Sexual dimorphism in birds: why are there so many different forms of dimorphism?

Variation in the extent of sexual dimorphism among bird species is traditionally attributed to differences in social mating system. However, there are many different forms of dimorphism among birds, and not all of them show an obvious correlation with social mating system. For example, recent work has shown that many highly polygamous species are, in fact, monomorphic, whereas many putatively monogamous species are dimorphic. In this paper we break up sexual dimorphism into subcomponents and then use comparative analyses to examine the pattern of covariation between these subcomponents and various aspects of sexual, social, and parental behaviour. Our first finding is that size dimorphism and plumage-colour dimorphism do not show the same pattern of covariation. Differences in size dimorphism are associated with variation in social mating system and sex differences in parental care, whereas differences in plumage-colour dimorphism are associated with variation in the frequency of extra-bond paternity. These results suggest that size dimorphism is associated with the sort of intrasexual competition described by traditional classifications of social mating system, whereas plumage-colour dimorphism is associated with cryptic female choice. However, when we break up plumage-colour dimorphism according to whether it is due to melanins, carotenoids or structural colours, we find that each category of plumage-colour dimorphism shows a different pattern of covariation. The correlation between overall plumage-colour dimorphism and the rate of extra-bond paternity is due to structural colours, whereas melanin-based dimorphism is associated with sex differences in parental care. The former result is particularly interesting given that new work suggests structural colours are associated with active sexual displays and the reflection of ultraviolet light.     

Sexual dimorphism in a Lower Miocene moronid?

A series of questionable elements in certain specimens ofMorone cf.aequalis (Koken 1891) from Lower Miocene deposits near the village of Berkersheim, N of Frankfurt a. M. (Hessen, Germany) is described, which has not been known from any other percoid before. These elements are fully ossified and cover the cheek and the preopercular region. Even within well-preserved material, they are only present in some specimens. Therefore, they may be specialized structures that are indicative for sexual dimorphism. Nevertheless, they clearly differ from all other respective structures that have been described from teleosts: Multicellular epidermal horny tubercles (“breeding tubercles”) mainly consist of keratine and not of calciumphosphate. By contrast, contact organs consist of bone and are located mainly at the surface of the fin rays and scales, respectively. At present, “breeding tubercles” are the favorite interpretation and the original substance may have been replaced via post-mortem phosphatization.     

Establishing sexual dimorphism: conservation amidst diversity?

The molecular mechanisms that control sexual dimorphism are very different in distantly related animals. Did sex determination arise several times with different regulatory mechanisms, or is it an ancient process with little surviving evidence of ancestral genes? The recent identification of related sexual regulators in different phyla indicates that some aspects of sexual regulation might be ancient. Studies of sex-determining mechanisms are beginning to reveal how sexual dimorphism arises and evolves.     

Does sexual dimorphism in human faces signal health?

Evolutionary psychologists suggest that a preference for sexually dimorphic traits in human faces is an adaptation for mate choice, because such traits reflect health during development. For male faces, this claim rests on the immunocompetence-handicap hypothesis, which states that the increased testosterone levels needed to develop large masculine traits stress the immune system. We examined whether masculine traits in adolescent male faces are associated with health during development, and also whether feminine traits in adolescent female faces signal health. Feminine traits are attractive, but it is less clear whether they should signal health. Rated masculinity in adolescent male faces correlated modestly with actual health, and was perceived as healthy, but not as attractive. Rated femininity in adolescent female faces did not correlate with actual health, although it was perceived as healthy and attractive. These results support the immunocompetence-handicap hypothesis for male faces in that masculine traits signalled health during adolescence. However, they suggest that any health-related evolutionary benefits obtained from preferences for attractive facial traits may be weak.     

β-1,3-Glucanase and dimorphism in Paracoccidioides brasiliensis

Mycelial and yeast forms of P. brasiliensis were tested for several glucohydrolases. In addition to high levels of -blucanases, low amounts of -glucanase, chitinase and maltase were found. Tests for invertase, amylase and lactase were negative. The levels of -1,3-glucanase were higher in the mycelial form. The shift to the mycelial phase correlated with an increase in the levels of -1,3-glucanase. The enzyme was present in the cytoplasm, cell wall and culture medium. The extracellular enzyme was purified 42 fold by ammonium sulphate precipitation and gel filtration. Maximal activity was obtained at 60°C and pH of 5.0 acetate buffer or pH 6.0 (phosphate buffer). Its K _m was 0.205 mg/ml. The cell wall-bound enzyme showed a higher temperature optimum. Optimum pH and K _m were also slightly different. Following treatment of the cell walls with chitinase, -1,3-glucanase was released into the medium.     

Sexual dimorphism: can you smell the difference?

A powerful new technique for visualizing neurons in the fly brain has uncovered fine neuroanatomical differences between the olfactory circuitries of male and female Drosophila.     

The evolution of male-female dimorphism: Older than sex?

This contribution considers the evolution of a dimorphism with respect to cell fusion characteristics in a population of primitive cells. These cells reproduce exclusively asexually. The evolution towards asymmetric fusion behaviour of cells is driven by selection promoting horizontal transfer of an endosymbiontic replicator. It is concluded that evolution of asymmetric cell fusion in this scenario is more likely than evolution of sexual differentiation in a sexually reproducing population. Pre-existing dimorphism with respect to cell fusion may thus have been the basis for the establishment of sexual differentiation at the level of gamete fusion, and this in turn is fundamental to the evolution of two different sexes, male and female.     

Can evolution of sexual dimorphism be triggered by developmental temperatures?

Genetic prerequisites for the evolution of sexual dimorphism, sex-specific heritabilities and low or negative genetic correlations between homologous traits in males and females are rarely found. However, sexual dimorphism is evolving rapidly following environmental change, suggesting that sexual dimorphism and its genetic background could be environmentally sensitive. Yet few studies have explored the sensitivity of the genetic background of sexual dimorphism on environmental variation. In this study, on Drosophila melanogaster, we used a large nested full-sib-half-sib breeding design where families were split into four different developmental temperatures: two constant temperature treatments of 25 and 30 °C and two cycling temperatures with means of 25 and 30 °C, respectively. After emergence, we tested heat shock tolerance of adult flies. We found that sexual dimorphism was strongly affected by temperature during development. Moreover, we found that female heritability was significantly lower in flies developing at hot temperature and more so under hot and cycling temperatures. Interestingly, most of the genetic variation for heat shock tolerance was orthogonal (i.e. noncorrelated) between sexes, allowing independent evolution of heat shock tolerance in males and females. These findings give support to the hypothesis that the evolution of sexual dimorphism can be influenced by the environments experienced during development.     

Why is there so much genetic variation for wing dimorphism?

Wing dimorphism appears in general to be determined either by a single locus, 2 allele system in which brachyptery is dominant, or by the additive action of numerous loci. In the latter case studies indicate that the heritability is typically quite large. It is generally postulated that wing dimorphism is under strong selection: why then is genetic variation not eroded? In this paper I consider three possible explanations. First, genetic variation may not be exposed to selection because environmental heterogeneity effectively makes heritability zero. Because wing dimorphisms are known to evolve it seems unlikely that this is the primary factor. Second, directional selection on a threshold trait may push the population almost to monomorphism but erodes genetic variance at a very slow rate. This mechanism cannot preserve variation but makes it possible for other factors to more easily maintain variability. Finally, I demonstrate that in a heterogeneous environment spatio-temporal variation in fitness will itself maintain a genetic polymorphism for wing dimorphism.     

Which proximate factor determines sexual size dimorphism in tiger snakes?

Diverse interactions between factors that influence body size complicate the identification of the primary determinants of sexual size dimorphism. Using data from a long‐term field study (1997–2009), we examined the contributions of the main proximate factors potentially influencing sexual size dimorphism from birth to adulthood in tiger snakes (Notechis scutatus). Data on body size, body mass and body condition of neonates, juveniles and adults were obtained by mark–recapture. Frequent recaptures allowed us to monitor reproductive status, diet and food intake, and to estimate survival and growth rates in age and sex classes. Additional data from females held briefly in captivity enabled us to assess reproductive output and the body mass lost at parturition (proxies for reproductive effort). From birth to maturity, individuals of both sexes experienced similar growth and mortality rates. We found no difference in diet, feeding and survival rates between the sexes, nor between juveniles and adults. On maturity, despite comparable diet and food intake by both sexes, the high energy requirements of vitellogenesis and gestation were responsible for a depletion of body reserves and probably resulted in a marked decrease in growth rates. Males were largely exempt from such costs of reproduction, and so could grow faster than females and attain larger body sizes. The absence of niche divergence between the sexes (uniformity of habitat, lack of predators) suggests that the impact of differential energetic investment for reproduction on growth rate is probably the main proximate factor influencing sexual size dimorphism in this species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 668–680.     

Sexual dimorphism in immunocompetence: what does life-history theory predict?

Sexual dimorphism in immunocompetence, usually in the directionof inferior male immunocompetence, has historically been explainedas the result of proximate physiological mechanisms such asthe immunosuppressive effects of the male hormone testosterone.More recently, it has been argued that this pattern is bestunderstood as a result of resource-based trade-offs betweenmale mating effort and immune defense, a trade-off that femalesdo not make. The central prediction of this hypothesis is thatas the strength of sexual selection on males increases, themagnitude of the sex differences in immunocompetence will increase.Two implicit assumptions of this argument are that 1) longevityis of more importance for female than for male fitness and 2)that the primary benefit of immunocompetence is increased longevity.However, both of these assumptions may not be as broadly applicableas has been argued. We have modeled the optimal allocation toimmunocompetence for males and females without making theseassumptions. We find that the optimal allocation to immune defensefor males decreases as the strength of sexual selection increases,as predicted. However, males may still invest more, relativeto females, into immunocompetence if the impact of parasiteson condition differs for the sexes and/or if the relationshipbetween condition and reproduction differs for the sexes. Weargue that these previously overlooked assumptions may be criticalfor predicting sex-specific patterns of immunocompetence.     

Is sexual dimorphism in the femur a "population specific phenomenon"?

The patterns of sexual dimorphism as well as the differences in amount between the populations were studied on a sample of 162 male and 159 female left femora, which were classified as Zulu, Sotho, Xosa and South Africans of European extraction. Multivariate analyses revealed that even adjacent African tribes exhibit a different pattern of sexual dimorphism, but there were similarities between Zulu and European femora. Furthermore, relative size differences, i.e. shape, discriminated more clearly between the sexes than did absolute size. Bicondylar width yielded a statistically significant higher degree of sexual dimorphism in Europeans when compared to African populations. This finding was interpreted in terms of the biomechanical demands on the femur under different living conditions. On the other hand, sexual dimorphism of femoral length did not differ among the populations. This was unexpected since femoral length correlates highly with stature, which was reported to show a lesser degree of sexual dimorphism in Africans than in Europeans. Detailed analyses of the results of the present study led to suggest that different living conditions may affect bones in complex ways of which linear growth is only one aspect.     

Is sexual body shape dimorphism consistent in aquatic and terrestrial chelonians?

Comparisons between aquatic and terrestrial species provide an opportunity to examine how sex-specific adaptations interact with the environment to influence body shape. In terrestrial female tortoises, selection for fecundity favors the development of a large internal abdominal cavity to accommodate the clutch; in conspecific males, sexual selection favors mobility with large openings in the shell. To examine to what extent such trends apply in aquatic chelonians we compared the body shape of males and females of two aquatic turtles (Chelodina colliei and Mauremys leprosa). In both species, females were larger than males. When controlled for body size, females exhibited a greater relative internal volume and a higher body condition index than males; both traits potentially correlate positively with fecundity. Males were more streamlined (hydrodynamic), and exhibited larger openings in the shell providing more space to move their longer limbs; such traits probably improve mobility and copulation ability (the males chase and grab the female for copulation). Overall, although the specific constraints imposed by terrestrial and aquatic locomotion shape the morphology of chelonians differently (aquatic turtles were flatter, hence more hydrodynamic than terrestrial tortoises), the direction for sexual shape dimorphism remained unaffected. Our main conclusion is that the direction of sexual shape dimorphism is probably more consistent than sexual size dimorphism in the animal kingdom.     

Does inbreeding avoidance maintain gender dimorphism in Wurmbea dioica (Colchicaceae)?

The maintenance of females in gender dimorphic populations requires that they have a fitness advantage to compensate for their loss of male reproductive function. We assess whether inbreeding avoidance provides this advantage in two subdioecious Wurmbea dioica populations by estimating seed production, outcrossing rates and inbreeding depression. Fruiting males produced less than half as many seeds as females, owing to low outcrossing rates and early acting inbreeding depression. Inbreeding coefficients of fruiting males demonstrated that progeny were more inbred than their parents, implying that few selfed progeny reach maturity, as confirmed by inbreeding depression estimates that exceeded 0.85. In a glasshouse experiment, open-pollinated females exhibited a fitness advantage of 3.7 relative to fruiting males, but when we increased fruiting male outcrossing rate, female advantage was only 1.4. This reduced advantage is insufficient to maintain females if nuclear genes control sex. Thus, inbreeding avoidance could maintain females at high frequencies, although this is contingent upon high frequencies of fruiting males, which can be altered by environmentally determined gender plasticity.     

Sexual dimorphism and the differential mortality model: is behaviour related to survival?

There are numerous hypotheses to explain the evolution of sexual dimorphism in spiders. One of the most controversial is the differential mortality model (DMM) which proposes that differing rates of (adult) male and female mortality can result in a skewed operational sex ratio and lead to the evolution of small males. This hypothesis has been examined using a comparative approach which assumes that the behaviour of males and females could be used as a surrogate measure of mortality. We tested this assumption using two model species, Hogna helluo and Pardosa milvina (Araneae: Lycosidae) that differ in the degree of sexual dimorphism both in terms of body size and level of activity. Our data demonstrate that differences in male and female behaviour are not predictive of differences in mortality. Rather, as in other organisms, mortality is a complex phenomenon dependent on activity as well as size. These data call into question the methods previously used to test the DMM and suggest that understanding sexual size dimorphism (SSD) in spiders will require evaluation of historical constraints as well as how size currently influences fitness in each sex.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 97−103.     

The evolution of sexual dimorphism in parasitic cuckoos: sexual selection or coevolution?

Sexual dimorphism is ubiquitous in animals and can result from selection pressure on one or both sexes. Sexual selection has become the predominant explanation for the evolution of sexual dimorphism, with strong selection on size-related mating success in males being the most common situation. The cuckoos (family Cuculidae) provide an exceptional case in which both sexes of many species are freed from the burden of parental care but where coevolution between parasitic cuckoos and their hosts also results in intense selection. Here, we show that size and plumage differences between the sexes in parasitic cuckoos are more likely the result of coevolution than sexual selection. While both sexes changed in size as brood parasitism evolved, we find no evidence for selection on males to become larger. Rather, our analysis indicates stronger selection on parasitic females to become smaller, resulting in a shift from dimorphism with larger females in cuckoos with parental care to dimorphism with larger males in parasitic species. In addition, the evolution of brood parasitism was associated with more cryptic plumage in both sexes, but especially in females, a result that contrasts with the strong plumage dimorphism seen in some other parasitic birds. Examination of the three independent origins of brood parasitism suggests that different parasitic cuckoo lineages followed divergent evolutionary pathways to successful brood parasitism. These results argue for the powerful role of parasite-host coevolution in shaping cuckoo life histories in general and sexual dimorphism in particular.