Dangerous liaisons: the predation risks of receiving social signals

Individuals are at risk when communicating because conspicuous signals attract both conspecifics and eavesdropping predators. This predation cost of communicating has typically been attributed to signalling individuals because of their conspicuous role, and is a core concept within sexual selection and communication ecology. But, if predators are attracted to signals, then receivers, both intended or otherwise, may also find themselves at risk of predation. Here, we review the theoretical basis and empirical evidence that receiving also carries a risk of predation. We distinguish between the risks of receiving and responding to signals, and we argue that receivers of signals that are long lived, are highly predictable in time or place and/or cannot be received quickly are likely to be at greater risk of predation compared to receivers of signals without these properties. We review recent empirical evidence from a variety of taxa that supports the hypothesis that receivers (including heterospecific prey) are aware of these risks and that they modify their behaviour to balance the risks against the benefits of receiving under predation threat. We also discuss the wider implications of risky receiving for receiving and signalling behaviour in prey, as well as for the prey's predators.     

Baboon sexual swellings: information content of size and color

Primate sexual swellings are hormone-dependent sexual signals that play a key role in determining patterns of behavior. They are among the most conspicuous signals exhibited by any mammal, and their large size and bright coloration have fascinated evolutionary biologists for well over a century. A number of different adaptive hypotheses have been proposed for the evolution of sexual swellings, and there have been several recent attempts to test some of these using precise swelling measurements made in the field. Most of these studies have focused only on the size element of the swelling, and those that have measured other aspects of swellings, such as color, have done so only crudely. A focus solely on swelling size is inconsistent with most theoretical models of mate choice, which emphasize the importance of multiple cues within sexual signals. Here, we present data on baboon (Papio hamadryas anubis) sexual swellings, including measures of both swelling size and color, measured objectively using digital photography at Gashaka-Gumti National Park, Nigeria. We combined these measurements with detailed data on fecal progestogen and estrogen levels, and estimates of the timing of ovulation and the fertile period around ovulation based on those levels. We show that swelling color and size vary independently, and that, consistent with results in other species, swelling size contains information about the timing of ovulation and the fertile period. However, we show that swelling color does not contain such information. In addition, swelling size contains information about female parity, and we found some evidence to suggest that color may also contain such information. These results indicate that baboon sexual swellings may contain information about multiple aspects of female fertility. We discuss the implications of these results for understanding the nature of swellings as behavioral signals, and the role of swellings in mate choice.     

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

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

Sexual signalling in female crested macaques and the evolution of primate fertility signals

ABSTRACT: BACKGROUND: Female signals of fertility have evolved in diverse taxa. Among the most interesting study systems are those of multimale multifemale group-living primates, where females signal fertility to males through multiple signals, and in which there is substantial inter-specific variation in the composition and reliability of such signals. Among the macaques, some species display reliable behavioural and/or anogenital signals while others do not. One cause of this variation may be differences in male competitive regimes: some species show marked sexual dimorphism and reproductive skew, with males fighting for dominance, while others show low dimorphism and skew, with males queuing for dominance. As such, there is variation in the extent to which rank is a reliable proxy for male competitiveness, which may affect the extent to which it is in females' interest to signal ovulation reliably. However, data on ovulatory signals are absent from species at one end of the macaque continuum, where selection has led to high sexual dimorphism and male reproductive skew. Here we present data from 31 cycles of 19 wild female crested macaques, a highly sexually dimorphic species with strong mating skew. We collected measures of ovarian hormone data from faeces, sexual swelling size from digital images, and male and female behaviour. RESULTS: We show that both sexual swelling size and female proceptivity are graded-signals, but relatively reliable indicators of ovulation, with swelling size largest and female proceptive behaviours most frequent around ovulation. Sexual swelling size was also larger in conceptive cycles. Male mating behaviour was well timed to female ovulation, suggesting that males had accurate information about this. CONCLUSION: Though probabilistic, crested macaque ovulatory signals are relatively reliable. We argue that in species where males fight over dominance, male dominance rank is surrogate for competitiveness. Under these circumstances it is in the interest of females to increase paternity concentration and assurance in dominants beyond levels seen in species where such competition is less marked. As such, we suggest that it may be variation in male competitive regimes that leads to the evolution of fertility signalling systems of different reliability.     

Why some rails have white tails: the evolution of white undertail plumage and anti-predator signaling

Conspicuous plumage patches have evolved in birds as conspecific signals for mate attraction and assessment, intersexual competition or to signal alarm. Signals may alternatively be directed at potential predators to discourage pursuit. Rails (Family Rallidae) are ground-dwelling birds, many of which inhabit wetlands, while others occur in forests and grasslands. They are renown for their secretive nature and the tendency to flick their tails when observed. This behavior is more conspicuous in species with white undertail coverts that contrast sharply with darker body plumage. Using species comparisons and controlling for phylogeny, we investigated four hypotheses for the evolution of white undertail coverts in rails. We found little support for the hypothesis that white tails are sexually selected: white tails were not more common in species with polygamous as opposed to monogamous mating systems, species with sexual dimorphism, nor species that display their tails in courtship. Nor did our results support the hypothesis that white tail plumage evolved for intersexual competition during territorial interactions. Instead, we found that species that flock for at least part of the year and species found in open as opposed to concealing habitats were significantly more likely to have white undertail coverts. Rail species inhabiting concealing habitats are less commonly gregarious and more likely selected for crypsis. Using phylogenetically-controlled statistical inference we found that adaptation to open wetland habitats significantly precedes the evolution of white undertails, whereas gregariousness likely evolved later in some lineages. The inferred order of trait evolution suggests that this plumage characteristic could have been selected primarily for enhancement of an anti-predator signal rather than a social signal for conspecifics.     

Coevolution of senders and receivers of sexual signals: Genetic coupling and genetic correlations

The parallel evolution of senders and receivers of sexual signals has been a topic of research in both neuroethology and evolutionary quantitative genetics. Neuroethologists have debated whether the same physiological mechanism underlies both production and reception of a signal, and whether the same genes affect the physiology of communication in each sex. Quantitative geneticists have discussed the possibility that particular types of signals, and preferences for those types, are inherited together. Studies of communication by a variety of insect species do not provide strong support for a common physiological mechanism, but do not rule out the genetic effect. The neuroethological perspective may be of assistance in understanding the evolution of sexual communication because it offers a way to subdivide communication into units for genetic analysis.     

Exposure to sexual signals during rearing increases immune defence in adult field crickets

Increased investment in immunity is expected to be beneficial under crowded conditions because of the greater risk of pathogen and parasite transmission, but the evolution of this facultative response relies on the ability to accurately assess social cues in the environment and adjust immune defences accordingly. Because of their highly conspicuous nature, long-range sexual signals are prime candidates to be used in evaluating the social conditions likely to be experienced upon adulthood in continuously breeding species; however, their role in mediating immune responses is unknown. We tested whether exposure to acoustic sexual signals in the field cricket Teleogryllus oceanicus affects immunity by manipulating male juvenile experience of acoustic signals, and measuring the effect on adult immunity. Adult males exposed to song during rearing showed stronger immune responses than males reared in silence: they were better able to encapsulate artificial nylon implants and showed higher levels of antimicrobial lysozyme-like activity in their haemolymph. Experience of sexual signals thus translates into increased immunity, which suggests that such signals may play a role in conveying information about population demography and shaping density-dependent responses in unintended receivers.     

Variations in sexual behavior among capuchin monkeys function for conspecific mate recognition: a phylogenetic analysis and a new hypothesis for female proceptivity in tufted capuchins

Researchers of capuchin monkeys have noted stereotyped body postures, facial expressions, and vocalizations that accompany copulations in this genus. Notable variations in these sexual behaviors are observed across capuchin species. Although several hypotheses exist to explain variation in the duration and vigor of sexual behaviors across species, there is no proposed explanation for variation in the forms of these behaviors. I hypothesized that the forms of sexual behaviors function as recognition signals of conspecific mates. Such signals are adaptive when F1 hybrids exhibit reduced fitness compared with nonhybrid offspring. Recent evidence from nonprimate taxa supports the existence of species recognition signals during mating. Using newly observed sexual behaviors for Cebus albifrons and a recent phylogeny of capuchins, I found significant support for a key prediction of the conspecific mate recognition hypothesis: evolutionary changes in sexual behaviors were associated with speciation. Given the resultant best model for evolution of sexual behaviors, I reconstructed the ancestral pattern of sexual behaviors for extant capuchin species (genera Cebus and Sapajus). This reconstruction suggests that the extreme female proceptivity of tufted capuchin monkeys may function to increase female reproductive choices in the context of sperm-limited males (genus Sapajus).     

Warning signals evolve to disengage Batesian mimics

Prey that are unprofitable to attack are typically conspicuous in appearance. Conventional theory assumes that these warning signals have evolved in response to predator receiver biases. However, such biases might be a symptom rather than a cause of warning signals. We therefore examine an alternative theory: that conspicuousness evolves in unprofitable prey to avoid confusion with profitable prey. One might wonder why unprofitable prey do not find a cryptic means to be distinct from profitable prey, reducing both their risk of confusion with profitable prey and their rate of detection by predators. Here we present the first coevolutionary model to allow for Batesian mimicry and signals with different levels of detectability. We find that unprofitable prey do indeed evolve ways of distinguishing themselves using cryptic signals, particularly when appearance traits can evolve in multiple dimensions. However, conspicuous warning signals readily evolve in unprofitable prey when there are more ways to look different from the background than to match it. Moreover, the more unprofitable the prey species, the higher its evolved conspicuousness. Our results provide strong support for the argument that unprofitable species evolve conspicuous signals to avoid confusion with profitable prey and indicate that peak shift in conspicuousness-linked traits is a major factor in its establishment.     

Functional significance of behavioral,morphological, and endocrine correlates across the ovarian cycle in semifree ranging female Tonkean macaques

The role of sexual displays in mating strategies and their reliability in indicating the time of ovulation has given rise to multiple explanations in nonhuman primates. In order to discriminate among hypotheses, socio-sexual behaviors were recorded in a semifree ranging group of Tonkean macaques (Macaca tonkeana), together with sexual skin swelling volumes and measurements of urinary concentrations of estrone conjugates and pregnanediol glucuronide. A clear preovulatory peak of urinary estrogen levels occurred 2 days before a defined rise in pregnanediol glucuronide concentrations, indicating that both hormones pointed out the female's ovulatory period. The concept of estrus rightly could be applied to female Tonkean macaques since fluctuations in estrogen levels correlated with cyclic changes in genital swelling sizes and rates of female behavioral attractivity and proceptivity. Males proved to be capable of recognizing the optimal conception period as judged from the occurrence of maximal rates of following behavior, serial matings, and ejaculations during the peri-ovulatory phase. During this time, males succeeded in maintaining exclusive and enduring associations with females. However, consortships occurred precociously, with males starting to affiliate with females, follow, and mount them 1 week before the presumed time of ovulation. These long-lasting consortships appear to be a consequence of the female extended follicular phase. This presumably sexually selected character allowed females to extend conspicuous sexual displays: genital swelling and utterance of an estrous call, which might attract males' attention and arouse them. With regard to female mating tactics, the combination of reliably indicating the time of ovulation to the male and durable periods of competitor exclusion led to reject explanations assuming manipulation about paternity or long-lasting intermale competition incitement in Tonkean macaques. Competition for mates between females also turned out to be an irrelevant factor as it was very low in the species. We conclude that the main function of sexual displays is to herald the approach of ovulation toward available mates. Am. J. Primatol. 46:285–309, 1998. © 1998 Wiley-Liss, Inc.     

Primate copulation calls and postcopulatory female choice

Females in some species of Old World monkeys and apes vocalizeafter copulation, but the function of these vocalizations isnot clear. In this article, we examine the hypothesis that copulationcalls are a form of postcopulatory female choice. Accordingto this hypothesis, copulation calls are honest signals of fertility(i.e., ovulation) that are used by females to encourage mateguarding by their preferred mating partners and reduce the likelihoodof sperm competition. Evidence in favor of this hypothesis isreviewed and discussed in relation to other hypotheses. We suggestthat the evolution of female copulation calls in primates islinked to the evolution of other female mating signals suchas exaggerated sexual swellings, the potential for sperm competition,and the opportunity for precopulatory female mate choice.     

Dynamics of Male–Female Multimodal Signaling Behavior across the Estrous Cycle in Giant Pandas (Ailuropoda melanoleuca)

Giant panda courtship behavior includes multimodal signaling assemblages consisting of olfactory, vocal, and postural elements. While signaling is generally conspicuous, successful copulation is inconsistently achieved in captivity, even when female behavioral and physiological data indicate that ovulation is imminent. We set out to characterize these complex patterns of social behavior by observing interactions between 26 unique pairs of giant pandas housed in adjoining pens throughout the females' reproductive cycle. We categorized social behaviors from a transactional perspective and examined social exchanges via analyses of the relative frequency of social behaviors, and via the sequential relationship between male and female social behavior. From non‐estrus to peak‐estrus, we found that the relative frequency of female affiliative and sexual behavior increased and that the relative frequency of ambivalent and aggressive behavior decreased. Male behavior was fairly constant, except for sexual behavior, which increased during peak‐estrus, when it was facilitated by female sexual behavior. Sequential analysis of social interactions showed that preceding behavior had a significant influence over the other panda's response behavior primarily during peak‐estrus, suggesting that pandas are most responsive to conspecific signaling during the peri‐ovulatory period. However, behavioral momentum was a dominant feature of the intra‐individual transitions. Females maintained sexual, ambivalent, and neutral behavior during interactions significantly more than would be expected by chance, with male behavior bearing little influence once the behavior was initiated. A similar pattern was also observed in males, who maintained affiliative, interested, and neutral behaviors. Overall, our data suggest that the multimodal signals used by giant pandas during courtship do not consistently evoke a discrete, immediate response from receivers. Instead, signals appear to advertise reproductive condition and influence potential mates over longer timeframes, suggesting the potential tonic role of communication.     

Conspicuous male coloration impairs survival against avian predators in Aegean wall lizards,Podarcis erhardii

Animal coloration is strikingly diverse in nature. Within‐species color variation can arise through local adaptation for camouflage, sexual dimorphism and conspicuous sexual signals, which often have conflicting effects on survival. Here, we tested whether color variation between two island populations of Aegean wall lizards (Podarcis erhardii) is due to sexual dimorphism and differential survival of individuals varying in appearance. On both islands, we measured attack rates by wild avian predators on clay models matching the coloration of real male and female P. erhardii from each island population, modeled to avian predator vision. Avian predator attack rates differed among model treatments, although only on one island. Male‐colored models, which were more conspicuous against their experimental backgrounds to avian predators, were accordingly detected and attacked more frequently by birds than less conspicuous female‐colored models. This suggests that female coloration has evolved primarily under selection for camouflage, whereas sexually competing males exhibit costly conspicuous coloration. Unexpectedly, there was no difference in avian attack frequency between local and non‐local model types. This may have arisen if the models did not resemble lizard coloration with sufficient precision, or if real lizards behaviorally choose backgrounds that improve camouflage. Overall, these results show that sexually dimorphic coloration can affect the risk of predator attacks, indicating that color variation within a species can be caused by interactions between natural and sexual selection. However, more work is needed to determine how these findings depend on the island environment that each population inhabits.     

Receiver psychology and the evolution of multicomponent signals

Many animals produce and respond to signals made up of multiple components. For example, many avian sexual displays are highly extravagant combinations of visual and acoustic elements, and are described as being 'multicomponent'. One possible reason for the evolution of such complex signals is that they provide more reliable information for receivers. However, receivers also influence signal evolution in another important way, by how they perceive and process signals: signallers will be selected to produce signals that are more easily received. The potential role of receiver psychology in the evolution of multicomponent signals has not previously been considered; in this review I present psychological results that support the notion that two components are better received than one alone. Detection can be improved by producing two components together, thus reducing the reaction time, increasing the probability of detection and lowering the intensity at which detection occurs. Discriminability of multicomponent stimuli is also made easier through better recognition, faster discrimination learning and multidimensional generalization. In addition, multicomponent stimuli also improve associative learning. I show that multicomponency does indeed improve signal reception in receivers, although the benefits of producing components in two sensory modalities (bimodal multicomponent signals) may be larger and more robust than producing them in just one (unimodal multicomponent signals). This highlights the need for consideration of receiver psychology in the evolution of multicomponent signals, and suggests that where signal components do not appear to be informative, they may instead be performing an important psychological function. Copyright 1999 The Association for the Study of Animal Behaviour.     

Primate sexual swellings as coevolved signal systems

Many female catarrhine primates possess visually conspicuous organs that apparently function to increase the sexual interest of adult male conspecifics around the time the female is ovulating—i.e. sexual swellings. The hypothesized functional benefits for both sexes of these sexual swellings are reviewed (honest signaling; paternity confusion; paternity confidence and paternal investment; protection; incitement of precopulatory male-male competition; and postcopulatory sexual selection), as well as an additional hypothesis that has not yet been applied to this problem (sensory exploitation). Currently available evidence is presented that supports or fails to support each of these hypotheses. Predictions associated with broad groupings of these hypotheses, which could be tested in noninvasive field studies, are then presented. Ecological circumstances are discussed that could have led to differential mating success among female primates, and hence to sexual selection on females and directional evolution of sexual swellings. It is concluded that the available evidence does not support the paternity confidence-paternal investment hypothesis; that the paternity confusion hypothesis lacks empirical support, but could still be viable; and that insufficient data exists at present to rigorously test the other hypotheses. The ecological factors that may have led to differential reproductive success among females as a function of mating frequency or mate choice likewise require further empirical investigation.     

Exaggerated Sexual Swellings and the Probability of Conception in Wild Sanje Mangabeys (<Emphasis Type="Italic">Cercocebus sanjei</Emphasis>)

Females of several catarrhine primate species exhibit exaggerated sexual swellings that change in size and coloration during the menstrual cycle and, in some species, gestation. Although their function remains under debate, studies indicate that swellings may contain information males could use to discern ovulation and the probability that a cycle will be conceptive. Here we combine visual ratings of swellings with hormonal data for a group of Sanje mangabeys (18 adult, 3 adolescent females) to determine if their swellings provide reliable information on female fertility. In all cases where ovulation was detected (N = 7), it occurred during maximum tumescence, and in 83.3% during the first two days of the “shiny phase,” a period during maximum tumescence when the swelling was brightest. There were no significant differences in maximum tumescence and shiny phase duration among cycles of different probability of conception, although there was a trend toward conceptive cycles exhibiting shorter shiny phases than nonconceptive ones. Only 25% (N = 4) of postconceptive swellings developed the shiny phase, and adolescents displayed the longest maximum tumescence and shiny phases. The conspicuous nature of the shiny phase and the frequent overlap between its onset and ovulation suggest that its presence serves as a general signal of ovulation and that the cycle has a high probability of being conceptive. It also suggests that swellings in some Sanje mangabeys are more accurate signals of fertility than in other primates.     

Information,influence, and the causal-explanatory role of content in understanding receiver responses

Sceptics of informational terminology argue that by attributing content to signals, we fail to address nonhuman animal communication on its own terms. Primarily, we ignore that communication is sender driven: i.e. driven by the intrinsic physical properties of signals, themselves the result of selection pressures acting on signals to influence receivers in ways beneficial for senders. In contrast, information proponents argue that this ignores the degree to which communication is, in fact, receiver driven. The latter argue that an exclusive focus on the intrinsic mechanical properties of signals cannot explain why receivers respond as they do. This is because receivers are not prisoners of sender influence. They possess response flexibility, and so we can only explain why receivers respond to signals as they do by positing that receivers ‘derive information’ from signals. I argue that, while basically true, this response flexibility can take one of two forms depending on the causal-explanatory role of information in understanding the response of the receiver: diachronic, on the one hand; and synchronic, on the other. In species with diachronic response flexibility only, information is derived by receivers from signals in a minimal sense. In such cases, information is an ultimate explanatory construct: one underpinned by historical facts at the population level. Alternatively, in species with synchronic response flexibility, information is derived by receivers from signals in a richer sense. Here, information is a proximate explanatory construct: one underpinned by cognitive-mechanistic facts at the level of the individual organism. Without recognising the different ways information can be derived from signals, and the different causal-explanatory roles (ultimate vs proximate) information can play in understanding alternate kinds of receiver flexibility (diachronic vs synchronic), proponents of information leave themselves open to the charge of anthropomorphising some signalling systems.     

Aposematic signal variation predicts male-male interactions in a polymorphic poison frog

Many species use conspicuous "aposematic" signals to communicate unpalatability/unprofitability to potential predators. Although aposematic traits are generally considered to be classic examples of evolution by natural selection, they can also function in the context of sexual selection, and therefore comprise exceptional systems for understanding how conspicuous signals evolve under multifarious selection. We used males from a highly territorial poison frog species in a dichotomous choice behavioral test to conduct the first examination of how aposematic signal variation influences male-male interactions. Our results reveal two behavioral patterns: (1) male dorsal brightness influences the behaviors of male conspecifics such that males approach and call to brighter males more frequently and (2) a male's dorsal brightness predicts his own behavior such that bright males approach stimulus frogs faster, direct more calls to bright stimulus frogs, and exhibit lower advertising call pulse rates (a fitness-related trait). These findings indicate the potential for sexual selection by male-male competition to impact aposematic signal evolution.     

Does spatial variation in predation pressure modulate selection for aposematism?

It is widely believed that aposematic signals should be conspicuous, but in nature, they vary from highly conspicuous to near cryptic. Current theory, including the honest signal or trade‐off hypotheses of the toxicity–conspicuousness relationship, cannot explain why adequately toxic species vary substantially in their conspicuousness. Through a study of similarly toxic Danainae (Nymphalidae) butterflies and their mimics that vary remarkably in their conspicuousness, we show that the benefits of conspicuousness vary along a gradient of predation pressure. Highly conspicuous butterflies experienced lower avian attack rates when background predation pressure was low, but attack rates increased rapidly as background predation pressure increased. Conversely, the least conspicuous butterflies experienced higher attack rates at low predation pressures, but at high predation pressures, they appeared to benefit from crypsis. Attack rates of intermediately conspicuous butterflies remained moderate and constant along the predation pressure gradient. Mimics had a similar pattern but higher attack rates than their models and mimics tended to imitate the signal of less attacked model species along the predation pressure gradient. Predation pressure modulated signal fitness provides a possible mechanism for the maintenance of variation in conspicuousness of aposematic signals, as well as the initial survival of conspicuous signals in cryptic populations in the process of aposematic signal evolution, and an alternative explanation for the evolutionary gain and loss of mimicry.     

Sexual swellings in wild white-handed gibbon females (Hylobates lar) indicate the probability of ovulation

Conspicuous sexual swellings in the females of some primate species have been a focus of scientific interest since Darwin first wrote about them in 1871. To understand these visual signals, research focused on exaggerated sexual swellings of Old World primates. However, some primate species develop much smaller sexual swellings and it is as yet unclear if these smaller swellings can serve similar functions as those proposed for exaggerated swellings, i.e. advertising fertility to attract mates. We studied the temporal patterns of sexual swellings, timing of ovulation and female reproductive status in wild white-handed gibbons (Hylobates lar) at Khao Yai National Park, Thailand, where this species has a variable social organization. We established fecal progestogen profiles in fifteen cycles of eight cycling females and, to detect swellings outside the menstrual cycle, five pregnant and six lactating females. In 80% of menstrual cycles, ovulation and maximum swelling phase (duration: ? 9.3 days; 42.8% of cycle length), overlapped tightly. The probability of ovulation peaked on day 3 of the maximum swelling period. Nevertheless, the temporal relationship between maximum swelling and probability of ovulation varied from day -1 to day 13 of the swelling period and three times ovulations fell outside the maximum swelling phase. The different swellings phases occurred in similar proportions in cycling and pregnant, but not lactating females, which were rarely swollen. Despite their smaller size, gibbons' sexual swellings probably serve functions similar to those suggested for exaggerated swellings by the graded-signal hypothesis, which predicts that sexual swellings indicate the probability of ovulation, without allowing males to pinpoint its exact time.