Intimate Rendezvous in a Tritrophic Context? Nothing but the Girls for Male Lysiphlebus testaceipes

In insects, mating often occurs after natal dispersal, and hence relies on a coevolved combination of sexual communication and movement allowing mate encounter. Volatile sex pheromones are widespread, generally emitted by females and triggering in‐flight orientation of conspecific males. In parasitoid wasps, unmated females can start laying unfertilized eggs via parthenogenesis so that host patches could serve as sites of rendezvous for mating. Males could therefore use cues associated with host patches to focus their search on females that have successfully found oviposition sites. We hypothesized that in parasitoids exploiting herbivorous hosts, sex pheromones, and herbivore‐induced plant volatiles (HIPV) should act in synergy, triggering male orientation toward ovipositing females. We tested this hypothesis with the aphid parasitoid Lysiphlebus testaceipes. Results from both field and laboratory experiments show that males are strongly attracted to virgin females, but that volatiles from aphid‐infested plants have no effect on male orientation, neither has a cue, nor in interaction with the female sex pheromone. The absence of synergy between sex pheromones and HIPV contrasts with results on other species and raises interesting questions on mating systems and sexual selection in parasitoid wasps.     

Navigation by Male Crab Spiders <Emphasis Type="Italic">Misumenoides formosipes</Emphasis> (Araneae: Thomisidae): Floral Cues May Aid in Locating Potential Mates

The crab spider Misumenoides formosipes is an ambush predator whose males search for relatively sedentary females within a heterogeneous habitat. Females are receptive to mating immediately after their adult molt and a male biased adult sex ratio together with precopulatory guarding places a premium on male ability to locate females quickly. It is unknown what cues males use to find females; we report here on male movements in association with floral cues. Males in field trials moved towards inflorescences more frequently when both visual and chemical cues were available, than when chemical cues were eliminated. Males in lab trials chose an inflorescence over leaf substrates even in the absence of visual cues. These findings support the hypothesis that M. formosipes males could utilize floral chemistry as a navigational cue in mate searches.     

Host plant volatiles serve to increase the response of male European grape berry moths, <Emphasis Type="Italic">Eupoecilia ambiguella,</Emphasis> to their sex pheromone

The European grape berry moth is an important pest in vineyards. Males respond to the female-produced sex pheromone released from a piezo nebulizer in a dose-dependent manner in a wind tunnel: <50% arrive at the source at 5–50 pg/min (underdosed), 80% arrive at 100 pg/min to 10 ng/min (optimal) and <20% arrive at 100 ng/min (overdosed). Males responding to overdosed pheromone show in flight arrestment at 80 cm from the source. Host plant chemostimuli for Eupoecilia ambiguella increase the responses of males to underdosed and overdosed pheromone. (Z)-3-hexen-1-ol, (+)-terpinen-4-ol, (E)-β-caryophyllene and methyl salicylate released with the underdosed pheromone cause a significant increase in male E. ambiguella flying to the source. Time–event analysis indicates a positive correlation between faster activation and probability of source contact by the responding males. The four host plant compounds added to the overdosed pheromone permitted males to take off faster and with a higher probability of flying to the source. This suggests that perception of host plant products with the sex pheromone facilitates male E. ambiguella to locate females on host plants, lending credence to the hypothesis that plant products can signal rendezvous sites suitable for mating. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

The evolution of the mating system of the Carpenter bee Xylocopa varipuncta(Hymenoptera: Anthophoridae)

The Carpenter bee, Xylocopa varipuncta , appears to have evolved a dispersed lek mating system. Males leave the natal nest in the late afternoon in March and April to hover for periods of up to two hours in the crowns of non-flowering trees growing in desert washes. Females and other males occasionally visit a hovering male, probably drawn by a sex pheromone. Resident males repel intruders of the same sex and attempt to induce females to alight on foliage, where mating occurs. The hovering territories do not contain nests, or potential nest sites, or food resources for females. Lek territoriality may have evolved in this species because nests are difficult to locate and the food-plants are evenly dispersed over a large area, making economical defence of patches of this resource unfeasible. Males are left with the option of demonstrating social dominance to potential mates through control of a superior landmark territory.     

Circadian Reproductive Rhythms, Pair-Bonding, and Evidence for Sex-Specific Pheromones in Nadezhdiella cantori (Coleoptera: Cerambycidae)

Mating and oviposition rhythms and evidence for sex-specific pheromones of the gray-black citrus longicorn beetle Nadezhdiella cantori were investigated. The reproductive and flight activities of this insect mainly occurred between 1700 and 2400 h, and each activity had two peaks with the first one taking place within the 3 h before sunset (2100 h) and the second occurring within the 3 h after sunset. Circadian rhythms of mating and oviposition were significantly correlated but neither mating nor oviposition was significantly correlated with flight. It is suggested that mating and oviposition repeat in turn during pair-bonding and that flight is an indication of termination of reproductive activities in some individuals of a population. Males differentiated between sexes by touching. Mating behavior of males was stimulated by a contact female sex pheromone on the female body surface. Fighting behavior of males was mediated by a chemical on the male body surface. Females left trails of the contact sex pheromone on the trees when walking, which males used to locate them. Implications of these findings are discussed.     

Sexual differences in the flight performance of the oriental fruit moth, Cydia molesta

Cydia molesta Busck (Lepidoptera: Tortricidae), a major pest of stone fruits and an increasingly important late season pest of apple, is predominately monitored by pheromone trapping of male moths. We investigated flight performance in relation to sex, age, and mating status using computer-linked flight mills, and also examined the relationship between female flight and reproduction. The crepuscular flight pattern of the experimental moths in relation to photointensity was very similar to that reported from field studies. Female moths significantly outperformed males in all measured flight parameters including total distance flown, distance of longest single flight, and velocity. The proportion of long-flying females (categorised as those completing an unbroken flight of greater than 1 km) was three to six times greater than that of males. Female flight performance was not related to mating status, but mated males displayed significantly greater flight than unmated males. The maximal flight period of mated females commenced on the third day after eclosion, following 30% egg deposition. Male flight was not significantly related to age. The data suggest that a limited proportion of the population, in particular females, may have the capacity to make inter-orchard flights. The limitations of monitoring C. molesta populations in apple orchards solely by pheromone trapping of males are discussed.     

Inter- and Intraspecific Effects of Volatile and Nonvolatile Sex Pheromones on Males, Mating Behavior, and Hybridization in Eretmocerus mundus and E. eremicus (Hymenoptera: Aphelinidae)

Eretmocerus species (Hym. Aphelinidae) are solitary parasitoids of Bemisia tabaci (Gennadius). Mate finding and mating behavior of two species, E. mundus and E. eremicus, were studied under laboratory conditions. We used three populations of Eretmocerus: typical arrhenotokous populations of E. eremicus (from USA) and E. mundus (from Spain), and an atypical thelytokous population of E. mundus (from Australia). We studied the intra- and interspecific responses of males to volatile and nonvolatile components of the female sex pheromones, mating behavior, and hybridization between populations and species. In both arrhenotokous populations, males reacted to volatile pheromones by walking toward conspecific virgin females. Males also reacted to nonvolatile pheromones by spending more time on and around patches on leaves of poinsettia plants that had been exposed to virgin females. Males of E. eremicus showed the same reaction to the nonvolatile sex pheromone of E. mundus females, but E. mundus males did not show any reaction to the nonvolatile sex pheromone of E. eremicus. There was no response of males of both species to thelytokous females of E. mundus. In both species three phases were distinguished in the mating behavior: premating, mating, and postmating. The duration of the phases differed between the three populations. Successful copulation between the two Eretmocerus species did not occur. In contrast, we recorded some successful copulations between Australian males and Spanish females of E. mundus, but they did not produce any hybrid females.     

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.     

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.     

Sex pheromone production and perception in the mating disruption-resistant strain of the smaller tea leafroller moth, Adoxophyes honmai

Mating disruption is an environmentally safe plant protection strategy that uses a synthetic copy of an insect pheromone to interfere with sexual communication and hence reproduction. To date, a number of pest moths have been controlled with applications of formulated pheromones as mating disruptants. Recently, however, the first example of resistance to mating disruption was documented in one of the major tea pests in Japan, the smaller tea leafroller moth, Adoxophyes honmai Yasuda (Lepidoptera: Tortricidae). To avoid other such cases, it is important to elucidate the mechanism(s) by which the disruptant lost its effectiveness. To this end, we imposed further selection by rearing field‐collected resistant insects with a synthetic pheromone in the laboratory. After more than 70 generations of selection, a strain with quite strong resistance was established, males of which could find and copulate with their mates even in the presence of 1 mg l^?1 of disruptant. Although the mating ability of this strain was greatly increased, the composition and blend ratio of the sex pheromone produced and emitted by females were not obviously changed in comparison with those of females sensitive to mating disruption. However, male response to the pheromone blend was markedly broadened after selection so that resistant males could locate a synthetic pheromone source even when it lacked a pheromone component that is normally necessary for attraction. Males capable of locking onto off‐ratio pheromone blends may be better able to find calling females in pheromone‐treated environments than narrowly tuned males because of greater capability of overcoming sensory imbalance.     

Nocturnal Calling Behavior in Mantids

Females of Mantis religiosa and Empusa pennata were video taped for several 24 h periods to determine if they showed behaviors associated with pheromone release. In the photophase the abdomen of both species was motionless and rested in continuous contact with the wings. However, at the beginning of the scotophase the females bent the abdomen ventrally so that the space between the abdomen and the wings increased significantly with respect to the daytime posture. Calling behavior (abdominal bending) was maintained throughout the 8 h scotophase and ended abruptly at lights on. Females of M. religiosa did not start calling until they were 30 days of age. Calling disappeared in mated females, but it reappeared two weeks later. Males stayed motionless in response to the odors emitted by other males or by noncalling females, but walked when a calling female was placed in the air flow. These observations suggest that female mantids bend their abdomens at night to release a sex pheromone. The adaptive function of nocturnal sex pheromone release in sexually cannibalistic species that rely strongly on visual cues for mating is discussed.     

同种雄性竞争对手的存在对星豹蛛雄蛛求偶和交配行为的影响

越来越多的研究发现,雄性产生精子(精液)也需付出代价。雄性除了依据配偶质量和竞争对手的竞争强度适应性调整生殖投入外,雄性在求偶和交配行为上也相应产生适应性反应,求偶和交配行为具有可塑性。目前雄性求偶和交配行为可塑性研究主要集中于雌性多次交配的类群中,在雌性单次交配的类群中研究甚少。以雌蛛一生只交配一次而雄蛛可多次交配的星豹蛛为研究对象,比较:(1)前一雄性拖丝上信息物质对后续雄蛛求偶和交配行为的影响,(2)雌雄不同性比对雄蛛求偶和交配行为的影响。研究结果表明,星豹蛛前一雄蛛拖丝上的信息物质对后续雄蛛求偶潜伏期、求偶持续时间和交配持续时间都没有显著影响,但前一雄蛛拖丝上的信息物质对后续雄蛛求偶强度有显著抑制作用。同时,性比对星豹蛛雄蛛求偶和交配行为都没有显著影响。可见,星豹蛛雄蛛对同种雄性拖丝上的化学信息可产生求偶行为的适应性调整,而对性比不产生适应性反应。     

Effects of sex pheromone in electrostatic powder on mating behaviour by Lobesia botrana males

Abstract:  In laboratory trials, mating behaviour of males of Lobesia botrana Den. and Schiff. (Lep., Tortricidae) was investigated after treatment with an electrostatically chargeable powder, Entostat^TM. Male moths were powdered with either blank Entostat or Entostat containing sex pheromone. Significantly more Entostat adhered to L. botrana males when sex pheromone was added to the powder compared with blank Entostat. Powdering male moths with Entostat, with or without sex pheromone, caused a significant reduction in antennal response when antennae were placed 2 cm away in downwind direction, while there was little effect when antennae were placed 25–100 cm from the EAG in downwind direction. In a flight study, powdering males of L. botrana caused significant increase in time before flight initiation and reduction in proportion of males making contact with calling females compared with untreated males. In a mating experiment in Petri dishes (to minimize required search time to locate female), significantly fewer females mated successfully (based on dissection of bursa copulatrix) when males had been treated with pheromone-loaded powder. Overall, powdering males of L. botrana caused considerable suppression of mating behaviour on various levels, and these suppressing effects were increased after adding sex pheromone to Entostat.     

Flirtation reduces males’ fecundity but not longevity

Theory predicts that due to limited resources males should strategically adjust their investment in reproduction and survival. Based on different conceptual framework, experimental designs, and study species, many studies support while others contradict this general prediction. Using a moth Ephestia kuehniella whose adults do not feed and thus have fixed resources for their lifetime fitness, we investigated whether males adjusted their investment in various life activities under dynamic socio‐sexual environment. We allowed focal males to perceive rivals or additional females without physical contact. We show that males do not adjust the number of sperm they transfer to mates in a given copulation at different immediate or both immediate and mean sperm competition levels. Contradictory to general predictions, our results demonstrate that cues from additional females increase males’ investment in courtship and reduce their lifetime number of copulations and sperm ejaculated, whereas cues from rivals have no effect on these parameters. Males have similar longevity in all treatments. We suggest that the sex pheromone produced by multiple females overstimulate males, increasing males’ costly flirtations, and reducing their lifetime copulation frequency and fecundity. This finding offers a novel explanation for the success of mating disruption strategy using sex pheromones in pest management.     

Host odours enhance the responses of adult Rhyzopertha dominica to male-produced aggregation pheromone

The behavioural responses of adult male and female Rhyzopertha dominica (Coleoptera, Bostrichidae) to blends of host volatiles and male-produced aggregation pheromone were observed in a four-arm airflow olfactometer. The odour sources used were five pheromone-releasing males each on a single maize grain (lower maize-volatiles blend), five pheromone-releasing males on 500 g of maize (higher maize-volatiles blend) or the host volatiles emanating from 500g of maize (maize volatiles alone). Multiple-choice tests, in which volatiles from all three odour sources were presented in the exposure chamber at the same time, were used to study odour preferences of the males and females. Both sexes showed strongest attraction to the higher maize volatile blend but there were significant sex differences in response to the odour sources. Males spent significantly more time than females in the zone with only maize volatiles, and females spent significantly more time in the higher maize-volatiles zone. However, when odour sources were offered singly, females gave numerically greater responses than males to all sources although this difference was statistically significant for only the lower maize-volatiles blend. As males are more attracted than females to host odours alone it is suggested that they may be more highly adapted to seek out new hosts while females are more inclined to locate a food source by following the pheromone signals produced by males. However, both sexes responded most strongly to the odour source comprising aggregation pheromone with the higher proportion of maize volatiles.     

Pheromones and body coloration affect mate recognition in the Japanese nine-spotted moth Amata fortunei (Lepidoptera: Arctiidae)

Adults of the Japanese nine-spotted moth, Amata fortunei, are diurnal and have white-spotted black wings and a black-and-yellow striped body pattern. We evaluated whether this species uses sex pheromones and whether visual cues from the female body are used in mate recognition. We introduced extracts of potentially scent-bearing abdominal tips of females to males. Males responded more to the extracts than to dried female specimens, suggesting the presence of sex pheromones in the extracts. Indeed, no males responded to the dried female specimens. To evaluate the importance of visual cues, we conducted the experiments with crude extract and an additional model stimulant. Males responded more to model females with the same number and similar area of yellow bands to the original conspecific females than to those with more bands and greater total band area, suggesting that dissimilarity in band number and area to conspecific females could interrupt male mating behavior. Males of A. fortunei likely find mating partners using olfactory cues over relatively long distances, while using both olfactory and visual cues over short distances. These results suggest that olfactory cues play a major role in mate recognition, whereas visual cues play a supplementary role.     

Observations on the mating behavior of the alfalfa blotch leafminer,Agromyza frontella (Rondani) (Diptera: Agromyzidae), and evidence of a female sex pheromone

The mating behavior of Agromyza frontella was studied under laboratory conditions. Adults were able to mate on the day of emergence, with no evident periodicity throughout the photophase. The host plant was essential for mating to occur, its presence affecting female receptivity rather than male copulatory behavior. Males generally entered a stationary phase once in the proximity of a female, before undertaking a final approach. This stationary behavior frequently resulted in male aggregations around a female, and under such conditions males exhibited a characteristic wing vibrating behavior. As male wing vibration was not an essential behavior for successful mating, and rarely occurred during male encounters in the absence of females or when only one male was near a female, it was considered as being primarily a male-male signal. The majority of females that mated exhibited an ovipositor pumping behavior that stimulated the male approach. However, such behavior was not essential to attract mates, as dead females elicited the entire sequence of male mating behavior. This suggested the presence of a cuticular sex pheromone, as reported for other species of higher Diptera. Whole virgin female (<24- h or 3- day-old) hexane extracts applied to male cadavers increased the time males spent on the plant, the number of contacts with the treated cadavers, the incidence of attempted copulations, and the wing vibrating behavior between males compared with controls using untreated cadavers. The results obtained indicate that females control copulation in A. frontella and that both semiochemical and visual cues are important in eliciting male mating behavior.     

Effects of social information on life history and mating tactics of males in the orb‐web spider Argiope bruennichi

Informed mating decisions are often based on social cues providing information about prospective mating opportunities. Social information early in life can trigger developmental modifications and influence later mating decisions. A high adaptive value of such adjustments is particularly obvious in systems where potential mating rates are extremely limited and have to be carried out in a short time window. Males of the sexually cannibalistic spider Argiope bruennichi can achieve maximally two copulations which they can use for one (monogyny) or two females (bigyny). The choice between these male mating tactics should rely on female availability that males might assess through volatile sex pheromones emitted by virgin females. We predict that in response to those female cues, males of A. bruennichi should mature earlier and at a smaller body size and favor a bigynous mating tactic in comparison with controls. We sampled spiders from two areas close to the Southern and Northern species range to account for differences in mate quality and seasonality. In a fully factorial design, half of the subadult males from both areas obtained silk cues of females, while the other half remained without female exposure. Adult males were subjected to no‐choice mating tests and could either monopolize the female or leave her (bigyny). We found that Southern males matured later and at a larger size than Northern males. Regardless of their origin, males also shortened the subadult stage in response to female cues, which, however, had no effects on male body mass. Contrary to our prediction, the frequencies of mating tactics were unaffected by the treatment. We conclude that while social cues during late development elicit adaptive life history adjustments, they are less important for the adjustment of mating decisions. We suggest that male tactics mostly rely on local information at the time of mate search.     

Mating behavior and thermoregulation of the reindeer warble fly,Hypoderma tarandi L. (Diptera: Oestridae)

Hypoderma (=Oedemagena) tarandi L. (Diptera: Oestridae) is characterized by a mating strategy in which both sexes meet and mate at two types of distinct topographical landmarks. In the expansive, treeless vidda (= tundra-like) biome, mating places are unique, rocky areas located along rivers and streams or in rocky areas of drying river and stream beds. In wooded valleys below the vidda, flies mated at certain topographical areas along dirt road tracks/paths. Thermoregulatory activities of males occupying perches at mating places included selection of substratum at perch site, orientation of body to sun's rays, crouching, stilting, and flights into upper cooler air. On warm sunny days males perched for just 1–2 min before flying up into cooler air to promote cooling. Laboratory and field studies revealed that flies could not metabolically cool down when held at 25–38°C. Time spent at mating places depended on temperature, duration of sunshine, and wind velocity. Males were very aggressive in pursuing allHypoderma-sized objects that passed by them or that landed near them, but they did not defend specific perch sites. Males either pursued and caught females in flight, or they hopped onto females that landed near them. During 5 years, 74 males and 14 females were seen at mating places. Dissection of six females caught at mating places revealed them to be recently eclosed flies full of fat body and with all eggs intact; two not paired with males were non-inseminated. Three experimentally paired females remainedin copulo for 10, 13, and 19.5 min.