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PubMed1997

Correlates versus predictors of courtship success: courtship song in Drosophila silvestris and D. heteroneura

Males of the picture-winged flies Drosophila silvestris and D. heteroneura produce a sound during courtship wing vibration; successful courtships include more time spent in wing vibration. We clipped male wings to test whether this sound is a necessary signal for the success of male courtship. We tested females in no-choice tests and also in choice tests in which each female was provided with a clipped and normal male simultaneously. We found no differences in the probability of copulation for either species in no-choice tests, and no difference for the choice tests with D. heteroneuraSurprisingly, clipped males were significantly more successful than control males in the choice tests with D. silvestrisThe latencies to copulate did not differ by treatment in any tests. Consequently, the sound produced by courtship wing vibration does not influence male mating success and thus does not meet the most important criterion for being a sexual signal. We propose that the vibration of thoracic muscles, transmitted through the substrate or through the male's forelegs, could provide a sexual signal in these species.1997The Association for the Study of Animal Behaviour

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PubMed1997

Social mating system affects the frequency of extra-pair paternity in house wrens

We tested two hypotheses regarding the effect of the social mating system on extra-pair paternity in an Illinois population of house wrens, Troglodytes aedonAccording to the genetic-quality hypothesis, polygynous males are of higher quality than monogamists, and monogamously paired females, in an attempt to obtain high-quality genes, should have a greater proportion of extra-pair nestlings in their nests than polygynously paired females. According to the trade-off hypothesis, polygynists, with temporally overlapping nests, will have a greater proportion of extra-pair nestlings in their nests than monogamists, because polygynists have difficulty guarding one or both of their social mates. DNA fingerprinting revealed that extra-pair paternity was most frequent in secondary nests of polygynists. The proportion of secondary broods with extra-pair nestlings increased with the temporal overlap of polygynists' nests, although this trend was not significant. Both results are consistent with the trade-off hypothesis but not with the genetic-quality hypothesis. We did not address the effects of genetic quality on male success at siring nestlings in the nests of other males. Although the trade-off hypothesis focuses on male mate guarding, female behaviours may also affect frequencies of extra-pair paternity. Secondary females may compensate for reduced male defence by engaging in extra-pair copulations with neighbours to reduce the likelihood that neighbours destroy their nests. Thus, in house wrens, female participation in extra-pair copulations in combination with male mate-guarding constraints may generate higher levels of extra-pair paternity in secondary broods than in primary polygynous or monogamous broods.Copyright 1997 The Association for the Study of Animal Behaviour1997The Association for the Study of Animal Behaviour

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PubMed1998

A test of how well the repeatability of courtship predicts its heritability.

We compared repeatability and heritability estimates for courtship traits in the housefly Musca domestica L., to test the expectation that the repeatability of a behaviour is an upper-bound estimate of its heritability. The repeatabilities across and within mating partners for two courtship traits, male wing buzzing (Buzz) and female wing extension (Wing-out), were compared to heritabilities calculated from the divergence among subpopulations due to founder-flush events or artificial selection. The male's Buzz showed highly significant repeatabilities within and across female partners, but the values ranged widely (0.3-0.5). The intensity of Buzz decreased significantly with successive courtship attempts to the same female, apparently as a male strategy to optimize his efforts to gain copulation by modulating behaviour for specific females. Repeatabilities for female Wing-out within and across male partners were not significant, but this trait yielded significant heritabilities in two assessments (based on successful courtships). We attribute this contradiction to biased sampling because only the subset of reluctant females met the necessary restriction to refuse copulation throughout four successive courtships. These results demonstrate that repeatabilities are labile within individuals and may actually underestimate heritabilities when there is a genetic predisposition not to repeat performance. We concur with related studies that repeatabilities can identify evolutionary potential, but we advise caution in interpreting low values or comparing their absolute values.Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

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PubMed1998

Lifetime mating opportunities and male mating behaviour in sexually cannibalistic praying mantids.

I examined the number of lifetime mating opportunities and mating behaviour of males in two sexually cannibalistic species, the Mediterranean, Iris oratoria, and bordered, Stagmomantis limbata, praying mantids (Mantodea: Mantidae). Two approaches estimated the number of lifetime mating opportunities: direct observations of intersexual encounters in the field, and an encounter model. I collected behavioural observations, together with ecological data for use in the model, over three field seasons. The ecological data included an assessment of the feeding condition of S. limbata females in nature; the females fed at a level comparable to females maintained on an abundant diet in the laboratory. As for the number of mating opportunities, individual males of both species encountered two or more females, as predicted by the model. I observed no male, however, in more than one copulation. This result could reflect individual variation in the times and places of sexual activity or an actual low number of mating opportunities in the field. Furthermore, a higher percentage of I. oratoria males encountered two or more females than S. limbata males, as the model indicates. Fewer mating opportunities could lead to greater selection upon S. limbata males to ensure paternity at each mating, which can explain the longer copulation times observed for S. limbata males. I considered two hypotheses about male behaviour in light of the number of lifetime encounters with females: male suicide and male reduction of the risk of cannibalism. Behavioural observations do not strongly support male suicide in either species. Certain male behaviours, such as the nature of copulatory position and, in captivity, mounting females from the rear, are consistent with the idea that males behave so as to reduce the probability that they are cannibalized during intersexual encounters. Moreover, male I. oratoria preferentially mount well-fed, fecund females in captivity. Taken together, these results suggest that the prospect of multiple mating opportunities has influenced male mating behaviour over evolutionary time. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

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PubMed1998

Male and female behaviour and extra-pair paternity in the wheatear

Behavioural observations and DNA fingerprinting were used to determine the relationship between male and female behaviours and levels of extra-pair paternity in the wheatear, Oenanthe oenanthe. Behavioural observations were consistent with the hypothesis that males attempted to ensure paternity by mate guarding, while pursuing extra-pair copulations (EPCs) primarily outside the fertile period of their pair female. The intensity of guarding varied with time of season and was greater at late nests. However, although males on territories with late nests also experienced high intrusion rates, the intensity of guarding was influenced more by the operational sex ratio (which was female skewed at early nests) than by intrusion rates per se. We suggest that early breeding males adopted a strategy of territorial defence to ensure paternity, as opposed to guarding their female directly (which late breeding males did), to capitalize on the increased opportunities to pursue EPCs in neighbouring territories. Females were less conspicuous than males in their pursuit of EPCs, were never seen off territory or observed to solicit extra-pair males directly, and rejected the majority of EPCs. The frequency of extra-pair paternity was 11% of 73 offspring, in 29% of 17 broods, and was not correlated with the intensity of guarding. Female cooperation appeared to be important for successful copulation, and extra-pair paternity is therefore likely to be a consequence of solicited, or at least accepted, EPCs. We discuss why females might have participated in EPCs. Copyright 1998 The Association for the Study of Animal Behaviour Copyright 1998 The Association for the Study of Animal Behaviour.

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PubMed1998

Orienting asymmetries in rhesus monkeys: the effect of time-domain changes on acoustic perception.

Humans exhibit left-hemisphere dominance for processing spoken language, a species-specific acoustic signal characterized by a suite of spectro-temporal parameters. Some nonhuman primates (genus Macaca) also exhibit left-hemisphere dominance for processing their species-specific vocalizations, as evidenced by right-ear biases in orienting and reaction-time studies, and more damaging effects from left- than right-hemisphere lesions. Little, however, is known about the acoustic features underlying such biases. We conducted field playback experiments on adult rhesus monkeys, Macaca mulatta, to determine whether asymmetries in perception (measured as an orienting bias) are sensitive to changes in the temporal characteristics of their calls. If the observed right-ear bias for perceiving conspecific calls (Hauser & Andersson 1994, Proceedings of the National Academy of Sciences, U.S.A., 91, 3946-3948) depends upon particular acoustic parameters, then experimental manipulations beyond the species-typical range of signal variation will cause a change in perceptual asymmetry, either reversing the pattern (i.e. right to left ear ) or wiping it out (i.e. no asymmetry). We presented manipulated and unmanipulated exemplars of three pulsatile call types within the rhesus repertoire: an affiliative signal 'grunt', an alarm signal 'shrill bark', and a mating signal 'copulation scream'. Signal manipulations involved either (1) a reduction of the interpulse interval to zero or the population minimum or (2) an expansion of the interpulse interval to the population maximum, or two times the maximum. For the grunt and shrill bark, manipulations of interpulse interval outside the range of natural variation either eliminated the orienting bias or caused a shift from right- to left-ear bias. For the copulation scream, however, a right-ear bias was observed in response to all stimuli, manipulated and unmanipulated. Results show that for some call types within the repertoire, temporal properties such as interpulse interval provide significant information to listeners about whether the signal is from a conspecific or not. We interpret the orienting bias as evidence that hemispheric asymmetries underly this perceptual effect.Copyright 1998 The Association for the Study of Animal Behaviour.

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PubMed1998

Alternative reproductive strategies in the ruff, Philomachus pugnax: a mixed ESS?

In the ruff, there are two alternative male reproductive strategies. The majority of males of this lekking bird attempt to establish and defend territories on leks, and are referred to as Independents. Other males, referred to as Satellites, forego this behaviour and instead attempt to get access to the territories defended by Independents by acting submissively. The system is thought to be an example of a mixed evolutionarily stable strategy (ESS), where the two strategies have equal fitness payoffs and are maintained by negative frequency-dependent selection. Satellites visited leks at the same time as females, and were associated with territory-holding Independents which were successful in attracting females. This appeared to be an effect both of Satellites following females, and of females being attracted to Independents that dominated submissive Satellites. Males pursuing the two strategies benefited from the presence of each other, at least to some extent. In this study, Satellites got fewer copulations than expected by their proportion in the population. Satellites on leks might have increased longevity or reproductive life span, and gained copulations off leks and while migrating, to compensate for their low observed mating success on leks. The Satellite strategy may be a low-cost, low-benefit strategy, which may have equal average lifetime reproductive success as the territorial strategy Copyright 1998 The Association for the Study of Animal Behaviour

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PubMed1998

A role for pheromones in mate choice in a lekking sandfly.

There is increasing evidence to suggest that pheromone-mediated communication may play an important role in mate choice in many insects. We investigated the possible role of pheromones in mate choice in the lekking sandfly, Lutzomyia longipalpis, and explored whether males or females display any behavioural traits that predict patterns of male mating success. Pairs of virgin males were introduced into a small mating chamber and allowed to interact for approximately 10 min prior to the introduction of a single virgin female. The trial continued until the female copulated with one of the males. We recorded trials on video for later analysis. After a successful copulation, we determined the quantity of pheromone remaining in the glands of both males and measured the size of the pheromone glands and tergites. We corrected behaviours and measurements for body size (tergite width). Mated males had significantly more pheromone present in their glands after a trial than unmated males. Males differed in how much time they invested in wing fanning. Members of each pair of males spent similar amounts of time wing fanning but mated males and males with more residual pheromone spent more time wing fanning than their counterparts throughout the trial. The time spent wing fanning and the amount of residual pheromone were not correlated, however. A male's ability to win fights did not relate to his reproductive success. Finally, a female was equally likely to wing-fan directly at, or walk away from, either male in a given pair. The possible relationships between the amount of pheromone released and the residual pheromones left in the glands, and the level of male wing fanning and pheromone productivity, are discussed. Copyright 1998 The Association for the Study of Animal Behaviour.

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PubMed1998

Sexual selection as a side-effect of sexual conflict in the seaweed fly, Coelopa ursina (Diptera: Coelopidae).

The mating system of the seaweed fly involves a premating struggle. When mounted, females violently try to remove the male. In this study 48% of premating struggles resulted in successful rejection of the male, 46% in copulation and 6% were terminated by the male. Large males had a mating advantage. However, contrary to what would be predicted if this sexual selection occurred as a result of active female mate choice, we found a positive association between the duration of premating struggles and male size. A positive association was also found between the duration of premating struggles and male mating success, suggesting that large males may benefit through their superior ability to withstand female rejection. Large females rejected males more easily than small females, suggesting that the premating struggle has not evolved to allow mate assessment by females. We conclude that sexual selection is occurring as a side-effect of the female rejection response, which has probably evolved in order to avoid costs associated with copulation. Nevertheless, a sexual size dimorphism has evolved with males being larger and much more variable in size than females. (c) 1998 The Association for the Study of Animal Behaviour.

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PubMed1999

The effect of experimental male removals on extrapair paternity in the wheatear, Oenanthe oenanthe.

To examine the role of the pair male in ensuring paternity in the wheatear, we removed pair males for 24 h during the fertile period (days 0 to +1 in 1992 and days -5 to -1 in 1993, where day 0=first egg date). Control males were removed during incubation. The frequency and duration of intrusions, and the frequency of extrapair copulations (EPCs), increased during removals in the fertile period, but not during incubation. The frequency of extrapair paternity (EPP) was marginally higher (25%) in experimental than control broods and the presence of the pair male seemed particularly important in ensuring paternity in those days immediately preceding laying. Females were selective over which males they copulated with in the absence of their mate and rejected the majority of attempted EPCs. Only extrapair males in better body condition than the removed pair male eventually gained successful matings. Experimental males in poorer body condition were also more likely to have EPP in their brood than experimental males in better body condition. Female copulatory behaviour was the most important factor in determining patterns of paternity, and mate-guarding behaviours by the pair male appeared to limit the females' opportunities to engage in EPCs by reducing the frequency and duration of intrusions, rather than to control female behaviour directly. Copyright 1999 The Association for the Study of Animal Behaviour.

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PubMed1999

Sexual selection resulting from extrapair paternity in collared flycatchers.

Extrapair paternity has been suggested to represent a potentially important source of sexual selection on male secondary sexual characters, particularly in birds with predominantly socially monogamous mating systems. However, relatively few studies have demonstrated sexual selection within single species by this mechanism, and there have been few attempts to assess the importance of extrapair paternity in relation to other mechanisms of sexual selection. We report estimates of sexual selection gradients on male secondary sexual plumage characters resulting from extrapair paternity in the collared flycatcher Ficedula albicollis, and compare the importance of this form of sexual selection with that resulting from variation in mate fecundity. Microsatellite genotyping revealed that 15% of nestlings, distributed nonrandomly among 33% of broods (N=79), were the result of extrapair copulations. Multivariate selection analyses revealed significant positive directional sexual selection on two uncorrelated secondary sexual characters in males (forehead and wing patch size) when fledgling number was used as the measure of fitness. When number of offspring recruiting to the breeding population was used as the measure of male fitness, selection on these traits appeared to be directional and stabilizing, respectively. Pairwise comparisons of cuckolded and cuckolding males revealed that males that sired young through extrapair copulations had wider forehead patches, and were paired to females that bred earlier, than the males that they cuckolded. Path analysis was used to partition selection on these traits into pathways via mate fecundity and sperm competition, and suggested that the sperm competition pathway accounted for between 64 and 90% of the total sexual selection via the two paths. The selection revealed in these analyses is relatively weak in comparison with many other measures of selection in natural populations. We offer some explanations for the relatively weak selection detected. Copyright 1999 The Association for the Study of Animal Behaviour.

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PubMed1999

Mate choice copying versus preference for actively displaying males by female pied flycatchers.

Should a female copy the mate choice of other females? A female may rank a mated male higher in quality than an unmated male because the former has demonstrated that he is able to attract a female. However, a prospecting female may also avoid a mated male because of the risk that she has to compete with the male's initial mate over access to copulations, breeding resources and male parental care. We studied the mate choice of female pied flycatchers Ficedula hypoleuca, in aviaries divided into three compartments, two for males, and one for a female. A female was allowed to choose (build a nest in a nestbox) between two males after a period when she had been kept in a cage behind a one-way mirror and had presumably seen that one of the males was in the company of another female for 5 h. There was no evidence that females copied the mate choice of conspecific females, or that they avoided males that had recently been in the company of another female. Instead, females apparently chose a mate independently of others, choosing the male showing most courtship display. The latter result explains the consistency in mate choice observed when different females chose between the same pair of males in repeat trials. The ability of females to pick out the same male independently may also explain why a few males obtain most copulations in lekking species. Copyright 1999 The Association for the Study of Animal Behaviour.

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PubMed1999

Affiliative preferences are stable and predict mate choices in both sexes of Japanese quail, Coturnix japonica.

Most laboratory studies of mate choice use affiliation as a surrogate measure for actual choice of a partner for copulation. In a series of recent studies using affiliation as a dependent variable, we showed that the tendency of both male and female Japanese quail to affiliate with members of the opposite sex is profoundly affected by seeing them mate with others: female quails show an increased tendency to affiliate with males that they have seen mating; and male quails show a decreased tendency to affiliate with females that they have seen mating. In the present experiments, we examined: (1) the consistency of affiliative preferences of male and female quail for individuals of both the same and opposite sex; and (2) the validity of measures of affiliation as predictors of choice of an individual with whom to mate. We found that: (1) both male and female quail display consistency from trial to trial in their affiliative preferences for members of the opposite but not the same sex, and (2) affiliative preference is a powerful predictor of choice of a partner for copulation in both male and female quail. In Japanese quail, affiliative behaviour is consistent over time and predicts mate choice. Copyright 1999 The Association for the Study of Animal Behaviour.

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PubMed2000

Female mate guarding: no evidence in a socially monogamous species.

To understand the behavioural aspects of sperm competition, the costs and benefits to both sexes should be considered. However, few studies have addressed the costs to females of their social mate engaging in extrapair copulations (EPCs). Measures of female mate guarding have concentrated on female solicitation and copulation; however, females may also control access to their mate by maintaining close proximity, as is common in males. I recorded the maintenance of pair proximity behaviour of an urban population of the socially monogamous European blackbird, Turdus merula, over three breeding seasons. There was no evidence that females guarded their mates to prevent them from engaging in EPCs, nor were there any effects of the potential quality indicators of age, body size or male bill colour on the intensity of mate guarding between individuals. The study adds to a small body of literature suggesting that female mate guarding may be found in (facultatively) polygynous species, but not in socially monogamous ones. Copyright 2000 The Association for the Study of Animal Behaviour.

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PubMed2000

Courtship role reversal and deceptive signals in the long-tailed dance fly, Rhamphomyia longicauda.

We examined the function of secondary sexual characters in the role-reversed, lekking behaviour of female long-tailed dance flies, Rhamphomyia longicauda Loew (Empididae), to test the hypothesis that the degree of abdominal distention is an honest female signal about the state of egg development. Female Rhamphomyia cannot hunt for prey and they receive all of their protein from males by exchanging copulations for nuptial prey gifts. Females compete for male gifts within leks that are organized for a brief period each evening before dark. Before hovering within leks, females swallow air, inflating expandable pouches on the pleural margins of the abdomen. The result is a large saucer-like abdomen which is further exaggerated by wrapping scaled pro-, meso- and metathoracic legs along its pleural margins. Male preference for an enlarged abdomen was confirmed by suspending plastic models of varying size from monofilament lines and recording which models attracted the most males. There was a positive relationship between egg development and abdominal distention in a related species, R. sociabilis (Williston), which lacks inflatable abdominal pouches. Multiple regression showed that in R. longicauda, abdominal inflation completely masks the state of egg development. We conclude that female R. longicauda deceive mate-seeking males with the unreliable message that eggs are nearing maturation in order to obtain a protein meal in exchange for copulation. Males that fail to identify a female bearing mature eggs risk near-certain cuckoldry and an increased probability that the female will die before oviposition. Copyright 2000 The Association for the Study of Animal Behaviour.

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PubMed2000

Female-female aggression and female mate choice on black grouse leks.

We studied female-female aggression in relation to female mate choice in black grouse, Tetrao tetrix, in central Finland, in 1994-1998. Aggression occurred on average every other minute when there was more than one female on a territory, and aggressive behaviour was most prominent when several females attended the lek. Interactions tended to be proportionally most frequent on the territories of the highest-ranking males, although not significantly so. Females that were chased by other females did not mate with lower-ranking males than their aggressors did. Furthermore, chased females were only rarely (6% of cases) forced to move off the territory by agonistic interactions and copulations were disrupted by other females even less often (3% of cases). The choice of a mating territory did not depend on the outcome of aggression even though the aggressors were more likely to mate on the territory where aggression occurred than elsewhere. There was a marginally significant tendency for aggressors to mate earlier in the season. Females placed themselves further away from other females on the territory when soliciting a copulation than just before aggression. Our results suggest that aggression between females does not effectively constrain female choice in black grouse. Its function may be to aid females to secure undisturbed mating opportunities for themselves rather than to prevent others from mating with a particular male. Copyright 2000 The Association for the Study of Animal Behaviour.

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PubMed2000

The evolution of postcopulatory displays in dabbling ducks (Anatini): a phylogenetic perspective.

Although postcopulatory displays are widespread in animals, little is known about their function or the intended receiver. The postcopulatory displays of dabbling ducks are among the best described for any animal group. We documented the presence of initial and additional postcopulatory displays in nearly all dabbling duck species. We then reconstructed the evolution of postcopulatory displays in dabbling ducks using a phylogeny derived from mitochondrial DNA sequences. The display immediately following copulation (the initial display) is highly stereotyped in most species and shows extreme phylogenetic conservation. In contrast, the performance of additional displays is less stereotyped and less phylogenetically conservative. We review the possible functions of postcopulatory displays. Using evidence from display orientation, display form and phylogenetic reconstruction, we suggest that the most likely functions of postcopulatory displays in dabbling ducks are pair bond maintenance, individual identification, or signalling a successful copulation. Copyright 2000 The Association for the Study of Animal Behaviour.

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PubMed2000

Territoriality as a paternity guard in the European robin, Erithacus rubecula.

To investigate the relative importance of paternity defences in the European robin we used behavioural observations, simulated intrusions and temporary male removal experiments. Given that paired males did not increase their mate attendance, copulation rate or territory size during the female's fertile period, the most frequently quoted paternity assurance strategies in birds were absent. However, males with fertile females sang and patrolled their territories more regularly, suggesting that territorial motivation and vigilance were elevated when the risk of cuckoldry was greatest. In addition, there was a significant effect of breeding period on response to simulated intrusions: residents approached and attacked freeze-dried mounts more readily in the fertile period. During 90-min removals of the pair male in the fertile period, neighbours trespassed more frequently relative to prefertile and fertile period controls and appeared to seek copulations with unattended females. When replaced on their territories, males immediately increased both song rate and patrolling rate in comparison with controls. We propose that male robins sing to signal their presence, and increase their territorial vigilance and aggression in the fertile period to protect paternity. Copyright 2000 The Association for the Study of Animal Behaviour.

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