Female‐Biased Helping in a Cooperatively Breeding Bird: Female Benefits or Male Costs?

There is often a sex bias in helping effort in cooperatively breeding species with both male and female helpers, and yet this phenomenon is still poorly understood. Although sex‐biased helping is often assumed to be correlated with sex‐specific benefits, sex‐specific costs could also be responsible for sex‐biased helping. Cooperatively breeding brown jays (Cyanocorax morio) in Monteverde, Costa Rica have helpers of both sexes and dispersal is male‐biased, a rare reversal of the female‐biased dispersal pattern often seen in birds. We quantified helper contributions to nestling care and analyzed whether there was sex‐biased helping and if so, whether it was correlated with known benefits derived via helping. Brown jay helpers provided over 70% of all nestling feedings, but they did not appear to decrease the workload of breeders across the range of observed group sizes. Female helpers fed nestlings and engaged in vigilance at significantly higher levels than male helpers. Nonetheless, female helpers did not appear to gain direct benefits, either through current reproduction or group augmentation, or indirect fitness benefits from helping during the nestling stage. While it is possible that females could be accruing subtle future direct benefits such as breeding experience or alliance formation from helping, future studies should focus on whether the observed sex bias in helping is because males decrease their care relative to females in order to pursue extra‐territorial forays. Explanations for sex‐biased helping in cooperative breeders are proving to be as varied as those proposed for helping behavior in general, suggesting that it will often be necessary to quantify a wide range of benefits and costs when seeking explanations for sex‐biased helping.     

Absence of Mate Choice and Postcopulatory Benefits in a Species with Extreme Sexual Size Dimorphism

Most hypotheses related to the evolution of female‐biased extreme sexual size dimorphism (SSD) attribute the differences in the size of each sex to selection for reproduction, either through selection for increased female fecundity or selection for male increased mobility and faster development. Very few studies, however, have tested for direct fitness benefits associated with the latter – small male size. Mecaphesa celer is a crab spider with extreme SSD, whose males are less than half the size of females and often weigh 10 times less. Here, we test the hypotheses that larger size in females and smaller size in males are sexually selected through differential pre‐ and postcopulatory reproductive benefits. To do so, we tested the following predictions: matings between small males and large females are more likely to occur due to mate choice; females mated to small males are less likely to accept second copulation attempts; and matings between small males and large females will result in larger clutches of longer‐lived offspring. Following staged mating trials in the laboratory, we found no support for any of our predictions, suggesting that SSD in M. celer may not be driven by pre‐ or post‐reproductive fitness benefits to small males.     

Seasonal Variation in Mate Choice of Photinus ignitus Fireflies

Mate choice by either sex may vary with changes in the associated costs and benefits, determined by factors such as the availability of potential mates and variation in mate quality. We examined seasonal variation in operational sex ratio, courtship behavior, spermatophore mass, egg count, and the relationship between morphological traits and mating success in Photinus ignitus fireflies to determine if mate choice in either sex varied with the availability and relative reproductive investment of fertilizable females and sexually active males. Successfully mating males had larger lanterns than unsuccessful males when the operational sex ratio was male‐biased. In addition, female responsiveness to male signals increased as the number of courting males decreased, and male spermatophore mass decreased with body size across the mating season. Successfully mating females had larger body mass than unsuccessful females. Female body mass predicted egg count and female rejection by males increased as the season progressed and female size decreased. These results suggest that both male and female P. ignitus exhibit mate choice, and that such choice is influenced by seasonal variation in the abundance and quality of potential mates.     

Male-female affiliative relationships in naturally occurring ringtailed lemurs (Lemur catta) at the Beza-Mahafaly Reserve,Madagascar

Affiliative behavior between adult male and female ringtailed lemurs was examined as part of a project concerning male affiliation with conspecifics of all age/sex classes. Males in three social groups were studied over a 12 month period. Male-female preferred partnerships existed, and were variable according to reproductive season. Dominance rank, age, or tenure of the male did not appear to affect either the number of partnerships or frequency of affiliative behaviors that males had with females. However, males residing in groups with fewer males exhibited both higher frequencies of affiliative interactions with females and were nearest neighbors to females more often than males living in a group containing more males. Females were found to be responsible for proximity maintenance of male-female dyads in the majority of cases. Neither reproductive season nor seasonal availability of food resources strongly affected the frequency of affiliative interactions between males and females, it is proposed that an important aspect of successful group membership for male ringtailed lemurs relates to the development of social relationships with adult females. Males can benefit from such relationships in terms of greater centrality to the spatial core of the group, which can result in enhanced predator protection, greater opportunities for social contact, and potentially greater access to estrous females. © 1996 Wiley-Liss, Inc.     

Testing Functional Hypotheses for the Behavior of Resident Pine Voles, Microtus pinetorum, Toward Non-Residents

Members of a social group should attempt to maximize their fitness by maintaining an optimal group composition. Allowing an immigrant into the group may be beneficial or costly depending on the characteristics of the immigrant as well as the composition of the group. Therefore, we examined behavioral interactions between pine voles to test three functional hypotheses proposed to explain behavior of residents toward non‐residents: the resource defense, mate defense, and benefit of extra‐pair copulation hypotheses. To test these, we examined the effects of age, sexual experience and sex of non‐residents as well as the effects of sex of residents on the behavior of residents. Neither male nor female residents showed affiliative behavior toward non‐residents. Residents were more aggressive toward non‐residents than vice versa. The frequency of same‐sex aggression was greater than opposite‐sex aggression for male residents and this aggression was directed at adult male non‐residents to a greater degree than at subadult males. Resident males were least aggressive toward adult females. We found no differences in the behavior of females toward subadults, sexually naive adult non‐residents or sexually experienced adult non‐residents. Females also displayed similar amounts of aggression toward male and female non‐residents. Therefore, for males, aggression may function in defense of a mate while for females, aggression functions in resource defense. For both sexes, aggression is likely to play a role in the regulation of group composition.     

Sexual dimorphism,survival, and parental investment in relation to offspring sex in a precocial bird

Differential growth rate between males and females, owing to a sexual size dimorphism, has been proposed as a mechanism driving sex‐biased survival. How parents respond to this selection pressure through sex ratio manipulation and sex‐biased parental investment can have a dramatic influence on fitness. We determined how differential growth rates during early life resulting from sexual size dimorphism affected survival of young and how parents may respond in a precocial bird, the black brant Branta bernicla nigricans. We hypothesized that more rapidly growing male goslings would suffer greater mortality than females during brood rearing and that parents would respond to this by manipulating their primary sex ratio and parental investment. Male brant goslings suffered a 19.5% reduction in survival relative to female goslings and, based on simulation, we determined that a female biased population sex ratio at fledging was never overcome even though previous work demonstrated a slight male‐biased post‐fledging survival rate. Contrary to the Fisherian sex ratio adjustment hypothesis we found that individual adult female brant did not manipulate their primary sex ratio (50.39% male, n = 645), in response to the sex‐biased population level sex ratio. However, female condition at the start of the parental care period was a good predictor of their primary sex ratio. Finally, we examined how females changed their behavior in response to primary sex ratio of their broods. We hypothesized that parents would take male biased broods to areas with increased growth rates. Parents with male biased primary sex ratios took broods to areas with higher growth rates. These factors together suggest that sex‐biased growth rates during early life can dramatically affect population dynamics through sex‐biased survival and recruitment which in turn affects decisions parents make about sex allocation and sex‐biased parental investment in offspring to maximize fitness.     

The effect of size and sex ratio experiences on reproductive competition in Nicrophorus vespilloides burying beetles in the wild

Male parents face a choice: should they invest more in caring for offspring or in attempting to mate with other females? The most profitable course depends on the intensity of competition for mates, which is likely to vary with the population sex ratio. However, the balance of pay‐offs may vary among individual males depending on their competitive prowess or attractiveness. We tested the prediction that sex ratio and size of the resource holding male provide cues regarding the level of mating competition prior to breeding and therefore influence the duration of a male's biparental caring in association with a female. Male burying beetles, Nicrophorus vespilloides were reared, post‐eclosion, in groups that differed in sex ratio. Experimental males were subsequently translocated to the wild, provided with a breeding resource (carcass) and filmed. We found no evidence that sex ratio cues prior to breeding affected future parental care behaviour but males that experienced male‐biased sex ratios took longer to attract wild mating partners. Smaller males attracted a higher proportion of females than did larger males, securing significantly more monogamous breeding associations as a result. Smaller males thus avoided competitive male–male encounters more often than larger males. This has potential benefits for their female partners who avoid both intrasexual competition and direct costs of higher mating frequency associated with competing males.     

FEMALE PREFERENCE FOR MALE COURTSHIP EFFORT CAN DRIVE THE EVOLUTION OF MALE MATE CHOICE

The evolution of male mate choice is constrained by costs of choice in species with a male‐biased operational sex ratio (OSR). Previous theoretical studies have shown that significant benefits of male choice are required, for example, by mating with more fecund females, in order for these costs to be offset and a male preference to spread. In a series of population genetic models we show the novel effect that male mating preference, expressed as a bias in courtship, can spread when females prefer, and thus are more likely to mate with, males who court more. We explore two female preference functions for levels of male courtship, one representing a threshold and the other a weighted female preference. The basic finding generally holds for both preference functions. However, the preference function greatly affects the spread of a male preference allele after the addition of competing males who can court more in total. Our results thus stress that a thorough understanding of the response of females to male courtship is a critical component to understanding male preference evolution in polygynous species.     

Sperm competition generates evolution of increased paternal investment in a sex role‐reversed seed beetle

When males provide females with resources at mating, they can become the limiting sex in reproduction, in extreme cases leading to the reversal of typical courtship roles. The evolution of male provisioning is thought to be driven by male reproductive competition and selection for female fecundity enhancement. We used experimental evolution under male‐ or female‐biased sex ratios and limited or unlimited food regimes to investigate the relative roles of these routes to male provisioning in a sex role‐reversed beetle, Megabruchidius tonkineus, where males provide females with nutritious ejaculates. Males evolving under male‐biased sex ratios transferred larger ejaculates than did males from female‐biased populations, demonstrating a sizeable role for reproductive competition in the evolution of male provisioning. Although larger ejaculates elevated female lifetime offspring production, we found little evidence of selection for larger ejaculates via fecundity enhancement: males evolving under resource‐limited and unlimited conditions did not differ in mean ejaculate size. Resource limitation did, however, affect the evolution of conditional ejaculate allocation. Our results suggest that the resource provisioning that underpins sex role reversal in this system is the result of male–male reproductive competition rather than of direct selection for males to enhance female fecundity.     

Costs of rearing and sex‐ratio variation in southern grey shrike Lanius meridionalis broods

Several non‐mutually exclusive hypotheses predict adaptive variation in the offspring sex ratio. When conditions for breeding are adverse, parents are predicted to produce more offspring of the less costly sex to rear (‘the cost‐of‐reproduction hypothesis’). Moreover, they also should produce the more dispersing sex in order to diminish future competition (‘the local‐resource‐competition hypothesis’). Here, we analyse brood sex ratio according to rearing conditions in the southern shrike Lanius meridionalis, a species with moderately reversed sexual dimorphism. Our results suggest that females are more costly to rear than males in this species. Adult females proved heavier than males, and female nestling tended to be heavier than male nestlings. Moreover, the greater brood reduction, the more male‐biased was the brood, suggesting that brood reduction implied higher mortality in female nestlings. Consistent with these findings, the brood sex ratio was biased to the less costly sex (males) when breeding conditions were adverse (bad years or low‐quality male parents), supporting the cost‐of‐reproduction hypothesis. By contrast, these findings did not support the local‐resource‐competition hypothesis, which predicted female‐biased brood sex ratio under adverse conditions. As a whole, our results support the idea that birds adaptively modulate sex ratio in order to minimize reproduction costs.     

Within‐Species Mate Preferences Do Not Contribute to the Maintenance of Sexually Monomorphic Mating Signals in Green Lacewings

We know much less about the evolutionary forces and constraints that maintain similar mating displays in females and males than we do about sexually dimorphic mating displays. Both female and male green lacewings have sexually monomorphic vibrational mating signals and are equally choosy against heterospecific mating signals. This similarity in between‐species sex roles may explain a large part of the presence of species‐specific female signals in these species, but does not necessarily predict why female and male signals are similar. We tested for within‐species sex‐specific similarities in mate preferences in Chrysoperla lucasina that may contribute to the maintenance of sexually monomorphic mating signals in this species. We found weak preferences and low levels of discrimination for signals with varying fine‐scale temporal features (volley duration, period, and volley‐duty cycle). The longer signals that both sexes produced in response to playback were sexually monomorphic, but some females and most males also produced shorter signals with significantly reduced volley durations and periods. Notably, all of these signals had indistinguishable volley‐duty cycles, the ratio of volley duration to volley period. We propose that mating signals in C. lucasina are maintained in both sexes because of similar between‐species mate preferences, but the sexually monomorphic mating signals cannot be attributed to significant within‐species mate preferences. What differences are present in within‐species sex roles may be resolved by a male‐biased signal polymorphism, in long and short signals that are hypothesized to have distinct functions during mate calling and courtship.     

Getting that female glance: Patterns and consequences of male nonverbal behavior in courtship contexts

Females apparently are the choosier sex in courtship contexts, but there still is limited information about female selection criteria in real courtship settings. Given that a female knows little about a heretofore unacquainted male, upon what dimensions can (and do) females base their initial courtship decisions? Here, we report findings from observational studies that investigated male nonverbal behavior in a bar context. Study 1 documented the body movements of males prior to making contact with a female. It was found that males who successfully made “contact” courtship initiation with females exhibited different body language in this precontact phase than did males who did not make contact with females, including significantly more glancing behaviors, space-maximization movements, intrasexual touching, and less closed-body movements. The findings from a second within-subject study comparing the behavior of men in a bar when women were present or not present supported the initial study's findings and showed that males' emphasis on these behaviors increases in a mate-relevant context. We suggest that certain aspects of male nonverbal behavior in courtship contexts can serve as self-presentation and mate-value signals.     

Male chimpanzee behavior in relation to female ano-genital swelling

This study examined the relationships between male agonistic, affiliative, and sexual behaviors and female estrus condition in captive adolescent and young-adult chimpanzees (Pan troglodytes). Data on agonistic, affiliative, and sexual behaviors of 11 males living in three social groups were collected during daily 45 minute observations over a 5 month period. Female estrus condition was assessed daily using the relative size of the female's ano-genital swelling. It was hypothesized that the presence of maximally tumescent females would generate conflicts between males, so an increase in inter-male agonism was predicted. Males exhibited higher rates of agonism toward other males when at least one female in the group was maximally tumescent. Male affiliative behavior directed toward other males and social play with males were affected by the presence and number of maximally tumescent females. Male sexual behavior increased when maximally tumescent females were present.     

Testing the predictions of sex allocation hypotheses in dimorphic,cooperatively breeding riflemen

Evolutionary theory predicts that parents should invest equally in the two sexes. If one sex is more costly, a production bias is predicted in favour of the other. Two well‐studied causes of differential costs are size dimorphism, in which the larger sex should be more costly, and sex‐biased helping in cooperative breeders, in which the more helpful sex should be less costly because future helping “repays” some of its parents’ investment. We studied a bird species in which both processes should favor production of males. Female riflemen Acanthisitta chloris are larger than males, and we documented greater provisioning effort in more female‐biased broods indicating they are likely costlier to raise. Riflemen are also cooperative breeders, and males provide more help than females. Contrary to expectations, we observed no male bias in brood sex ratios, which did not differ significantly from parity. We tested whether the lack of a population‐wide pattern was a result of facultative sex allocation by individual females, but this hypothesis was not supported either. Our results show an absence of adaptive patterns despite a clear directional hypothesis derived from theory. This appears to be associated with a suboptimal female‐biased investment ratio. We conclude that predictions of adaptive sex allocation may falter because of mechanistic constraint, unrecognized costs and benefits, or weak selection.     

Mate choice and sexual conflict in <Emphasis Type="Italic">Brachionus plicatilis</Emphasis> (Rotifera)

Male mate choice in Brachionus plicatilis is based on information from coronal receptors and contact chemoreception of a glycoprotein signal on the body surface of females. Male mating behavior is modulated by mating signal quality and strength, which determines male mating persistence and circling intensity. We probe the sensory abilities of males by better resolving their discrimination of female age. Males preferentially initiate mating with young females, but not too young. Males circle 0.2 h old females just hatched from resting eggs only about 1/2 as frequently as 12 h old females. Males are even more discriminating of females with whom they copulate, preferring 3 h old females significantly more than 0.2 h or 6 h old females. In contrast, males cannot distinguish between virgin females and those who have already copulated. Similarly, males circled and copulated with females hatching from resting eggs with equal frequency as those hatching from amictic eggs. The counterpoint to male mate choice is female resistance to mating. When males contact females, the females respond with one of four behaviors: no response, acceleration, foot flipping, or coronal retraction. In 65% of male–female encounters, there was no initial response by females. However, when males began circling females, females accelerated 11.1 times more often than when males were absent. The second type of evasive female behavior was foot-flipping, which tends to knock off males attempting to circle the female. In the presence of circling males, females performed foot-flipping behavior three times more often than in the absence of males. Coronal retraction, where they stop swimming and withdraw their corona, was observed less frequently than acceleration or foot-flipping, and there was no difference in the presence or absence of males. These data are interpreted in the context of sexual conflict, where the behaviors that optimize male and female fitness differ. Guest editors: S. S. S. Sarma, R. D. Gulati, R. L. Wallace, S. Nandini, H. J. Dumont & R. Rico-Martínez Advances in Rotifer Research     

Sex Differences in Territorial Behavior Exhibited by the Spotted Hyena (Hyaenidae, Crocuta crocuta)

Spotted hyenas ( Crocuta crocuta ) are gregarious carnivores that defend group territories against encroachment by neighboring conspecifics. Here we monitored the behavior of members of one clan of free-ranging spotted hyenas during border patrols, 'wars' with neighboring clans, and other interactions with alien intruders, to document differences between the sexes in territorial behavior in this species. We also examined the possibility that the probability or rate of attack on alien hyenas encountered within the clan's territory would vary with the sex of the intruders. Initiation and leadership of most cooperative territorial behaviors were by adult female clan members, although border patrols were occasionally conducted by groups composed exclusively of resident males. The vast majority of alien intruders into the territory of the study clan were males. Resident females were more likely to attack intruding females than intruding males, but hourly rates of aggression directed by females towards aliens did not vary with intruder sex. Resident males were more likely than resident females to attack alien males, and resident males directed significantly higher hourly rates of aggression towards intruding males than females. Although female leadership in most cooperative territorial behaviors distinguishes spotted hyenas from many mammalian carnivores, other sex differences in the territorial behavior of spotted hyenas resemble those documented in other gregarious predators. Sex differences observed in hyena territoriality are consistent with the hypothesis that male and female clan members derive different selective benefits from advertisement and defense of group territories.     

COSTLY SIGNALS IN A FIELD CRICKET CAN INDICATE HIGH‐ OR LOW‐QUALITY DIRECT BENEFITS DEPENDING UPON THE ENVIRONMENT

The handicap hypothesis proposes that male signals provide reliable information to females because only males of high condition provide high‐quality mating benefits and can afford the costs of producing attractive signals. In the context of direct benefits, the handicap hypothesis predicts that benefit quality and signal attractiveness will positively covary among genotypes, positively covary among environments, or be affected by congruent genotype–environment interactions. The latter should occur if the relative condition of a genotype is environment‐dependent. We tested these predictions in the variable field cricket, Gryllus lineaticeps. An interaction between male family and nutritional environment affected the expression of a costly signal preferred by females, while only male family affected direct benefit quality. These noncongruent effects of family and nutritional environment are inconsistent with the handicap hypothesis, and appear to have resulted from variation among nutritional environments in the relationship between signal attractiveness and benefit quality. Surprisingly, signal attractiveness was positively correlated with benefit quality when males experienced a low nutrition environment but negatively correlated with benefit quality when males experienced a high nutrition environment. As a result, female choice for direct benefits may be difficult, particularly in heterogeneous environments, unless females can assess the environmental histories of males.     

Incidental Sanctions and the Evolution of Direct Benefits

Recent research has shown that a variety of traits that increase male success in mating and sperm competition can impose costs on females, resulting in antagonistic coevolution between the sexes. Yet, in many animals, females are known to receive direct benefit from their mates, including many in which female multiple mating results in intense sperm competition among males. The most common explanation for the evolution of male‐provided direct benefits is pre‐mating female choice based on benefit quality. This explanation is insufficient, however, for those direct benefits that females cannot directly assess prior to mating. Given that intrasexual selection will often favor male traits that increase female mating costs, many types of direct benefits can thus be difficult to explain. In this paper, we review four additional hypotheses for the evolution of male‐provided direct benefits, and present a fifth hypothesis that has received little attention. This latter hypothesis proposes that selection often favors female reproductive tactics that are conditional upon the past costs and benefits of mating. These conditional female reproductive tactics should evolve because the quality of the benefit provided by a previous mate can change the costs and benefits of alternative reproductive decisions. Furthermore, many of the conditional reproductive tactics we might expect females to express should incidentally penalize males which provide lower quality direct benefits. These conditional reproductive tactics may thus play an important role in determining whether females incur costs or receive benefits from their mates. In addition to favoring the evolution of direct benefits, we argue that conditional female reproductive tactics may also favor reliable signaling of benefit quality. The most common explanation for reliable signaling is the handicap mechanism, which proposes that differential costs of signaling prevent low quality males from deceptively producing attractive signals. For direct benefits, however, there is a second type of deception: males which produce attractive signals and can afford to provide high quality direct benefits may choose to cheat on the advertised benefit. The handicap mechanism does nothing to prevent cheating on direct benefits by males which can afford to produce attractive signals, and is thus insufficient for ensuring reliable signaling of benefit quality. In contrast, conditional female reproductive tactics that incidentally penalize low benefit males should also penalize males which cheat on the benefits advertised by their signals.     

Influences of Mating Group Composition on the Behavioral Time-Budget of Male and Female Alpine Ibex (Capra ibex) during the Rut

During the rut, polygynous ungulates gather in mixed groups of individuals of different sex and age. Group social composition, which may vary on a daily basis, is likely to have strong influences on individual’s time-budget, with emerging properties at the group-level. To date, few studies have considered the influence of group composition on male and female behavioral time budget in mating groups. Focusing on a wild population of Alpine ibex, we investigated the influence of group composition (adult sex ratio, the proportion of dominant to subordinate males, and group size) on three behavioral axes obtained by Principal Components Analysis, describing male and female group time-budget. For both sexes, the first behavioral axis discerned a trade-off between grazing and standing/vigilance behavior. In females, group vigilance behavior increased with increasingly male-biased sex ratio, whereas in males, the effect of adult sex ratio on standing/vigilance behavior depended on the relative proportion of dominant males in the mating group. The second axis characterized courtship and male-male agonistic behavior in males, and moving and male-directed agonistic behavior in females. Mating group composition did not substantially influence this axis in males. However, moving and male-directed agonistic behavior increased at highly biased sex ratios (quadratic effect) in females. Finally, the third axis highlighted a trade-off between moving and lying behavior in males, and distinguished moving and female-female agonistic behavior from lying behavior in females. For males, those behaviors were influenced by a complex interaction between group size and adult sex ratio, whereas in females, moving and female-female agonistic behaviors increased in a quadratic fashion at highly biased sex ratios, and also increased with increasing group size. Our results reveal complex behavioral trade-offs depending on group composition in the Alpine ibex, and emphasize the importance of social factors in influencing behavioral time-budgets of wild ungulates during the rut.     

Sequential Analysis Reveals Behavioral Differences Underlying Female‐Biased Predation Risk in Stalk‐Eyed Flies

Stalk‐eyed flies are classic models of how sexual selection can drive morphological and behavioral elaboration. Exaggerated ornaments born by stalk‐eyed flies could impose locomotor costs and increase susceptibility to predation; however, a previous study determined that behavior, not eye span, was the major influence on predation risk. Despite the importance of behavior, relatively little is known about how these flies avoid and deter predators. We created an ethogram of behaviors and used it to score individual interactions of male and female Teleopsis dalmanni paired with an actively foraging, generalist arachnid predator (Phidippus audax). Sequential analysis was employed to identify temporal patterns in behavior and determine how males and females differ in their approaches to avoiding predation. Our results indicate that males and females significantly differ when specific behaviors were employed. Patterns in the behavioral transitions suggest that males are more aggressive than females and are more likely to approach a predator to jab, abdomen bob, or display. Males elicited more retreat responses from the predator, whereas females elicited more attacks. Although the behavioral repertoires of male and female stalk‐eyed flies are indistinguishable, their uses of the behaviors differ, particularly the sequential order of presentation, suggesting a strong sex difference in anti‐predatory behavior.