Courtship and mating by the sandfly Phlebotomus duboscqi, a vector of zoonotic cutaneous leishmaniasis in the Afrotropical region

Courtship behaviour of males of the Afrotropical sandfly Phlebotomus duboscqi Neveu-Lemaire (Diptera: Psychodidae) involved mounting the female and clasping her 'waist' with the male coxites placed between the female's thorax and abdomen. This behaviour, which we call 'piggy-backing', was preceded by male wing beating, perhaps involving mate recognition and contact pheromones. It did not seem to be pre- or postcopulatory mate guarding. Piggy-backing was attempted by P. duboscqi males on females of other species (P. papatasi and P. perniciosus) and even on other male P. duboscqi. The majority of female P. duboscqi piggy-backed by males were already inseminated, and most of the courting did not lead to copulation. This, coupled with the presence of a mating plug (semen) in each spermatheca of inseminated females, suggests that female P. duboscqi are monogamous for at least the first gonotrophic cycle. Male courtship with piggy-backing was more intense when females could feed on a hamster than when a hamster was present but the females were denied access to the host. It is suggested that, when a hamster was available to the females, the conditions in the laboratory are similar to those in rodent holes, the natural habitat of P. duboscqi.     

Male competition is disruptive to courtship in the Pacific blue‐eye

The present study examined the effects of competition on male courtship in the Pacific blue‐eye Pseudomugil signifer , a species of fish where females have previously been shown to use courtship, but not male fighting prowess, as an important mate choice cue. Courtship bouts directed to a stimulus female were shortest when two males were allowed to freely interact (contact treatment) and longest when there was only one male (non‐interaction). Courtship length in trials where one of two males was confined to a clear cylinder (visual) was intermediate between the other two treatments. Courtship in visual and contact treatments was constantly disrupted. The percentage of interrupted courtships was higher for the contact compared to visual treatment where aggressive interactions were also longer in duration and took place more frequently. Within contact trials, dominant males courted longer than subordinates despite both males experiencing comparable rates of courtship disruption. These results suggest that male‐male competition can have important implications for adaptive female choice particularly in circumstances where the benefits being sought by females are unrelated to male fighting ability.     

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 Comparative Analysis of Mating Recognition Signals in Graylings: Hipparchia statilinus vs. H. semele (Lepidoptera: Nymphalidae, Satyrinae)

Courtship is an important character linked to fitness and can evolve in response to female mate choice, to ensure sex identity and quality of a potential mate. It can also be a major pre-mating isolating mechanism in butterflies and many other species. In this paper, aspects of the reproductive behaviour of Hipparchia statilinus are described. First, the ethogram and the sequential structure of courtship leading to copulation and female refusal behaviour are showed; second, the results are compared with the behaviour of the sympatric and syntopic species, H. semele. Sequential analysis of H. statilinus courtship showed that, once a male has started the sequence, the following steps most likely follow up to the end. Courtship seems to reflect female behaviour: the male can repeat the whole sequence or part of it more than once, waiting for female signals. The general pattern of sexual behaviour in H. semele and in H. statilinus is mainly identical: perching strategy, flight pursuit, courtship. In both species, courtship consists of a highly stereotypic sequence, and differences between sequences are primarily related to presence/absence, order and performance of steps. This study is the first step of an experimental design addressed to a better understanding of mating recognition signals in Hipparchia and provides more details to define the relative roles of phylogeny and environment in shaping boundaries among species.     

Changes in courtship behaviour following rejection: The influence of female phenotype in Drosophila melanogaster

Courtship can be costly and so selection should favour individual males that reduce courtship towards female types that have a low probability of resulting in copulation. One way males can do this is by associating previous courtship failure with the traits of particular rejecting females. We characterised changes in male Drosophila melanogaster courtship behaviour following a failed mating attempt with one of the four female phenotypes that varied in size, age or mating status. To do this, we assessed individual courtship behaviour for each male presented again with a female of the same phenotype that previously rejected him. Males reduced subsequent courtship most strongly for recently mated (sexually non‐receptive) females. More interestingly, males also significantly reduced courtship activity following a failed mating experience from old females but did not do so for control (large, young, virgin) or small females. As such, males significantly reduced courtship towards both female types possessing chemical cues associated with their phenotype (age and mating status), but not towards a female phenotype based on physical characteristics (body size). Our results suggest that males are able to modify their courtship behaviour following experience, but that they are better prepared to associate chemical traits that may be more reliable indicators of the likelihood of courtship failure.     

Species and sex-specific songs and courtship behaviour in the genus Zaprionus (Diptera-Drosophilidae)

Courtship behaviour and songs of six Zaprionus species show differences not only between species but also between sexes. Courtship behaviour differs from that in the related genus Drosophila. In most species, males produce two song types which may alternate or be repeated. Singing occurs during courtship displays, mounting, after copulation and briefly during male-to-male interactions. Females produce a loud whine and body rocking movements in refusal but have a species-specific pulsed song produced during normal courtship which differs from the conspecific male song. One species produces an irregular male song but a regular female song. Sex-specific songs may have selective advantages but pose problems for the sensory template hypothesis.     

Copulatory courtship and cryptic female choice in red flour beetles Tribolium castaneum

Males of many animal species engage in courtship behaviours during and after copulation that appear to be solely aimed at stimulating the female. It has been suggested that these behaviours have evolved by cryptic female choice, whereby females are thought to impose biases on male postmating paternity success. Males of the red flour beetle Tribolium castaneum rub the lateral edges of the females' elytra with their tarsi during copulation. We manipulated female perception of this behaviour by tarsal ablation in males, thus preventing males from reaching the edge of the female elytra with their manipulated legs, and by subsequently performing a series of double-mating experiments where the copulatory behaviour was quantified. We found a positive relationship between the intensity of the copulatory courtship behaviour and relative fertilization success among unmanipulated males. This pattern, however, was absent in manipulated males, where female perception of male behaviour differed from that actually performed. Thus, female perception of male copulatory courtship behaviour, rather than male behaviour per se, apparently governs the fate of sperm competing over fertilizations within the female, showing that copulatory courtship is under selection by cryptic female choice.     

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.     

Comparative courtship behaviour in two species of the parasitic chalcid wasp Melittobia (Hymenoptera: Eulophidae)

Timing and form of courtship behaviour elements constitute a major isolating mechanism for two morphologically and ecologically similar parasitic wasps. Because males are blind, the complex courtship of Melittobia chalybii and a new as yet undescribed species previously confused with it, designated as M. sp. A, depends upon tactile and chemical cues. In both species, after the female orients to the male's abdomen, the male turns in the direction of contact and mounts. However, chalybii males antennate females continuously, lifting mesothoracic legs at regular intervals, while in sp. A male antennation alternates with metathoracic leg pumping. Courtship and copulation duration are greater in Chalybii than in sp. A.     

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.      

Copulatory Courtship in Drosophila: Behavior and Songs of D. birchii and D. serrata

D. birchii and D. serrata, two endemic Australian Drosophila species, have a copulatory courtship. The males of these species begin to court the female after mounting her and often go on with the courtship after the copulation is over. In the present paper we have described behavioral interactions between the male and the female and analyzed acoustic signals produced by the flies during courtship. Species differences were more pronounced in female than in male behavior. Variation within the species was obvious in the relative proportions of time the flies spent in different behaviors. Even though courtship took place nearly solely during copulation, some remains of precopulatory courtship were observed in both species. It is suggested that copulatory courtship exhibited by D. birchii and D. serrata flies is a derived rather than a primitive character.     

EVIDENCE FOR WIDESPREAD COURTSHIP DURING COPULATION IN 131 SPECIES OF INSECTS AND SPIDERS,AND IMPLICATIONS FOR CRYPTIC FEMALE CHOICE

Male courtship behavior is generally thought to function prior to copulation, as an inducement to the female to allow the male to copulate with her; this study indicates however, that male courtship during and following copulation (“copulatory courtship”) is common in insects and spiders (81% of 131 species in 102 genera and 49 families, mostly Coleoptera, Hemiptera, Diptera, and Araneioidea). Copulatory courtship is apparently evolutionarily labile, as expected if it is under sexual selection; intrageneric variation occurred in all 17 genera in which more than one species was observed. In 81% of 94 species with copulatory courtship, the male abandoned the female soon after copulation ended; thus, copulatory courtship appears not to function generally to induce acceptance of further copulatory attempts. The most likely explanation for copulatory courtship is that it represents attempts by males to influence cryptic female choice. This suggests that an aspect of sexual selection by female choice not considered by Darwin may be more important than previously appreciated and that the common practice in evolutionary studies of measuring male reproductive success by counting numbers of copulations may sometimes be misleading because of cryptic female choice during and after copulation.     

Mating Behavior of Cephalonomia tarsalis (Ashmead) (Hymenoptera: Bethylidae) and the Effect of Female Mating Frequency on Offspring Production

The courtship behavior of Cephalonomia tarsalis, a solitary semiectoparasitoid of Oryzaephilus surinamensis, was investigated in the laboratory. Courtship behavior includes a series of stereotypic movements. Males play the most active role, executing the majority of courtship action, and females respond with relatively limited observable behaviors. Males typically keep antennae still during encounters with females prior to mounting, which may be correlated with recognition of the female's sexual status. After mounting, males display a series of movements on females, such as antennae touching female's antennae, antennae or mouth touching female's head or thorax, and walking around on female, which may serve to stimulate females towards increased receptivity. Females signal receptivity by assuming a stereotypical posture of remaining stationary, with head down, and antennae still in front of the body. The male then inserts his aedeagus and the pair copulates. After an average of 40.4 s of copulation, females signal the end of copulation by waving the antennae and moving away from the copulation site. Males continue copulating for a short time after females start moving but dismount soon thereafter. After dismounting, the two wasps move away from each other immediately, and they typically begin grooming. Neither males nor females exhibit mating preference based on mate's mating status in both choice and no-choice tests. The male is polygynous and the mated female can mate multiple times within the first 3 days after starting oviposition. However, female mating frequency does not affect the production of female progeny.     

Singing on the wings! Male wing fanning performances affect female willingness to copulate in the aphid parasitoid Lysiphlebus testaceipes (Hymenoptera: Braconidae: Aphidiinae)

Lysiphlebus testaceipes (Hymenoptera: Braconidae: Aphidiinae) is a generalist endoparasitoid attacking more than 100 aphid species. In L. testaceipes, wing fanning is a main male courtship display evoked by a female‐borne sex pheromone. However, no information is available on the characteristics and behavioral role of male fanning during courtship in this parasitoid. Here, the courtship behavior of a wild strain of L. testaceipes was quantified and the male wing fanning performances were analyzed through high‐speed video recordings and examined in relation to mating success. Courtship sequence of wild L. testaceipes did not substantially differ from that previously reported for other populations mass reared on aphids. We observed that the male courtship duration did not affect mating success. However, video analysis revealed that the males producing high‐frequency fanning signals achieved higher mating success over those that display low‐frequency fanning. Wing fanning before successful and unsuccessful courtship differed in amplitude of wing movements and alignment toward the mate, highlighting that frontal courtship positively influence the female mating decisions. This study increases knowledge on sexual behavior in a key parasitoid of aphids, highlighting the importance of wing fanning among the range of sensory modalities used in the sexual communication of L. testaceipes. From a practical point of view, this information is useful in L. testaceipes‐based biocontrol strategies, since it can help to establish parameters for quality checking of mass‐reared wasps over time.     

Competition-induced switch in young of the year perch,Perca fluviatilis: an experimental test of resource limitation

Synopis Reproductively developed male fathead minnows, Pimephales promelas, exhibited courtship behaviour in the presence of female conspecifies under laboratory conditions. Male courtship consisted of several distinctive and visually conspicuous behaviours directed toward females, including approach, display, and two contact behaviours, as well as leading behaviour from the female to a suitable spawning site. An ovulated condition in females was not necessary to generate male courtship behaviour; in fact, the amount of courtship exhibited by males may depend inversely on the readiness of females to spawn.     

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.     

Copulation behaviour of Glossina pallidipes (Diptera: Muscidae) outside and inside the female, with a discussion of genitalic evolution

If species-specific male genitalia are courtship devices under sexual selection by cryptic female choice, then species-specific aspects of the morphology and behaviour of male genitalia should often function to stimulate the female during copulation. The morphology and behaviour of the complex, species-specific male genitalia of the tsetse fly, Glossina pallidipes Austen, were determined from both direct observations and dissections of flash-frozen copulating pairs; we found that some male genitalic traits probably function to stimulate the female, while others function to restrain her. The male clamps the ventral surface of the female's abdomen tightly with his powerful cerci. Clamping does not always result in intromission. Clamping bends the female's body wall and her internal reproductive tract sharply, posteriorly and dorsally, and pinches them tightly. The male performed sustained, complex, stereotyped, rhythmic squeezing movements with his cerci that were not necessary to mechanically restrain the female and appeared instead to have a stimulatory function. Six different groups of modified setae on and near the male's genitalia rub directly against particular sites on the female during squeezing. The designs of these setae correlate with the force with which they press on the female and the probable sensitivity of the female surfaces that they contact. As expected under the hypothesis that these structures are under sexual selection by female choice, several traits suspected to have stimulatory functions have diverged in G. pallidipes and its close relative, G. longipalpis. Additional male non-genitalic behaviour during copulation, redescribed more precisely than in previous publications, is also likely to have a courtship function. The elaborate copulatory courtship behaviour and male genitalia may provide the stimuli that previous studies showed to induce female ovulation and resistance to remating.     

Pheromones and Courtship Behavior in Butterflies

The courtship of butterflies begins with a general attractionof males to moving objects. If the object of the attractionis a receptive female, her response wil1 elicit further courtshipbehavior from the male. Male courtship pheromones are disseminatedduring the aerial phases of courtship and after the female haslanded. While female butterflies generally lack specific courtshippheromone glands, females of some groups do possess these. Courtship pheromones of males of four members of the subfamilyDanainae in the family Nymphalidae have now been identified.Three of these share the same active component, but each hasspecific additional components. Although the courtship pheromonesecretions of butterflies have odors which are very distinctiveto humans, the active compound of these pheromones is odorlessto humans and elicits responses from antennal receptors of allbutterflies tested. The pheromone receptors are short thin-walledpegs on the antennae of both male and female butterflies.     

Courtship behavior of the mosquitoSabethes cyaneus (Diptera: Culicidae)

Unlike any other mosquito reported, Sabethes cyaneus(Fabricius) displays an elaborate courtship before and during copulation. A male approaches a female suspended from a horizontal stick, suspends himself in front of her as he grasps her folded wings, and proceeds with a series of discrete stereotyped behaviors that involve proboscis vibration and movement of iridescent blue paddles on his midlegs. The sequence of these behaviors is as follows: freeleg waving, swinging, copulation attempt, superficial coupling, waving, genital shift, waggling, and release. Insemination occurs after genital shift. The only overt reciprocation by the female is abdomen lowering during the male's swinging. Courtship is often unsuccessful, and males are usually rejected during freeleg waving. The relation between male performance and mating success remains obscure.     

Courtship herding in the fiddler crab <Emphasis Type="Italic">Uca elegans</Emphasis>

Male and female animals are not always complicit during reproduction, giving rise to coercion. One example of a system that is assumed to involve sexual coercion is the mate herding behaviour of fiddler crabs: males push females towards the home burrow with the goal of forcing copulation at the burrow entrance. We recorded and analysed in detail the courtship behaviour of a North Australian species of fiddler crab Uca elegans. Courtship was composed of four main phases: broadcast waving, outward run, herding and at burrow display. During interactions males produced claw-waving displays which were directed posteriorly towards the female and which varied in timing and structure depending on the courtship phase. We suggest that courtship herding in U. elegans is driven primarily by mate choice for the following reasons, (1) females can evade herding, (2) no other reproductive strategies were observed, (3) males broadcast their presence and accompany courtship with conspicuous claw waves, and (4) the behaviour ends with the female leading the male into the home burrow. As an alternative function for herding in U. elegans we suggest that the behaviour represents a form of courtship guiding, in which males direct complicit females to the correct home burrow.