Sex-specific trait architecture in a spider with sexual size dimorphism

Sexual dimorphism, or sex-specific trait expression, may evolve when selection favours different optima for the same trait between sexes, that is, under antagonistic selection. Intra-locus sexual conflict exists when the sexually dimorphic trait under antagonistic selection is based on genes shared between sexes. A common assumption is that the presence of sexual-size dimorphism (SSD) indicates that sexual conflict has been, at least partly, resolved via decoupling of the trait architecture between sexes. However, whether and how decoupling of the trait architecture between sexes has been realized often remains unknown. We tested for differences in architecture of adult body size between sexes in a species with extreme SSD, the African hermit spider (Nephilingis cruentata), where adult female body size greatly exceeds that of males. Specifically, we estimated the sex-specific importance of genetic and maternal effects on adult body size among individuals that we laboratory-reared for up to eight generations. Quantitative genetic model estimates indicated that size variation in females is to a larger extent explained by direct genetic effects than by maternal effects, but in males to a larger extent by maternal than by genetic effects. We conclude that this sex-specific body-size architecture enables body-size evolution to proceed much more independently than under a common architecture to both sexes.     

Sexual selection in sticklebacks in the field: correlates of reproductive, mating, and paternal success

Male sticklebacks display multiple ornaments, and these ornamentshave been shown to be preferred by females in laboratory experiments.However, few field data exist, and it is not known whether thesepreferences are simultaneously or sequentially operative ina single population. We report correlates of reproductive successin two stickleback populations that differ in their ecology,over several periods within their breeding season. In both populationslarger males had higher reproductive success, but not in all periodsof the breeding season. Reproductive success increased withredness of the throat only in the Wohlensee population, andonly in one period that was characterized by low average success.In the Wohlensee population, the parasitic worm Pomphorhynchuslaevis is abundant, and reproductive success decreased withthe presence of the parasite. In the Roche population, maleswith nests concealed in a plant had higher mating success. Thesenests were less likely to fail, suggesting that females preferredto spawn in concealed nests because of higher offspring survivorship.The different sexual traits appear to reveal different aspectsof male quality (multiple message hypothesis): females probablyfind large males attractive because of their higher paternalquality, but it seems more likely that red males are preferred forbetter genetic qualities. Females also discriminate on territoryquality, and male traits may be important in competition forthese territories. The correlates of reproductive success werenot consistent during the season, probably due to changes inthe availability of ripe females. Such fluctuating selectionpressures will contribute to the maintenance of genetic variationin sexual traits.     

The genetics of adaptive shape shift in stickleback: pleiotropy and effect size

The distribution of effect sizes of genes underlying adaptation is unknown ( Orr 2005 ). Are suites of traits that diverged under natural selection controlled by a few pleiotropic genes of large effect (major genes model), by many independently acting genes of small effect (infinitesimal model), or by a combination, with frequency inversely related to effect size (geometric model)? To address this we carried out a quantitative trait loci (QTL) study of a suite of 54 position traits describing body shapes of two threespine stickleback species: an ancestral Pacific marine form and a highly derived benthic species inhabiting a geologically young lake. About half of the 26 detected QTL affected just one coordinate and had small net effects, but several genomic regions affected multiple aspects of shape and had large net effects. The distribution of effect sizes followed the gamma distribution, as predicted by the geometric model of adaptation when detection limits are taken into account. The sex‐determining chromosome region had the largest effect of any QTL. Ancestral sexual dimorphism was similar to the direction of divergence, and was largely eliminated during freshwater adaptation, suggesting that sex differences may provide variation upon which selection can act. Several shape QTL are linked to Eda, a major gene responsible for reduction of lateral body armor in freshwater. Our results are consistent with predictions of the geometric model of adaptation. Shape evolution in stickleback results from a few genes with large and possibly widespread effects and multiple genes of smaller effect.     

Body condition and reproductive status in sticklebacks exposed to a single wave of Schistocephalus solidus infection

This paper describes the results of a study of the effects of Schistocephalus solidus on one of its intermediate hosts, the stickleback Gasterosteus aculeatus , in an annual population in which infection is known to occur in one major wave in autumn. Weight, as a function of length, was lower in infected sticklebacks compared with uninfected fish during autumn and spring; in winter and summer, both categories of fish were in equally poor condition. In early autumn, the hepatosomatic indices of newly infected fish were higher than those of uninfected fish, perhaps due to a pathological response. Thereafter, relative liver size of uninfected and infected sticklebacks was comparable until spring, when it increased sharply in uninfected sticklebacks but remained stable at a low level in infected sticklebacks. Few infected fish reached maturity. The only males to attain maturity whilst sustaining an infection of S. solidus were in particularly good condition. Nuptial coloration, kidney hypertrophy and testes size were unimpaired in these mature infected males, but whether they were capable of successful reproduction remains debatable.     

Population variation in sexual selection and its effect on size allometry in two dung fly species with contrasting sexual size dimorphism

Body size is one of the most important quantitative traits under evolutionary scrutiny. Sexual size dimorphism (SSD) in a given species is expected to result if opposing selection forces equilibrate differently in both sexes. We document variation in the intensity of sexual and fecundity selection, male and female body size, and thus SSD among 31 and 27 populations of the two dung fly species, Scathophaga stercoraria and Sepsis cynipsea, across Switzerland. Whereas in S. cynipsea females are larger, the SSD is reversed in S. stercoraria. We comprehensively evaluated Fairbairn and Preziosi's (1994) general, three-tiered scenario, hypothesizing that sexual selection for large male size is the major driving force of SSD allometry within these two species. Sexual selection intensity on male size in the yellow dung fly, S. stercoraria, was overall positive, greater, and more variable among populations than fecundity selection on females. Also, sexual selection intensity in a given population correlated positively with mean male body size of that population for both the field-caught fathers and their laboratory-reared sons, indicating a response to selection. In S. cvnipsea, sexual selection intensity on males was lower overall and significantly positive, about equal in magnitude, but more variable than fecundity selection on females. However, there was no correlation between the intensity of sexual selection and mean male body size among populations. In both species, the laboratory-reared offspring indicate genetic differentiation among populations in body size. Despite fulfillment of all key prerequisites, at least in S. stercoraria, we did not find hypoallometry for SSD (Rensch's rule, i.e., greater evolutionary divergence in male size than female size) for the field-caught parents or the laboratory-reared offspring: Female size was isometric to male size in both species. We conclude that S. cynipsea does not fit some major requirements of Fairbairn and Preziosi's (1994) scenario, whereas for S. stercoraria we found partial support for it. Failure to support Rensch's rule within the latter species may be due to phylogenetic or other constraints, power limitations, erroneous estimates of sexual selection, insufficient genetic isolation of populations, or sex differences in viability selection against large size.     

Nest-site selection in a fish species with paternal care

Hydrobiologia - Fish that perform paternal care may increase their fitness by choosing nest sites that enhance survival and development of embryos. We studied nest-site choice with respect to...     

It is not always about body size: evidence of Rensch's rule in a male weapon

In many species, sexual dimorphism increases with body size when males are the larger sex but decreases when females are the larger sex, a macro-evolutionary pattern known as Rensch''s rule (RR). Although empirical studies usually focus exclusively on body size, Rensch''s original proposal included sexual differences in other traits, such as ornaments and weapons. Here, we used a clade of harvestmen to investigate whether two traits follow RR: body size and length of the fourth pair of legs (legs IV), which are used as weapons in male–male fights. We found that males were slightly smaller than females and body size did not follow RR, whereas legs IV were much longer in males and followed RR. We propose that sexual selection might be stronger on legs IV length than on body size in males, and we discuss the potential role of condition dependence in the emergence of RR.     

Sex-specific plasticity of growth and maturation size in a spider: implications for sexual size dimorphism

Sex-specific plasticity in body size has been recently proposed to cause intraspecific patterns of variation in sexual size dimorphism (SSD). We reared juvenile male and female Mediterranean tarantulas (Lycosa tarantula) under two feeding regimes and monitored their growth until maturation. Selection gradients calculated across studies show how maturation size is under net stabilizing selection in females and under directional selection in males. This pattern was used to predict that body size should be more canalized in females than in males. As expected, feeding affected male but not female maturation size. The sex-specific response of maturation size was related to a dramatic divergence between subadult male and female growth pathways. These results demonstrate the existence of sex-specific canalization and resource allocation to maturation size in this species, which causes variation in SSD depending on developmental conditions consistent with the differential-plasticity hypothesis explaining Rensch's Rule.     

Long-term paternity skew and the opportunity for selection in a mammal with reversed sexual size dimorphism

Most mammalian groups are characterized by male-biased sexual size dimorphism, in which size-dependent male-male competition and reproductive skew are tightly linked. By comparison, little is known about the opportunity for sexual selection in mammalian systems without male-biased dimorphism, where the traits under sexual selection might be less obvious. We examined 10 years of parentage data in a colony of greater horseshoe bats (Rhinolophus ferrumequinum) to determine the magnitude of male reproductive skew and the opportunity for sexual selection in a mammal in which females are the larger sex. Annual paternity success was weakly skewed but consistent patterns led to strong longitudinal paternity skew among breeders. Just three males accounted for a third of all paternity assignments, representing at least a fifth of all colony offspring born in a decade. Paternity success was in part determined by age but was not influenced by dispersal status. Our results show that paternity skew and the opportunity for sexual selection in a species with reversed sexual size dimorphism can approach levels reported for classical examples of species with polygyny and male-biased dimorphism, even where the traits under sexual selection are not known.     

Sexes of a monomorphic species differ in preference for mates with a novel trait

We investigated the different roles of the sexes in the originationof novel traits in the sexually monomorphic Javanese mannikinLonchura leucogastroides. We introduced a red feather as anevolutionarily novel trait in both sexes and tested their preferencesfor prospective mates with this trait. Males rejected femalesbearing the red feather and preferred to court unadorned females.In contrast, females partly preferred adorned males. Specifically,previously unattractive males gained in attractiveness and could increasetheir reproductive success when bearing the ornament, whereas previouslyattractive males lost in attractiveness, but this did not affect theirreproductive success. We introduced two other novel traits inmales and investigated the females' response to these in matechoice tests. Each of the three new traits interacted with thenatural attractiveness of males. The more attractive a malewas before ornamentation, the more it lost in attractiveness afterornamentation and vice versa. Thus, the position of the traitdid not affect the interaction. Because males rejected adornedfemales and females partly preferred adorned males, novel traitsmight evolve by intersexual selection in males rather than infemales. This can lead to a sexual dimorphism with conspicuoustraits in males. Our study reveals a new insight into the mechanismof the evolution from monomorphism to dimorphism with ornamentaltraits in males.     

The anal fin complex in a weakly discharging electric fish, Gnathonemus petersii(Mormyridae)

An examination of the permanent bony structures of the anal fin complex in the mormyrid fish, Gnathonemus petersii , revealed two new structural sexual dimorphisms: longer proximal pterygiophores and wider anal fin rays in males than in females. Both structures are thought to facilitate the male's courtship‐associated anal fin reflex. Adult male mormyrid fishes are characterized by a dorsally directed indentation of the posterior body wall (anal fin indentation). The expression of this indentation in males, presumably driven by anal fin musculature, was correlated with the fish's gonadal state: large indentations were associated with high gonado‐somatic indices and small indentations with low indices.     

Sexual dimorphism in chelicerae size in three species of nuptial-gift spiders: a discussion of possible functions and driving selective forces

Positive allometric patterns observed for intersexual signalling characters are related to directional sexual selection, and supported by theoretical and empirical data. Recent models have shown that positive allometry may not hold as a rule if the influence of natural selection is added to the model. Here we tested these models applying traditional morphometrical techniques for the analysis of chelicerae sexual dimorphism and allometric patterns within the genus Paratrechalea : Paratrechalea azul , Paratrechalea galianoae and Paratrechalea ornata . Spider chelicerae are basically used for prey capture, but males of Paratrechalea also use the chelicerae to offer a nuptial gift during courtship, also presenting a clear size and colour sexual dimorphism supporting a possible role as a signal. Chelicerae size was male biased for all the variables studied and showed an isometric pattern, while females showed a higher variation. Our findings are in accordance with models of viability-related function for prey capture, questioning some statements proposed by the positive allometry model.     

Sexual selection explains sex-specific growth plasticity and positive allometry for sexual size dimorphism in a reef fish

In 1950, Rensch noted that in clades where males are the larger sex, sexual size dimorphism (SSD) tends to be more pronounced in larger species. This fundamental allometric relationship is now known as ‘Rensch''s rule’. While most researchers attribute Rensch''s rule to sexual selection for male size, experimental evidence is lacking. Here, we suggest that ultimate hypotheses for Rensch''s rule should also apply to groups of individuals and that individual trait plasticity can be used to test those hypotheses experimentally. Specifically, we show that in the sex-changing fish Parapercis cylindrica, larger males have larger harems with larger females, and that SSD increases with harem size. Thus, sexual selection for male body size is the ultimate cause of sexual size allometry. In addition, we experimentally illustrate a positive relationship between polygyny potential and individual growth rate during sex change from female to male. Thus, sexual selection is the ultimate cause of variation in growth rate, and variation in growth rate is the proximate cause of sexual size allometry. Taken together, our results provide compelling evidence in support of the sexual selection hypothesis for Rensch''s rule and highlight the potential importance of individual growth modification in the shaping of morphological patterns in Nature.     

Evolution of sexual dimorphism in bill size and shape of hermit hummingbirds (Phaethornithinae): a role for ecological causation

Unambiguous examples of ecological causation of sexual dimorphism are rare, and the best evidence involves sexual differences in trophic morphology. We show that moderate female-biased sexual dimorphism in bill curvature is the ancestral condition in hermit hummingbirds (Phaethornithinae), and that it is greatly amplified in species such as Glaucis hirsutus and Phaethornis guy, where bills of females are 60 per cent more curved than bills of males. In contrast, bill curvature dimorphism is lost or reduced in a lineage of short-billed hermit species and in specialist Eutoxeres sicklebill hermits. In the hermits, males tend to be larger than females in the majority of species, although size dimorphism is typically small. Consistent with earlier studies of hummingbird feeding performance, both raw regressions of traits and phylogenetic independent contrasts supported the prediction that dimorphism in bill curvature of hermits is associated with longer bills. Some evidence indicates that differences between sexes of hermit hummingbirds are associated with differences in the use of food plants. We suggest that some hermit hummingbirds provide model organisms for studies of ecological causation of sexual dimorphism because their sexual dimorphism in bill curvature provides a diagnostic clue for the food plants that need to be monitored for studies of sexual differences in resource use.     

Sex differences in phenotypic plasticity of a mechanism that controls body size: implications for sexual size dimorphism

The degree and/or direction of sexual size dimorphism (SSD) varies considerably among species and among populations within species. Although this variation is in part genetically based, much of it is probably due to the sexes exhibiting differences in body size plasticity. Here, we use the hawkmoth, Manduca sexta, to test the hypothesis that moths reared on different diet qualities and at different temperatures will exhibit sex-specific body size plasticity. In addition, we explore the proximate mechanisms that potentially create sex-specific plasticity by examining three physiological variables known to regulate body size in this insect: the growth rate, the critical weight (which measures the cessation of juvenile hormone secretion from the corpora allata) and the interval to cessation of growth (ICG; which measures the time interval between the critical weight and the secretion of the ecdysteroids that regulate pupation and metamorphosis). We found that peak larval mass of males and females did not exhibit sex-specific plasticity in response to diet or temperature. However, the sexes did exhibit sex-specific plasticity in the mechanism that controls size; males and females exhibited sex-specific plasticity in the growth rate and the critical weight in response to both diet and temperature, whereas the ICG only exhibited sex-specific plasticity in response to diet. Our results suggest it is important for the sexes to maintain the same degree of SSD across environments and that this is accomplished by the sexes exhibiting differential sensitivity of the physiological factors that determine body size to environmental variation.     

Variation in selection pressure acting on body size by age and sex in a reverse sexual size dimorphic raptor

Body size strongly influences fitness, with larger individuals benefiting in terms of both greater productivity and survivorship; for reverse sexual size dimorphic (RSD) species, this relationship may be more complex. We examined the selection pressures acting on body size in male and female Merlins Falco columbarius to assess whether larger or smaller individuals of this RSD species were favoured in terms of survival and breeding performance. For males and females there were clear links between body size and survival but the exact relationship varied by sex. Among males, birds that survived each year class were larger than those that died and yearlings were on average smaller than older birds, but there were no measurable differences among adult males (age 2+). Among females, larger individuals aged 1 and 2 years were more likely to survive, but this size‐based pattern was not apparent in older age classes. Size early in life predicted the lifespan in male Merlins but not as strongly as for females and not for the largest individuals. Reproductive performance based on brood size was not associated with body size in either males or females, but there was a weak positive relationship between female body size and lifetime reproductive success. Selection appears to favour larger males and females but there is no indication that the population is evolving towards bigger individuals, perhaps in part due to selection against the largest birds. Increased survival may allow larger and higher quality individuals to occupy higher quality territories as they age and thereby to accrue greater lifetime reproductive success in the process.