Sexual Swellings Research Articles

Testing for links between face color and age, dominance status, parity, weight, and intestinal nematode infection in a sample of female Japanese macaques.

Studies of the role of secondary sexual ornaments in mate choice tend to focus on colorful traits in males, but females of many animal species express colorful ornamentation too. Among non-human primates, investigations into the role of female secondary sexual traits as indicators of life history characteristics, reproductive success, and health status have mostly focused on sexual swellings, whereas only few studies have been conducted on the role of facial color. Recent studies on rhesus macaques and mandrills suggested that female ornamentation might provide information about female life history characteristics, but not on disease resistance factors and parasite infection, which have been shown to affect male ornamentation in some non-primate species. In Japanese macaques (Macaca fuscata), females have brightly colored faces that are indicative of their reproductive status. Here, we aimed to determine whether female facial color might also convey information about age, dominance rank, parity, weight, and intestinal nematode infection in free-ranging individuals. We analyzed whether female facial parameters (luminance and redness) were linked to these individual characteristics, using digital photography and data on intestinal parasite infection collected systematically during 1 month for each of seven free-ranging females. We found no evidence to suggest that female facial color is an indicator of any of these measures in Japanese macaques. Considering our small data set, it is still preliminary to draft any clear conclusions. Future studies combining digital, hormonal, parasitological and behavioral data are needed to assess the possible role of female face color on male preferences and mating choice in Japanese macaques.

Mixed messages: wild female bonobos show high variability in the timing of ovulation in relation to sexual swelling patterns.

BackgroundThe evolution of primate sexual swellings and their influence on mating strategies have captivated the interest of biologists for over a century. Across the primate order, variability in the timing of ovulation with respect to females’ sexual swelling patterns differs greatly. Since sexual swellings typically function as signals of female fecundity, the temporal relation between ovulation and sexual swellings can impact the ability of males to pinpoint ovulation and thereby affect male mating strategies. Here, we used endocrine parameters to detect ovulation and examined the temporal relation between the maximum swelling phase (MSP) and ovulation in wild female bonobos (Pan paniscus). Data were collected at the Luikotale field site, Democratic Republic of Congo, spanning 36 months. Observational data from 13 females were used to characterise female swelling cycles (N = 70). Furthermore, we measured urinary oestrone and pregnanediol using liquid chromatography–tandem mass spectrometry, and used pregnanediol to determine the timing of ovulation in 34 cycles (N = 9 females).ResultsWe found that the duration of females’ MSP was highly variable, ranging from 1 to 31 days. Timing of ovulation varied considerably in relation to the onset of the MSP, resulting in a very low day-specific probability of ovulation and fecundity across female cycles. Ovulation occurred during the MSP in only 52.9 % of the analysed swelling cycles, and females showed regular sexual swelling patterns in N = 8 swelling cycles where ovulation did not occur. These findings reveal that sexual swellings of bonobos are less reliable indicators of ovulation compared to other species of primates.ConclusionsFemale bonobos show unusual variability in the duration of the MSP and in the timing of ovulation relative to the sexual swelling signal. These data are important for understanding the evolution of sexual signals, how they influence male and female mating strategies, and how decoupling visual signals of fecundity from the periovulatory period may affect intersexual conflict. By prolonging the period during which males would need to mate guard females to ascertain paternity, the temporal variability of this signal may constrain mate-guarding efforts by male bonobos.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-016-0691-3) contains supplementary material, which is available to authorized users.

Sex differences in inhibitory control in socially-housed baboons (Papio papio)

Inhibitory control is an important component of executive function. An emerging literature in humans suggests that inhibitory control is sexually dimorphic and modulated by sex steroids, but evidence for such a link in nonhuman animals is scarce. In this study, we examined the effects of menstrual cycle and biological sex on response inhibition, as measured by a Stop-Signal task, in the baboon (Papio papio). The monkeys (n=13) were socially-housed, with voluntary access to multiple touchscreen computerized stations. The task required monkeys to inhibit prepotent responses (touching a target, “Go” trials) following the appearance of a visual stop signal on 25% of the trials (“Stop” trials). The cognitive data, consisting of computerized records of the monkeys’ performance on the Stop-Signal task over a year of testing, were matched to records of female sexual swellings. Same-day menstrual and cognitive data were available for 5 females, aged 5–18 years. These data were compared to those of 8 males (5–14 years old) performing the Stop-Signal task over the same time period. Contrary to our hypothesis, performance on the task was not significantly affected by the phase (ovulatory vs. luteal) of the cycle in females. However, males were slower than females on Go trials and were less efficient in inhibiting responses on Stop trials. Slower responses in males were indicative of a speed-accuracy trade-off, as overall accuracy was also better in males than in females. Analyses of trial history indicated that males did not speed as much as females following a successful Go trial, but did not differ from females in post-error slowing or post-inhibiting responses. Overall, the data show that biological sex modulates Stop-Signal performance in the baboon, with males exhibiting slower response execution overall, less efficient inhibition, but greater accuracy than females. This pattern of sex differences may reflect motivational sex differences in which males emphasize accuracy rather than speed. Interestingly, these sex differences do not seem to vary as a function of ovarian hormones in females. Males’ greater focus on accuracy is possibly due to enhanced sensitivity to reward mediated by testosterone levels.

Female Reproductive Seasonality and Male Influxes in Wild Mandrills (Mandrillus sphinx)

The correlates of variation in the number of males in primate groups form a long-standing question in primatology. We investigated female reproductive seasonality and the numbers of males in groups of wild mandrills (Mandrillus sphinx) in a 25-month camera-trap survey with 160 camera locations in Moukalaba-Doudou National Park, Gabon. We used 1760 videos to analyze group composition, including the presence of females with newborn infants and with sexual swellings, the number of males present in groups, and male spatial positioning in groups. Female reproduction was seasonal, with a peak in the number of newborns in the mid-rainy season and a peak in the number of females with sexual swellings in the early dry season. The number of males in the group increased in the dry season, with a much greater increase in the number of mature males (sevenfold) than in submature males (twofold). The peak number of mature males, but not submature males, in the group lagged significantly behind the peak in the number of females with sexual swellings, suggesting that submature males enter groups before mature males and/or that mature males stay in the group longer after the peak of females with sexual swellings. Mature, but not submature, males appeared frequently near females with sexual swellings. In conclusion, we found a clear relationship between the presence of receptive females and the number of males in the group, and evidence that male competitive ability influences male strategies.

Growth trajectories in wild geladas (Theropithecus gelada).

Life history and socioecological factors have been linked to species-specific patterns of growth across female vertebrates. For example, greater maternal investment in offspring has been associated with more discrete periods of growth and reproduction. However, in primates it has been difficult to test such hypotheses because very few studies have obtained growth measurements from wild populations. Here we utilize a promising noninvasive photogrammetric method-parallel lasers-to examine shoulder-rump (SR) growth in a wild primate, the gelada (Theropithecus gelada, Simien Mountains National Park, Ethiopia). In this species, a graminivorous diet coupled with high extrinsic infant mortality risk suggests that maternal investment in neonates is low. Therefore, in contrast with other closely related papionins, we expected female geladas to exhibit less discrete periods of growth and reproduction. For both sexes, we compared size-for-age patterns (N = 154 females; N = 110 males) and changes in growth velocity relative to major life history milestones. Female geladas finished 88.5% of SR growth by first sexual swelling, and 97.2% by first reproduction, reaching adult body size by 7.72 years of age. Compared to closely related papionins, gelada females finished more growth by first reproduction, despite producing relatively small, and presumably "cheap," neonates. Male geladas finished 85.4% of growth at dispersal, and 96.0% at estimated first birth. Contrary to other polygynous primates, males are larger than females because they grow for a longer period of time (not because they grow faster), surpassing females around 6 years of age when female growth slows. Our results demonstrate that parallel lasers are an easy and promising new method that can be used to construct comprehensive life history perspectives that were once out of reach for wild populations. Am. J. Primatol. 78:707-719, 2016. © 2016 Wiley Periodicals, Inc.

Exaggerated sexual swellings in female nonhuman primates are reliable signals of female fertility and body condition

In some species of Old World monkeys and apes, females exhibit exaggerated swellings of the anogenital region that vary in size across the ovarian cycle. Exaggerated swellings are typically largest around the time of ovulation, and swelling size has been reported to correlate positively with female quality, supporting the hypothesis that exaggerated swellings are honest signals of both female fecundity and quality. However, the relationship between swelling size and timing of ovulation is weak in some studies, and the relationship between swelling size and female quality has also not been consistently reported. Here, we collated empirical studies that have reported either swelling size and estimated timing of ovulation ( N = 26) or swelling size and measures of individual quality ( N = 7), to assess the strength of these relationships using meta-analytical methods. Our analyses confirmed that the period of maximal swelling size is closely associated with the most fertile period of the ovarian cycle and that a large proportion of ovulations occur during the maximal swelling period. A small, positive effect size was also found for the relationship between swelling size and body condition. In contrast, the relationships with age and social rank were not significant. Swelling size, therefore, potentially signals both female condition and timing of the fertile phase. Males are likely to benefit from allocating mating effort according to swelling size, while females with large swellings potentially benefit from exerting control over matings in species in which female control is compromised by male mating strategies.

A review on safety and efficacy of products containing Longifolia

Eurycoma longifolia (commonly called tongkat ali ) is a flowering plant in the family Simaroubaceae , native to Indonesia, Malaysia, and, to a lesser extent, Thailand, Vietnam and also Laos. The roots extract of E. longifolia , is a well-known traditional herbal medicine in Asia used for many purposes such as sexual dysfunction, aging, malaria, cancer, diabetes, anxiety, aches, constipation, exercise recovery, fever, increased energy, increased strength, leukemia, osteoporosis, stress, syphilis and glandular swelling. The roots are also used as an aphrodisiac, antibiotic, appetite stimulant and health supplement. It is very important to conserve this valuable medicinal plant for the health benefit of future generations. The purpose of this review article is to evaluate and summarize the existing literatures regarding the efficacy and safety of products which contain E. longifolia as its main ingredient. In summary, based on the literature evaluated in this review article, products which contain tongkat ali showed a clinical benefit on improving erectile dysfunction as well as a good safety profile. We recommend consumers to check the level of the bioactive compound “eurycomanone” as their guide before choosing any E. longifolia product.

Coevolution of female ovulatory signals and male–male competition in primates

Visual signals of ovulation vary among primate species. Although slight ovulatory signals are considered primate ancestral traits of which some species still exhibit, some show prominent swelling of their perineal skin (exaggerated sexual swellings) and others do not exhibit any signals of ovulation (concealed ovulation). These signals strongly affect male mating behaviors. I develop an evolutionary model of female ovulatory signals and male–male competition. I assume that each male allocates his effort between attraction of females and male–male competition for dominance. Each female gains a benefit if she is fertile and free from the alpha male who always guards one of the most fertile females in the group, but suffers a cost if she expresses a different ovulatory signal from an ancestral trait. I show that various types of ovulatory signals may evolve and can be multistable. Male–male competition becomes intense when the signal honestly indicates ovulation. Ovulatory signals may evolve to be less exaggerated in unimale groups than in multimale groups and monogamy is more likely to evolve when ovulation is concealed. These results may partly explain why various types of primate ovulatory signals evolved and how they have affected primate societies.

Changes in Women's Facial Skin Color over the Ovulatory Cycle are Not Detectable by the Human Visual System.

Human ovulation is not advertised, as it is in several primate species, by conspicuous sexual swellings. However, there is increasing evidence that the attractiveness of women’s body odor, voice, and facial appearance peak during the fertile phase of their ovulatory cycle. Cycle effects on facial attractiveness may be underpinned by changes in facial skin color, but it is not clear if skin color varies cyclically in humans or if any changes are detectable. To test these questions we photographed women daily for at least one cycle. Changes in facial skin redness and luminance were then quantified by mapping the digital images to human long, medium, and shortwave visual receptors. We find cyclic variation in skin redness, but not luminance. Redness decreases rapidly after menstrual onset, increases in the days before ovulation, and remains high through the luteal phase. However, we also show that this variation is unlikely to be detectable by the human visual system. We conclude that changes in skin color are not responsible for the effects of the ovulatory cycle on women’s attractiveness.

Exaggerated sexual swellings and male mate choice in primates: testing the reliable indicator hypothesis in the Amboseli baboons

The paradigm of competitive males vying to influence female mate choice has been repeatedly upheld, but, increasingly, studies also report competitive females and choosy males. One female trait that is commonly proposed to influence male mate choice is the exaggerated sexual swelling displayed by females of many Old World primate species. The reliable indicator hypothesis posits that females use the exaggerated swellings to compete for access to mates, and that the swellings advertise variation in female fitness. We tested the two main predictions of this hypothesis in a wild population of baboons (Papio cynocephalus). First, we examined the effect of swelling size on the probability of mate-guarding ('consortship') by the highest-ranking male and the behavior of those males that trailed consorshipts ('follower males'). Second, we asked whether a female's swelling size predicted several fitness measures. We found that high-ranking males do not prefer females with larger swellings (when controlling for cycle number and conception) and that females with larger swellings did not have higher reproductive success. Our study-the only complete test of the reliable indicator hypothesis in a primate population-rejects the idea that female baboons compete for mates by advertising heritable fitness differences. Furthermore, we found unambiguous evidence that males biased their mating decisions in favor of females who had experienced more sexual cycles since their most recent pregnancy. Thus, rather than tracking the potential differences in fitness between females, male baboons appear to track and target the potential for a given reproductive opportunity to result in fertilization.

Urinary C-peptide levels in male bonobos (Pan paniscus) are related to party size and rank but not to mate competition

Within- and between-species variation in male mating strategies has been attributed to a multitude of factors including male competitive ability and the distribution of fertile females across space and time. Differences in energy balance across and within males allow for the identification of some of the trade-offs associated with certain social and mating strategies. Bonobos live in groups with a high degree of fission–fusion dynamics, there is co-dominance between the sexes and a linear dominance hierarchy among males. Males compete over access to females, breeding is aseasonal, and females exhibit sexual swellings over extended time periods. In this study we use urinary C-peptide (UCP) levels in male bonobos (Pan paniscus) obtained from 260 urine samples from a wild bonobo community, to quantify male energy balance during mate competition and levels of gregariousness in the species. Although high ranking males are more aggressive, spend more time in proximity to maximally tumescent females, and have higher mating frequencies, we found no indication that mate guarding or mate competition affected male energy balance. Our results showed a positive correlation between monthly mean UCP levels and mean party size. When traveling in large parties, high ranking males had higher UCP levels than those of the low ranking males. These results support the hypothesis that patterns of fission–fusion dynamics in bonobos are either linked to energy availability in the environment or to the energetic costs of foraging. The finding of a rank-bias in UCP levels in larger parties could also reflect an increase in contest competition among males over access to food.

Prolonged maximal sexual swelling in wild bonobos facilitates affiliative interactions between females

Perineal sexual skin swelling in relation to menstrual cycle occurs in a variety of primate taxa. However, sexual swelling with exaggerated size and colour is found only in some Old World monkeys and the two Pan species. Although several hypotheses have been proposed (e.g., reliable indicator hypothesis and graded signal hypothesis), it seems unlikely that a single explanation can account for the significance of the sexual swelling in all of these species. Bonobos (Pan paniscus) provide an excellent opportunity for studying sexual swelling since they have the most prolonged maximal swelling periods among primates. In this study we propose a new hypothesis that sexual swelling in female bonobos increases their attractiveness to other females and thereby facilitates affiliative social interaction with them. We found that free-ranging female bonobos with maximal sexual swelling engaged in affiliative social interactions with other females, including genito-genital rubbing, staying in close proximity and grooming, more frequently than females without maximal swelling. These tendencies suggest that females with maximal swelling were attractive to other females. The results also suggest that the benefits of maximal swelling might vary among females depending on their life-history stage. In particular, young females may get more benefits from prolonged maximal swelling through increased grooming reciprocity and staying in close proximity to other females. Thus our study supported the hypothesis that one function of prolonged maximal swelling in bonobos is to increase attractiveness to other females, thereby enhancing affiliative relationships between females in a male-philopatric social system.

Mating behavior of adolescent male chimpanzees (Pan troglodytes) at Ngogo, Kibale National Park, Uganda.

Male mating tactics vary extensively in many primates. Some variation occurs because adolescent males often are sexually active but cannot invest heavily in mating effort because of their limited ability to compete directly with adults and because they are still investing in growth; consequently, most of their mating attempts may be surreptitious and/or with females whose fecundity is low. Chimpanzees (Pan troglodytes) have a complex mating system: most copulations occur between estrous females with full sexual swelling and multiple males in group settings where the potential for sperm competition is high, but males sometimes mate-guard females, and sometimes male-female pairs mate exclusively with each other while avoiding other males during "consortships." Among other factors, dominance ranks, coalition formation, and variation in male-female association influence male mating and reproductive success. Mating effort increases from adolescence into prime adulthood. At Gombe and Mahale, adolescent males copulated more with nulliparous than with parous females, and mostly when females were unlikely to be ovulating, partly because of low adult male interest in nulliparous females and partly because of aggression from or avoidance of adult males. Adolescents thus had low probabilities of siring infants. However, adolescents are known to have gained some paternity at Gombe and in other populations, and their mating behavior deserves more study. I present data on mating by adolescent males in an unusually large chimpanzee community at Ngogo, Kibale National Park, Uganda. Adolescents at Ngogo also copulated more with nulliparous than parous females and mostly copulated outside of periovulatory periods. Also, they directed less aggression at estrous females than did adult males. However, they gained lower shares of copulations than reported for Gombe and Mahale, regardless of female parity, and received more aggression from adult males. These differences might partly reflect the influence of variation in the number of males per community on male mating tactics.

Multimodal signalling of ovulation in human and non-human primates

Communication in primates is based on multiple elements of signals, often expressed in different modalities. While some individual primate signals are relatively well described for a particular cue (e.g. copulation calls, sexual swellings) or for one specific dimension of a signal (e.g. the size of sexual swellings), few studies have simultaneously investigated the role of multiple signals and cues in the same communicative context. This paper reviews comparative evidence on primate sexual communication, specifically concerning multimodal sexual signalling of ovulation and mating strategies. A comparison is made across primate species, including humans, of the characteristics of different female sexual cues and signals (behavioural, visual, auditory and olfactory) and of their potential roles in the discrimination of the timing of ovulation. The possible selective reasons for such multimodal signalling are discussed.

Influence of personality, age, sex, and estrous state on chimpanzee problem-solving success

Despite the importance of individual problem solvers for group- and individual-level fitness, the correlates of individual problem-solving success are still an open topic of investigation. In addition to demographic factors, such as age or sex, certain personality dimensions have also been revealed as reliable correlates of problem-solving by animals. Such correlates, however, have been little-studied in chimpanzees. To empirically test the influence of age, sex, estrous state, and different personality factors on chimpanzee problem-solving, we individually tested 36 captive chimpanzees with two novel foraging puzzles. We included both female (N=24) and male (N=12) adult chimpanzees (aged 14-47 years) in our sample. We also controlled for the females' estrous state-a potential influence on cognitive reasoning-by testing cycling females both when their sexual swelling was maximally tumescent (associated with the luteinizing hormone surge of a female's estrous cycle) and again when it was detumescent. Although we found no correlation between the chimpanzees' success with either puzzle and their age or sex, the chimpanzees' personality ratings did correlate with responses to the novel foraging puzzles. Specifically, male chimpanzees that were rated highly on the factors Methodical, Openness (to experience), and Dominance spent longer interacting with the puzzles. There was also a positive relationship between the latency of females to begin interacting with the two tasks and their rating on the factor Reactivity/Undependability. No other significant correlations were found, but we report tentative evidence for increased problem-solving success by the females when they had detumescent estrous swellings.

Correlations between Social Context and Fecal Glucocorticoid Metabolite Concentrations in Free-ranging Female Gray-cheeked Mangabeys (Lophocebus albigena) in Kibale National Park, Uganda

An elevated concentration of glucocorticoids is an indicator of stress, and chronically high glucocorticoid levels are often associated with poor health and reduced fertility. We explored conditions that might be stressors in the lives of adult female gray-cheeked mangabeys (Lophocebus albigena) by measuring fecal glucocorticoid metabolites (fGCM) concentrations. During a six-month study we collected 109 fecal samples from 28 adult females from five groups in Kibale National Park, Uganda. We examined fecal fGCM levels of individual females relative to their own reproductive status (cycling or not cycling) and that of other females and to the presence of newcomer (immigrant) males. We found elevated fGCM concentrations in females when other females in their groups were at the peak of sexual swelling, and when immigrant males joined the group.

A New Method for the Analysis of Soft Tissues with Data Acquired under Field Conditions

Analyzing soft-tissue structures is particularly challenging due to the lack of homologous landmarks that can be reliably identified across time and specimens. This is particularly true when data are to be collected under field conditions. Here, we present a method that combines photogrammetric techniques and geometric morphometrics methods (GMM) to quantify soft tissues for their subsequent volumetric analysis. We combine previously developed methods for landmark data acquisition and processing with a custom program for volumetric computations. Photogrammetric methods are a particularly powerful tool for field studies as they allow for image acquisition with minimal equipment requirements and for the acquisition of the spatial coordinates of points (anatomical landmarks or others) from these images. For our method, a limited number of homologous landmarks, i.e., points that can be found on any specimen independent of space and time, and further distinctive points, which may vary over time, space and subject, are identified on two-dimensional photographs and their three-dimensional coordinates estimated using photogrammetric methods. The three-dimensional configurations are oriented by the spatial principal components (PCs) of the homologous points. Crucially, this last step orients the configuration such that x and y-information (PC1 and PC2 coordinates) constitute an anatomically-defined plane with the z-values (PC3 coordinate) in the direction of interest for volume computation. The z-coordinates are then used to estimate the volume of the tissue. We validate our method using a physical, geometric model of known dimensions and physical (wax) models designed to approximate perineal swellings in female macaques. To demonstrate the usefulness and potential of our method, we use it to estimate the volumes of Barbary macaque sexual swellings recorded in the field with video images. By analyzing both the artificial data and real monkey swellings, we validate our method's accuracy and illustrate its potential for application in important areas of biological research.

Multimodal Sexual Signaling and Mating Behavior in Olive Baboons (Papio anubis)

In primate species, mating decisions seem to be based on multiple signal elements with different roles in the signaling of female reproductive status. Whereas some primate signals are relatively well described (e.g., sexual swellings and copulation calls), studies that simultaneously assess visual, auditory, behavioral, and olfactory cues as signals of reproductive state are rarely undertaken. We used data on variation in sexual behaviors and sexual swellings in relation to the fertile period (estimated from the date of swelling detumescence) from a troop of semi-free ranging olive baboons (Papio anubis) to assess how different signals influence patterns of mate choice. Using an objective and quantitative measure of swelling size and color, along with detailed data on sexual behaviors from 13 cycles of nine adult females, we found that fine-scale variation in sexual swelling size, female behavior and copulation call rates could advertise the beginning of the fertile phase whereas swelling color did not. Rates of olfactory inspections by males also increased during the fertile phase, suggesting that olfactory signals were of interest to males and may contain information about ovulation. There was no relationship between female characteristics (age and rank) and the expression of sexual signals, except for proceptive behaviors which increased with female rank. Males displayed more sexual behaviors such as approaches and holding and tended to direct more ejaculatory mounts during the fertile phase. All together, we suggest that whereas all males could have information concerning the timing of ovulation through female proceptive behaviors and swelling size, consorting males may have access to additional signals (olfactory cues). Sexual communication in olive baboons is consistent with a multimodal framework for fertility signaling, potentially allowing males and females to establish different mating strategies. The possible selective pressures leading to multi-modal signaling are discussed.

Male mating behaviour in relation to female sexual swellings, socio-sexual behaviour and hormonal changes in wild Barbary macaques

In many cercopithecine primates females display probabilistic cues of fertility to indicate the periovulatory period to males. These cues may include female behaviour, acoustic signals, and morphological signs such as the anogenital swelling. However, the extent to which males can utilise this information varies between species. We describe male sexual behaviour in relation to changes in anogenital swelling size, timing of ovulation and female socio-sexual behaviour in wild Barbary macaques (Macaca sylvanus). We further compare male sexual behaviour during conception and post-conception cycles to evaluate if males differentiate between these qualitatively different cycle types. Our results show that during conception cycles male mating behaviour was concentrated around the fertile phase implying that males inferred information from more than swelling size alone. Male mating frequency increased in line with female socio-sexual behaviour, namely female presenting and the frequency of copulations with copulation calls. Most strikingly our results show that males invested equally in mating during fertile and non-fertile, i.e. post-conception, maximum swelling phases. Whether post-conception swellings were merely a result of changes in hormone concentrations during pregnancy or part of a female reproductive strategy remains elusive. In sum, this study adds to the body of research on the evolution of female sexual signals and how males may infer information from these cues.

Paternal care and the evolution of exaggerated sexual swellings in primates.

The exaggerated sexual swellings exhibited by females of some primate species have been of interest to evolutionary biologists since the time of Darwin. We summarize existing hypotheses for their function and evolution and categorize these hypotheses within the context of 3 types of variation in sexual swelling size: 1) variation within a single sexual cycle, 2) variation between the sexual cycles of a single female, and 3) differences between females. We then propose the Paternal Care Hypothesis for the function of sexual swellings, which posits that exaggerated sexual swellings function to elicit the right quantity and quality of male care for a female's infant. As others have noted, swellings may allow females to engender paternity confusion, or they may allow females to confer relative paternal certainty on one male. Key to our hypothesis is that both of these scenarios create an incentive for one or more males to provide care. This hypothesis builds on previous hypotheses but differs from them by highlighting the elicitation of paternal care as a key function of swellings. Our hypothesis predicts that true paternal care (in which males accurately differentiate and provide assistance to their own offspring) will be most common in species in which exaggerated swellings accurately signal the probability of conception, and males can monopolize females during the window of highest conception probability. Our hypothesis also predicts that females will experience selection to behave in ways that either augment paternity confusion or enhance paternal certainty depending on their social and demographic contexts.