Male position and calling effort together influence male attractiveness in leks of the medfly,Ceratitis capitata (Diptera: Tephritidae)

Despite the close association between lek mating systems and the study of female mate choice, male mating success in leks is often associated with other aspects of sexual selection as well as female choice of male display traits. Males of the medfly Ceratitis capitata form leks on the undersides of leaves of their host plants. By experimentally creating artificial leks, we show that male success at attracting females depends not only on male calling effort (pheromone production dispersed by wing movement), but also on the position of the male within a lek. Males in the highest position in the artificial lek (closest to the light) attracted more females, and received more visits from those females. In our experiment, we deliberately minimized the visual cues that females approaching a male could use and, under these conditions, found no associations between male attractiveness and male size, weight or fluctuating asymmetry, either of the wings or sex setae (a pair of bilateral supra‐fronto‐orbital bristles). The latter result contrasts with earlier studies showing a significant negative association between sex setae fluctuating asymmetry and mating success. Accordingly, we conclude that symmetry of the male sex setae has no role in nonvisual communication (e.g. through directing pheromone plumes). Mating patterns associated with this insect are therefore every bit as complex as those in vertebrate leks. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 479–487.     

Female alpine newts (Triturus alpestris) mate initially with males signalling fertility benefits

Gametic asymmetry implies that females invest more per gamete than males do and thus sperm is considered to be a relatively cheap resource. However, contrary to this classic view, sperm has been shown to be frequently in short supply; hence, selection favouring females that mate for fertility benefits should occur. For this reason, we determined whether males signalling fertility are preferred by female newts of the species Triturus alpestris . We performed paired female–male trials using unmated and previously inseminated females to determine potential criteria for female interest in a courting male, to establish what factors lead to successful mating and to assess the importance of female choice for direct and indirect benefits. We found that female interest in any potential mate and mating success decreased once mating had occurred. Furthermore, we detected an increase in spermatophore deposition rate and rapid spermatophore transfer in encounters that resulted in a successful mating. The results obtained indicate that female alpine newts are attracted to males showing signs of relatively high fertility and that females exhibit a decreased propensity to mate once initial sperm reserves have been acquired. Our results support the theory of initial female choice for fertility benefits.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 483–491.     

SIZE,SYMMETRY, AND SEXUAL SELECTION IN THE HOUSEFLY,MUSCA DOMESTICA

Relationships between measures of body size, asymmetry, courtship effort, and mating success were investigated in the housefly, Musca domestica (Diptera: Muscidae). A previous study indicated that both male and female flies with low fluctuating asymmetry enjoyed enhanced mating success. The aim of our investigations was to determine whether the greater success of symmetrical males is due to variation in male mating effort or to female choice and whether males exhibited mate choice. However, our study found directional rather than fluctuating asymmetry with both male and female flies having, on average, longer left wings than right. Also, asymmetry was not related to mating success in either sex. Rather, both males and females appeared to exhibit choice on the basis of the size of potential mates, with males preferring females with long bodies and females preferring heavy males. Possible benefits from choice of large mates are discussed. The initial mating strikes (in which the male leaps onto the back of the female) did not appear to be targeted according to female morphology, and their frequency did not vary according to male morphology. This indicates that mate choice by both sexes according to size probably occurs during the later stages of courtship, when the flies are in intimate contact. Possible reasons for the absence of choice according to asymmetry are discussed.     

Male courtship success and female discrimination in the mediterranean fruit fly (Diptera: Tephritidae)

The mating success of individually marked male Mediterranean fruit flies was monitored over 6 consecutive days in the laboratory. Mating frequency was nonrandom, as the numbers of both males that failed to mate and males that mated many (more than four) times were much higher than expected by chance alone. Differential mating success resulted in part from intermale variation in activity level. Male copulatory success was positively correlated with the numbers of courtships performed, attempted copulations (mountings), and females courted. Male-male aggression, on the other hand, and a negligible effect on male mating success. Female choice also appeared to influence male mating frequency. Fewer than 10% of courtships resulted in mating, and in most cases females terminated courtship by simply moving away from the male. Females that did mate generally selected males having higher copulatory scores than previously rejected males.     

Ectoparasitism in mothers causes higher positional fluctuating asymmetry in their sons: implications for sexual selection

Random deviations from perfect bilateral symmetry, fluctuating asymmetry, arise from developmental instability. I tested experimentally whether parasitism in female Drosophila nigrospiracula increases fluctuating asymmetry in male offspring. I also developed a novel measure for estimating developmental instability in a meristic trait called positional fluctuating asymmetry, which is based on the difference in the position of thoracic bristles between the right and left sternopleuron. I expected this measure to be a more sensitive indicator of developmental instability than the traditional numerical fluctuating asymmetry, because the latter is based on the difference in the actual presence or absence of bristles. Female flies burdened with hematophagous mites, Macrocheles subbadius (Macrochelidae), produced sons with significantly higher positional fluctuating asymmetry than did females carrying no mites. This effect, which may have resulted from impaired provisioning of oocytes by infested females, was dose dependent and magnified in the progeny of younger (18-20 d) versus older (30-32 d) females. This apparent magnification resulted from a slight but not significant increase in asymmetry of offspring of the older and unparasitized females. In contrast, the same mite loads had no effect on offspring numerical fluctuating asymmetry. If low-positional fluctuating asymmetry males enjoy a mating advantage, then with appropriate genetic variability, sexual selection could drive the evolution of host resistance in host populations. However, variability in neither kind of asymmetry influenced male mating success in nature. Thus, although male positional fluctuating asymmetry is causally associated with parasitism via maternal effects, asymmetry-based sexual selection is unlikely to influence the evolution of mite resistance in D. nigrospiracula. The value of the sensitivity afforded by positional fluctuating asymmetry is discussed in the context of sexual selection and conservation biology.     

Locomotor capacity and dominance in male lizards Lacerta monticola: a trade‐off between survival and reproductive success?

Trade-offs between reproduction and survival are important determinants of life-history characteristics of lizards. Organisms cannot increase the allocation of limited resources to reproduction without diverting a proportional amount of energy from another trait. Locomotor performance is an ecologically relevant trait that potentially influences survival by affecting the ability to escape from predators. Most studies have used female lizards as subjects because pregnancy is known to reduce their locomotor abilities, whereas little is known on costs of reproduction in males. In this study we suggest that in males of the lizard Lacerta monticola reproductive investment in morphological traits that confer dominance (i.e. head size) might lead to a low probability of survival by decreasing investment in other traits that affect locomotor performance (i.e. limb symmetry). We staged laboratory agonistic encounters between males and measured their morphology and burst speed on a race track to examine possible relationships between morphology, social dominance and locomotor capacity. Our results indicate that social dominance was positively related to relative head height, and that escape speed was negatively related to levels of fluctuating asymmetry in femur length, but also negatively related to relative head height. Males with greater relative head height also had more asymmetrical femurs, thus dominant males suffered a decrease in locomotor performance. Males with higher heads tend to dominate male–male interactions and hence may gain access to reproductive females, thus increasing their current reproduction success. However, this might occur at the expense of future survivorship mediated by a decrease in escape speed. Therefore, in male L. monticola there might be a trade-off between current reproductive success and survival.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002; 77 , 201–209.     

Fluctuating asymmetry and sexual selection in the Mediterranean fruitfly (Diptera, Tephritidae)

Sexual selection is known to operate at medfly leks with a few males gaining a high proportion of matings. However, specific male characteristics subject to sexual selection have not been identified. Here we report laboratory studies indicating that directional sexual selection operates on the level of fluctuating asymmetry (FA) in the superior frontal orbital setae (sex setae), with symmetrical males gaining more matings. Studies relating mating success with FA in a male trait have generally been taken as evidence of the operation of indirect sexual selection. For a male trait to acts as a reliable indicator of fitness, FA in the male trait should be negatively associated with trait size and females should mate with the males with the most exaggerated form of the trait. However no association was found between seta FA and mean seta length. In addition sexual selection did not appear to operate on mean trait size, although males with an intermediate sex seta to wing length ratio did achieve higher mating success, indicating that stabilizing sexual selection operates on this relative dimension. It is suggested that differences in male competitive ability may provide an alternative explanation of how such associations between mating success and FA in male characteristics can arise.     

Condition dependence of developmental stability in the sexually dimorphic fly Telostylinus angusticollis (Diptera: Neriidae)

Developmental stability is widely regarded as a condition‐dependent trait, but its relation to genotype and environment, and extent of developmental integration, remain contentious. In Telostylinus angusticollis, the dorsocentral bristles exhibit striking variation in developmental stability, manifested as fluctuating asymmetry (FA) in bristle position (‘positional FA’) and failure to develop some bristles (‘bristle loss’), in natural and laboratory populations. To determine whether this variation reflects condition, I tested for effects of genotype and environment (larval diet quality), and examined covariation with condition‐dependent traits. Positional FA was not affected by genotype or environment. However, positional FA covaried negatively with secondary sexual trait expression in males, and with sexual dimorphism in body shape, but covaried positively with body size in females. Bristle loss reflected both genotype and larval diet. Flies reared on poor‐quality diet exhibited a similar rate of bristle loss as wild flies. Both positional FA and bristle loss were greater in males. These results suggest that the relation between developmental stability and condition is complex and sex dependent.     

Securing paternity: mating plugs in the dwarf spider Oedothorax retusus (Araneae: Erigoninae)

One of the various male strategies to prevent or impede female remating is the production of a mating plug that covers the female genital opening or remains inside of the female genital tract after mating. Such structures have been described for many species in many animal taxa; however, in most cases, we know little or nothing about their specific adaptive value. Our investigations demonstrate that females of the dwarf spider species Oedothorax retusus (Westring, 1851) (Linyphiidae, Erigoninae) exhibit a substance on one or both of her paired genital openings only after copulation. We performed double-mating trials and forced the second male to mate into the previously used or unused spermathecal duct of the female by amputating one of his paired male gonopods (pedipalps). Furthermore, to investigate whether the duration of the first mating has an effect on the size and efficiency of the mating plug, we interrupted first matings after either 1 or 3 min, categorized plug size and recorded mating behaviour of subsequent males. The amount of secretion transferred was larger in long compared to short copulations. A long first copulation successfully prevented subsequent males from mating into the used ducts, whereas mating success after short first matings was similar to matings into unused copulatory ducts of the females. The present study demonstrates that a male O. retusus can prevent a rival from transferring sperm into the same spermatheca by applying a mating plug, but only if he mates for long enough.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 574–583.     

A test for negative frequency-dependent mating success as a function of male colour pattern in the bluefin killifish

Rare male mating advantage (a form of negative frequency dependence) is frequently proposed as a mechanism for the maintenance of genetic variation within populations. This hypothesis is attractive for systems with pronounced male colour polymorphism because it can maintain particularly high levels of variation. We tested for negative frequency-dependent mating success between yellow and red male colour patterns in bluefin killifish, Lucania goodei . Lucania goodei populations harbour substantial colour pattern polymorphism, and a large proportion of this variation has a genetic basis. We established outdoor mesocosms with red and yellow males in three different ratios: yellow rare (one yellow ♂ : five red ♂), even (three yellow ♂ : three red ♂), and red rare (five yellow ♂ : one red ♂). We obtained eggs and used microsatellites to determine paternity. By contrast to expectations, we found no support for a rare male mating advantage. Red males had slightly higher spawning success than yellow males, particularly in replicates with large clutches and when red males were rare. However, yellow males did not have higher mating success when rare. We discuss alternative mechanisms for the maintenance of the polymorphism as well as the potential reasons for the lack of a rare male mating advantage.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 489–500.     

Female morphology,web design,and the potential for multiple mating in Nephila clavipes: do fat‐bottomed girls make the spider world go round?

In animal species where females mate with multiple males, female mating success might be expected to covary with aspects of female morphology, such as size or shape. Spiders are especially interesting in this regard, as the females of several spider groups weave intricate webs that often accommodate multiple male spiders, all of whom are potential mates. Because web design is likely to be dependent on female size/shape, we use multivariate methods to assess the relationships among female morphology, web design, and reproductive ecology over a range of body sizes in the orb-weaving spider Nephila clavipes . Of the measured variables, only abdomen size explained a significant amount of the variation in number of males on a web, and this relationship holds even after statistically accounting for body size. Because abdomen size is an indicator of body condition in spiders, we suggest that condition is likely to be an important factor relating to potential mating success in female spiders. We found no evidence for an association between web design and number of males on a web, although our data indicate that larger females build webs that are both larger and further from the ground than those of smaller females.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 87 , 95−102.     

Signal components, acoustic preference functions and sexual selection in a cricket

In many sexually reproducing organisms, females choose mates based on multiple male traits. This study examined how two temporal components of the male mating call – chirp rate and chirp duration – affect female mating preference in five populations of a widely distributed North American cricket, Allonemobius socius (Orthoptera, Gryllidae). Chirp rate and chirp duration of the A. socius mating call were varied independently, and the responses of virgin females to these experimentally manipulated calls were repeatedly measured using a sequential single-stimulus design. Significant among- and within-population variation in chirp-duration preferences of females were found. Contrary to many previous studies, call chirp rate had no effect on female phonotaxis. Also there was no evidence of an interaction between chirp rate and chirp duration on female response to male mating calls. Moreover, female responsiveness to average and above-average chirp duration appeared to decline with female (adult) age. Overall, these results suggest evolved differences among populations in chirp-duration preferences, and that selection can act within populations on female chirp-duration preference.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 461–472.     

Mating distribution and its temporal dynamics affect operational sex ratio: a simulation study

The present study investigated how variation in mating distribution in time and among males influences the operational sex ratio (OSR) with a simulation inspired by paternally caring fish. Varying (1) the potential reproductive rate of each sex, (2) the mating distribution among males, and (3) the length of male mating phase, we created different mating patterns. In each case, we searched for the adult sex ratio that resulted in an OSR of 50% (where sex-roles switch). This approach enabled a comparison with a previous model. We found that the OSR was influenced by the distribution of matings in time and among males when the male mating phase was limited by a parental phase. Furthermore, the mating dynamics were shaped by the fact that the numbers of males and females and their capacities for collateral investment affected OSR immediately from the start of the reproductive season, whereas their times-out had a delayed effect on OSR.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 551–559.     

Seasonal variation in genital and body size, sperm displacement ability, female mating rate, and male harassment in two calopterygid damselflies (Odonata: Calopterygidae)

Sperm competition is a pervasive force. One adaptation is the male ability to displace the rivals' sperm that females have stored from previous copulations. In the damselfly, Calopteryx haemorrhoidalis asturica , males with wider aedeagi displace more spermathecal sperm. The present study documents that the same mechanism operates in another damselfly, Hetaerina americana . However, this genital width in both species decreases along the season, but late-emerging females have more sperm displaced than early-emerging females. Because territorial males mated more and were larger in body and genital size than nonterritorial males, late-season females mated with considerably larger males with respect to female size and this produced higher sperm displacement. Assuming female benefits from storing sperm but that such benefit does not prevail if males displace sperm, it is predicted that, along the season, females will mate less and male harassment (in terms of male mating attempts and oviposition duration) will increase. These predictions were corroborated. In H. americana , it was also tested whether spermathecal sperm became less viable along the season. The results obtained did not corroborate this. This is the first evidence indicating that season affects sperm displacement ability and female mating frequency due to changes in male body and genital size.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 815–829.     

Testing the role of coadapted genes versus bet-hedging for mating strategies in colour polymorphic pygmy grasshoppers

Recent investigations of mate choice indicate that the genetic effect of sires on offspring fitness may depend on the interaction between maternal and paternal genotypes and the environmental conditions experienced by the offspring. Alternative colour morphs of the pygmy grasshopper, Tetrix subulata , represent ecological strategies that differ in body size, life history, thermoregulatory behaviour, and habitat selection. The hypothesis that selection promotes behaviours maintaining coadapted gene complexes predicts individuals to mate assortatively with respect to colour morph. On the other hand, the bet-hedging hypothesis predicts that the temporal variability of the environment inhabited by these animals may select for disassortative mating behaviour resulting in heterogeneous offspring. To distinguish between these competing hypotheses, we investigated mating behaviours using dual-choice experiments. Our results were not in agreement with the prediction of assortative mating but suggest instead that matings were random with regard to colour morph. Polyandry was common, and females mated with the second male regardless of whether the first mating was assortative or disassortative. Polyandry also was equally frequent among females in triads in which the two males belonged to different colour morphs as in triads where both males belonged to the same colour morph. A field experiment confirmed that polyandry occurred also among free-ranging individuals, and uncovered variation in mating success among male colour morphs, probably due to indirect effects of coloration on activity or habitat use. The consequences of this random and polyandrous mating strategy for the evolutionary dynamics of the colour polymorphism remain to be explored.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 491–499.     

Testing the adaptive significance of sex‐specific mating tactics in collared lizards (Crotaphytus collaris)

We tested whether territorial defence is adaptive in male collared lizards by examining the extent to which territory owners monopolize females. We also tested whether females benefit by mating with multiple males using alternative tactics when local sex ratios varied. Surprisingly, neither the number of offspring that males sired, nor the number of females that males mated with varied as a consequence of highly variable local sex ratios. Moreover, both the number of offspring sired and the number of female mates were independent of male social status. Courtship frequency was under positive directional sexual selection for mating success for territorial males. None of the phenotypic traits that we examined were targets of sexual selection in nonterritorial males. Although offspring survivorship decreased with the degree of multiple mating, females mated multiply with similar numbers of territorial and nonterritorial males during all three seasons. Females did not obtain material or genetic benefits that balanced the apparent offspring survival cost imposed by mating with multiple males. Instead, females appeared to be ‘making the best of a bad job’, perhaps because the abundance of hiding places used by subordinate males makes it difficult for females to avoid male harassment. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 423–436.     

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.     

Seasonal monogamy and multiple paternity in a wild population of a territorial reptile (tuatara)

Investigating the mating system of a population provides insight into the evolution of reproductive patterns, and can inform conservation management of threatened or endangered species. Combining behavioural and genetic data is necessary to fully understand the mating system and factors affecting male reproductive success, yet behavioural data are often difficult to collect for threatened species. In the present study, we use behavioural data and paternity analyses to characterize the mating system of a high density population of a long-lived, ancient reptile (tuatara, Sphenodon punctatus ). We further investigate the phenotypic traits (including body size, body condition, tail length, and ectoparasite load) that affect male reproductive success. Our behavioural data reflect a seasonally monogamous system with low levels of polyandry and polygyny that are consistent with male mate guarding. Male reproduction is highly skewed (only 25–30% of males are successful), and body size is the primary predictor of male reproductive success. Based on the genetic data, multiple paternity was found in only 8% of clutches, and the results of the paternity analyses showed monandrous clutches from socially polyandrous females. Our behavioural and genetic results revealed complexities in female mating patterns that support the potential for cryptic female choice or sperm competition. This warrants further experimental investigation into the mechanisms underlying reptile fertilization and the disparities between social and genetic polyandry in wild populations.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 161–170.     

Is size assortative mating in Paracalliope fluviatilis (Crustacea: Amphipoda) explained by male–male competition or female choice?

Field and laboratory studies were used to assess: (1) whether size assortative mating occurred in the New Zealand amphipod Paracalliope fluviatilis and (2) hypotheses developed to explain size assortative mating. We found that assortative mating occurred and that larger females carried more eggs, suggesting they may be more valuable as mates. Laboratory experiments were then used to determine whether: (1) male size influenced the size of the female selected (mechanical constraints hypothesis); (2) male size influenced pairing success in the presence of competition (intrasexual selection hypothesis); (3) take‐overs of females occurred and whether large males were more successful (intrasexual selection hypothesis); (4) guard duration varied relative to male and female size (guard duration hypothesis); and (5) females had control over pairing success and guard duration (intersexual selection hypothesis). Although there was evidence to suggest the existence of intrasexual competition for mates (i.e. both small and large males preferred large females), there was no evidence of overt competition (i.e. takeovers of paired females). There was also no difference with respect to how long small and large males guarded females, but large females were guarded longer by both male size classes. Females handicapped by having their mobility reduced were guarded for the same duration as control females but males were more likely to pair with handicapped females, suggesting that they were easier to amplex. Given the lack of evidence for direct male–male competition or female choice, we suggest that assortative mating may be the result of: (1) indirect competition (e.g. in situ large males may be better able to access and amplex the largest females) or (2) female resistance to small males in combination with higher costs that small males may incur in securing large females. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 173–181.     

Variation in energy reserves and role of body size in the mating system of Anopheles gambiae

Anopheles gambiae mates in flight. Males gather at stationary places at sunset and compete for incoming females. Factors that account for male mating success are not known but are critical for the future of any genetic control strategy. The current study explored variations in nutritional reserves (sugars, glycogen, lipids, and proteins) in wild‐caught swarming and resting males and evaluated the effect of body size and wing symmetry on male mating success. Our results showed that glycogen and sugar reserves are mobilized for flight. Males consume proportionally 5.9‐fold as much energy derived from sugars in swarming activities than when they are at rest. Mated males were on average bigger than unmated ones (P<0.0001). A strong correlation between the left and right wings in both mated and unmated males was found and additional analysis on fluctuating asymmetry did not show any indication of mated males being more symmetrical than unmated ones. The distribution of wing size of mated males was focused around a central value, suggesting that intermediate size of males is advantageous in the An. gambiae mating system. The results are discussed in the context of sexual selection.