Growth and food consumption of whiting Merlangius merlangus

In the western Baltic Sea (WBS), whiting Merlangius merlangus is the main piscivorous fish together with cod Gadus morhua. In the present study, we investigate the growth and food consumption rates of WBS M. merlangus and compare the growth rates of males and females with those of M. merlangus in the North Sea (NS). Food consumption rates are estimated directly from sampled stomach contents in the WBS using a gastric evacuation rate model and indirectly by using a static energy‐budget model together with the growth rates. The results indicate that male and female M. merlangus in the WBS have similar feeding and growth strategies, while in the NS M. merlangus show more pronounced differences in food consumption and growth dynamics between the sexes. Female WBS M. merlangus grow significantly slower than their conspecifics in the NS, but there is no significant difference for males. Sexual size dimorphism is seen in both areas, but for M. merlangus in the WBS the difference is less pronounced. Food consumption rates in the WBS differ between seasons, with the lowest food intake in the first 2 quarters of the year and the highest in the 3rd quarter. No differences in consumption rates were seen between males and females, which could be related to the more similar growth pattern seen for M. merlangus in the WBS.     

Sex‐specific food intake in whiting Merlangius merlangus

In this study, the topic of sexual growth dimorphism in whiting Merlangius merlangus is examined. To understand the magnitude and underlying mechanisms, North Sea International Bottom Trawl Survey (IBTS) data and two additional datasets from the third quarter of 2007 and the first quarter of 2012 were analysed. Merlangius merlangus displays distinct differences in growth parameters between males and females, with females reaching a higher asymptotic length (L_∞) than males. To identify the mechanisms which lead to higher growth in females, the quantity and the quality of the diet of M. merlangus in the North Sea were investigated to compare the sex‐specific energy uptake levels. The diet composition did not differ between the sexes, but females had higher stomach content masses than males of the same total length (L_T), and showed lower proportions of empty stomachs. Moreover, female M. merlangus had higher liver and empty stomach masses compared with males of the same size, which indicates additional sex‐specific differences in the metabolic costs and energy allocation patterns. Finally, interannual differences were found in the stomach contents, the share of empty stomachs and liver masses of M. merlangus in the North Sea.     

Species‐specific TaqMan probes for simultaneous identification of (Gadus morhua L.), haddock (Melanogrammus aeglefinus L.) and whiting (Merlangius merlangus L.).

We report the development of a multiplex, single tube (TaqMan) PCR assay for the identification of three commercially important gadoid species, the cod (Gadus morhua L.), the haddock (Melanogrammus aeglefinus L.) and the whiting (Merlangius merlangus L.). The assay unambiguously identifies known adult tissue samples, and ethanol‐preserved eggs from all three species with greater than 98% accuracy, providing a powerful tool for the development of catch independent stock assessment methods, mapping of spawning sites and the detection of commercial fraud in the fishing and food production industries.     

Sexual size dimorphism in caecilian amphibians: analysis, review and directions for future research

Sexual dimorphism, widespread in the animal kingdom, describes differences between the sexes in size, shape and many other traits. Sexual size dimorphism (SSD) plays a significant role in understanding life history evolution and mating systems. The snakelike morphology of limbless caecilian amphibians lacking obvious secondary sexual characters (in contrast to frogs and salamanders) impedes accurate intrasexual comparisons. In this study, sexual size dimorphism in the oviparous caecilian Ichthyophis cf. kohtaoensis, a phylogenetically basal caecilian, was analysed. Females were larger in all body and head characters tested. However, when adjusted to body size (total length), females differed only in their cloacal shape. Clutch volume was positively correlated to female body size, thus female fecundity increased with body size supporting the hypothesis of a fecundity-selected SSD in the oviparous Ichthyophis cf. kohtaoensis. A review of the present SSD data for caecilians shows that many species are monomorphic for body size but show dimorphism in head size, while other species demonstrate female-biased SSD. Male-biased SSD has not been reported for caecilians. To understand life history evolution in caecilians, further studies on the reproductive biology of other taxa are urgently needed, in particular for rhinatrematids and uraeotyphlids. New data will allow phylogenetically controlled comparative analyses to fully explore the pattern of SSD among caecilian lineages.     

The evolution of sexual size dimorphism in cottontail rabbits (Sylvilagus,Leporidae)

In mammals, ‘female‐biased’ sexual size dimorphism (SSD), in which females are larger than males, is uncommon. In the present study, we examined Sylvilagus, a purported case of female‐biased SSD, for evolutionary correlations among species between SSD, body‐size, and life‐history variables. We find that: (1) although most species are female‐biased, the degree and direction of SSD vary more than was previously recognized and (2) the degree of SSD decreases with increasing body size. Hence, Sylvilagus provides a new example, unusual for a female‐biased taxon, in which allometry for SSD is consistent with ‘Rensch's Rule’. As a corollary to Rensch's Rule, we observe that changes in SSD in Sylvilagus are typically associated with larger, more significant changes in males than females. Female‐biased SSD could be produced by selection for larger females, smaller males, or both. Although larger female size may be related to high fecundity and the extremely rapid fetal and neonatal growth in Sylvilagus, we find little evidence for a correlation between SSD and various fecundity‐related traits in among‐species comparisons. Smaller male size may confer greater reproductive success through greater mobility and reduced energetic requirements. We propose that a suite of traits (female dispersion, large male home ranges, reduced aggression, and a promiscuous mating system) has favoured smaller males and thus influenced the evolution of SSD in cottontails. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 141–156.     

Growth and survivorship as proximate causes of sexual size dimorphism in peninsula dragon lizards Ctenophorus fionni

Males and females differ in body size in many animals, but the direction and extent of this sexual size dimorphism (SSD) varies widely. Males are larger than females in most lizards of the iguanian clade, which includes dragon lizards (Agamidae). I tested whether the male larger pattern of SSD in the peninsula dragon lizard, Ctenophorus fionni, is a result of sexual selection for large male size or relatively higher mortality among females. Data on growth and survivorship were collected from wild lizards during 1991–1994. The likelihood of differential predation between males and females was assessed by exposing pairs of male and female lizards to a predator in captivity, and by comparing the frequency of tail damage in wild‐caught males and females. Male and female C. fionni grew at the same rate, but males grew for longer than females and reached a larger asymptotic size (87 mm vs. 78 mm). Large males were under‐represented in the population because they suffered higher mortality than females. Predation may account for some of this male‐biased mortality. The male‐biased SSD in C. fionni resulted from differences in growth pattern between the sexes. The male‐biased SSD was not the result of proximate factors reducing female body size. Indeed SSD in this species remained male‐biased despite high mortality among large males. SSD in C. fionni is consistent with the ultimate explanation of sexual selection for large body size in males.     

Sexual size and shape evolution in European newts (Amphibia: Caudata: Salamandridae) on the Balkan Peninsula

We used a phylogenetic perspective in an examination of the direction and extent of sexual dimorphism in body size and body shape in European newts from the Balkan Peninsula (alpine newts, Mesotriton alpestris; crested newts, Triturus cristatus superspecies; smooth newts, Lissotriton vulgaris). We found a strong, female‐biased sexual size dimorphism (SSD) in the analysed clades of alpine newt, whereas within crested newts we found a less stringent female‐biased SSD in Triturus carnifex, Triturus macedonicus and Triturus karelinii, and no significant SSD in T. cristatus or Triturus dobrogicus. Among the smooth newts, we found male‐biased SSD in Lissotriton vulgaris vularis and Lissotriton vulgaris greacus and no SSD in Lissotriton vulgaris meridionalis. Most of these newts also exhibit a significant sexual dimorphism in body shape, which varied more randomly than body size, regardless of SSD level. Female and male body size as well as the degree of SSD displayed statistically significant phylogenetic signal, while sexual dimorphism in body shape was phylogenetically independent. The relationship between independent contrast data for female size and male size indicated that SSD in European newts could be driven by a disproportionate increase in female size as increase in female size was not accompanied by a proportional increase in male size.     

Field evidence challenges the often‐presumed relationship between early male maturation and female‐biased sexual size dimorphism

Female‐biased sexual size dimorphism (SSD) is often considered an epiphenomenon of selection for the increased mating opportunities provided by early male maturation (i.e., protandry). Empirical evidence of the adaptive significance of protandry remains nonetheless fairly scarce. We use field data collected throughout the reproductive season of an SSD crab spider, Mecaphesa celer, to test two hypotheses: Protandry provides fitness benefits to males, leading to female‐biased SSD, or protandry is an indirect consequence of selection for small male size/large female size. Using field‐collected data, we modeled the probability of mating success for females and males according to their timing of maturation. We found that males matured earlier than females and the proportion of virgin females decreased abruptly early in the season, but unexpectedly increased afterward. Timing of female maturation was not related to clutch size, but large females tended to have more offspring than small females. Timing of female and male maturation was inversely related to size at adulthood, as early‐maturing individuals were larger than late‐maturing ones, suggesting that both sexes exhibit some plasticity in their developmental trajectories. Such plasticity indicates that protandry could co‐occur with any degree and direction of SSD. Our calculation of the probability of mating success along the season shows multiple male maturation time points with similar predicted mating success. This suggests that males follow multiple strategies with equal success, trading‐off access to virgin females with intensity of male–male competition. Our results challenge classic hypotheses linking protandry and female‐biased SSD, and emphasize the importance of directly testing the often‐assumed relationships between co‐occurring animal traits.     

Absence of Mate Choice and Postcopulatory Benefits in a Species with Extreme Sexual Size Dimorphism

Most hypotheses related to the evolution of female‐biased extreme sexual size dimorphism (SSD) attribute the differences in the size of each sex to selection for reproduction, either through selection for increased female fecundity or selection for male increased mobility and faster development. Very few studies, however, have tested for direct fitness benefits associated with the latter – small male size. Mecaphesa celer is a crab spider with extreme SSD, whose males are less than half the size of females and often weigh 10 times less. Here, we test the hypotheses that larger size in females and smaller size in males are sexually selected through differential pre‐ and postcopulatory reproductive benefits. To do so, we tested the following predictions: matings between small males and large females are more likely to occur due to mate choice; females mated to small males are less likely to accept second copulation attempts; and matings between small males and large females will result in larger clutches of longer‐lived offspring. Following staged mating trials in the laboratory, we found no support for any of our predictions, suggesting that SSD in M. celer may not be driven by pre‐ or post‐reproductive fitness benefits to small males.     

Function‐related Drivers of Skull Morphometric Variation and Sexual Size Dimorphism in a Subterranean Rodent,Plateau Zokor (Eospalax baileyi)

Sexual dimorphism is prevalent in most living organisms. The difference in size between sexes of a given species is generally known as sexual size dimorphism (SSD). The magnitude of the SSD is determined by Rensch's rule where size dimorphism increases with increasing body size when the male is the larger sex and decreases with increasing average body size when the female is the larger sex. The unique underground environment that zokors (Eospalax baileyi) live under in the severe habitat of the Qinghai‐Tibetan Plateau (QTP) could create SSD selection pressures that may or may not be supported by Rensch's rule, making this scientific question worthy of investigation. In this study, we investigated the individual variation between sexes in body size and SSD of plateau zokors using measurements of 19 morphological traits. We also investigated the evolutionary mechanisms underlying SSD in plateau zokors. Moreover, we applied Rensch's rule to all extant zokor species. Our results showed male‐biased SSD in plateau zokors: The body‐ and head‐related measurements were greater in males than in females. Linear regression analysis between body length, body weight, and carcass weight showed significant relationships with some traits such as skull length, lower incisor length, and tympanic bulla width, which might support our prediction that males have faster growth rates than females. Further, the SSD pattern corroborated the assumption of Rensch's rule in plateau zokors but not in the other zokor species. Our findings suggest that the natural underground habitat and behavioral differences between sexes can generate selection pressures on male traits and contribute to the evolution of SSD in plateau zokors.     

Sexual size dimorphism in greater mouse‐eared bat Myotis myotis (Chiroptera: Vespertilionidae) from a Mediterranean region

Although sexual size dimorphism (SSD) is common among mammals, there is no clear explanation for its maintenance in nature. Bats are one of the few groups of mammals where reverse SSD appears. In this group, the size of individuals may have very important ecological consequences related with flight. In this study, we examine sexual dimorphism in the wing measurements of 195 adult individuals (141 males and 54 females) of the greater mouse‐eared bat Myotis myotis in the south‐east of the Iberian Peninsula. We also investigated size differences between paired and single males in a swarming roost. The results indicate that there are significant differences in the wing measurements between sexes, females being bigger than males in this respect. While no significant differences in the wing measurements of paired and single males were observed, significant differences were found in their relative weight and fitness, single males being significantly heavier and having a better physical condition. We discuss the implications of SSD for the female of M. myotis in terms of reproductive advantages, trophic niche segregation and a greater ability to move, which may favour gene flow between populations.     

Proximate causes of sexual size dimorphism in Colorado pikeminnow, a long‐lived cyprinid

Evidence for sexual size dimorphism (SSD) and its possible causes were examined in the endangered Colorado pikeminnow Ptychocheilus lucius, a large, piscivorous, cyprinid endemic to the Colorado River system of North America. Individuals representing 18–24% of the upper Colorado River population were captured, measured, sexed and released in 1999 and 2000. Differing male and female total length‐(L_T) frequency distributions revealed SSD with females having greater mean and maximum sizes than males. Although both sexes exhibit indeterminate post‐maturity growth, growth trajectories differed. The point of trajectory divergence was not established, but slowed male growth might coincide with the onset of maturation. Differing growth rate was the dominant proximate cause of SSD, accounting for an estimated 61% of the observed difference in mean adult L_T. The degree of SSD in adults, however, was also related to two other factors. Evidence suggests males become sexually active at a smaller size and earlier age than females; a 2 year difference, suggested here, accounted for an estimated 12% of the between‐sex difference in mean adult L_T. Temporal shifts in gender‐specific survival accounted for an additional 27% of the observed between‐sex difference in mean adult L_T. Estimated age distributions indicated a higher number of older females than older males and more younger males than younger females in the population during the period of sampling. Dissimilarity of age distributions was an unexpected result because the male : female population sex ratio was 1 : 1 and estimates of long‐term annual survival for adult males and females were equal (88%). Future assessments of SSD in this population are apt to vary depending on the prior history of short‐term gender‐specific survival. Without recognizing SSD, non‐gender‐specific growth curves overestimate mean age of adult females and underestimate mean age of adult males of given L_T. Assuming age 8 years for first reproduction in males and age 10 years for females, the adult male : female ratio was estimated as 1·1 : 1 and mean adult age, or generation time, was estimated as 16·4 years for males and 18·4 years for females.     

Lack of support for Rensch's rule in an intraspecific test using red flour beetle (Tribolium castaneum) populations

Rensch's rule proposes a universal allometric scaling phenomenon across species where sexual size dimorphism (SSD) has evolved: in taxa with male‐biased dimorphism, degree of SSD should increase with overall body size, and in taxa with female‐biased dimorphism, degree of SSD should decrease with increasing average body size. Rensch's rule appears to hold widely across taxa where SSD is male‐biased, but not consistently when SSD is female‐biased. Furthermore, studies addressing this question within species are rare, so it remains unclear whether this rule applies at the intraspecific level. We assess body size and SSD within Tribolium castaneum (Herbst), a species where females are larger than males, using 21 populations derived from separate locations across the world, and maintained in isolated laboratory culture for at least 20 years. Body size, and hence SSD patterns, are highly susceptible to variations in temperature, diet quality and other environmental factors. Crucially, here we nullify interference of such confounds as all populations were maintained under identical conditions (similar densities, standard diet and exposed to identical temperature, relative humidity and photoperiod). We measured thirty beetles of each sex for all populations, and found body size variation across populations, and (as expected) female‐biased SSD in all populations. We test whether Rensch's rule holds for our populations, but find isometry, i.e. no allometry for SSD. Our results thus show that Rensch's rule does not hold across populations within this species. Our intraspecific test matches previous interspecific studies showing that Rensch's rule fails in species with female‐biased SSD.     

THE EVOLUTION OF FEMALE-BIASED SEXUAL SIZE DIMORPHISM: A POPULATION-LEVEL COMPARATIVE STUDY IN HORNED LIZARDS (PHRYNOSOMA)

Female-biased sexual size dimorphism is uncommon among vertebrates and traditionally has been attributed to asymmetric selective pressures favoring large fecund females (the fecundity-advantage hypothesis) and/or small mobile males (the small-male advantage hypothesis). I use a phylogenetically based comparative method to address these hypotheses for the evolution and maintenance of sexual size dimorphism among populations of three closely related lizard species (Phrynosoma douglasi, P. ditmarsi, and P. hernandezi). With independent contrasts I estimate evolutionary correlations among female body size, male body size, and sexual size dimorphism (SSD) to determine whether males have become small, females have become large, or both sexes have diverged concurrently in body size during the evolutionary Xhistory of this group. Population differences in degree of SSD are inversely correlated with average male body size, but are not correlated with average female body size. Thus, variation in SSD among populations has occurred predominantly through changes in male size, suggesting that selective pressures on small males may affect degree of SSD in this group. I explore three possible evolutionary mechanisms by which the mean male body size in a population could evolve: changes in size at maturity, changes in the variance of male body sizes, and changes in skewness of male body size distributions. Comparative analyses indicate that population differentiation in male body size is achieved by changes in male size at maturity, without changes in the variance or skewness of male and female size distributions. This study demonstrates the potential of comparative methods at lower taxonomic levels (among populations and closely related species) for studying microevolutionary processes that underlie population differentiation.     

SEX‐SPECIFIC SELECTION AND INTRASPECIFIC VARIATION IN SEXUAL SIZE DIMORPHISM

Sexual size dimorphism (SSD) is thought to evolve due to sex differences in selection on body size, but it is largely unknown whether intraspecific variation in SSD reflects differences in sex‐specific selection among populations. We addressed this question by comparing viability selection between two island populations of the brown anole lizard (Anolis sagrei) that differ in the magnitude of male‐biased SSD. On both islands, females experienced stabilizing selection favoring intermediate size whereas males experienced directional selection favoring larger size. Thus, sex‐specific selection matched the overall pattern of male‐biased SSD, but population differences in the magnitude of SSD were not associated with local differences in selection. Rather, population differences in SSD appear to result from underlying differences in the environmental potential for a rapid growth, coupled with sex‐specific phenotypic plasticity. Males grew more slowly on the island with low SSD whereas growth of females did not differ between islands. Both sexes had substantially lower mass per unit length on the island with low SSD, suggesting that they were in a relatively poorer energetic condition. We propose that this energetic constraint disproportionately impacts growth of males due to their greater absolute energy requirements, thus driving intraspecific variation in SSD.     

External morphology of the short-beaked common dolphin, Delphinus delphis: growth, allometric relationships and sexual dimorphism

Growth, allometric relationships and sexual dimorphism are described from measurements of 105 male, 149 female and 38 unsexed specimens of short‐beaked common dolphin, Delphinus delphis, stranded along the Irish coastline (53.8% of the sample) or by‐caught in fisheries (46.2% of the sample), from 1990 to 2003. For each dolphin, 24 external body length measurements were recorded. Ages were determined for 183 dolphins by analysis of growth layer groups in the dentine. Males ranged in total body length (TBL) from 105 to 231 cm and females from 93 to 230 cm, with a maximum age of 25 years obtained for both sexes. Using a single Gompertz growth curve, asymptotic values obtained for TBL were 211.6 cm and 197.4 cm for males and females, respectively. Asymptotic lengths were attained at 11 years in males and 9 years in females. The gestation period was estimated to last approximately 11.5 months. Sexual size dimorphism (SSD) was evident, with males being significantly larger than females for 20 of the characters measured, and an SSD ratio of 1.06 was obtained. Sexual shape dimorphism was lacking, except for the presence of prominent postanal humps in mature males.     

TRAPPING AND RINGING OF EGYPTIAN GEESE AND AFRICAN SHELDUCK AT VOGELVLEI,CAPE

Many small plovers Charadrius spp. have sexually monomorphic plumage and cryptic sexual size dimorphism. The objective of our study was to assess the variation in body sizes between male and female plovers breeding in Madagascar. We collected blood samples and data on adult body sizes of four small plovers (Madagascar Plover Charadrius thoracicus, Kittlitz's Plover C. pecuarius, White-fronted Plover C. marginatus and Three-banded Plover C. tricollaris), and used molecular genetic markers to sex the adults. We found significant differences in body size among the four species, and between sexes. Furthermore, individuals from the southern ecoregion tended to be larger than in the western ecoregion. The Madagascar Plover's body size was significantly more dimorphic than the Kittlitz's and White-fronted Plovers. Breeding Malagasy plovers' show significant sexual size dimorphism (SSD): Madagascar Plover females were heavier and had longer wings than males, whereas the males had longer tarsi; in White-fronted Plover only wing length was different between the sexes. Taken together, our work reports SSD in small African plovers that exhibit monomorphic plumage, and we propose that SSD may be more common than currently acknowledged; we term this 'cryptic sexual size dimorphism'. Our results also suggest sexual selection and/or natural selection exert different pressures on body size in different Malagasy plover species.     

Sexual selection on male size drives the evolution of male‐biased sexual size dimorphism via the prolongation of male development

Sexual size dimorphism (SSD) arises when the net effects of natural and sexual selection on body size differ between the sexes. Quantitative SSD variation between taxa is common, but directional intraspecific SSD reversals are rare. We combined micro‐ and macroevolutionary approaches to study geographic SSD variation in closely related black scavenger flies. Common garden experiments revealed stark intra‐ and interspecific variation: Sepsis biflexuosa is monomorphic across the Holarctic, while S. cynipsea (only in Europe) consistently exhibits female‐biased SSD. Interestingly, S. neocynipsea displays contrasting SSD in Europe (females larger) and North America (males larger), a pattern opposite to the geographic reversal in SSD of S. punctum documented in a previous study. In accordance with the differential equilibrium model for the evolution of SSD, the intensity of sexual selection on male size varied between continents (weaker in Europe), whereas fecundity selection on female body size did not. Subsequent comparative analyses of 49 taxa documented at least six independent origins of male‐biased SSD in Sepsidae, which is likely caused by sexual selection on male size and mediated by bimaturism. Therefore, reversals in SSD and the associated changes in larval development might be much more common and rapid and less constrained than currently assumed.     

Does ‘gliding while gravid’ explain Rensch's rule in flying lizards?

Among species with sexual size dimorphism (SSD), taxa in which males are the larger sex have increasing SSD with increasing body size, whereas in taxa in which females are the larger sex, SSD decreases with body size: Rensch's rule. We show in flying lizards, a clade of mostly female‐larger species, that SSD increases with body size, a pattern similar to that in clades with male‐biased SSD or more evenly mixed SSD. The observed pattern in Draco appears due to SSD increasing with evolutionary changes in male body size; specifically divergence in body size among species that are in sympatric congeneric assemblages. We suggest that increasing body size, resulting in decreased gliding performance, reduces the relative gliding cost of gravidity in females, and switches sexual selection in males away from a small‐male, gliding advantage and toward selection on large size and fighting ability as seen in many other lizards. Thus, selection for large females is likely greater than selection for large males at the smaller end of the body size continuum, whereas this relationship reverses for species at the larger end of the continuum. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 270–282.