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.     

Corpus al latum regulation of copulatory behaviour in the male grasshopper, Melanoplus sanguinipes

Abstract The relationship between corpus allatum (CA) regulation of copulatory behaviour and the CA's influence on the development of the accessory reproductive glands was studied in the male grasshopper, Melanoplus sanguinipes (Fabr.). Allatectomy slowed the development of copulatory behaviour; ≤80% of control males 7 days and older mated with females, but allatectomized males did not show comparable levels of copulation until day 19. In a 3 h observation period, the time taken for 50% of males to initiate copulation was c. 60 min for allatectomized males and only c. 10 min for controls. At the age when 50% of males copulated, the accessory glands of allatectomized males were less developed than those of controls; the accessory gland weight was 65%, soluble protein was 72% and reserves of long hyaline protein I were 40% those of controls. Treatment with Juvenile Hormone III on day 2 stimulated accumulation of long hyaline protein I in 9-day-old allatectomized males to the level found in untreated 19-day-old allactec-tomized males, but did not elicit heightened copulation. No positive feedback from the accessory glands appeared necessary to elicit copulation since vasectomy, ablation of the accessory glands or isolation of the terminal abdominal ganglion by ventral nerve cord transection did not inhibit copulatory behaviour in 7-day-old males. Because allatectomy inhibited copulatory behaviour in 7-day-old males in which the accessory glands were removed but not in 19-day-old males without accessory glands, the increased tendency of older allatectomized males to mate with females was not due to release of inhibition caused by continued growth of the accessory reproductive glands.     

Grasshopper sexual pheromone: a component of the defensive secretion in Taeniopoda eques

ABSTRACT. Both male and female Taeniopoda eques (Burmeister) emit a defensive secretion from their thoracic spiracles. The secretion from mature females was found to act as a sex pheromone, eliciting mating behaviour in males. Females became chemically attractive to males about 16–18 days after eclosion. The antennae are shown to be the site of pheromone reception in males. Feeding upon natural host plants was not a prerequisite for pheromone production.     

Mate discrimination in Littorina littorea (L.) and L. saxatilis (Olivi) (Mollusca:Prosobranchia)

The ability of males of Littorina littorea and L. saxatilis to discriminate between mates of different sex, species and size was examined. In partner choice experiments males of L. littorea had the possibility to initiate a copulation with either a female or a male. The males did not show a preference for either sex. There was therefore no evidence that they could determine the sex of a conspecific prior to copulation. The duration of intrasexual copulation was considerably shorter than for intersexual copulation, both in the field and in laboratory experiments. For the two species, intersexual copulations were far more frequent than intrasexual ones. This can partly be explained by the difference in copulation time.Few interspecific copulating pairs were found on the shore. This may reflect a low interspecific encounter rate rather than a mechanism of species recognition. On all of these occasions, however, the active male was of L. saxatilis. It is argued that selection against precopulatory species and sex recognition is a more likely explanation than an hypothesis that states that the required mutations for precopulatory mate identification has not yet occurred. L. littorea males copulated longer with large than with small females. Copulation time was short with parasitized females, which are sterile or of low fecundity. The allocation of mating effort by males is discussed.     

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.     

Pheromonal inhibition of male courtship behaviour in the brown tree snake, Boiga irregularis: a mechanism for the rejection of potential mates

Pheromones play a central role in coordinating the events leading up to copulation in snakes. We report here a novel pheromone system in the brown tree snake in which females release a pheromone that inhibits male courtship behaviour. In a previous study, we made observations of female brown tree snakes releasing cloacal secretions (CS) during courtship that appeared to cause courting males to cease courtship. All snakes have glands that release CS through ducts located along the cloacal orifice. Although CS have been studied for many years, their function in the mediation of snake behaviour has not been experimentally well determined. We examined the role of CS in the reproductive behaviour of male and female brown tree snakes. We conducted four experiments to test the effect of both male and female CS on brown tree snake behaviour under two behavioural contexts, courtship and male-male ritualized combat. Within each experiment, we compared the effects of CS to a control. Female CS caused a decrease in the time that males spent courting females and a decrease in the intensity of courtship compared with the control treatment. Male CS did not, however, affect the time that males spent displaying courtship or the intensity of that courtship. Neither male nor female CS had significant effects on male ritualized combat behaviour, including time that males spent in combat or the intensity of combat behaviours displayed. Furthermore, neither female nor male CS had an effect on female courtship versus controls. The inhibition of brown tree snake reproductive behaviours is specific to female CS inhibiting male courtship behaviour. This pheromone acts in concert with the female sex pheromone to regulate the events leading to copulation.Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Antennal glands in Psylliodes chrysocephala, and their possible role in reproductive behaviour

Abstract. The antennal morphology of Psylliodes chrysocephala L. (Coleoptera: Chrysomelidae), an important pest of oilseed rape ( Brassica napus ), was studied. Two types of tricellular, integumentary glands were found. The common antennal glands are distributed under all sensilla-bearing parts of male and female antennae. The male-specific antennal glands are only located under a glabrous area found on antennomeres 6–10 of male beetles. The common antennal glands are synthetically active in both pre- and post-diapause adults, but the male specific antennal glands are only active in post-diapause (reproductively active) males. During studies of mating behaviour, the antennae of the males were highly active at the beginning and end of copulation, and in response to increased female activity. The male specific antennal glands may secrete a sex pheromone, and the glabrous area on the male antennae could be the release site for such a pheromone.     

The adaptive significance of mate guarding in the soapberry bug,Jadera Haematoloma (Hemiptera: Rhopalidae)

Male soapberry bugs (Jadera haematoloma)face severe mating competition at the northern edge of their range due to male-biased adult sex ratios. Copulations lasting up to 11 days may serve a mate guarding function (encompassing four or more ovipositions), but copulation duration is highly variable, with some pairings lasting as little as 10 min. Data were gathered to describe factors that influence the reproductive costs and benefits of prolonged copulation. Estimated copulation durations (mean ± SD) were 20 ± 23 h in the lab and 50 ± 8 h in the field and were only weakly affected by sex ratio. Females mated for 5 min produced as many fertile eggs as those mated for 600 min laid; they became depleted of fertile sperm after about 25 days. In twicemated females, the first male's paternity was reduced by about 60%, and all females (N = 13) whose mates were removed experimentally mated again within an average of 6 min. The outcome of sperm competition on a perclutch basis was not highly predictable. The possibility of increased sperm displacement in longer copulations was not tested. Males often guarded females during oviposition and successfully defended them from intruding single males by recopulating. Such intrusions occurred in the majority of oviposition attempts observed in nature. Even though most females mated promiscuously, in a focal aggregation with a mean sex ratio of 2.2 ± 0.4 males/female, the interval between matings by males was commonly several days. Males appeared to respond facultatively to several aspects of the distribution and availability of females. The intensities of mating competition and sperm competition indicate that monogamous mate guarding should be favored over nonguarding in nature. Unpredicted brief. pairings may result from assessment by males of female reproductive value or of their own physical condition, or from female resistance.     

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.     

Mating behaviour and reproductive isolation of three species of reptile tick

The mating behaviour of three species of reptile tick, Aponomma hydrosauri, Amblyomma albolimbatum and Amb. limbatum is similar and involves a female sex pheromone which activates males to detach from their hosts and search. After contacting females, a stereotyped six-phase courtship sequence occurs. There are qualitative differences between the species in courtship behaviour at phase 3 (reversal of position by dorsally mounted males) and phase 6 (the copulation position). On-host observations of non-conspecific mating show that females of the three species have species-specific activation pheromones, which is contrary to reports in other species of tick. Such specificity should result in reproductive isolation of the three species; however, under certain circumstances it may not prevent non-conspecific contacts between the sexes. Off-host observations of courtship behaviour show that once males of the three species contact non-conspecific females, they attempt courtship and are persistent with their courtship. Rarely did non-conspecific courtships proceed beyond phase 4 of the courtship sequence, as non-conspecific females did not lift their bodies to allow males venter contact. Differences between the species in leg orientation in the copulation position, together with body size differences, are responsible for a complete barrier to successful non-conspecific copulation. The observations illustrate the role that behavioural mechanisms play in reproductively isolating these three species of tick.     

Reproductive biology of Pseudococcus maritimus (Hemiptera: Pseudococcidae)

This study characterized several parameters associated with reproductive behaviors of a Californian population of the grape mealybug, Pseudococcus maritimus (Ehrhorn). Experiments with caged adults confirmed that females from this population could not reproduce parthenogenetically. Both sexes were capable of mating multiple times on the same day and on sequential days. Median times between copulations were short (< 10 min) on the first day that males were presented with females, but tended to increase with sequential copulation events. Unmated females lived for up to 19 wk, whereas mating and oviposition resulted in reduced longevity. We hypothesized that constant exposure to pheromone might stimulate males to leave their cocoons sooner and increase their activity levels compared with males that were not exposed to pheromone, resulting in shortened life-spans. However, whereas exposure to pheromone did stimulate males to emerge slightly sooner than control males, pheromone exposure had no effect on the longevity of males. The implications of the results of these experiments for pheromone-based methods of detection and control of mealybugs are discussed.     

Differential response of Trichogramma wasps to extreme sex pheromone types of the noctuid moth Heliothis virescens

1. Chemical espionage in nature may occur when predators or parasitoids home in on animal or plant communication signals. Parasitoid wasps are known to use pheromones emitted by adults hosts to locate host eggs, larvae or pupae. The response of Trichogramma egg parasitoids to a synthetic sex pheromone blend of moths has been shown in a number of studies over the past 40 years. 2. Trichogramma pretiosum (Hymenoptera, Trichogrammatidae) is a tiny parasitic wasp, attacking the eggs of the noctuid moth Heliothis virescens (Lepidoptera, Noctuidae). This study investigated whether T. pretiosum homes in on the sex pheromone of H. virescens at close range. The arrestment response of the wasps to sex pheromone gland extracts of two types of female moths, so‐called high and low females, was also tested, referring to two selected extreme pheromone types of H. virescens. The study also investigated whether the wasps would mount females, possibly to hitchhike with them. 3. The wasps were arrested by the common, ‘low’ pheromone, but not by the rare, ‘high’ pheromone or by extracts from male hairpencils. The wasps did not show a preference for separate sex pheromone compounds, but when pre‐exposed to the major sex pheromone component of H. virescens before the tests together with H. virescens eggs, they did show a preference, indicating learning behaviour. In the mounting experiments, mated females were mounted significantly more than virgin females or males, suggesting that hitchhiking is a strategy used by these wasps to locate moth eggs. 4. This represents the first study to show a differential response of parasitoid wasps to two different sex pheromone types in a single host species. The results warrant further investigations into the potential role of parasitic wasps in the evolution of sexual communication in moths.     

Towards the identification and synthesis of the sex pheromone of the citrus leafminer, Phyllocnistis citrella Stainton (Lepidoptera: Gracillariidae)

The objective of this work was to characterize the sexual behavior of the citrus leafminer, Phyllocnistis citrella Stainton, as the foundation for the isolation, identification, and synthesis of the complete sex pheromone of this species. Mating occurred in a time window of 2h, starting 1h before the onset of photophase. The large majority of tested insects mated in the first two days after emergence, with no significant difference between mating at day 1 and day 2. A stereotypical courtship and copulation behavior were described for this species. When mating was successful, the copulation was recorded in average for 49.6 min. In Y-olfactometer tests conducted at the time of mating activity, males were strongly attracted to caged virgin females as well as to extracts from putative pheromone glands.     

Behavioural evidence for a female sex pheromone in Cotesia flavipes (Hymenoptera: Braconidae)

Abstract.  The mating behaviour of Cotesia flavipes Cameron, 1891 (Hymenoptera: Braconidae) was observed under laboratory conditions. The results demonstrate that chemical stimuli play an important role in bringing males and females together for mating. Females from this species release chemical compounds to attract males, which exhibit a distinct courtship behaviour divided into three main steps: (i) antennation, (ii) lateral bouncing and (iii) wing fluttering. Ethological tests using hexane extracts from two different parts of the female's body (head plus thorax and abdomen) demonstrate that males are more attracted and also display a courtship behaviour when a filter paper containing hexane extracts from the abdomen is presented to them, indicating that the abdomen is the source of the sex pheromone of this species. Naive males of this species respond to hexane extracts of this gland by displaying courtship behaviour. Furthermore, additional tests show that C. flavipes females mate only once, whereas virgin males do not show any preference for virgin or mated females, suggesting that mated females of this species continue to produce sex attractants after mating.     

Esterase 6 of Drosophila melanogaster: Kinetics of transfer to females,decay in females and male recovery

Esterase 6 (E.C. 3.1.1.1) is an enzyme in the reproductive system of Drosophila melanogaster and is transferred from males to females at mating. This enzyme is conveyed during the first 3 min of a copulation lasting approximately 20 min and remains active in the female reproductive tract for 1–2 hr after mating. Males require 24–48 hr to recover premating levels of esterase 6 activity. These results suggest that the effect of esterase 6 on the rate of sperm loss in females, which extends to 4 days after mating, is indirect since activity in the female falls rapidly after insemination. It is suggested that esterase 6 may also be a part of a pheromone system which influences female reproductive behaviour.     

Floral evolution and pollinator mate choice in a sexually deceptive orchid

Sexually deceptive orchids mimic sex pheromones and appearance of female insects to attract males, which pollinate the flowers in an attempted mating. This study examines the effects of pollinator mate choice on orchid floral evolution using the Thynnine wasp Neozeleboria cryptoides (Smith) (Hymenoptera: Tiphiidae), which pollinates the sexually deceptive orchid Chiloglottis trapeziformis Fitzg. (i) When male wasps were given the choice between two female dummies of different sizes and identical amount of synthetic pheromone, they preferentially attempted to copulate with medium-sized dummies over small dummies. (ii) When given the choice between two dummies of identical size but different amounts of pheromone, males preferred the larger amount of pheromone. Larger amounts of pheromone generally attracted more males than smaller amounts. (iii) Orchid flower labella, which mimic a female body, were significantly longer and broader than female wasp bodies, and the flowers also produced on average 10 times more 'pheromone' than females. The evolution and maintenance of these exaggerated mating signals is likely to be mediated by the male pollinator behaviour demonstrated here. (iv) When five dummies were offered simultaneously in a 10 cm circular array, males rarely attempted copulation on more than one dummy during a single visit. This behaviour may foster the evolution or maintenance of clonality in C. trapeziformis, as it will minimize pollen exchange within clones.     

Reproductive behaviour in the harpacticoid copepod Tigriopus fulvus

The results of several experiments on reproductive behaviour of Tigriopus fulvus, indicate that females release a sex pheromone promoting sexual recognition and attraction in males. This compound has been verified to be not only species-specific, but also slightly modified in specimens belonging to different geographical populations. The compound possesses a mass of between 100 and 1000 daltons. A male can invest from one to 15 days in the precopulatory phase; delayed fertilization causes a strong reduction in the average number of offspring produced by parental specimens belonging to the oldest agegroups, but does not have any effect on the sex ratio.     

The effect of mating behaviour on progeny sex ratio of Mastrus ridens (Hymenoptera: Ichneumonidae), a biological control agent of codling moth

Mastrus ridens (Horstmann) (Hymenoptera: Ichneumonidae) was imported into quarantine as a potential biocontrol agent for codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae). Mating behaviour of the parasitoid and its effect on progeny sex ratio (as a proportion of males) were studied to help sustain the laboratory culture. Both females and males were reproductively active soon after emergence. Unmated females produced only male progeny, confirming males developed from unfertilized eggs. The proportion of males in a progeny was independent of the copulation period (24–40 s) of the parents. The progeny sex ratios from three parent ratios (f:m 2:1, 1:1 and 1:2) were not significantly different. Females effectively mated only once but males mated frequently in their lifetime. The progeny sex ratios from single females with a known single mating and possible multiple matings (through exposure to two males for 18 days) were not significantly different. However, when males copulated with five virgin females in sequence over a 2 h period, the fifth female produced more males than the preceding four, presumably due to sperm depletion. Results of this study provide data on progeny sex allocation of M. ridens that help to prevent the development of a male-biased sex ratio that could threaten the maintenance of the culture over time.     

Male-specific (Z)-9-tricosene stimulates female mating behaviour in the spider Pholcus beijingensis

Chemical signals play an important role in spider sexual communication, yet the chemistry of spider sex pheromones remains poorly understood. Chemical identification of male-produced pheromone-mediating sexual behaviour in spiders has also, to our knowledge, not been reported before. This study aimed to examine whether chemically mediated strategies are used by males of the spider Pholcus beijingensis for increasing the probability of copulation. Based on data from gas chromatography–mass spectrometry analysis, electroantennography assay and a series of behavioural tests, we verified that (Z)-9-tricosene is a male-specific compound in the spider P. beijingensis. This compound acts as an aphrodisiac: it increases the likelihood that a female will mate. Mate-searching males release (Z)-9-tricosene to stimulate sexual behaviour of conspecific females. In the two-choice assay, however, sexually receptive females show no preference to the chambers containing (Z)-9-tricosene. This indicates that the male pheromone of P. beijingensis is not an attractant per se to the conspecific females. This is, to our knowledge, the first identification of a male-produced aphrodisiac pheromone in spiders.     

The mating behaviour of the egyptian cotton leafworm moth, spodoptera littoralis (Boisd.)

A detailed study of courtship in Spodoptera littoralis showed that there were four significant behaviour patterns. The male flew to a calling female and hovered above her with his brushes fully extended. In response, the female lifted her wings, curved her abdomen and withdrew her pheromone gland. The male settled beside the female to pair and then moved to hang head downwards during copulation. Thirty percent of successful courtships lacked one of the main behaviour patterns. Nearly half the courtships observed did not end in copulation: none of these included all of four major behaviour patterns and the majority lacked two or three. Females often rejected males with a rapid flick of the wings. Antennaless males did not mate or extend the brushes in response to a calling female. Just over half of the antennaless females observed during 135-min tests mated with normal males, but courtship was abnormal. Olfactory cues appeared to be important to females in recognizing the courting male, since antennaless females did not wing flick, were significantly more likely to take to flight as the result of the male brush display and frequently failed to retract the pheromone gland during the latter states of courtship. The courtship behaviour of S. littoralis is compared with published accounts for other Noctuids.