Pathways to elaboration of sexual dimorphism in bird plumage patterns

Patterns, such as bars and spots, are common in birds. Some patterns can function in camouflage and/or communication and can benefit both males and females, paving the way for elaboration in sexual dimorphism. Historically, sexual dichromatism was predominantly considered to be a consequence of mating systems. However, the distribution of traits between the sexes is not always indicative of function; genetic correlation may cause traits to evolve in both sexes and traits may serve a social function in males and/or females. In addition, sexual dichromatism in bird plumage patterns can be composed of multiple types of patterns within and/or between the sexes. Therefore, there can be more than one type of dimorphism and some are more elaborate than others. Under classical models of genetic correlation, patterns evolve in both sexes followed by a loss of patterning in one sex. Elaborate types of sexual dimorphism in plumage patterns may be due to selection acting on existing patterns and are perhaps derived. Waterfowl (Anseriformes) and gamebirds (Galliformes) arguably have the most striking plumage patterns. Using 288 species from these orders I reconstructed the evolutionary history of plumage pattern dimorphism. There was little support for genetic correlation but elaborate types of dimorphism are probably derived. Backward and forward evolutionary transitions between different types of dimorphism can occur by loss or elaboration. These results demonstrate that plumage patterns are evolutionary labile and current forms may represent shifting adaptations to a changing environment. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 262–273.     

Macroevolutionary patterns of sexual size dimorphism in copepods

Major theories compete to explain the macroevolutionary trends observed in sexual size dimorphism (SSD) in animals. Quantitative genetic theory suggests that the sex under historically stronger directional selection will exhibit greater interspecific variance in size, with covariation between allometric slopes (male to female size) and the strength of SSD across clades. Rensch''s rule (RR) also suggests a correlation, but one in which males are always the more size variant sex. Examining free-living pelagic and parasitic Copepoda, we test these competing predictions. Females are commonly the larger sex in copepod species. Comparing clades that vary by four orders of magnitude in their degree of dimorphism, we show that isometry is widespread. As such we find no support for either RR or for covariation between allometry and SSD. Our results suggest that selection on both sexes has been equally important. We next test the prediction that variation in the degree of SSD is related to the adult sex ratio. As males become relatively less abundant, it has been hypothesized that this will lead to a reduction in both inter-male competition and male size. However, the lack of such a correlation across diverse free-living pelagic families of copepods provides no support for this hypothesis. By comparison, in sea lice of the family Caligidae, there is some qualitative support of the hypothesis, males may suffer elevated mortality when they leave the host and rove for sedentary females, and their female-biased SSD is greater than in many free-living families. However, other parasitic copepods which do not appear to have obvious differences in sex-based mate searching risks also show similar or even more extreme SSD, therefore suggesting other factors can drive the observed extremes.     

Sexual selection and mating patterns in a mammal with female-biased sexual size dimorphism

In mammals, species with highly male-biased sexual size dimorphismtend to have high variance in male reproductive success. However,little information is available on patterns of sexual selection,variation in male and female reproductive success, and bodysize and mating success in species with female-biased size dimorphism.We used parentage data from microsatellite DNA loci to examinethese issues in the yellow-pine chipmunk (Tamias amoenus), asmall ground squirrel with female-biased sexual size dimorphism.Chipmunks were monitored over 3 years in the Kananaskis Valley,Alberta, Canada. We found evidence of high levels of multiplepaternity within litters. Variation in male and female reproductivesuccess was equal, and the opportunity for sexual selectionwas only marginally higher in males than females. Male and femalereproductive success both depended on mating success. We foundno evidence that the number of genetic mates a male had dependedon body size. Our results are consistent with a promiscuousmating system in which males and female mate with multiple partners.Low variation in male reproductive success may be a generalfeature of mammalian species in which females are larger thanmales.     

Inferential assessment of the MI index of sexual dimorphism: A comparative study with some other sexual dimorphism measures

In order to provide inferential support to the MI measure of sexual dimorphism we proposed for populations distributed as mixture models with two normal components, an interval estimate is constructed. There do not appear to exist measures of sexual dimorphism that possess inferential properties other than some statistics used with this purpose. The use of these sample functions in such a context as well as the purported inferential support of some other sexual dimorphism indices are discussed. A biological case study illustrates the distinct inferential conclusions that can be obtained when the indices here discussed and the one we proposed are considered.     

Sniffing out patterns of sexual dimorphism in floral scent

1.   A major transition in flowering plants has been the evolution of separate sexes (dioecy) from combined sexes (hermaphroditism). This transition is often, but not always, accompanied by the evolution of sexual dimorphism in attractive traits, and floral scent is no exception.
2.   In this review I aim to improve our understanding of variation in sexual dimorphism in floral scent characteristics by first explicating the relevant hypotheses, and then deriving explicit predictions for the pattern of floral scent from each.
3.   Next, I synthesize and qualitatively review published data on floral volatile emission rate and composition in 33 gender dimorphic species to identify emerging patterns, and evaluate these in light of predictions derived from the hypotheses.
4.   Although conclusions must be viewed as preliminary, a handful of strong patterns were revealed: (1) in the majority of the species studied males emit more volatiles per flower than females, (2) in over half of the species studied the sexes differed in at least one aspect of scent composition, (3) sexual dimorphism in scent composition was less common in species with rewardless females and/or brood site pollination than those that offered nectar and/or pollen, (4) a one-to-one correspondence between sex differences in overall scent composition and male organ-specific scent production is largely not found.
5.   This review has highlighted gaps in our understanding of the genesis of patterns of sexual dimorphism in floral scent, and makes clear that to move the field forward we need to shift our focus from pattern to process, and this will be best achieved by simultaneously testing alternative hypotheses at the same level of analysis.     

Sexual dimorphism in human skulls. A comparison of sexual dimorphism in different populations

Application and comparison of sex discriminant functions in different populations led to the conclusion that a certain combination and weighting of a few sex dimorphism variables (in this study we only used craniometric variables) can give a good discrimination between male and female individuals, independent of the racial group to which this function is applied. In our study, the sex-discriminatory power of five discriminant functions which were based on different ordination and selection procedures (e.g. professional knowledge, stepwise discriminant analysis, literature) of the cranial variables is compared. These discriminant functions were applied to three different data sets, the first being skull measurements from an Amsterdam series (Europids), the second skull measurements of a Zulu series (Negrids) and the third skull measurements of a Japan series (Mongolids). Our decision as to whether a function is a good or less good sex-discriminating function is determined by the Dt values (these values give an idea about the discriminatory value of the discriminant function when applied to a new test sample), the number of variables necessary to obtain this Dt and the location of the sectioning point (i.e. comparison between the estimation of the sectioning point and the ”real” sectioning point). These discriminant functions were compared withGiles Elliot's (1962, 1963) “race-independent” sex function.     

Characterization of 12 highly variable tetranucleotide microsatellite loci for Arctic grayling (Thymallus arcticus) and cross amplification in other Thymallus species

We describe the isolation and development of 12 polymorphic tetranucleotide microsatellite loci for Arctic grayling (Thymallus arcticus). In a sample of 46 fish, we observed between three and 20 alleles per locus. All 12 of these loci were also polymorphic in at least one of the following Thymallus species and subspecies: T. burejensis, T. amurensis, T. thymallus, T. brevirostris, T. grubii, T. arcticus baicalensis and T. arcticus pallasi. These loci will aid in our understanding of the population genetics, behaviour and conservation of grayling species throughout the northern hemisphere.     

Degrees of sexual dimorphism in Cebus and other new world monkeys

Sexual dimorphism in primate species expresses the effects of phylogeny, life history, behavior, and ontogeny. The causes and implications of sexual dimorphism have been studied in several different primates using a variety of morphological databases such as body weight, canine length, and coat color and ornamentation. In addition to these different patterns of dimorphism, the degree to which a species is dimorphic results from a variety of possible causes. In this study we test the general hypothesis that a species highly dimorphic for one size-based index of dimorphism will be equally dimorphic (relative to other species) for other size-based indices. Specifically, the degree and pattern of sexual dimorphism in Cebus and several other New World monkey species is measured using craniometric data as a substitute for the troublesome range of variation in body weight estimates. In general, the rank ordering of species for dimorphism ratios differs considerably across neural vs. non-neural functional domains of the cranium. The relative degree of sexual dimorphism in different functional regions of the cranium is affected by the independent action of natural selection on those regions. Regions of the cranium upon which natural selection is presumed to have acted within a species show greater degrees of dimorphism than do the same regions in closely related taxa. Within Cebus, C. apella is consistently more dimorphic than other Cebus species for facial measurements, but not for neural or body weight measurements. The pattern in C. apella indicates no single best measurement of the degree of dimorphism in a species; rather, the relative degree of dimorphism applies only to the region being measured and may be enhanced by other selective pressures on morphology. Am J Phys Anthropol 107:243–256, 1998. © 1998 Wiley-Liss, Inc.     

Macroecological patterns of sexual size dimorphism in turtles of the world

Sexual size dimorphism (SSD) is a well‐documented phenomenon in both plants and animals; however, the ecological and evolutionary mechanisms that drive and maintain SSD patterns across geographic space at regional and global scales are understudied, especially for reptiles. Our goal was to examine geographic variation of turtle SSD and to explore ecological and environmental correlates using phylogenetic comparative methods. We use published body size data on 135 species from nine turtle families to examine how geographic patterns and the evolution of SSD are influenced by habitat specialization, climate (annual mean temperature and annual precipitation) and climate variability, latitude, or a combination of these predictor variables. We found that geographic variation, magnitude and direction of turtle SSD are best explained by habitat association, annual temperature variance and annual precipitation. Use of semi‐aquatic and terrestrial habitats was associated with male‐biased SSD, whereas use of aquatic habitat was associated with female‐biased SSD. Our results also suggest that greater temperature variability is associated with female‐biased SSD. In contrast, wetter climates are associated with male‐biased SSD compared with arid climates that are associated with female‐biased SSD. We also show support for a global latitudinal trend in SSD, with females being larger than males towards the poles, especially in the families Emydidae and Geoemydidae. Estimates of phylogenetic signal for both SSD and habitat type indicate that closely related species occupy similar habitats and exhibit similar direction and magnitude of SSD. These global patterns of SSD may arise from sex‐specific reproductive behaviour, fecundity and sex‐specific responses to environmental factors that differ among habitats and vary systematically across latitude. Thus, this study adds to our current understanding that while SSD can vary dramatically across and within turtle species under phylogenetic constraints, it may be driven, maintained and exaggerated by habitat type, climate and geographic location.     

Reproductive skew drives patterns of sexual dimorphism in sponge-dwelling snapping shrimps

Sexual dimorphism is typically a result of strong sexual selection on male traits used in male–male competition and subsequent female choice. However, in social species where reproduction is monopolized by one or a few individuals in a group, selection on secondary sexual characteristics may be strong in both sexes. Indeed, sexual dimorphism is reduced in many cooperatively breeding vertebrates and eusocial insects with totipotent workers, presumably because of increased selection on female traits. Here, we examined the relationship between sexual dimorphism and sociality in eight species of Synalpheus snapping shrimps that vary in social structure and degree of reproductive skew. In species where reproduction was shared more equitably, most members of both sexes were physiologically capable of breeding. However, in species where reproduction was monopolized by a single individual, a large proportion of females—but not males—were reproductively inactive, suggesting stronger reproductive suppression and conflict among females. Moreover, as skew increased across species, proportional size of the major chela—the primary antagonistic weapon in snapping shrimps—increased among females and sexual dimorphism in major chela size declined. Thus, as reproductive skew increases among Synalpheus, female–female competition over reproduction appears to increase, resulting in decreased sexual dimorphism in weapon size.     

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.     

Offspring sex ratio in the Common Tern Sterna hirundo, a species with negligible sexual size dimorphism

In many vertebrates, male offspring are affected more than female offspring by adverse conditions during growth, resulting in facultative adjustment of offspring sex ratio by parents in response to social and environmental conditions during breeding. The greater vulnerability of male offspring is generally attributed to their higher energy requirements associated with their larger size, although greater sensitivity to adverse conditions could be related to other factors such as negative effects of androgens on male physiology. To control for sexual differences in body size, we examined variation in offspring sex ratio in the Common Tern Sterna hirundo , a species with negligible sexual size dimorphism. In this species, the last-laid egg (termed the c-egg) is smaller than the first two and hatches last, so that the chick obtains relatively little food and hence has a low probability of survival to fledging. This species thus provides a powerful model for examining sex-linked mortality and sex ratio variation under natural conditions. We found that the sex ratio of c-eggs, but not of earlier laid eggs, was significantly biased in favour of females. Chicks hatched from c-eggs (termed c-chicks) had low survival but female c-chicks had significantly higher survival than male c-chicks. These data provide strong evidence that factors other than sexual size dimorphism are responsible for producing greater vulnerability of male offspring to adverse conditions during growth.     

A comparison of altitudinal species richness patterns of bryophytes with other plant groups in Nepal, Central Himalaya

Aim To explore species richness patterns in liverworts and mosses along a central Himalayan altitudinal gradient in Nepal (100–5500 m a.s.l.) and to compare these patterns with patterns observed for ferns and flowering plants. We also evaluate the potential importance of Rapoport’s elevational rule in explaining the observed richness patterns for liverworts and mosses. Location Nepal, Central Himalaya. Methods We used published data on the altitudinal ranges of over 840 Nepalese mosses and liverworts to interpolate presence between maximum and minimum recorded elevations, thereby giving estimates of species richness for 100‐m altitudinal bands. These were compared with previously published patterns for ferns and flowering plants, derived in the same way. Rapoport’s elevational rule was assessed by correlation analyses and the statistical significance of the observed correlations was evaluated by Monte Carlo simulations. Results There are strong correlations between richness of the four groups of plants. A humped, unimodal relationship between species richness and altitude was observed for both liverworts and mosses, with maximum richness at 2800 m and 2500 m, respectively. These peaks contrast with the richness peak of ferns at 1900 m and of vascular plants, which have a plateau in species richness between 1500 and 2500 m. Endemic liverworts have their maximum richness at 3300 m, whereas non‐endemic liverworts show their maximum richness at 2700 m. The proportion of endemic species is highest at about 4250 m. There is no support from Nepalese mosses for Rapoport’s elevational rule. Despite a high correlation between altitude and elevational range for Nepalese liverworts, results from null simulation models suggest that no clear conclusions can be made about whether liverworts support Rapoport’s elevational rule. Main conclusions Different demands for climatic variables such as available energy and water may be the main reason for the differences between the observed patterns for the four plant groups. The mid‐domain effect may explain part of the observed pattern in moss and liverwort richness but it probably only works as a modifier of the main underlying relationship between climate and species richness.