Sexual selection for male dominance reduces opportunities for female mate choice in the European bitterling (Rhodeus sericeus)

Sexual selection involves two main mechanisms: intrasexual competition for mates and intersexual mate choice. We experimentally separated intrasexual (male-male interference competition) and intersexual (female choice) components of sexual selection in a freshwater fish, the European bitterling (Rhodeus sericeus). We compared the roles of multiple morphological and behavioural traits in male success in both components of sexual competition, and their relation to male reproductive success, measured as paternity of offspring. Body size was important for both female choice and male-male competition, though females also preferred males that courted more vigorously. However, dominant males often monopolized females regardless of female preference. Subordinate males were not excluded from reproduction and sired some offspring, possibly through sneaked ejaculations. Male dominance and a greater intensity of carotenoid-based red colouration in their iris were the best predictors of male reproductive success. The extent of red iris colouration and parasite load did not have significant effects on female choice, male dominance or male reproductive success. No effect of parasite load on the expression of red eye colouration was detected, though this may have been due to low parasite prevalence in males overall. In conclusion, we showed that even though larger body size was favoured in both intersexual and intrasexual selection, male-male interference competition reduced opportunities for female choice. Females, despite being choosy, had limited control over the paternity of their offspring. Our study highlights the need for reliable measures of male reproductive success in studies of sexual selection.     

Mate preference in female electric fish, Brachyhypopomus pinnicaudatus

Weakly electric fish communicate with brief electrostatic field pulses called electric organ discharges (EODs). EOD waveforms are sexually dimorphic in most genera, a condition thought to result from mate choice acting to shape the electric signal's constituent action potentials. We have no direct behavioural evidence that sexual selection by either mate choice or intrasexual competition is responsible for sex differences in the EOD waveforms of electric fish. We explored sexual selection in electric fish by conducting two-choice unforced preference tests with live, unaltered gymnotiform electric fish,Brachyhypopomus pinnicaudatus , which are sexually dimorphic. In the initial test, gravid females selected males over females only when the males were larger than average. Gravid females in later tests preferred larger males to smaller males in a significant majority of those trials in which they showed a preference. In about one-third of those trials, females spawned with their preferred male, confirming their preference. We concluded that passage through the choice apparatus was related to mate choice. The signals of chosen males had larger EOD amplitudes and longer EOD durations. These findings show that femaleB. pinnicaudatus do have a preference for a certain male phenotype. The system requires additional study to dissociate correlated male phenotypic characters to identify which male traits the female prefers. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Diet of characin,Brycon falcatus (Muller and Troschel, 1844) in the Amazon Basin: a case study on an attractant for fish in the Teles Pires River

The use of fresh soybeans as an attractant in fishing has been altering the diet of the characiform Brycon falcatus, which are thriving at fishing spots. This has subsequently affected the amount of abdominal fat in the species from the Tapajós River basin, Southern Amazon. In the Teles Pires River basin region, one of the most common capture techniques employed by fishermen is the use of attractant feeds (soybeans, corn, cassava) at certain locations to attract mostly omnivorous fish. Tourists also feed fish chips and breadcrumbs. In the experimental design, rivers were mapped according to attractant availability to develop an attractant‐density classification system comprising four rivers with different attractant densities (low, medium, and high), plus a river serving as control (no attractants). Monthly collections were carried out during the 2012 and 2013 dry seasons. For diet analyses, methods of frequency of occurrence and relative volume were used to calculate the feeding index (IAi) of 97 specimens. For analyses of abdominal fat, an index was calculated by dividing the wet weight of the abdominal fat by the total wet weight of each individual. Standard length ranged from 15.0 to 48.5 cm, and weight between 0.18 and 4.40 kg. Composition and diversity of diet items changed with the increasing density of attractants. In the river with a high attractant density, fresh soybeans and corn were dominant diet items; in the control river, the diet was natural (e.g. seeds, fruits, leaves, and insects) and therefore similar to those described for B. falcatus in undisturbed environments. Fish collected from rivers with high attractant densities were in better condition than those from the river with no attractants. Although use of attractants is prohibited by State legislation, there are no inspections. This study clearly shows an alteration in the natural diet and abdominal fat of B. falcatus resulting from an imbalanced, high‐calorie feed via an attractant. It is recommended that these areas be monitored for the preservation of B. falcatus.     

Attunement to the fertility status of same-sex rivals: women's testosterone responses to olfactory ovulation cues

Evolutionary theories of mating suggest that changes in fertility across the menstrual cycle play an important role in sexual selection. In line with this framework, the current research examined whether olfactory cues to the fertility of a same-sex rival would prompt hormonal signs of intrasexual competition in women. Women exposed to the scent of another woman close to ovulation subsequently displayed higher levels of testosterone than women exposed to the scent of a woman far from ovulation. Whereas women exposed to the scent of a woman in the mid-luteal phase displayed sizable decreases in testosterone over time, no such decline was observed among women exposed to the scent of a woman near ovulation. Thus, olfactory cues signaling a rival's heightened level of fertility were associated with endocrinological responses in women that could be linked to intrasexual competition.     

Do Red Deer Stags (Cervus elaphus) Use Roar Fundamental Frequency (F0) to Assess Rivals?

It is well established that in humans, male voices are disproportionately lower pitched than female voices, and recent studies suggest that this dimorphism in fundamental frequency (F0) results from both intrasexual (male competition) and intersexual (female mate choice) selection for lower pitched voices in men. However, comparative investigations indicate that sexual dimorphism in F0 is not universal in terrestrial mammals. In the highly polygynous and sexually dimorphic Scottish red deer Cervus elaphus scoticus, more successful males give sexually-selected calls (roars) with higher minimum F0s, suggesting that high, rather than low F0s advertise quality in this subspecies. While playback experiments demonstrated that oestrous females prefer higher pitched roars, the potential role of roar F0 in male competition remains untested. Here we examined the response of rutting red deer stags to playbacks of re-synthesized male roars with different median F0s. Our results show that stags’ responses (latencies and durations of attention, vocal and approach responses) were not affected by the F0 of the roar. This suggests that intrasexual selection is unlikely to strongly influence the evolution of roar F0 in Scottish red deer stags, and illustrates how the F0 of terrestrial mammal vocal sexual signals may be subject to different selection pressures across species. Further investigations on species characterized by different F0 profiles are needed to provide a comparative background for evolutionary interpretations of sex differences in mammalian vocalizations.     

Sexually Dimorphic Intrasexual Duetting in an Otherwise Monomorphic Green Lacewing (Neuroptera, Chrysopidae, Chrysoperla plorabunda): Sexual Selection or Sex Recognition?

Songs produced during heterosexual duets in a green lacewing, C. plorabunda, are sexually monomorphic. However, individuals of either sex will also engage in intrasexual duets, which can exhibit sexual dimorphism. We confined males and females together in various combinations in a small arena to study the phenotypes, behavioral interactions, and functional roles of duetting in this species. The goal was to test whether sexual selection or sex recognition provided the better explanation of song sexual dimorphism. We determined that the monomorphic form of the intrasexual duet was long and stable, and could take place either between males or between females. Such “standard” intrasexual duetting songs were acoustically indistinguishable from heterosexual songs. However, males could also engage other males in special “fast duets” that sped up and terminated abruptly. Equivalent fast duets were not part of the female repertory. Fast duetting songs between males differed significantly from other types of male or female duetting songs in every measurable characteristic, but their role in the mating system was ambiguous. Contrary to one prediction of the sexual selection hypothesis, fast duetting between males occurred less often in situations where it might be the most useful to males in securing mates, i.e., during male-male-female interactions (trios). In addition, fast songs that started, ended, both started and ended, or neither started nor ended duets were acoustically indistinguishable, making it unlikely that females were choosing males based on such variation. However, songs that “both started and ended” fast duets were associated with a significant mating advantage, indicating a possible role for fast duetting in male-male sexual competition. Because the alternative hypothesis of sex recognition was also supported by some of our results, we conclude that aggressive qualities of male-male fast duets probably mediate intrasexual selection, while their increasing tempo serves as an adaptive response to promote rapid sex recognition by truncating unproductive and potentially dangerous intrasexual duetting.     

The relative importance of intra- and intersexual selection on human male sexually dimorphic traits

Recent evidence suggests that in sexual selection on human males, intrasexual competition plays a larger role than female choice. In a sample of men (N?=?164), we sought to provide further evidence on the effects of men's physical dominance and sexual attractiveness on mating success and hence in sexual selection. Objective measures and subjective ratings of male sexually dimorphic traits purportedly under sexual selection (height, vocal and facial masculinity, upper body size from 3D scans, physical strength, and baseline testosterone) and observer perceptions of physical dominance and sexual attractiveness based on self-presentation video recordings were assessed and associated with mating success (sociosexual behaviour and number of potential conceptions) in a partly longitudinal design. Results from structural equation models and selection analyses revealed that physical dominance, but not sexual attractiveness, predicted mating success. Physical dominance mediated associations of upper body size, physical strength, as well as vocal and facial physical dominance and attractiveness with mating success. These findings thus suggest a greater importance of intrasexual competition than female choice in human male sexual selection.     

The role of sex ratios and resource availability on the pre-mating behavior of a monogamous fish

Convict cichlid fish (Archocentrus nigrofasciatus) are one of the few monogamous species where the female appears more colorful than the male. We examined whether this sexual dimorphism was reflected in a female-biased actual sex ratio (ASR) and whether convict cichlid females would exhibit behavioral traits typified in monogamous males when they possessed the extravagant colors and/or morphology. Our field observations revealed that females are marginally, but not significantly, more abundant than males but they did initiate more intersexual social interactions (i.e. approach behavior) than males. They also showed more intrasexual aggression. Furthermore, males more commonly chased females rather than vice versa. Our laboratory experiment also indicated that differences in intrasexual aggression were not related to differences in the ASR but did appear with the addition of a breeding site. Thus, while intrasexual competition was more frequently observed between females in the field it was probably related to the availability of breeding sites rather than our estimates of the ASR.     

Sexual Selection in the Loud Calls of Male Primates: Signal Content and Function

Researchers have used sexual selection theory and hypotheses based on intersexual mate choice and intrasexual mate competition to explain the role of spontaneous long-distance vocalizations emitted by adult male primates, relying on the tacit assumption that assessment or identity cues are encoded in the vocalizations. I review the published literature and aim to substantiate a relationship between sexual selection and long-distance vocal communication in primates. First, I review findings from nonprimate taxa to determine the relative importance of inter- and intrasexual selection and to provide a background for examining primates. Next, I describe several hypotheses for signal content and function in adult male loud calls. Then, I examine the available data across Primates for evidence to support or to refute these hypotheses and to determine if they meet proposed criteria for demonstrating sexual selection [Snowdon, C. T. (2004). Sexual Selection in Primates: New and Comparative Perspectives]. Signal content refers to patterns of acoustic features within vocalizations from which listeners might extract cues or information about the signaler. I interpret signal function, in turn, from behavioral responses of receivers and assume it has ultimate effects on the evolution and design of acoustic signals if direct fitness consequences exist. After the general review across primates, I propose orangutans as a candidate species for further evaluation of sexual selection in vocal communication. The available evidence corroborates a demonstrable relationship between sexual selection and adult male loud calls based on individual recognition, but it is necessary to obtain additional data to affirm a direct benefit to reproductive success.     

Intra‐ and Intersexual swim bladder dimorphisms in the plainfin midshipman fish (Porichthys notatus): Implications of swim bladder proximity to the inner ear for sound pressure detection

The plainfin midshipman fish, Porichthys notatus , is a nocturnal marine teleost that uses social acoustic signals for communication during the breeding season. Nesting type I males produce multiharmonic advertisement calls by contracting their swim bladder sonic muscles to attract females for courtship and spawning while subsequently attracting cuckholding type II males. Here, we report intra‐ and intersexual dimorphisms of the swim bladder in a vocal teleost fish and detail the swim bladder dimorphisms in the three sexual phenotypes (females, type I and II males) of plainfin midshipman fish. Micro‐computerized tomography revealed that females and type II males have prominent, horn‐like rostral swim bladder extensions that project toward the inner ear end organs (saccule, lagena, and utricle). The rostral swim bladder extensions were longer, and the distance between these swim bladder extensions and each inner‐ear end organ type was significantly shorter in both females and type II males compared to that in type I males. Our results revealed that the normalized swim bladder length of females and type II males was longer than that in type I males while there was no difference in normalized swim bladder width among the three sexual phenotypes. We predict that these intrasexual and intersexual differences in swim bladder morphology among midshipman sexual phenotypes will afford greater sound pressure sensitivity and higher frequency detection in females and type II males and facilitate the detection and localization of conspecifics in shallow water environments, like those in which midshipman breed and nest.     

Social plasticity in non-territorial male African cichlid fish <Emphasis Type="Italic">Astatotilapia burtoni</Emphasis>

The African cichlid fish Astatotilapia burtoni represents a valuable model system for studies of social decision-making due to its socially mediated phenotypic plasticity. The males reversibly transition in social status from reproductively dominant and territorial (DOM) to submissive and non-territorial (SUB). Males are traditionally categorized into these two behavioral phenotypes by observational scoring. There is evidence, however, that this dichotomous categorization might not capture the behavioral plasticity displayed by individuals transitioning between SUB and DOM status. To test this concept, we used focal observations of intrasexual conflict behavior in fish communities combined with a modified analysis of the ethogram typically used in A. burtoni. Results revealed a cluster of males close to the crossover point between SUB and DOM status as defined by the traditional dominance index. These intermediate males showed the highest frequency of intrasexual conflict behaviors, distinct behavioral responses to threats, and body pigment signaling displays that distinguish them from prototypical SUBs and DOMs. As such, our results provide a noninvasive behavioral metric to categorize A. burtoni males into three groups, thus further capturing the complex social dynamic of this model organism.     

Intrasexual competition underlies sexual selection on male breeding coloration in the orangethroat darter,Etheostoma spectabile

Elaborate, sexually dimorphic traits are widely thought to evolve under sexual selection through female preference, male–male competition, or both. The orangethroat darter (Etheostoma spectabile) is a sexually dichromatic fish in which females exhibit no preferences for male size or coloration. We tested whether these traits affect individual reproductive success in E. spectabile when multiple males are allowed to freely compete for a female. The quality and quantity of male coloration were associated with greater success in maintaining access to the female and in spawning as the primary male (first male to participate). On the other hand, sneaking behavior showed little correlation with coloration. Male breeding coloration in E. spectabile may therefore demonstrate how intrasexual competition can be a predominant factor underlying the evolution of male ornaments.     

Olfactory sensitivity to conspecific bile fluid and skin mucus in the European eel Anguilla anguilla (L.)

The present study assessed the olfactory potency of conspecific bile fluid and skin mucus in the European eel Anguilla anguilla by the electro-olfactogram. Immature males showed high olfactory sensitivity to conspecific bile, giving large amplitude responses in a concentration-dependent manner with estimated thresholds of detection of <1:10⁷ ( n = 6). Mucus also proved to contain highly potent odorants with thresholds of detection of c . 1:10⁶ ( n = 6). Crude solid-phase extraction of bile fluid (C-18 and C-2/ENV+ cartridges) showed that the majority of olfactory activity in bile fluid was contained in the eluate of C-18 cartridges ( n = 6). There were quantitative differences, however, between the sexes; female bile fluid had a higher proportion of activity in this fraction. Similar solid-phase extraction of mucus showed that it contains a higher proportion of odorants in the C-18 filtrate than bile fluid. Mucus from mature eels, however, had a higher proportion of olfactory activity in the eluate than immature fish ( n = 6). Cross-adaptation experiments suggest that there are qualitative differences in the odorants contained in bile and mucus depending on both the sex and state of sexual maturation of the donor ( n = 6). These results are consistent with a role for chemical communication in the reproduction of the European eel and suggest that both bile and mucus are potential sources of the odorants involved.     

Relationship of sexual dimorphism in canine size and body size to social,behavioral, and ecological correlates in anthropoid primates

Among anthropoid primates there are interspecific differences in the degree of sexual dimorphism in both body size and canine size. Within the suborder body size dimorphism and canine size dimorphism are positively correlated,r=0.76. This correlation suggests that the two dimorphisms are equally developed in some species, while in other species there is a differential degree of sexual dimorphism. An analysis of these results and their relation to social organization and other ecological variables reveals: (1) the degree of canine size dimorphism is closely related to the amount of male intrasexual selection in a given mating system; and (2) the degree of body size dimorphism is also related to male intrasexual selection, but may be modified (either enhanced or diminished) by selection pressure from factors such as habitat, diet, foraging behavior, antipredator behavior, locomotory behavior, and female preference.     

The evolution of sexually dimorphic earwig forceps: social interactions among adults of the toothed earwig, Vostox apicedentatus

Earwigs (Insecta, Dermaptera) are characterized by uniquelyelaborated cerci, commonly called forceps, the function of whichremains unclear. We studied intrasexual and intersexual interactionsin the laboratory to examine the context and pattern of forcepsuse in the toothed earwig. Vostox apictdenlatus (Caudell). Interactionsbetween pairs of earwigs were recorded in four social situations:(1) two males, (2) two males plus a virgin female, (3) two females,and (4) one male and one female. Forceps were used as both weaponsand display structures by males and females in all of thesesocial contexts. During pairwise male-male interactions, onemale clearly dominated the other male. Dominant males were moreactive and more likely to use their forceps in intrasexual interactionsthan were subordinate males. In interactions where there weretwo males and one female present, the male that dominated male-maleinteractions was able to maintain exclusive access to the female.There was no indication of active female choice during or aftercourtship. During intersexual interactions, only males usedtheir forceps during courtship. The behavioral repertoire involvingforceps was greater for males than for females, especially inintrasexual contests. There was no clear outcome of intrasexualinteractions among females. These results suggest that forcepsfunction mainly as weapons in male-male interactions and mayhave evolved, at least in part, as a result of sexual selection.Further research is required to test for female mate choiceand to separate the various mechanisms of sexual selection ifmate choice exists. Comparative studies are needed to determineif sexual selection was the original evolutionary mechanismleading to the development of these unusual structures or ifsexual selection is relegated to a secondary effect, leadingto the elaboration and sexualdimorphism of these structuresin selected groups of earwigs.     

Length–weight relationship and condition factor of the Characidae matrinxã, Brycon falcatus (Müller & Troschel, 1844), in the Teles Pires River,southern Amazon

The aim of this study was to verify whether the in situ supply of soybean as a fish attractant bears any influence on the length–weight relationship and condition factor of the omnivorous fish species, Brycon falcatus. In addition, estimated was the length–weight relationship, standard and total length, condition factor, and size of B. falcatus in the spawning period in the Teles Pires River Basin. In the experimental design, rivers of the Teles Pires Basin containing soybean attractant (Verde River, Celeste River, Teles Pires River and the Tapaiuna River) were mapped. Fish were also collected from the Cristalino River, which contained no soybean attractant (treatment control). Samples were collected from August 2012 to July 2013. Length–weight relationship of B. falcatus in the Teles Pires River Basin can be obtained by the expression: Wt = 0.0091*SL^3.376, r² = .97, n = 102, and length‐length relationship: TL = 1.128*SL + 3.978, r² = .96. The condition factor of all specimens collected in the Teles Pires Basin was k = 1.055. Size at capable phase (L50) of spawning was estimated at 23 cm standard length. Results showed no significant difference in the length–weight relationship or condition factor of B. falcatus collected from rivers with or without soybean attractant. Concluded is that the in situ supply of soybean as a fish attractant does not interfere with the wellbeing of B. Falcatus in the Teles Pires River Basin.     

Genetic evidence for the mating system and reproductive success of black sea bream (Acanthopagrus schlegelii)

Understanding the mating system and reproductive success of a species provides evidence for sexual selection. We examined the mating system and the reproductive success of captive adult black sea bream (Acanthopagrus schlegelii), using parentage assignment based on two microsatellites multiplex PCR systems, with 91.5% accuracy in a mixed family (29 sires, 25 dams, and 200 offspring). Based on the parentage result, we found that 93.1% of males and 100% of females participated in reproduction. A total of 79% of males and 92% of females mated with multiple partners (only 1 sire and 1 dam were monogamous), indicating that polygynandry best described the genetic mating system of black sea bream. For males, maximizing the reproductive success by multiple mating was accorded with the sexual selection theory while the material benefits hypothesis may contribute to explain the multiple mating for females. For both sexes, there was a significant correlation between mating success and reproductive success and the variance in reproductive success of males was higher than females. Variation in mating success is the greatest determinant to variation in reproductive success when the relationship is strongly positive. The opportunity for sexual selection of males was twice that of females, as well as the higher slope of the Bateman curve in males suggested that the intensity of intrasexual selection of males was higher than females. Thus, male–male competition would lead to the greater variation of mating success for males, which caused greater variation in reproductive success in males. The effective population number of breeders (N_b) was 33, and the N_b/N ratio was 0.61, slightly higher than the general ratio in polygynandrous fish populations which possibly because most individuals mated and had offspring with a low variance. The relatively high N_b contributes to the maintenance of genetic diversity in farmed black sea bream populations.     

Quantitative Evaluation of Erotactin Secretion in Eggs of Hormosira banksii (Fucales,Phaeophyceae)

The release of the C₁₁-hydrocarbon spermatozoid attractant hormosirene by eggs of the Australian fucoid Hormosira banksii has been determined quantitatively using a closed-loop extraction technique. The eggs secrete erotactin for more than 2 weeks with an exponentially decreasing rate. The initial secretion rate was estimated to be 75 fmol h^?1 egg^?1 and the total amount of hormosirene produced per egg to be 2.35 pmol (0.35 ng). The results were used to calculate the threshold range for spermatozoid chemotaxis in the pheromone gradient around eggs.     

Sexual Selection in the Three-Spined Stickleback

The purpose of this study was to seek evidence for the influence of sexual selection on both male and female three-spined sticklebacks, a species in which the ♂ alone has parental responsibilities. We studied the intrasexual relationships of ♂♂ and ♀♀ prior to courtship, and the intrasexual and intersexual relationships of both sexes during courtship. Results demonstrated that both ♂♂ and ♀♀ compete intrasexually, and suggested that both sexes are discriminative in mate selection. These results are compatible with Trivers' (1972) model of the role of parental investment in sexual selection.     

Sexual selection for low male voice pitch among Amazonian forager-horticulturists

Pitch is the most perceptually salient feature of the voice, yet it is approximately five standard deviations lower in men than in women, a degree of sexual dimorphism exceeding that of all extant nonhuman apes. Evidence from Western samples suggests that low-frequency vocalizations may have augmented male mating success ancestrally by intimidating competitors and/or attracting mates. However, data are lacking from small-scale societies. We therefore investigated sexual selection on male pitch (measured by fundamental frequency, f_o) in a population of Bolivian forager-horticulturists, the Tsimané. We found that experimentally lowering f_o in audio clips of men speaking increased perceptions of fighting ability but did not affect perceptions of prestige and decreased their attractiveness to women. Further, men with lower speaking f_o reported higher numbers of offspring, and this was mediated by the reproductive rates of men's wives, suggesting that men with lower f_o achieved higher reproductive success by having access to more fertile mates. These results thus provide new evidence that men's f_o has been shaped by intrasexual competition.