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

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

Territory quality,male quality,courtship intrusions,and female nest-choice in the threespine stickleback, Gasterosteus aculeatus

In a recent study (Sargent & Gebler 1980), we allowed male sticklebacks to compete for a limited number of clay flowerpots as nest sites in circular wading pools. Although the males who nested in post enjoyed higher reproductive success (Sargent & Gebler 1980), a further analysis of these data revealed the males in these two classes did not differ significantly with regard to territory size, total courtships, and female rejections of the nest. In the present study, for the first half of the experiment, flowerpots were assigned randomly to half of the males nesting in wading pools; the pots were later taken from those males who first had them and were given to the remaining males for the second half of the experiment. Using paired comparisons, each male's performance was examined with and without a flowerpot. The males received more female nest-choices, and fewer intrusions from other males, when their nests were in flowerpots; however, the females appeared to engage in courtships with males independently of the males' nest site concealment. For the two classes of nest site concealment, the males were not significantly heterogeneous for the number of nest-choices that they received: thus male quality did not appear to directly affect female nest-choice. These results suggest that territory quality, in the form of nest site concealment, was the most important determinant of female nest-choice and male reproductive success in the experiments of Sargent & Gebler (1980). Females appear to minimize the risk of egg-cannibalism by spawning with those males who experience the least amount of courtship interference.     

Male Courtship Behavior and Weapon Trait as Indicators of Indirect Benefit in the Bean Bug,Riptortus pedestris

Females prefer male traits that are associated with direct and/or indirect benefits to themselves. Male–male competition also drives evolution of male traits that represent competitive ability. Because female choice and male–male competition rarely act independently, exploring how these two mechanisms interact is necessary for integrative understanding of the evolution of sexually selected traits. Here, we focused on direct and indirect benefits to females from male attractiveness, courtship, and weapon characters in the armed bug Riptortus pedestris. The males use their hind legs to fight other males over territory and perform courtship displays for successful copulation. Females of R. pedestris receive no direct benefit from mating with attractive males. On the other hand, we found that male attractiveness, courtship rate, and weapon size were significantly heritable and that male attractiveness had positive genetic covariances with both courtship rate and weapon traits. Thus, females obtain indirect benefits from mating with attractive males by producing sons with high courtship success rates and high competitive ability. Moreover, it is evident that courtship rate and hind leg length act as evaluative cues of female choice. Therefore, female mate choice and male–male competition may facilitate each other in R. pedestris. This is consistent with current basic concepts of sexual selection.     

Group mating among Norway rats II. The social dynamics of copulation: Competition,cooperation, and mate choice

The social interactions within groups of Norway rats (Rattus norvegicus) had a strong impact on the individual pattern of copulation which, in turn, affects sperm precedence and the probability of implantation in this species. Males alternated uninterrupted ejaculatory series, augmenting each others' copulatory investment. Females took turns mating after receiving an intromission, collectively potentiating the males' copulatory behaviour; increasing the number of oestrous females increased the number of intromissions and ejaculations achieved by each male but did not affect the amount of copulation experienced by each female. These turn-taking patterns within each sex provided the opportunity to change partners and permitted the emergence of different sex-typical patterns of copulation. Furthermore, the dominant male contributed more intromissions and tended to give each female more ejaculations than the subordinates did. Dominant males were also more likely to inhibit the subordinates' sperm transport. Females competed among themselves for the opportunity to mate with a male as he approached ejaculation and were likely to protect more of the dominant male's sperm transport than the subordinate male's.     

Sexual conflict over mating in Gnatocerus cornutus? Females prefer lovers not fighters

Female mate choice and male–male competition are the typical mechanisms of sexual selection. However, these two mechanisms do not always favour the same males. Furthermore, it has recently become clear that female choice can sometimes benefit males that reduce female fitness. So whether male–male competition and female choice favour the same or different males, and whether or not females benefit from mate choice, remain open questions. In the horned beetle, Gnatocerus cornutus, males have enlarged mandibles used to fight rivals, and larger mandibles provide a mating advantage when there is direct male–male competition for mates. However, it is not clear whether females prefer these highly competitive males. Here, we show that female choice targets male courtship rather than mandible size, and these two characters are not phenotypically or genetically correlated. Mating with attractive, highly courting males provided indirect benefits to females but only via the heritability of male attractiveness. However, mating with attractive males avoids the indirect costs to daughters that are generated by mating with competitive males. Our results suggest that male–male competition may constrain female mate choice, possibly reducing female fitness and generating sexual conflict over mating.     

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.     

Visual signals in the courtship of Drosophila melanogaster: Mutant analysis

Visual cues are necessary for optimal mating success in Drosophila melanogaster. The male's most important visually guided behaviour is tracking. It is shown here that tracking requires intact visual receptor cells R1–6 and the presence of screening pigments in the eye. Thus flies carrying the mutation ebony as well as wild type flies affected in receptor cell R1–6 are unable to use visual cues when they track females. A similar defect was obseved in white-eyed flies lacking screening pigments. Female receptivity depends on visual signals provided by the male flies. Most important cues are the light reflection from and the shape of the male's eyes. No influence of the light reflected from the thorax could be seen. Absence of eyes in the male, however, does not depress female receptivity as much as white eyes. Some evidence is provided that male courtship behaviour is evaluated visually by the female.     

Juvenile social experience affects pairing success at adulthood: congruence with the loser effect?

Social interactions with adults are often critical for the development of mating behaviours. However, the potential role of other primary social partners such as juvenile counterparts is rarely considered. Most interestingly, it is not known whether interactions with juvenile females improve males’ courtship and whether, similar to the winner and loser effects in a fighting context—outcome of these interactions shapes males’ behaviour in future encounters. We investigated the combined effects of male quality and juvenile social experience on pairing success at adulthood in zebra finches (Taeniopygia guttata). We manipulated brood size to alter male quality and then placed males in either same- or mixed-sex juvenile dyads until adulthood. We found that males from reduced broods obtained more copulations and males from mixed-sex dyads had more complete courtships. Furthermore, independent of their quality, males that failed to pair with juvenile females, but not juvenile males, had a lower pairing success at adulthood. Our study shows that negative social experience with peers during adolescence may be a potent determinant of pairing success that can override the effects of early environmental conditions on male attractiveness and thereby supports the occurrence of an analogous process to the loser effect in a mating context.     

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.     

Male displays and female preferences in the courtship of a gregarious cricket

The courtship of males of the gregarious cricket Amphiacusta maya involves a variety of signals. The quantitative aspects of both successful and unsuccessful courtship sequences were examined to determine whether certain aspects of male displays were correlated with female mating preferences regardless of which male performed them. Although variability among males was high for most courtship components measured, I found no evidence of female choice with respect to the courtship variables studied. About 35% of the duration of each male courtship sequence is devoted to chirping, but there were no differences in either the likelihood of copulation or the latency to copulation between normal males and experimentally silenced males. The possibility that intrinsic differences in male quality explained the variability in courtship duration was examined with a two-way analysis of variance. The variance in courtship duration was attributable to variance among females, not to variance among males. Thus the courtship behaviour of male A. maya is variable enough to allow females to exert stabilizing or disruptive selection on displays, but there is no evidence that females use the available information.     

Female Choice or Male Sex Drive? The Advantages of Male Body Size during Mating in Drosophila Melanogaster

The mating success of larger male Drosophila melanogaster in the laboratory and the wild has been traditionally been explained by female choice, even though the reasons are generally hard to reconcile. Female choice can explain this success by virtue of females taking less time to mate with preferred males, but so can the more aggressive or persistent courtships efforts of large males. Since mating is a negotiation between the two sexes, the behaviors of both are likely to interact and influence mating outcomes. Using a series of assays, we explored these negotiations by testing for the relative influence of male behaviors and its effect on influencing female courtship arousal threshold, which is the time taken for females to accept copulation. Our results show that large males indeed have higher copulation success compared to smaller males. Competition between two males or an increasing number of males had no influence on female sexual arousal threshold;—females therefore may have a relatively fixed ‘arousal threshold’ that must be reached before they are ready to mate, and larger males appear to be able to manipulate this threshold sooner. On the other hand, the females’ physiological and behavioral state drastically influences mating; once females have crossed the courtship arousal threshold they take less time to mate and mate indiscriminately with large and small males. Mating quicker with larger males may be misconstrued to be due to female choice; our results suggest that the mating advantage of larger males may be more a result of heightened male activity and relatively less of female choice. Body size per se may not be a trait under selection by female choice, but size likely amplifies male activity and signal outputs in courtship, allowing them to influence female arousal threshold faster.     

Multiple male traits interact: attractive bower decorations facilitate attractive behavioural displays in satin bowerbirds

Sexually selected male courtship displays often involve multiple behavioural and physical traits, but little is known about the function of different traits in mate choice. Here, we examine female courtship behaviours to learn how male traits interact to influence female mating decisions. In satin bowerbirds (Ptilonorhynchus violaceus), successful males give highly aggressive, intense behavioural displays without startling females. Males do this by modulating their displays in response to female crouching, which signals the display intensity that females will tolerate without being startled. Females typically visit multiple males for multiple courtships before choosing a mate, and females show differing tolerance for intense displays during their first courtship with each male. We test three hypotheses that may explain this: (i) familiarity with the courting male; (ii) the order of the courtship in mate-searching; and (iii) the attractiveness of the courting male. We found that females are more tolerant of intense displays during first courtships with attractive males; this increased female tolerance may allow attractive males to give higher intensity courtship displays that further enhance their attractiveness. We then examined why this is so, finding evidence that females are less likely to be startled by males with better physical displays (bower decorations), and this reduced startling then contributes to male courtship success. This role of physical displays in facilitating behavioural displays suggests a novel mechanism by which multiple physical and behavioural traits may influence female choice.     

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.     

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.     

Male–male competition for large nests as a determinant of male mating success in a Japanese stream goby, Rhinogobius sp. DA

Males of the stream goby Rhinogobius sp. DA (dark type) court females in deep pools and care for the eggs under stones in shallow riffles. We studied male–male competition for access to females and nest sites to understand how male size influences the mating success of this species. In field observations, larger males won in fighting with other males. However, large males did not tend to monopolize courtship opportunities, and the frequency of successful courtships, after which males led the females to the nests, was not related to male body size. The fact that courted females always escaped from the fighting sites once males began fighting likely explains why male size was not positively related to courtship success. Large males occupied large nest stones, and the number of eggs received in the nest was correlated positively with nest size. In aquarium experiments with two tiles of different sizes provided as nesting materials, males always chose the larger nest and, when two males were introduced simultaneously, the larger one occupied the larger nest. These results suggested that male mating success of this goby is determined by male–male competition for large nests rather than for access to females. Received: June 9, 2000 / Revised: September 2, 2000 / Accepted: October 4, 2000     

Courtship processing in Drosophila melanogaster. I. Selection for receptivity to wingless males

The first experiment was a selection procedure designed to increase the receptivity of Drosophila melanogaster females to males whose courtship had become less effective through attenuation of wing vibration. The clearest response was in the percentage of females receptive per generation, but changes were also detected in courtship duration and latency to courtship. The two selected and two control lines were hybridized in experiment 2, and the hybrid of the selected lines (Selx) was found to maintain the adaptation to selection of its parents. The relative receptivity of all female types was compared in experiment 3. The results demonstrated that adaptations had occurred in the females' courtship processing, rather than merely in the males' ability to court.     

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.     

Dominance is not always an honest signal of male quality,but females may be able to detect the dishonesty

Females prefer dominant males as mating partners in numerous species. Male dominance rank is considered as an honest signal of male quality, because only healthy males in good condition are thought to be able to win fights with other males. Here, we tested whether activation of the immune system influences the success of males in male–male competition and mating in the field cricket, Gryllus integer. We activated the immune system of males with a nylon monofilament (to mimic a parasitoid larva), and arranged fights between male pairs to assess male dominance and associated mating success. Activation of the immune system with nylon monofilament substantially enhanced the fighting success of males during male–male competition but had no effect on mating success. However, sham-manipulation (a wound only) did not have any effect on fighting success although females mated more often with dominant males. Our study suggests that when male crickets meet an apparent survival threat they may behave more dominantly, probably owing to terminal investment. Male success during male–male competition is not always an honest signal of males’ quality, but females may be able to detect this dishonesty.     

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.