Comparative tests of reproductive skew in male primates: the roles of demographic factors and incomplete control

Reproductive skew models have been proposed as a unifying framework for understanding animal social systems, but few studies have investigated reproductive skew in a broad evolutionary context. We compiled data on the distribution of mating among males for 31 species of primates and calculated skew indices for each study. We analyzed the determinants of mating skew with phylogenetic comparative methods to investigate two models from reproductive skew theory, the concession model and the tug-of-war model. Mating skew decreased as the number of males increased in multimale groups, suggesting that monopolization of females becomes more difficult when there are more rivals, and therefore supporting the tug-of-war model. We predicted that single males are unable to monopolize receptive females as overlap in female receptivity increases (estrous synchrony) and, as a result, that mating skew decreases. However, we did not find any evidence for a link between female estrous synchrony and male mating skew. Finally, the concession model predicts high skew in male philopatric species relative to species in which males disperse, yet our measures of mating skew showed no significant associations with qualitative scores of male dispersal. More definitive tests of the concession model will require more quantitative measures of relatedness, which are presently unavailable for most primate species in our study. Overall, our results provide support for the tug-of-war model in primates, and the approach developed here can be applied to study comparative patterns of skew in other biological systems.Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible to authorized users.     

Mechanisms of sexual fidelity in the monogamous California mouse,Peromyscus californicus

Summary Peromyscus californicus are exclusively monogamous in the wild. We examined in the laboratory whether established pairs of male and female P. californicus would remain faithful when given the opportunity to mate with an opposite sex stranger, either in the presence or absence of their partner. When the partner was present she or he was tethered and could not intervene. Females in postpartum estrus (day 0 postpartum) preferred to associate with their pairmate rather than the unfamiliar virgin male. Females also preferred to mate with their partner, but 15–20% of the females mated with the unfamiliar male both in the presence and absence of their partner. Paired males were tested with an unfamiliar, sexually inexperienced estrous female on days 4 and 8 postpartum with their partners either absent or present but not in postpartum estrus, and on day 0 in the absence of their partner. Males preferred to associate with their pairmate. Contrary to theoretical expectations, males did not copulate with the estrous female when given the opportunity regardless of whether their partner was present or not. Furthermore, few males (2/13) tested in the absence of their partner mated with a sexually experienced, postpartum estrous female. These results indicate that monogamy in P. californicus is maintained by a strong attraction and preference of pairmates for each other and by self-restraint from mating with others. Male mate guarding may further ensure female faithfulness. Males exhibit apparently more sexual fidelity than females.Correspondence to: D.J. Gubernick     

Arousal by sexual stimuli accelerates the re-entrainment of hamsters to phase advanced light-dark cycles

Summary The arousal caused by the 3 h presentation of female socio-sexual cues to male hamsters given an 8 h advance of the light-dark cycle caused an acceleration in the rate of re-entrainment to this new light-dark cycle. This accelerating effect was seen most clearly in animals allowed to run in their wheels in response to a nearby female hamster in estrous condition. However, wheel-running was not a necessary condition for enhancement. In one experiment, males with their wheels prevented from rotating also showed an accelerated rate of re-entrainment; in another experiment, the effect was almost significant (P=0.058). Mating itself seemed to counteract this accelerating effect. Males aroused by nearby but inaccessible estrous females re-entrained quicker than males allowed to mate with an estrous female. Lastly, male hamsters placed in a novel wheel but not allowed to run showed a variable rate of re-entrainment to an 8 h light-dark cycle advance; individuals judged to be sleeping during most of the 3 h in the novel wheel took longer to re-entrain than those judged to be awake during most of the 3 h. These results suggest that there is a general arousal mechanism through which non-specific, non-photic events can enhance the phase advancing actions of a lightentrainable clock.     

Cycle synchrony and probability of conception in female hamadryas baboonsPapio hamadryas

We investigated the factors affecting conception in a captive population ofPapio hamadryas. We collected data on reproductive states from 16 females over 12 years. Probability of conception was related to the outcome of the immediately preceding pregnancy and the fate of the preceding offspring. After abortions or when an infant survived more than 6 months, a female needed more cycles to conceive than when an infant died within the first 6 months post partum. The degree of estrus synchrony within, but not between, one-male units influenced the probability of conception. Females experiencing conceptive estrus showed less synchrony than those experiencing nonconceptive estrus. The number of females simultaneously in estrus within a one-male unit was negatively correlated with the probability of conception. Within our captive population, group size and male and female age had no influence on the probability of conception. The results indicate that sperm may be a limited resource in the one-male reproductive units of hamadryas baboons. Female-female competition for conception may exist and should influence the demography of one-male units.     

Dominance rank,female reproductive synchrony,and male reproductive skew in wild Assamese macaques

In groups with multiple males, direct mate competition may select for the evolution of dominance hierarchies that sort males into a queue for access to fertile females. The priority-of-access (PoA) model proposed by Altmann in 1962 makes explicit predictions about the resulting paternity distribution based on an interaction between male dominance rank and the overlap of female receptive phases. Here, we investigated whether the logic of the PoA model predicted the distribution of paternity across ranks in a seasonal breeder with high reproductive synchrony over six consecutive mating seasons. We studied 18 males that resided in a group of wild Assamese macaques (Macaca assamensis) in their natural habitat at Phu Khieo Wildlife Sanctuary, Thailand, between 2006 and 2011 with 5 to 13 conceptions per season. We assessed whether mate guarding increased paternity success, described “short-term” deviations from predicted paternity distribution, and examined how these are related to the number of competitors and fertile females. We determined genetic paternity of 43 (93 %) offspring born into the study group and found reproductive skew to be relatively low with 29 % alpha male paternity in accordance with the high degree of female reproductive synchrony observed. Short-term deviations from expected paternity distribution over ranks were not explained by the number of resident males or the number of conceiving females or their interaction. Within the limits of this study, these results suggest that even if males cannot discern female fertile phases, if reproduction is seasonal, and if reproductive synchrony is high, males may also compete directly over access to females.     

Variability in adult group compositions of a prosimian primate

Many species fall into specific mating-system categories, and that category is usually associated with a suite of behavioral and morphological characteristics. Several lemur species, including Propithecus diadema edwardsi, have been labeled "idiosyncratic" because variation in socionomic sex ratios among groups is consistent with wide variation in social structure. We used several hypotheses founded in behavioral ecology to assess variability in P. d. edwardsi. First we examined 46 group-mating seasons to quantify variability. We then tested predictions that the number of males per group would increase as the number of adult females increased, and the number of males would increase as female mating synchrony increased. Examining variation in offspring survival relative to the number of adult males in a group may tell us which composition is likely to persist into the future, so we also hypothesized that as the number of males in a group increased, fertility and offspring survival would increase. We found an equal distribution of polygynous, polygynandrous, pairs, and polyandrous groups. Furthermore, female distribution and mating synchrony did not predict the number of males, and offspring survival was not correlated with the number of males. Since infants survived equally well in groups of all compositions, sifakas experienced no pressure to maintain a particular number of adult males per adult female. The small number of adults per group (mean=3.2) may result from balancing feeding competition against predator detection. Augmenting the mate pool available from the group with mates from neighboring groups may promote the notable variability seen in the adult group compositions of sifakas.     

Breeding synchrony and paternity in the barn swallow (Hirundo rustica)

The barn swallow (Hirundo rustica) is a socially monogamous passerine which usually breeds in colonies where extra-pair copulations are frequent. Males intensively guard their mates during the female fertile period. Since males are more likely to be available for extra-pair copulations when their mate is not fertile, synchrony in timing of breeding may affect paternity of individual males. In this study, we analysed the change in mate-guarding rate by males in relation to the fertility condition of the female, and the relationships between breeding synchrony and density with paternity in first broods of 52 male barn swallows. Paternity (proportion of nestlings fathered in own brood) was assessed by typing of three highly polymorphic microsatellite loci. Mate guarding by males peaked during the fertile period of their mates. Paternity increased as breeding synchrony in the colony increased. Paternity of barn swallows is positively associated with the degree of exaggeration of male tail ornaments. The relationship between male ornamentation and paternity was partly mediated by an effect of ornament size on breeding synchrony. We suggest that females might delay breeding with low-quality males to enhance their opportunities for being fertilised by high-quality extra-pair males. Received: 16 January 1998 / Accepted after revision: 25 October 1998     

The number of males in primate social groups: a comparative test of the socioecological model

As applied to polygynous mammals, the socioecological model assumes that environmental risks and resources determine the spatial and temporal distribution of females, which then sets male strategies for monopolizing fertile matings. The effects of female spatial distribution (i.e., female number) and temporal overlap (female mating synchrony) have been examined in comparative studies of primates, but the relative influence of these two factors on male monopolization potential (the number of males) remains unclear. One particular problem is that female synchrony is more difficult to estimate than female number. This paper uses multivariate statistical methods and three independent estimates of female synchrony to assess the roles of spatial and temporal effects in the context of a phylogenetically corrected dataset. These analyses are based on sensitivity analyses involving a total of four phylogenies, with two sets of branch length estimates for each tree, and one nonphylogenetic analysis in which species values are used (because male behavior may represent a facultative response to the distribution of females). The results show: (1) that breeding seasonality predicts male number (statistically significant in six out of nine sensitivity tests); (2) that expected female overlap, after controlling for female group size using residuals, also accounts for the number of males in primate groups (significant in eight out of nine tests), and (3) that actual estimates of female mating synchrony predict male number, again after correcting for female group size (significant in five out of nine tests). Nonsignificant results are in the predicted direction, and female group size is significant in all statistical tests. These analyses therefore demonstrate an independent influence of female temporal overlap on male monopolization strategies in mammalian social systems. Received: 24 July 1998 / Received in revised form: 5 February 1999 / Accepted: 7 February 1999     

Extra-pair paternity in eastern bluebirds: effects of manipulated density and natural patterns of breeding synchrony

The causes of variation in rates of extra-pair paternity among avian populations remain unclear, but could include environmental factors such as breeding density and synchrony. By experimentally manipulating nest site availability, we tested the effects of breeding density on the frequency of extra-pair paternity in eastern bluebirds (Sialia sialis). We also examined the role of breeding synchrony on extra-pair paternity using natural timing of nests. Microsatellite analysis revealed 34 of 305 nestlings (11.2%) were the result of extra-pair fertilizations; and 21 of 79 broods (26.6%) had at least one extra-pair nestling. Several measures of breeding density had independent effects on extra-pair paternity. First, experimental plot type affected extra-pair paternity, with 28 of 34 (82.4%) extra-pair young from nests in high density areas, and only six (17.6%) from nests in low density areas. Independently of plot type, the number of breeding neighbors within a 320-m radius was a significant predictor of the likelihood of extra-pair paternity at the nest. Extra-pair paternity was associated with temporal factors such as absolute timing of breeding and natural levels of local breeding synchrony, but only in bivariate comparisons. We found a positive interaction between density of neighbors within a 320-m radius and local breeding synchrony; this term reduced the main effects of synchrony and number of neighbors, but not experimental treatment. Our results demonstrate the importance of utilizing multiple aspects of proximity in breeding density analyses and testing for interactions between ecological factors that can influence the behavioral events leading to extra-pair fertilizations.     

Altered responses to female odors in parasitized male mice: neuromodulatory mechanisms and relations to female choice

There is accumulating evidence that females may preferentially select parasite-free or -resistant males. Minimal attention has, however, been paid to the mate preferences and responses of the parasitized male hosts themselves. Here, we considered the effects of parasitic infection on male host mate responses, the neuromodulatory correlates of these responses, and the relations of these responses to female mate choice. Using an odor “preference” test, we examined the effects of different stages of an acute, sub-clinical infection with the naturally occurring, enteric, single host, protozoan parasite, Eimeria vermiformis, on the responses of male mice, Mus musculus domesticus, to the odors of estrous females along with the responses of uninfected females to the parasitized males. At 4 days post-infection (non-infective, pre-patent stage) E. vermiformis-infected male mice showed a significantly decreased preference for the odors of estrous females, whereas at 10 days post-infection (infective, patent stage) infected males showed a significantly increased preference for the odors of estrous females. Parasitized males displayed no significant changes in their responses to the odors of non-estrous females, supporting effects on the reproductively related responses of the host. In parallel, estrous females displayed a reduced interest in the odors of infected males. Least interest was expressed in the odors of the patent, infective males, consistent with the avoidance of contagion. Using selective opioid peptide receptor agonists and antagonists we found evidence that enhanced kappa opioid peptide (e.g., dynorphin) activity was related to the decreased sexual interest of the pre-infective males, while augmented delta opioid peptide (e.g., enkephalin) activity was associated with the enhanced responses of the infective males to females. We further showed that acute kappa opiate administration reduced the responses of uninfected males to females and that uninfected females displayed modified responses to the odors of uninfected males subject to acute modifications of opioid activity. We suggest that these differential shifts in endogenous opioid activity in the parasitized males are associated with and, or related to alterations in neuro-immune and endocrine functions. These findings show that parasitic infection can have, depending on the stage of infection and associated neuromodulatory changes, either significant facilitatory or inhibitory effects on male host preferences for and responses to females. Received: 22 April 1996 / Accepted after revision: 15 March 1997     

Mating skew in Barbary macaque males: the role of female mating synchrony,female behavior,and male–male coalitions

A fundamental question of sexual selection theory concerns the causes and consequences of reproductive skew among males. The priority of access (PoA) model (Altmann, Ann NY Acad Sci 102:338–435, 1962) has been the most influential framework in primates living in permanent, mixed-sex groups, but to date it has only been tested with the appropriate data on female synchrony in a handful of species. In this paper, we used mating data from one large semi-free ranging group of Barbary macaques: (1) to provide the first test of the priority-of-access model in this species, using mating data from 11 sexually active females (including six females that were implanted with a hormonal contraceptive but who showed levels of sexual activity comparable to those of naturally cycling females) and (2) to determine the proximate mechanism(s) underlying male mating skew. Our results show that the fit of the observed distribution of matings with sexually attractive females to predictions of the PoA model was poor, with lower-ranking males mating more than expected. While our work confirms that female mating synchrony sets an upper limit to monopolization by high-ranking individuals, other factors are also important. Coalitionary activity was the main tactic used by males to lower mating skew in the study group. Coalitions were expressed in a strongly age-related fashion and allowed subordinate, post-prime males to increase their mating success by targeting more dominant, prime males. Conversely, females, while mating promiscuously with several males during a given mating cycle, were more likely to initiate their consortships with prime males, thus reducing the overall effectiveness of coalitions. We conclude that high-ranking Barbary macaque males have a limited ability to monopolize mating access, leading to a modest mating skew among them.     

The significance of hotspots to lekking topi antelopes (<Emphasis Type="Italic">Damaliscus lunatus</Emphasis>)

Controversy has surrounded the question of why lek-breeding has evolved in certain ungulate species. Can the behavior be explained simply by males mapping onto a female distribution that is determined by factors unrelated to mating? Or are leks created because estrous females distinguish between males and favor males who cluster? Here I address these questions by looking at spatial distribution in lekking topi antelopes (Damaliscus lunatus). Contrary to the predictions of a model assuming male clustering in the zone of maximum female range overlap, territories were highly clustered also within this zone, and lek size correlated positively with population density. In support of models derived from the ideal free distribution of males onto female dispersion, leks were in areas with high female density during the rut. However, models not taking into account both individual variation in male quality and female mate preferences failed to explain the extreme male clumping also within high density areas, which was revealed by a strongly male-biased sex-ratio on leks. Additional support for the female preference-based model came from the finding that estrous females concentrated onto leks. Female preference for clustered males may develop if males initially follow an ideal free distribution of unequal competitors with high quality males slightly clustered at density hotspots; positive feedback between female benefits of preference for clustered males and male benefits of clustering could lead to contraction of the territorial network and lek behavior. Thus only the female preference-based model correctly predicted a negative correlation between male mating rate and resource density.     

Spatial synchrony in coral reef fish populations and the influence of climate

We investigated spatial patterns of synchrony among coral reef fish populations and environmental variables over an eight-year period on the Great Barrier Reef, Australia. Our aims were to determine the spatial scale of intra- and interspecific synchrony of fluctuations in abundance of nine damselfish species (genus Pomacentrus) and assess whether environmental factors could have influenced population synchrony. All species showed intraspecific synchrony among populations on reefs separated by < or =100 km, and interspecific synchrony was also common at this scale. At greater spatial scales, only four species showed intraspecific synchrony, over distances ranging from 100-300 km to 500-800 km, and no cases of interspecific synchrony were recorded. The two mechanisms most likely to cause population synchrony are dispersal and environmental forcing through regionally correlated climate (the Moran effect). Dispersal may have influenced population synchrony over distances up to 100 km as this is the expected spatial range for ecologically significant reef fish dispersal. Environmental factors are also likely to have synchronized population fluctuations via the Moran effect for three reasons: (1) dispersal could not have caused interspecific synchrony that was common over distances < or =100 km because dispersal cannot link populations of different species, (2) variations in both sea surface temperature and wind speed were synchronized over greater spatial scales (>800 km) than fluctuations in damselfish abundance (< or =800 km) and were correlated with an index of global climate variability, the El Ni?o-Southern Oscillation (ENSO), and (3) synchronous population fluctuations of most damselfish species were correlated with ENSO; large population increases often followed ENSO events. We recorded regional variations in the strength of population synchrony that we suspect are due to spatial differences in geophysical, oceanographic, and population characteristics, which act to dilute or enhance the effects of synchronizing mechanisms. We conclude that synchrony is common among Pomacentrus populations separated by tens of kilometers but less prevalent at greater spatial scales, and that environmental variation linked to global climate is likely to be a driving force behind damselfish population synchrony at all spatial scales on the Great Barrier Reef.     

Parental investment, adult sex ratios, and sexual selection in a socially monogamous seabird

Although most birds are monogamous, theory predicts that greater female parental investment and female-biased adult sex ratios will lower the polygyny threshold. This should result in polygynous mating, unless obligate biparental care or the spatial and temporal distribution of fertilizable females constrains a male’s ability to take advantage of a lowered polygyny threshold. Here we present data on the extent of male sexually dimorphic plumage, adult sex ratios and breeding season synchrony in three populations of a socially monogamous seabird, the brown booby Sula leucogaster. For one of these populations, San Pedro Mártir Island, we also present data on differences in male and female parental investment, mortality and probability of pairing. The extent of plumage dimorphism varied among populations. Sex ratios were female biased in all populations. On San Pedro Mártir Island, parental investment was female biased, females failed more often than males to find a mate, but there was no polygyny. We suggest that on San Pedro Mártir: (1) a period of obligate biparental care coupled with a relatively synchronous breeding season constrained the ability of males to take advantage of a high environmental polygamy potential and (2) the resulting socially monogamous mating system, in combination with the female-biased adult sex ratio, caused females to be limited by the availability of males despite their greater parental investment. Received: 18 November 1999 / Accepted: 24 January 2000     

Birth synchrony in squirrel monkeys (Saimiri oerstedi)

Summary The vigilance behaviors and association patterns of adult female squirrel monkeys (Saimiri oerstedi) in Parque Nacional Corcovado, Costa Rica were studied over the annual reproductive cycle to identify a possible adaptive function associated with the unusually close within-group birth synchrony prevalent in Saimiri populations. Raptor association with the troops, predation attempts, and monthly changes in food abundance were also documented. Seasonal fluctuations in food abundance probably constrained the timing of births among three contiguous troops to a two month period. The close within-troop birth synchrony (majority of births occurring within a week) is suggested to be advantageous because of intense predation pressure on neonate squirrel monkeys. Adult females with neonates cooperate in anti-predator vigilance and defense of neonates. Participation in these maternal associations may increase the probability of neonate survivorship. Variable gestation length in squirrel monkeys may reflect a lability in the duration of gestation to allow coordination of the timing of births among females in a troop.     

Reproductive seasonality is a poor predictor of receptive synchrony and male reproductive skew among nonhuman primates

Among nonhuman primates, male reproductive skew (i.e., the distribution of reproductive success across males) appears to be affected primarily by receptive synchrony and the number of males per group. These factors have been assumed to depend on reproductive seasonality, with strong seasonality increasing receptive synchrony, which in turn reduces the strength of male monopolization associated with more males and lower skew. Here we tested the importance of reproductive seasonality for 26 populations representing 15 species living in multimale groups. We obtained data from the literature on paternity, number of males per group, receptive synchrony, and three measures of seasonality of reproduction. We analyzed these data using bivariate regressions and hierarchical regression by sets and controlled for the effect of evolutionary relationships using phylogenetic generalized least squares. As expected, alpha male paternity decreased as the number of males per group increased as well as with increasing female receptive synchrony. Reproductive seasonality did not explain variation in reproductive skew over and above the variation explained by synchrony and the number of males. Reproductive seasonality alone only explained a small proportion of the variation in skew, and there was no strong association between reproductive seasonality and synchrony. The effects of receptive synchrony and reproductive seasonality as well as their link were reduced if we excluded captive populations. These results indicate that across primates male reproductive skew is related to the number of competitors in a group and that seasonality does not reliably predict synchrony or male reproductive skew.     

Mate attraction by females in a sexually cannibalistic praying mantis

We studied mate attraction by females of the praying mantid, Tenodera aridifolia sinensis, testing honest signaling of mate availability versus deceptive signaling to attract males for sexual cannibalism. We experimentally varied female diet and mating history and measured the rate of attraction of a wild population of males to caged females. Honest signaling theory predicts that virgin females will attract males at the greatest rate whereas deceptive signaling predicts that hungry females (which are more likely to cannibalize males) will attract more males, particularly among non-virgin females. Our results show that hungry females did not attract more males than well-fed females. Indeed, the opposite was true: hungry females attracted significantly fewer males. Moreover, hungry females were no more likely than well-fed females to attract males subsequent to mating, and mated females attracted males at a lower rate than did virgin females. We also observed female T. aridifolia sinensis and male Mantis religiosa arriving at the caged females and we discuss the significance of these observations. The results refute the hypothesis of deceptive signaling and show that mate attraction signals of female T. aridifolia sinensis are honest indicators of female mate availability and a lower risk of sexual cannibalism.     

Mating competition in an asocial ground squirrel,Spermophilus tridecemlineatus

Summary Male thirteen-lined ground squirrels (Spermophilus tridecemlineatus) are subject to three forms of intrasexual competition: competition over location of spatially scattered females, overt conflict over access to those females, and sperm competition. The likelihood of both overt conflict and sperm competition varies between seasons; changes in male density, female density, and breeding synchrony were investigated as possible sources of that variability. Male density was artificially reduced mid-way through one mating season. The results of that experiment, as well as subsequent between-season comparisons, indicate that fluctuations of male density have hardly any effect on levels of overt conflict and sperm competition. By contrast, those forms of intrasexual competition are influenced strongly by female density and/or breeding synchrony.     

Female choice in the opportunistic mating of wild chimpanzees (Pan troglodytes schweinfurthii) at Mahale

For female chimpanzees (Pan troglodytes schweinfurthii) in the Mahale Mountains National Park, Tanzania, the most common mating pattern is opportunistic. In such opportunistic matings, females copulated promiscuously but not randomly. This study describes female mate choice during 1-year observation of six females who exhibited regular genital-swelling cycles. During the study period, 169 opportunistic matings and four restrictive matings were recorded over the course of 51 days. As female estrus progressed, mating frequency and the number of adult male mating partners increased, although the number of potential mating partners did not change. Criteria of female choice examined were the direction and consent/rejection of courtship, proximity maintenance, and female grooming. Adult-male courtships were successful more often than those of adolescent males. During the earlier phase of estrus, females copulated rather promiscuously with many males. But during the later phase of estrus when the likelihood of conception is expected to be highest, they copulated repeatedly with high-ranking adult males. There was a positive correlation between female grooming frequency and mating frequency when the likelihood of conception was greatest. Female chimpanzees are thought to choose high-ranking males as fathers of their offspring. Moreover, female chimpanzees may adopt one or both of two mating strategies, i.e., a many-male strategy and a best-male strategy. Received: 23 November 1998 / Received in revised form: 12 April 1999 / Accepted: 26 April 1999     

Postcopulatory mate guarding by vocalization in the Formosan squirrel

The Formosan squirrel, Callosciurus erythraeus thaiwanensis, emitted different vocalizations in response to terrestrial and aerial predators and snakes. Each vocalization caused nearby individuals to adopt a different type of anti-predator behaviour. In mating bouts, males produced two types of loud calls: precopulatory calls, emitted before copulations, and postcopulatory calls, emitted after copulations. The latter continued for 17 min on average. The estrous female and other males attending the mating bouts stopped moving during the postcopulatory call, so that the calling male was able to tend the female without interruption. The sound characteristics of anti-terrestrial-predator and postcopulatory calls recorded in the captivity were compared, and none of the ten characters of duration and frequency measured differed between the two calls. Playback experiments also showed that responses to the sounds in two different contexts, escape behaviour and defensive immobility, did not differ. The similarity between anti-predator and postcopulatory calls is discussed with reference to the possibility of manipulation and other explanatory hypotheses.