Evolution Of Female Ornaments Research Articles

Female ornamentation and the fecundity trade-off in a sex-role reversed pipefish.

Sexual ornaments found only in females are a rare occurrence in nature. One explanation for this is that female ornaments are costly to produce and maintain and, therefore, females must trade‐off resources related to reproduction to promote ornament expression. Here, we investigate whether a trade‐off exists between female ornamentation and fecundity in the sex‐role reversed, wide‐bodied pipefish, Stigmatopora nigra. We measured two components of the disk‐shaped, ventral‐striped female ornament, body width, and stripe thickness. After controlling for the influence of body size, we found no evidence of a cost of belly width or stripe thickness on female fecundity. Rather, females that have larger ornaments have higher fecundity and thus accurately advertise their reproductive value to males without incurring a cost to fecundity. We also investigated the relationship between female body size and egg size and found that larger females suffer a slight decrease in egg size and fecundity, although this decrease was independent of female ornamentation. More broadly, considered in light of similar findings in other taxa, lack of an apparent fecundity cost of ornamentation in female pipefish underscores the need to revisit theoretical assumptions concerning the evolution of female ornamentation.

Evolution of a mating preference for a dual-utility trait used in intrasexual competition in genetically monogamous populations.

The selection pressures by which mating preferences for ornamental traits can evolve in genetically monogamous mating systems remain understudied. Empirical evidence from several taxa supports the prevalence of dual‐utility traits, defined as traits used both as armaments in intersexual selection and ornaments in intrasexual selection, as well as the importance of intrasexual resource competition for the evolution of female ornamentation. Here, we study whether mating preferences for traits used in intrasexual resource competition can evolve under genetic monogamy. We find that a mating preference for a competitive trait can evolve and affect the evolution of the trait. The preference is more likely to persist when the fecundity benefit for mates of successful competitors is large and the aversion to unornamented potential mates is strong. The preference can persist for long periods or potentially permanently even when it incurs slight costs. Our results suggest that, when females use ornaments as signals in intrasexual resource competition, males can evolve mating preferences for those ornaments, illuminating both the evolution of female ornamentation and the evolution of male preferences for female ornaments in monogamous species.

Intrasexual competition and throat color evolution in female three-spined sticklebacks

Interest in the evolution of female ornaments has been growing but explaining their presence remains a challenge for many animal taxa. Although female ornaments may be sexually selected via male choice, they can also function in female–female competition. Here, we studied a stream-resident population of three-spined sticklebacks (Gasterosteus aculeatus), in which females possess male-typical red throat coloration, and a nearby anadromous population that lacks such coloration and is thought to possess ancestral character states for most traits in this species complex. We asked first whether there were population-level differences in agonism that might be associated with the evolution of female ornaments. We tested if the derived population possessing some red-throated females exhibited relatively higher levels and clearer patterns of within-population aggression and dominance in dyadic trials and more frequent behavioral dominance over individuals of the population lacking red females. Second, to address the relationship between the derived coloration and female–female competition, we tested whether red-throated stream females were more aggressive and dominant toward dull-throated stream females in both dyadic and socially complex contexts. We found that clear dominance was established more often in pairs of stream females, and stream females were more often dominant in stream-anadromous dyads. However, we found no clear evidence that the red throat coloration was directly linked to a female competitive advantage within the stream population, suggesting alternative untested signaling or perhaps nonadaptive functions. Our study represents the first evaluation of the potential relationship between a male-typical nuptial trait and intrasexual competition in female three-spined sticklebacks.

Reversible Trait Loss: The Genetic Architecture of Female Ornaments

The evolution of female ornamentation has been subject to considerable debate. Although originally thought to be by-products of selection on males, recent studies have emphasized the adaptive nature of female ornaments. Here, I review current knowledge on the genetic architecture of ornamental traits and how their expression is (or is not) restricted to one sex. Ornament development typically involves deeply conserved developmental genes. Their expression is modified by pre-existing sexually dimorphic signals, such as hormones in vertebrates or doublesex gene products in insects. Changes in dosage and binding affinities in these systems apparently allow frequent switches between sexually dimorphic and monomorphic ornament expression. However, early-developing ornaments may be difficult to modulate, as this would result in severe pleiotropic effects. Comparative studies on the molecular structure of the sex-specific modulating signals and their receptors should provide detailed insight into the mechanisms underlying the loss and gain of female ornaments.

Wake up and smell the conflict: odour signals in female competition.

Odour signals used in competitive and aggressive interactions between males are well studied in the context of sexual selection. By contrast, relatively little is known about comparable signals used by females, despite current interest in the evolution of female ornaments and weaponry. Available evidence suggests that odour signals are important in competitive interactions between female mammals, with reductions or reversals of male-biased sexual dimorphism in signalling where female competition is intense. Scent marking is often associated with conflict between females over access to resources or reproductive opportunities. Female scent marks may therefore provide reliable signals of competitive ability that could be used both by competitors and potential mates. Consistent with this hypothesis, we report that aggressive behaviour of female house mice is correlated with the amount of major urinary protein (MUP) excreted in their urine, a polymorphic set of proteins that are used in scent mark signalling. Under semi-natural conditions, females with high MUP output are more likely to produce offspring sired by males that have high reproductive success, and less likely to produce offspring by multiple different sires, suggesting that females with strong MUP signals are monopolized by males of particularly high quality. We conclude that odour signals are worthy of more detailed investigation as mediators of female competition.

Female ornaments revisited – are they correlated with offspring quality?

The evolution and signalling content of female ornamentation has remained an enduring challenge to evolutionary biologists, despite the fact that secondary sexual characters are widespread in females. While females usually invest significant amounts of their resources, including carotenoids, in offspring, all the resources allocated to elaborate ornamentation reduce resources available for other purposes. This may in turn constrain female fitness leading to dishonest female signalling. We review the literature for empirical studies on mutually ornamented species with conventional sex roles, by focusing on the association between female ornaments and quality of their offspring. We found 43 papers where 33 (77%) are bird-studies, nine (21%) are on fishes, and one (2%) is a lizard-study. Nine of these report negative, 14 non-existing, and 20 positive associations between female ornament and offspring quality. Eighteen of the bird studies (55%) show a positive association between the two traits investigated, whereas five (15%) of the studies report a negative association. The number of fish studies, although few, is skewed in the opposite direction with two (22%) and four (44%) studies supporting positive and negative association, respectively. A minority of studies on carotenoids-based ornaments reports a positive association (4 of 18 studies, or 22%) between the traits, which is low compared to studies on non-carotenoids-based ornaments (16 of 25 studies, or 64%). The above-mentioned relative large number of studies with negative association, especially common in studies on fishes and in carotenoids-based-ornaments, challenges the generality of the direct selection hypothesis to account for female fineries. This is important because this hypothesis seems to have strong support in recent literature on the topic. In the present paper, we also propose possible explanations for the observed differences between taxa and suggest directions and ideas for future research on the evolution of female ornamentation.

The evolution of female ornaments and weaponry: social selection, sexual selection and ecological competition.

Ornaments, weapons and aggressive behaviours may evolve in female animals by mate choice and intrasexual competition for mating opportunities-the standard forms of sexual selection in males. However, a growing body of evidence suggests that selection tends to operate in different ways in males and females, with female traits more often mediating competition for ecological resources, rather than mate acquisition. Two main solutions have been proposed to accommodate this disparity. One is to expand the concept of sexual selection to include all mechanisms related to fecundity; another is to adopt an alternative conceptual framework-the theory of social selection-in which sexual selection is one component of a more general form of selection resulting from all social interactions. In this study, we summarize the history of the debate about female ornaments and weapons, and discuss potential resolutions. We review the components of fitness driving ornamentation in a wide range of systems, and show that selection often falls outside the limits of traditional sexual selection theory, particularly in females. We conclude that the evolution of these traits in both sexes is best understood within the unifying framework of social selection.

Safe betting: males help dull females only when they raise high-quality offspring

In evolutionary biology, whether parents should enhance or reduce parental care according to mate ornamentation is a subject of great debate. However, the evolution of female ornaments can shed light on this question. In theory, female ornamentation should be traded off against fecundity and thus cannot be wholly informative to males without a direct indication of fecundity. Hence, direct cues of offspring quality should affect the relationship between male investment and female ornamentation. Under this hypothesis, we manipulated two direct cues of offspring quality (egg size and color) after first egg laying in the blue-footed booby and registered male incubation patterns. In this species, foot color is a dynamic signal of current condition and in females is traded off with egg size. We found that males spent more time incubating when paired with dull females but only in nests with large eggs. Males also spent less time incubating small dull eggs. Results indicate that egg size, a direct cue of reproductive value, affected the relationship between male effort and female ornamentation. Males may be willing to help females that have invested in offspring at the expense of ornamentation, which suggests compensation when females are in low condition. Another possibility is that males relax their effort when paired with highly ornamented and fecund females because they have high parenting abilities. Our findings suggest that the information conveyed by female ornaments may depend on direct cues of fecundity. Results also highlight that parental decisions are complex, modulated by a combination of information sources.

Male, but not female, preference for an ornament expressed in both sexes of the polygynous mosquito Sabethes cyaneus

The evolution of female ornaments in species with a female-biased operational sex ratio (OSR) and intense female competition is well understood. In contrast, the adaptive value of female ornaments in species with a male-biased OSR and male competition remains largely unresolved. Mutual mate choice is one proposed explanation for the evolution of ornaments expressed in both males and females, a hypothesis supported by the increasing empirical evidence of mutual mate choice in species with a male-biased OSR. None the less, the evolution of female ornaments remains constrained, as investment in ornaments may detract from any direct benefits being signalled to males and females may fail to reap benefits sufficient to outweigh the costs of signalling. We used phenotypic engineering (i.e. manipulation of ornament size) to ask whether both sexes show a preference for sexually homologous ornaments in the polygynous mosquito, Sabethes cyaneus . We found a directional male preference for ornamented females, but no female preference for ornamented males. There was no evidence of assortative mating based on ornament size. We discuss these results within the framework of current sexual selection theory, addressing implications for both the evolution of male mate choice and the evolution of female ornaments.

Female ornamentation, parental quality, and competitive ability in the rock sparrow

The evolution of female ornaments is poorly understood. Recent evidence suggests not only that female ornaments may be genetic correlates of selection on males but may also have evolved through male mate choice and/or through female–female aggressive interactions. In the rock sparrow, Petronia petronia, both sexes have a carotenoid-based yellow patch that is sexually selected by both sexes. The benefits that male may gain from choosing an attractive female remain unidentified. Both parents participate in caring for the young, so there should be mutual mate choice because males and females should both benefit from choosing a good parent (good parent hypothesis; GPH). Moreover, it has already been demonstrated that the yellow patch in males is also a badge of status (armament). Therefore, the yellow patch could also serve as both ornament and armament in females (dual utility hypothesis; DUH). We investigated the hypothesis that male and female yellow patch size signals parental quality in the field. We tested by an experiment in captivity the signal function of the yellow patch in female–female aggressive interactions for access to food. Yellow patch size correlated with paternal, but not maternal, feeding rates. Thus, this study supports the hypothesis that yellow patch dimension signals male parental quality, but there is no evidence for the GPH to explain female ornamentation. In the experiment females with relatively large yellow patches had earlier access to food than those with small patches. These results seem to suggest that a sexually selected carotenoid-feather signal may be used in female–female competition, in agreement with the DUH. Males may benefit from choosing well ornamented females because these may be superior competitors.

Fishes as models in studies of sexual selection and parental care

Fishes are by far the most diverse group of vertebrates. This fact is in no way, however, reflected in their use as model organisms for understanding sexual selection or parental care. Why is this so? Is it because fishes are actually poor models? The usefulness of fishes as models for sexual selection and parental care is discussed by emphasizing some problems inherent in fish studies, along with a number of reasons why fishes are indeed excellently suited. The pros and cons of fishes as models are discussed mainly by comparison with birds, the most popular model organisms in animal behaviour. Difficulties include a lack of background knowledge for many species, and the problems of marking and observing fishes in their natural environment. Positive attributes include the diversity of lifestyles among fishes, and the ease with which they can be studied experimentally in the laboratory. How useful fish models can be is briefly illustrated by the impressive and broadly relevant advances derived from studies of guppies Poecilia reticulata and three‐spined sticklebacks Gasterosteus aculeatus. A selection of topics is highlighted where fish studies have either advanced or could greatly enhance, the understanding of processes fundamental to animal reproductive dynamics. Such topics include sex role dynamics, the evolution of female ornamentation and mate choice copying. Finally, a number of potential pitfalls in the future use of fish as models for sexual selection and parental care are discussed. Researchers interested in these issues are recommended to make much more extensive use of fish models, but also to adopt a wider range of models among fishes.