Mature sex roll reversal-Wild sex: when sex roles get reversed, some females develop a 'penis'

Our understanding of sexual selection has greatly improved during the last decades. The focus is no longer solely on males, but also on how female competition and male mate choice shape ornamentation and other sexually selected traits in females. At the same time, the focus has shifted from documenting sexual selection to exploring variation and spatiotemporal dynamics of sexual selection, and their evolutionary consequences. Here, I review insights from a model system with exceptionally dynamic sexual selection, the two-spotted goby fish Gobiusculus flavescens. The species displays a complete reversal of sex roles over a 3-month breeding season.

Mature sex roll reversal

Mature sex roll reversal

Thomas, G. Yasukawa Sed, Searcy WA, By contrast, realized sexual selection expresses the degree to which specific traits e. The sex experiencing the strongest mating competition would be expected to Mature sex roll reversal courtship more often. This is likely to be mostly true in many species, but is less likely in mutual choice systems. Empirical work on birds have largely supported best-of-n models, as females have often been found to revisit and mate with previously se males. Mesquite: a modular system for evolutionary analysis v. Sex-role reversal in vertebrates: behavioural and endocrinological accounts.

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Help us improve our products. Sign up to take part. A Nature Research Journal. Sex-role reversal represents a formidable challenge for evolutionary biologists, since it is not clear which ecological, life-history or social factors facilitated conventional sex roles female care and male-male competition for mates to be reversed male care and female-female competition.

Classic theories suggested ecological or life-history predictors of role reversal, but most studies failed to support these hypotheses. Recent theory however predicts that sex-role reversal should be driven by male-biased adult sex ratio ASR. Here we test this prediction for the first time using phylogenetic comparative analyses. Consistent with theory, both mating system and parental care are strongly related to ASR in shorebirds: conventional sex roles are exhibited by species with female-biased ASR, whereas sex-role reversal is associated with male-biased ASR.

These results suggest that social environment has a strong influence on breeding systems and therefore revealing the causes of ASR variation in wild populations is essential for understanding sex role evolution. Access provided by.

One of the fundamental patterns in animal social behaviour is that females tend to be the caring sex, whereas males compete for access to females 1 , 2 , 3. Our understanding of what determines these conventional sex roles is challenged by the reversal of sex roles in a number of organisms: the males contribute more to care than females, whereas the females compete for males 1 , 2 , 4.

In sex-role reversed species, the females are often larger and more ornamented than males, whereas the males may have specific adaptations for caring for eggs and young 2 , 4 , 5. Sex-role reversal is taxonomically widespread occurring in insects, fishes, amphibia and birds 1 , 4.

Sex-role reversal has been a formidable puzzle for evolutionary biologists ever since Darwin 6 , because it is not clear why males under some circumstances provide most or all parental care, and why competition for mates should be stronger among females than among males 1 , 2 , 7 , 8.

Previous hypotheses of sex-role reversal focused on specific ecological and life-history characteristics, such as temporal and spatial variation in food resources, offspring predation and breeding dispersal 1 , 9. Empirical evaluations, however, almost uniformly rejected these hypotheses 1 , 9 , 10 , Indeed, the life histories and ecology of sex-role reversed species are so diverse that it is hard to imagine common environmental circumstances that have led to the evolution and maintenance of sex-role reversal.

Species with reversed and conventional sex roles may breed side-by-side sharing much of the environment. Examples include habitats as diverse as the Arctic tundra phalaropes Phalaropus spp. Higher potential reproductive rates of females have been shown to correlate with more intense mating competition among females in species where only males care for the offspring 8 , although this relationship does not reveal the ecological, life-history or social predictors that have facilitated the evolution of male care in the first place.

Recent theoretical models put breeding-system evolution in a different perspective by showing that ASR expressed here as the proportion of adult males in the adult population has a major influence on mating competition, mating systems and parental behaviour 14 , These models predict that the rarer sex is under selection to provide less care; for instance, male-biased ASR should facilitate male-biased parental care henceforth, male care and thus reversal of conventional parental roles, whereas female-biased ASR is predicted to favour female-biased care henceforth, female care 14 , Evolutionary changes in mating and parental behaviour are predicted to respond to ASR because if there are substantially more males in the population than females, males have low chances of finding a new mate.

Under such circumstances, the best strategy for a male may be to provide care for the offspring, rather than desert the female after copulation and face stiff competition in acquiring a new mate.

Given that the male cares and the ASR is male-biased, the females can desert the brood and acquire new mates. Testing these predictions in wild populations, however, has been challenging. The predictions are difficult to test in a single species, because most species do not exhibit sufficient variation in sex roles and ASR, although one component of sex roles, female social mating system, has been shown to correlate with ASR in dunnock Prunella modularis A multi-species comparative approach is needed, in which the variation in sex roles is compared across a set of species that differs in ASRs.

However, such tests have to date been limited by the lack of data on ASR, mating system and parental care from a group of organisms that exhibit both reversed and conventional sex roles. Here we provide the first evidence that ASR correlates with parental care and social mating system consistently with the theoretical predictions using shorebirds Scolopaci and Charadrii, sandpipers, plovers and allies.

Shorebirds are eminently suitable for testing theoretical predictions of breeding-system evolution, because they exhibit unusual diversity in mating system and parental care, including some of the textbook examples of sex-role reversal 1 , 2 , We carried out a comprehensive search in primary publications, reference books and online resources for data on ASR, social mating system and parental care, with special attention to species that have been reported to exhibit sex-role reversal.

Although data on ASR from wild populations are difficult to obtain 19 , the information now available for shorebirds permits tests of the theoretical predictions using statistically robust sample sizes.

ASR is significantly associated with social mating system: sex-role reversed species like most jacanas Jacanidae and greater painted-snipe Rostratula benghalensis that exhibit female polygamy and female—female competition for mates typically have strongly male-biased ASR, whereas species with male polygamy such as Northern lapwing Vanellus vanellus and ruff Philomachus pugnax have female-biased ASR.

The relationships between social mating system and ASR are significant when we use polygamy frequencies Fig. Consistent with theoretical expectations, ASR also correlates with the relative contribution of sexes to parental care, because male care is associated with male-biased ASR Fig. In addition, differences in the duration of care provided by males and females, another proxy for parental roles, are also significantly related to ASR Fig.

These results are not sensitive to a specific phylogenetic hypothesis, or potentially confounding variables. The aforementioned results are highly consistent between alternative phylogenetic hypotheses and different branch-length assumptions: the four key tests remain highly significant by using randomly selected trees from the most recent avian phylogeny 20 Supplementary Fig.

S1 , or using alternative phylogenies of shorebirds Supplementary Table S1. We ascertained whether the genetic mating system of shorebirds may confound the relationships between ASR, social mating system and care. We also tested whether breeding density, the only ecological correlate of male care demonstrated previously 21 , could influence the mating system, parental care and ASR relationships.

However, ASR remains strongly associated with both mating system and parental care when breeding density is added to the models Supplementary Table S2.

ASR has been estimated using different methods in the field see Methods , and we tested whether different estimation methodology may have biased the results. Nevertheless, by splitting the analyses into two subsamples either using direct counts of breeding birds or using ASRs estimated by all other methods, see Methods both effect sizes and the direction of relationships remain consistent with those for the whole species set.

Furthermore, different detectability of the sexes, a potential confound of field estimates of ASR 19 , is not likely to bias our results: the more polygamous sex is expected to be more conspicuous because of elaborate plumage, displays and general activity 2 , which would potentially bias ASR estimates towards the direction opposite to our findings that is, biasing ASR estimates towards the polygamous sex. The relationships between mating system, parental care and ASR may be due to changes in behaviour of males, females or both sexes.

We investigated these propositions by focusing on the behaviour of males and females in separate analyses. Intriguingly, the behaviour of both sexes responds to variation in ASR, because male-biased ASRs are associated with female polygamy and male care, whereas female-biased ASRs are associated with male polygamy and female care Fig.

Our results also reveal that both male and female behaviour show evolutionary responses to ASR, suggesting evolutionary flexibility in both mating and parental behaviour in both sexes. We propose that the evolutionary flexibility of both sexes to provide full care on their own and variation in ASR among species are among the key factors that facilitate the evolution of diverse sex roles 11 , Although in this paper we focused on sex-role reversal, our results also show that ASR is related to sex roles in general: it is associated with mating and parental behaviour through the whole range of avian sex roles, from conventional to role reversed.

We conjecture that ASR may influence other aspects of social behaviour. For example, in populations with biased sex ratios homosexual pairings may be more common, and biased sex ratios may also lead to cooperative breeding where the more common sex in the population postpones dispersal, stay in the family and provide help. Further studies are needed to identify why ASR is variable across species.

Biased ASRs may arise in several ways: there may be a bias in the primary sex ratio that is, sex ratio at conception , or males and females may have differential survival during development and maturation, or as adults. Recent studies suggest that offspring sex ratio at hatching is approximately in many birds 25 , therefore sex biases are likely to emerge after birth.

It is important to emphasize that mating behaviour, parenting and sex ratios may have more dynamic relationships than currently acknowledged 15 , 26 , First, ASR can affect sex roles see above , and conversely, reproductive behaviours can also influence mortalities and thus ASR. On the one hand, if mortality from care provisioning is high in a population with male-biased or female-biased care, this would reduce the extent of ASR bias in the population. On the other hand, if sexual selection is costly, then this may generate a positive feedback between ASR and sex roles, so that ASR may shift towards more extreme bias It is conceivable that populations can be locked in an unusual breeding system, because it is the best response to a biased ASR as generated by the breeding system itself.

Intense sexual competition and care provisioning have substantial energetic and mortality costs 29 , 30 , and thus likely that ASR and sex roles can evolve quickly and concurrently in ecological time scales, rather than in a sequential manner over evolutionary time scales for example, changes in ASR precedes changes in sex roles or vice versa. We propose that these relationships have a complex dynamics, and the dynamics itself may contribute to the immense diversity of sex roles and breeding systems in nature.

Operational sex ratio OSR, the ratio of sexually active males to receptive females is often used in the same context as ASR, although it has been suggested that this is mistaken OSR is only equal to ASR if the sexually active periods of adult males are identical with those of adult females. A significance of our present analyses is therefore to point out that a demographic property, the ratio of adult males and females, is closely correlated with mating and parenting behaviour in wild populations.

ASR on its own, however, is unlikely to explain all subtle variation in mating system and parenting of animals, because these may also depend on a suite of other factors.

We propose two further lines of studies to investigate the influence of ASR on sex roles. First, taxa with variable sex roles for example, pipefish Syngnathidae, poison dart frogs Dendrobatidae and tinamous Tinamidae 4 , 5 , 8 are ideal groups to separate the effects of phylogenetic history, ASR, life history and ecological traits on sex-role reversal: ASR may predict sex roles in these organisms once ecology and life-history differences have been controlled for.

Second, experiments are needed to manipulate ASR and investigate the corresponding changes in sex roles. Although ASR has been manipulated in the lab, experiments in natural populations, preferably in species with flexible sex roles, are required. We calculated ASR as the ratio of adult males to all adults males plus females in the populations.

When several estimates were available for a species, we used their mean value. In intensively studied breeding populations, ASR was often based on censuses of individually marked breeding adults. From the non-breeding period, we only included data if the ASR estimates were consistent among studies 31 , 32 , For 14 species, ASR data were taken from the original source, whereas for an additional 4 species, ASR was calculated using the data from the original sources.

By restricting the analyses to the former 14 species, our results do not change qualitatively Supplementary Table S4. In two species Jacana spinosa and Metopidius indicus , separate estimates were available for i breeding birds and ii breeders plus non-breeders; we repeated the analyses using both sets of data and the results remained highly consistent Supplementary Table S4.

We aimed at obtaining ASR for as many shorebird species as possible, including both sex-role reversed and non-reversed species. In the main analyses Fig. All data and references are provided in Supplementary Tables S5 and S6. We used two variables to describe social mating systems. First, we recorded the percentages of socially polygamous individuals separately for males and females 30 , using reference works and primarily literature Supplementary Tables S5 and S6.

Both simultaneous and sequential polygamy were included for both sexes, and if both types of social polygamy occurred within a sex, we used their sum. If several estimates of polygamy were reported for a species, we used their mean. We considered males or females monogamous if social polygamy was not reported for the given sex.

Lekking birds two species P. We did not find data on polygamy frequency for two species Charadrius nivosus and Rostratula benghalensis , so the maximal sample size for mating system bias tests is 16 species. Second, we also used mating system scores as a proxy variable of social mating systems for two reasons: i these scores are robust to observer errors in frequency estimates and ii to include the two species in the analyses see above that did not have frequency data available.

For C. Mating score bias was then calculated as the difference between the male and female scores. We used two variables to estimate the role of the sexes in care provisioning. First, we scored the participation of males on a five-point scale 0—4 for five types of parental behaviour: nest building, incubation, nest guarding guarding and defending the nest during incubation , chick brooding and chick guarding guarding and defending of the brood after hatching 30 ,

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Mature sex roll reversal

Mature sex roll reversal

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Sex roles and sexual selection: lessons from a dynamic model system

In the pipefish Syngnathus typhle sex roles are reversed, that is, females compete more intensely than males over mates. However, competition over mates among individuals of one sex does not necessarily prevent members of that same sex from being choosy, and choosiness in the other sex does not prevent competition within it. In an experiment we allowed a female pipefish to choose freely between two males, after which we released the males and let the three interact.

Comparisons with earlier results show that both sexes courted partners and competed with consexuals. However, females courted more often than did males, and courtship was more frequent in treatments involving large individuals than in treatments with small individuals.

Males competed among themselves for access to mates but for a shorter duration than females in the same situation. Males displayed an ornament towards females but not to males during mating competition. Females, however, used their ornament in both contexts. Females did not always mate with the male of their previously made choice, which we interpret as females being constrained by male-male competition, male motivation to mate, or both.

Thus, in this sex-role reversed species, mate choice in the more competitive sex may be circumvented and even overruled by mate competition and mating willingness in the least competitive sex. Hence, sex roles should not be considered as sexes being either choosy or competitive but rather that males and females may exhibit different combinations of choice and competition.

Mate choice and competition over mates are well-studied processes causing sexual selection Andersson, Usually, males compete among themselves over access to females, which are in short supply due to their lower potential reproductive rate. Females are typically more choosy than males when selecting a mate from the pool of available partners. In some species, however, the pattern for various reasons is the opposite: the parental investment has evolved to burden males more than females, decreasing male potential reproductive rate below that of females.

These species have reversed sex roles, where females compete more intensely over mates Berglund et al. One such species is the deep-snouted pipefish, Syngnathus typhle L. In nature, sex roles may commonly be less straightforward: it is well known that females in many species with conventional sex roles, while being choosy, may very well also compete with other females over males or mating status. For instance, in pied flycatchers and great tits, females may compete with other females to remain monogamously mated Breiehagen and Slagsvold, ; Slagsvold, In the blue tit, a female-biased sex ratio may cause severe competition for breeding opportunities among females Kempenaers, In dunnocks, females may use song to compete for males Langmore and Davies, , and in the alpine accentor, females attract males by song Langmore et al.

Furthermore, in red-winged blackbirds female-female aggression may ensure paternal investment Yasukawa and Searcy, , and in lekking topis, female compete over central males Bro-Jorgensen, , Consequently, females may show adaptations aiding in such competition, such as various displays e. Moreover males, while still being the more competitive sex, may also be choosy when given the opportunity e.

Furthermore, sex roles may change over the season in response to changes in mating competition Forsgren et al. However, even if conventional sex roles thus may be somewhat modified, there is yet no demonstration of such modifications in species with reversed sex roles.

Are the forces producing sex-role reversal so strong that such modifications do not exist? Mutual mate choice is expected to occur in species where potential reproductive rates of the two sexes are more similar Owens and Thompson, and mutual choice may have consequences for selection pressures on both males and females Bergstrom and Real, ; Deutsch and Reynolds, ; Johnstone, ; Johnstone et al. Mutual mate choice has been demonstrated in several species e.

Individual behaviors may be governed by factors beyond the individual's control, such as by competition from others, and sometimes an individual may even be forced to act in conflict with its own interest due to, for instance, manipulation by the other sex. For example, individuals in good condition may be better competitors and are able to pursue their interest to a larger extent than low-quality individuals. In the deep-snouted pipefish, females compete intensely over mates Berglund and Rosenqvist, b ; Vincent et al.

Furthermore, males, while preferring larger over smaller Berglund et al. Moreover, mate choice decisions are adjustable to circumstances: male choice in S. Both sexes benefit directly from mating with larger individuals of the opposite sex, so larger size means higher quality in terms of reproductive returns in both males and females of this species Berglund et al.

In this paper, we report a study of male-male competition and female choice in a species with reversed sex roles. We did this by investigating mutual mate choice and mating competition in S. We also investigated whether individual quality here body size affected choice and competition. For S. After females had chosen between two males in separate compartments, we released all three fish into the same compartment and recorded matings. A comparison with an earlier published, almost identical experiment on male choice allowed us to discuss differences between the sexes in this respect.

Female S. Males will compete for access to females but less so than females placed in the same situation: this prediction stems from the same logic as in 1 above. Courtship will be more intense in treatments involving large individuals than in treatments involving small individuals: if courtship reveals individual quality, large individuals should be more eager to display their superior quality.

Large males will engage more in competition than small males: large individuals are usually stronger than small individuals and thus may have more to gain from competitive interactions. We ran the female choice experiment two consecutive summers, from 20 May to 8 June in and from 26 May to 18 June in Stock tanks contained plastic plants and continuously renewed seawater temperature, salinity, and light regime following natural conditions.

The fish were fed live brine shrimps Artemia , small, wild-caught crustaceans, and frozen mysids ad lib twice daily. Each of 73 trials was run for 8 h, with one female choosing between two enclosed males during the first 4 h. After that, the enclosures were removed to allow male-male competition and actual mating to occur for the rest of the time. Enclosed males could not see or smell each other but both could see the female. The female could see and smell the two males as water flowed from the male compartments into the female compartment and then out of the aquarium.

The aquaria were continuously provided with seawater. Temperature and light followed natural conditions. We planted fresh eelgrass Zostera marina L.

Fish were not fed during trials. After each trial, we scored the degree to which the male's brood pouch was filled with eggs and gently removed and counted them. After recovery, a quick process, males as well as females were released back into the wild. No mortality occurred during trials. We chose males deliberately to differ in overall color green versus brown so that we could recognize individuals.

Brown males did not appear darker than green ones, so color did not interfere with our estimate of contrast or ornament display. New males and females were used in each replicate.

We have previously shown that color does not affect male choice of females Berglund and Rosenqvist, a , b. We ran two treatments, one involving only large fish and the other involving only small fish. All males had fully developed brood pouches. For females, size in the large treatment was Intense light a W spot and a 20 W luminescent lamp placed 0.

Video scoring was done by assistants naive to the purpose of the study. During video analysis we measured the time the female spent in front of each male's compartment; the time the males and the female were resting, swimming, or dancing; and the time a male or a female displayed the ornament.

A male and a female were recorded as dancing when they simultaneously bobbed up and down in close proximity while facing one another. We considered males to be competing when they pursued each other in the aquarium for more than a minute. We used the Observer 4. In trials where no dancing or copulation occurred, males were assigned a value of 4 h latency for dancing and copulation, respectively, that is, the time a trial took.

During video analysis a few tapes could not be scored, so the N values reported may be lower than the total number of replicates. Statistical probabilities reported from these experiments are two tailed.

Nonparametric tests were used whenever the assumptions of parametric tests were not met. We tested for evidence that females showed preferences for particular males.

If they did not, and distributed themselves randomly between the two compartments, then the difference in the time spent in front of the left side minus the time spent in front of the right would be normally distributed around a mean of zero. If females did exhibit preferences, then the distribution of differences should have been bimodal, with the absolute values of those differences consequently being normally distributed see Rowe et al.

Thus, females did not distribute themselves randomly before the two males, most likely due to female choice for a particular male. Within replicates, males displayed their ornament much less than did the female. While enclosed, females displayed on average for 2. When set free, females displayed for Number of replicates with a ornament displays or no displays by enclosed individuals to either potential partner in left: a previously made male choice experiment Berglund and Rosenqvist, a , using a similar design as in this study but letting a focal male choose between two object females, and right: the female choice experiment reported here b intrasexual aggression or no aggression in the form of chasing between free consexuals in the male left and female right choice experiments.

A comparison between a previous male choice experiment Berglund and Rosenqvist, a , where a focal male could choose between two object females, and the female choice experiment reported here, where a focal female could choose between two object males. The number of replicates where specific events occurred is shown.

The focal fish is the choosing fish in front, the object fish are the two chosen same-sex fish. A Fisher's exact probability test was used where expected frequencies were too small to allow a chi-square test. Only large males displayed an ornament, never small ones. When the partitions in the aquaria were removed and all interactions between individuals were allowed, males who displayed their ornament directed their display only towards the female during courtship and not to the other male during male-male competition except once for a few seconds only.

Large and small males did not differ in the duration of chases large males: 7. The male preferred by the focal female while enclosed did not always succeed i. However, a more powerful analysis reveals that the choice made by the enclosed females predicted which male she subsequently got to mate or dance with in the large but not in the small treatment Figure 2.

Enclosed males that were first chosen by the female and also successful when free tended to be larger than males first chosen but unsuccessful later on The fate of chosen and rejected males when set free in the small top and large bottom treatments. Success is the proportion of males that participated in dances or matings when free, where each male could score either 1 success, i. Average square with standard error box and standard deviation whisker shown. These and previous results Berglund and Rosenqvist, a show that in the sex-role reversed pipefish, S.

However, males compete less than females, and, while displaying an ornament towards females, males never use this signal in mating competition. Females, however, use their ornament in both contexts.

Mature sex roll reversal

Mature sex roll reversal

Mature sex roll reversal