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.     

Sexual size dimorphism and male combat in snakes

Summary This paper reviews published literature on snakes to test the hypothesis that large male size, relative to female size, evolves because of the advantage it confers in male combat. Analysis of the data reveals a high correlation between the occurrence of male combat, and sexual dimorphism in which the male is the larger sex. This correlation holds (i) within the total sample of snake species (n=224), (ii) within the family Colubridae (n=134), and (iii) in a comparison between the eight families of snakes for which data are available. These results strongly support the hypothesis that large male size is an adaptation to intrasexual competition. The analysis also shows that females are larger than males in about 66% of snake species, that male combat is known in only about 15% of species, and that both sexual size dimorphism and the incidence of male combat tend to be distributed along taxonomic lines.     

Sexual selection explains Rensch's rule of allometry for sexual size dimorphism

In 1950, Rensch first described that in groups of related species, sexual size dimorphism is more pronounced in larger species. This widespread and fundamental allometric relationship is now commonly referred to as 'Rensch's rule'. However, despite numerous recent studies, we still do not have a general explanation for this allometry. Here we report that patterns of allometry in over 5300 bird species demonstrate that Rensch's rule is driven by a correlated evolutionary change in females to directional sexual selection on males. First, in detailed multivariate analysis, the strength of sexual selection was, by far, the strongest predictor of allometry. This was found to be the case even after controlling for numerous potential confounding factors, such as overall size, degree of ornamentation, phylogenetic history and the range and degree of size dimorphism. Second, in groups where sexual selection is stronger in females, allometry consistently goes in the opposite direction to Rensch's rule. Taken together, these results provide the first clear solution to the long-standing evolutionary problem of allometry for sexual size dimorphism: sexual selection causes size dimorphism to correlate with species size.     

Sexual size dimorphism in anurans

Several hypotheses have been proposed to explain the direction and extent of sexual size dimorphism in anurans (in which males are usually smaller than females) as a result of sexual selection. Here, we present an analysis to test the hypothesis that sexual dimorphism in anurans is largely a function of differences between the sexes in life-history strategies. Morphological and demographic data for anurans were collected from the literature, and the mean size and age in each sex were calculated for 51 populations, across 30 species and eight genera. Comparisons across 14 Rana species, eight Bufo species and across the genera showed a highly significant relationship between size dimorphism, measured using the female-male size ratio, and mean female-male age difference. A comparison of a subset of 17 of these species for which phylogenetic information was available, using the method of independent contrasts, yielded a similar result. These results indicate that most of the variation in size dimorphism in the anura can be explained in terms of differences in the age structure between the sexes in breeding populations. If sexual selection has an effect on size dimorphism in anurans, it is likely to be only a secondary one.     

Sexual size dimorphism in northern giant petrels: ecological correlates and scaling

Northern giant petrels ( Macronectes halli ) are among the largest and most sexually size dimorphic species of seabirds, with females being only 80% the mass of males. Both sexes scavenge on seal and penguin carrion in the sub-Antarctic ecosystem, but during the breeding season females also feed extensively on other marine food resources and show more pelagic habits than males. The outstanding sexual segregation in foraging and feeding ecology in northern giant petrels suggests that mechanisms maintaining sexual size dimorphism by ecological factors may be operating. I evaluated this possibility by examining ecological correlates with body size and by static allometry analyses. Fledging sex ratio in four consecutive years did not depart from parity. There was no assortative mating by size neither association between the male size with the breeding performance. By contrast, smaller females raised their chick in better condition. Moreover, bill size showed a size dimorphism beyond that expected by body size dimorphism, i.e. when controlling for body mass, males showed relatively longer bill than females. This trait did not deviate from isometry with respect to body size and its phenotypic variability was low, suggesting that the disproportionately large bill of males is related to their more scavenging life style compared to females. In general, the increase and maintenance of sexual size dimorphism in giant petrels is more consistent with an ecological causation rather than a result of sexual selection.     

Sexual selection, sexual size dimorphism and Rensch's rule in Odonata

Odonata (dragonflies and damselflies) exhibit a range of sexual size dimorphism (SSD) that includes species with male-biased (males > females) or female-biased SSD (males < females) and species exhibiting nonterritorial or territorial mating strategies. Here, we use phylogenetic comparative analyses to investigate the influence of sexual selection on SSD in both suborders: dragonflies (Anisoptera) and damselflies (Zygoptera). First, we show that damselflies have male-biased SSD, and exhibit an allometric relationship between body size and SSD, that is consistent with Rensch's rule. Second, SSD of dragonflies is not different from unit, and this suborder does not exhibit Rensch's rule. Third, we test the influence of sexual selection on SSD using proxy variables of territorial mating strategy and male agility. Using generalized least squares to account for phylogenetic relationships between species, we show that male-biased SSD increases with territoriality in damselflies, but not in dragonflies. Finally, we show that nonagile territorial odonates exhibit male-biased SSD, whereas male agility is not related to SSD in nonterritorial odonates. These results suggest that sexual selection acting on male sizes influences SSD in Odonata. Taken together, our results, along with avian studies (bustards and shorebirds), suggest that male agility influences SSD, although this influence is modulated by territorial mating strategy and thus the likely advantage of being large. Other evolutionary processes, such as fecundity selection and viability selection, however, need further investigation.     

Sexual size dimorphism in seabirds: sexual selection, fecundity selection and differential niche-utilisation

Seabirds exhibit a range of sexual size dimorphism (SSD) that includes both male-biased (males>females) and female-biased SSD (males相似文献   

Sexual size dimorphism and sexual selection in turtles (order testudines)

Summary This paper combines published and original data on sexual size dimorphism, reproductive behavior, and habitat types in turtles. Our major finding is that observed patterns of sexual size dimorphism correlate with habitat type and male mating strategy. (1) In most terrestrial species, males engage in combat with each other. Males typically grow larger than females. (2) In semiaquatic and bottom-walking aquatic species, male combat is less common, but males often forcibly inseminate females. As in terrestrial species, males are usually larger than females. (3) In truly aquatic species, male combat and forcible insemination are rare. Instead, males utilize elaborate precoital displays, and female choice is highly important. Males are usually smaller than females.We interpret these correlations between sexual behavior and size dimorphism in terms of sexual selection theory: males are larger than females when large male size evolves as an adaptation to increase success in male combat, or to enable forcible insemination of females. In contrast, males are usually smaller than females where small size in males evolves to increase mobility (and hence, ability to locate females), or because selection for increased fecundity may result in increased female size. In turtle species with male combat or forcible insemination, the degree of male size superiority increases with mean species body size.     

Sexual size dimorphism in parasitoid wasps

Sexual dimorphism in body length and proportion of overlap between the ranges of body length for males and females were estimated for 361 species of parasitoid wasps from 21 families. In most species, females are generally larger than males, though the range of male and female sizes overlap. Species in the family Ichneumonidae differ significantly from species in other families in three ways: (1) ichneumonids on average are larger, (2) in most species, females are generally smaller than males, and (3) on average, proportion overlap between the ranges of body length for males and females is greater. At present, there is a paucity of life history data on parasitoid wasp species for which size dimorphism is known. Thus it is not clear why ichneumonids differ from species in other families. Possible evolutionary explanations for variation in dimorphism among parasitoid wasp species are discussed.     

Sexual size dimorphism within species increases with body size in insects

Studies examining interspecific differences in sexual size dimorphism (SSD) typically assume that the degree of sexual differences in body size is invariable within species. This work was conducted to assess validity of this assumption. As a result of a systematic literature survey, datasets for 158 insect species were retrieved. Each dataset contained adult or pupal weights of males and females for two or more different subsets, typically originating from different conditions during immature development. For each species, an analysis was conducted to examine dependence of SSD on body size, the latter variable being used as a proxy of environmental quality. A considerable variation in SSD was revealed at the intraspecific level in insects. The results suggest that environmental conditions may strongly affect the degree, though not the direction of SSD within species. In most species, female size appeared to be more sensitive to environmental conditions than male size: with conditions improving, there was a larger relative increase in female than male size. As a consequence, sexual differences in size were shown to increase with increasing body size in species with female-biased SSD (females were the larger sex in more than 80% of the species examined). The results were consistent across different insect orders and ecological subdivisions. Mechanisms leading to intraspecific variation in SSD are discussed. This study underlines the need to consider intraspecific variation in SSD in comparative studies.     

Sexual size dimorphism and selection in the wild in the waterstrider Aquarius remigis: Body size,components of body size and male mating success

Sexual size dimorphism is assumed to be adaptive and is expected to evolve in response to a difference in the net selection pressures on the sexes. Although a demonstration of sexual selection is neither necessary nor sufficient to explain the evolution of sexual size dimorphism, sexual selection is generally assumed to be a major evolutionary force. If contemporary sexual selection is important in the evolution and maintenance of sexual size dimorphism then we expect to see concordance between patterns of sexual selection and patterns of sexual dimorphism. We examined sexual selection in the wild, acting on male body size, and components of body size, in the waterstrider Aquarius remigis, as part of a long term study examining net selection pressures on the two sexes in this species. Selection was estimated on both a daily and annual basis. Since our measure of fitness (mating success) was behavioral, we estimated reliabilities to determine if males perform consistently. Reliabilities were measured as ? statistics and range from fair to perfect agreement with substantial agreement overall. We found significant univariate sexual selection favoring larger total length in the first year of our study but not in the second. Multivariate analysis of components of body size revealed that sexual selection for larger males was not acting directly on total length but on genital length. Sexual selection for larger male body size was opposed by direct selection favoring smaller midfemoral lengths. While males of this species are smaller than females, they have longer genital segments and wider forefemora. Patterns of contemporary sexual selection and sexual size dimorphism agree only for genital length. For total length, and all other components of body size examined, contemporary sexual selection was either nonsignificant or opposed the pattern of size dimporhism. Thus, while the net pressures of contemporary selection for the species may still act to maintain sexual size dimorphism, sexual selection alone does not.     

Sexual selection for small size in male mosquitofish (Gambusia holbrooki)

In many poeciliid fishes, large males which court females coexist with small males which sneak-copulate. It is unclear whether these two tactics represent two evolutionarily stable strategies or if sneaking is a conditional strategy adopted by small, unattractive males. We studied the success of sneaky copulation by looking for sperm in the gonoduct of females after they were kept for 48 h with a male. A logistic regression analysis showed that the probability of a female being inseminated increased with female length and decreased with male length. The length of the male relative to that of the female was the best predictor of success. This result was confirmed using virgin females, thereby excluding any possible confounding effect due to the release of sperm from previous copulations. Sperm counts suggested that large males do not compensate for their reduced copulatory success by releasing larger sperm numbers. Behavioural data indicate that the advantages to small males are twofold: they have a greater chance to approach females from behind without being detected, and manoeuvre better when inserting the gonopodium into the female''s gonoduct. The selective advantage of small size might explain male dwarfism in poeciliids. Our results also suggest that small males adopting the sneaky tactic may be as successful as large males adopting courtship, and that alternative mating strategies may be maintained by negative density-dependent selection.     

Sexual dimorphism of tooth size in anthropoids

We have examined the size of the canine and postcanine teeth of cebid and catarrhine primates in relation to each other, to jaw size and to body weight. We have found that the canine size of males is large enough to be limited by jaw shape and size. A large contribution of P4 to the postcanine row is associated with smaller canines in males. Neither factor seems to limit canine size in females. The females of a small number of species possess enlarged canines. Much of the variation of the postcanine row can be described by the ratio of the (nominal) crown areas of M1 to M3. This ratio is monomorphic which conforms with the general lack of dietary dimorphism in primates. A brief discussion of the evolution of canine size is offered with a new suggestion to account for canine reduction in male hominids.     

Sexual dimorphism in the size of nuclei with dioecious plants

Measurements of the size of the nuclei of dioecious plants showed that the nuclei of male and female plants differ in agreement with the larger quantity of chromatin. The male. plants ofRumex acetosella andMelandrium album had larger nuclei, their Y chromosome being larger than the X chromosome, the same is true forRumex acetosa where the Y chromosome is smaller but there are two in the set.Ginkgo biloba had larger female nuclei because the Y chromosome was smaller than the X. The curves obtained by grouping all the nuclei of both sexes never had two peaks with regard to the small differences between the classes of maximum frequency.     

Sexual size dimorphism in species with asymptotic growth after maturity

If animals mature at small sizes and then grow to larger asymptotic sizes, many factors can affect male and female size distributions. Standard growth equations can be used to study the processes affecting sexual size dimorphism in species with asymptotic growth after maturity. This paper first outlines the effects of sex differences in growth and maturation patterns on the direction and degree of sexual dimorphism. The next section considers the effects of variation in age structure or growth rates on adult body sizes and sexual size dimorphism. Field data from a crustacean, fish, lizard and mammal show how information on a species' growth and maturation patterns can be used to predict the relationships between male size, female size and sexual size dimorphism expected if a series of samples from the same population simply differed with respect to their ages or growth rates. The last section considers ecological or behavioural factors with different effects on the growth, maturation, survival or movement patterns of the two sexes. This study supports earlier suggestions that information on growth and maturation patterns may be useful, if not essential, for comparative studies of sexual size dimorphism in taxa with asymptotic growth after maturity.     

Sexual selection maintains whole-body chiral dimorphism in snails

Although the vast majority of higher animals are fixed for one chiral morph or another, the cause for this directionality is known in only a few cases. In snails, for example, rare individuals of the opposite coil are unable to mate with individuals of normal coil, so directionality is maintained by frequency-dependent selection. The snail subgenus Amphidromus presents an unexplained exception, because dextral (D) and sinistral (S) individuals occur sympatrically in roughly equal proportions (so-called 'antisymmetry') in most species. Here we show that in Amphidromus there is sexual selection for dimorphism, rather than selection for monomorphism. We found that matings between D and S individuals occur more frequently than expected by chance. Anatomical investigations showed that the chirality of the spermatophore and the female reproductive tract probably allow a greater fecundity in such inter-chiral matings. Computer simulation confirms that under these circumstances, sustained dimorphism is the expected outcome.     

Sexual dimorphism and human tooth size differences

Current male/female differences in tooth size are due to the male/female differences in body bulk that exist in any given human population. These differences are residues of the sexual dimorphism that was maintained for adaptive reasons during the Middle Pleistocene. Late in the Pleistocene the development of food processing techniques led to the reduction of both male and female dental dimensions. Dental sexual dimorphism, however, was maintained until the very end of the Pleistocene when the hunting of large game animals by crude techniques was replaced by a focus on great numbers of small game caught by more sophisticated means and by an increasing utilization of plant foods. The subsequent reduction in dimorphism represents the actions of the Probable Mutation Effect operating under conditions of relaxed selection. The conclusion offered is that the smallest degree of sexual dimorphism visible in the modern world is to be found among those populations that are separated by the greatest interval of time from precursors who depended for their survival on a Pleistocene big game hunting mode of subsistence.     

Sexual selection: harem size and the variance in male reproductive success

Sexual selection is potentially stronger than natural selection when the variance in male reproductive fitness exceeds all other components of fitness variance combined. However, measuring the variance in male reproductive fitness is difficult when nonmating males are absent, inconspicuous, or otherwise difficult to find. Omitting the nonmating males inflates estimates of average male reproductive success and diminishes the variance, leading to underestimates of the potential strength of sexual selection. We show that, in theory, the proportion of the total variance in male fitness owing to sexual selection is approximately equal to H, the mean harem size, as long as H is large and females are randomly distributed across mating males (i.e., Vharem=H). In this case, mean harem size not only provides an easy way to estimate the potential strength of sexual selection but also equals the opportunity for sexual selection, I(mates). In nature, however, females may be overdispersed with VharemH. We show that H+(k-1) is a good measure of the opportunity for sexual selection, where k is the ratio Vharem/H. A review of mating system data reveals that in nature the median ratio for Vharem/H is 1.04, but as H increases, females tend to become more aggregated across mating males with V(harem) two to three times larger than H.