Costs of female odour in males of the parasitic wasp <Emphasis Type="Italic">Lariophagus distinguendus</Emphasis> (Hymenoptera: Pteromalidae)

The display of female traits by males is widespread in the animal kingdom. In several species, this phenomenon has been shown to function adaptively as a male mating strategy to deceive sexual rivals (female mimicry). Freshly emerged males of the parasitic wasp Lariophagus distinguendus (Hymenoptera: Pteromalidae) are perceived by other males as if they were females because of a very similar composition of cuticular hydrocarbons which function as a sex pheromone in this species inducing courtship behaviour in males. Within 32 h, however, males deactivate the pheromone and are no longer courted by other males. In this paper, behavioural experiments were performed to test hypotheses on potential costs and benefits associated with the female odour in young males. We did not find any benefits, but demonstrated that young males were significantly more often outrivaled in male-male contests when competing with two older males for a female. Also, young males were significantly more often mounted in homosexual courtship events during these contests. Thus, display of female traits by males is not necessarily beneficial, and in fact, can be disadvantageous. We suggest that these costs have favoured the evolution of the pheromone deactivation mechanism in L. distinguendus males. The function of cuticular hydrocarbons as a female courtship pheromone in L. distinguendus might have evolved secondarily from a primary function relevant for both genders, and the deactivation of the signal in males might have caused a shift of specificity of the chemical signal from the species level to the sex level.     

A contact sex pheromone component of the emerald ash borer Agrilus planipennis Fairmaire (Coleoptera: Buprestidae)

Analyses of the elytral hydrocarbons from male and female emerald ash borer, Agrilus planipennis Fairmaire, that were freshly emerged vs. sexually mature (>10 days old) revealed a female-specific compound, 9-methyl-pentacosane (9-Me-C₂₅), only present in sexually mature females. This material was synthesized by the Wittig reaction of 2-decanone with (n-hexadecyl)-triphenylphosphonium bromide followed by catalytic reduction to yield racemic 9-Me C₂₅, which matched the natural compound by gas chromatography/mass spectrometry (retention time and EI mass spectrum). In field bioassays with freeze-killed sexually mature A. planipennis females, feral males spent significantly more time in contact and attempting copulation with unwashed females than with females that had been washed in n-hexane to remove the cuticular lipids. Hexane-washed females to which 9-Me-C₂₅ had been reapplied elicited similar contact time and percentage of time attempting copulation as unwashed females, indicating that 9-methyl-pentacosane is a contact sex pheromone component of A. planipennis. This is the first contact sex pheromone identified in the Buprestidae.     

Sexual selection on receptor organ traits: younger females attract males with longer antennae

Sexual selection theory predicts that female choice may favour the evolution of elaborate male signals. Darwin also suggested that sexual selection can favour elaborate receiver structures in order to better detect sexual signals, an idea that has been largely ignored. We evaluated this unorthodox perspective by documenting the antennal lengths of male Uraba lugens Walker (Lepidoptera: Nolidae) moths that were attracted to experimentally manipulated emissions of female sex pheromone. Either one or two females were placed in field traps for the duration of their adult lives in order to create differences in the quantity of pheromone emissions from the traps. The mean antennal length of males attracted to field traps baited with a single female was longer than that of males attracted to traps baited with two females, a pattern consistent with Darwin’s prediction assuming the latter emits higher pheromone concentrations. Furthermore, younger females attracted males with longer antennae, which may reflect age-specific changes in pheromone emission. These field experiments provide the first direct evidence of an unappreciated role for sexual selection in the evolution of sexual dimorphism in moth antennae and raise the intriguing possibility that females select males with longer antennae through strategic emission of pheromones.     

Evidence for contact sex recognition pheromone of the Asian longhorned beetle,<Emphasis Type="Italic"> Anoplophora glabripennis</Emphasis> (Coleoptera: Cerambycidae)

Field observations of the Asian longhorned beetle (Anoplophora glabripennis) mating behavior in China suggested that a female-produced contact pheromone was almost certainly involved in sex recognition. Gas chromatography–mass spectrometry (GC-MS) analysis of A. glabripennis adults' whole body cuticular extracts indicates that a series of long-chain hydrocarbons comprise the cuticular waxes of both sexes. Although for the most part the GC profiles are similar for the two sexes, five monounsaturated compounds were consistently more abundant in samples from females than in those from males. These compounds were identified as (Z)-9-tricosene, (Z)-9-pentacosene, (Z)-7-pentacosene, (Z)-9-heptacosene, and (Z)-7-heptacosene in the approximate ratio of 1:2:2:8:1, respectively. Antennal and palpi contact to a polypropylene micro-centrifuge tube coated with a synthetic mixture of the five compounds stimulated copulatory behavior in males.Electronic Supplementary Material  Supplementary material is available for this article if you access the article at . A link in the frame on the left on that page takes you directly to the supplementary material.     

Female sex pheromone and male behavioral responses of the bombycid moth <Emphasis Type="Italic">Trilocha varians</Emphasis>: comparison with those of the domesticated silkmoth <Emphasis Type="Italic">Bombyx mori</Emphasis>

Analysis of female sex pheromone components and subsequent field trap experiments demonstrated that the bombycid moth Trilocha varians uses a mixture of (E,Z)-10,12-hexadecadienal (bombykal) and (E,Z)-10,12-hexadecadienyl acetate (bombykyl acetate) as a sex pheromone. Both of these components are derivatives of (E,Z)-10,12-hexadecadienol (bombykol), the sex pheromone of the domesticated silkmoth Bombyx mori. This finding prompted us to compare the antennal and behavioral responses of T. varians and B. mori to bombykol, bombykal, and bombykyl acetate in detail. The antennae of T. varians males responded to bombykal and bombykyl acetate but not to bombykol, and males were attracted only when lures contained both bombykal and bombykyl acetate. In contrast, the antennae of B. mori males responded to all the three components. Behavioral analysis showed that B. mori males responded to neither bombykal nor bombykyl acetate. Meanwhile, the wing fluttering response of B. mori males to bombykol was strongly inhibited by bombykal and bombykyl acetate, thereby indicating that bombykal and bombykyl acetate act as behavioral antagonists for B. mori males. T. varians would serve as a reference species for B. mori in future investigations into the molecular mechanisms underlying the evolution of sex pheromone communication systems in bombycid moths.     

When males whistle at females: complex FM acoustic signals in cockroaches

Male cockroaches of the species Elliptorhina chopardi expel air through a pair of modified abdominal spiracles during courtship. This air expulsion simultaneously produces air and substrate-borne vibrations. We described and compared in details these two types of vibrations. Our analysis of the air-borne signals shows that males can produce three categories of signals with distinct temporal and frequency parameters. “Pure whistles” consist of two independent harmonic series fast frequency modulated with independent harmonics that can cross each other. “Noisy whistles” also possess two independent voices but include a noisy broad-band frequency part in the middle. Hiss sounds are more noise-like, being made of a broad-band frequency spectrum. All three call types are unusually high in dominant frequency (>5 kHz) for cockroaches. The substrate-borne signals are categorised similarly. Some harmonics of the substrate-borne signals were filtered out, however, making the acoustic energy centered on fewer frequency bands. Our analysis shows that cockroach signals are complex, with fast frequency modulations and two distinct voices. These results also readdress the question of what system could potentially receive and decode the information contained within such complex sounds.Electronic supplementary material  Supplementary material is available for this article at and is accessible for authorized users.     

Male scent-marking pheromone of <Emphasis Type="Italic">Bombus ardens ardens</Emphasis> (Hymenoptera; Apidae) attracts both conspecific queens and males

To explore the role of the volatiles emitted from male labial gland (LG) of the bumblebee Bombus ardens ardens, we investigated the responses of virgin queens and males to volatiles using a gas chromatography–electroantennographic detector (GC–EAD) system and Y-tube olfactometer. GC–EAD analysis revealed that citronellol, the main compound detected in the male LG, caused clear electrophysiological responses in the antennae of B. a. ardens virgin queens and males although two minor compounds elicited antennal responses when applied in a high concentration. Behavioral tests using a Y-tube olfactometer showed that queens and males were significantly attracted to both LG extracts and citronellol more than to the solvent alone. This is the first study to demonstrate that citronellol as a major compound of male scent-marking pheromone in B. a. ardens functions as a sex attractant for queens. The results also suggest that this compound has another function as a trail marker used by males.     

Chirality determines pheromone activity for flour beetles

Olfactory perception and orientation behaviour of female and male flour beetles (Tribolium castaneum, T. confusum) to single stereoisomers of their aggregation pheromone revealed maximal receptor potentials and optimal attraction in response to 4R,8R-(?)-dimethyldecanal, whereas its optical antipode 4S,8S-(+)-dimethyldecanal was found to be inactive in this respect. Female flour beetles of both species were ≈ 10³ times less attracted to 4R,8S-(+)- and 4S,8R-(?)-dimethyldecanal than to 4R,8R-(?)-dimethyldecanal, while male flour beetles failed to respond to the R,S-(+)- and S,R-(?)-stereoisomers. Pheromone extracts of prothoracic femora from unmated male flour beetles elicited higher receptor potentials in the antennae of females than in those of males. The results suggest that the aggregation pheromone emitted by maleT. castaneum as well as maleT. confusum has the stereochemical structure of 4R,8R-(?)-dimethyl-decanal, which acts as sex attractant for the females and as aggregant for the males of both species.     

Male behaviors reveal multiple pherotypes within vine mealybug Planococcus ficus (Signoret) (Hemiptera; Pseudococcidae) populations

The vine mealybug (VM) females collected in Israel produce two sex pheromone compounds: lavandulyl senecioate (LS) and (S)-lavandulyl isovalerate (LI). The males display ambiguous behavior to LI: repulsion in the vineyard and attraction of laboratory-reared males. We addressed the question of individual male behavior, i.e., do males respond to both LS and LI, or might they display a distinct response to each of the two pheromone compounds. We compared male pherotype frequencies between wild-caught and laboratory-reared populations. Then, we examined the relationship between pherotype composition and male capture rates in pheromone traps. Finally, we addressed the heredity of the pherotypes. The Israeli VM populations contain nine different male pherotypes, as defined according to the male behavior to pheromone compounds. The studied Portuguese populations included five of the nine pherotypes; none of the Portuguese males were attracted to LI. It seems that the high frequency of males that were attracted to LI is related to dense VM populations. It is hypothesized that selection for the male pherotypes, I males, those that respond to LI, occur under high-density rearing conditions. This may result from shorter development times of males and females that produce more I male pherotypes. The lower relative frequency of trapping of males in LI-baited traps than expected from the percentage determined in a Petri dish arena suggests that males that respond solely to LS (S males) are better fliers. The results also suggest that the pherotype trait is inherited by both sexes of the VM.     

Behavioural evidence of male volatile pheromones in the sex-role reversed wolf spiders Allocosa brasiliensis and Allocosa alticeps

The use of chemical signals in a sexual context is widespread in the animal kingdom. Most studies in spiders report the use of female pheromones that attract potential sexual partners. Allocosa brasiliensis and Allocosa alticeps are two burrowing wolf spiders that show sex-role reversal. Females locate male burrows and initiate courtship before males perform any detectable visual or vibratory signal. So, females of these species would be detecting chemical or mechanical cues left by males. Our objective was to explore the potential for male pheromones to play a role in mate detection in A. brasiliensis and A. alticeps. We designed two experiments. In Experiment 1, we tested the occurrence of male contact pheromones by evaluating female courtship when exposed to empty burrows constructed by males or females (control). In Experiment 2, we tested the existence of male volatile pheromones by evaluating female behaviour when exposed to artificial burrows connected to tubes containing males, females or empty tubes (control). Our results suggest the occurrence of male volatile pheromones that trigger female courtship in both Allocosa species. The sex-role reversal postulated for these wolf spiders could be driving the consequent reversal in typical pheromone-emitter and detector roles expected for spiders.     

Surface lipids of queen-laid eggs do not regulate queen production in a fission-performing ant

In animal societies, most collective and individual decision making depends on the presence of reproductive individuals. The efficient transmission of information among reproductive and non-reproductive individuals is therefore a determinant of colony organization. In social insects, the presence of a queen modulates multiple colonial activities. In many species, it negatively affects worker reproduction and the development of diploid larvae into future queens. The queen mostly signals her presence through pheromone emission, but the means by which these chemicals are distributed in the colony are still unclear. In several ant species, queen-laid eggs are the vehicle of the queen signal. The aim of this study was to investigate whether queen-laid eggs of the ant Aphaenogaster senilis possess queen-specific cuticular hydrocarbons and/or Dufour or poison gland compounds, and whether the presence of eggs inhibited larval development into queens. Our results show that the queen- and worker-laid eggs shared cuticular and Dufour hydrocarbons with the adults; however, their poison gland compounds were not similar. Queen-laid eggs had more dimethylalkanes and possessed a queen-specific mixture of cuticular hydrocarbons composed of 3,11?+?3,9?+?3,7-dimethylnonacosane, in higher proportions than did worker-laid eggs. Even though the queen-laid eggs were biochemically similar to the queen, their addition to experimentally queenless groups did not prevent the development of new queens. More studies are needed on the means by which queen ant pheromones are transmitted in the colony, and how these mechanisms correlates with life history traits.     

Male-produced anti-sex pheromone in a plant bug

In plant bugs (Miridae), females produce sex pheromones in the metathoracic scent gland, which in most other true bugs (Heteroptera) is responsible for chemical defense. The possibility that the metathoracic gland secretion of male plant bugs plays a role other than defense has been largely overlooked. Here we show that in a pine-inhabiting mirid, Phytocoris difficilis Knight, hexyl butyrate and (E)-2-hexenyl butyrate are abundantly produced only in males; we demonstrate that these metathoracic gland compounds elicit strong antennal responses in conspecific males, and that these butyrates totally interrupt attraction of males to the female-produced sex pheromone. Our results suggest that in at least some plant bugs the male metathoracic scent gland esters have a natural communicative function as anti-sex pheromones, probably to interrupt further mating attempts by other males.     

Does the mechanism of sex determination constrain the potential for sex manipulation? A test in geckos with contrasting sex-determining systems

The concentration of yolk steroids was suggested to influence offspring gender in oviparous animals subject to both temperature-dependent sex determination (TSD) and genotypic sex determination (GSD). However, the proposed mechanisms of steroid effects are thought to differ between TSD and GSD: a direct effect of oestrogens on gonad feminisation in TSD species vs a differential induction of male-producing or female-producing gametes in GSD species. Geckos offer an ideal opportunity for testing these suggested mechanisms. Closely related gecko species differ in their modes of sex determination. They lay clutches of two synchronously formed eggs; both eggs share equal steroid levels. If identical hormonal composition and environment during vitellogenesis, gravidity and incubation determine the sex of the progeny, siblings should share the same gender in both TSD and GSD geckos. We found strong support for this prediction in a TSD gecko species. Among clutches that were incubated at the temperature that produced both sexes, there were no clutches with siblings of the opposite sex. On the other hand, about half of the clutches yielded siblings of the opposite sex in four GSD species. These results suggest that sex-determining systems constrain the ability of the female to produce single-sex siblings and, hence, it seems that the GSD mechanism constrains the opportunities for sex ratio manipulation in geckos via yolk steroid manipulation.     

Habitat influences on diversity of bacteria found on German cockroach in Beijing

Cockroaches are worldwide indoor pests carrying microorganisms of medical importance.German cockroaches (Blattella germanica) were sampled in five habitats (hospital, restaurant, office home, and market) in Beijing, and the bacteria were isolated from their external surface and alimentary tract and identified using a Biolog identification system.Cockroach densities significantly differed among habitats (market > home > office > restaurant > hospital).However, no significant differences in bacterial abundance carried by individual German cockroaches (of either sex) were found among habitats.The bacterial abundance in the gut was significantly higher than that on the surface.There were no significant differences in bacterial species richness observed among habitats, sex, carrying position or their interaction.Cluster analysis showed that cockroach densities and bacterial abundance found in the market differed significantly from the other four habitats.The bacterial diversity was not significantly reduced in sensitive facilities such as hospital and restaurant, even though pesticide and bactericide were more frequently applied there.The implications of these findings were discussed in this article.     

Methyl 2-(methylthio)benzoate: the unique sulfur-containing sex pheromone of<Emphasis Type="Italic"> Phyllophaga crinita</Emphasis>

The female-produced sex pheromone of Phyllophaga crinita (Burmeister) (Coleoptera: Scarabaeidae: Melolonthinae; the adult has no common name) is identified as methyl 2-(methylthio)benzoate. This is the first identification of a sulfur-containing, long-distance, female-produced sex attractant from any insect taxa. The root-feeding larvae of this species are serious pests in many crops in Texas and Mexico. In field tests, many P. crinita males were captured in traps baited with the authentic compound. Interestingly, a heteroatom analog, methyl 2-methoxybenzoate, also captured P. crinita males, but only at a dose 10,000 times higher than the lowest tested dose of the authentic pheromone.In memory of M.G. Villani, who died on 15 May 2001. His enthusiasm for soil-dwelling scarab beetles and contributions to their research are greatly missed.     

Attraction of two lacewing species to volatiles produced by host plants and aphid prey

It is well documented that host-related odors enable many species of parasitoids and predatory insects to locate their prey and prey habitats. This study reports the first characterization of prey and prey host odor reception in two species of lacewings, Chrysoperla carnea (Say) and Chrysopa oculata L. 2-Phenylethanol, one of the volatiles emitted from their preys host plants (alfalfa and corn) evoked a significant EAG response from antennae of C. carnea. Traps baited with this compound attracted high numbers of adult C. carnea, which were predominantly females. One of the sex pheromone components (1R,4aS,7S,7aR)-nepetalactol of an aphid species, Acyrthosiphon pisum (Harris) attracted only C. oculata adults. Single sensillum recordings showed that the olfactory neurons of C. carnea responded to both 2-phenylethanol and aphid sex pheromone components, but those of C. oculata only responded to the latter.     

A new class of mealybug pheromones: a hemiterpene ester in the sex pheromone of Crisicoccus matsumotoi

Mealybugs, which include several agricultural pests, are small sap feeders covered with a powdery wax. They exhibit clear sexual dimorphism; males are winged but fragile and short lived, whereas females are windless and less mobile. Thus, sex pheromones emitted by females facilitate copulation and reproduction by serving as a key navigation tool for males. Although the structures of the hitherto known mealybug pheromones vary among species, they have a common structural motif; they are carboxylic esters of monoterpene alcohols with irregular non-head-to-tail linkages. However, in the present study, we isolated from the Matsumoto mealybug, Crisicoccus matsumotoi (Siraiwa), a pheromone with a completely different structure. Using gas chromatography–mass spectrometry and nuclear magnetic resonance spectroscopy, we identified the pheromone as 3-methyl-3-butenyl 5-methylhexanoate. Its attractiveness to males was confirmed in a series of field trapping experiments involving comparison between the isolated natural product and a synthetic sample. This is the first report of a hemiterpene mealybug pheromone. In addition, the acid moiety (5-methylhexanoate) appears to be rare in insect pheromones.     

Ultrasonic courtship song in the Asian corn borer moth, Ostrinia furnacalis

Although sex pheromone communication in the genus Ostrinia (Lepidoptera: Crambidae) has been studied intensively, acoustic communication in this genus has not been explored. In this study, we report that male-produced ultrasound serves as a courtship song in the Asian corn borer moth, O. furnacalis. Upon landing close to a pheromone-releasing female, a male showed a series of courtship behaviors involving emission of ultrasound. The sounds were produced when the wings were vibrated quickly in an upright position. The male song was composed of chirps, i.e., groups of pulses (duration of a chirp = 58.9 ms, 8.8 pulses/chirp), with a broadband frequency of 25–100 kHz. In flight tunnel experiments, deaf and hearing females showed a significant difference in the incidence of three behavioral responses to courting males, i.e., immediate acceptance, acceptance after walking, and rejection. Deaf females showed more ‘rejection’ and less ‘acceptance after walking’ than hearing females, indicating that the detection of male-produced ultrasound plays an important role in the acceptance of a male. The findings are discussed in the context of exploitation of receiver bias and mate choice.