Environmental effects on sexual size dimorphism of a seed-feeding beetle

Sexual size dimorphism is widespread in animals but varies considerably among species and among populations within species. Much of this variation is assumed to be due to variance in selection on males versus females. However, environmental variables could affect the development of females and males differently, generating variation in dimorphism. Here we use a factorial experimental design to simultaneously examine the effects of rearing host and temperature on sexual dimorphism of the seed beetle, Callosobruchus maculatus. We found that the sexes differed in phenotypic plasticity of body size in response to rearing temperature but not rearing host, creating substantial temperature-induced variation in sexual dimorphism; females were larger than males at all temperatures, but the degree of this dimorphism was smallest at the lowest temperature. This change in dimorphism was due to a gender difference in the effect of temperature on growth rate and not due to sexual differences in plasticity of development time. Furthermore, the sex ratio (proportion males) decreased with decreasing temperature and became female-biased at the lowest temperature. This suggests that the temperature-induced change in dimorphism is potentially due to a change in non-random larval mortality of males versus females. This most important implication of this study is that rearing temperature can generate considerable intraspecific variation in the degree of sexual size dimorphism, though most studies assume that dimorphism varies little within species. Future studies should focus on whether sexual differences in phenotypic plasticity of body size are a consequence of adaptive canalization of one sex against environmental variation in temperature or whether they simply reflect a consequence of non-adaptive developmental differences between males and females.     

The relationship between ecological segregation and sexual body size dimorphism in large herbivores

Ecological segregation (sexual differences in diet or habitat use) in large herbivores has been intimately linked to sexual body size dimorphism, and may affect both performance and survival of the sexes. However, no one has tested comparatively whether segregation occurs at a higher frequency among more dimorphic species. To test this comparatively, data on sex-specific diet, habitat use and body size of 40 species of large herbivores were extracted from the literature. The frequency of ecological segregation was higher among more dimorphic herbivores; however, this was only significant for browsers. This provides the first evidence that segregation is more common among more dimorphic species. The comparative evidence supported the nutritional-needs hypothesis over the incisor breadth hypothesis, as there was no difference in frequency of segregation between seasons with high and low resource levels, and since segregation was also evident among browsers. Whether the absence of a correlation between ecological segregation and level of sexual body size dimorphism for intermediate feeders and grazers is due to biological differences relative to browsers or to the fact that the monomorphic species included in the analysis were all browsers is discussed. Received: 18 August 1999 / Accepted: 31 January 2000     

Pelvic dimorphism in relation to body size and body size dimorphism in humans

Many mammalian species display sexual dimorphism in the pelvis, where females possess larger dimensions of the obstetric (pelvic) canal than males. This is contrary to the general pattern of body size dimorphism, where males are larger than females. Pelvic dimorphism is often attributed to selection relating to parturition, or as a developmental consequence of secondary sexual differentiation (different allometric growth trajectories of each sex). Among anthropoid primates, species with higher body size dimorphism have higher pelvic dimorphism (in converse directions), which is consistent with an explanation of differential growth trajectories for pelvic dimorphism. This study investigates whether the pattern holds intraspecifically in humans by asking: Do human populations with high body size dimorphism also display high pelvic dimorphism? Previous research demonstrated that in some small-bodied populations, relative pelvic canal size can be larger than in large-bodied populations, while others have suggested that larger-bodied human populations display greater body size dimorphism. Eleven human skeletal samples (total N: male = 229, female = 208) were utilized, representing a range of body sizes and geographical regions. Skeletal measurements of the pelvis and femur were collected and indices of sexual dimorphism for the pelvis and femur were calculated for each sample [ln(M/F)]. Linear regression was used to examine the relationships between indices of pelvic and femoral size dimorphism, and between pelvic dimorphism and female femoral size. Contrary to expectations, the results suggest that pelvic dimorphism in humans is generally not correlated with body size dimorphism or female body size. These results indicate that divergent patterns of dimorphism exist for the pelvis and body size in humans. Implications for the evaluation of the evolution of pelvic dimorphism and rotational childbirth in Homo are considered.     

Size-sex relationship and sexual dimorphism in JapaneseArisaema (Araceae)

The size-sex relationship and sexual differences of sixArisaema species native to Japan were investigated. The size-sex relationship showed almost the same pattern in all species. When the plant was small in size, the sex expression was male, and sex expression changed from male to female as the plant grew larger. Male ratios decreased rapidly around a critical size, but this critical size differed from one species to another. Sexual differences were detected in reproductive structures and behavior, although no difference was detected in vegetative structures. The stoutness, longevity and inner tissue of the scape showed remarkable differences between males and females, and this difference was represented most clearly as the size-weight relationship. Earlier initiation of flowering in males was also observed. No difference was found in resource allocation to reproductive structures between male and female plants at the flowering stage. However, a broad variation in the amount of resource allocation to reproductive structures was found at the fruiting stage in female individuals, which was attributed to differences in the setting rate of mature fruits.     

Sexual size dimorphism,growth, and maturity of the fluvial eight-barbel loach in the Kako River,Japan

The maturation and growth pattern of the fluvial eight-barbel loach Lefua sp. (Japanese name: nagare-hotoke-dojo), an endangered species, was investigated using an individual identification-recapture method from 1995 to 1998 in an upper reach of a headwater tributary of the Kako River, Hyogo Prefecture, Japan. Based on observations of the gonads through the abdominal skin, the loach was estimated to breed mostly from May to July. All the males matured by age 1⁺, and all the females matured by age 2⁺. Gamete release in all individuals of both males and females was predicted from recaptured loaches during each breeding season. The standard length of mature females was significantly larger than that of males, showing sexual size dimorphism (SSD). The maximum sizes recorded were 75.4 mm SL for females and 61.2 mm SL for males. Both males and females of immature specimens grew mainly from May to November, including the breeding season, with no significant differences in growth rates between them. After sexual maturity, both males and females grew mainly from July to October (or November), after the breeding season, and the females exhibited higher growth rates than males. Therefore, SSD of the species seems to be attributable to the different growth rates after maturity. The longevity of the loach was estimated to exceed ten years based on individual growth patterns of various sizes during the survey period. It is likely that the loach has an iteroparous life history, breeding every year, and moderate growth rates after maturity.     

Numerical analysis of sexual dimorphism inSaguinus dentition

Among New World monkeys, more or less sexual dimorphism exists in the dentition, especially in the Cebidae. On the other hand, the Callitrichidae includingSaguinus are said to be characterized by a broad lack of sexual dimorphism with the exception of the reproductive organs. In the present article, sexual dimorphism in the dentition of someSaguinus species was reconfirmed using univariate and multivariate analytical methods. The results of the analysis were as follows: (1) there is no sexual dimorphism in the canine tooth size, except for the upper canine ofS. geoffroyi and lower canine ofS. mystax; (2) the overall tooth size difference between males and females is slight or none inS. geoffroyi, S. leucopus, andS. fuscicollis, relatively small inS. oedipus andS. mystax, and rather larger inS. midas; (3) an overall difference in shape factor between both sexes exists in all species ofSaguinus to a greater or lesser extent; (4) although only slight sexual dimorphism is recognized in the canine tooth itself, sexual dimorphism does exist in some adjacent teeth of the canine in a few species; and (5) there are some interspecific differences in the magnitude of the sexual dimorphism of theSaguinus dentition and these differences are more evident in species inhabiting the peripheral regions of the distribution areas of this genus. Taking all the evidence obtained into account, the sexual dimorphism in theSaguinus dentition must be re-investigated in comparison with other genera of the Callitrichidae.     

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.     

Climate change and sexual size dimorphism in an Arctic spider

Climate change is advancing the onset of the growing season and this is happening at a particularly fast rate in the High Arctic. However, in most species the relative fitness implications for males and females remain elusive. Here, we present data on 10 successive cohorts of the wolf spider Pardosa glacialis from Zackenberg in High-Arctic, northeast Greenland. We found marked inter-annual variation in adult body size (carapace width) and this variation was greater in females than in males. Earlier snowmelt during both years of its biennial maturation resulted in larger adult body sizes and a skew towards positive sexual size dimorphism (females bigger than males). These results illustrate the pervasive influence of climate on key life-history traits and indicate that male and female responses to climate should be investigated separately whenever possible.     

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.     

Factors influencing the degree of sexual size dimorphism within and among calanoid copepod species

Populations of Diaptomus leptopus (Copepoda: Calanoida) and other calanoid copepods exhibit varying degrees of sexual size dimorphism. We examined whether intraspecific or interspecific variation in dimorphism could be explained by allometry, and we examined the relationship between adult size attained and development rate to determine any relationship between the two. We compared the degree of sexual size dimorphism in D. leptopus and in other calanoid copepods inhabiting temporary and permanent habitats. Allometry did not explain variation in sexual size dimorphism within or among populations or among species. Permanence of habitat affected the degree of dimorphism: dimorphism was greater within and among species inhabiting temporary environments. Non-significant differences in development rate were found among populations and significant differences were found between sexes of D. leptopus when reared under identical laboratory conditions: males developed more rapidly than females but there was no general relationship between development rate and adult size. Potential adaptive hypotheses to explain the differences between populations inhabiting temporary and permanent habitats are discussed.     

Sexual size dimorphism in the ontogeny of the solitary predatory wasp Symmorphus allobrogus (Hymenoptera: Vespidae)

Sex-specific patterns of individual growth, resulting in sexual size dimorphism (SSD), are a little studied aspect of the ontogeny related to the evolutionary history and affected by the ecology of a species. We used empirical data on the development of the predatory wasp Symmorphus allobrogus (Hymenoptera, Vespidae) to test the hypotheses that sexual differences of growth resulting in the female-biased SSD embrace the difference in (1) the egg size and the starting size of larva, (2) the larval development duration, and (3) the larval growth rate. We found that eggs developing into males and females have significant differences in size. There was no significant difference between the sexes in the duration of larval development. The relative growth rate and the food assimilation efficiency of female larvae were significantly higher than compared to those of male larvae. Thus, the SSD of S. allobrogus is mediated mainly by sexual differences in egg size and larval growth rate.     

Contrasting patterns of body size for Daphnia species that segregate by habitat

Summary We investigate how body size of two coexisting Daphnia species varies among 7 lakes that represent a gradient of predation risk. The two species segregate vertically in stratified lakes; D. galeata mendotae is typically smaller and more eplimnetic than D. pulicaria. The extent of vertical habitat partitioning, however, varies seasonally within and among lakes in apparent response to predation intensity by epilimnetic planktivorous fishes. Daphnia pulicaria uses the epilimnion at low levels of fish predation but is restricted to the hypolimnion under high fish predation, whereas D. galaeta mendotae always utilizes the epilimnion. The species display contrasting patterns of genetic variation in neonate size and size at maturity. D. pulicaria is larger in lakes with higher fish and Chaoborus densities whereas D. galeata mendotae is smaller. This contrast in body size in lakes with high predation is associated with greater habitat segregation in those lakes. In lakes with low predation risk, the two species are similar in body size at birth and maturity.Authorship order alphabetical     

Sexual behaviour and evolution of sexual dimorphism in body size in Jaera (Isopoda Asellota)

Individuals of the genus Jaera do not mate at random. In the species from the Mediterranean group, J. italica and. J. nordmanni, large males and medium sized females are at an advantage and their sizes are positively assorted. These effects are attributable to sexual competition between males. In the Ponlo-caspian species J. istri, no advantage of large males exists, but sexual selection could be the cause for a long passive phase prior to copulation and for normalizing selection upon female size at pairing. In the Atlantic species, J. albifrons, no selection can be ascertained.
Differential mating success in males appears as one of the causes of the evolution of sexual dimorphism in body size, which makes males larger, of equal size, or smaller than females according to the species. The reason for this reversal in dimorphism seems to differ in the two sexes. Sexual selection provides an explanation for the evolution of male size, while the interspecific changes in female length are more likely due to ecological factors.     

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. Received: 20 March 1996 / Accepted: 10 March 1998     

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.     

Causes of secondary sexual differences in plants — Evidence from extreme leaf dimorphism in Leucadendron (Proteaceae)

In extreme cases leaves in male plants of the dioecious genus Leucadendron (Proteaceae) are up to an order of magnitude smaller than female leaves. This secondary sexual dimorphism (SSD) in leaf size has previously been suggested to be due to intra-male sexual selection, leading to an increase in male allocation to reproduction in dimorphic species. After critically evaluating previous data provided to support this hypothesis, I suggest on both theoretical grounds and on re-analysis that this argument is unlikely and unsupported. Leaf size dimorphism could theoretically evolve directly due to disruptive ecological selection between genders, leading to niche dimorphism either within or between habitats. I test this ecological causation hypothesis by providing data on specific leaf area (sla) and water use efficiency (δ ¹³C) of leaves from males and females of several Leucadendron species. Results confirm the expectation of minimal gender differences. I argue that leaf dimorphism is a consequence of selection on flower size and architecture.