The multiple signals assessed by female satin bowerbirds: could they be used to narrow down females' choices of mates?

Female choice based on multiple male traits has been documented in many species but the functions of such multiple traits are still under debate. The satin bowerbird has a polygynous mating system in which males attract females to bowers for mating; females choose mates based on multiple aspects of males and their bowers. In this paper, we demonstrate that females use some cues to decide which males to examine closely and other cues to decide which males to mate with. Female visitation rates to bowers were significantly related to male size and the males' 'solitary' display rates, and, to a lesser extent, to the numbers of bower decorations. After controlling for female visitation rates, it was found that a male's mating success was significantly related to his size and the rate at which he 'painted' his bower with saliva and chewed up plant material.     

Male southern water skinks (Eulamprus heatwolei) use both visual and chemical cues to detect female sexual receptivity

A wide range of organisms use chemical and visual cues in mate attraction and courtship; however, chemical discrimination relevant to reproduction and the interplay between these two types of communication are poorly understood in reptiles. We experimentally tested the ability of male Eulamprus heatwolei, a scincoid lizard, to discriminate between sexually receptive and non-receptive females in two ways. First, we conducted 155 staged encounters between males and females over 29 days to determine the start and the duration of the female receptive period based on the date of copulations. These data suggest that the receptive period lasted for approximately 7 days in late October under controlled laboratory conditions. We also recorded 6,330 individual male and female behaviours during these trials to evaluate the frequency of female courtship and rejection behaviours and the intensity of male courtship behaviour. Female courtship increased sharply during the receptive period and then diminished. The disparity between female courtship behaviours and female rejection behaviours was greatest during the receptive period. Male courtship intensity increased sharply prior to the receptive period, peaked during the receptive period and thereafter declined rapidly. To determine if males were using visual cues, chemical cues or both from females, we conducted an experiment during and after the receptive period wherein male lizards were presented with a choice of two retreat sites treated either with the odour of large sexually receptive females, odour of small sexually non-receptive females or no odour (control). Males preferred the scent of females over the odourless control, and analysis using a special form of a generalized linear model, the Bradley–Terry model, showed a clear order of retreat site preferences, with large sexually receptive females favoured over small non-receptive females over the odourless control. We speculate that males use vision to find females and then use their chemosensory ability to chemically evaluate female sexual receptivity once the pair are in close proximity.     

Mate preference for multiple cues: interplay between male and nest size in the sand goby, Pomatoschistus minutus

Female mating preferences are often based on more than one cue.In empirical studies, however, different mate choice cues aretypically treated separately ignoring their possible interactions.In the current work, we studied how male body size and sizeof the male's nest jointly affect mate preferences of femalesand gobies, Pomatoschistus minutus. The females were givena binary choice between males that differed either in body sizeor size of their nest or both. We found that neither body sizenor size of the nest alone affected male attractiveness, buttogether these 2 cues had a significant effect. Specifically,large males were more popular among females when they had alarge nest than when they occupied a small nest. The resultssuggest that if interaction effects between multiple mate choicecues are not considered, there is a danger of ignoring or underestimatingthe importance of these cues in sexual selection by female choice.     

Sex Recognition via Chemical Cues in the Sex-Role-Reversed Gulf Pipefish (Syngnathus scovelli)

Sexual selection theory predicts that members of the choosy sex, usually females, should employ multiple sensory systems to obtain information about potential mates. Such predictions should also apply to systems in which sexual selection acts most strongly on females (i.e. sex-role-reversed species), and males of these taxa should be the individuals employing multiple cues to assess female attractiveness. Very little work has been directed toward mate choice involving multiple sensory systems in sex-role-reversed taxa, but fishes of the family Syngnathidae (pipefishes, seahorses, and sea dragons) provide an excellent opportunity to contribute to this research enterprise. While much is known about visual communication in pipefish, the role of chemical communication has not been investigated. Using dichotomous choice tests, we found that male, but not female, Gulf pipefish attended to chemical cues of opposite sex conspecifics. Given that males distinguished sex on the basis of chemical cues, we also tested whether males could assess female body size, an important trait with respect to mate choice in pipefish, on the basis of chemical cues alone. When given the choice between chemical cues produced by large vs. small females, males exhibited no preference. Our results suggest that male pipefish can use chemical cues to distinguish between males and females but not to differentiate females of different body size.     

Environmental change mediates mate choice for an extended phenotype,but not for mate quality

Sexual cues, including extended phenotypes, are expected to be reliable indicators of male genetic quality and/or provide information on parental quality. However, the reliability of these cues may be dependent on stability of the environment, with heterogeneity affecting how selection acts on such traits. Here, we test how environmental change mediates mate choice for multiple sexual traits, including an extended phenotype–‐the structure of male‐built nests – in stickleback fish. First, we manipulated the dissolved oxygen (DO) content of water to create high or low DO environments in which male fish built nests. Then we recorded the mate choice of females encountering these males (and their nests), under either the same or reversed DO conditions. Males in high DO environments built more compact nests than those in low DO conditions and males adjusted their nest structure in response to changing conditions. Female mate choice for extended phenotype (male nests) was environmentally dependent (females chose more compact nests in high DO conditions), while female choice for male phenotype was not (females chose large, vigorous males regardless of DO level). Examining mate choice in this dynamic context suggests that females evaluate the reliability of multiple sexual cues, taking into account environmental heterogeneity.     

Scent marking and mate choice in the prairie vole, Microtus ochrogaster

Scent marking is common among male and female rodents and might be used in male-male competition and as a mechanism for mate attraction. I tested the hypotheses that females would choose males based on their frequency and placement of scent marks, and that a female would advertise interest in a particular male by placing her scent marks on or near those of a preferred mating partner. In a series of experiments conducted with prairie voles, Microtus ochrogaster, females did not choose mates based on the frequency or placement of scent marks by males nor did they advertise their interest in a particular male through the frequency or placement of scent marks. The number of males chosen that scent-marked more than their opponents did not differ significantly between females exposed (11 of 15) and not exposed (10 of 15) to scents of males. Females exposed and not exposed to scents of males preferred seven of the same males that had scent-marked more than their opponents. When a third group of females was exposed to four times more scent of the less preferred than preferred males, they still chose the preferred males. Thus, the frequency and placement of scent marks by males were not used to assess males for mate choice nor did female prairie voles use scent to advertise their preference for a mating partner. In that scent marking is common in male and female mammals, scent quality might be more important than quantity in male-male competition and mate attraction.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

The scent of your enemy is my friend? The acquisition of large carnivore scent by a smaller carnivore

Scent marking is critical to intraspecific communication in many mammal species, but little is known regarding its role in communication among different species. We used 4 years of motion-triggered video to document the use of scent marking areas—termed “community scrapes”—by pumas (Puma concolor) (http://www.momo-p.com/showdetail-e.php?movieid=momo160812pc01a) and other carnivore species. We found that gray foxes (Urocyon cinereoargenteus) routinely rubbed their cheeks on puma scrapes (http://www.momo-p.com/showdetail-e.php?movieid=momo160812uc01a), and tested a series of hypotheses to determine its function. We found that gray foxes selected puma scrapes over other objects, and cheek rubbing by foxes was also correlated with how recently a puma had visited the scrape, suggesting that foxes were intent upon accumulating fresh puma scent. Cheek rubbing by foxes was not correlated with their breeding season or with how recently another fox had visited the site. Finally we found a cascading pattern in the occurrence of pumas, coyotes (Canis latrans) and gray foxes at community scrapes, suggesting that gray foxes may use puma scent to deter predation. This is the first published study to find evidence of a subordinate species using the scent of a dominant species to communicate with heterospecifics. The behavioral cascade we found in scent marking patterns also suggests that scent marking could be a mechanism that impacts the distribution and abundance of species. Additional videos pertaining to this article include http://www.momo-p.com/showdetail-e.php?movieid=momo160812uc02a, and http://www.momo-p.com/showdetail-e.php?movieid=momo160812uc03a.     

The Effect of Variation in Male Sexually Dimorphic Traits on Female Behaviour in Pronghorn (Antilocapra americana)

Several sexually dimorphic morphological, behavioural, and territory characteristics of male pronghorn, Antilocapra americana, are highly variable. To test the hypothesis that females respond to variation among males for mate choice, I examined the relationship between these male characteristics and female congregation size and variability. Increased movement up to and during rut and differential female congregation on the territories of males with favoured characteristics were detected as predicted. Individual females increased their home ranges nearly fourfold from summer to rut and visited 88% of the territories in the study area. Female congregation size was related to male phenotype but not to territory characteristics. A canonical correlation analysis revealed that not only were several male traits simultaneously related to female congregation size and variability, but the importance of these traits varied across years. However, male courtship, cheek-patch size, and scent marking behaviours were consistently correlated to female congregation patterns across years. Female congregation was significantly related to male courtship, but there was no correlation between former territory owner's and newcomer's attraction of females. This finding is consistent with the hypothesis that females were primarily assessing males instead of territories.     

Chemoreception, symmetry and mate choice in lizards

Research on fluctuating asymmetry (FA)-mediated sexual selection has focused almost exclusively on visual signals and ignored chemical communication despite the fact that many species rely on chemical signals for attracting mates. Female mate choice based on visual traits appears to be rare in lizards. However, the femoral glands of male lizards produce pheromones which might transmit chemical information about an individual's developmental stability. Therefore, we hypothesized that mate choice may be based on chemical cues. We analysed the effect of the developmental stability levels of males on the attractiveness of males' scents to females in a laboratory experiment with the lizard Lacerta monticola. When we offered two males of similar body size, females preferentially associated with the scents of males with low FA in their femoral pores and also with the scents of males with a higher number of femoral pores. This suggested that the females were able to discriminate the FA of the males by chemical signals alone and that the females preferred to be in areas marked by males of high quality, thus increasing their opportunities of mating with males of high quality. We suggest that the quality and/or amount of male pheromones could communicate the heritable genetic quality of a male to the female and thereby serve as the basis for adaptive female choice in lizards.     

Reproductive strategies in snakes

Snakes of both sexes display remarkable flexibility and diversity in their reproductive tactics. Many features of reproduction in female snakes (such as reproductive mode and frequency, seasonality and multiple mating) allow flexible maternal control. For example, females can manipulate not only the genotypes of their offspring (through mate choice or enhanced sperm competition) but also the phenotypes of their offspring (through allocation 'decisions', behavioural and physiological thermoregulation, and nest-site selection). Reliance on stored energy ('capital') to fuel breeding results in low frequencies of female reproduction and, in extreme cases, semelparity. A sophisticated vomeronasal system not only allows male snakes to locate reproductive females by following scent trails, but also facilitates pheromonally mediated mate choice by males. Male-male rivalry takes diverse forms, including female mimicry and mate guarding; combat bouts impose strong selection for large body size in males of some species. Intraspecific (geographical) variation and phenotypic plasticity in a wide array of reproductive traits (offspring size and number; reproductive frequency; incidence of multiple mating; male tactics such as mate guarding and combat; mate choice criteria) provide exceptional opportunities for future studies.     

Honest olfactory ornamentation in a female‐dominant primate

Sexual selection theory predicts that potential mates or competitors signal their quality to conspecifics. Whereas evidence of honest visual or vocal signals in males abounds, evidence of honest signalling via scent or by females is scarce. We previously showed that scent marks in male lemurs seasonally encode information about individual heterozygosity – a reliable predictor of immunocompetence and survivorship. As female lemurs dominate males, compete over resources, and produce sexually differentiated scent marks that likely evolved via direct selection, here we tested whether females also advertise genetic quality via olfactory cues. During the breeding season specifically, individual heterozygosity correlated negatively with the diversity of fatty acids (FAs) expressed in labial secretions and positively with the diversity of heavy FA esters. As odour–gene relationships predictive of health and survivorship emerged during a period critical to mate choice and female competition, we posit that genital scent marks function as honest olfactory ornaments in females.     

Female choice for heterozygous mates changes along successive matings in a lizard

Female mate choice and female multiple mating are major focuses of studies on sexual selection. In a multiple mating context, the benefits of mate choice can change along successive matings, and female choice would be expected to change accordingly. We investigated sequential female mate choice in the moderately polyandrous common lizard (Zootoca vivipara, synonym Lacerta vivipara). Along successive mating opportunities, we found that females were relatively unselective for the first mate, but accepted males of higher heterozygosity for subsequent mating, consistent with the trade-up choice hypothesis. We discuss the evidence of trade-up mate choice in squamates and generally trade-up for mate heterozygosity in order to motivate new studies to fill gaps on these questions.     

The relative importance of orange spot coloration and total length of males in female guppy mate preference

Female mate choice by multiple male traits is an important current topic in animal behavior. However, the relative importance among the multiple cues in female choice is not explored in most cases. Female guppies Poecilia reticulata use both the color saturation of orange spots and the total length of males as mate choice criteria. In the present study, we used digitally modified video playbacks to examine the relative importance of these two male traits to female mate preferences. We initially examined the effective difference in the color saturation of orange spots as well as that in total length between two stimulus male images. Females only showed a strong preference for a bright male image (compared to the dull image) when the difference in color saturation was large (91% versus 25%). Conversely, females only exhibited a preference for larger size when they were presented a choice between two relatively small male images (total length 26.0 mm versus 23.0 mm). When two male images in which both the two traits were modified were presented to females, they prioritized male images possessing higher color saturation of orange spots, indicating the color saturation of male orange spots to be a more important factor than the total length in their mate choice. The color saturation of orange spots may convey more reliable information about the males to the females than their total lengths. These findings imply that females may rank multiple male criteria depending on relative benefits or costs derived from their mate choice based on each criterion.     

The role of model female quality in the mate choice copying behaviour of sailfin mollies

Female mate choice copying is a socially mediated mate choice behaviour, in which a male's attractiveness to females increases if he was previously chosen by another female as a mate. Although copying has been demonstrated in numerous species, little is known about the specific benefits it confers to copying females. Here we demonstrate that the mate choice behaviour of female sailfin mollies (Poecilia latipinna) is influenced by the phenotypic quality of model females with whom males are observed consorting. Test females choosing between two males of similar body length were found to significantly increase time spent with previously non-preferred males after having observed them with a relatively high-quality female. Conversely, females were found to significantly decrease time spent with previously preferred males after having observed them with a relatively low-quality female. Female mate choice copying might be maintained by selection based on the heuristic value it provides females choosing between males whose quality differences are not easily distinguishable.     

On the function of female ornaments: male bluethroats prefer colourful females

Female ornaments in animals with conventional sex roles have traditionally been considered non-functional, being merely a genetically correlated response to selection for male ornamentation. Alternatively, female ornaments may be influenced by selection acting directly on the females, either through female–female competition or male choice. We tested the latter hypothesis in mate choice experiments with bluethroats (Luscinia s. svecica), a passerine bird in which females vary considerably in coloration of an ornamental throat patch. In outdoor aviaries placed in prime breeding habitat, males were allowed to choose between a colourful and a drab female. We found that males associated more with, and performed more sexual behaviours towards, colourful females. Female coloration was not age-related, but correlated significantly with body mass and tarsus length. Thus, we have demonstrated both a male preference for female ornamentation, and a relationship between ornament expression and female body size, which may be indicative of quality. Our results refute the correlated response hypothesis and support the hypothesis that female ornamentation is sexually selected.     

Indications for female mate choice in grey-cheeked mangabeys <Emphasis Type="Italic">Lophocebus albigena johnstoni</Emphasis> in Kibale National Park,Uganda

Female sexual strategies affect male strategies and can play an important role in shaping mating systems. We investigated female sexual behaviour within five groups of grey-cheeked mangabeys in Kibale National Park, Uganda, and tested the hypothesis that females exhibit mate choice using as indications the prevalence of (1) females soliciting matings by presenting to males and (2) females refusing to mate with approaching males. In addition, we describe how these behaviours as well as grooming and copulation calls are distributed over high-ranking, low-ranking and migrating males and discuss these patterns with regard to trade-offs that could play a roll in female mate choice in multi-male groups. Females were promiscuous and initiated almost half of the matings, with both resident and migrating males. More than half of male mating approaches were refused by peak females. Female mate choice in this species may depend on individual female preferences, oestrus phase and male tactic.     

Tactical release of a sexually-selected pheromone in a swordtail fish

Background

Chemical communication plays a critical role in sexual selection and speciation in fishes; however, it is generally assumed that most fish pheromones are passively released since most fishes lack specialized scent glands or scent-marking behavior. Swordtails (genus Xiphophorus) are widely used in studies of female mate choice, and female response to male chemical cues is important to sexual selection, reproductive isolation, and hybridization. However, it is unclear whether females are attending to passively produced cues, or to pheromones produced in the context of communication.

Methodology/Principal Findings

We used fluorescein dye injections to visualize pulsed urine release in male sheepshead swordtails, Xiphophorus birchmanni. Simultaneous-choice assays of mating preference showed that females attend to species- and sex-specific chemical cues emitted in male urine. Males urinated more frequently in the presence and proximity of an audience (conspecific females). In the wild, males preferentially courted upstream of females, facilitating transmission of pheromone cues.

Conclusions/Significance

Males in a teleost fish have evolved sophisticated temporal and spatial control of pheromone release, comparable to that found in terrestrial animals. Pheromones are released specifically in a communicative context, and the timing and positioning of release favors efficient signal transmission.     

Chemical communication of reproductive status in female cotton-top tamarins (Saguinus oedipusoedipus)

Among Callitrichids, scent secretions have been identified as carrying information regarding species, subspecies, gender, social status, individuality, hormonal status, and timing of ovulation. We propose that information regarding familiarity and reproductive status is also communicated. Seven male–female pairs of cotton-top tamarins (Saguinus oedipus oedipus) were presented with several drops of distilled water, a scent secretion from the female of that pair, scent secretions from unfamiliar, reproductively mature but noncycling females, and scent secretions from unfamiliar, reproductively cycling females. Behavioral responses from both males and females were recorded over a 10 min period. Differences in behavioral response for both males and females were significant across all four conditions. This indicates that cotton-top tamarins are capable of discriminating a familiar scent from an unfamiliar scent as well as the reproductive status of an unfamiliar female. The communication of this information may play an important role in regulating mate selection, reducing reproductive competition, and stabilizing transfers in and out of groups in the wild. Am. J. Primatol. 45:337–349, 1998. © 1998 Wiley-Liss, Inc.     

A comparison of scent marking between a monogamous and promiscuous species of peromyscus: pair bonded males do not advertise to novel females

Scent marking can provide behavioral and physiological information including territory ownership and mate advertisement. It is unknown how mating status and pair cohabitation influence marking by males from different social systems. We compared the highly territorial and monogamous California mouse (Peromyscus californicus) to the less territorial and promiscuous white-footed mouse (P. leucopus). Single and mated males of both species were assigned to one of the following arenas lined with filter paper: control (unscented arena), male scented (previously scent-marked by a male conspecific), or females present (containing females in small cages). As expected, the territorial P. californicus scent marked and overmarked an unfamiliar male conspecific's scent marks more frequently than P. leucopus. Species differences in responses to novel females were also found based on mating status. The presence of unfamiliar females failed to induce changes in scent marking in pair bonded P. californicus even though virgin males increased marking behavior. Pair bonding appears to reduce male advertisement for novel females. This is in contrast to P. leucopus males that continue to advertise regardless of mating status. Our data suggest that communication through scent-marking can diverge significantly between species based on mating system and that there are physiological mechanisms that can inhibit responsiveness of males to female cues.     

Mutual Mate Assessment in Wolf Spiders: Differences in the Cues Used by Males and Females

When males engage in conspicuous courtship displays, it seems obvious that females would use characteristics of that display in mating decisions. However, males must also have a way to identify and evaluate females prior to engaging in what might be a costly mating ritual. Although it was known that female wolf spiders of the species Pardosa milvina (Araneae; Lycosidae) attract males using volatile chemical cues, the nature of the cues used by males and females in mate selection had not been investigated. Specifically we determined whether males could detect the mating status of the female and if chemotactile cues from the female played a role in that process. In addition, we quantified conspicuous aspects of the male courtship (leg raises and body shakes) to determine if courtship intensity was related to female choice. Although repeated mating occurred in our studies, males were more likely to court and mate with virgin females. Males used substrate‐borne cues deposited by females to discriminate between mated and virgin females. Females used the conspicuous behaviors of males during courtship, body shakes and leg raises, in mate selection. Thus males and females use different kinds of information and different sensory modalities to assess the suitability of a potential mate.