Testosterone,growth and the evolution of sexual size dimorphism

The integration of macroevolutionary pattern with developmental mechanism presents an outstanding challenge for studies of phenotypic evolution. Here, we use a combination of experimental and comparative data to test whether evolutionary shifts in the direction of sexual size dimorphism (SSD) correspond to underlying changes in the endocrine regulation of growth. First, we combine captive breeding studies with mark‐recapture data to show that male‐biased SSD develops in the brown anole lizard (Anolis sagrei) because males grow significantly faster than females as juveniles and adults. We then use castration surgeries and testosterone implants to show that castration inhibits, and testosterone stimulates, male growth. We conclude by reviewing published testosterone manipulations in other squamate reptiles in the context of evolutionary patterns in SSD. Collectively, these studies reveal that the evolution of SSD has been accompanied by underlying changes in the effect of testosterone on male growth, potentially facilitating the rapid evolution of SSD.     

Life-history theory, fertility and reproductive success in humans

According to life-history theory, any organism that maximizes fitness will face a trade-off between female fertility and offspring survivorship. This trade-off has been demonstrated in a variety of species, but explicit tests in humans have found a positive linear relationship between fitness and fertility. The failure to demonstrate a maximum beyond which additional births cease to enhance fitness is potentially at odds with the view that human fertility behaviour is currently adaptive. Here we report, to our knowledge, the first clear evidence for the predicted nonlinear relationship between female fertility and reproductive success in a human population, the Dogon of Mali, West Africa. The predicted maximum reproductive success of 4.1+/-0.3 surviving offspring was attained at a fertility of 10.5 births. Eighty-three per cent of the women achieved a lifetime fertility level (7-13 births) for which the predicted mean reproductive success was within the confidence limits (3.4 to 4.8) for reproductive success at the optimal fertility level. Child mortality, rather than fertility, was the primary determinant of fitness. Since the Dogon people are farmers, our results do not support the assumptions that: (i) contemporary foragers behave more adaptively than agriculturalists, and (ii) that adaptive fertility behaviour ceased with the Neolithic revolution some 9000 years ago. We also present a new method that avoids common biases in measures of reproductive success.     

Reproductive success, mortality and sexual size dimorphism in the adder, Vipera berus

From 1981 to 1986 an isolated adder population was studied in the extreme south of Sweden. During this period 48 adult males and 44 adult females were marked. Male adders did not grow as large as the females. Large males had a significantly higher annual mating success and were engaged in more combats than smaller males. The mean length of recaptured males was significantly lower than that of those not recaptured, indicating a higher motality of larger males. Females brood size was positively correlated with body size. In females there was no difference in mean length of recaptured vs not recaptured individuals. The adder is one of the few snake species with male combat where males are smaller than females. I suggest that this is due to stronger selective advantages for large body size in females than in males.     

The evolution of condition-dependent sexual dimorphism

Theory suggests that the net benefit of allocating resources to a sexual trait depends both on the strength of sexual selection on that trait and on individual condition. This predicts a tight coevolution between sexual dimorphism and condition dependence and suggests that these patterns of within-sex and between-sex variation may share a common genetic and developmental basis. Although condition-dependent expression of sexual traits is widely documented, the extent of covariation between condition dependence and sexual dimorphism remains poorly known. I investigated the effects of condition (larval diet quality) on multivariate sexual dimorphism in the fly Telostylinus angusticollis (Neriidae). Condition determined the direction of sexual size dimorphism and modulated sexual shape dimorphism by affecting allometric slopes and/or intercepts of sexually homologous traits in both sexes. Although the greatest responses to condition manipulation were observed in male sexual traits, both sexual and nonsexual traits exhibited substantial variation in the nature and magnitude of condition effects. Nonetheless, condition dependence and sexual dimorphism were remarkably congruent: variation in the strength of condition effects on male traits explained more than 90% of the variation in the magnitude of sexual dimorphism, whether quantified in terms of trait size or allometric slope. The genetic mechanisms that give rise to multivariate sexual dimorphism in body shape thus function in a strongly condition-dependent manner in this species, suggesting a common genetic basis for body shape variation within and between sexes.     

A note on the evolution of sexual dimorphism

Sexual dimorphism is discussed as a diffusion problem. The establishment of different gametic size is favoured through Brownian motion.     

Variable preferences for sexual dimorphism in height as a strategy for increasing the pool of potential partners in humans

Human mate preferences are known to be related to a number of morphological traits. Those relating to female waist-to-hip ratio or body mass index and to male height appear to be distinctive mate-choice criteria and are known to be related to reproductive success. In addition to absolute height, a possibly important mate-choice criterion may be relative height, i.e. the extent of sexual dimorphism in stature (SDS = male height/female height) between oneself and a potential partner. Here, I demonstrate that people adjust their preferences for SDS in relation to their own height in order to increase the potential pool of partners. This causes nonlinearity in assortative mating in relation to height and shows that in relation to intrapopulational SDS both men and women are responsible for stabilizing selection.     

Testing some hypotheses on human evolution and sexual dimorphism

Research on human evolution and sexual dimorphism motivates an interesting test problem. In studying hominid phylogeny it is of interest to test whether parallel evolution plays a role. With regard to sexual dimorphism it is of interest to known whether the directions of sexual dimorphism in the populations being compared are the same. We show that testing these two problems gives rise to the same type of hypothesis testing, viz. the problem of testing the hypothesis that the means of independent, normally distributed random vectors with unit covariance matrices are situated on a straight line through the origin. A test is proposed and applied to study the sexual dimorphism of 20 recent skull populations. In this example the hypothesis of equal directions of sexual dimorphism is rejected. The classical theory of constructing multiple discriminant functions (canonical variates) is adapted to the problem of comparing sexual dimorphisms.     

Annual reproductive cycle and sexual dimorphism in the dragonet,repomucenus valenciennei, in tokyo bay, japan

The annual reproductive cycle of the dragonet,Repomucenus valenciennei, from Tokyo Bay, Japan, was studied by analysis of seasonal trends in gonadosomatic indices and histological observations of gonads. Sexual dimorphism in the growth of several body parts relative to standard length (SL) and changes in color pattern of the first dorsal fin with growth were also investigated. The spawning season lasted from spring (April [1991] or February [1992]) to autumn (October) with two spawning peaks, in spring and autumn. In spring, only one-year-old (age 1+) fish spawned, age 0+ females not spawning until autumn, at which time they had reached age 1+. Likewise, histological observations of males indicated that testes had reached full maturity by 80 mm SL (age 1+). The minimum mature size of females was estimated as 60 mm SL. In males 45–80 mm SL, the first spine of the first dorsal fin, last ray of the second dorsal fin, last ray of the anal fin, and caudal fin ray showed strongly positive allometry, indicating rapid growth of these structures relative to SL. Subsequently, their growth returned to an isometric pattern in males>80 mm SL. In females, on the other hand, the same body parts showed slightly positive allometry throughout their growth. The color pattern on the first dorsal fin also changed in males 45–80 mm SL.     

Human fertility variation, size-related obstetrical performance and the evolution of sexual stature dimorphism

In several animal species, change in sexual size dimorphism is a correlated response to selection on fecundity. In humans, different hypotheses have been proposed to explain the variation of sexual dimorphism in stature, but no consensus has yet emerged. In this paper, we evaluate from a theoretical and an empirical point of view the hypothesis that the extent of sexual dimorphism in human populations results from the interaction between fertility and size-related obstetric complications. We first developed an optimal evolutionary model based on extensive simulations and then we performed a comparative analysis for a total set of 38 countries worldwide. Our optimization modelling shows that size-related mortality factors do indeed have the potential to affect the extent of sexual stature dimorphism. Comparative analysis using generalized linear modelling supports the idea that maternal death caused by deliveries and complications of pregnancy (a variable known to be size related) could be a key determinant explaining variation in sexual stature dimorphism across populations. We discuss our results in relation to other hypotheses on the evolution of sexual stature dimorphism in humans.     

Genetic constraints and the evolution of display trait sexual dimorphism by natural and sexual selection

The evolution of sexual dimorphism involves an interaction between sex-specific selection and a breakdown of genetic constraints that arise because the two sexes share a genome. We examined genetic constraints and the effect of sex-specific selection on a suite of sexually dimorphic display traits in Drosophila serrata. Sexual dimorphism varied among nine natural populations covering a substantial portion of the species range. Quantitative genetic analyses showed that intersexual genetic correlations were high because of autosomal genetic variance but that the inclusion of X-linked effects reduced genetic correlations substantially, indicating that sex linkage may be an important mechanism by which intersexual genetic constraints are reduced in this species. We then explored the potential for both natural and sexual selection to influence these traits, using a 12-generation laboratory experiment in which we altered the opportunities for each process as flies adapted to a novel environment. Sexual dimorphism evolved, with natural selection reducing sexual dimorphism, whereas sexual selection tended to increase it overall. To this extent, our results are consistent with the hypothesis that sexual selection favors evolutionary divergence of the sexes. However, sex-specific responses to natural and sexual selection contrasted with the classic model because sexual selection affected females rather than males.     

The evolution of sexual dimorphism in animals: Hypotheses and tests

Three major hypotheses, based upon mechanisms of sexual selection, intersexual food competition and reproductive role division, have been advanced to explain the evolution of sexual dimorphism in body size and morphology of animals. Genetic models suggest that all of the hypotheses are plausible, and empirical studies demonstrate that each of the three mechanisms operates in natural populations. However, problems arise in testing hypotheses for the evolution of sexual dimorphism: more than one mechanism may be operating simultaneously, and the demonstrated occurrence of a mechanism does not indicate that it actually results in selection for dimorphism. A recent statistical technique offers a solution to these problems and provides a promising new approach to the study of sexual dimorphism, in which researchers can assess the relative importance of each mechanism in present-day selection for sexual dimorphism within a species.     

Physical attractiveness influences reproductive success of modern men

Theory suggests that reproductive success is positively associated with an individual’s genetic quality. However, the association between physical attractiveness and reproductive success (i.e., number of offspring) in modern humans remains less clear. Here we examined associations between men’s reproductive success and physical attractiveness from retrospective data obtained from married, divorced, and single samples of Slovakian men. As predicted, facially more attractive and taller men were more likely to engage in marriage. In turn, married men had higher reproductive success than single men. Even when men’s marital status was considered, facially more attractive men had higher reproductive success than their less attractive counterparts. This supports the importance of physical attractiveness in sexual selection in modern humans.     

Adaptive dynamics in diploid, sexual populations and the evolution of reproductive isolation

Evolutionary branching is the process whereby an initially monomorphic population evolves to a point where it undergoes disruptive selection and splits up into two phenotypically diverging lineages. We studied evolutionary branching in three models that are ecologically identical but that have different genetic systems. The first model is clonal, the second is sexual diploid with additive genetics on a single locus and the third is like the second but with an additional locus for mate choice. Evolutionary branching occurred under exactly the same ecological circumstances in all three models. After branching the evolutionary dynamics may be qualitatively different. In particular, in the diploid, sexual models there can be multiple evolutionary outcomes whereas in the corresponding clonal model there is only one. We showed that evolutionary branching favours the evolution of (partial) assortative mating and that this in turn effectively restores the results from the clonal model by rendering the alternative outcomes unreachable except for the one that also occurs in the clonal model. The evolution of assortative mating during evolutionary branching can be interpreted as the initial phase of sympatric speciation with phenotypic divergence and partial reproductive isolation.     

Frequency-dependent variation in reproductive success in Narcissus: implications for the maintenance of stigma-height dimorphism

Negative frequency-dependent selection is a major selective force maintaining sexual polymorphisms. However, empirical demonstrations of frequency-dependent reproductive success are rare, particularly in plants. We investigate this problem by manipulating the frequencies of style morphs in a natural population of Narcissus assoanus, a self-incompatible herb with style-length dimorphism and intra-morph compatibility. We predicted that the reproductive success of morphs would vary negatively with their frequency because of the effects of morph-specific differences in sex-organ position on patterns of pollen transfer. This prediction was generally supported. The fruit and seed set of the two morphs did not differ significantly in plots with 1 : 1 morph ratios. However, short-styled plants produced significantly fewer seeds than long-styled plants in monomorphic plots, and significantly more seeds than long-styled plants in plots with 'long-biased' morph ratios. These patterns indicate that in the absence of physiological barriers to intra-morph mating, negative frequency-dependent selection contributes to the maintenance of stylar polymorphism through inter-morph pollen transfer. Our experimental results also provide insights into the mechanisms governing the biased style-morph ratios in populations of Narcissus species.     

Unpredictable environments,nuptial gifts and the evolution of sexual size dimorphism in insects: an experiment

Many insects have a mating system where males transfer nutrients to females at mating, which are often referred to as ''nuptial gifts''. Among butterflies, some of the characteristic features of these species are polyandry (females mate multiple times), and relatively large male ejaculates. When males produce part of the resources used for offspring, the value of body size might then increase for males and decrease for females. The male/female size ratio is also observed to increase when the degree of polyandry and gift size increase. Butterfly species where gift-giving occurs are generally more variable in body size, suggesting that food quality/quantity fluctuates during juvenile stages. This will cause some males to have much to provide and some females to be in great need, and could be conducive to the evolution of a gift-giving mating system. In such a system, growing male and female juveniles should react differently to food shortage. Females should react by maturing at a smaller size since their own lack of reproductive resources can partly be compensated for by male contributions. Males have to pay the full cost of decreased reproduction if they mature at a small size, making it more important for males to keep on growing, even when growth is costly. An earlier experiment with the polyandrous and gift-giving butterfly, Pieris napi, supported this prediction. The pattern is expected to be absent or reversed for species with small nuptial gifts, where females do not benefit from mating repeatedly, and will thus be dependent on acquiring resources for reproduction on their own. To test this prediction, we report here on an experiment with the speckled wood butterfly, Pararge aegeria. We find that growth response correlates with mating system in the two above species, and we conclude that differences in environmental conditions between species may act as an important factor in the evolution of the mating system and sexual size dimorphism.     

Age related evolution of sexual dimorphism in the Red-necked NightjarCaprimulgus ruficollis

Some morphological and sexual characters were measured in juvenile, first-year and adult Red-necked Nightjars in SW Spain. We found significant age-related growth in wing, tail and body mass, first-year birds being heavier than adults. Adults showed sex differences in tail and wing lengths, and the size of tail and wing spots differed between males and females of all age classes. There was not any difference in the number of wing spots, and juvenile males had a lower number of tail spots than older males. An age-related increase in sexual dimorphism was found to exist by size increase of wing spots in males. Since wing and tail spots are exhibited in courtship displays, the elongation of these secondary sexual characters could be related to mate choice mechanisms.

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Zusammenfassung Einige morphologische Merkmale juveniler, einjähriger und adulter Rothals-Ziegenmelker (Caprimulgus ruficollis) in SW Spanien wurden gemessen. Es ergaben sich signifikante altersbedingte Unterschiede in Flügel- und Schwanzlänge sowie in der Körpermasse; einjährige Vögel waren schwerer als erwachsene. Adulte Vögel wiesen geschlechtsabhängige Unterschiede in der Flügel- und Schwanzlänge auf; die Größe der Schwanz- und Flügelflecken unterschieden sich zwischen Männchen und Weibchen aller Altersklassen. Es gab keine Unterschiede in der Anzahl der Flügelflecken; juvenile Männchen hatten aber weniger Schwanzflecken als ältere. Eine durch das Wachstum der Flügelflecken der Männchen bedingte Zunahme des Sexualdimorphismus wurde gefunden. Da die Flügel und Schwanzflecken bei der Balz eine Rolle spielen, könnte die Vergrößerung dieser sekundären Geschlechtsmerkmale mit Paarwahlmechanismen zusammenhängen.

     

Female reproductive success and the evolution of mating-type frequencies in tristylous populations

In tristylous populations, mating-type frequencies are governed by negative frequency-dependent selection typically resulting in equal morph ratios at equilibrium. However, Narcissus triandrus generally exhibits long-styled (L)-biased populations with a deficiency of the mid-styled (M)-morph. Here we used a pollen-transfer model and measurements of female fertility in natural populations to investigate whether these uneven morph ratios were associated with variation in female reproductive success. Our theoretical analysis demonstrated that morph ratio bias can result from maternal fitness differences among the morphs, and that these effects were magnified by asymmetrical mating. In nine out of 15 populations of N. triandrus, seed set differed significantly among the morphs, but pollen limitation occurred in only two of 11 populations investigated. Average seed set of the M-morph was positively associated with its frequency in populations. Flower size was negatively correlated with the seed set of the M-morph. Our results suggest that interactions between mating patterns and female fertility are responsible for variation in morph frequencies and loss of the M-morph from tristylous populations of N. triandrus.