Correspondence between rates of host plant consumption and responses to the Acalymma vittatum male-produced aggregation pheromone

Abstract In field studies, wind-directed traps were used to investigate the resource-based properties of the male-produced aggregation pheromone of Acalymma vittatum (F.) (Coleoptera: Chrysomelidae). Responses to early season overwintered field-collected male A. vittatum without food were compared with those feeding on pollen, and in turn these were compared with those feeding on cucurbit seedlings. These comparisons were modelled after transitions as they would occur at the initiation of A. vittatum host plant colonization.
In another experiment, colony-reared male A. vittatum that had been fed only near isogenic lines of potted cucumber ( Cucumis sativus ) seedlings that either lacked or contained cucurbitacin were placed as bait in the traps. Using the inverse relationship, which is understood for A. vittatum , between previous exposure and the phagostimulant response to cucurbitacin, rates of foliage consumption could be manipulated. The interaction between relative rates of host plant consumption of the males used as lures and responses to the pheromone could thus be tested. The use of A. vittatum reared entirely without dietary cucurbitacin as bait also enabled evaluation of the potential role of cucurbitacin in pheromone biosynthesis. Trap responses were closely correlated with the rates of feeding of the beetles in the trap lures, but cucurbitacin was neither a precursor nor a prerequisite for the pheromone. Implications for early season field colonization and the adaptive significance of this pheromone are discussed.     

Effects of age, sex, and dietary history on response to cucurbitacin in Acalymma vittatum

The chrysomelid Acalymma vittatum is stenophagous, subsisting almost entirely on plants in the Cucurbitaceae, which generally contain cucurbitacins. Cucurbitacins are extremely bitter tetracyclic triterpenoids that are toxic to most organisms. As do other diabroticite beetles, A. vittatum sequester cucurbitacins, which have been shown to act as phagostimulants and arrestants. Our results reveal, however, that for A. vittatum the response to cucurbitacin diminishes with continued sequestration. Colony-reared A. vittatum were fed only roots (as larvae) and foliage of either `Marketmore 76' (which contains a normal amount of cucurbitacin, `bitter') or `Marketmore 80' (a near isogenic line that contains no cucurbitacin, `non-bitter') cucumber. Over 1200 individual beetles from the day of adult emergence to 15 days following emergence were placed in choice and no-choice arenas containing potted cotyledons of the two cucumber varieties for 24 h. In choice tests, overall preference for the bitter cucumber cultivar was maintained, but degree of preference changed with age and became significantly less for beetles reared on bitter diets. Furthermore, in no-choice tests, age, sex, dietary history, and interactions among these variables all significantly affected the feeding response to cucurbitacin. For A. vittatum reared without cucurbitacin, total consumption of the bitter cultivar increased over time. For beetles reared with cucurbitacin, total foliage consumption of the bitter cultivar declined, within nine days, to equal that of the non-bitter cultivar. Feral A. vittatum, unexpectedly, consumed more of the non-bitter than the bitter cultivar in no-choice tests. Ecological and applied implications of this variation in response to cucurbitacin are discussed.     

Attraction of pepper weevil to volatiles from damaged pepper plants

Pioneer herbivorous insects may find their host plants through a combination of visual and constitutive host‐plant volatile cues, but once a site has been colonized, feeding damage changes the quantity and quality of plant volatiles released, potentially altering the behavior of conspecifics who detect them. Previous work on the pepper weevil, Anthonomus eugenii Cano (Coleoptera: Curculionidae), demonstrated that this insect can detect and orient to constitutive host plant volatiles released from pepper [Capsicum annuum L. (Solanaceae)]. Here we investigated the response of the weevil to whole plants and headspace collections of plants damaged by conspecifics. Mated weevils preferred damaged flowering as well as damaged fruiting plants over undamaged plants in a Y‐tube olfactometer. They also preferred volatiles from flowering and fruiting plants with actively feeding weevils over plants with old feeding damage. Both sexes preferred volatiles from fruiting plants with actively feeding weevils over flowering plants with actively feeding weevils. Females preferred plants with 48 h of prior feeding damage over plants subjected to weevil feeding for only 1 h, whereas males showed no preference. When attraction to male‐ and female‐inflicted feeding damage was compared in the Y‐tube, males and females showed no significant preference. Wind tunnel plant assays and four‐choice olfactometer assays using headspace volatiles confirmed the attraction of weevils to active feeding damage on fruiting plants. In a final four‐choice olfactometer assay using headspace collections, we tested the attraction of mated males and virgin and mated females to male and female feeding damage. In these headspace volatile assays, mated females again showed no preference for male feeding; however, virgin females and males preferred the headspace volatiles of plants fed on by males, which contained the male aggregation pheromone in addition to plant volatiles. The potential for using plant volatile lures to improve pepper weevil monitoring and management is discussed.     

Mate finding in the forest cockchafer, Melolontha hippocastani, mediated by volatiles from plants and females

The response of the forest cockchafer, Melolontha hippocastani F. (Coleoptera, Scarabaeidae), towards volatiles emitted by different host plants and conspecifics was tested in field experiments during the flight period at dusk. Funnel traps containing artificially damaged leaves from the host plants Carpinus betulus L. and Quercus rubra L., as well as from the non‐host plant Prunus serotina Ehrh. caught significantly more beetles than empty control traps. On the other hand, traps baited with undamaged leaves from Q. rubra did not catch significantly more beetles than empty controls. Leaves from C. betulus damaged by beetle feeding did not attract more beetles than artificially damaged leaves. By use of gas chromatography coupled with electroantennographic detection (GC‐EAD) electrophysiological responses of males and females were shown for 18 typical plant volatiles. A synthetic mixture of selected typical green plant volatiles was also highly attractive in the field. A total of 9982 beetles was caught during the field experiments, among them only 33 females. This suggests that attraction to damaged foliage during flight period at dusk is male‐specific. Field experiments testing the attractiveness of female M. hippocastani towards conspecific males by employing caged beetles and beetle extracts indicated that males of M. hippocastani use a female‐derived sex pheromone for mate location. On wired cages containing either unmated feeding females, or unmated females without access to foliage, or feeding males in combination with extracts from unmated females, significantly more males landed during the flight period than on comparable control cages containing feeding males or male extracts. A possible scenario of mate location in M. hippocastani involving feeding‐induced plant volatiles and a female‐derived sex pheromone is discussed.     

Functional response of Nesidiocoris tenuis (Hemiptera: Miridae) to Trialeurodes vaporariorum (Hemiptera: Aleyrodidae): effect of different host plants

The influence of three host plants, namely cucumber, tomato and eggplant, on functional response of male, virgin and mated female predatory bug Nesidiocoris tenuis was investigated on different densities of Trialeurodes vaporariorum nymphs. The 24-h experiment conducted at laboratory conditions revealed that N. tenuis exhibited a type II functional response to T. vaporariorum on host plants. There were no significant differences between attack rates, as well as handling times estimated for each adult stage of the predator between host plants. However, on each host plant, the handling time estimated for the mated female in comparison with two other adult stages had lower values (0.7952, 0.6827 and 0.8884?h^?1 on cucumber, tomato and eggplant, respectively). Handling time estimated for the mated female on cucumber was significantly lower than that estimated for the male predator. The highest maximum handling rate (T/T_h) was estimated for the mated female followed by the virgin female and male on all host plants. For three adult stages of the predator, the highest value of this parameter was determined on tomato followed by cucumber and eggplant. Unlike virgin and mated females, the host plant significantly affected prey consumption by the male. Prey consumed by mated females was higher than those obtained for two other adult stages of the predator on each host plant. The difference in trichome density between three host plants may be responsible for the obtained results. These results revealed that N. tenuis is more effective in the biological control of T. vaporariorum on tomato in comparison with cucumber and eggplant.     

Sexual Differences in the Behavior of the Harvestman Leiobunum vittatum (Opiliones, Sclerosomatidae) Towards Conspecific Cues

Preliminary observations of the harvestman Leiobunum vittatum found that individuals rub their bodies against the substrate, presenting the possibility of chemical marking. To determine whether or not L. vittatum individuals can detect substrate-borne chemical cues, we compared responses of L. vittatum males and females to substrate-borne male and female cues. We found that individuals of L. vittatum do respond to conspecific cues and that their responses are sex-specific. In response to substrate-borne conspecific cues, male L. vittatum spent more time, engaged in more scraping with their sensory legs I, and engaged in pedipalpal tapping more often in the presence versus absence of conspecific cues (male and female equally). Furthermore, in the presence of conspecific cues, males engaged in two behaviors never observed in females—(a) “fast approach” and (b) “jerking”, the latter of which was never observed in the presence of cricket cues. In contrast to males, females did not spend more time on conspecific cues, but did spend more time tapping their pedipalps in the presence of male vs female cues, suggesting an ability to distinguish between them. A final experiment explored the possibility that females could discriminate among males of varying histories of agonistic interactions based upon their chemical cues. We found no support for this hypothesis. Our results demonstrate that L. vitattum do respond to conspecific cues, and introduce the possibility that intraspecific communication may be mediated in part by chemical cues.     

Effects of host plant on life‐history traits in the polyphagous spider mite Tetranychus urticae

Studying antagonistic coevolution between host plants and herbivores is particularly relevant for polyphagous species that can experience a great diversity of host plants with a large range of defenses. Here, we performed experimental evolution with the polyphagous spider mite Tetranychus urticae to detect how mites can exploit host plants. We thus compared on a same host the performance of replicated populations from an ancestral one reared for hundreds of generations on cucumber plants that were shifted to either tomato or cucumber plants. We controlled for maternal effects by rearing females from all replicated populations on either tomato or cucumber leaves, crossing this factor with the host plant in a factorial design. About 24 generations after the host shift and for all individual mites, we measured the following fitness components on tomato leaf fragments: survival at all stages, acceptance of the host plant by juvenile and adult mites, longevity, and female fecundity. The host plant on which mite populations had evolved did not affect the performance of the mites, but only affected their sex ratio. Females that lived on tomato plants for circa 24 generations produced a higher proportion of daughters than did females that lived on cucumber plants. In contrast, maternal effects influenced juvenile survival, acceptance of the host plant by adult mites and female fecundity. Independently of the host plant species on which their population had evolved, females reared on the tomato maternal environment produced offspring that survived better on tomato as juveniles, but accepted less this host plant as adults and had a lower fecundity than did females reared on the cucumber maternal environment. We also found that temporal blocks affected mite dispersal and both female longevity and fecundity. Taken together, our results show that the host plant species can affect critical parameters of population dynamics, and most importantly that maternal and environmental conditions can facilitate colonization and exploitation of a novel host in the polyphagous T. urticae, by affecting dispersal behavior (host acceptance) and female fecundity.     

Response of the pollen beetle Meligethes aeneus to volatiles emitted by intact plants and conspecifics

The response of the pollen beetle Meligethes aeneus Fab. (Coleoptera, Nitidulidae) to the volatiles of undamaged plants and conspecifics was tested in a Y-tube-olfactometer-bioassay. Beetles that had hibernated preferred significantly the volatiles emitted by their most important host plant, oilseed-rape (Brassica napus L. ‘Lorar’) in the early bud-stage. However, the odour emitted by rye (Secale cereale L.), tomato plants (Solanum lycopersicum L.), and yarrow (Achillea millefolium L.) were also attractive when tested against the corresponding growing-medium. Dock plants (Rumex obtusifolius L.) and touch-me-not (Impatiens parviflora L.) possessed no attractive properties. When tested against each other, oilseed-rape was preferred significantly by M. aeneus above all other plants, with the exception of tomato. The results indicate that M. aeneus is able to locate its host plant by olfactory stimuli in the early bud-stage, i.e. in that stage in which the infestation begins in the field and when the typical yellow colour and floral scent of oilseed-rape are absent. Female pollen beetles avoided significantly the volatiles emitted by female conspecifics and an ether extract of conspecifics of mixed sex, while volatiles from males had no significant effect on females. Furthermore, males showed no preferences when responding to conspecific odour in the olfactometer. These results suggest the existence of an epideictic pheromone for M. aeneus.     

INFLUENCE DU SUPERPARASITISME SUR LA DURÉE DU DÉVELOPPEMENT LARVAIRE ET LE POIDS DU PARASITOÏDE LIXOPHAGA DIATRAEAEÉLEVÉ DANS UN HÔTE DE SUBSTITUTION GALLERIA MELLONELLA

Résumé G. mellonella infestée au dernier stade larvaire avec 1, 2, 3 ou 5 planidia/hôte (ph/H) produit 1 à 5 pupes/hôte (pu/H). La mortalité des chenilles augmente avec le nombre de pl/H. Le poids des pupes et décroît avec un nombre croissant de pu/H (18,2 à 12,9 mg pour les et 12,5 à 9,7 mg pour les ). Le développement larvaire dure 8,7 j. chez les et 8,3 chez les ; il est peu affecté par le superparasitisme. Avec 1, 2, 3 et 5 pl/H nous obtenons 0,84–1,61–2,17 et 3,43 pu/H et 0,81–1,48–2,10 et 3,11 imagos/H. L'optimum est de 3 pl/H ou 1 à 2=pl/H pour obtenir des parasitoïdes plus lourds.

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Summary The influence of superparasitism on larval and pupal development is investigated. Last-larval instar G. mellonella (200±5 mg) were infected with 1, 2, 3 or 5 planidia/host (pl/H) producing 1 to 5 pupae per host (pu/H). Host mortality (8.6–8.3–14.3 and 22.2%) increased with the number of planidia. The planidia may transmit a bacteriosis. Pupal weight ( and ) decreased as number of pu/H increased. pupae were heavier than ones: 12.9 to 18.2 mg for against 9.7 to 12.5 mg for . Larval development lasted slightly longer for (8.7 d.) than for (8.3 d.), and its duration was little affected by superparasitism. 1, 2, 3 or 5 pl/H yielded 0.84–1.61–2.17 and 3.43 pu/H and 0.81–1.48–2.10 and 3.11 adults/H. An optimum was obtained with 3 planidia of L. diatraeae on G. mellonella or 1 to 2 to obtain heavier parasitoids.

     

PHOTOSYNTHESIS RESPONSE OF ‘CAROLINA’ CUCUMBER TO SIMULATED AND ACTUAL STRIPED CUCUMBER BEETLE (COLEOPTERA: CHRYSOMELI-DAE) DEFOLIATION*

Abstract Field studies were conducted in 1995 to compare the photosynthesis response between simulated striped cucumber beetle, Acalymma vittatum (F.), defoliation and actual A. vittatum defoliation in ‘Carolina’ cucumber. Six simulated defoliation levels (0%–100%)were used over 5 timings of defoliation (first true leaf to harvest). Plots were defoliated with scissors twice each week throughout each timing interval, and defined as continuous defoliation. In addition to continuous defoliation, one-time simulated defoliation treatments were imposed using the same timing treatments but only 4 defoliation levels (0, 25, 50, and 100%). Two cage studies, with A. vittatum adults, were used to provide estimates of actual insect defoliation injury. Beetles were placed in cages for 2 weeks at densities of 0, 1, 3, 5, and 10/plant and net photosynthetic rates were measured. No significant differences (P <0. ⁰⁵ were detected in pho-tosynthetic rates between simulated and actual insect defoliation treatments. Also, few significant differences in photosynthetic rates were detected between damaged and undamaged leaves in the simulated insect defoliation study. Results indicated that the simulated insect defoliation procedure appeared to accurately reflect the growth response of ‘Carolina’ cucumber subjected to actual A. vittatum defoliation, and that simulated defoliation results should be appropriate for developing economic injury levels for A. vittatum on cucumber.     

RELEVANCE OF PLANT VOLATILES TO SEX PHERO‐MONE IN LURING APHIDS IN THE FIELD1

Abstract Outdoor trials were made to test the responses of male aphids to pheromone lures in fields with different host‐plants, in order to know whether host‐plants would affect catches of males to pheromone lures. In peach orchard, males of three aphid species, i.e. Myzus persicae, Hyalopterus amygdali and Rhopalosiphum nymphaeae, were found in traps baited with aphid sex pheromone nepetalactone and nepetalactol. The most numerous numbers was M, persicae and it was also attracted by plant‐derived nepetalactone alone in peach orchard and cabbage garden. Schizaphis graminum males were caught in water traps in wheat field. In apple orchard, males of two aphid species, Aphis cifricola and Sappaphis sinipiricola were caught in significantly larger numbers in the pheromone traps than in the control traps. Field study indicated that the sex pheromone of H. amygduli comprises either one or both of nepetalactone and nepetalactol, and nepetalactone may be one component of sex pheromones of A cifricola and S sinipiricola Another field trial had shown that catches were increased if volatiles from a steam‐distilled extract of peach leaves were released simultaneously with nepetalactone. Elec‐troantennograms (EAGs) were recorded from males of M. persicae to volatiles of peach leaves and buds.     

Sex‐specific responses of Asian citrus psyllid to volatiles of conspecific and host‐plant origin

Diaphorina citri Kuwayama (Hemiptera: Psyllidae) is the primary vector of Candidatus Liberibacter spp. bacteria that cause citrus greening, a disease of worldwide importance. Olfactometry was employed to test responses of D. citri to odours from intact citrus plants (Mexican lime, Citrus aurantifolia, sour orange, Citrus aurantium, Marsh grapefruit, Citrus paradisi and Valencia orange, Citrus sinensis), citrus plants previously infested with D. citri, and odours of conspecifics including nymphs, adult insects of same and opposite sex, and their products (honeydew), both alone and in combination. In contrast to other studies, psyllids of both sexes were attracted to volatiles of undamaged Mexican lime leaves, whereas undamaged grapefruit attracted only females, and leaves of Valencia and sour orange did not attract either sex. All four plant species attracted female psyllids when previously infested, but only Mexican lime and sour orange‐attracted males. Thus, Citrus species appear to vary in the production of both constituitive and induced volatiles that attract adult psyllids. Volatiles emitted by nymphs did not attract either sex, but psyllid honeydew was attractive to males, likely due to female pheromone residues. Males oriented to the odour of females, whereas the reverse was not true, and neither males nor females oriented to same‐sex volatiles. The addition of conspecific cues (adults, nymphs or honeydew) did not increase female attraction to previously infested leaves, but male response was increased by the presence of adults and honeydew, regardless of plant species. Thus, female psyllids appear to orient more strongly to volatiles of plant origin, whereas males respond more strongly to cues emanating from females and conspecific excretions. These results suggest that female psyllids drive the initial colonization of host plants, whereas males orient to females and infested plants. Identification of the specific volatiles involved may permit their use in monitoring and management of this pest.     

Behavioral evidence for a female-produced sex attractant in Diaphorina citri

Diaphorina citri Kuwayama (Hemiptera: Psyllidae) is an important worldwide pest of citrus. It vectors three phloem-restricted bacteria in the genus Candidatus Liberibacter that cause huanglongbing (citrus greening disease). Studies were conducted to examine the behavioral responses of male and female D. citri to conspecifics of the same and opposite sex, with and without associated citrus host plants, in both open-air arena choice assays and Y-tube olfactometer assays. Virgin and mated male D. citri colonized citrus plants that were currently or had been previously colonized by virgin or mated female D. citri in greater numbers than control plants without females. However, males or females did not accumulate more on plants colonized by conspecifics of the same sex compared to uninfested plants, and females showed no preference for plants pre-infested with males compared with uninfested controls. In complementary Y-tube olfactometer assays, virgin and mated males chose arms with odor sources from mated females compared with blank controls in the absence of associated citrus host plant volatiles. In both behavioral assays, mated female D. citri appeared more attractive than virgin females. The vibrational calling behavior of male D. citri was reduced when males were challenged by the odors of conspecific mated females relative to when males were challenged by the odor of other males. Collectively, our results provide behavioral evidence for a female-produced volatile sex attractant pheromone in D. citri . Future identification and synthesis of a sex attractant pheromone will be an important contribution to current monitoring and management practices for D. citri .     

A Glandular Gift in the German Cockroach, Blattella germanica (L.) (Dictyoptera: Blattellidae): The Courtship Feeding of a Female on Secretions from Male Tergal Glands

In the courtship behavior of the German cockroach, the male presents tergal glands to the female and feeds her with glandular secretions to place her in the appropriate precopulatory position. The phagostimulant activity of the secretions was quantitatively examined using the polyethylene glycol film method. The methanol extract of the glands on the eighth tergite induced a potent feeding response in 6-day-old virgin females (EC ₅₀ = 0.0037 male equivalent/40 g PEG spot). However, there was no temporal relation between the feeding response and the sexual receptivity of the females. Moreover, besides virgin females, the extract induced a feeding response in gravid or mated females, males, and the last-instar nymphs. These results strongly suggest that the secretions function as a dietary feeding stimulant in principle but as a courtship pheromone in the context of courtship behavior where the stimulants are offered as a nuptial gift.     

Influence of host plant,geography and pheromone strain on genomic differentiation in sympatric populations of Ostrinia nubilalis

Patterns of mating for the European corn borer (Ostrinia nubilalis) moth depend in part on variation in sex‐pheromone blend. The ratio of (E)‐11‐ and (Z)‐11‐tetradecenyl acetate (E11‐ and Z11‐14:OAc) in the pheromone blend that females produce and males respond to differs between strains of O. nubilalis. Populations also vary in female oviposition preference for and larval performance on maize (C4) and nonmaize (C3) host plants. The relative contributions of sexual and ecological trait variation to the genetic structure of O. nubilalis remains unknown. Host‐plant use (¹³C/¹⁴C ratios) and genetic differentiation were estimated among sympatric E and Z pheromone strain O. nubilalis males collected in sex‐pheromone baited traps at 12 locations in Pennsylvania and New York between 2007 and 2010. Among genotypes at 65 single nucleotide polymorphism marker loci, variance at a position in the pheromone gland fatty acyl‐reductase (pgfar) gene at the locus responsible for determining female pheromone ratio (Pher) explained 64% of the total genetic differentiation between males attracted to different pheromones (male response, Resp), providing evidence of sexual inter‐selection at these unlinked loci. Principal coordinate, Bayesian clustering, and distance‐based redundancy analysis (dbRDA) demonstrate that host plant history or geography does not significantly contribute to population variation or differentiation among males. In contrast, these analyses indicate that pheromone response and pgfar‐defined strain contribute significantly to population genetic differentiation. This study suggests that behavioural divergence probably plays a larger role in driving genetic variation compared to host plant‐defined ecological adaptation.     

Consumptive and non‐consumptive effects of wolf spiders on cucumber beetles and cucumber plant damage

Biological control research often focuses on the ability of predators to reduce pest densities and protect crops through consumption. Less studied is their ability to protect crops by altering pest behaviour (non‐consumptive effects). Lab experiments were conducted to test predation rates of striped cucumber beetles (Acalymma vittatum; Coleoptera: Chrysomelidae) and spotted cucumber beetles (Diabrotica undecimpunctata howardi; Coleoptera: Chrysomelidae) by large (>10 mm) wolf spiders (Araneae: Lycosidae). Field experiments were conducted to examine how the physical presence and/or cues of spiders impact the behaviour and mortality of A. vittatum (specialist) and D. undecimpunctata (generalist) cucumber beetles as well as growth and damage of cucumber plants (Cucumis sativus; Cucurbitaceae). A. vittatum and D. undecimpunctata adults were added to caged cucumber plants without a spider, with spider cues only (spider removed before beetle inclusion), with spider only (spider introduced to plants immediately before beetle inclusion), and with spiders and their cues present (spiders introduced 24 hr in advance of beetle inclusion). A. vittatum responded to spider cues primarily by emigrating from plants. Contrarily, D. undecimpunctata did not display obvious responses, such as reduced feeding or increased emigration, to spider foraging and/or cues. Actively foraging lycosids increased A. vittatum mortality and reduced densities of D. undecimpunctata in the field when cucumber plants were flowering. This study highlights how non‐consumptive and consumptive effects can play a role in modifying pest populations, and how these effects can vary across species and plant growth stages.     

Behavioural response of the invasive Halyomorpha halys (Hemiptera: Pentatomidae) to host plant stimuli augmented with semiochemicals in the field

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Male moth songs tempt females to accept mating: the role of acoustic and pheromonal communication in the reproductive behaviour of Aphomia sociella

Background

Members of the subfamily Galleriinae have adapted to different selective environmental pressures by devising a unique mating process. Galleriinae males initiate mating by attracting females with either chemical or acoustic signals (or a combination of both modalities). Six compounds considered candidates for the sex pheromone have recently been identified in the wing gland extracts of Aphomia sociella males. Prior to the present study, acoustic communication had not been investigated. Signals mediating female attraction were likewise unknown.

Methodology/Principal Findings

Observations of A. sociella mating behaviour and recordings of male acoustic signals confirmed that males initiate the mating process. During calling behaviour (stationary wing fanning and pheromone release), males disperse pheromone from their wing glands. When a female approaches, males cease calling and begin to produce ultrasonic songs as part of the courtship behaviour. Replaying of recorded courting songs to virgin females and a comparison of the mating efficiency of intact males with males lacking tegullae proved that male ultrasonic signals stimulate females to accept mating. Greenhouse experiments with isolated pheromone glands confirmed that the male sex pheromone mediates long-range female attraction.

Conclusion/Significance

Female attraction in A. sociella is chemically mediated, but ultrasonic communication is also employed during courtship. Male ultrasonic songs stimulate female sexual display and significantly affect mating efficiency. Considerable inter-individual differences in song structure exist. These could play a role in female mate selection provided that the female''s ear is able to discern them. The A. sociella mating strategy described above is unique within the subfamily Galleriinae.