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.     

Reproductive behaviour of the white-tailed zygaenid moth,Elcysma westwoodii (Lepidoptera,Zygaenidae). II. Female mating strategy

Male white-tailed zygaenid moths.Elcysma westwoodii, often gather to a female to mate her. One of the factors that seems to cause male gathering is the females' tendency to refuse males. Field observations indicated that there was a relationship between the courtship duration, the copulation duration and egg fertilization: short courtship was followed by long copulation and oviposition of fertile eggs in the same day, while long courtship was followed by short copulation and failure of oviposition. This phenomenon may be the result of differences in male activity. More “fertile” males can reach copulation more quickly and continue it longer than males by limiting receptivity. This “passive female choice” seems to function in high male density. Long copulation usually continued until the female began oviposition.     

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 ofPhytoecia rufiventris Gautier (Coleoptera: Cerambycidae)

Sexual behavior between males and females, as well as between males, is described and discussed for the cerambycid beetlePhytoecia rufiventris. The beetles' taxis toward plants taller than average height brings the sexes together from a distance. A male may mount another individual (male or female) and attempt copulation without sex discrimination. The male can discern the sex of another individual only when the terminal part of his abdomen touches the ventral surface of the fifth visible sternite of the latter. No evidence of a sex pheromone is found in this species. Within 1.5–5.5 cm the substrateborne vibrations produced by a moving individual may be the important factor which elicits males to approach a moving individual and attempt copulation. If a female is receptive when a male touches her, he can copulate with her without any courtship display. However, if the female runs away and appears unreceptive, the male will perform courtship displays. Copulation is usually terminated by males. Homosexual behavior between males is discussed.     

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.     

Female control in yellow-legged gulls: trading paternity assurance for food

Females in many socially monogamous birds copulate hundreds of times more than necessary for fertilization, although little is known about the benefits of this excess. Females may not directly benefit from high copulation rates, but instead may exploit male interest in copulating to obtain benefits. In species with courtship feeding, females may trade copulations for food (immediate benefits hypothesis). I tested this hypothesis by analysing female behaviour during courtship in yellow-legged gulls, Larus cachinnans. Female gulls to some extent controlled sperm transfer, because they moved during copulation bouts, and this behaviour influenced the number of cloacal contacts per mounting that the male achieved. Female control was related to previous feeding by the male, and hence the male courtship feeding rate correlated with the cloacal contact rate. Males that give more food probably enhance their chances of fathering offspring. By analysing within-individual female behaviour, I also found that the number of cloacal contacts was higher when the male fed the female than when he did not, which indicates that female gulls followed a decision rule to resist copulation when food is not given. Overall, these results support the hypothesis that female gulls manipulate their mates to obtain food.     

Wing-flick Signals in the Courtship of the African Cave Cricket,Phaeophilacris spectrum

The male of the African cave cricket Phaeophilacris spectrum (Saltatoria: Grylloidea: Phalangopsidae) possesses tegmina without stridulatory organs, and both females and males lack tympanal organs. Therefore acoustic communication in the usual sense, which is typical for most crickets, is absent in this species. However, adaptations of the wing articulation allow the males to flick their wings forward over their heads. During courtship these movements are performed in series of 4–5 wing-flicks at a rate of 8–12 Hz (called wing-flick series = WFS) which elicit low-frequency air movements. Such signals may have taken over the functional significance of an acoustic ‘courtship song’. A quantitative analysis of the cricket's behaviour showed that in terms of frequency and total duration, ‘wing-flick series’ and ‘rocking’ behaviour are the most prominent signals of the male's courtship display. Moreover, analysis of the spatial relationships between male and female demonstrates that the wing-flicking is directed towards the female. The response of a female to a wing-flicking, courting male seems mainly to be calmed down and to reach a passive, receptive state, necessary for subsequent copulation. Females confronted with a male's courtship display exhibited a significantly higher threshold to disturbing stimuli than uncourted ones.     

Copulation Song in Drosophila: Do Females Sing to Change Male Ejaculate Allocation and Incite Postcopulatory Mate Choice?

Drosophila males sing a courtship song to achieve copulations with females. Females were recently found to sing a distinct song during copulation, which depends on male seminal fluid transfer and delays female remating. Here, it is hypothesized that female copulation song is a signal directed at the copulating male and changes ejaculate allocation. This may alter female remating and sperm usage, and thereby affect postcopulatory mate choice. Mechanisms of how female copulation song is elicited, how males respond to copulation song, and how remating is modulated, are considered. The potential adaptive value of female signaling during copulation is discussed with reference to vertebrate copulation calls and their proposed function in eliciting mate guarding. Female copulation song may be widespread within the Drosophila genus. This newly discovered behavior opens many interesting avenues for future research, including investigation of how sexually dimorphic neuronal circuits mediate communication between nervous system and reproductive organs.     

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.     

Temporal patterning of courtship behaviour in some parasitic Hymenoptera,with special reference to Melittobia acasta

In many parasitic hymenoptera copulation is preceded by elaborate courtship displays which include species-specific characteristics. Other features, shared by related species, may be used for defining higher taxa. The male's repertoire consists of movements involving the wings, legs, antennae, and mouthparts. These movements are performed continuously, or intermittently, depending on the species involved. The elements of a repertoire are repeated over and over again until the female indicates her readiness to copulate. Temporal patterning of various displays, and the timing of the female response are described. In Melittobia acasta (Walker) (Eulophidae) the male display is composed not only of repeating elements, but also includes new elements, introduced along the way; the display progresses towards a climactic finale. The timing of the female's copulation signal is accurately predictable. The morphology of Melittobia males is discussed in relation to this behaviour. Courtship of a related species, M.chalybii, is compared to the courtship of M.acasta.     

Short‐Term Nutritional Limitation Affects Mating Behaviour and Reproductive Output in Dwarf Spiders

Mating decisions can vary considerably depending on individual experience, mate availability and nutritional status. Here, we applied short‐term dietary restrictions to adult female spiders that were well fed during the juvenile stage in an effort to understand whether and how brief periods of food shortage can influence male and female mating decisions and mating behaviour. To assess whether responses vary between closely related species, we conducted the same experiment on the dwarf spiders Oedothorax retusus and O. apicatus. During courtship and mating, males of both species offer secretion to females from glandular tissue in their prosoma. Females were subject to food shortage over a period of 3 wks (‘low‐diet’ treatment, LD) or fed regularly (‘high‐diet’ treatment, HD). We compared courtship probability, mating probability/behaviour, and reproductive output between dietary groups and species. In both Oedothorax species, females in the LD treatment were less likely to mate and more aggressive towards males. Furthermore, LD females produced egg sacs that were significantly lighter than were those of the HD females. Effects of food deprivation on copulation duration, gustatory behaviour and oviposition latency differed between species. Our study shows that short periods of dietary restriction during the adult stage can strongly affect mating behaviour and reproductive output with differences between closely related species.     

The courtship behaviour of the pipefish Nerophis lumbriciformis: reflections of an adaptation to intertidal life

The courtship behaviour of Nerophis lumbriciformis (Pisces: Syngnathidae) consists of three distinct phases (initial courtship, spawning, and embrace) marked by prominent behavioural changes. The first courtship phase is characterised by female quivering. Courtship activity increases from low to high levels of quivering, causing undulatory head movements in the female. In the second phase, the female transfers her eggs onto the male incubating ventral surface. During the final phase, the male wraps his body around the female (embrace). Females actively initiate courtship indicating that these pipefishes are courtship role reversed. The complete lack of swimming and vertical movements in the courtship behaviour of N. lumbriciformis, unlike in the behaviour of other syngnathid species, suggests an adaptation to intertidal conditions.     

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.     

The biology of ant-like jumping spiders: intraspecific interactions of Myrmarachne lupata (Araneae, Salticidae)

Myrmarachne lupata is an ant-like salticid in which males have very large chelicerae. The display repertoire of this species is unusually large and complex for a salticid spider. Each individual male uses one of three different mating tactics depending on the female's maturity and location. With adult females outside nests type 1 courtship occurs which seems to be a form of visual communication and includes specialized movements and postures of the legs, palps and body. With adult females inside nests, males use type 2 courtship, which seems to be a form of non-visual communication and consists primarily of probing with the legs on the silk; males mate with receptive females inside the nests. With subadult females, males first use type 2 courtship then spin an adjacent silken chamber and cohabit. After she moults and matures, mating occurs inside the nest. Vacant nests of conspecific females, but not those of another sympatric salticid species, elicit courtship behaviour from males. During male-male interactions, embracing occurs with the large chelicerae spread apart. Females and subadults also display, and different displays occur in interactions depending on the sex/age classes of the spiders involved. Despite the unusual morphology of these spiders, their individual displays are similar to those of more typical salticids. During copulation males stand beside the female instead of over or on her as occurs with typical salticids.     

Male Performance and Body Size Affect Female Re-Mating Occurrence in the Orb-Web Spider Leucauge mariana (Araneae, Tetragnathidae)

Females can affect male probabilities of paternity success through behavioural, morphological and/or physiological processes occurring during or after copulation. These processes under female-control include the acceptance or rejection of mating attempts by subsequent males. Leucauge mariana is an orb weaving spider that shows male mate guarding of penultimate females, male–male competition on female webs and copulatory plugs, suggesting a polyandric mating system. The aim of the present study was to ascertain whether male behaviour during courtship and copulation in L. mariana relate with female re-mating decisions. Forty-three virgin females were exposed to up to three males until they mated. In 24 cases, the copulatory plug was absent after mating and females were exposed the next day to up to three other males. Eighteen females accepted a second mating. Relatively larger females were more receptive to second matings and were more likely to copulate if the second male was smaller. Longer duration of female tapping and abdominal bobbing during courtship, and first copulations with less short insertions and more flubs, were associated with increased female acceptance to second matings. The results indicate cryptic female choice on male courtship and copulatory performance and suggest female-control over the determination of male mating success in this spider species.     

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.     

Male singing behaviour and female presence in the territory in whitethroats Sylvia communis

Unmated male songbirds usually change their vocal behaviour when females enter their territories. Either the males court the females by changing the rate or pattern with which their normal long-ranging full songs are emitted, or they shift to special displays and long- or short-ranging vocalisations. In this study we quantified how female presence and behaviour affect the singing behaviour of male whitethroats. In the presence of a female the male frequently performed song flights, maybe to locate the female before it was courted, with sequences of diving-song displays. The courtship was interrupted by periods of perch songs. Female dscharp calls and short movements made the males initiate or resume courtship, whereas short horizontal jumps made the males intensify their courtship. Overall, the males changed their dual-function song activity in females' presence by emitting fewer perch songs and more flight songs. The quiet diving songs were only emitted during direct courtship of a female. The song types emitted immediately before, during, and after courtship are all highly variable, thus allowing for a quick assessment of the male's song repertoire. The courtship was also interrupted by periods of male woid calling, a call that is known to have a deterring effect on rival males. Bouts of woid calls were usually followed by song flights, again probably to locate the female that might have moved out of sight, or maybe to locate potential rival intruders. The latter was supported by an increased intrusion rate during female presence. Communicated by P. McGregor     

Time spent close to a sexual partner as a measure of female mate preference in a sex-role-reversed population of the blenny<Emphasis Type="Italic"> Salaria pavo</Emphasis> (Risso) (Pisces: Blenniidae)

In most species females do not exhibit conspicuous sexual behaviours and female mate preferences are often measured by means of the time spent close to males. In spite of its widespread use, in only a few studies has this measure been validated as a reliable indicator of female mate preference. Sex-role-reversed species offer a better opportunity to test female preferences, as females usually court males. We tested in a sex-role-reversed population of the blenny Salaria pavo (Risso) if the time spent by females close to males related to the number of courtship displays directed towards males, and whether these measures of female preference reliably predicted matings. Females were simultaneously presented with two males behind a glass partition and the time spent close to each male was measured. We then allowed females to have access to the males' compartments and measured the courtship behaviours performed by females towards each male and recorded with which of the males females spawned. Females spent more time close to the male that subsequently received more courtship displays, and the preferred male had a more developed head crest. However, both measures of female preference failed to predict matings. Females were often attacked by males and probably had, in some occasions, to spawn with the less preferred male. Females that spawned with the previously preferred male had more swollen bellies than females that spawned with the less preferred male, suggesting a male preference towards more ripe females. These results validate the use of "time spent close to a sexual partner" as a measure of female preference in S. pavo. We argue that matings may not always reliably indicate mate preferences, particularly in the less choosy sex.Communicated by R. Serrão Santos