Searching behaviour and mate recognition by males of the two-spot ladybird beetle, Adalia bipunctata

Abstract. 1. Adult males of the two-spot ladybird beetle, Adalia bipunctata , did not show a functional response to increase in aphid abundance and consumed markedly fewer aphids than do the females.
2. At high densities of prey, females spent more time in area-restricted search than when prey was scarce. Males were always less active than females and they did not respond to an increase in prey abundance by a change in searching behaviour.
3. After a brief encounter with a female, a male showed area-restricted searching behaviour. This behaviour occurred in response to encountering a female's elytra and in particular to a chloroform-soluble component (sex pheromone) present on or in the elytra.
4. Males needed to encounter a female in order to respond to her presence, which indicated the pheromone is a contact pheromone.
5. The searching behaviour of males appeared to be mainly directed towards locating females; that of females towards locating aphids. This difference between the sexes should be taken into account when quantifying the predatory response of ladybirds to aphid abundance in the field.     

Production and perception of pheromones by the beetle Tribolium confusum

Adults of Tribolium confusum secrete two pheromones. The first, produced by the male, is attractive to both sexes and the second, produced by the female, is attractive to the male only. Pheromone production and perception was studied in relation to habituation, beetle age, time of day and previous mating. A living source of each pheromone habituates the responding beetles, the male pheromone habituating more strongly; female pheromone habituates only in the absence of the male pheromone. Habituation to one pheromone was always accompanied by an enhanced response to the other.Five days after emergence, production of male pheromone reaches a peak that is maintained. Production of female pheromone peaks after 3 days. Both sexes are responsive to male pheromone immediately upon eclosion, males reaching maximum response at 14 days, females at 8 days. Males are also responsive to female pheromone upon eclosion reaching maximum response at 8 days; female response to female pheromone is imperceptible. Males but not females display a 24 hr rhythm in pheromone production. Mated beetles did not differ significantly from unmated beetles in their ability to perceive pheromones. Alteration in male pheromone production after mating was detected by females but not males; this pheromone may, therefore, act as both a sex and aggregation pheromone.     

Vibratory courtship in a web-building spider: signalling quality or stimulating the female?

Courtship behaviour in spiders in the form of premating vibrations by males may function (1) as a male identity signal used for species recognition, (2) in suppression of female aggressiveness, (3) to stimulate female mating behaviour, or (4) as a quality signal used in female choice. We investigated the function of web vibration by male Stegodyphus lineatus in a series of experiments. Regardless of vibratory performance, all males mated successfully with virgin females but only 56.4% of males mated with nonvirgin females. Vibratory performance did not influence male mating success, but heavier males had a higher probability of mating with mated females. Males vibrated less often and produced fewer vibrations when introduced on the web of a mated female. Males that vibrated webs of virgin females mated faster than nonvibrating males, but there was no effect of vibration rate or body mass. There was no effect of male vibratory effort or vibration rate on female reproductive success measured as time to egg laying, clutch size, number of hatched young, number of dispersed young and offspring body mass after a single mating. Males vibrated on abandoned virgin female webs but the response decreased with increasing duration of female absence, suggesting that females produce a web-borne pheromone, which elicits male vibrating behaviour. Mated females were less receptive and not stimulated by male vibrating behaviour. We conclude that male premating vibrations in S. lineatus do not function as a male quality signal selected via female choice. Rather, the primary function of this behaviour may be to stimulate a receptive female to mate. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

A female‐produced short‐range sex pheromone in the egg parasitoid Trissolcus brochymenae

The potential for short‐range sex pheromone communication by the egg parasitoid wasp Trissolcus brochymenae (Hymenoptera: Platygastridae) was investigated in closed arena bioassays. Males of this parasitoid showed more antennal drumming and more frequent mounting behaviour on 1‐ to 2‐d‐old virgin females compared with 8‐d‐old virgin females. Male copulation attempts were fewer with previously mated females than with virgin females. Males courted and made copulation attempts with 1‐ to 2‐d‐old female cadavers, but not with male cadavers or with female cadavers rinsed in organic solvents of different polarities. Male attraction to female cadavers was re‐established by treating cadavers with acetone extracts of females, but not with ether or hexane extracts. In experiments using female cadavers dissected into head, mesosoma, and gaster, and then reassembled using one unwashed body section and two body sections washed in acetone, males were attracted only to the reassembled cadavers with an unwashed mesosoma. These findings suggest that (1) courtship behaviour in males of T. brochymenae is triggered by a short‐range sex pheromone produced by females; (2) the age and the physiological condition of females (virgin/mated) influence pheromone release or production; (3) the female's mesosoma is the source of the sex pheromone; and (4) polar components of the sex pheromone play a major role in influencing male behaviour. Our results suggest that quasi‐gregarious egg parasitoids are selected for short‐range rather than long‐range sex pheromones.     

Responses of Second-Instar Male Nymphs of Four Mealybug Species (Hemiptera: Pseudococcidae) to Conspecific and Heterospecific Female Sex Pheromones

The response of the late second-instar male nymphs of the mealybug species (Hemiptera, Pseudococcidae), Planococcus citri (Risso), Planococcus ficus (Signoret), Pseudococcus cryptus Hempel Nipaecoccus viridis (Newstead), to their conspecific and heterospecific female pheromone was studied. Males that exhibited the typical appearance of late-second-instar nymphs were tested. The male behavior was monitored soon after their exposure to the tested female sex pheromone in glass Petri dish arenas. Male nymph behavior toward the pheromone source was characterized based on their aggregation on the disks in the arena. Males of all four tested mealybug species were attracted to their conspecific female pheromone. By contrast, almost no interceptions of male nymphs with disks impregnated with a heterospecific female pheromone were observed. The mode of attraction of each of male nymphs of P. ficus, among most of the tested individuals (>80%), to the conspecific female sex pheromone, (S)-lavandulyl senecioate and or (S)-lavandulyl isovalerate, was the same as the mode of attraction latter on as adult. We suggest that by being attracted to the conspecific pheromone these males may direct themselves to a suitable pupation site near conspecific non-sibling mature females, thus preventing inbreeding. The repellency of heterospecific sex pheromone to males that are looking for a pupation site suggests that the latter try to avoid close contact with heterospecific females.     

Mating behaviour and chemical communication in the invasive Chinese mitten crab Eriocheir sinensis

Mating in the Chinese mitten crab (Eriocheir sinensis) was examined; in particular the nature of mating and the role of sex pheromones. A semi-lunar periodicity (16.8 days and 14.5 days, respectively) was observed in the mating frequencies in two consecutive breeding seasons (2001-2002 and 2002-2003). This semi-lunar rhythm coincided with spring tides (full and new moon), and activity peaked in November. Observation of the progression of specific behaviour types in mating and non-mating pairs revealed that pairs which would go on to complete mating progressed from fighting to mating behaviour significantly faster than non-mating pairs. These findings indicate that mate recognition occurs only after physical contact. Reproductively active pairs (ascertained from mating experiments) were then used for several bioassays aiming to assess under which conditions pheromones may be released by females. Firstly, male E. sinensis were exposed to female smell in an actograph experiment and secondly, male antennule flick rate was recorded before and after exposure to the urine of a sexually active female. In both cases no change in male E. sinensis behaviour was observed. Both experiments used females which had not had immediately prior exposure to males. However, in further experiments using water where mating had occurred, a significant response in antennule flick rate was triggered. Finally, a sponge assay was used in order to test the male attraction to a sponge injected with a water sample of varying concentrations (0.5×, 1×, 3×, 4.5×, 9×) of female smell. These samples were conditioned using a female immediately following a mating attempt. Males tried to grasp the sponge at 3× increased concentrations or higher. In conclusion, this study found no indication that E. sinensis females release a distance pheromone, but instead that mate recognition occurs after physical contact between male and female, most likely via a contact pheromone.     

The role of signals of different modalities in initiating vibratory communication in Nezara viridula

Signals of different modalities are involved in the behaviour of the green stink bug, Nezara viridula (L.) (Pentatomidae, Heteroptera). Long range attraction is mediated by male pheromones, resulting in aggregation of bugs on the same plant where vibratory signals, vision and various chemical signals become important. Both males and females sing spontaneously. When both are on the plant, males start vibratory communication as often as females. Females induce the exchange of vibratory signals spontaneously or triggered by the male pheromone while males initiate the duet either spontaneously or after seeing the female. Males and females sing spontaneously and respond to signals of different modalities more often in the daylight than in the dark. Long lasting autonomous emission of the female calling song is present when triggered by the male pheromone and males respond to female calling predominantly by the emission of the courtship song.     

Evidence for male- and juvenile-specific contact pheromones of the common bed bug Cimex lectularius

Males and females of the common bed bug, Cimex lectularius L. (Heteroptera: Cimicidae), have been shown to produce and respond to an aggregation pheromone. We tested whether juvenile C. lectularius also produce and respond to aggregation pheromone, and whether the pheromone is perceived by contact chemoreception. In dual‐choice laboratory experiments, juveniles, but not males or females, preferred juvenile‐exposed paper discs to control discs. Unlike juveniles, males and females preferred male‐exposed paper discs to control discs. Neither juveniles, males, nor females preferred female‐exposed discs to control discs. When test stimuli were inaccessible, C. lectularius failed to show any preference. Male‐ and juvenile‐specific contact pheromones may have contrasting functions of marking shelters as safe refugia for development and growth (juveniles) or mate encounter (adults), but result in the same phenomenon, the aggregation of conspecifics.     

Dermacentor variabilis and Dermacentor andersoni: Genital sex pheromones

There is evidence for the existence of a previously undescribed sex pheromone (or pheromones) in the ticks Dermacentor variabilis and D. andersoni. In addition to 2,6-dichlorophenol, which attracts mate-seeking males, a pheromone released on the cuticle of the female genital area enables the sexually excited male to locate the gonopore. The compound (or compounds) appears to act as a contact pheromone; male copulation responses are greatly reduced when the female genital surface is washed with solvents, especially hexane. It is also a potent excitant; males will puncture or dislodge barriers placed over the gonopore to copulate. However, the response is eliminated if the genital area is washed (hexane or acetone) prior to sealing the gonopore, suggesting the reproductive system as the source of the pheromone. A species specific copulation-eliciting pheromone appears necessary to excite the male to form and implant its spermatophore in the vulva of a conspecific female. Males encountering trans-specific females probe their gonopores, but mating attempts are almost always aborted within 5–10 min. The copulation-eliciting pheromone may be the same, or similar, to that used to locate the gonopore. Physical differences in the shape of the female gonopore in the two species, although slight, may contribute to the male's ability to identify conspecific females. Using this pheromone-guided process of attraction and identification, females present in mixed species populations will almost always be distinguished and inseminated by conspecific mate-seeking males.     

Evidence for a female sex pheromone in the brassica pod midge Dasineura brassicae

ABSTRACT. The responses of Dasineura brassicae Winn. (Diptera, Cecidomyiidae) to odours from both male and female midges and from their host ( Brassica napus ) and non-host ( Vicia faba ) plants were studied using a Y-tube olfactometer. Female but not male D. brassicae are attracted to crushed B. napus leaves but not to those of V. faba. Males are attracted to live virgin females, to crushed virgin and mated females and to hexane washes of females, suggesting that the female produces a sex pheromone that attracts males. Males were also attracted to hexane washes of female ovipositors, but not to washes of the bodies of females without ovipositors, suggesting that the ovipositor is the site of pheromone release or production. Males were not attracted to live mated females or to other live or crushed males and mated females were not attracted to other mated females.     

The effect of female body size on male mounting behaviour in Dermacentor variabilis and D. andersoni

Abstract. This study was undertaken to determine if body size of female ticks was an important factor in eliciting the mating behaviour of Dermacentor variabilis and D. andersoni males. Dummy female ticks (DFTs) representing varying sizes of feeding females were prepared from plastic beads. Size of DFT was shown to be important in determining both the time spent by males in contact with DFTs and the extent of male mating response released. Dermacentor varlabilis males preferred the smaller-sized DFT, whereas D. andersoni males preferred the larger sizes. Males of both species were able to discriminate between size of DFT only when mounting sex pheromone (MSP) was present.
Size of the DFT was more important than the composition of the MSP extract in determining the time males spent in contact with DFTs.
Males of both species were sensitive to variations in concentration of extract, and spent 2–6 times longer in contact with DFTs when the concentration was optimal than when it was not. No significant difference in the male's mating response was seen in response to variation in concentration of heterospecific extract.
Male mating response, i.e. the behaviour of the male as it progresses through the initial contact, climbing onto the dorsal surface and turning onto the venter, was shown not to be dependent upon how long the male spent in contact with the DFT. Male D. andersoni had a much lower mating response to DFTs than D. variabilis males, suggesting that some further stimulus may be required.     

Male Courtship Pheromones Affect Female Behaviour in the Swordtail Characin (Corynopoma riisei)

Pheromones constitute an important cue used by both males and females during courtship. Here, we investigate the effect of male pheromones on female behaviour in the swordtail characin (Corynopoma riisei), a species of fish where males have a caudal pheromone gland which has been suggested to affect female behaviour during courtship. We subjected female C. riisei to male courtship pheromones and investigated the effect on both female behaviour and brain serotonergic activity levels compared to a control group. While no difference in serotonergic activity was found, the pheromone‐treated females showed lower stress levels compared to the control group. Furthermore, pheromone‐treated females increased locomotor activity over time, while a decrease in locomotor activity was observed in the control group. These results suggest that the male courtship pheromones may serve to reduce female stress and increase female activity, possibly to aid males in gaining access to females and facilitating sperm transfer.     

Copulation behaviour of Lygocoris pabulinus under laboratory conditions

As a prerequisite to the elucidation of the sex pheromone of the green capsid bug Lygocoris pabulinus (L.), we studied the bug's reproductive development and behaviour. When kept under long-day conditions (L18:D6) at 25 °C, both males and females start to copulate 4 days after the final moult. Second matings occur in some females, even on consecutive days. The premating period is 10 min on average and copulation lasts 1 to 2 min. At long range, when males are attracted to traps baited with live females, we did not observe a specific calling position of the females. At short range, a characteristic courtship behaviour of the females was not observed either. Males showed a characteristic vibration of the abdomen, which was repeated several times before copulation. This behaviour can be used as a bioassay to test potential sex pheromone compounds.     

Male position and calling effort together influence male attractiveness in leks of the medfly,Ceratitis capitata (Diptera: Tephritidae)

Despite the close association between lek mating systems and the study of female mate choice, male mating success in leks is often associated with other aspects of sexual selection as well as female choice of male display traits. Males of the medfly Ceratitis capitata form leks on the undersides of leaves of their host plants. By experimentally creating artificial leks, we show that male success at attracting females depends not only on male calling effort (pheromone production dispersed by wing movement), but also on the position of the male within a lek. Males in the highest position in the artificial lek (closest to the light) attracted more females, and received more visits from those females. In our experiment, we deliberately minimized the visual cues that females approaching a male could use and, under these conditions, found no associations between male attractiveness and male size, weight or fluctuating asymmetry, either of the wings or sex setae (a pair of bilateral supra‐fronto‐orbital bristles). The latter result contrasts with earlier studies showing a significant negative association between sex setae fluctuating asymmetry and mating success. Accordingly, we conclude that symmetry of the male sex setae has no role in nonvisual communication (e.g. through directing pheromone plumes). Mating patterns associated with this insect are therefore every bit as complex as those in vertebrate leks. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 479–487.     

Linyphid spider courtship: Releaser and attractant functions of a contact sex pheromone

Experiments with male bowl-and-doily spiders (Frontinella communis) on methanol-washed and unwashed female webs indicate that the courtship of males is chemically released. Parallel experiments on the webs of females of another linyphiid species and on immature F. communis webs indicate that the releaser pheromone is species specific and that it is produced only by mature females. The pheromone that releases male courtship also functions as an attractant, changing the male's orientation from negatively to positively geotactic. The presence or absence of a female on the test web has only a slight effect on the male's behaviour.     

松墨天牛是否存在雌性接触信息素?

将松墨天牛MonochamusalternatusHope的身体不同部位切除之后的交配试验显示,分别剪去头部、鞘翅以及前胸部后仍然可以正常交配,用雄虫的腹部代替雌虫的腹部后,仍然能够引起雄虫的交配趋势动作,但不能完成真正交配,因此推测雌性接触信息素如果存在,主要集中在雌虫身体后半部,尤其是中后胸部位。但是冻死后的雌虫不再引起雄虫的交配行为,而且冻死的雌虫与活雌虫同时放入培养皿中,雄虫喜欢与活雌虫交配,不选择冻死的雌虫,因此推测视觉在其松墨天牛的交配过程中扮演着重要的角色,而接触性信息素有可能并不存在。将雄虫的触角分别从第5~6节和第1节剪去的试验结果显示,雄虫仍然可以很快地找到雌虫,发生交配行为。这有2种可能的解释一是雄虫不仅仅靠触角来感受雌虫的接触信息素,头部下颚须和下唇须上或者足上也有可以感受信息素的感受器;二是雌虫不存在接触信息素,雄虫通过视觉就可以找到对方,完成交配过程。浸泡致死的雌虫不再引起雄虫的交配行为,并且将正己烷和乙醚的浓缩液滴加在浸泡致死的雌虫以及玻璃棒和鹅卵石的生测试验结果显示,都没有引起雄虫的交配行为。这进一步说明,松墨天牛可能并不存在体表接触信息素。性成熟后的松墨天牛雌雄成虫是靠衰弱寄主的强烈的挥发性气味聚集到寄主树干上,在树干这个较为狭小的环境里,有可能是通过视觉找到配偶,完成交配过程。     

An analysis of anemotactic flight in female moths stimulated by host odour and comparison with the males' response to sex pheromone

ABSTRACT. The flight pattern of mated female navel orangeworm moths, Amyelois transitella (Walker), responding to odour from potential larval hosts is zigzagging upwind flight. However, at times these moths are capable of flying nearly directly upwind towards the odour source (track angles near 0). This response indicates that these females are capable of very accurate anemotactic control of their heading or course angle, since small angular errors in this measure would translate into larger deviations from direct upwind flight. Males of this species exhibit flight patterns similar to those of females, including track angles clustered about 0 when flying upwind to a source of the female-produced pheromone, but under these experimental conditions they flew with a higher average airspeed than the females. When females lose contact with an odour plume they initiate a well-defined programme of cross-wind counterturning or casting, which may normally increase their chances of retrieving contact with that plume when the wind direction shifts. The resultant track angles of females increase significantly by 0.8 s after plume loss, indicating that the female has initiated changes in both her course angle and airspeed. By 1 s after plume loss the females' track angles are no longer unimodally distributed about 0, but are bimodally distributed about -90 and +90. Males responded more rapidly to the loss of a pheromone plume, demonstrating a significant change in track angle 0.4 s after plume loss. Overall, female and male A.transitella exhibited remarkably similar anemotactic flight manoeuvres during upwind flight to odour sources as well as after plume loss.     

Chemosensory stimulation of luteinizing hormone secretion in male Siberian hamsters (Phodopus sungorus)

Male Siberian hamsters (Phodopus sungorus) housed in long days (LD), but not short days (SD) release luteinizing hormone (LH) when exposed to females. This study examined whether this response is specific to a female and identifies the source of a stimulus that induces LH release. Serum concentrations of LH, testosterone (T), follicle stimulating hormone (FSH), and cortisol were examined in all experiments. T concentrations mirrored the LH response; FSH and cortisol were unchanged in response to all stimuli. Exposure to an LD female, irrespective of her reproductive status, but not an SD female, elicited LH release. Exposure to another male did not trigger LH release. Males released LH when allowed physical contact with an anesthetized female, but not when separated from a normally active female, suggesting that tactile or nonvolatile chemosensory stimuli elicit LH release. Urine and secretions collected from the vagina as well as oral, midventral, perineal, and rectal glands, elicited marked behavioral responses in male P. sungorus. Despite these behavioral responses, only feces from females elicited LH release in males. Males released LH in response to feces extracted from the rectum and to cotton swabs that had been rubbed against the rectal mucosa, suggesting that a component of rectal secretions may trigger LH release in male Siberian hamsters. Taken together, these data and previous data from our laboratory indicate that both the production of and the response to a pheromone that triggers the selective release of LH is regulated by day length.     

Inhibition of mating behaviour before feeding in the tick Aponomma hydrosauri

When on their host, both male and female adults of the reptile tick Aponomma hydrosauri require at least five days of feeding before normal mating behaviour will start. In off-host trials, fed females attract significantly more male contact than do unfed females, but unfed males show normal mating behaviour. All aspects of female mating behaviour are inhibited before feeding. Male mating behaviour has two stages: one comprises detachment and searching, the other, courtship behaviour. Only the former is inhibited before male feeding. This suggests the female signal has two components: one initiating searching, and the other initiating courtship. Males will always respond to the second component, but only after feeding will they respond to the first.