Patterns Of Sexual Size Dimorphism Research Articles

Lifetime selection on adult body size and components of body size in a waterstrider: opposing selection and maintenance of sexual size dimorphism.

Sexual size dimorphism (SSD), the difference in body size between males and females, is common in almost all taxa of animals and is generally assumed to be adaptive. Although sexual selection and fecundity selection alone have often been invoked to explain the evolution of SSD, more recent views indicate that the sexes must experience different lifetime selection pressures for SSD to evolve and be maintained. We estimated selection acting on male and female adult body size (total length) and components of body size in the waterstrider Aquarius remigis during three phases of life history. Opposing selection pressures for overall body size occurred in separate episodes of fitness for females in both years and for males in one year. Specific components of body size were often the targets of the selection on overall body size. When net adult fitness was estimated by combining each individual's fitnesses from all episodes, we found stabilizing selection in both sexes. In addition, the net optimum overall body size of males was smaller than that of females. However, even when components of body size had experienced opposing selection pressures in individual episodes, no components appeared to be under lifetime stabilizing selection. This is the first evidence that contemporary selection in a natural population acts to maintain female size larger than male size, the most common pattern of SSD in nature.

Statistics of sexual size dimorphism

In comparative studies of sexual size dimorphism (SSD), the methods used to quantify dimorphism are controversial. SSD is commonly expressed as a ratio between species mean values of males and females, such as M/F or (M−F)/([M+F]/2), but a number of investigators have suggested that ratios should not be used, mainly because their distributions usually violate the assumptions of parametric statistical tests, or because they lead to spurious relationships that invalidate the interpretation and statistical significance of regressions and correlations. As an alternative to ratios, the comparative study of SSD can be conducted by a combination of regression with sex-specific data and residuals from this regression. Twenty-five data sets were selected from the literature and used to duplicate a variety of statistical procedures commonly employed in studies of SSD. All analyses were repeated with five different ratios and with methods that avoid the calculation of any ratios. These data and a review of the statistical properties of ratios and residuals indicate that: (1) most of the ratios used in the SSD literature are unnecessary, and several commonly used ratios are statistically inferior to others. Only two ratios are needed, one on a logarithmic scale and one on a linear scale; (2) there is no problem with spurious correlation or non-normality when ratios are used in several types of statistical procedures commonly employed in studies of SSD; (3) residuals cannot replace ratios for the evaluation of many questions regarding the pattern of SSD among species; and (4) residuals usually are used incorrectly, leading to misspecified regression equations. Most of the questions for which residuals are used should be addressed by multiple regression. These results apply to studies using comparative methods with or without adjustments for phylogenetic effects.

Patterns of growth and sexual size dimorphism in two species of box turtles with environmental sex determination.

An adaptive explanation for environmental sex determination is that it promotes sexual size dimorphism when larger size benefits one sex more than the other. That is, if growth rates are determined by environment during development, then it is beneficial to match developmental environment to the sex that benefits more from larger size. However, larger size may also be a consequence of larger size at hatching or growing for a longer time, i.e., delayed age at first reproduction. Therefore, the adaptive significance of sexual size dimorphism and environmental sex determination can only be interpreted within the context of both growth and maturation. In addition, in those animals that continue to grow after maturation, sexual size dimorphism at age of first reproduction could differ from sexual size dimorphism at later ages as growth competes for energy with reproduction and maintenance. I compared growth using annuli on carapace scales in two species of box turtles (Terrapene carolina and T. ornata) that have similar patterns of environmental sex determination but, reportedly, have different patterns of sexual size dimorphism. In the populations I studied, sexual size dimorphism was in the same direction in both species; adult females were, on average, larger than adult males. This was due in part to males maturing earlier and therefore at smaller sizes than females. In spite of similar patterns of environmental sex determination, patterns of growth differed between the species. In T. carolina, males grew faster than females as juveniles but females had the larger asymptotic size. In T. ornata, males and females grew at similar rates and had similar asymptotic sizes. Sexual size dimorphism was greatest at maturation because, although males matured younger and smaller, they grew more as adults. There was, therefore, no consistent pattern of faster growth for females that may be ascribed to developmental temperature.

Sexual selection and tail streamers in the barn swallow

The functional significance of elongated, narrow tips of the tail feathers of certain birds, so-called tail streamers, has recently been discussed from an aerodynamic point of view, and the effects of sexual selection on such traits have been questioned. We review our long-term field studies using observational and experimental approaches to investigate natural and sexual selection in the barn swallow, Hirundo rustica, which has sexually size-dimorphic outermost tail feathers. Experimental manipulation of the length of the outermost tail feathers has demonstrated sexual selection advantages of tail elongation and disadvantages of tail shortening, with opposite effects for natural selection in terms of foraging efficiency, haematocrit and survival. These findings are contrary to the prediction of a general deterioration from both shortening and elongation, if the tail trait was determined solely by its effects on aerodynamic efficiency and flight manoeuvrability. Patterns of sexual selection in manipulated birds conform with patterns in unmanipulated birds, and selection differentials for different components of sexual selection in manipulated birds are strongly positively correlated with differentials in unmanipulated birds. Age and sex differences in tail length, and geographical patterns of sexual size dimorphism, are also consistent with sexual selection theory, but inconsistent with a purely natural selection advantage of long outermost tail feathers in male barn swallows.

Body size and sexual size dimorphism in calanoid copepods

Patterns of sexual size dimorphism and body size in calanoid copepods are examined. We hypothesize that favorable conditions for development will result in large body size and high sexual size dimorphism among populations of a given species and that differences in this allometric relationship among species is governed by the male's role in insemination. We confirm that there is a greater advantage to large female size, normally the larger sex, when compared to males, hence leading to selection for developmental patterns favoring high size dimorphism. Individuals from populations of four centropagid copepod species were measured; other sizes were obtained from published sources. In the four species we examined, the relationships between prosome length and both clutch size and the ability to produce multiple clutches with one insemination were determined. Results show a trend toward hyperallometry in all centropagid species examined: sexual size dimorphism increases with increasing size. Large females produce larger clutches and more additional clutches on one insemination. That hyperallometry is not observed in diaptomid copepods may result from the greater role the male plays in reproduction. Males are needed for each clutch produced, hence the selective pressure to be larger is greater than that in the centropagidae.

SEXUAL SELECTION AND THE EVOLUTION OF SEXUAL SIZE DIMORPHISM IN THE WATER STRIDER, AQUARIUS REMIGIS.

Sexual size dimorphism (SSD) is often attributed to sexual selection, particularly when males are the larger sex. However, sexual selection favoring large males is common even in taxa where females are the larger sex, and is therefore not a sufficient explanation of patterns of SSD. As part of a more extensive study of the evolution of SSD in water striders (Heteroptera, Gerridae), we examine patterns of sexual selection and SSD in 12 populations of Aquarius remigis. We calculate univariate and multivariate selection gradients from samples of mating and single males, for two sexually dimorphic traits (total length and profemoral width) and two sexually monomorphic traits (mesofemoral length and wing form). The multivariate analyses reveal strong selection favoring larger males, in spite of the female-biased SSD for this trait, and weaker selection favoring aptery and reduced mesofemoral length. Selection is weakest on the most dimorphic trait, profemoral width, and is stabilizing rather than directional. The pattern of sexual selection on morphological traits is therefore not concordant with the pattern of SSD. The univariate selection gradients reveal little net selection (direct + indirect) on any of the traits, and suggest that evolution away from the plesiomorphic pattern of SSD is constrained by antagonistic patterns of selection acting on this suite of positively correlated morphological traits. We hypothesize that SSD in A. remigis is not in equilibrium, a hypothesis that is consistent with both theoretical models of the evolution of SSD and our previous studies of allometry for SSD. A negative interpopulation correlation between the intensity of sexual selection and the operational sex ratio supports the hypothesis that, as in several other water strider species, sexual selection in A. remigis occurs through generalized female reluctance rather than active female choice. The implications of this for patterns of sexual selection are discussed.

Geographic Variation and Sexual Size Dimorphism in Mauremys mutica (Cantor, 1842) (Reptilia: Bataguridae), with Description of a New Subspecies from the Southern Ryukyus, Japan

Abstract The batagurid turtle, Mauremys mutica, is widely distributed in tropical to temperate East Asia. Analyses of morphometric characters and coloration revealed that the southern Ryukyu populations of this species are much diverged from the other populations, presumably as a result of their long geographical isolation. We describe those populations as a new subspecies, M. m. kami. Analysis of geographic variation also suggested that distinctly isolated populations of the central and northern Ryukyus, and Kyoto and Shiga Prefectures of central Japan have originated from animals artificially introduced from the Yaeyama Group, and Taiwan, respectively. We confirmed the absence of “larger female” sexual size dimorphism (SSD) in M. mutica unlike most other aquatic batagurids, and further demonstrated variation in SSD pattern between the subspecies: in M. m. kami, the adult male has a significantly greater carapace length than adult females, whereas the adult carapace length does not differ significantly b...

Sex ratios, mating behavior and sexual size dimorphism of the northern water snake, Nerodia sipedon

Competition among males to mate is generally associated with male-biased size dimorphism. In this study we examine mating behavior in the northern water snake (Nerodia sipedon), a species in which males are much smaller than females despite substantial competition among males to mate. Competition among males was a consequence of a male-biased operational sex ratio due to slightly higher female mortality from a birth sex ratio of 1 : 1, and, in 1 year, more synchronous and longer mating activity by males. Approximately one-third of both males and females appeared not to mate in a given year. Larger males were generally more likely to attempt mating, but size did not explain the variance in the number of aggregations in which individual males participated. Within aggregations, males that were successful at achieving intromission were larger than unsuccessful males in 1 of 2 years. Variation in condition (mass relative to length) and relative tail length were not generally useful predictors of either mating effort or success in males. Because large size was often advantageous to males, sexual size dimorphism appeared not to be a consequence of sexual selection favoring smaller males. Because sexual dimorphism was evident at birth, and both males and females matured sexually at about 4 years, sexual dimorphism was not simply a consequence of one sex growing at the maximum rate for longer. Female fecundity increased with size, and sex differences in size-fecundity relations may underly the pattern of sexual size dimorphism. However, because multiple mating by females is common, sperm competition is likely to be important in determining male reproductive success. Therefore, allocation of energy to sperm rather than growth may also prove to be an important influence on male growth rates and sexual size dimorphism.

Sexual selection in the barn swallow (Hirundo rustica). V. Geographic variation in ornament size

AbstractModels of sexual selection in a cline predict the patterns of clinal variation in female mate preference and male secondary sexual characters. These predictions were tested for the nominate subspecies of the barn swallow Hirundo rustica which demonstrates clinal variation in morphology, with several characters in both sexes showing increasing size at higher latitudes. Sexual size dimorphism in the length of the tail ornament and the short, central tail feathers increase with increasing latitude while size dimorphism in other morphological characters is independent of latitude. The main reason for the two divergent patterns of sexual size dimorphism appears to be the higher foraging cost of having a long tail ornamental at low latitudes. The control of development decreases with increasing latitude as demonstrated by an increasing latitudinal cline in fluctuating asymmetry of tail length. Phenotypic variance in tail length increases with latitude in males, but not in females, as shown by the coefficients of variation. Clinal variation in morphology is not due to natural selection associated with a latitudinal increase in the distance between breeding and wintering areas. The geographic patterns of morphological variation suggest that the tail character has diverged geographically as a result of a sexual process of reliable signalling.

Patterns of Sexual Size Dimorphism in Seabirds of the Southern Hemisphere

A review of published data on 99 populations (60 species) of seabirds shows that the degree of sexual size dimorphism varies considerably among species, with males larger than females in some species but smaller in others. Males tend to be larger, relative to females, in populations (1) that have large average body mass, and (2) that feed in areas of the ocean (especially, high latitudes) where primary productivity is high. These correlations are not due to phylogenetic conservatism, because the relationships hold even when the data are analysed cladistically. These patterns are similar in some respects to those seen in other avian lineages, but their adaptive significance remains unclear

Sexual size dimorphism in relation to resource partitioning in North American dabbling ducks

Data from museum specimens were used to examine patterns of sexual size dimorphism in bill morphology in relation to resource partitioning in seven species of North American dabbling ducks (Anas sp.). All species were dimorphic with respect to bill length. Male and female bill length distributions were skewed in opposite directions in five of seven species (P < 0.01). Four of seven species were dimorphic with respect to lamellar density, but male and female distributions were skewed in opposite directions in only two species. The four dimorphic species were positioned between other species along a prey size dimension; species which were not dimorphic had only one neighbour on the prey size dimension. These results are consistent with the hypothesis that intersexual competition for food promotes niche divergence between sexes and contributes to sexual size dimorphism in ducks.

Variation in Age and Size at Maturity of the Slider Turtle (Pseudemys scripta)

The adaptive strategy of the slider turtle, Pseudemys scripta, in terms of age and size at sexual maturity is discussed. Populations from a natural aquatic habitat and from a cooling reservoir on the US DOE Savannah River Plant are compared. (HCR)

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

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 SELECTION AND BODY SIZE IN MALE RED-WINGED BLACKBIRDS.

In most species of birds adult males are larger than adult females. The two major hypotheses proposed to explain this pattern of sexual size dimorphism are appropriately termed the sexual selection and competition avoidance hypotheses. The sexual selection hypothesis states that large size gives males an advantage in intrasexual competition for females or in epigamic displays, activities not engaged in by females. Males will thus be larger than females, and the difference should be greater among polygynous and promiscuous species where there is greater variability in male reproductive success and thus stronger sexual selection (Darwin, 1871; Huxley, 1938; Fisher, 1958). The sexual selection hypothesis, then, explains the relationship between pronounced sexual dimorphism in size and nonmonogamous mating systems in birds (Darwin, 1871; Selander, 1958, 1972; Amadon, 1959; Orians, 1961), a relationship particularly evident among the Icteridae (Selander, 1958). The competition avoidance hypothesis states that sexual size dimorphism is favored because it reduces competition between males and females of the same species (Selander and Giller, 1963; Selander, 1966, 1972; Johnston and Selander, 1973). If size affects the range of resources used (e.g., larger individuals eat larger food items), then those males differing most in size from females would overlap in resource utilization with the fewest individuals and thus have the most resources available. This hypothesis may be especially applicable to monogamous, ter-

Parent Cannibalism of Offspring and Egg Raiding as a Courtship Strategy

In fish, and possibly other groups, egg-guarding males may benefit from the consumption of some of their own genetic offspring by remaining in good condition for subsequent brood cycles. This parasitism of the female's great cytoplasmic commitment to reproduction generates conflicts of interest which provide an evolutionary explanation of several previously enigmatic behavioral patterns. The fundamental conflict obtains from females trying to minimize their egg loss by laying for males that will receive multiple clutches; this means males without eggs are handicapped in courtship. Egg raiding in sticklebacks, wherein eggs are stolen and deposited in the thief's nest, is interpreted as a courtship strategy, partly deceptive and partly honest, designed to convince the female that loss of her eggs to the fasting male will be minimal. The canopied nests of sticklebacks may conceal the absence of eggs and make courtship fanning serve to deceptively entice females to proceed with courtship, because fanning normally indicates that the male has eggs. The shift from the courtship to the parental phase by guardian males occurs because females should refuse to lay for males with perceptibly advanced eggs yet will benefit from eating such eggs. Finally, differences among species in the relative importance of filial cannibalism and heterocannibalism may explain patterns of sexual size dimorphism in fish with parental care. Where heterocannibalism is more important, females should prefer larger males as better brood defenders; thus, epigamic and agonistic sexual selection are reinforcing. But where filial cannibalism is more important, females should prefer small males because they need to consume fewer eggs for their own maintenance; thus, intramale selection for larger body size is countered by female choice of small males.