Possible links between phenotypic variability,habitats and connectivity in the killifish Aphaniops stoliczkanus in Northeast Oman

There is a significant gap in our knowledge of the intraspecific morphological variability in freshwater fish, although such data are crucial for understanding species diversity. Here we use the killifish Aphaniops stoliczkanus (Day, 1872; Aphaniidae: Cyprinodontiformes), which is a widespread but poorly known freshwater species in the Middle East, to investigate variability in morphological traits within and between its populations. As otolith morphology is known to evolve on ecological timescales and can signal the presence of cryptic lineages, a special focus lies on otolith variability. Based on samples from six populations in northern Oman, we found that variation in pigmentation, disparities in body shape and otolith variability can be associated with distinctive environmental conditions. The unique otolith shape of A. stoliczkanus from a hot sulphuric spring (Nakhal) suggests that a cryptic lineage may have emerged there. Our new data can serve as a benchmark for future studies on the diversity of Aphaniops and other Aphaniidae and help to clarify whether cryptic diversity is present in some lineages. Moreover, our data can serve as an actualistic model for studies on fossil fishes, in which morphological characters provide the only accessible data source for taxonomic and phylogenetic interpretations.     

CONVERGENT EVOLUTION OF SEXUAL DIMORPHISM IN SKULL SHAPE USING DISTINCT DEVELOPMENTAL STRATEGIES

Studies integrating evolutionary and developmental analyses of morphological variation are of growing interest to biologists as they promise to shed fresh light on the mechanisms of morphological diversification. Sexually dimorphic traits tend to be incredibly divergent across taxa. Such diversification must arise through evolutionary modifications to sex differences during development. Nevertheless, few studies of dimorphism have attempted to synthesize evolutionary and developmental perspectives. Using geometric morphometric analysis of head shape for 50 Anolis species, we show that two clades have converged on extreme levels of sexual dimorphism through similar, male‐specific changes in facial morphology. In both clades, males have evolved highly elongate faces whereas females retain faces of more moderate proportion. This convergence is accomplished using distinct developmental mechanisms; one clade evolved extreme dimorphism through the exaggeration of a widely shared, potentially ancestral, developmental strategy whereas the other clade evolved a novel developmental strategy not observed elsewhere in the genus. Together, our analyses indicate that both shared and derived features of development contribute to macroevolutionary patterns of morphological diversity among Anolis lizards.     

Sexual differences in locomotor performance in Tropidurus catalanensis lizards (Squamata: Tropiduridae) – body shape,size and limb musculature explain variation between males and females

Sexual dimorphism (SD) is the evolutionary outcome of selection acting differently on males and females. Several studies describe sexual differences in body size, although other morphological traits might be allometric between sexes and imply functional consequences. Here we test whether morphological differences between sexes in size and shape in the lizard Tropidurus catalanensis explain variation in performance of four locomotor traits. Our results show that males are larger than females and also exhibit longer limbs, longer muscles and larger muscle cross‐sectional areas, while females have longer trunks and more sharped anterior claws; males outperform females in all locomotor performances measured. Sexual differences in sprinting and climbing is related with body size, and climbing performance is also explained by limb lengths, by differences in lengths and cross‐sectional areas of specific muscles, and by interlimb distances. Between‐sex differences in exertion are also related to SD, despite associations with sharper posterior claws that are independent of sex. Grasping performance, however, is associated with some muscle and morphological parameters that are not sexually dimorphic. Together our results suggest that morphology might be under sexual selection in T. catalanensis, given that better locomotor performance likely favours male lizards in typical activities of this polygenic species, such as territory defence and female acquisition. Moreover, the longer trunks that characterize females may confer more space to accommodate eggs. On the other hand, territory defence by males probably increases their exposure to predators, resulting in a synergistic effect of sexual and natural selection in the evolution of SD in T. catalanensis.     

Sex-linked variations in the sagittal otolith biometry of Nemipterus randalli (Russell, 1986) from the eastern Mediterranean Sea

Few studies have been conducted on the sagittal otolith shape and morphometry of Nemipterus randalli, and none of these studies has examined the effect of sexual dimorphism on the otolith morphology of this species, therefore this study aimed to contribute to knowledge about the otolith morphology of N. randalli, an invasive fish species for the Mediterranean Sea. For this purpose, a total of 132 samples (51 female and 81 male) were obtained from İskenderun Bay with the help of commercial fishermen in November 2018. Relationships between otolith measurements and fish size were determined. Shape indices and elliptic Fourier coefficients were calculated. Significant differences were detected between males and females in all analysis. The sexes were separated from each other using both shape indices and elliptic Fourier coefficients. However, shape analysis was more effective in distinguishing sexes than traditional morphometric analysis. Asymmetry in the otolith morphology of sexes has been attributed to differences in the growth and sexual maturity of male and female fish. The results of this study indicated that sexual dimorphism in Nemipterus randalli was also reflected in the otolith morphology.     

Ontogenetic pattern,morphological sexual and side dimorphism in the saccular otolith of a scaleless killifish Aphanius furcatus (Teleostei: Aphaniidae)

The relationships between ontogenetic and sexual factors with otolith morphology are poorly studied in killifish genus Aphanius Nardo, 1827. We studied the ontogeny, morphological sexual and side dimorphism in the saccular otolith of a scaleless killifish Aphanius furcatus inhabiting a high‐salinity environment in Southern Iran. The results highlighted growth‐dependent variability in the otolith of A. furcatus, which might be useful to define its certain developmental stages. We underlined those characters that were consistent during fish development, which could be valuable to identify the species. Considering the ontogenetic pattern of otolith in A. furcatus, it was found that the otoliths of young individuals have largely been influenced by sexes and that it was reduced during the fish development. Morphological side dimorphism in the otolith of A. furcatus tends to be increased during fish development. The outcomes of this study are essential since they have provided new information on ontogenetic changes and the sexual and side dimorphism of Aphanius otolith.     

SEX‐SPECIFIC PATTERNS OF MORPHOLOGICAL DIVERSIFICATION: EVOLUTION OF REACTION NORMS AND STATIC ALLOMETRIES IN NERIID FLIES

The consequences of sex‐specific selection for patterns of diversification remain poorly known. Because male secondary sexual traits are typically costly to express, and both costs and benefits are likely to depend on ambient environment and individual condition, such traits may be expected to diversify via changes in reaction norms as well as the scaling of trait size with body size (static allometry). We investigated morphological diversification within two species of Australian neriid flies (Telostylinus angusticollis, Telostylinus lineolatus) by rearing larvae from several populations on larval diets varying sixfold in nutrient concentration. Mean body size varied among populations of T. angusticollis, but body size reaction norms did not vary within either species. However, we detected diversification of reaction norms for body shape in males and females within both species. Moreover, unlike females, males also diversified in static allometry slope and reaction norms for static allometry slope of sexual and nonsexual traits. Our findings reveal qualitative sex differences in patterns of morphological diversification, whereby shape–size relationships diversify extensively in males, but remain conserved in females despite extensive evolution of trait means. Our results highlight the importance of incorporating plasticity and allometry in studies of adaptation and diversification.     

Sexual dimorphism in traits related to locomotion: ontogenetic patterns of variation in Podarcis wall lizards

Sexual dimorphism in body size and shape in animals is normally linked to sexual selection mechanisms that modify the morphological properties of each sex. However, sexual dimorphism of ecologically relevant traits may be amplified by natural selection and result in the ecological segregation of both sexes. In the present study, we investigated patterns of sexual dimorphism of morphological traits relevant for locomotion in two lacertid lizards, Podarcis bocagei and Podarcis carbonelli, aiming to identify ontogenetic sources of variation. We analysed trunk and limb variation in relation to total body size, as well as the covariation of different traits, aiming to shed light on the proximate causation of adult sexual dimorphism. We find that, although immatures are generally monomorphic, adult females have a longer trunk, and adult males have longer fore and hind limbs. Both sexes differ substantially with respect to their growth trajectories and relationships between traits, whereas, in some cases, there are signs of morphological constraints delimiting the observed patterns. Because of the direct connection between limb size/shape and locomotor performance, which is relevant both for habitat use and escape from predators, the observed patterns of sexual dimorphism are expected to translate into ecological differences between both sexes. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 530–543.     

Allometric variation and sexual dimorphism in <Emphasis Type="Italic">Lycodes yamatoi</Emphasis> Toyoshima, 1985 (Perciformes: Zoarcidae) from the Sea of Japan

A detailed analysis of allometric variation and sexual dimorphism in Lycodes yamatoi (Perciformes: Zoarcidae) was carried out using an original method developed by the authors. An allometric pattern of growth was revealed not only in plastic but in some meristic characters, such as the number of teeth on the bones of the mouthparts and the number of gill rakers on the first gill arc. It was determined that most of the morphological characters of L. yamatoi were subjected to positive allometric variability (the proportions of the head and the anterior part of the body), while growth of the remaining characters was specified by negative allometry (the length of the rays of pair and caudal fins, interorbital distance, and isthmus width) and isometry (the standard length, ante-anal distance, and head width on the vertical neck). The juveniles of L. yamatoi have no significant sexual distinctions either in plastic or in meristic characters. Adult females and males of L. yamatoi significantly differed by characters composing the head complex, the actual length of the head, and correlatively related with it ante-dorsal and ante-pectoral distances. The dimensional indexes of antedorsal and ante-pectoral distances, and the relative sizes of maxillary bones (lmx/c) are of the greatest diagnostic value for sex differentiation.     

Insights into the genetic architecture of sexual dimorphism from an interspecific cross between two diverging Silene (Caryophyllaceae) species

The evolution of sexual dimorphism in species with separate sexes is influenced by the resolution of sexual conflicts creating sex differences through genetic linkage or sex‐biased expression. Plants with different degrees of sexual dimorphism are thus ideal to study the genetic basis of sexual dimorphism. In this study we explore the genetic architecture of sexual dimorphism between Silene latifolia and Silene dioica. These species have chromosomal sex determination and differ in the extent of sexual dimorphism. To test whether QTL for sexually dimorphic traits have accumulated on the sex chromosomes and to quantify their contribution to species differences, we create a linkage map and performed QTL analysis of life history, flower and vegetative traits using an unidirectional interspecific F2 hybrid cross. We found support for an accumulation of QTL on the sex chromosomes and that sex differences explained a large proportion of the variance between species, suggesting that both natural and sexual selection contributed to species divergence. Sexually dimorphic traits that also differed between species displayed transgressive segregation. We observed a reversal in sexual dimorphism in the F2 population, where males tended to be larger than females, indicating that sexual dimorphism is constrained within populations but not in recombinant hybrids. This study contributes to the understanding of the genetic basis of sexual dimorphism and its evolution in Silene.     

Sexual size dimorphism in Ophisops elegans (Squamata: Lacertidae) in Iran

Twenty-three morphological features of 140 specimens of Ophisops elegans were analysed in order to identify sexual dimorphism in west and northwestern populations of Iran. Sexual dimorphism is significant (P<0.05) in nearly all metric features except for trunk length (TL) and length of widest part of belly (LWB), and in only two meristic characters, the number of dorsal scales around mid-body (DSN) and the number of femoral pores (FPN). Males have a relatively longer snout-vent length (SVL) than females and males have generally relatively larger heads compared to females.     

Sex‐linked inheritance,genetic correlations and sexual dimorphism in three melanin‐based colour traits in the barn owl

Theory states that genes on the sex chromosomes have stronger effects on sexual dimorphism than genes on the autosomes. Although empirical data are not necessarily consistent with this theory, this situation may prevail because the relative role of sex‐linked and autosomally inherited genes on sexual dimorphism has rarely been evaluated. We estimated the quantitative genetics of three sexually dimorphic melanin‐based traits in the barn owl (Tyto alba), in which females are on average darker reddish pheomelanic and display more and larger black eumelanic feather spots than males. The plumage traits with higher sex‐linked inheritance showed lower heritability and genetic correlations, but contrary to prediction, these traits showed less pronounced sexual dimorphism. Strong offspring sexual dimorphism primarily resulted from daughters not expressing malelike melanin‐based traits and from sons expressing femalelike traits to similar degrees as their sisters. We conclude that in the barn owl, polymorphism at autosomal genes rather than at sex‐linked genes generate variation in sexual dimorphism in melanin‐based traits.     

Reproduction and sexual dimorphism in two populations of the polymorphic spiny lizard Sceloporus minor from Hidalgo,México

Male‐biased sexual dimorphism is extensive in New World spiny lizards (Sceloporus: Phrynosomatidae) and is particularly prominent in the polymorphic minor lizard Sceloporus minor. However, the possible relationship between patterns of sexual dimorphism and gonadal development is little known for most species. In this study, we explored aspects of sexual dimorphism in noncolor morphological traits in S. minor and characterized the gonadal cycle in males and females from each of two sites (El Enzuelado and La Manzana) in Hidalgo, México, differing in key ecological aspects linked to life‐history trait expression in other lizards. Males were generally larger than females in each population and expressed larger forms of several other morphological traits, although not all comparisons attained statistical significance. Both sexes attained reproductive maturity at a larger size in El Enzuelado relative to La Manzana, and females from El Enzuelado had larger litters. Gonadal cycles differed substantially between the two populations and suggest that reproductive activity of males and females was synchronous at El Enzuelado and asynchronous at La Manzana, an unusual pattern. Geographic variation in sex‐specific responses to environmental variables may be at least partly responsible for the exceptional species diversity of spiny lizards in México.     

SEX CHROMOSOME LINKED GENETIC VARIANCE AND THE EVOLUTION OF SEXUAL DIMORPHISM OF QUANTITATIVE TRAITS

Theory predicts that sex chromsome linkage should reduce intersexual genetic correlations thereby allowing the evolution of sexual dimorphism. Empirical evidence for sex linkage has come largely from crosses and few studies have examined how sexual dimorphism and sex linkage are related within outbred populations. Here, we use data on an array of different traits measured on over 10,000 individuals from two pedigreed populations of birds (collared flycatcher and zebra finch) to estimate the amount of sex‐linked genetic variance (h²_z). Of 17 traits examined, eight showed a nonzero h²_Z estimate but only four were significantly different from zero (wing patch size and tarsus length in collared flycatchers, wing length and beak color in zebra finches). We further tested how sexual dimorphism and the mode of selection operating on the trait relate to the proportion of sex‐linked genetic variance. Sexually selected traits did not show higher h²_Z than morphological traits and there was only a weak positive relationship between h²_Z and sexual dimorphism. However, given the relative scarcity of empirical studies, it is premature to make conclusions about the role of sex chromosome linkage in the evolution of sexual dimorphism.     

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.     

大山雀身体大小的性二态及其季节变化

动物中普遍存在雌雄个体身体大小的性二态现象。了解近缘种之间身体大小性二态现象的差异，可为深入探讨身体大小性二态现象的潜在驱动机制提供证据。国外对欧亚大山雀（Parus major）的研究发现，其喙长、跗跖长、翅长等 6 项身体大小指标存在着明显的性二态，且喙长的性二态存在季节间差异。大山雀（P. cinereus）曾被作为欧亚大山雀的一个亚种，其形态和行为与欧亚大山雀存在着诸多相似之处。为探讨大山雀是否也存在身体大小性二态及季节性差异，本研究分析了 2018 至 2020 年间在河南董寨国家级自然保护区捕捉的 226 只（雌性 96 只和雄性 130 只）大山雀的喙长、头喙长、跗跖长、翅长、尾长和体长这 6 项体征指标的两性差异及其季节变化。结果显示，大山雀上述 6 项身体大小指标均存在不同程度的性二态现象，且雄性个体仅喙长与雌性的差异不显著，其余 5 项指标均显著大于雌性。此外，身体大小指标的两性差异不随季节显著变化，但两性的跗跖长在秋季均显著短于冬季和繁殖季，尾长在繁殖季均显著长于秋季和冬季。上述结果表明，大山雀身体大小的性二态及其季节性差异与欧亚大山雀并不完全相似。无论其身体大小存在性二态和季节变化的原因，还是其与欧亚大山雀在身体大小性二态模式上的差别，都有待今后进一步的研究。     

SEXUAL DIMORPHISM IN SPOTTED DOLPHINS (STENELLA ATTENUATA) IN THE EASTERN TROPICAL PACIFIC OCEAN

We analyzed 4 meristic and 32 morphometric cranial traits of 612 adult specimens of spotted dolphins ( Stenella attenuata ) from the eastern tropical Pacific Ocean for sexual dimorphism. Eleven 5" latitude/longitude blocks with five or more of each sex were assessed with a two-way ANOVA for sex and geographic differences. Interaction between these two factors was found for six measurements, suggesting that the degree of sexual dimorphism varies geographically for a few characters (although in no discernible geographic pattern). Sexual dimorphism was demonstrated for 23 of the 36 characters, with differences ranging from 0.00 to 5.88 percent. Females characteristically had a longer rostrum, while males generally had larger skulls overall. In terms of number of characters, the extent of sexual dimorphism demonstrated for skulls of spotted dolphins goes considerably beyond that shown for any other small delphine. A discriminant function involving a combination of 10 characters enabled us to identify correctly the sex of more than 75 percent of the specimens. A procedure for correcting specimen measurements is outlined that would enable an investigator to combine male and female specimens in geographic variation studies. A term (zwitter) is proposed for use when referring to specimens where measurements have been corrected to take into account differences between the sexes.     

Sexual dimorphism in a trophically polymorphic cichlid fish?

Sexual dimorphism in ecologically relevant traits is ubiquitous in animals. However, other types of intraspecific phenotypic divergence, such as trophic polymorphism, are less common. Because linkage to sex should often lead to balancing selection, understanding the association between sex and phenotypic divergence could help explain why particular species show high morphological variability. To determine if sexual dimorphism could be helping to maintain ecomorphological variation in a classic case of intraspecific trophic polymorphism, we examined the association between sex and morphological divergence in the cichlid Herichthys minckleyi. Although H. minckleyi with enlarged molariform teeth on their pharyngeal jaws have been reported to more commonly be male, we did not find an association between sex and pharyngeal morphotype. Sex was associated with divergence in body size (as measured through standard length). But, sex was not associated with any of the other trophic traits examined. However, pharyngeal morphotype did show an association with gut length, gape, and tooth number. Sexual dimorphism is not playing a central role in enhancing trophic diversity within H. minckleyi. J. Morphol. 276:1448–1454, 2015. © 2015 Wiley Periodicals, Inc.     

Ontogenetic stage‐specific quantitative trait loci contribute to divergence in developmental trajectories of sexually dimorphic fins between medaka populations

Sexual dimorphism can evolve when males and females differ in phenotypic optima. Genetic constraints can, however, limit the evolution of sexual dimorphism. One possible constraint is derived from alleles expressed in both sexes. Because males and females share most of their genome, shared alleles with different fitness effects between sexes are faced with intralocus sexual conflict. Another potential constraint is derived from genetic correlations between developmental stages. Sexually dimorphic traits are often favoured at adult stages, but selected against as juvenile, so developmental decoupling of traits between ontogenetic stages may be necessary for the evolution of sexual dimorphism in adults. Resolving intralocus conflicts between sexes and ages is therefore a key to the evolution of age‐specific expression of sexual dimorphism. We investigated the genetic architecture of divergence in the ontogeny of sexual dimorphism between two populations of the Japanese medaka (Oryzias latipes) that differ in the magnitude of dimorphism in anal and dorsal fin length. Quantitative trait loci (QTL) mapping revealed that few QTL had consistent effects throughout ontogenetic stages and the majority of QTL change the sizes and directions of effects on fin growth rates during ontogeny. We also found that most QTL were sex‐specific, suggesting that intralocus sexual conflict is almost resolved. Our results indicate that sex‐ and age‐specific QTL enable the populations to achieve optimal developmental trajectories of sexually dimorphic traits in response to complex natural and sexual selection.     

Cross‐sex genetic correlations and the evolution of sex‐specific local adaptation: Insights from classical trait clines in Drosophila melanogaster

Natural selection varies widely among locations of a species’ range, favoring population divergence and adaptation to local environmental conditions. Selection also differs between females and males, favoring the evolution of sexual dimorphism. Both forms of within‐species evolutionary diversification are widely studied, though largely in isolation, and it remains unclear whether environmental variability typically generates similar or distinct patterns of selection on each sex. Studies of sex‐specific local adaptation are also challenging because they must account for genetic correlations between female and male traits, which may lead to correlated patterns of trait divergence between sexes, whether or not local selection patterns are aligned or differ between the sexes. We quantified sex‐specific divergence in five clinally variable traits in Drosophila melanogaster that individually vary in their magnitude of cross‐sex genetic correlation (i.e., from moderate to strongly positive). In all five traits, we observed parallel male and female clines, regardless of the magnitude of their genetic correlation. These patterns imply that parallel spatial divergence of female and male traits is a reflection of sexually concordant directional selection imposed by local environmental conditions. In such contexts, genetic correlations between the sexes promote, rather than constrain, local adaptation to a spatially variable environment.     

When do meadow vipers (Vipera ursinii) become sexually dimorphic? – ontogenetic patterns of sexual size dimorphisms

Contrary to an increasing number of papers that document sexual dimorphism in size (and/or shape) in adults, studies dealing with sex differences in newborn and juvenile snakes are surprisingly scarce. Data about ontogenetic shifts in sexual dimorphism are generally lacking and hence, it is unclear whether sex differences are set at birth or arise post‐natally. In this study, we analyzed patterns of sexual dimorphism in body size, head dimensions and tail length (TL) among newborn, subadult and adult meadow vipers (Vipera ursinii) from the Bjelasica Mt. in Montenegro. Patterns of sexual size dimorphisms differed among traits. There was no significant difference in head dimension of males and females, but adult snakes were sexually dimorphic in body size. Sexual differences in TL were evident since birth but changed in degree throughout ontogeny. Neonate meadow vipers presented highly significant inter‐litter variation in the sexual dimorphism of all traits we have measured. Such family effects may have an important influence on extent of inter‐sexual differences in snakes and should be included in analyses of sexual dimorphism.