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.     

Firefly Courtship: Behavioral and Morphological Predictors of Male Mating Success in Photinus greeni

Differences in male mating success can generate selection on male morphological traits and courtship behaviors involved in male–male competition or female mate choice. In Photinus fireflies (Coleoptera: Lampyridae), courtship is based on bioluminescent flash signals produced by both sexes. We conducted field observations of Photinus greeni fireflies engaged in competitive courtships, in which females are able to simultaneously assess several males, to identify male morphological traits and courtship behaviors that might predict male mating success. Male morphological traits did not differ between males that successfully mated compared with unsuccessful males (dialoging males that did not mate). However, courtship behavioral interactions differed: successful males tended to have higher flash pattern rates (number of flash patterns per minute), and their courtship flashes were more likely to be answered by females. We also examined how the risk of predation by Photuris fireflies altered courtship behavior of their Photinus prey. When predatory Photuris fireflies were present, P. greeni females were less likely to mate, and showed decreased flash responses to most males. However, P. greeni males that did successfully mate in spite of Photuris presence were males that maintained high flash pattern rates that elicited female responses. These results suggest that both female mate choice and Photuris predation exert strong selective pressures on the evolution of courtship signals in Photinus fireflies.     

Competitive Mating Success and Attractiveness of Sterile and Fertile Males of Drosophila montana

The attractiveness and the mating success of fertile (normal and brownie, br) and sterile (rudimentary testis, rt) Drosophila montana males were studied in female-choice experiments (one female + two males). The normal males courted the sterile rt males nearly as much as they courted the females, while they courted other males less frequently. The females required a long courtship with the rt males and accepted them as their mating partner less often than normal males. The courtship behaviour of normal, br and rt males was recorded in single pair situations with a normal female. The courtships of the males of all types proceeded in a similar manner, but the successful courtships of the rt males included more licking and touching than those of the normal and br males. The licking and touching periods are probably the critical phases at which the females are able to detect differences in male fertility.     

Mate choice by wild and mass‐reared females of the Mediterranean fruit fly

The sterile insect technique (SIT) is used to control Mediterranean fruit fly, Ceratitis capitata (Wiedemann), but its effectiveness is limited by low sexual competitiveness of mass‐reared males. This study investigated whether wild and mass‐reared [from a temperature sensitive lethal (tsl) genetic sexing strain] females display similar mate preferences and thus exert similar selective forces on the evolution of male courtship behaviour. Wild females preferred wild males over tsl males, whereas tsl females mated indiscriminately. The probability that mounting resulted in copulation was related to the duration of pre‐mount courtship for wild females, and wild males performed longer courtships than tsl males. Copulation occurred independently of courtship duration in tsl females. Counter to the aim of the SIT, female choice by tsl females appears to promote the evolution of male behaviour disfavoured by wild females.     

Mating with sperm-depleted males does not increase female mating frequency in the parasitoid Lariophagus distinguendus

Sexual conflicts due to divergent male and female interests in reproduction are common in parasitic Hymenoptera. The majority of parasitoid females are monandrous, whereas males are able to mate repeatedly. Thus, accepting only a single mate might be costly when females mate with a sperm‐depleted male, which may not transfer a sufficient amount of sperm. In the present study, we investigated the reproductive performance in the parasitoid Lariophagus distinguendus Först. (Hymenoptera: Pteromalidae) and studied whether mating with experimentally sperm‐depleted males increases the tendency of females to remate. Males were able to mate with up to 17 females offered in rapid succession within a 10‐h test period. The resulting female offspring, as an indirect measure of sperm transfer, remained constant during the first six matings and then decreased successively with increasing number of copulations by the males. Experimentally sperm‐depleted males continued to mate even if they transferred only small amounts or no sperm at all. Unlike males, the majority of females mated only once during a 192‐h test period. A second copulation was observed only in a few cases (maximum 16%). The frequency of remating was not influenced by the mating status of the first male the females had copulated with, suggesting that these events are not controlled by sperm deficiency of the females. Furthermore, we investigated male courtship behaviour towards mated females. Male courtship intensity towards mated females decreased with increasing time. However, females that had mated with an experimentally sperm‐depleted male did not elicit stronger or longer‐lasting behavioural responses in courting males than those that had mated with a virgin male. As the observed behaviours in L. distinguendus are known to be elicited by a courtship pheromone, these results suggest that females no longer invest in pheromone biosynthesis after mating (as indicated by ceasing behavioural responses of courting males), irrespective of whether they have received a sufficient amount of sperm or not. We discuss the results with respect to a possible mating strategy of sperm‐depleted males.     

Courtship behavior of Heliconius erato phyllis (Lepidoptera, Nymphalidae) towards virgin and mated females: conflict between attraction and repulsion signals?

Previous observations of Heliconius erato phyllis females being intensively courted a few days after mating contradict the proposal of sexual repulsion caused by a male-transferred antiaphrodisiac. Furthermore, data on courtship in Heliconius butterflies are almost absent from the literature. In this work we aim to describe the courtship behavior of H. erato phyllis and to compare it towards virgin and mated females. Sexual interactions using both kinds of females were observed and filmed in seminatural conditions for subsequent analysis and quantification. Courtship of virgin and mated females differed qualitatively and quantitatively, but the results do not agree with the proposal of sexual repulsion. The differences found indicate that the courtship sequence towards mated females is interrupted in the transition that would lead to the behavior immediately prior to copulation, in such a way that the male continues courting the female in an intermediate phase. Preliminary tests even showed that older females, virgin or not, are much less courted by males. We suggest that there is a trade-off between an attraction pheromone, important for males to locate female pupae, and the male-transferred antiaphrodisiac, so that young, mated females may have both signals active. .     

FUNCTIONAL ASPECTS OF DROSOPHILA COURTSHIP

1. The elements that make up the courtship behaviour of males and of females are briefly described. It is pointed out that some of the terms used, such as female ‘repelling’ behaviour, are misleading as they do not reflect the known functions of the behaviours. 2. Evidence has been presented for a number of distinct pheromones with different functions during courtship. These claims are critically examined as the evidence is incomplete and at times conflicting. It seems unlikely that any pheromones other than those acting over a very short distance are involved in courtship. There is sound evidence for an aphrodisiac pheromone produced by all females which stimulates male courtship. A pheromone, which may be the same one, is produced by males less than 12 h old, which also stimulates male courtship. No function is ascribed to this pheromone. Fertilized females either produce less aphrodisiac pheromone or they may, in addition, produce one that inhibits male courtship. Mature males may also produce an inhibitory pheromone. Females produce a contact pheromone which is species-specific and involved in sexual isolation. It is not at present clear whether this is different from the aphrodisiac pheromone. 3. There is considerable variability in the importance of vision in courtship. Many species will mate satisfactorily in the dark, suggesting that visual stimuli are not critical. Most species use vision to orient towards one another and for males to track and follow females. Even in light-independent species such as D. melanogaster, specific visual signals may be used in courtship although they are not obligatory. Thus the red eye of the male is a sexual signal for females. Conversely, some light-dependent species do not appear to make use of visual signals as a major factor in courtship. Some, however, do perform behaviours that are clearly visual and which may act to emphasize markings on wings, head or body. 4. The majority of Drosophila species perform courtship songs by vibrating one or both wings. The songs produced by males sexually stimulate the females. They are species specific and there is considerable indirect and some direct evidence that the songs are involved in sexual isolation. Males of many species produce two different songs during courtship and it is probable that one is concerned mainly with sexual stimulation and the other with species recognition. Females of certain species of Drosophila and Zaprionus also sing during courtship and these songs may aid species recognition by males. In addition males and unreceptive females perform ‘aggressive’ songs. 5. Almost all studies of Drosophila courtship have been made in very confined conditions in the laboratory. Interpretation of some of the results obtained in this way may require modification in the light of ecological research and observation of courtships under more natural conditions.     

Interactions of the Males and Females of Three Sympatric Drosophila virilis-Group Species, D. montana, D. littoralis, and D. lummei, (Diptera: Drosophilidae) in Intra- and Interspecific Courtships in the Wild and in the Laboratory

Four species of the Drosophila virilis group, D. montana, D. littoralis, D. lummei, and D. ezoana, occur sympatrically in several locations in northern Europe. Courtship interactions between the flies of the three first-mentioned species were observed at malt baits in Kemi, northern Finland, to find out how the flies of different species recognize conspecific individuals and how interspecific courtships differ from intraspecific ones in the wild. Intraspecific courtships (including females of different reproductive stages) and interspecific courtships were also videotaped and analyzed in laboratory. In the wild the males courted both conspecific and allospecific females, even though the species varied in how much the males were attracted to females of different species. Interspecific courtships usually broke off when the male touched the female or when the male and/or the female vibrated his/her wings, producing acoustic cues. In the laboratory males courted conspecific females irrespective of the reproductive stage of the female, even though the courtships directed toward immature and fertilized females usually included only orienting and touching (no licking and singing). D. littoralis, and very rarely D. montana and D. lummei, males courted also allospecific females. In the few interspecific courtships between these three species, where the male proceeded to singing, females responded to male singing by vibrating their wings. This ended the courtship. It is suggested that both the chemical cues affecting female attractivity and the acoustic signals of males and females, which are produced by wing vibration, function in maintaining sexual isolation between these three species.     

The courtship behavior of the cabbage moth,Mamestra brassicae (Lepidoptera: Noctuidae), and the role of male hair-pencils

The courtship behavior of the cabbage moth, Mamestra brassicae(L.), was studied in moving air conditions in a wind tunnel, using video techniques. Quantitative analyses were undertaken to determine the behavioral sequence occurring in the courtship. Comparison of successful and unsuccessful courtship suggested that courtship success was more dependent on the behavior of the female than that of the male. In an attempt to elucidate the function of the male hair-pencils (HPs), courtships involving males without HPs were also studied. HP removal did not affect the overall courtship success rate of males, but detailed analysis showed significant changes infernale behavior during such courtships. HP removal also affected female behavior following pair formation, with females struggling more when paired with males without HPs. Consequently, it is proposed that the HP volatiles act as an arrestant for the female, both during courtship and after pair formation, to increase female acceptance and to prevent premature termination of copulation. Experiments were also conducted to test previous hypotheses for HP function. However, no evidence was found to suggest that the HP volatiles in M. brassicaeact to attract females, affect female calling behavior, or affect the behavior of other males. A further possible function of HPs in defense is discussed.     

Courtship Behavior of the Cumberland Plateau Woodland Salamander,Plethodon kentucki (Amphibia: Plethodontidae), with a Review of Courtship in the Genus Plethodon

Complete catalogs of courtship behavior are available for only seven of the 42 currently recognized species of Plethodon. Additional detailed studies of courtship behavior in Plethodon species are needed to analyze the evolution of courtship behavior in this genus. We investigated the courtship behavior of the Cumberland Plateau woodland salamander (Plethodon kentucki) and compared it to previous accounts of courtship in other Plethodon species. In the laboratory, we videotaped the complete courtship of 30 different P. kentucki pairs, which included 46 tail-straddling walks that resulted in spermatophore deposition. From a transition matrix of observed motor patterns, we constructed a flow diagram of significant motor-pattern transitions during courtship. In general, the courtship behavior of this species is very similar to that of other large eastern Plethodon; however, there are some notable differences. Prior to the tail-straddling walk, the male transfers courtship pheromone from his mental gland to the female's nasolabial grooves (via mental-gland tapping and ‘chin-to-chin’ behavior patterns) more frequently than in other Plethodon. In most courtships, the female initiates contact leading to the tail-straddling walk. Males exhibit a greater propensity to deposit multiple spermatophores per courtship (two deposited in 33% of courtships, three deposited in 10% of courtships, X = 1.53 spermatophores per courtship) than males of other plethodontids. Mean size of the spermatophore is smaller than in other large eastern Plethodon. Overall, the courtship behavior of this species is most similar to that of P. jordani.     

Information transmission during intra- and interspecific courtship inEphestia elutella andCadra figulilella

Using information theory, courtship posturing in the moths Ephestia elutella(Hübner) and Cadra figulilella(Gregson) was analyzed for information transmission, which was partitioned into autocovariability (intraindividual transmission) and cross-covariability (interindividual transmission). This two-factor analysis was sufficient to account for more than 60% of the behavioral variance in males of E. elutellaand in both sexes of C. figulilelladuring intraspecific courtships; however, there were large residual variances in the behavior of male and female C. figulilelladuring interspecific courtships and in E. elutellafemales during both inter- and intraspecific courtships. In E. elutella,significant levels of transmission were attributable to both inter- and intraindividual effects, whereas in C. figulilella,only autocovariability was high and no interindividual communication could be assigned to courtship postures. Although courtship in these two species was qualitatively very similar and males readily courted nonconspecific females, high levels of reproductive isolation resulted from courtship. Male C. figulilellahad 94% fewer copulations with E. elutellafemales than with conspecific females and E. elutellamales had 78% fewer copulations with C. figulilellafemales than with conspecifics. These reductions were due to a differential response in both females and males, causing inter-specific courtships to be terminated much earlier than intraspecific courtships. This discrimination indicates that interindividual communication was indeed occurring during courtship and was only partially measured by analysis of postures. Thus, communication took place largely in some other modality, most likely the chemical modality, where species specificity is suggested for both male and female pheromones.     

Visual and Olfactory Female-Borne Cues Evoke Male Courtship in the Aphid Parasitoid Aphidius colemani Viereck (Hymenoptera: Braconidae)

We investigated the courtship and mating behavior of the pan-tropical polyphagous endoparasitoid Aphidius colemani Viereck. The courtship and mating displays, the magnitude of male-male sexual approaches and the role of female-borne cues evoking male courtship behavior were quantified. The sequence of events leading to copulation in this parasitoid did not differ from that found for other braconids. Females refused to copulate more than once. Same-sex courtships were observed among males and their possible role in an adaptive context is discussed. Olfactory female-borne cues played a key role in eliciting the courtship responses of males. Males were attracted by freshly dead females, but not by dead females soaked in hexane, nor by visual cues from females alone. Intense male wing fanning behavior was elicited by crushed abdomens of virgin females, suggesting that the female abdomen is the source of a short-distance pheromone crucial in evoking male courtship. Further studies are required to clarify the exact nature of the chemicals involved.     

The nature and development of sex attractant specificity in cockroaches of the genus Periplaneta. I. Sexual dimorphism in the distribution of antennal sense organs in five species

Sexual dimorphism in the distribution of antennal sense organs is common among adults of the genus Periplaneta. In three out of the four strains of Periplaneta americana examined, adult males had more contact chemoreceptors than females. In the fourth strain of P. americana and in P. australasiae, P. brunnea, P. fuliginosa, and P. japonica, no statistically supportable sexual dimorphism of contact chemoreceptors was found. However, in all strains and species of Periplaneta examined, sexual dimorphism was found in the total number and/or density of olfactory sensilla. Male adults had nearly twice as many olfactory sensilla as female adults. These observations are consistent with the behavioral observation that males within the genus Periplaneta rely on the reception of an airborne pheromone for the initiation of courtship behavior. In P. americana, where sexual dimorphism was found in the contact chemoreceptors, contact stimuli release the full wing raising display and presentation in males during courtship.     

Analysis of the Courtship Behavior of the Navel Orangeworm, Amyelois transitella (Walker) (Lepidoptera: Pyralidae), with a Commentary on Methods for the Analysis of Sequences of Behavioral Transitions

The courtship behavior of the navel orangeworm, Amyelois transitella, was examined in a wind tunnel. Sixty nine courtship sequences were analyzed and successful sequences divided into two categories: rapid courtship sequences, which involved few breaks in contact, short or no periods of male/female chasing and lasted <10 s between initial contact and mating; and prolonged courtship sequences, which involved many breaks in contact, extended periods of male/female chasing and lasted >10 s. Fifty six (81%) courtships were successful (50.7% rapid courtship and 30.4% prolonged courtship); the remaining 13 (18.8%) sequences were failed courtships. Of failed courtships, 9 (13.0%) were due to males losing contact with females during courtship chases and 4 (5.8%) due to females flying away immediately after male contact. Of all courtship sequences involving a break in contact during a chase, 38.5% resulted in an unsuccessful mating attempt. These findings contrast with previous studies of the courtship behavior of the navel orangeworm, potentially indicating that the type of bioassay used to study courtship may have a large effect on the behavioral sequences displayed. We evaluate several diagnostic techniques for the analysis of sequences of behavioral transitions.     

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.     

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.     

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.     

Generalization of courtship learning in Drosophila is mediated by cis-vaccenyl acetate

Reproductive behavior in Drosophila has both stereotyped and plastic components that are driven by age- and sex-specific chemical cues. Males who unsuccessfully court virgin females subsequently avoid females that are of the same age as the trainer. In contrast, males trained with mature mated females associate volatile appetitive and aversive pheromonal cues and learn to suppress courtship of all females. Here we show that the volatile aversive pheromone that leads to generalized learning with mated females is (Z)-11-octadecenyl acetate (cis-vaccenyl acetate, cVA). cVA is a major component of the male cuticular hydrocarbon profile, but it is not found on virgin females. During copulation, cVA is transferred to the female in ejaculate along with sperm and peptides that decrease her sexual receptivity. When males sense cVA (either synthetic or from mated female or male extracts) in the context of female pheromone, they develop a generalized suppression of courtship. The effects of cVA on initial courtship of virgin females can be blocked by expression of tetanus toxin in Or65a, but not Or67d neurons, demonstrating that the aversive effects of this pheromone are mediated by a specific class of olfactory neuron. These findings suggest that transfer of cVA to females during mating may be part of the male's strategy to suppress reproduction by competing males.     

The role of silk in courtship and chemical communication of the false widow spider, <Emphasis Type="Italic">Steatoda grossa</Emphasis> (Araneae: Theridiidae)

In spiders, sex pheromones are often associated with silk produced by females, and function in mate attraction, recognition, and evaluation. Silk-bound pheromones typically elicit courtship behaviour in web-building spiders. Here we (1) describe courtship interactions of Steatoda grossa males with virgin or mated females, and (2) show that silk and methanol extracts of silk produced by virgin females trigger courtship behaviour (silk production) by males, whereas silk of mated females does not. Our results indicate that (1) virgin females produce a silk-bound sex pheromone, (2) males discriminate between virgin and mated females based on silk cues, and (3) male silk likely functions in sexual communication.     

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.