length were small between individuals captured in pitfall versus pheromone traps, indicating that dispersal is rarely a condition-dependent response in O. eremita. Individuals captured in pheromone traps were consistently lighter, indicating that most dispersal events occur relatively late in life, which agrees with trap catch data. In addition, most (79%) females captured in pheromone traps were mated, showing that females typically mate before leaving their natal tree. Our data show that integrating odour attractants into insect conservation biology provides a means to target dispersing individuals and could greatly improve our knowledge of dispersal biology in threatened species.     

Evidence of female-produced sex pheromone of Adelphocoris suturalis (Hemiptera: Miridae): effect of age and time of day

After the reduction in the use of broad-spectrum insecticides, Adelphocoris suturalis (Hemiptera: Miridae) has become an important pest of transgenic cotton, Gossypium hirsutum L., in China. A. suturalis is a highly mobile insect, moving rapidly in and out of crop fields, and traps baited with sex pheromones may be useful in monitoring and controlling its numbers. However, little is currently known regarding the chemical communication based on sex pheromones by this species. To use the synthesized sex pheromone effectively as part of the integrated pest management, some fundamental biological parameters under which the pheromone is produced must be understood. In this study, attractiveness of adult females or males of A. suturalis to conspecific individuals was examined by Y-tube bioassay; we also investigated the female sexually mature age and circadian rhythm of male attraction to females of A. suturalis in the field in 2008 and 2010. Y-tube bioassay results indicated that only males were attracted to odors from sexually mature virgin females. Two-year field trapping experiments showed that 4-18-d-old virgin females effectively attracted males, and males were attracted to virgin females during the scotophase, with a peak between 1900 and 2300 hours. These results improve our understanding of the sexual communication behavior of A. suturalis and provide evidence of female-produced sex pheromone in this species.     

Temporal variation in abundance of male and female spruce budworms at combinatory associations of light traps and pheromone traps

A 3‐year study (2014–2016) was conducted at Rocky Harbour near the west coast of Newfoundland, Canada, to record the abundance and phenology of adult spruce budworms captured at traps, using a factorial design (light traps and pheromone traps deployed contiguously or segregated spatially). Budworms were most abundant and occurred seasonally earlier in 2014 than in 2015 and 2016; these findings held generally true for males and females. The geographic setting of Newfoundland (large island isolated from the mainland by an oceanic barrier of >100 km across) provides an ideal location to discriminate local flight from long‐range immigrations; in our study, however, immigrations cannot be ruled out for any single day of trapping due to broad overlap in emergence patterns at Rocky Harbour relative to forest stands with known populations of budworms on the mainland. Based on moderate daily variation in adult abundance, however, major immigration events (defined as external deposition of budworms with large numerical amplitude) likely did not take place at Rocky Harbor between 2014 and 2016. Males were more abundant at light traps coupled with pheromone traps, whereas abundance of males at pheromone traps was similar with or without contiguous light traps. This outcome may be mediated by lower range of attraction for light traps (usually <100 m) and (generally assumed to be several hundreds of meters). Females were equally abundant at light traps with or without pheromone traps. As expected, males were captured earlier in the season at pheromone traps than at light traps, and females occurred later in the season due to protandry. The onset of flight observed at light traps or pheromone traps in 2015 and 2016 occurred 10–15 days later than simulated predictions; caution is thus warranted as to conclusions derived on computer modeling of adult emergence.     

Identification of the Aggregation Pheromone of the Melon Thrips,Thrips palmi

The objective of this study was to identify the aggregation pheromone of the melon thrips Thrips palmi, a major pest of vegetable and ornamental plants around the world. The species causes damage both through feeding activities and as a vector of tospoviruses, and is a threat to world trade and European horticulture. Improved methods of detecting and controlling this species are needed and the identification of an aggregation pheromone will contribute to this requirement. Bioassays with a Y-tube olfactometer showed that virgin female T. palmi were attracted to the odour of live males, but not to that of live females, and that mixed-age adults of both sexes were attracted to the odour of live males, indicating the presence of a male-produced aggregation pheromone. Examination of the headspace volatiles of adult male T. palmi revealed only one compound that was not found in adult females. It was identified by comparison of its mass spectrum and chromatographic details with those of similar compounds. This compound had a structure like that of the previously identified male-produced aggregation pheromone of the western flower thrips Frankliniella occidentalis. The compound was synthesised and tested in eggplant crops infested with T. palmi in Japan. Significantly greater numbers of both males and females were attracted to traps baited with the putative aggregation pheromone compared to unbaited traps. The aggregation pheromone of T. palmi is thus identified as (R)-lavandulyl 3-methyl-3-butenoate by spectroscopic, chromatographic and behavioural analysis.     

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.     

Mate Finding Via a Trail Sex Pheromone by <Emphasis Type="Italic">Aphytis melinus</Emphasis> DeBach (Hymenoptera: Aphelinidae) Males

Finding mates is frequently problematic for parasitoid wasps. In some parasitoid species, males rely on volatile, airborne sex pheromones for locating mates, while in others they rely on contact, trail sex pheromones. This study sought to shed light on the mate finding mechanism of males of Aphytis melinus. Specifically, the goal was to determine whether A. melinus males use airborne or contact pheromones, or both, for locating mates. The study showed that A. melinus males rely on a contact, trail sex pheromone for locating mates: A. melinus males responded to substrate-borne cues left by virgin females, while they did not respond to airborne cues from virgin females. Specifically, males more frequently encountered virgin females when the females walked across an arena to a fixed encounter point compared to when they were manually placed at the encounter point, and spent greater than expected time on surfaces previously visited by virgin females compared to control surfaces not visited by females. In contrast, males did not respond to airborne cues from virgin females in an airflow olfactometer nor to traps baited with virgin females in the field, and spent similar lengths of time on surfaces visited by newly-mated or 24-h mated females versus control surfaces not visited by females. The main effect of the trail sex pheromone on the behavior of A. melinus males was to direct their search and, so, increase the likelihood of encountering mates. This effect apparently is not preceded by longer-range attraction of males via an airborne female sex pheromone. Overall, the results of this study support a hypothesis in which A. melinus males searching on substrates on which females may be present rely exclusively on a trail sex pheromone to locate mates.