Sexually selected traits are larger and more variable in male than female beach-spawning capelin,Mallotus villosus

We evaluated whether morphological traits in capelin, Mallotus villosus, that appear to be sexually selected (pectoral fin, pelvic fin, anal fin, lateral ridge) were larger and more variable in males than females compared with naturally selected morphological traits (eyes, dorsal fin). Photographs were obtained of 136 capelin captured at two spawning sites and standardised measurements were taken of six morphological traits. Males had larger traits than females for a given body size and this was most pronounced in the traits thought to be sexually selected. Body size explained much of the variation in female traits but less variation in male traits, suggesting alternative selection pressures are involved. We suggest that larger male body size aids in endurance rivalry and sexually dimorphic traits help males to remain in physical contact with females while spawning on the beach.     

A new species of Astyanax Baird & Girard (Characiformes: Characidae) from the Rio Paraguaçu basin,Chapada Diamantina,Bahia, Brazil,with comments on bony hooks on all fins

A new species of Astyanax from the Rio Paraguaçu basin, Bahia, Brazil, is described. This new species is distinguished from its congeners known to occur in Brazilian drainages by the presence of small bony hooks on all fins of mature males. Furthermore, the new species can be diagnosed by its colour pattern, which consists of a unique vertically elongated humeral blotch, a conspicuous dark drop‐shaped horizontal blotch over the caudal peduncle, tapering anteriorly and not extending to the median caudal‐fin rays, and by the absence of a conspicuous broad dark midlateral stripe. The new species differs further by having the greatest body depth just anterior to the dorsal‐fin origin, 32–35 perforated scales in the lateral line and a reduced number of branched anal‐fin rays (16–20). Apparently, the new species does not fit into any species complex of Astyanax. The occurrence of bony hooks on all fins of Astyanax is discussed.     

Female Preference for Male Lateral Projection Area in the Shortfin Molly, Poecilia mexicana: Evidence for a Pre-existing Bias in Sexual Selection

Sailfin molly (Poecilia latipinna) males possess a large dorsal fin (sailfin) and perform an elaborate courtship display. Females prefer to associate and mate with males of greater body and sailfin size. Evidence supports a single origin for the sailfin species complex from a shortfin ancestor. Unlike sailfin species, males of the shortfin species complex are sexually monomorphic in fin size and exhibit little or no courtship behavior. In this study, we tested the pre‐existing bias and lateral projection area (LPA) hypotheses for sexual selection by examining female mating preferences in the shortfin molly, P. mexicana. Specifically, we presented females with pairs of dummy males differing in: (1) dorsal fin and body size together (holding fin:body size ratio constant); (2) body size (holding dorsal fin size constant); (3) dorsal fin size (holding body size constant); and dorsal fin:body size ratio (holding total LPA constant). Females spent more time near dummies of greater body and dorsal fin size. The preference functions based on the first three sets of stimuli showed a similar pattern: the greater the LPA difference between paired dummies, the stronger the preference for the larger of the two. However, in the fourth experiment, neither fin size, body size, nor any particular dorsal fin + body size combination was preferred. These findings support the LPA hypothesis suggesting that increased LPA is more stimulating to sexually receptive females and that females consequently prefer larger males. Moreover, these data are consistent with results obtained in an identical series of experiments conducted on P. latipinna. The preference for increased male dorsal fin size/LPA by both female P. latipinna and P. mexicana supports the pre‐existing bias hypothesis. Thus, a bias for increased male LPA and consequent selection for enlarged dorsal fins may have preceded the appearance of the sailfin trait within the Molliensia lineage.     

Temperature-dependent ovariole and testis maturation in the yellow dung fly

Temperature is one of the abiotic environmental factors most strongly affecting animal behaviour, physiology, and life history. In insects, lower temperatures generally slow down most physiological processes, reducing growth rate and prolonging the juvenile period. Here, we investigate temperature‐dependent ovariole and testis maturation in the anautogenous yellow dung fly, Scathophaga stercoraria L. (Diptera: Scathophagidae), and relate it to corresponding temperature effects on pre‐adult development time and the adult pre‐reproductive period. Flies were reared in the laboratory at three constant temperatures (18, 22, and 26 °C), and the size of the developing ovarioles and testes (reflecting sperm production) was measured over time (i.e., age). Ovariole size increased asymptotically over the first 12 days of adult life, while the testes continued to fill after day 10. In accordance with the temperature‐size rule, warmer temperatures resulted in smaller ovarioles (eggs) and smaller testes, independent of body size. Warmer temperatures also greatly reduced pre‐adult development time by more than half, from 12 to 25 °C, the larger males always taking 1–3 days longer than the females. Corresponding temperature effects on the adult pre‐reproductive period were small (<1 day between 15 and 25 °C), with males taking 5–6 days and females 10–13 days to first reproduction. Time lost by males during the pre‐adult stage, when ovaries and testes are produced, can thus be more than compensated‐for by time gained during the pre‐reproductive period, when eggs and sperm are produced, so males can nevertheless start reproducing sooner than females.     

Sexual selection versus alternative causes of sexual dimorphism in teiid lizards

Summary The presence and extent of sexual dimorphisms in body form (size and shape) of adult macroteiid lizards were investigated. Males were significantly larger than females in the temperate species, Cnemidophorus tigris, and in the tropical species, Ameiva ameiva and C. ocellifer. Young adult C. tigris males grew faster than young adult females within and between reproductive seasons. Adult males of all species had larger heads than adult females of the same body size; this difference increased with body size. Moreover, male C. tigris were heavier than females of the same snout-vent length. The causes and consequences of the sexual dimorphisms were also examined. The possible causes of body size are especially numerous, and distinguishing the relative influences of the various causal selection factors on body size is problematical. Nevertheless, observational field data were used to tentatively conclude that intrasexual selection was the cause of larger body size of C. tigris males relative to females because (1) larger males won in male aggressive interactions, (2) the winning males gained access to more females by repelling competitors and by female acceptance, (3) larger males consequently had higher reproductive success, and (4) other hypothetical causes of larger male size were unsupported.     

Sex-specific niche partitioning and sexual size dimorphism in Australian pythons (Morelia spilota imbricata)

Sexual dimorphism is usually interpreted in terms of reproductive adaptations, but the degree of sex divergence also may be affected by sex-based niche partitioning. In gape-limited animals like snakes, the degree of sexual dimorphism in body size (SSD) or relative head size can determine the size spectrum of ingestible prey for each sex. Our studies of one mainland and four insular Western Australian populations of carpet pythons ( Morelia spilota ) reveal remarkable geographical variation in SSD, associated with differences in prey resources available to the snakes. In all five populations, females grew larger than males and had larger heads relative to body length. However, the populations differed in mean body sizes and relative head sizes, as well as in the degree of sexual dimorphism in these traits. Adult males and females also diverged strongly in dietary composition: males consumed small prey (lizards, mice and small birds), while females took larger mammals such as possums and wallabies. Geographic differences in the availability of large mammalian prey were linked to differences in mean adult body sizes of females (the larger sex) and thus contributed to sex-based resource partitioning. For example, in one population adult male snakes ate mice and adult females ate wallabies; in another, birds and lizards were important prey types for both sexes. Thus, the high degree of geographical variation among python populations in sexually dimorphic aspects of body size and shape plausibly results from geographical variation in prey availability.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 113–125.     

Repeated matings and sperm depletion in the freshwater crayfish Austropotamobius italicus

1. In sexually reproducing organisms, the energetic costs of spermatogenesis can be considerable, and can limit the reproductive potential of the males. In species where males mate more than once during the reproductive season, the costs of sperm production are generally predicted to result in a decrease of ejaculate size and quality with successive fertilizations. 2. In this study we examined the variation in ejaculate size among successive fertilizations in a long‐lived freshwater crayfish species, Austropotamobius italicus. 3. Sexually active adult males of various sizes were allowed to mate repeatedly with different females on consecutive days. Trials for a given male ended when he copulated but did not release any sperm or refused to mate. 4. Males fertilized between 0 and 4 females, and most (42.5%) fertilized a single female. The overall number of females fertilized by a given male decreased with increasing male body size. Ejaculate size decreased markedly with consecutive fertilizations in a similar fashion among both large and small males, while simultaneously increasing with female body size. The total ejaculate size over successive fertilizations decreased with increasing male size. 5. Our study indicates that either sperm production or release involves non‐trivial costs in freshwater crayfish, and suggests that large/old males may face greater difficulties in gamete release than small/young ones, as shown by the lower number of females fertilized by large compared with small males, which may reflect the ongoing senescence of their reproductive performance.     

Sexual selection on female ornaments in the sex‐role‐reversed Gulf pipefish (Syngnathus scovelli)

Understanding how selection acts on traits individually and in combination is an important step in deciphering the mechanisms driving evolutionary change, but for most species, and especially those in which sexual selection acts more strongly on females than on males, we have no estimates of selection coefficients pertaining to the multivariate sexually selected phenotype. Here, we use a laboratory‐based mesocosm experiment to quantify pre‐ and post‐mating selection on female secondary sexual traits in the Gulf pipefish (Syngnathus scovelli), a sexually dimorphic, sex‐role‐reversed species in which ornamented females compete for access to choosy males. We calculate selection differentials and gradients on female traits, including ornament area, ornament number and body size for three episodes of selection related to female reproductive success (number of mates, number of eggs transferred and number of surviving embryos). Selection is strong on both ornament area and ornament size, and the majority of selection occurs during the premating episode of selection. Interestingly, selection on female body size, which has been detected in previous studies of Gulf pipefish, appears to be indirect, as evidenced by a multivariate analysis of selection gradients. Our results show that sexual selection favours either many bands or larger bands in female Gulf pipefish.     

Sexual dimorphism in four species of rockfish genusSebastes (Scorpaenidae)

Synopsis Sexual dimorphisms, and factors influencing the evolution of these differences, have been investigated for four species of rockfish:Sebastes melanops, S. flavidus, S. mystinus, andS. serranoides. These four species, which have similar ecology, tend to aggregate by species with males and females staying together throughout the year. In all four species adult females reach larger sizes than males, which probably relates to their role in reproduction. The number of eggs produced increases with size, so that natural selection has favored larger females. It appears males were subjected to different selective pressures than females. It was more advantageous for males to mature quickly, to become reproductive, than to expend energy on growth. Other sexually dimorphic features include larger eyes in males of all four species and longer pectoral fin rays in males of the three piscivorous species:S. melanops, S. flavidus, andS. serranoides. The larger pectoral fins may permit smaller males to coexist with females by increasing acceleration and, together with the proportionately larger eye, enable the male to compete successfully with the female to capture elusive prey (the latter not necessarily useful for the planktivoreS. mystinus). Since the size of the eye is equivalent in both sexes of the same age, visual perception should be comparable for both sexes.     

Host stage selection of the mealybug parasitoid Anagyrus spec. nov near sinope

The mealybug parasitoid Anagyrus spec. nov near sinope (Hymenoptera: Encyrtidae) is an undescribed parasitoid of the Madeira mealybug, Phenacoccus madeirensis Green (Homoptera: Pseudococcidae). We investigated the preference of Anagyrus spec. nov near sinope for six developmental stadia (first‐ and second‐instar nymphs, third‐instar immature females, third‐ or fourth‐instar immature males, pre‐reproductive adult females, and ovipositing adult females) of P. madeirensis and the fitness consequences of the host stage selection behavior. In the no‐choice test, Anagyrus spec. nov near sinope parasitized and completed development in all host stadia except third‐instar immature males. When all host stadia were offered simultaneously, the parasitoids preferred third‐instar immature and pre‐reproductive adult females. Dissection of the stung mealybugs revealed that the clutch size (number of eggs per host) was approximately four and three in the third‐instar and pre‐reproductive females, respectively, and one egg per first‐instar nymph. Parasitoids emerged from P. madeirensis parasitized at third‐instar or pre‐reproductive adult female completed development in the shortest duration, achieved a higher progeny survival rate, larger brood and body size, and the lowest proportion of males. We showed that the continued development of mealybugs had significant influence on the fitness of the parasitoids. Although deposited as eggs in first‐ or second‐instar nymphs, parasitoids emerged from mummies that had attained third‐instar or adult development achieved similar progeny survival rate, brood size, body size, and sex ratio as those parasitoids deposited and developed in third‐instar or adult mealybugs. By delaying larval development in young mealybugs, Anagyrus spec. nov near sinope achieved higher fitness by allowing the parasitized mealybugs to grow and accumulate body size and resources. We suggest that the fitness consequence of host stage selection of a koinobiont parasitoid should be evaluated on both the time of parasitism and the time of mummification.     

棒花鱼形态特征的两性异形和雌性个体生育力

测定了棒花鱼(Abbottina rivularis)繁殖期形态特征包括体长、头长、头宽、头高、眼间距、鼻间距、背鳍基长、胸鳍长、胸鳍腹鳍间距、尾柄长、尾鳍长和体重的两性异形和雌性个体生育力。结果表明,雄性个体的数量显著多于雌性个体,雄性个体的体长显著大于雌性个体。特定体长的雌性个体的胸鳍腹鳍间距显著大于雄性个体,头长、头宽、头高、眼间距、鼻间距、背鳍基长、胸鳍长、尾柄长和尾鳍长显著小于雄性个体,雌雄两性体重不存在显著差异。棒花鱼的怀卵数量与体长和体重回归关系显著。偏相关分析显示,当控制第三者恒定时,怀卵数量与体长和体重呈正相关但不显著。棒花鱼存在个体大小和其他局部特征显著的两性异形,雌性个体主要通过腹腔容积的增加提高个体生育力。棒花鱼形态特征的两性异形是性选择和生育力选择共同作用的结果。     

乐山棒花鱼形态特征的两性异形和雌性个体生育力

测定了乐山棒花鱼(Abbottina kiatingensis)繁殖期形态特征包括体长、头长、头宽、头高、吻长、眼后头长、眼径、眼间距、体高、尾柄长、尾柄高、尾鳍长、背鳍基前距、背鳍基长、腹鳍基前距、腹臀间距、体重和去内脏体重的两性异形和雌性个体生育力。繁殖期雄性个体的数量显著多于雌性个体,雌雄两性个体的体长差异不显著。特定体长的雌性个体的头长、头宽、头高、吻长、眼后头长、尾柄高、背鳍基前距、背鳍基长和去内脏体重显著小于雄性个体,其余指标不存在明显的差异。回归分析表明,乐山棒花鱼的怀卵数量与体长和体重回归关系显著,雌性通过个体大小(体长和体重)的增加来提高个体生育力。     

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.     

Sexual dimorphism in the tusked frog, Adelotus brevis (Anura:Myobatrachidae): the roles of natural and sexual selection

At least two adaptive processes can lead to the evolution of sexual dimorphism: sexual selection (e.g. male-male combat) or natural selection (e.g. dietary divergence). We investigated the adaptive significance of a distinctive pattern of sexual dimorphism in a south-eastern Australian frog, Adelotus brevis. Male Adelotus grow larger than female conspecifics, have larger heads relative to body size, and have large paired projections (‘tusks’) in the lower jaw. All of these traits are rare among anurans. We quantified the degree of dimorphism in Adelotus, and gathered data on diets and mating systems of this species to evaluate the possible roles of sexual selection and dietary divergence in favoring die evolution of these sexually dimorphic traits. Analysis of prey items in alimentary tracts revealed significant sex differences in prey types. For example, females ate proportionally more arthropods and fewer molluscs than did males. However, this difference is likely to be a secondary consequence of habitat differences between the sexes (due in turn to their different reproductive roles) rather than a selective force for the evolution of sexual dimorphism. Calling males spend their time in moist habitats where pondsnails are abundant, whereas females are more often encountered in the drier arthropod-rich woodlands. A three-year behavioural ecology study on a field population revealed that reproductive males engage in agonistic interactions, with the sexually dimorphic tusks used to attack rivals. Larger body size contributed to male reproductive success. Small males were excluded from calling sites and, among the calling males, larger animals had higher reproductive success (numbers of matings) than did smaller individuals. Hence, the atypical pattern of sexual dimorphism in Adelotus brevis seems to have resulted from sexual selection for larger body size and tusk size in males, in the context of male-male agonistic behaviour, rather than natural selection for ecological divergence between the sexes.     

Morphological and histological examination of short‐wing formation in the winter moth Protalcis concinnata (Insecta: Lepidoptera,Geometridae)

Winter geometrid moths exhibit sexual dimorphism in wing length and female‐specific flightlessness. Female‐specific flightlessness in insects is an interesting phenomenon in terms of sexual dimorphism and reproductive biology. In the winter geometrid moth, Protalcis concinnata (Wileman), adult females have short wings and adult males have fully developed wings. Although the developmental process for wing reduction in Lepidoptera is well studied, little is known about the morphology and the developmental pattern of short‐winged flightless morphs in Lepidoptera. To clarify the precise mechanisms and developmental processes that produce short‐winged morphs, we performed morphological and histological investigations of adult and pupal wing development in the winter geometrid moth P. concinnata. Our findings showed that (a) wing development in both sexes is similar until larval‐pupal metamorphosis, (b) the shape of the sexually dimorphic wings is determined by the position of the bordering lacuna (BL), (c) the BL is positioned farther inward in females than in males, and (d) after the short pupal diapause period, the female pupal wing epithelium degenerates to approximately two‐thirds its original size due to cell death. We propose that this developmental pattern is a previously unrecognized process among flightless Lepidoptera.     

Relationships among male sexually selected traits in the bean bug,Riptortus pedestris (Heteroptera: Alydidae)

Current concepts of sexual selection suggest that male reproductive success is determined by multiple sexual traits. As expression and production of multiple sexual traits are frequently associated with each other, positive or negative correlations among multiple sexual traits ensue. These relationships among traits associated with male reproductive success may be crucial in the evolution of male reproductive strategies. Here, we investigate phenotypic relationships among sexually selected traits in the armed bean bug Riptortus pedestris. In this insect, males with a larger body and weapon are more likely to win male–male competitions, and males with a larger weapon or higher courtship rate are more attractive to females. There was a significant positive correlation between body size and weapon size, whereas the courtship rate had significant negative correlations with body size and weapon size. Our results suggested that there was a phenotypic trade‐off between courtship rate and male morphology. In this insect, smaller males may make more effort in courtship behavior as an alternative mating tactic.     

Mate Choice and Spawning Success in the Fighting Fish Betta splendens: the Importance of Body Size, Display Behavior and Nest Size

Courtship displays should be exaggerated enough to attract mates and yet tempered so as not to deter them. We tested this hypothesis in the fighting fish Betta splendens by studying courtship displays and body size and their relationships with male parental quality and female fecundity, as well as the effects of display behavior and body size on mate choice decisions and spawning success. Because of their high degree of parental investment, males are expected to be discriminating in their choice of mates. Males who displayed more frequently built larger nests, a measure of parental quality, but larger males did not. When females were paired with males with high display rates, however, the pair had fewer eggs in their nest, even when accounting for female body mass. In a mate choice test using computer‐generated male stimuli that differed only in display behavior, females showed no preferences for displaying males vs. non‐displaying males, or for males with higher display rates vs. lower display rates. In similar tests in which the computer‐generated males differed only in size, females preferred larger males, but also preferred males that differed with respect to body size (negative assortative mating). Males preferred computer‐generated females that performed courtship displays over non‐displaying females, but showed no preferences for female body size. Neither a female's body size nor her display behavior was a significant predictor of her fecundity as estimated by the number of eggs released during spawning. Thus, our results suggest that female B. splendens must balance male parental quality (nest size) with the risk of potentially disruptive or dangerous behavior during spawning, and that females may minimize these risks through negative size‐assortative mating. Female display behavior, while unrelated to fecundity in our study, may attract males because it indicates reproductive readiness or serves a species‐recognition function.     

Male-male competition and mating success in the orb-web spider,Nephila clavata,with reference to temporal factors

Seasonal occurrence patterns of adults of both sexes, intensity of male-male interactions, and mating success in the spider,Nephila clavata, were examined in the field. Adult males began to attend female webs about 2 weeks before female maturation. Large adult males were abundant in the early breeding season, but small males increased later in the season. From the distribution of males among female webs and size relationship of males within a web, male-male interactions seemed to be more intense when most females were still subadult. This was verified by a field experiment in which males were artificially introduced to female webs that were attended by other males. It was found that the probability of introduced males remaining on subadult female webs was lower than that on adult webs. As mating occurred mostly in the period shortly after the female final molt and first male sperm precedence was known in all spiders reported so far, intense male-male competition on subadult female webs seemed to be reasonable. Male longevity had an important influence on the mating success of males with just-molted females. Mating success was also affected by the relative body size of males present in a given period. Since larger males occupied the position closest to females within a web and stayed there longer, relative body size appeared to influence mating success through male-male competition. Female body size at maturation declined with time; hence, males that attained sexual maturity earlier had the advantage of mating with larger and more fecund females. Therefore, early maturation as well as larger size seem to be two important trairs influencing the reproductive success of males.     

Size and sex of cricket prey predict capture by a sphecid wasp

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Morphological correlates of river velocity and reproductive development in an ornamented stream fish

Adaptive phenotypic divergence can arise when environments vary in ways favoring alternative phenotypic optima. In aquatic habitats, the costs of locomotion are expected to increase with water velocity, generally favoring a more streamlined body and the reduction of traits that produce drag. However, because streamlining in fish may come at the cost of maneuverability, the net benefits of drag reduction can differ not only among habitats, but also among individuals (or classes of individuals) that rely on locomotion for different uses (e.g., males vs. females or adults vs. juveniles). We tested these predictions by exploring relationships among river velocity, body streamlining, ornamental fin size, and male reproductive condition in the steelcolor shiner (Cyprinella whipplei), a small-bodied North American cyprinid. Overall, males in peak reproductive condition (defined by the development of sexually dimorphic tubercles) had less streamlined bodies and larger ornamental fins than males in lower reproductive condition or individuals lacking these secondary sexual characters (females and immature males). There was a relationship between river velocity and body streamlining only for males in peak reproductive condition, but it was in the opposite direction of our predictions: these males were less streamlined in faster rivers. We found only weak support for the prediction that ornamental fin size would be negatively associated with river velocity. Overall, these results suggest either that drag is not an important selective pressure in these habitats, or that the sexual selection advantages of a deep body and large fin compensate any natural selection costs for C. whipplei males. This study highlights the often overlooked diversity of selective pressures acting on streamlining in fishes, and can offer novel insights and predictions allowing a more nuanced understanding of fish ecomorphology.