Calling Behaviour Influences Mating Success in Male Painted Reed Frogs,Hyperolius marmoratus

A detailed study was made of the natural calling behaviour of male painted reed frogs in the wild. Comprehensive minute by minute records of the vocal activity of males calling simultaneously in the breeding chorus were generated over a 19-night period using a multi-channel event recorder. With few exceptions, the period over which males called either spanned or extended beyond the time window when females sought mates in the chorus. Males varied little in calling persistence and the majority produced advertisement calls during each minute of the calling period. Aggressive interactions between males curtailed advertisement call production but fights were short-lived, seldom repeated and limited to a small proportion (12.6%) of males. Call rate was independent of body size and varied widely among males active in the chorus at the same time and under the same environmental conditions. The relative call rate ranks of individuals were sustained over time. The call rates of mated males were high in relation to the rates of other males monitored in the chorus at, and prior to, the time of mating. Females usually preferred the higher of two call rate alternatives presented in a range of two-choice phonotaxis experiments.     

Fighting for females in the toad Bufo calamita is affected by the operational sex ratio

Fighting between male natterjack toads, Bufo calamita, for access to females was studied in relation to both the influence of male and female density each night and the operational sex ratio. Fighting males were larger than other males and were present in the breeding area on more nights but their average mating success was the same. Fighting between males was associated with not strongly male-biased operational sex ratios. The density of breeding individuals, however, was irrelevant. Take-over rate was very low with an insignificant effect on mating success.     

Opportunity for sexual selection arises at moderate densities in the frog Rana chensinensis: an experimental approach

Heavily male-biased operational sex ratio and high male density characterizing explosively breeding anurans are expected to enhance the opportunity for sexual selection through large-male mating advantage brought about by severe male–male competition. However, our previous study on the temperate frog Rana chensinensis suggested that these two factors might restrict the opportunity under natural conditions for females to be grasped by any male they meet first, and scramble competition between males occurred less frequently because amplectant pairs hid on the bottom of ponds. In the present experimental study, in which the operational sex ratios and male densities were much lower and individual interactions were sufficiently longer than in natural population, we found a large-male pairing advantage in female mate choice, multiple mate choice, and male displacement experiments. These findings lend circumstantial support for the predication, although there was no evidence of large-female mating advantage in a male mate choice experiment.     

Individual Male Calling Pattern and Male Mating Success in the European Treefrog (Hyla arborea): Is there Evidence for Directional or Stabilizing Selection on Male Calling Behaviour?

In anurans, call properties are commonly classified based on within‐male variability as being either static or dynamic. Numerous playback experiments in the laboratory have indicated that female preferences based on dynamic call properties are usually strongly directional, while female preferences based on static call properties are often stabilizing or weakly directional. However, there are only few studies demonstrating that female preferences for high values of dynamic call properties indeed exert directional selection on male calling behaviour in natural populations. Moreover, field studies investigating whether female preferences for values of static call properties around the mean of the population lead to currently operating stabilizing selection on male calling patterns in natural populations are completely lacking. Here I investigate for two consecutive breeding seasons male calling patterns and male mating success in a population of individually marked European treefrogs (Hyla arborea), a hylid frog with prolonged breeding season and a lek mating system. Individual male calling pattern as analysed in terms of seven temporal and spectral call properties did not differ between males that survived from one breeding season to the next and those not surviving. None of the seven call properties investigated differed significantly between mated and unmated males, indicating that there is no strong directional selection on male calling behaviour in the study population. However, in one study season males that produced calls with a number of pulses around the mean of the population were significantly more likely to obtain matings than males that produced calls with a number of pulses at the low or high end of the distribution. Thus, this study provides preliminary evidence for the operation of stabilizing selection on a static call property (i.e. the number of pulses per call) in a natural population of an anuran amphibian.     

Limited male dispersal in a social spider with extreme inbreeding

Cooperatively breeding animals commonly avoid incestuous mating through pre-mating dispersal. However, a few group-living organisms, including the social spiders, have low pre-mating dispersal, intra-colony mating, and inbreeding. This results in limited gene flow among colonies and sub-structured populations. The social spiders also exhibit female-biased sex ratios because survival benefits to large colonies favour high group productivity, which selects against 1 : 1 sex ratios. Although propagule dispersal of mated females may occasionally bring about limited gene flow, little is known about the role of male dispersal. We assessed the extent of male movement between colonies in natural populations both experimentally and by studying colony sex ratios over the mating season. We show that males frequently move to neighbouring colonies, whereas only 4% of incipient nests were visited by dispersing males. Neighbouring colonies are genetically similar and movement within colony clusters does not contribute to gene flow. Post-mating sex ratio bias was high early in the mating season due to protandry, and also in colonies at the end of the season, suggesting that males remain in the colony when mated females have dispersed. Thus, male dispersal is unlikely to facilitate gene flow between different matrilineages. This is consistent with models of non-Fisherian group-level selection for the maintenance of female biased sex ratios, which predict the elimination of male dispersal.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society 2009, 97 , 227–234.     

Female choice of large males in the treefrog Hyla ebraccata

The mating behaviour and male mating success of Hyla ebraccata were examined over three study periods. Mated males were larger than unmated males on a significant number of nights and for one of the three study periods. In field observations of pair formation, female behaviour was consistent with choice of large males: females moved freely through the chorus, remaining within 10 cm of males larger than the nightly mean, before the male initiated amplexus. In 27% (n = 3) of these observations, males chased and fought over the female. However, the females removed two of these three males from amplexus, suggesting that females can also exercise choice after amplexus. There was a significant negative correlation between male size and dominant frequency of the primary note, indicating that the male's advertisement call contained size-related information. Comparisons of the size of mated and unmated males suggest that two factors may have affected the degree to which female choice influenced male mating success. First, the distance between calling males may have limited the opportunity for females to express a mating preference. Secondly, an increase in mean and a decrease in the variance of male size in one of the three study periods also may have limited the ability of females to express a preference for large males.     

The effect of interspecific mating on sex ratios in the twospotted spider mite and the Banks grass mite (Acarina: Tetranychidae)

Mixed populations of the twospotted spider mite (TSM),Tetranychus urticae (Koch), and the Banks grass mite (BGM),Oligonychus pratensis (Banks), occur on corn and sorghum plants in late summer in the Great Plains. Interspecific matings between these arrhenotokous species occur readily in the laboratory but yield no female offspring. The effect of interspecific mating on female: male sex ratios was measured by examining the F₁ progeny of females that mated with both heterospecific and conspecific males in no-choice situations. TSM females that mated first with BGM males and then with TSM males produced a smaller percentage of female offspring than TSM females that mated only with TSM males (43.1±5.8 and 78.9±2.8% females, respectively). Similarly, BGM females mated with heterospecific males and then with conspecific males produced fewer female offspring than females mated only with BGM males (55.7±5.2 and 77.5±2.5%, respectively). Lower female: male sex ratios were produced also by BGM females that mated with TSM males after first mating with conspecifics (62.4±3.4%). In mixed populations containing males of both species, females also produced lower female: male sex ratios, but these ratios were not as low as expected based on mating propensities and progeny sex ratios observed in no-choice tests. These data suggest that interspecific mating may substantially reduce female fitness in both mite species by reducing the output of female offspring, but in mixed populations this effect is mitigated by unidentified behavioral mechanisms.     

Familiarity and flexible mating strategies of a solitary rodent, Dipodomys ingens

Because mating is a product of individual reproductive strategies that may vary with changing conditions, we predicted variable mating behaviour in an arid-adapted, territorial rodent, the giant kangaroo rat, Dipodomys ingens. We also predicted that familiarity would facilitate nonaggressive courtship and mating in this solitary rodent. Through direct observations in the field, we found that mating varied from exclusive to multiple partners. Where densities were low, and on nights when multiple females were in oestrus, each animal mated with one member of the opposite sex. In conditions where the operational sex ratio was skewed towards multiple males, males footdrummed and competed for females. Males were able to mate with one or two females in adjacent territories, and they successfully competed for these same females throughout the breeding season. Females that mated exclusively with one male had more pups emerge from the burrow compared with females that experienced male competition. Females allowed nearest neighbour males to enter their burrows, and they engaged in more nonaggressive contact with close neighbours than with other males. Paired encounters in the field showed less aggression towards neighbours than strangers. In laboratory tests, females were less aggressive towards and allowed more contact with familiar than unfamiliar males. These results show that D. ingens can alter mating strategies as conditions change. Familiarity is an important factor in nonaggressive interactions between males and females and may be important to mate preferences in females during reproduction. The less aggressive behaviour to neighbours than to strangers (‘dear enemy’ phenomenon) is consistent with other solitary animals that defend multipurpose territories. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

The adaptiveness of intense contact mate guarding by males of the emerald damselfly,Lestes sponsa (Odonata, Lestidae): The male’s perspective

We studied the mating system of the emerald damselflyLestes sponsa. All males showed intense contact mate guarding by holding the female in tendem during the entire oviposition period. Our findings support the predictions made by Alcock (1994) about the occurrence of intense mate guarding: (1) a high female receptivity after copulation, (2) a high male capacity to resist takeovers, (3) sperm precedence, (4) a high operational sex ratio, (5) a high male density, (6) high access by rivals to mated females, (7) low energy expenditure, (8) a low risk of guarding, and (9) a short interval between copula and oviposition. This indicates a positive cost-benefit balance for this behavior, at least in males. A comparison within the genusLestes suggests that the male-biased sex ratios and the ease with which mated females are detected have been strong selection pressures in the evolution of intense contact mate guarding.     

Mate selection in Litoria chloris and Litoria xanthomera: Females prefer smaller males

It is generally accepted that high quality males are those that succeed in male–male competition: in either aggression or rivalry to attract and be selected by females. Previous studies of amphibians have suggested that the main characters influencing male mating success include variation in call characteristics (e.g. call rate, call intensity), calling behaviour, body condition, age and chorus tenure. In the present paper, several of the characters influencing female mate choice (male body size, body condition, call rate, call frequency and chorus tenure) are investigated in two closely related, explosive breeding frog species Litoria chloris and Litoria xanthomera. Smaller males of both species are shown to be more successful than larger males and this success is attributed to the increased chorus tenure of smaller males in L. xanthomera. This increased chorus tenure was attributed to the lower total energy used per call by a small male calling at a higher frequency. Whether increased chorus tenure explains female mate choice in L. chloris is uncertain but is highly probable given the strong similarity between the two species in both ecology and call characteristics.     

The effects of sex ratio on sexual competition in the European lobster

During the breeding season an individual's access to mates may be affected by operational sex ratios, causing strong variation in mating success. We manipulated adult sex ratios of the European lobster, Homarus gammarus, to test the predictions of models that relate sexual competition to (1) the sex ratio, (2) the time that an individual is not available to mate and (3) 'collateral investment', whereby two males contribute to a single clutch. The model predictions proved to be relatively insensitive to collateral investment. Male-male competition predominated in the male-biased but not in the female-biased sex ratio. This matches the predictions of one model that incorporates an extended period of female receptivity because the time that a male was unavailable to mate was small compared to the time spent by females in cohabitation and parental care. Although females increased their competitiveness when males were in the minority, male competition remained high. The insensitivity of male-male competition to sex ratios may be due to an upper limit to the costs that males can afford when there is a serious risk of injury, preventing males from increasing their aggression when females are in short supply. Copyright 1999 The Association for the Study of Animal Behaviour.     

Influence of male mating history on female reproductive success among monandrous Naryciinae (Lepidoptera: Psychidae)

1. Multiple male copulations can have detrimental effects on female fitness due to sperm limitation. 2. Monandrous Naryciinae females are immobile while the males are short‐lived and do not feed. Multiple male mating is therefore expected to lead to sperm limitation in females. Sperm limitation and male limitation are hypothesised as causes of the repeated evolution of parthenogenetic reproduction in the Psychidae. 3. In this study, the effects of multiple male mating on female reproduction are investigated in several species of Naryciinae by allowing males multiple copulations. The results for two species, Siederia listerella and Dahlica lichenella, are compared. The sex ratios of 53 natural populations are examined for indications of male limitation. 4. Previous copulations by the male increased the female's risk of remaining unfertilised. However, contrary to expectations, those unfertilised females were capable of successful re‐mating. 5. In S. listerella, the number of previous copulations of males negatively influenced female fitness. Females produced 30% fewer offspring if they mated with a previously mated male. In D. lichenella, the older the male and the lower its number of total lifetime copulations, the higher the female's reproductive success. 6. Only a fraction of the investigated populations had a female‐skewed sex ratio, but differences in development time between males and females could lead to reproductive asynchrony. 7. In conclusion, male mating history did not lead to strong sperm limitation in Naryciinae as had been suggested by their life history.     

Constrained sex allocation in a parasitoid due to variation in male quality

The theory of constrained sex allocation posits that when a fraction of females in a haplodiploid population go unmated and thus produce only male offspring, mated females will evolve to lay a female-biased sex ratio. I examined evidence for constrained sex ratio evolution in the parasitic hymenopteran Uscana semifumipennis. Mated females in the laboratory produced more female-biased sex ratios than the sex ratio of adults hatching from field-collected eggs, consistent with constrained sex allocation theory. However, the male with whom a female mated affected her offspring sex ratio, even when sperm was successfully transferred, suggesting that constrained sex ratios can occur even in populations where all females succeed in mating. A positive relationship between sex ratio and fecundity indicates that females may become sperm-limited. Variation among males occurred even at low fecundity, however, suggesting that other factors may also be involved. Further, a quantitative genetic experiment found significant additive genetic variance in the population for the sex ratio of offspring produced by females. This has only rarely been demonstrated in a natural population of parasitoids, but is a necessary condition for sex ratio evolution. Finally, matings with larger males produced more female-biased offspring sex-ratios, suggesting positive selection on male size. Because the great majority of parasitic hymenoptera are monandrous, the finding of natural variation among males in their capacity to fertilize offspring, even after mating successfully, suggests that females may often be constrained in the sex allocation by inadequate number or quality of sperm transferred.     

Asynchronous arrival pattern, operational sex ratio and occurrence of multiple paternities in a territorial breeding anuran, Rana dalmatina

Understanding why females mate multiply is a major issue in evolutionary ecology. We investigated the consequences of an asynchronous arrival pattern on male competition and multiple paternity in the apparently monoandrous agile frog ( Rana dalmatina ). The largest frogs arrived first and both males and females lost weight significantly during the spawning period. Asynchronous arrival at breeding sites resulted in a male-biased operational sex ratio (OSR). The OSR was more strongly male-biased at the beginning and at the end of the breeding period when the number of satellite males increased. All females mated only once, but multiple paternity within clutches occurred at the beginning and the end of the breeding period. The influence of asynchronous arrival and biased sex ratio suggests that reduced variance or bet-hedging promoting female fitness had only a reduced role in the evolution of polyandry, and polyandry is likely to be associated with male benefits. Polyandry in frogs can be explained either by forced mating as a result of sexual conflict or by clutch piracy. By modifying intrasexual competition, asynchronous arrival and changes in OSR may have a decisive influence upon the evolution of mating systems and favour both polyandry and stable coexistence of alternative mating behaviour.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 191–200.     

An ecological analysis of mating biology of Xylocopa virginica in southern Ontario

1. Xylocopa virginica virginica Linnaeus is a wide‐ranging species with plastic nesting behaviour that appears to represent an intermediary between solitary and social nesting species. Over 3 years, a natural population was studied with the objective of quantifying the relationship among population dynamics, climate, female nest provisioning behaviour, and male mating strategy. 2. Males in the population congregated around female‐occupied nesting sites before the beginning of nest provisioning by females; both resident and satellite male mating strategies were observed. Overall, the present results are consistent with female defence polygyny. 3. Male mating strategies were consistent across the three breeding seasons of our study, in spite of annual variation in population size, sex ratio, and weather. Male mating behaviour was also consistent with that seen in other populations with longer breeding seasons. 4. Adult non‐breeding females that never leave nests are observed in nests throughout the breeding season and we hypothesise that males continue to defend territories after breeding females have mated because of a small probability they can mate with one of these non‐breeding females. 5. These results are important to our understanding of the relationship between mating systems and the evolution of sociality, contributing data on the role of ecological factors to male mating behaviour. Collection of such data for a variety of species that differ in sociality is necessary for the comparative analysis that is required to fully elucidate coevolution of mating systems and sociality.     

Mating patterns of Minshan's toad (Bufo minshanicus) from three populations along an altitudinal gradient

The large-male mating advantage and size-assortative mating are two different size-based patterns, which deviate from random mating in toads. These two pairing patterns may arise due to female choice, male-male competition, male choice, or a combination of these. This study investigated the mating system of Minshan's toad (Bufo minshanicus) from three populations along an altitudinal gradient during two breeding reasons in the northeastern Tibetan plateau. Our study shows that males found in amplexus with females were larger on average than non-amplectant males in two sites with higher operational sex ratios. Similarly, in those sites, males and females found in amplexus maintained an optimal size ratio. These data suggest that male-male competition leads to size-assortative mating in the lack of mate choice (female and male mate choice) by Minshan's toad, as larger males performed higher frequencies for taking-over other low quality ones with amplectant females.     

Evolutionarily stable oviposition and sex ratio in parasitoid wasps with single‐sex broods

Abstract 1. The flexibility of hymenopteran sex ratios is well documented, particularly in structured populations featuring sib mating. 2. Using game theoretic models, the present study examines species producing single‐sex broods in which sib mating is unlikely, and focuses on the role of population density in determining evolutionarily stable oviposition strategies. 3. Since only mated females can produce offspring of both sexes while unmated females produce only male offspring, mated females are under selection to produce more females overall to balance the primary sex ratio. 4. As the proportion of all females that are mated should increase with density, offspring sex ratio of mated females is strongly linked to density at low to moderate densities. The present study shows that when density becomes low enough for fewer than half of all females to have mated, then female offspring generate higher fitness. 5. In this low density situation, females may gain a fitness benefit from waiting at their emergence site or from using other costly means to find and mate with males before ovipositing. 6. The predicted correspondence between females waiting at the emergence site and fewer than half of females in the population containing sperm, can be tested empirically, as can the somewhat counter‐intuitive prediction that greater access to males should yield a more male‐biased sex ratio in the offspring of mated females. 7. The present study also indicates how measuring the variance in giving up times by females waiting for males at low density, can provide insight into mechanisms determining waiting times.     

同种雄性竞争对手的存在对星豹蛛雄蛛求偶和交配行为的影响

越来越多的研究发现,雄性产生精子(精液)也需付出代价。雄性除了依据配偶质量和竞争对手的竞争强度适应性调整生殖投入外,雄性在求偶和交配行为上也相应产生适应性反应,求偶和交配行为具有可塑性。目前雄性求偶和交配行为可塑性研究主要集中于雌性多次交配的类群中,在雌性单次交配的类群中研究甚少。以雌蛛一生只交配一次而雄蛛可多次交配的星豹蛛为研究对象,比较:(1)前一雄性拖丝上信息物质对后续雄蛛求偶和交配行为的影响,(2)雌雄不同性比对雄蛛求偶和交配行为的影响。研究结果表明,星豹蛛前一雄蛛拖丝上的信息物质对后续雄蛛求偶潜伏期、求偶持续时间和交配持续时间都没有显著影响,但前一雄蛛拖丝上的信息物质对后续雄蛛求偶强度有显著抑制作用。同时,性比对星豹蛛雄蛛求偶和交配行为都没有显著影响。可见,星豹蛛雄蛛对同种雄性拖丝上的化学信息可产生求偶行为的适应性调整,而对性比不产生适应性反应。     

Turtle mating patterns buffer against disruptive effects of climate change

For organisms with temperature-dependent sex determination (TSD), skewed offspring sex ratios are common. However, climate warming poses the unique threat of producing extreme sex ratio biases that could ultimately lead to population extinctions. In marine turtles, highly female-skewed hatchling sex ratios already occur and predicted increases in global temperatures are expected to exacerbate this trend, unless species can adapt. However, it is not known whether offspring sex ratios persist into adulthood, or whether variation in male mating success intensifies the impact of a shortage of males on effective population size. Here, we use parentage analysis to show that in a rookery of the endangered green turtle (Chelonia mydas), despite an offspring sex ratio of 95 per cent females, there were at least 1.4 reproductive males to every breeding female. Our results suggest that male reproductive intervals may be shorter than the 2-4 years typical for females, and/or that males move between aggregations of receptive females, an inference supported by our satellite tracking, which shows that male turtles may visit multiple rookeries. We suggest that male mating patterns have the potential to buffer the disruptive effects of climate change on marine turtle populations, many of which are already seriously threatened.     

Mating Behavior and Reproductive Potential in the Turnip Moth Agrotis segetum (Lepidoptera: Noctuidae)

We investigated the lifetime mating potential and the reproductive behavior of male and female turnip moths Agrotis segetum (Schiff.) under field and laboratory conditions. The sex ratio was 1 : 1 in a lab-reared population as well as in two wild populations. Males were capable of mating repetitively a relatively large number of times (mean of 6.7 ± 2.7 matings) when given access to new virgin females throughout their lifetimes. Females seldom mated more than once (mean ± 1.3 ± 0.6 matings), indicating a male-biased operational sex ratio. The mean potential lifetime mating was five times higher in males, while the coefficient of variance was lower in males. There was no differences in longevity between animals that were allowed to mate and animals not allowed to mate, indicating no direct costs or benefits of mating in physiological terms. In males, the number of matings was positively correlated with longevity, but this was not the case in females. Nor was there a correlation between the number of female matings and the number of fertilized eggs. There was a negative correlation between the number of eggs fertilized and the number of times males had previously mated, indicating that male ejaculates were limited. Male spermatophore size also decreased with number of achieved matings. Laboratory-reared females attracted males in the field throughout their lifetimes, with a peak at 3–7 days of age. Wild males, allowed to choose between pairs of caged females in the field, were attracted in equal numbers to females of different ages. Females did not show any mate-rejection behavior in the field. They mated with the first male that courted them. No incidence of mate replacement by males arriving later to already courted females were recorded.