Conspicuous Female Ornamentation and Tests of Male Mate Preference in Threespine Sticklebacks (Gasterosteus aculeatus)

Sexual selection drives the evolution of exaggerated male ornaments in many animal species. Female ornamentation is now acknowledged also to be common but is generally less well understood. One example is the recently documented red female throat coloration in some threespine stickleback (Gasterosteus aculeatus) populations. Although female sticklebacks often exhibit a preference for red male throat coloration, the possibility of sexual selection on female coloration has been little studied. Using sequential and simultaneous mate choice trials, we examined male mate preferences for female throat color, as well as pelvic spine color and standard length, using wild-captured threespine sticklebacks from the Little Campbell River, British Columbia. In a multivariate analysis, we found no evidence for a population-level mate preference in males, suggesting the absence of directional sexual selection on these traits arising from male mate choice. Significant variation was detected among males in their preference functions, but this appeared to arise from differences in their mean responsiveness across mating trials and not from variation in the strength (i.e., slope) of their preference, suggesting the absence of individual-level preferences as well. When presented with conspecific intruder males, male response decreased as intruder red throat coloration increased, suggesting that males can discriminate color and other aspects of phenotype in our experiment and that males may use these traits in intrasexual interactions. The results presented here are the first to explicitly address male preference for female throat color in threespine sticklebacks.     

Male Sexual Signals and Female Choice in Drosophila grimshawi (Diptera: Drosophilidae)

Sexual selection and sexual signaling have been prominent topics in recent behavioral studies, but limited data have led to controversy regarding these topics. For example, the Hawaiian Drosophila are often cited as examples in which female choice has resulted in the evolution of elaborate male courtship signals, but relatively few data exist to test these claims adequately. We studied D. grimshawi, a lek-forming Hawaiian Drosophila, to determine whether there was evidence for female choice without male competition and to elucidate the possible cues females use to discriminate. Male mating success was found to be nonrandom and males that courted females intensely and deposited many pheromone-containing streaks on the substrate were the most successful. Hence, multiple cues seem to be involved in male mating success in this species. Some males performed only one display, however, and may represent an alternate male mating tactic. The protein content of the adult male diet significantly influenced the level of pheromone streak deposition, and thus, foraging environment may affect the outcome of sexual selection.     

Coevolution of Male and Female Response Preferences to Sexual Signals in Music Frogs

Male signaling influences both female choice and male-male competition. Although male signaling characteristics and female preferences have been shown to coevolve in many species, few studies have examined whether male signal characteristics and male receiver responses related to male-male competition also coevolve. The present study tested the hypothesis that male and female signal receiver preferences may coevolve in parallel for frogs in the genus Babina by comparing the acoustic structure of male advertisement calls of four closely related and geographically isolated Babina species. Then we assessed the behavioral responses of both male and female B. daunchina(Emei music frog) to male call playbacks from each of the four species. The results support the hypothesis that male and female signal receiver preferences have coevolved in this species. Specifically, both male and female B. daunchina respond strongly to the heterospecific calls of B. hainanensis, suggesting that preexisting biases exist in both females and males. Both male and female individuals showed a slight response to the calls of B. adenopleura while no response was evoked by the calls of B. lini. The manifestation of similar response profiles in male and female B. daunchina to the calls of the four species support the idea that male and female signal receiver preferences evolved in parallel and that the origin of these receiver biases reflects adaptations dependent on the same neural and cognitive systems in both sexes.     

Sexual Dichromatism and Female Preference in Eulemur fulvus Subspecies

We experimentally tested the hypothesis that sexual dichromatism in the subspecies of Eulemur fulvus is the evolutionary result of female preference for brightly colored males. Ten female lemurs representing 6 different subspecies of Eulemur fulvus were subjects in the experiment; controls were 4 females of non-sexually dichromatic lemurid taxa. For each taxon we presented photographs of the face of a male of that taxon whose colors had been digitally altered to make him less and more colourful. Median viewing times of the pooled female Eulemur fulvus are significantly correlated with colorfulness. Viewing times in the control females are not correlated with color or brightness of the stimulus photographs. We concluded that the females of the Eulemur fulvus sspp. preferred to view photographs of more colorful males, which is consistent with the predictions of sexual selection theory.     

Male Ornamentation in the European Minnow (Phoxinus phoxinus) Signals Swimming Performance

The association between male secondary sexual ornamentation and male quality has been under intensive study for decades. It has been shown that male ornaments in fish indicate a variety of physical and genetic qualities. However, only few studies have investigated whether male ornaments signal locomotor performance, a trait that is expected to be vital for individual fitness in the nature. In this study, we sampled two populations of European minnow (Phoxinus phoxinus), a cyprinid fish species with spectacular and colourful male breeding ornamentation, and examined the association between male ornaments (tubercle number, lateral darkness and abdominal redness), swimming performance, and three fitness‐related traits, body length (L_T), condition factor (K) and gonadosomatic index (GI). We found that male tubercle number and abdominal redness were positively correlated with their swimming performance. In addition, these two ornaments were correlated with male fitness‐related traits, which in turn were also directly associated with the swimming performance. In general, these results suggest that in the European minnow male ornamentation may act as an honest signal of several fitness‐related traits and locomotor performance in the nature.     

The Role of Visual Ornamentation in Female Choice of a Multimodal Male Courtship Display

The courtship behavior of male Schizocosa uetzi wolf spiders incorporates both visual and seismic signals into a multimodal display. These two signals have been shown to interact in such a manner that the seismic signal alters a female's response to the visual signal, leading to a putative increased importance of visual signaling in the presence of a seismic signal. Experiments leading to this attention‐focusing hypothesis relied in part on the video playback technique, eliciting the question of its significance under more biologically relevant conditions. Here, we directly examine female mate choice of males with differing visual signals (foreleg pigmentation) both in the presence and absence of a seismic courtship signal. We first quantified the natural variation of male foreleg pigmentation within a population of S. uetzi. The proportion of the tibia covered in pigmentation was found to be positively correlated with male weight, suggesting that this signal may convey reliable information about male size. Visual signals of live males were then manipulated into two treatments: black and brown male foreleg tibias, representing the extreme ends of the natural variation found. The seismic signaling environment was also manipulated into two treatments: seismic signal present and absent. Mating frequency was higher in the presence of a seismic signal than in its absence, but there was no interaction between the seismic and visual signaling treatments. Females mated with black and brown males equally whether a seismic signal was present or absent. This study suggests that inexperienced females do not distinguish between males of different manipulated foreleg pigmentations in mate‐choice decisions, even when in the presence of a seismic courtship signal.     

Androgen Induction of Male Sexual Behaviors in Female Goldfish

The effectiveness of testosterone (T) and 11-ketotestosterone (K) in inducing male-typical sex behaviors in goldfish was examined by implanting intact adult females with one empty (blank) Silastic implant (B females), one implant containing T or K, or one T and one K implant (T + K females). Behavior of the four female groups was compared to that of untreated males and males containing a blank implant. Male-typical behaviors (coutship, spawning) and associated behavioral changes (increased activity, reduced spontaneous feeding) were assessed 3.5 and 4.5 months after implant in 30-min tests in which the test female or male was allowed to interact with a stimulus female in which sexual receptivity and attractivity had been induced by acute prostaglandin F_2αinjection. Prostaglandin-induced female-typical spawning behavior in the test females and males was also assessed 4.5 months after implant in a 60-min test for female-typical behavior in which the test fish was injected with prostaglandin and placed immediately with a sexually active male. Blood samples 5 months postimplant showed that implants generated physiological levels of T and K. In both tests for male-typical behaviors, K and T + K females exhibited the full suite of behaviors shown by spawning males, e.g., male-typical courtship and spawning, increased swimming activity, and reduced spontaneous feeding. Although behaviors of K and T + K females did not differ, those of T + K females were more often equivalent to those of males and significantly different from those of B females. T females exhibited marginal male-typical behaviors which never differed significantly from those of B females. Androgen-treated females exhibited female-typical; spawning behaviors equivalent to that of males and B females. The results show that adult female goldfish can be behaviorally masculinzed without behavioral defeminization, and suggest that male-typical sex behaviors in goldfish are dependent on K, although other steroids also may be required. The inducible behavioral bisexuality of goldfish, a gonochoristic species, is discussed in terms of the prevalence of hermaphroditism in teleosts.     

Brain Serotonin Signaling Does Not Determine Sexual Preference in Male Mice

It was reported recently that male mice lacking brain serotonin (5-HT) lose their preference for females (Liu et al., 2011, Nature, 472, 95–100), suggesting a role for 5-HT signaling in sexual preference. Regulation of sex preference by 5-HT lies outside of the well established roles in this behavior established for the vomeronasal organ (VNO) and the main olfactory epithelium (MOE). Presently, mice with a null mutation in the gene for tryptophan hydroxylase 2 (TPH2), which are depleted of brain 5-HT, were tested for sexual preference. When presented with inanimate (urine scents from male or estrous female) or animate (male or female mouse in estrus) sexual stimuli, TPH2-/- males show a clear preference for female over male stimuli. When a TPH2-/- male is offered the simultaneous choice between an estrous female and a male mouse, no sexual preference is expressed. However, when confounding behaviors that are seen among 3 mice in the same cage are controlled, TPH2-/- mice, like their TPH2+/+ counterparts, express a clear preference for female mice. Female TPH2-/- mice are preferred by males over TPH2+/+ females but this does not lead to increased pregnancy success. In fact, if one or both partners in a mating pair are TPH2-/- in genotype, pregnancy success rates are significantly decreased. Finally, expression of the VNO-specific cation channel TRPC2 and of CNGA2 in the MOE of TPH2-/- mice is normal, consistent with behavioral findings that sexual preference of TPH2-/- males for females is intact. In conclusion, 5-HT signaling in brain does not determine sexual preference in male mice. The use of pharmacological agents that are non-selective for the 5-HT neuronal system and that have serious adverse effects may have contributed historically to the stance that 5-HT regulates sexual behavior, including sex partner preference.     

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.     

Infanticide as Sexual Conflict: Coevolution of Male Strategies and Female Counterstrategies

One of the earliest recognized forms of sexual conflict was infanticide by males, which imposes serious costs on female reproductive success. Here I review two bodies of evidence addressing coevolved strategies of males and females. The original sexual selection hypothesis arguing that infanticide improves male mating success by accelerating the return of females to fertilizable condition has been generally supported in some taxa—notably, some primates, carnivores, rodents, and cetaceans—but not in other taxa. One result of recent research has been to implicate other selective benefits of infanticide by males in various taxa from insects to birds to mammals, such as acquisition of breeding status or improvement of the female breeding condition. In some cases, however, the adaptive significance of male infanticide remains obscure. The second body of data I review is arguably the most important result of recent research: clarifying the possible female counterstrategies to infanticide. These potential counterstrategies span diverse biological systems, ranging from sexual behavior (e.g., polyandrous mating), to physiology (e.g., the Bruce effect), to individual behavior (e.g., maternal aggression), to social strategies (e.g., association with coalitionary defenders of either sex). Although much remains to be studied, these current data provide compelling evidence of sexually antagonistic coevolution surrounding the phenomenon of infanticide.At its most elemental level, infanticide is the killing of a newborn individual by a conspecific. With the growing appreciation of its biological significance, however, infanticide came to be defined more broadly as any “behavior that makes a direct and significant contribution to the immediate death of an embryo or newly hatched or born member of the performer’s own species” (Mock 1984, p. 4) or “any form of lethal curtailment of parental investment in offspring brought about by conspecifics” (Hrdy and Hausfater 1984, p. xv). These definitions highlight the heterogeneous and variable nature of the phenomenon, which can be perpetrated by either sex, by parents or other kin, by individuals unrelated to the victim, in a wide variety of social and mating systems, under a range of seasonal or aseasonal breeding regimes, and across diverse taxa straddling vertebrates and invertebrates.One adaptive form of infanticide—the killing of infants by unrelated males—is arguably the archetype of sexual conflict. In 450 BCE, Herodotus not only documented the behavior among Egyptian cats, but explained it as a male “trick” to obtain sexual access to females otherwise preoccupied with maternal duties (Delibes et al. 2012). Among the myriad ideas inaugurating sociobiology in the 1970s, the hypothesis that infanticide is a male strategy that improves reproductive success at the expense of female fitness (Hrdy 1974) constituted one of the first demonstrations of the “battle of the sexes” theory developed by Williams (1966) and Trivers (1972). Partly because of the controversy surrounding the appearance of this hypothesis (Rees 2009), however, subsequent research focused more on male strategy than on the other party in this sexual dialectic, the female. Thus, field and laboratory research has helped to establish its many forms and conditional occurrence, describe its distribution across taxa, and clarify its adaptive significance, but it is only relatively recently that female counterstrategies have become the subjects of rigorous study, even though their potential importance was grasped early on (Hrdy 1979).In this article, I review selected aspects of this body of data and analysis. My focus is on nonparental male infanticide targeting dependent young—in mammals, nursing individuals—as opposed to older, weaned offspring, the killing of which is variably rendered “juvenilicide,” “pedicide,” or “filicide” (e.g., Agoramoorthy and Mohnot 1988; Palombit 2014, in press).     

Reproductive Signals of Female Lizards: Pattern of Trait Expression and Male Response

Relative to the volume of studies concerning the function and evolution of male‐biased sexually dimorphic traits, instances of female‐biased sexual dimorphisms remain largely unstudied, especially in species with conventional sex roles. I investigated the signal function of a female‐specific ornamental trait using the striped plateau lizard (Sceloporus virgatus, Phrynosomatidae) as a model system. During the reproductive season, female S. virgatus develop orange color on their throats that is absent in conspecific males. I established the relationship between color expression and female reproductive state, and determined male response to female color. I show that dynamic changes occurring within the color patch can potentially identify each stage of the female reproductive cycle, largely because of a lag in patch growth relative to color intensification. Sexual receptivity is associated with intense patches rapidly growing in size; ovulation occurs near peak color expression; and the unreceptive period is associated with large patches fading in intensity. Because females express orange color during both the receptive and unreceptive periods, the pattern of color expression is consistent with the courtship‐stimulation and courtship‐rejection hypotheses of signal function. Males may preferentially associate with females that have more highly developed color patches during the courtship season, and/or ignore such females when they are unreceptive. An examination of male behavior towards unfamiliar females indicates that female color has a role in courtship stimulation but has little, if any, role in courtship rejection. During the pre‐mating season, males maintained significantly closer affiliation with, and tended to perform more social behavior towards females with more intense color. During the post‐mating season, female color had no apparent effect on male behavior. The evolution and current function of female ornaments may vary among taxonomically‐related species as a result of differences in ecology, social system, and life‐history.     

Sexual Partner Preference in Female Zebra Finches: The Role of Early Hormones and Social Environment

This experiment investigated the effects of early estrogen treatment and sex composition of the social environment on sexual partner preference in female zebra finches, a pair-bonding socially monogamous species. Birds were injected daily with estradiol benzoate (EB) or the steroid vehicle for the first 2 weeks posthatch and then lived in either a unisex (all-female) or a mixed sex group from 40 to 100 days. After 100 days birds were implanted with testosterone propionate and given three kinds of tests: tests with a stimulus female, two-choice mate preference tests with male and female stimuli, and colony tests to assess pairing preference in a more naturalistic context. Both EB and unisex housing independently resulted in a preference for females (masculinized preference) in the two-choice tests, but only females with both EB treatment and unisex living were more likely to pair with females in the colony tests. Sexual partner preference, a key sexually dimorphic component of mate choice, appears to be organized by sex steroids in this pair-bonding species, but in a manner that may be mediated by the social environment.     

Sexual Selection in the Water Spider: Female Choice and Male–Male Competition

The water spider Argyroneta aquatica is the only spider spending its whole life under water, and one of the few spider species in which males are larger than females. Previous studies indicated that males can cannibalize females, which is uncommon among spiders. Here we aimed to further test for a potential influence of sexual selection on male body size. We examined the importance of female choice by testing whether females prefer the larger of two simultaneously presented males as mating partners. Further, we examined the influence of male–male competition by comparing the fighting behaviour between large and small males when alone or when together with a female, and we determined the outcome of fights. We found that females approach and choose large males as mating partners, despite the risk of male cannibalism. Additionally, males intensively compete for females, and large males clearly win against smaller ones. Hence sexual selection seems to be important for the evolution of the peculiar sexual size dimorphism of water spiders, as large size is beneficial for males in both the intra‐ and intersexual context. Previous studies have suggested an important role of natural selection in the sex‐specific body size of water spiders, but natural and sexual selection mechanisms apparently work in the same direction, favouring large male size.     

Effect of Early Sucrose Diet on Ethanol Preference and Behavior in Male and Female Wistar Rats

The effect of adolescent sucrose diet on ethanol preference was studied in female and male Wistar rats. Our data show less pronounced ethanol preference in females. In males, pubertal exposure to sucrose was crucial for further ethanol preference, suggesting that cross-sensitization is more inherent to males than females, for whom the increased anxiety factor proved to be more significant. Finally, it was shown that in rats of both sexes habitual alcohol intake determins ethanol preference in the two-bottle test. Overall, our study demonstrates sex differences in ethanol preference as well as its anxiolytic properties.     

Geographic Variation in Male Sexual Signals in Strawberry Poison Frogs (Dendrobates pumilio)

In this paper, we compare the advertisement calls of 207 neotropical strawberry poison frogs (Dendrobates pumilio) collected in 21 localities along a transect from northern Costa Rica to western Panama. Populations varied most in call duration and call rate, while pulse rate and duty cycle were less variable. Multivariate analyses showed that call variation followed a cline with higher call rates, shorter calls, lower duty cycles and higher pulse rates in the southeast. Body size decreased towards the southeast and explained most variation in dominant frequency, as well as some residual variation in call rate. We conclude that a combination of geography and morphology is largely responsible for call variation within this species. Two inferred bio‐acoustic groups were roughly in accordance with two genetic groups, geographically separated in central Costa Rica. However, genetic distances among populations did not co‐vary with call dissimilarity after correction for geographic distances. Thus, differences in calls between genetic groups are probably mainly a result of clinal variation. These findings agree with the general observation that bio‐acoustic variation is often not (highly) associated with genetic divergence. Moreover, colour polymorphism observed among Panamanian populations was not reflected in a higher variability in call parameters relative to the monomorphic Costa Rican populations.     

Agonistic Encounters Among Male Elephant Seals: Frequency, Context, and the Role of Female Preference

SYNOPSIS. Aggressive interactions between male northern elephantseals, Mirounga angustirostris, observed during the course ofthree breeding seasons are analyzed from the perspective ofcosts and benefits to individuals. Males spend less than onepercent of their time in aggressive activities and the overwhelmingmajority of agonistic interactions consist only of visual andvocal threats rather than physical combat. Males are most likelyto threaten other males when the likelihood that they will besuccessful in displacing the opponent is high. Males who aremounting females, or who are in close proximity to females,are threatened more frequently than males who are otherwiseoccupied. However, high ranking bulls do regularly issue threatswhich are not provoked by the location or behavior of the recipient.Despite the obvious costs of such threats, this behavior maybe favored as a result of female choice. Estrous females aremore receptive when mounted by a male who has just dominatedanother than when mounted by a male of similar rank who hasbeen resting or recently displaced by a stillmore dominant male. A general argument is made that in species where social statusof males is correlated with their genetic fitness, female choiceis likely to be based on social signals which are used incompetitionbetween males.     

Female Preferences Based on Male Quality in House Mice: Interaction between Male Dominance Rank and t-Complex Genotype

Considerable evidence indicates that female house mice (Mus domesticus) prefer dominant over subordinate males as mates. In addition, male genotype at the t-complex seems to be an important characteristic used by females in mate choice. Specifically, female mice that carry a t-haplotype at the t-complex prefer +/+ over +/t males as mates. The purpose of the present study was to examine the relative contributions of male dominance rank and male t-complex genotype to female mating preference when both factors were systematically varied. We tested females of three genotypes (+/+, +/t, and t/t) in a preference apparatus using pairs of stimulus males varying in relative dominance status and t-complex genotype. In general, when given the choice, females preferred dominant over subordinate males regardless of the male's t-complex genotype. The preference for dominant males was manifested when both stimulus males were of the same t-complex genotype but differed in dominance rank. In addition, when forced to choose between a dominant +/+ and subordinate +/t or between a dominant +/t and subordinate +/+, females continued to prefer the dominant male. Preference for dominant males was independent of female genotype. Only when both males were dominant but differed in t-complex genotype (i.e. one male was +/+ and the other +/t) or when males were unranked (i.e. had not been used in aggressive encounters to determine dominance rank) did females carrying t-haplotypes manifest preferences for +/+ males. Quite unexpectedly, when both males were subordinate but differed in t-complex genotype, preferences of all females shifted in the direction of the +/t male. It is not clear from present data whether the propensity of females to give greater weight to male dominance rank than to t-complex genotype in choosing mates results in greater fitness. However, if these trends are found in natural populations, it would indicate that the role of mating preference in regulating the frequency of t-haplotypes in wild populations is less straightforward than had been previously thought.     

Sexual Traits as Quality Indicators in Lekking Male Great Bustards

We studied the development of two sexual traits, whiskers and neck plumage, in relation to sexual selection in 41 free‐living great bustard, Otis tarda, males radio‐tracked at nine leks in central Spain in 1998–2001. During the pre‐breeding male–male competition period (Feb.) prior to female arrival, number and length of whiskers correlated with weight, but not with body size or age. Whiskers may thus have evolved as an intrasexual indicator of weight, which in the absence of other weapons in this species is decisive in male–male combats. Signalling through whiskers contributes to minimizing dangerous aggressive interactions in the lek. During the mating period (Apr.), both whisker and neck development were correlated with weight and age. Males reaching higher expression of both traits exhibited higher display intensity, a more prolonged display period through the mating season, and a higher estimated mating success. Moreover, interannual changes in a male’s expression of both traits were associated with changes in its display intensity and estimated mating success. Our results resolve earlier debates and contradictory results from previous authors, suggesting that these two secondary sexual traits, whiskers and neck, may function as reliable indicators of age and weight, the two main factors determining social rank of males in great bustard leks, during both rival assessment and mate choice. Their dual functions provide support for the pre‐existing trait and redundant signal hypotheses and suggest that multiple ornaments functioning as redundant signals might be more widespread than previously acknowledged.