Mate guarding and territorial aggression vary with breeding synchrony in golden whistlers (<Emphasis Type="Italic">Pachycephala pectoralis</Emphasis>)

Male paternity assurance behaviour during the female fertile period has been widely documented amongst birds. In contrast, how sex-specific behavioural strategies vary with local breeding synchrony levels remains largely unknown. This is important because, in many species, intra-population patterns of extra-pair fertilisation rates, and hence cuckoldry risk, are known to vary with the number of simultaneously fertile females. Each sex may therefore differ in how they behave towards male conspecifics during different degrees of breeding synchrony. Here I provide evidence of such sex-specific differences in the golden whistler (Pachycephala pectoralis), a species in which within-pair paternity assurance is negatively associated with breeding synchrony. Via simulated territorial intrusions using decoy males, I show that males, but not females, increase levels of aggression to male intruders during periods of low synchrony, possibly because cuckoldry risk is greatest during this period. In addition, males appear to invest more effort into mate guarding after, but not before, territorial intrusions during this period. These inter-sexual differences may reflect conflicts in interest between the sexes, with females consistently showing interest in males during the fertile period regardless of synchrony levels and males investing more resources into expelling intruders when the risk of paternity loss is greatest. This study thus provides evidence that males may be able to detect variation in breeding synchrony and cuckoldry risk and adjust their paternity assurance behaviour accordingly.     

Parental care compromises feeding in the pumpkinseed (<Emphasis Type="Italic">Lepomis gibbosus</Emphasis>)

Providing parental care is potentially costly. Costs can arise through elevated energy expenditure or from an increased risk of mortality. A cost of parental care can also occur because a parent is compromised in their ability to forage. We used pumpkinseed Lepomis gibbosus, a fish with an alternative male mating strategy, to test whether parental males differed in their feeding in comparison with females and cuckolder males. To address this question, we examined the stomach contents of female, cuckolder male, and parental male pumpkinseed during the breeding season over an entire diel cycle. We showed that parental males had a lower total weight of food in their stomachs in comparison with females, while cuckolder males did not. Parental males also had a lower weight and number of chironomids in their stomachs. The temporal pattern of feeding of parental males diverged from that of females, and they had a lower probability of pupal chironomids in their stomachs, which implies spatial segregation in foraging. Parental males had a greater probability of conspecific eggs in their stomachs than females, while the probability of egg cannibalism did not differ between cuckolder males and females. Overall, these finding meet predictions in accordance with an assumption that parental care and territoriality can compromise feeding.     

Female ornamentation and territorial conflicts in collared flycatchers (Ficedula albicollis)

Female ornaments in species with conventional sex roles often indicate individual quality, but the evolutionary forces maintaining them are less clear. Sexual competition for breeding opportunities may represent an important role for female signals, especially in polygynous species, but there is little experimental evidence for this. The wing patch size (WPS) of female collared flycatchers indicates age and body condition and predicts social mating patterns. We challenged nest-building females with decoy females of varying WPS and found that the aggressive response of residents increased with decoy WPS, suggesting a role for this female ornament in territorial competition. Our results explain why female WPS predicts territorial distances when mated to a polygynous male and indicate that the role of WPS in female competitive interactions is similar to that in males of the same population.     

Territorial meadow pipit males (<Emphasis Type="Italic">Anthus pratensis</Emphasis>; Passeriformes) become more aggressive in female presence

Although mate guarding as prevention of extra-pair copulation is common among birds, evidence for aggressive behaviour involving physical contact related to mate guarding in passerines is scarce and cases of the presence of one partner directly influencing the aggressiveness of the other are lacking. We investigated the intra-specific territorial behaviour of male meadow pipits (Anthus pratensis; Passeriformes: Motacillidae) at the beginning of the breeding season by placing a pipit model accompanied by an intra-specific song playback in the territory of socially paired males and compared the responses of males whose mates were physically present during trials with those whose females were out of sight. The level of aggression of males was significantly higher in the presence of the female; half of the males in this group physically attacked the model (the most intense and risky aggressive behaviour). Physical attacks did not occur among males whose female was absent during the trial; response to the playback by most of these males was only weak. This pattern may be related to the prevention of extra-pair copulation; if the risks involved in the conflict are outweighed by potential loss of paternity, such aggressive mate guarding may pay off. The apparently overlooked effect on the territorial behaviour of a partner’s passive physical presence during conflict should be further evaluated because it may be important for the design and interpretation of results of behavioural experiments.     

Interannual differences in the relative timing of southward migration of male and female western sandpipers (Calidris mauri)

In the shorebird subfamily Calidridinae, one of the parents shortens parental care and initiates southward migration before the other. We estimated the difference in passage date between male and female western sandpipers (Calidris mauri) at their first major stopover on the southward migration from breeding areas in Alaska, in 18 years between 1978 and 2000. Overall, adult females preceded adult males by 1.22 days. A novel finding was that among juveniles, which migrate approximately a month later than adults, females preceded males by similar magnitude (1.14 days). There was wide variation among years, however, and males actually preceded females in years with late hatch. We relate these findings to hypotheses for female-first southward migration in sandpipers.     

The effect of female wings on male courtship behavior in the cricket Gryllus bimaculatus

Sexual dimorphism exists in the shape and the structure of the forewings of the cricket (Gryllus bimaculatus). However, the functional significance of the wings in the G. bimaculatus female has been unclear. In common blue butterflies (Polyommatus icarus), wings in females have been suggested as being important for attracting males. To test whether female crickets need wings for conspecific males to recognize them and initiate mating behavior, we removed all wings from females and observed the behavior of males towards them. Most males (87.5%) showed mating behavior towards the wingless females: they produced courtship song and transferred spermatophores to the wingless females. Similarly, 88.5% of the males showed mating behavior towards intact females. When males were placed with both a wingless female and an intact female, no significant difference was detected in male mate choice. The findings demonstrate that the wing of the G. bimaculatus female is not necessary for female recognition by conspecific males and the initiation of male mating behavior, and that it is not important in male mate choice.     

Temporal variation in size-assortative mating and male mate choice in a spider with amphisexual care

Males should be more selective when they have a high investment in reproduction, especially in species with biparental or paternal care. In this context, male mate choice can promote size-assortative mating (SAM) when (1) large males win intrasexual disputes, (2) large females are more fecund, and (3) males prefer larger females to smaller ones. In the spider Manogea porracea, males exhibit high reproductive investment by building their webs above those of females and exhibiting extended care of offspring in the absence of females. Under these circumstances, we expect the occurrence of SAM and male preference for large females. Herein, we performed observations and experiments in the field to evaluate the hypotheses that (1) M. porracea mates assortatively by size and (2) SAM is influenced by male mate choice. Furthermore, we measured variables that could affect mating patterns, the sex ratios, and densities of both sexes. Pairing in M. porracea was positively size-assortative in 2012, but not in 2013. Large males won most disputes for mates and preferred larger females, which produced more eggs. The inconsistency in detection of SAM was due to population dynamics, namely variations in sex ratio and population density across the breeding season. Furthermore, we found that the significance of male mate choice on sexual selection of body size in M. porracea strongly depends on the competition intensity for mating opportunities. The traditional sexual selection hypothesis of SAM needs to be reviewed and must include measures of competition intensity.     

Postnatal nutrition influences male attractiveness and promotes plasticity in male mating preferences

Poor early-life nutrition could reduce adult reproductive success by negatively affecting traits linked to sexual attractiveness such as song complexity. If so, this might favor strategic mate choice, allowing males with less complex songs to tailor their mating tactics to maximize the reproductive benefits. However, this possibility has been ignored in theoretical and empirical studies. By manipulating the micronutrient content of the diet (e.g., low or high) during the postnatal period of male zebra finches, we show for the first time (1) that males reared on a poor (low) micronutrient diet had less complex songs as adults; (2) that these males, in contrast to the high micronutrient diet group, were more selective in their mating strategies, discriminating against those females most likely to reduce their clutch size when paired with males having less complex songs; and (3) that by following different mating strategies, males reared on the contrasting diets obtained similar reproductive benefits. These results suggest that early-life dietary conditions can induce multiple and long-lasting effects on male and female reproductive traits. Moreover, the results seem to reflect a previously unreported case of adaptive plasticity in mate choice in response to a nutritionally mediated reduction in sexual attractiveness.     

Females prefer carotenoid colored males as mates in the pentamorphic livebearing fish,<Emphasis Type="Italic"> Poecilia parae</Emphasis>

The first results of female preference and chosen male mating success in a new model organism, the pentamorphic livebearing fish, Poecilia parae, are presented. Poecilia parae is a relative of the guppy, P. reticulata, and is assumed to have similar reproductive behavior. We tested the hypothesis that P. parae females, like female guppies, prefer caretenoid colored males as mates. Here we show that the time a female spent with males was significantly greater for carotenoid coloration in red and yellow melanzona, but time with these two morphs did not differ. The preferred red and yellow males mated significantly more often with their choosing females than did the non-preferred blue and parae males. The few blue melanzona and parae males that mated did so without performing courtship displays. Some females mated with all phenotypes including immaculata males during open group trials. Female P. parae clearly preferred males with carotenoid coloration, thereby corroborating the hypothesis. Alternative male mating tactics by blue melanzona, parae, and immaculata morphs and promiscuous mating by females also resembled features of reproductive behaviors exhibited by guppies.     

Silk wrapping of nuptial gifts as visual signal for female attraction in a crepuscular spider

An extensive diversity of nuptial gifts is known in invertebrates, but prey wrapped in silk is a unique type of gift present in few insects and spiders. Females from spider species prefer males offering a gift accepting more and longer matings than when males offered no gift. Silk wrapping of the gift is not essential to obtain a mating, but appears to increase the chance of a mating evidencing a particularly intriguing function of this trait. Consequently, as other secondary sexual traits, silk wrapping may be an important trait under sexual selection, if it is used by females as a signal providing information on male quality. We aimed to understand whether the white color of wrapped gifts is used as visual signal during courtship in the spider Paratrechalea ornata. We studied if a patch of white paint on the males’ chelicerae is attractive to females by exposing females to males: with their chelicerae painted white; without paint; and with the sternum painted white (paint control). Females contacted males with white chelicerae more often and those males obtained higher mating success than other males. Thereafter, we explored whether silk wrapping is a condition-dependent trait and drives female visual attraction. We exposed good and poor condition males, carrying a prey, to the female silk. Males in poor condition added less silk to the prey than males in good condition, indicating that gift wrapping is an indicator of male quality and may be used by females to acquire information of the potential mate.     

Male wing color properties predict the size of nuptial gifts given during mating in the Pipevine Swallowtail butterfly (Battus philenor)

In many animals, males bear bright ornamental color patches that may signal both the direct and indirect benefits that a female might accrue from mating with him. Here we test whether male coloration in the Pipevine Swallowtail butterfly, Battus philenor, predicts two potential direct benefits for females: brief copulation duration and the quantity of materials the male passes to the female during mating. In this species, males have a bright iridescent blue field on the dorsal hindwing surface, while females have little or no dorsal iridescence. Females preferentially mate with males who display a bright and highly chromatic blue field on their dorsal hindwing. In this study, we show that the chroma of the blue field on the male dorsal hindwing and male body size (forewing length) significantly predict the mass of material or spermatophore that a male forms within the female’s copulatory sac during mating. We also found that spermatophore mass correlated negatively with copulation duration, but that color variables did not significantly predict this potential direct benefit. These results suggest that females may enhance the material benefits they receive during mating by mating with males based on the coloration of their dorsal hindwing.     

Sexually dichromatic coloration reflects size and immunocompetence in female Spanish terrapins, Mauremys leprosa

Many studies have shown the importance of colorful ornamentation in mate choosiness or intrasexual conflict. However, research on color ornaments has focused mainly on birds, lizards or fish, but remains practically unknown in other animal groups such as turtles. In addition, female ornaments and their relation with sexual selection also remain almost unknown. Here, we measured the coloration of the shell and the limb stripes of male and female Spanish terrapins Mauremys leprosa and explored the existence of sexual dichromatism and the relation of color characteristics with body size and health state estimated from the immune response to the injection of an antigen (phytohaemagglutinin test). Our results showed that shell coloration, which could be constrained by natural selection to be cryptic, changed with body size, but did not differ between sexes. In contrast, females had brighter and less ultraviolet-saturated and more orange-saturated limb stripes than males. In females, interindividual variation in limb stripe coloration was related with body size and immune response suggesting that this coloration may inform honestly about multiple traits that could be important in sexual selection. In contrast, coloration of limb stripes of males was duller than in females, and was not related with any trait suggesting that coloration is not important in sexual selection for males.     

Gestational exposure to hydroxyprogesterone caproate suppresses reproductive potential in male rats

Hydroxyprogesterone caproate was administered to pregnant rats at a dose level of 10 and 25 mg/kg body weight on 1st, 7th and 14th gestational day and the male pups (F1 generation) were allowed to grow for 90 days. The effect of gestational exposure to hydroxyprogesterone caproate on fertility was assessed by breeding F1 male rats with control female rats besides analyzing sperm quality and quantity in F1 male rats. The number of implantation sites and viable fetuses was significantly reduced in females mated with F1 males that were exposed to hydroxyprogesterone caproate during embryonic development. The decrease in sperm function was associated with a decrease in sperm motility, sperm viability and sperm count in F1 rats. The study clearly indicates that in utero exposure to hydroxyprogesterone caproate affects fertility in male rats.     

Body size phenotypes are heritable and mediate fecundity but not fitness in the lepidopteran frugivore Cydia pomonella

The inheritance and functional roles of quantitative traits are central concerns of evolutionary ecology. We report two sets of experiments that investigated the heritability and reproductive consequences of body size phenotypes in a globally distributed lepidopteran frugivore, Cydia pomonella (L.). In our first set of experiments, we tested the hypotheses that (1) body size is heritable and (2) parental body size mediates egg production and offspring survival. Midparent–offspring regression analyses revealed that body mass is highly heritable for females and moderately heritable for males. The contribution of fathers to estimates of additive genetic variance was slightly greater than for mothers. Egg production increased with mean parental size, but offspring survival rates were equivalent. Based on this result, we tested two additional hypotheses in a second set of experiments: (3) male size moderates female egg production and egg fertility and (4) egg production, egg fertility, and offspring survival rate are influenced by female mating opportunities. Females paired with large males produced more eggs and a higher proportion of fertile eggs than females paired with small males. Females with multiple mating opportunities produced more fertile eggs than females paired with a single male. However, egg production and offspring survival rates were unaffected by the number of mating opportunities. Our experiments demonstrate that body mass is heritable in C. pomonella and that size phenotypes may mediate fecundity but not fitness. We conclude that male size can influence egg production and fertility, but female mate choice also plays a role in determining egg fertility.     

A "spare" compensates for the risk of destruction of the elongated penis of earwigs (Insecta: Dermaptera)

Male animals in several groups have multiple intromittent organs that outnumber the corresponding female gonopore. In Dermaptera (earwigs), males of the family Anisolabididae have paired, elongated male intromittent organs (virgae), while females have a single sperm-storage organ (spermatheca). Several authors have assumed that one of the paired virgae is non-functional, because it points in the "wrong" direction. We investigated the mating success of handicapped males of Euborellia plebeja in which one of their paired virgae was removed experimentally. These handicapped males succeeded in inseminating a mate. Males with genital damage are found in the field, suggesting that the "spare" functions under natural conditions. Based on phylogenetic information on earwigs, we discuss possible evolutionary scenarios for this genital peculiarity.     

Behavioural evidence of male volatile pheromones in the sex-role reversed wolf spiders Allocosa brasiliensis and Allocosa alticeps

The use of chemical signals in a sexual context is widespread in the animal kingdom. Most studies in spiders report the use of female pheromones that attract potential sexual partners. Allocosa brasiliensis and Allocosa alticeps are two burrowing wolf spiders that show sex-role reversal. Females locate male burrows and initiate courtship before males perform any detectable visual or vibratory signal. So, females of these species would be detecting chemical or mechanical cues left by males. Our objective was to explore the potential for male pheromones to play a role in mate detection in A. brasiliensis and A. alticeps. We designed two experiments. In Experiment 1, we tested the occurrence of male contact pheromones by evaluating female courtship when exposed to empty burrows constructed by males or females (control). In Experiment 2, we tested the existence of male volatile pheromones by evaluating female behaviour when exposed to artificial burrows connected to tubes containing males, females or empty tubes (control). Our results suggest the occurrence of male volatile pheromones that trigger female courtship in both Allocosa species. The sex-role reversal postulated for these wolf spiders could be driving the consequent reversal in typical pheromone-emitter and detector roles expected for spiders.     

Sexual selection on receptor organ traits: younger females attract males with longer antennae

Sexual selection theory predicts that female choice may favour the evolution of elaborate male signals. Darwin also suggested that sexual selection can favour elaborate receiver structures in order to better detect sexual signals, an idea that has been largely ignored. We evaluated this unorthodox perspective by documenting the antennal lengths of male Uraba lugens Walker (Lepidoptera: Nolidae) moths that were attracted to experimentally manipulated emissions of female sex pheromone. Either one or two females were placed in field traps for the duration of their adult lives in order to create differences in the quantity of pheromone emissions from the traps. The mean antennal length of males attracted to field traps baited with a single female was longer than that of males attracted to traps baited with two females, a pattern consistent with Darwin’s prediction assuming the latter emits higher pheromone concentrations. Furthermore, younger females attracted males with longer antennae, which may reflect age-specific changes in pheromone emission. These field experiments provide the first direct evidence of an unappreciated role for sexual selection in the evolution of sexual dimorphism in moth antennae and raise the intriguing possibility that females select males with longer antennae through strategic emission of pheromones.     

Costs of female odour in males of the parasitic wasp <Emphasis Type="Italic">Lariophagus distinguendus</Emphasis> (Hymenoptera: Pteromalidae)

The display of female traits by males is widespread in the animal kingdom. In several species, this phenomenon has been shown to function adaptively as a male mating strategy to deceive sexual rivals (female mimicry). Freshly emerged males of the parasitic wasp Lariophagus distinguendus (Hymenoptera: Pteromalidae) are perceived by other males as if they were females because of a very similar composition of cuticular hydrocarbons which function as a sex pheromone in this species inducing courtship behaviour in males. Within 32 h, however, males deactivate the pheromone and are no longer courted by other males. In this paper, behavioural experiments were performed to test hypotheses on potential costs and benefits associated with the female odour in young males. We did not find any benefits, but demonstrated that young males were significantly more often outrivaled in male-male contests when competing with two older males for a female. Also, young males were significantly more often mounted in homosexual courtship events during these contests. Thus, display of female traits by males is not necessarily beneficial, and in fact, can be disadvantageous. We suggest that these costs have favoured the evolution of the pheromone deactivation mechanism in L. distinguendus males. The function of cuticular hydrocarbons as a female courtship pheromone in L. distinguendus might have evolved secondarily from a primary function relevant for both genders, and the deactivation of the signal in males might have caused a shift of specificity of the chemical signal from the species level to the sex level.     

Sperm-depleted males influence the reproductive behaviour of conspecifics

In many insect species, sperm-depleted males (SDMs, i.e. males that have exhausted their sperm after a given number of matings) remain sexually active and continue to mate females. Here, we investigated the behavioural modifications that occur in both sexes of the parasitoid Asobara tabida Nees (Hymenoptera: Braconidae), after matings by fertile males and sperm-depleted males. We show that (i) virgin females, mated females and females mated to a SDM exhibited different behaviours and that (ii) males responded differently to females depending on whether the females had previously mated with an SDM or not. Our findings demonstrate that SDM influenced the reproductive behaviour of both males and females, especially with regard to male responsiveness and female attractiveness. These findings are discussed in the context of adaptive behaviour and fitness maximization in both males and females.