Foraging Cost of a Long Tail Ornament: An Experiment with Sand Martin Females

The handicap hypothesis assumes that sexual ornaments impose a viability cost upon the bearers. There have been few empirical tests of this assumption. Previous studies show evidence for the cost of a tail ornament in male birds: a negative relationship between an experimentally increased tail ornament (long tail streamers) and efficiency at foraging for nestlings. However, it must be admitted, that the apparent impairing effect of an elongated tail could be a result of a decrease in male parental effort in response to an increase of female parental effort, which might have occurred in response to increased male attractiveness (differential allocation of female parental effort). In this study, the effect of differential parental expenditure was eliminated by lengthening the tail in female, rather than male, sand martins ( Riparia riparia ). Tail-elongated females decreased the rate at which they fed nestlings, and captured more but smaller insects. There was no simultaneous increase of feeding rate in the males that could explain the decrease of feeding rate in the females. These results confirm the existence of a cost of a tail ornament in birds feeding in flight, as is expressed in terms of impaired flight and foraging capacity.     

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.     

Offspring sex ratios in relation to mutual ornamentation and extra-pair paternity in the Black Swan Cygnus atratus

In sexually dichromatic birds, females may adaptively adjust the sex ratio of their offspring prior to hatching in relation to male ornamentation, for example, by producing more sons when paired to a highly attractive partner. However, to our knowledge no studies have investigated offspring sex ratio modification in species in which both sexes are ornamented, and it is unknown whether such a process would be adaptive. Here we examine variation in offspring sex ratio in the mutually ornamented Black Swan Cygnus atratus . Brood sex ratio was not related to the degree of ornament elaboration in either parent, or to extra-pair paternity. We suggest that parental attractiveness may not be inherited in a sex-linked manner, or may be largely non-heritable. Thus, females may not benefit from biasing the sex ratio of their offspring in relation to parental attractiveness.     

Nestling sex ratio is associated with both male and female attractiveness in rock sparrows

According to theory, in species in which male variance in reproductive success exceeds that of the females, sons are more costly to produce; females mated with high quality males or those in better condition should produce more sons. In monogamous species, however, the variance in the reproductive success of the two sexes is often similar and mate choice is often mutual, making predictions regarding sex allocation more difficult. In the rock sparrow Petronia petronia, both males and females have a sexually selected yellow patch on the breast, whose size correlates with individual body condition. We investigated whether the brood sex ratio co‐varies with the size of the yellow patch of the father and the mother in a sample of 173 broods (818 chicks) over 8 breeding seasons. While the size of the yellow patch of the mother and the father did not predict per se a deviation from the expected 1:1 sex ratio, brood sex ratios were predicted by the interaction of male and female yellow patch size. This result is surprising, as the ornament is sexually selected by both males and females as an indicator of quality in both sexes and should therefore be inherited by all offspring irrespective of their sex. It indirectly suggests that other sex‐specific traits associated with patch size (e.g. polygyny in males and fecundity in females) may explain the sex allocation bias observed in rock sparrows. Thus, female individual quality alone, as expressed through the size of the yellow patch, was not associated with the biases in sex ratios reported in this study. Our results rather suggest that sex allocation occurs in response to male attractiveness in interaction with female attractiveness. In other words, females tend to preferentially allocate towards the sex of the parent with more developed ornament within the pair.     

Examining the Cost of Experimentally Imitated Ornaments

Long forked tail ornament in male barn swallows (Hirundo rustica) were suggested to impose a condition‐dependent viability cost ( Møller 1989 ; reviewed in Møller 1994 ; Møller & de Lope 1994 ; Møller et al. 1995 ): long tails impair male flight and foraging ability in terms of mean size of prey captured. In a recent study, we ( Matyjasiak et al. 1999 ) showed such a foraging cost of an experimentally imitated tail ornament in the sand martin (Riparia riparia), which have no tail ornament. We lengthened the tail in females, instead of in males, and thus controlled experimentally for the possible effect of differential allocation of female parental expenditure (differential‐allocation hypothesis, Burley 1986 ). Cuervo (2000) raises important issues in his comments about our paper. He questions the method used in our study ( Matyjasiak et al. 1999 ). He points out that ‘in order to test the cost of flight of an ornament, the trait to be experimentally manipulated has to be an ornament’, and that we should have not only elongated but also shortened tail feathers. Secondly, he suggests that we did not provide, in our article, the exclusive evidence for the handicap model of sexual selection. Thirdly, he disagrees with our suggestion that the results from the barn swallow experiments could be confounded by the differential allocation of female parental expenditure. Here, we outline the areas of agreement and disagreement between Cuervo's critique and our paper. Firstly, we agree with Cuervo regarding the importance of tail shortening in studies of fully developed tail ornaments, which has already been pointed out by other authors ( Thomas & Rowe 1997 ; Evans & Thomas 1997 ). However, shortening of an ornament that is at equilibrium would always produce a decrease in costs. Therefore it will only confirm that an ornament is costly. However, if shortening is done to individuals of various, known condition, it may result in crucial information for distinguishing between hypotheses of sexual selection. We agree that for a character that is not an ornament, as in our experiments, shortening would not give new insights apart from another way of confirming the measures used in the barn swallow studies. However, we disagree with Cuervo's claims of irrelevance of our experiment to the issues of the evolution of signals. We have chosen sand martins as a model species because the shape of its tail resembles that in ancestors of modern tail‐ornamented swallows, from which tail feather elongation under sexual selection may have started ( Matyjasiak et al. 2000 ). Possible scenarios of the early evolution of a tail ornament may be examined by experimentally adding such an ornament, which requires manipulating original, non‐ornamental traits (e.g. Goötmark 1994, 1996 ). We disagree with Cuervo that the existence of traits that may reduce the costs of ornaments (e.g. Møller 1996 ) eliminates the validity of tail manipulation studies in species without ornaments. We believe that such cost‐reducing traits may not have existed during the early stages of evolution of tail ornaments (as well as other sexual traits) but may have developed later, and our experiment may represent such a situation very well. Hence, by imitating the initial development of a forked tail ornament in sand martins, we performed a biologically relevant manipulation. Secondly, we do not state in our paper that we have tested or proved the handicap principle. We only mention in the last paragraph of the Discussion, that our results are consistent with this principle. However, we do admit that mentioning only the handicap hypothesis in the Introduction and Discussion only, and omitting other processes by which costly tail ornaments might arise, was unwarranted, as Cuervo properly argued; this might have left a false impression that our goal was to test the handicap hypothesis. We do believe that we have properly measured the costs of tail elongation, and this was our goal, as stated in the Abstract and in the last paragraph of the Introduction. Nowhere in the paper did we state that we actually aimed to test the differential costs of ornaments that are crucial to the handicap hypothesis. This issue was the objective of a different paper ( Matyjasiak et al. 2000 ) in which we consider conditions important for the early evolution of tail ornaments, with special emphasis on the handicap principle. Thirdly, we are more cautious at interpreting the barn swallow studies and we do not exclude a possibility that differential allocation of parental expenditure can affect the size of prey brought by males. Differential allocation does exist in this species ( Møller 1992 ; de Lope & Møller 1993 ), as shown by the higher feeding rate by females responding to male higher attractiveness (longer tails). We believe that even though simple logical, intuitive reasoning may suggest that prey size should be unaffected by a differential allocation mechanism, it is not just reasoning but empirical proof that is needed here. Because there was no proof for a lack of the effect of differential allocation on the size of prey, we thought of an independent way of illustrating the costs of tail elongation in swallows, using a species without possible differential allocation effects. We thought that if we obtained results confirming the barn swallow studies by Møller and collaborators, we could help in validating the measures of tail elongation costs used in those studies. This was possible using the sand martin females for reasons discussed in the paper, all of which point to the lack of any differential allocation effects in this species. From this point of view, experimentally coupling tail elongation with tail shortening in our study appears unnecessary. Even though, as argued by Cuervo (2000) , it seems likely that the prey size in male swallows is unaffected by a change in feeding rate of females in response to male attractiveness (an assumption inherent in the barn swallow studies), another scenario is also possible. Imagine a female that has increased the amount of food for nestlings, in response to increased sexual attractiveness of her male partner due to experimental elongation of his tail. In such a situation the male has been relieved from a considerable duty of providing the young with a large amount of food. Hence, it is possible that such a male shifts from maximising the energy brought to nestlings per unit time (i.e. maximising foraging rate) to the criterion of obtaining the daily energetic needs at the least expense (i.e. minimising foraging costs). Such shifts may lead to including some non‐preferred, small insects that can be captured quickly and inexpensively in terms of energy, because it does not require the use of expensive flapping flight, which is required to capture larger, preferred insects ( Waugh 1978 ; Bryant & Turner 1982 ; Turner 1980, 1982 ). By including more small insects in their catch at the expense of a reduced foraging rate, attractive males could save energy for future use. Barn swallows appear to compromise between maximising their foraging efficiency (maximising foraging gains per costs) and maximising energy intake per unit time (see fig. 5.7 in Turner 1980 and table IV in Turner 1982 ). Hence, if sexually attractive males aim at minimising foraging costs rather than maximising foraging rate for nestlings, then we cannot exclude the possibility that this may result in smaller insects being caught, on average, by males with experimentally longer tails. By a similar reasoning, it is theoretically possible that differential allocation effects could lead to larger mean prey size in males with experimentally shortened tails ‐ an effect actually shown in the barn swallow studies ( de Lope & Møller 1993 ; Møller & de Lope 1994 ; Møller et al. 1995 ). Hence, whether shortening, elongating or performing both experimental manipulations, in our view we cannot be entirely sure whether the results are affected by differential allocation. This was sufficient motivation for us to investigate, independently, the usefulness of the change in prey size as an indicator of foraging costs due to elongated tails. In contrast to Cuervo's opinion, we believe that our results are relevant to the issues important for the evolution of forked tail ornaments. We have measured the costs of a character that imitates the hypothetical early stages of the evolution of sexual ornaments, and we may use these results to discuss the early evolution of sexual ornaments (see Matyjasiak et al. 2000 ). We also confirmed the validity of measures of tail elongation costs used in the barn swallow studies.     

Incubation effort in relation to male attractiveness in zebra finches Taeniopygia guttata

The division of labour in parental care between the two sexes varies between and within species. In birds, parents have been shown to invest more into egg production and nestling care when paired with an attractive rather than an unattractive mate, as predicted by the differential allocation hypothesis. Here we investigate variation in the female's and male's share of incubation behaviour, a vital, and costly, period of parental care during which the embryo is vulnerable to perturbations in developmental conditions. We manipulated the attractiveness of male zebra finches Taeniopygia guttata , using red or green leg-rings. To simulate their natural social environment we allowed them to breed in outdoor aviaries. All males within an aviary were given the same coloured ring to avoid ring-colour related assortative mating. Males within a colony, however, were still expected to show some variation in attractiveness with the earliest laying females possibly pairing with the most attractive males. Indeed we found that both factors played a role in explaining female incubation effort. Among females mated to red ringed males, earlier laying females contributed significantly more to incubation than late laying females, but no such pattern was found in females mated to green ringed males. Overall, there were no differences in the level of incubation provided by both parents between treatment groups, suggesting some compensation within the pair. Hatching success was correlated with a pair's total incubation effort. These results suggest that variation in the division of parental care between the sexes is in agreement with both increased effort of females mated with attractive males, and females compensating for the reduced effort of attractive males seeking further mating opportunities. These two factors can act at the same time in natural populations and both should be considered when explaining variation in division of labour between the sexes.     

Parental care and UV coloration in blue tits: opposite correlations in males and females between provisioning rate and mate’s coloration

Parental investment and sexually‐selected signals can be intimately related, either because the signals indicate the amount of investment that an individual is prepared to make, and hence its value as a mate (the ‘good parent process’), or because individuals are selected to vary their own investment in relation to their mate’s signals (‘differential allocation’ or ‘reproductive compensation’). Correlations between parental investment and the sexually selected signals of both an individual and its mate are therefore of central interest in sexual selection. Blue tits Cyanistes caeruleus are an ideal study species to investigate such correlations because they provide substantial amounts of biparental care and possess sexually‐selected structural UV coloration that seems to signal attractiveness in both sexes. We investigated whether feeding rates of male and female blue tits were correlated with either their own or their mate’s UV coloration, and whether any such correlation was affected by the sex ratio of the brood. We also investigated whether any such correlations were reflected in offspring phenotype. Feeding rates were not correlated with either sex of parent’s own UV coloration. However, they were correlated with the mate’s UV coloration, but in opposite directions in males and females: females had higher feeding rates when mated to bright UV males, implying differential allocation, while males had lower feeding rates when mated to bright UV females, implying reproductive compensation. These relationships were unaffected by the sex ratio of the brood. In addition, fledgling tarsus length, but not mass, was related to male UV coloration, and to female UV coloration in interaction with male age. These results suggest that both male and female attractiveness influence parental investment of the mate, and that this in turn affects offspring phenotype. We found no evidence for differential sex allocation.     

Sex ratio influences the motivational salience of facial attractiveness

The sex ratio of the local population influences mating-related behaviours in many species. Recent experiments show that male-biased sex ratios increase the amount of financial resources men will invest in potential mates, suggesting that sex ratios influence allocation of mating effort in humans. To investigate this issue further, we tested for effects of cues to the sex ratio of the local population on the motivational salience of attractiveness in own-sex and opposite-sex faces. We did this using an effort-based key-press task, in which the motivational salience of facial attractiveness was assessed in samples of faces in which the ratio of male to female images was manipulated. The motivational salience of attractive opposite-sex, but not own-sex, faces was greater in the own-sex-biased (high competition for mates) than in the opposite-sex-biased (low competition for mates) condition. Moreover, this effect was not modulated by participant sex. These results present new evidence that sex ratio influences human mating-related behaviours. They also present the first evidence that the perceived sex ratio of the local population may modulate allocation of mating effort in women, as well as men.     

Offspring sex ratios correlate with pair-male condition in a cooperatively breeding fairy-wren

We examined sex allocation patterns in island and mainland populationsof cooperatively breeding white-winged fairy-wrens. The markeddifferences in social structure between island and mainlandpopulations, in addition to dramatic plumage variation amongmales both within and between populations, provided a uniquesituation in which we could investigate different predictionsfrom sex allocation theory in a single species. First, we testthe repayment (local resource enhancement) hypothesis by askingwhether females biased offspring sex ratios in relation to theassistance they derived from helpers. Second, we test the malequality (attractiveness) hypothesis, which suggests that femalesmated to attractive high-quality males should bias offspringsex ratios in favor of males. Finally, we test the idea thatfemales in good condition should bias offspring sex ratios towardmales because they are able to allocate more resources to offspring,whereas females in poor condition should have increased benefitsfrom producing more female offspring (Trivers-Willard hypothesis).We used molecular sexing techniques to assess total offspringsex ratios of 86 breeding pairs over 2 years. Both offspringand first brood sex ratios were correlated with the pair-male'sbody condition such that females increased the proportion ofmales in their brood in relation to the body condition (masscorrected for body size) of their social partner. This relationwas both significant and remarkably similar in both years ofour study and in both island and mainland populations. Althoughconfidence of paternity can be low in this and other fairy-wrenspecies, we show how this finding might be consistent with themale quality (attractiveness) hypothesis with respect to malecondition. There was no support for the repayment hypothesis;the presence of helpers had no effect on offspring sex ratios.There was weak support for both the male quality (attractiveness)hypothesis with respect to plumage color and the maternal conditionhypothesis, but their influence on offspring sex ratios wasnegligible after controlling for the effects of pair-male condition.     

Strategic female reproductive investment in response to male attractiveness in birds

Life-history theory predicts that individuals should adjust their reproductive effort according to the expected fitness returns on investment. Because sexually selected male traits should provide honest information about male genetic or phenotypic quality, females may invest more when paired with attractive males. However, there is substantial disagreement in the literature whether such differential allocation is a general pattern. Using a comparative meta-regression approach, we show that female birds generally invest more into reproduction when paired with attractive males, both in terms of egg size and number as well as food provisioning. However, whereas females of species with bi-parental care tend to primarily increase the number of eggs when paired with attractive males, females of species with female-only care produce larger, but not more, eggs. These patterns may reflect adaptive differences in female allocation strategies arising from variation in the signal content of sexually selected male traits between systems of parental care. In contrast to reproductive effort, female allocation of immune-stimulants, anti-oxidants and androgens to the egg yolk was not consistently increased when mated to attractive males, which probably reflects the context-dependent costs and benefits of those yolk compounds to females and offspring.     

Male rock sparrows adjust their breeding strategy according to female ornamentation: parental or mating investment?

We investigated the relations between female quality and ornamentation and between male breeding investment and female ornamentation in the rock sparrow, Petronia petronia, a passerine in which both sexes have a yellow breast patch. Breast patch size in females was positively correlated with body mass and breeding status; double-brooding and primary females of polygynous males had a larger patch, and patch size could therefore be an indicator of female phenotypic quality. We conducted a field experiment to test whether males allocate their parental effort in relation to female quality, as predicted by the differential allocation hypothesis. We increased and reduced the ornament sizes of paired females and compared the behaviour of their males before and after manipulation. Frequency of brood feeding by the male was not affected by female ornament manipulation; there was a nonsignificant trend for females with enlarged ornaments, contrary to predictions, to increase their feeding rate. Reducing female ornaments resulted in a decrease in male nest attendance, a measure of passive brood defence, whereas enlarging the ornament had no effect. Males concurrently reduced their territorial (song output) and sexual activity (courtship and copulation). The reduction in sexual activity suggests that males may have changed their nest attendance in response to their mate's renesting probability. Whatever the interpretation, these results provide some of the first evidence that not only female, but also male, birds change breeding strategy according to their mate's phenotype in the wild. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

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.     

Genetic association between male attractiveness and female differential allocation

Differential allocation of reproductive effort towards offspring of attractive mates is a form of post-copulatory mate choice. Although differential allocation has been demonstrated in many taxa, its evolutionary implications have received little attention. Theory predicts that mate choice will lead to a positive genetic correlation between female preference and male attractiveness. This prediction has been upheld for pre-copulatory mate choice, but whether such a relationship between male attractiveness and female differential allocation exists has never been tested. Here, we show that both female pre-copulatory mate choice and post-copulatory differential allocation are genetically associated with male attractiveness in house crickets, Acheta domesticus. Daughters of attractive males mated sooner and laid more eggs when paired with larger males. These forms of mate choice are strongest in large females, suggesting that costs decrease with increasing female size. The genetic association between attractiveness and differential allocation suggests potential for differential allocation to become exaggerated by coevolutionary runaway processes in an analogous manner to pre-copulatory choice. Sexual selection is thus likely to be stronger than predicted by pre-copulatory choice alone.     

Morphological and genetic predictors of parental care in the North American barn swallow Hirundo rustica erythrogaster

Sexually dimorphic traits often signal the fitness benefits an individual can provide to potential mates. In species with altricial young, these signals may also predict the level of parental care an individual is expected to provide to shared offspring. In this study, we tested three hypotheses that traditionally relate sexually dimorphic traits to parental care in two populations of North American barn swallows Hirundo rustica erythrogaster. The good parent hypothesis predicts a positive relationship between an individual's ornamentation and his or her care whereas the differential allocation (more care given by individuals when paired to high quality mates) and reproductive compensation (more care given by individuals when paired to low quality mates) hypotheses predict that an individual's level of parental investment is relative to the quality of their mate. Male and female North American barn swallows have colorful ventral feathers and elongated tail streamers, but there is evidence that ventral color, not tail streamer length, predicts measures of seasonal reproductive success. Accounting for the positive correlation between within‐pair feeding rates and other potentially confounding variables in all of our models, we found no support for the good parent hypothesis because in both males and females, traits shown to be under sexual selection did not predict feeding rates in either sex. However, our data reveal that male coloration, and not streamer length, predicted a female's provisioning rate to shared offspring (females fed more when paired with darker individuals) in two separate populations, supporting the differential allocation, but not the reproductive compensation hypothesis. Because genetic traits have also been shown to affect parental investment, we evaluated this variable as well and found that a male's paternity did not have significant effects on either male or female feeding rates. Overall, our results suggest that females do not pair with darker males in order to gain direct benefits in terms of his expected levels of parental care to shared offspring, but do themselves invest greater levels of care when paired to darker males. Further, our results are consistent with previous studies which suggest that ventral feather color, not streamer length, is a target of sexual selection in North American populations of barn swallow because females invested more in their offspring when paired to darker mates.     

Male-Biased Sex Ratios in Offspring of Attractive Males in the Guppy

The attractiveness hypothesis predicts that females produce offspring with male-biased sex ratios when they mate with attractive males because their male offspring will inherit the paternal sexual attractiveness and may have high reproductive success. In this study, we examined the effect of the attractiveness of the male guppy Poecilia reticulata in terms of the conspicuousness of its orange spot patterns, important criteria affecting female choice in this species, on the offspring sex ratios. We found that food-manipulation treatment altered the conspicuousness of the orange spot patterns in a full-sibling male pair. When females were presented to these males, they showed a greater mate preference for males having brighter orange spots than for those having duller orange spots. Subsequently, half of the females were mated with the preferred males and the remaining females were mated with the less preferred males. When the females exhibited a greater preference for their mates, their offspring sex ratios were more male biased. These results appear to be consistent with the prediction of the attractiveness hypothesis. In the guppy, as male sexual attractiveness is heritable, the male-biased sex ratios of the broods of attractive males may be adaptive.     

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 COMPLEX INTERPLAY OF SEX ALLOCATION AND SEXUAL SELECTION

It is well recognized that sex allocation strategies can be influenced by sexual selection, when females adjust offspring sex ratios in response to their mates’ attractiveness. Yet the reciprocal influence of strategic sex allocation on processes of sexual selection has only recently been revealed. Recent theoretical work demonstrates that sex allocation weakens selection for female preferences, leading to the decline of male traits. However, these results have been derived assuming that females have perfect knowledge of mate attractiveness and precise control over cost‐free allocation. Relaxing these assumptions highlights the importance of another feedback: that adaptive sex allocation must become difficult to maintain as traits and preferences decline. When sex allocation strategies erode not only traits and preferences but also their own selective advantage, predictions can no longer be expressed as a simple linear correlation between ornament exaggeration and adaptive sex allocation. Instead, strongest sex ratio biases may be found at intermediate trait levels.     

Endocrine correlates of intra-specific variation in the mating system of the St. Peter's fish (Sarotherodon galilaeus)

The Challenge Hypothesis postulates that androgen levels are a function of the social environment in which the individual is living. Thus, it is predicted that in polygynous males that engage in social interactions, androgen levels should be higher than in monogamous animals that engage in parental care. In this study, we tested this hypothesis at the intra-specific level using a teleost species, Sarotherodon galilaeus, which exhibits a wide variation in its mating system. Experimental groups of individually marked fish were formed in large ponds with different operational sex-ratios (OSR) to study the effects of partner availability on blood plasma levels of sex steroids [11-ketotestosterone (11-KT), testosterone (T), and 17,20beta-dihydroxy-4-pregnen-3-one (17,20beta-P)] and gonadosomatic index (GSI). Polygyny mostly occurred in the female biased OSR groups. 17,20beta-P and gonadosomatic index did not differ among OSR groups. However, 11-KT was high in male biased OSR and positively correlated with aggressive challenges, thereby supporting the central postulate of the Challenge Hypothesis. The results of T were the inverse of those of 11-KT, probably because 11-KT is metabolized from T. 11-KT levels of polygynous males did not differ neither from those of monogamous males, nor from those of males that participated in parental care. These results do not support the expected relationships between polygyny, parental care, and androgen levels. The differences from expectations for 11-KT may be related to the fact that in S. galilaeus, the mating and the parenting phase are not clearly separated and thus, males may still fight and court while they are brooding.     

Sex allocation and sex‐specific parental investment in an endangered seabird

Biased offspring sex ratio is relatively rare in birds and sex allocation can vary with environmental conditions, with the larger and more costly sex, which can be either the male or female depending on species, favoured during high food availability. Sex‐specific parental investment may lead to biased mortality and, coupled with unequal production of one sex, may result in biased adult sex ratio, with potential grave consequences on population stability. The African Penguin Spheniscus demersus, endemic to southern Africa, is an endangered monogamous seabird with bi‐parental care. Female adult African Penguins are smaller, have a higher foraging effort when breeding and higher mortality compared with adult males. In 2015, a year in which environmental conditions were favourable for breeding, African Penguin chick production on Bird Island, Algoa Bay, South Africa, was skewed towards males (1.5 males to 1 female). Males also had higher growth rates and fledging mass than females, with potentially higher post‐fledging survival. Female, but not male, parents had higher foraging effort and lower body condition with increasing number of male chicks in their brood, thereby revealing flexibility in their parental strategy, but also the costs of their investment in their current brood. The combination of male‐biased chick production and higher female mortality, possibly at the juvenile stage as a result of lower parental investment in female chicks, and/or at the adult stage as a result of higher parental investment, may contribute to a biased adult sex ratio (ASR) in this species. While further research during years of contrasting food availability is needed to confirm this trend, populations with male‐skewed ASRs have higher extinction risks and conservation strategies aiming to benefit female African Penguin might need to be developed.     

Comparative evidence supports a role for reproductive allocation in the evolution of female ornament diversity

1. Sexually selected ornaments are highly variable, even among closely related species, and the ultimate causes of variation in ornament evolution are unclear, including in rare cases of female ornament expression. One hypothesis is that differences across species in female reproductive allocation may help to explain patterns of female ornament expression among insects with nuptial gifts. 2. Dance flies (Diptera: Empididae: Empidinae) vary considerably among species in the presence and extravagance of female ornaments, which probably evolved through female contests for mates. In most dance flies, adult females appear to acquire all their dietary protein from nuptial gifts provided by males during mating. The importance of nuptial feeding on egg development is not yet known. 3. To test the prediction that the presence of female ornaments reflects differences in the degree to which females rely on nuptial feeding for egg development, egg development was examined in wild females of two species, one ornamented and the other unornamented. An ageing technique based on cuticular bands was validated, which permitted a regression of egg size on adult age. 4. We found that egg development depended on mating status in the ornamented species alone, meaning the eggs of unmated females of the ornamented species did not develop. This contrast across species is consistent with expectations that females of different species vary in their dependence on nuptial gifts for egg development. 5. These findings provide preliminary support for the hypothesis that differences in reproductive allocation mediate the intensity of female contests for nuptial gifts.