SEX-RATIO SELECTION IN A BIVOLTINE THRIPS. I. CONDITIONAL SEX-RATIO MANIPULATION AND FITNESS VARIATION

Females of the bivoltine thrips Elaphrothrips tuberculatus (Hood) (Insecta: Thysanoptera) produce broods of either all males (by viviparity) or all females (by oviparity). Measurements of the sex-allocation ratio, ecological and physiological conditions affecting male and female offspring body size, and correlates of the relative fitnesses of adult males and females in relation to size indicate that female parents tend to be viviparous (produce males) if their offspring will become relatively large adults, and that males gain more in fitness from large size than do females. However, the conditions that link sex allocation with offspring fitness differ between the spring and summer generations. In spring, when breeding is synchronous, 1) oviparous and viviparous females do not differ in body size, 2) females tend to be viviparous where the fungus upon which they feed is relatively dense and where their offspring will become relatively large adults, and 3) fungus density is highly correlated with male and female offspring size. In summer, when breeding is relatively asynchronous, 1) viviparous females are much larger than oviparous females early (but not late) in the season, 2) large viviparous females begin breeding earlier than smaller ones, 3) offspring developing earlier in the season become larger adults, and 4) a higher proportion of females are viviparous earlier than later. Field experiments and field collections show that the covariation among sex allocation, conditions, and fitness is not caused by differential mortality by size or sex. Differences between the spring and summer generations in the cues used by females to adjust offspring sex ratio may be caused by seasonal variation in the factors that affect offspring size. However, in both generations, females tend to produce sons only when their offspring will become relatively large adults, whereas daughters are produced regardless of offspring size. These data suggest that females of E. tuberculatus avoid production of males (the sex with higher variance in expected fitness) when the size of their offspring is relatively uncertain.     

Extra-pair young in house wren broods are more likely to be male than female

Sex-allocation theory predicts that females should preferentially produce offspring of the sex with greater fitness potential. In socially monogamous animal species, extra-pair mating often increases the variance in fitness of sons relative to daughters. Thus, in situations where offspring sired by a female''s extra-pair mate(s) will typically have greater fitness potential than offspring sired by the within-pair mate, sex-allocation theory predicts that females will bias the sex of offspring sired by extra-pair mates towards male. We examined the relationship between offspring sex and paternity over six breeding seasons in an Illinois population of the house wren (Troglodytes aedon), a cavity-nesting songbird. Out of the 2345 nestlings that had both sex and paternity assigned, 350 (15%) were sired by extra-pair males. The sex ratio of extra-pair offspring, 0.534, was significantly greater than the sex ratio of within-pair offspring, 0.492, representing an increase of 8.5 per cent in the proportion of sons produced. To our knowledge, this is the first confirmed report of female birds increasing their production of sons in association with extra-pair fertilization. Our results are consistent with the oft-mentioned hypothesis that females engage in extra-pair mating to increase offspring quality.     

Female attractiveness affects paternal investment: experimental evidence for male differential allocation in blue tits

Introduction

The differential allocation hypothesis (DAH) predicts that individuals should adjust their parental investment to their current mate??s quality. Although in principle the DAH holds for both sexes, male adjustment of parental investment has only been tested in a few experimental studies, revealing contradictory results. We conducted a field experiment to test whether male blue tits (Cyanistes caeruleus) allocate their parental effort in relation to female ornamentation (ultraviolet colouration of the crown), as predicted by the DAH.

Results

We reduced the UV reflectance in a sample of females and compared parental care by their mates with that of males paired to sham-manipulated control females. As predicted by the DAH our results demonstrate that males paired with UV-reduced females invested less in feeding effort but did not defend the chicks less than males paired with control females.

Conclusions

To our knowledge, this is one of the first studies providing support for male differential allocation in response to female ornamentation.     

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 choice for good genes and sex-biased broods in guppies

In a population of guppies Poecilia reticulata females were found to prefer large males and the offspring of these males had a higher survival rate than those sired by smaller males, suggesting that females were selecting their mates on the basis of their good genes. The possibility that females differentially invested in male or female offspring depending on the size or attractiveness of their mate was also investigated, but no relationship was found between a male's attractiveness or body size and the sex ratio of the offspring he sired. However, a strong female-biased sex ratio was detected in the broods and the proportion of males produced decreased with the amount of time that a female was confined with a male. The possible reasons for this are discussed.     

Social common mole-rats enhance outbreeding via extra-pair mating

Females in many species engage in matings with males that are not their social mates. These matings are predicted to increase offspring heterozygosity and fitness, and thereby prevent the deleterious effects of inbreeding. We tested this hypothesis in a cooperative breeding mammal, the common mole-rat Cryptomys hottentotus hottentotus. Laboratory-based studies suggested a system of strict social monogamy, while recent molecular studies indicate extensive extra-pair paternity despite colonies being founded by an outbred pair. Our data show that extra-pair and within-colony breeding males differed significantly in relatedness to breeding females, suggesting that females may gain genetic benefits from breeding with non-resident males. Extra-colony male mating success was not based on heterozygosity criteria at microsatellite loci; however, litters sired by extra-colony males exhibited increased heterozygosity. While we do not have the data that refute a relationship between individual levels of inbreeding (Hs) and fitness, we propose that a combination of both male and female factors most likely explain the adaptive significance of extra-pair mating whereby common mole-rats maximize offspring fitness by detecting genetic compatibility with extra-pair mates at other key loci, but it is not known which sex controls these matings.     

Female Control of Offspring Sex Ratios Based on Male Attractiveness in the Guppy

The sex allocation hypothesis predicts that females manipulate the offspring sex ratios according to mate attractiveness. Although there is increasing evidence to support this prediction, it is possible that paternal effects may often obscure the relationship between female control of offspring sex ratios and male attractiveness. In the present study, we examined whether females played a primary role in the manipulation their offspring sex ratios based on male attractiveness, in the guppy Poecilia reticulata, a live‐bearing fish. We excluded the paternal effects by controlling the relative sexual attractiveness of the male by presenting them to the females along with a more attractive or less attractive stimulus male. The test male was perceived to be relatively more attractive by females when it was presented along with a less attractive stimulus male, or vice versa. Subsequently, test male was mated in two different roles (relatively more and less attractive) with two females. If females were responsible for offspring sex ratio manipulation, the sex ratio of the brood would be altered on the basis of the relative attractiveness of the test male. On the other hand, if males play a primary role in offspring sex ratio manipulation, the sex ratios would not differ with the relative attractiveness of the test male. We found that females gave birth to more male‐biased broods when they mated with test males in the attractive role than when they mated with males in the less attractive role. This finding suggests that females are responsible for the manipulation of offspring sex ratios based on the attractiveness of their mates.     

Change in offspring sex ratio over a very short season in Lincoln's Sparrows: the potential role of bill development

ABSTRACT The sex ratios of offspring are targets of natural selection that can affect parental energy expenditure and fitness, adult sex ratios, and population dynamics. Parents may manipulate offspring sex ratios based on sex differences in their offsprings' potential for reproductive success. In Lincoln's Sparrows (Melospiza lincolnii), male bill shape is associated with the quality of songs, and song quality predicts female preferences in a reproductive context. Males and females that hatch later relative to brood mates or later in the breeding season tend to develop bill shapes that are, for males, associated with low‐quality song. Because females do not sing and do not experience this selection pressure, we predicted that the sex of offspring produced late relative to their brood mates or relative to the season should be biased toward females. Using a molecular technique to sex nestlings, we found no effects of hatching order or any interaction between date of clutch initiation (season) and hatching order on offspring sex. However, we found a seasonal decline in the proportion of male offspring, from approximately 0.8 at the beginning to 0.4 at the end of a clutch initiation season only 19 d in duration. To our knowledge, this is the shortest period over which the offspring sex ratio has been shown to change in a bird population. Moreover, these findings are consistent with the hypothesis that sex differences in the potential attractiveness of offspring ultimately influence offspring sex ratios.     

Offspring sex ratio allocation in the parasitic jaeger: selection for pale females and melanic males?

The maintenance of plumage color polymorphism in the parasiticjaeger (Stercorarius parasiticus) is still not well understood.Earlier studies indicated that selection may favor pale femalesand melanic males. If so, females would maximize their fitness,producing pale female and melanic male offspring. We thereforepredicted that females might bias their offspring sex ratiotoward daughters in pale pairs and toward sons in melanic pairs.Females might also choose to mate assortatively in relationto plumage color, thereby maximizing the probability of producingeither pale or melanic offspring. Because females are largerthan males, differential rearing costs may affect the offspringsex ratio independent of parental plumage color. We examinedoffspring sex ratio allocation, breeding variables indicativeof parental quality, and mating pattern in relation to plumagecolor in a colony of parasitic jaegers in northern Norway. Jaegerstended to mate assortatively in relation to plumage color. Thereproductive performance declined with season, and matched pairsappeared to be of lower quality than mixed pairs. The proportionof male offspring increased with hatching date in matched paleand mixed pairs, whereas the situation was reversed in matchedmelanic pairs. Matched pale pairs produced an overall surplusof favorable pale but costly daughters despite their lower quality,while melanic pairs produced a surplus of favorable melanicsons. However, differential offspring rearing costs and parentalrearing capacity may have additionally affected the realizedoffspring sex ratio. Mixed pairs producing an overall surplusof pale and melanic daughters allocated their resources accordingto differential rearing costs and parental quality only. Wesuggest that both strategies of sex ratio allocation togetherwith differences in reproductive success in matched versus mixedpairs may have a balancing effect on the mating pattern betweenplumage morphs and may contribute to the maintenance of thecolor polymorphism in this species.     

Reproductive costs of mating with a sibling male: sperm depletion and multiple mating in Cephalonomia hyalinipennis

Polygynous parasitoid males may be limited by the amount of sperm they can transmit to females, which in turn may become sperm limited. In this study, I tested the effect of male mating history on copula duration, female fecundity, and offspring sex ratio, and the likelihood that females will have multiple mates, in the gregarious parasitoid Cephalonomia hyalinipennis Ashmead (Hymenoptera: Bethylidae: Epyrinae), a likely candidate for sperm depletion due to its local mate competition system. Males were eager to mate with the seven females presented in rapid succession. Copula duration did not differ with male mating history, but latency before a first mating was significantly longer than before consecutive matings. Male mating history had no bearing on female fecundity (number of offspring), but significantly influenced offspring sex ratio. The last female to mate with a given male produced significantly more male offspring than the first one, and eventually became sperm depleted. In contrast, the offspring sex ratio of first‐mated females was female biased, denoting a high degree of sex allocation control. Once‐mated females, whether sperm‐depleted or not, accepted a second mating after a period of oviposition. Sperm‐depleted females resumed production of fertilized eggs after a second mating. Young, recently mated females also accepted a second mating, but extended in‐copula courtship was observed. Carrying out multiple matings in this species thus seems to reduce the cost of being constrained to produce only haploid males after accepting copulation with a sperm‐depleted male. I discuss the reproductive fitness costs that females experience when mating solely with their sibling males and the reproductive fitness gain of males that persist in mating, even when almost sperm‐depleted. Behavioural observations support the hypothesis that females monitor their sperm stock. It is concluded that C. hyalinipennis is a species with a partial local mating system.     

Secondary sex ratio covaries with demographic trends and ecological conditions in the barn swallow

Parents are expected to invest more in the sex that benefits most from the local environment. When the quality of breeding sites varies spatially and natal dispersal of males and females differs, parents in high-quality habitats should skew their progeny sex ratio in favor of the less dispersing sex. We tested this prediction in the barn swallow (Hirundo rustica L.), by relating the proportion of male offspring around fledging (secondary sex ratio) of first and second broods to the ecological quality (presence of livestock farming and relative surface of hayfields in the foraging range) and local demographic trends of the farms where the colonies were located. Consistent with our predictions, the proportion of male offspring, which are more philopatric than females, increased with the extent of hayfields, which are high quality, preferred foraging habitats. Moreover, the proportion of male offspring in second broods was smaller in colonies with positive demographic trends, possibly indicating density-dependent effects on sex ratio. Independent of the mechanism generating uneven sex ratio (zygote sex ratio adjustment or sex-related pre-fledging mortality), barn swallows breeding under favorable conditions overproduced the sex that is more likely to benefit from such conditions.     

Male dominance determines female egg laying rate in crickets

A key prediction of theories of differential allocation and sexual conflict is that male phenotype will affect resource allocation by females. Females may adaptively increase investment in offspring when mated to high quality males to enhance the quality of their offspring, or males may vary in their ability to manipulate female investment post-mating. Males are known to be able to influence female reproductive investment, but the male traits underlying this ability have been little studied in taxa other than birds. We investigated the relationship between male dominance and female oviposition rate in two separate experiments using the field cricket, Gryllus bimaculatus. In both experiments, females mated to more dominant (but not larger) males laid more eggs. This reveals that either females allocate more effort to reproduction after mating with a dominant male or that dominance status is associated with male ability to manipulate their mates. This is the first evidence that dominance, rather than male attractiveness, has a post-copulatory effect on reproductive investment by females.     

Density-related changes in sexual selection in red deer.

In sexually dimorphic mammals, high population density is commonly associated with increased mortality of males relative to females and with female-biased adult sex ratios. This paper investigates the consequences of these changes on the distribution of male breeding success, the intensity of competition for females and the opportunity for sexual selection. After the red deer (Cervus elaphus L.) population of the North Block of Rum (Inner Hebrides) was released from culling, female numbers rose and male numbers declined, leading to an adult sex ratio of around one male to two females. This change was the result of increased mortality of males relative to females during the first two years of life; of increased emigration rates by young males; and of reduced immigration by males from outside the study area. The increasing bias in the adult sex ratio affected the timing of breeding as well as the distribution of mating success in males. As the adult sex ratio became increasingly biased towards females, the degree of skew in mating success (calculated across all harem-holders) increased, but mature males defended harems for shorter periods and a higher proportion of males held harems. In addition, a higher proportion of calves were fathered by immigrant males and the proportion fathered by males born in the study area declined. These results support the contention that, where high population density is associated with a female-biased adult sex ratio, competition for mates is likely to decline.     

Reproductive strategies of the larval parasitoid Microplitis croceipes

Mate choice may have important consequences for offspring sex ratio and fitness of haplodiploid insects. Mate preference of females of the solitary larval parasitoid Microplitis croceipes (Cresson) (Hymenoptera: Braconidae) for virgin and mated males, and vice versa, and the reproductive consequences (i.e., the sex ratio expressed as the proportion of male offspring) were examined in choice and non‐choice experiments. In addition, the effect of repeated rapid and daily copulation of an individual male on the sex ratio of offspring of the female mates was assessed. Males preferred virgins over mated females, whereas females copulated with a male irrespective of his mating status. In both the rapid and daily copulation assay, females copulating with a male that had copulated five times or more produced a higher sex ratio than females that had copulated with a virgin male. Females that copulated with virgin males once or twice produced a significantly and considerably lower sex ratio than females that first copulated with a sperm‐depleted male followed by a virgin male. This indicates that copulating with a sperm‐depleted male has costs and limits acquisition by the female of sperm from virgin males.     

Differential sex allocation in sand lizards: bright males induce daughter production in a species with heteromorphic sex chromosomes

In sand lizards (Lacerta agilis), males with more and brighter nuptial coloration also have more DNA fragments visualized in restriction fragment length polymorphism analysis of their major histocompatibility complex class I loci (and, hence, are probably more heterozygous at these loci). Such males produce more viable offspring, with a particularly strong viability effect on daughters. This suggests that females should adjust both their reproductive investment and offspring sex ratio in relation to male coloration (i.e. differential allocation). Our results show that experimental manipulation of partner coloration in the wild results in significantly higher maternal effort and a 10% higher proportion of daughters than sons. This supports the hypothesis that females increase their maternal energetic expenditure and adjust their offspring sex ratio in response to high-quality partners. However, it also suggests that this has probably evolved through natural selection for increased offspring viability (primarily through production of daughters), rather than through increased mate attraction (e.g. sexy sons).     

Temporal but not spatial environmental variation drives adaptive offspring sex allocation in a plural cooperative breeder

Cooperatively breeding birds have been used frequently to study sex allocation because the adaptive value of the sexes partly depends upon the costs and benefits for parents of receiving help. I examined patterns of directional sex allocation in relation to maternal condition (Trivers-Willard hypothesis), territory quality (helper competition hypothesis), and the number of available helpers (helper repayment hypothesis) in the superb starling, Lamprotornis superbus, a plural cooperative breeder with helpers of both sexes. Superb starlings biased their offspring sex ratio in relation to prebreeding rainfall, which was correlated with maternal condition. Mothers produced relatively more female offspring in wetter years, when they were in better condition, and more male offspring in drier years, when they were in poorer condition. There was no relationship between offspring sex ratio and territory quality or the number of available helpers. Although helping was male biased, females had a greater variance in reproductive success than males. These results are consistent with the Trivers-Willard hypothesis and suggest that although females in most cooperatively breeding species make sex allocation decisions to increase their future direct reproductive success, female superb starlings appear to base this decision on their current body condition to increase their own inclusive fitness.     

Female Cape sugarbirds (Promerops cafer) modify egg investment both for extra‐pair mates and for male tail length

The differential allocation hypothesis predicts that females should invest more in reproduction when paired with attractive males. We measured egg volume in Cape sugarbirds (Promerops cafer), a sexually dimorphic passerine, in relation to paternity of the offspring and in response to an experimental tail length treatment. We manipulated tail length, after pair formation, but before egg laying: males had their tails either shortened or left unmanipulated. Our manipulation was designed to affect female allocation in a particular breeding attempt rather than long‐term mate choice: males with shortened tails would appear to be signalling at a lower level than they should given their quality. We found that egg volume was smaller in the nests of males with experimentally shortened tails but larger when the offspring were the result of extra‐pair matings. Both these findings are consistent with the differential allocation hypothesis. We suggest that tail length may be used by females as a cue for mate quality, eliciting reduced female investment when breeding with social mates; and with males with shortened tails.     

The Effects of Copulatory Courtship on Differential Allocation in the Red Flour Beetle <Emphasis Type="BoldItalic">Tribolium castaneum</Emphasis>

Mate attractiveness is known to sometimes influence female reproductive investment (i.e. differential allocation) and the sex ratio of her offspring (i.e. sex allocation). Males of the red flour beetle Tribolium castaneum rub the lateral edges of the females elytra with their tarsi during copulation. This behavior is important for paternity success when females have mated with two males. We manipulated female perception of the leg rubbing behavior by tarsal ablation and tested whether this behavior is also favored through differential allocation and whether it affects sex allocation. We found some support for an increase in female oviposition rate in response to intensive leg rubbing but failed to find any support for an effect on sex allocation. The overall sex ratio of offspring was slightly male biased but females did not appear to regulate the sex ratio of their offspring.     

Female blue tits adjust parental effort to manipulated male UV attractiveness

The differential allocation hypothesis predicts that parents should adjust their current investment in relation to perceived mate attractiveness if this affects offspring fitness. It should be selectively advantageous to risk more of their future reproductive success by investing heavily in current offspring of high reproductive value but to decrease investment if offspring value is low. If the benefits of mate attractiveness are limited to a particular offspring sex we would instead expect relative investment in male versus female offspring to vary with mate attractiveness, referred to as 'differential sex allocation'. We present strong evidence for differential allocation of parental feeding effort in the wild and show an immediate effect on a component of offspring fitness. By experimentally reducing male UV crown coloration, a trait known to indicate attractiveness and viability in wild-breeding blue tits (Parus caeruleus), we show that females, but not males, reduce parental feeding rates and that this reduces the skeletal growth of offspring. However, differential sex allocation does not occur. We conclude that blue tit females use male UV coloration as an indicator of expected offspring fitness and adjust their investment accordingly.     

The significance of biparental care in the brown bullhead,Ictalurus nebulosus

Synopsis Parental care in the brown bullhead is characterized by variation in the participation of each sex. Most broods are attended by both sexes, but some are attended by a male alone, or rarely, a female alone. Two care-givers were more successful than one alone in fostering offspring survival. However, there was no significant difference between two care-givers and one alone in the proportion of time that broods were unattended. Potential brood predators were chased less frequently by one adult alone than by adults aided by their mate. This difference may be unimportant since two adults simultaneously attended their brood only 19% of the time. Males alone attended their broods a significantly greater proportion of time than did either males or females aided by their mates. This difference suggests that males alone sustain a greater cost of care-giving (starvation and therefore reduced future reproduction) than do males aided by their mates. Thus, males alone may more often leave broods (and not return) than males that are aided in care-giving. The differential success observed may be due to a difference in the likelihood that the male (the principal care-giver) leaves the brood permanently, rather than differences in the quality of care one or two adults provide. I suggest that two care-givers are more successful than one because the net benefits of care-giving exceed the net benefits of leaving for males when aided by their mates.