Personality trait structures across three species of Macaca, using survey ratings of responses to conspecifics and humans.

Comparative studies reliant on single personality surveys to rate wild primates are scarce yet remain critical for developing a holistic comparative understanding of personality. Differences in survey design, item exclusion, and factor selection impede cross-study comparisons. To address these challenges, we used consistently collected data to assess personality trait structures in wild rhesus (Macaca mulatta), bonnet (M. radiata), and long-tailed (M. fascicularis) macaques that varied in their degree of phylogenetic closeness, species-typical social styles, and anthropogenic exposure in urban or urban-rural environments. We administered 51-item personality surveys to familiar raters, and, after reliability and structure screenings, isolated 4-5 factor solutions among the species. Four consistent factors emerged: Confident, Sociable, Active, and Irritable/Equable. This latter factor had differential expression across species. Item composition of the Irritable/Equable factor was consistent with their anticipated differences in social styles, but confounded by cross-site anthropogenic variation. We also administered a 43-item survey confined to human-primate situations which paralleled our findings of social style variation, while also exhibiting variation that aligned with population differences in human density. Our findings indicate that macaque personality trait structures may be emergent outcomes of evolutionary and/or socioecological processes, but further research is needed to parse these processes' relative contributions.

The dynamics of sociality and glucocorticoids in wild male Assamese macaques.

For males of gregarious species, dominance status and the strength of affiliative relationships can have major fitness consequences. Social dynamics also impose costs by affecting glucocorticoids, mediators of homeostasis and indicators of the physiological response to challenges and within-group competition. We investigated the relationships between dominance, social bonds, seasonal challenges, and faecal glucocorticoid metabolite (fGC) measures in wild Assamese macaques (Macaca assamensis) at Phu Khieo Wildlife Sanctuary, Thailand, combining behavioural data with 4129 samples from 62 adult males over 15 years. Our previous work on this population suggested that increased competition during the mating season was associated with elevated fGC levels and that, unusually for male primates, lower rank position correlated with higher fGC levels. With a much larger dataset and dynamic measures of sociality, we re-examined these relationships and additionally tested the potentially fGC-attenuating effect of social support. Contrary to our previous study, yet consistent with the majority of work on male primates, dominance rank had a positive relationship with fGC levels, as high status correlated with elevated glucocorticoid measures. fGC levels were increased at the onset of the mating season. We demonstrated an fGC-reducing effect of supportive relationships in males and showed that dynamics in affiliation can correlate with dynamics in physiological responses. Our results suggest that in a system with intermediate contest potential, high dominance status can impose physiological costs on males that may potentially be moderated by social relationships. We highlight the need to consider the dynamics of sociality and competition that influence hormonal processes.

Necrophilic behaviour in wild stump-tailed macaques (Macaca arctoides).

Necrophilic behavior (attempted copulation with corpses) has been scarcely reported in non-human primates, especially in the wild. Here is the first case of necrophilic behavior observed in wild stump-tailed macaques in Thailand. Six groups of total N > 460 individuals have been identified and habituated. The corpse of an adult female was found and directly observed for 2 days and by camera trap for 3 days. The cause of death could not be identified, but no prominent physical injury was detected. Within 3 days of the observation, three different males attempted copulation with the corpse. Noteworthy for this observation was that not only males in the group of the dead female but also males from different groups interacted with the corpse. Taken together, these observations suggest that some cues emanating from the corpse coupled with a nonresistant/passive orientation may have triggered these responses in the males. Given that necrophiliac responses have been scarcely reported in non-human primates, our findings provide new insight into these behaviors and to comparative thanatology in general.

Simple visual stimuli are sufficient to drive responses in action observation and execution neurons in macaque ventral premotor cortex.

Neurons responding during action execution and action observation were discovered in the ventral premotor cortex 3 decades ago. However, the visual features that drive the responses of action observation/execution neurons (AOENs) have not been revealed at present. We investigated the neural responses of AOENs in ventral premotor area F5c of 4 macaques during the observation of action videos and crucial control stimuli. The large majority of AOENs showed highly phasic responses during the action videos, with a preference for the moment that the hand made contact with the object. They also responded to an abstract shape moving towards but not interacting with an object, even when the shape moved on a scrambled background, implying that most AOENs in F5c do not require the perception of causality or a meaningful action. Additionally, the majority of AOENs responded to static frames of the videos. Our findings show that very elementary stimuli, even without a grasping context, are sufficient to drive responses in F5c AOENs.

Twinning in wild, endangered lion-tailed macaques Macaca silenus in the Anamalai Hills of the Western Ghats, India.

Many primate species show various behavioural and ecological adaptations to provisioning, one of which is the unusual occurrence of twins. Here, we report observations on two pairs of surviving twins in lion-tailed macaques Macaca silenus in the Anamalai Hills of the Western Ghats, India. The Puthuthottam population of lion-tailed macaques has historically been restricted to a rainforest fragment measuring 92 ha, situated adjacent to human settlements. Over the last 10 years, however, several groups from this population have begun to directly interact with the local human communities, visiting settlements at a rate of 0.52 events/day and exploiting various anthropogenic food resources. We followed and opportunistically collected behavioural ad libitum data on two sets of twins for a year, between March 2019 and March 2020. Both of the mothers were primarily terrestrial, although the mother with the younger set of twins also used the tree canopy and other precarious substrates, such as cables. Although two previous cases of twinning have been reported in this population, one in the late 1990s and one between 2000 and 2002, neither of those sets of twins survived beyond a few weeks, with at least one infant in each pair dying of unknown causes. We discuss, but discount, the possibility that one of the infants in either set of twins was an adoptee. Our observations indicate that some lion-tailed macaque twins can survive under free-ranging conditions if they receive adequate care from their biological mother or another female. Our findings also provide further evidence of increased rates of twinning as a consequence of dietary changes in synanthropic non-human primate populations.

Visuo-frontal interactions during social learning in freely moving macaques.

Social interactions represent a ubiquitous aspect of our everyday life that we acquire by interpreting and responding to visual cues from conspecifics1. However, despite the general acceptance of this view, how visual information is used to guide the decision to cooperate is unknown. Here, we wirelessly recorded the spiking activity of populations of neurons in the visual and prefrontal cortex in conjunction with wireless recordings of oculomotor events while freely moving macaques engaged in social cooperation. As animals learned to cooperate, visual and executive areas refined the representation of social variables, such as the conspecific or reward, by distributing socially relevant information among neurons in each area. Decoding population activity showed that viewing social cues influences the decision to cooperate. Learning social events increased coordinated spiking between visual and prefrontal cortical neurons, which was associated with improved accuracy of neural populations to encode social cues and the decision to cooperate. These results indicate that the visual-frontal cortical network prioritizes relevant sensory information to facilitate learning social interactions while freely moving macaques interact in a naturalistic environment.

Effects of 90 dB pure tone exposure on auditory and cardio-cerebral system functions in macaque monkeys.

Excessive noise exposure presents significant health risks to humans, affecting not just the auditory system but also the cardiovascular and central nervous systems. This study focused on three male macaque monkeys as subjects. 90 dB sound pressure level (SPL) pure tone exposure (frequency: 500Hz, repetition rate: 40Hz, 1 min per day, continuously exposed for 5 days) was administered. Assessments were performed before exposure, during exposure, immediately after exposure, and at 7-, 14-, and 28-days post-exposure, employing auditory brainstem response (ABR) tests, electrocardiograms (ECG), and electroencephalograms (EEG). The study found that the average threshold for the Ⅴ wave in the right ear increased by around 30 dB SPL right after exposure (P < 0.01) compared to pre-exposure. This elevation returned to normal within 7 days. The ECG results indicated that one of the macaque monkeys exhibited an RS-type QRS wave, and inverted T waves from immediately after exposure to 14 days, which normalized at 28 days. The other two monkeys showed no significant changes in their ECG parameters. Changes in EEG parameters demonstrated that main brain regions exhibited significant activation at 40Hz during noise exposure. After noise exposure, the power spectral density (PSD) in main brain regions, particularly those represented by the temporal lobe, exhibited a decreasing trend across all frequency bands, with no clear recovery over time. In summary, exposure to 90 dB SPL noise results in impaired auditory systems, aberrant brain functionality, and abnormal electrocardiographic indicators, albeit with individual variations. It has implications for establishing noise protection standards, although the precise mechanisms require further exploration by integrating pathological and behavioral indicators.

Cellular and laminar architecture: A short history and commentary.

The feedforward/feedback classification, as originally stated in relation to early visual areas in the macaque monkey, has had a significant influence on ideas of laminar interactions, area reciprocity, and cortical hierarchical organization. In some contrast with this macroscale "laminar connectomics," a more cellular approach to cortical connections, as briefly surveyed here, points to a still underappreciated heterogeneity of neuronal subtypes and complex microcircuitries. From the perspective of heterogeneities, the question of how brain regions interact and influence each other quickly leads to discussions about concurrent hierarchical and nonhierarchical cortical features, brain organization as a multiscale system forming nested groups and hierarchies, connectomes annotated by multiple biological attributes, and interleaved and overlapping scales of organization.

Developmental and environmental modulation of fecal thyroid hormone levels in wild Assamese macaques (Macaca assamensis).

Thyroid hormones are key modulators of development, as well as mediators of environmental conditions, by regulating developmental processes and metabolism in primates. Hormone measurement in noninvasively collected samples, that is, feces and urine, is a valuable tool for studying the endocrine function of wildlife, and recent studies have demonstrated the feasibility of measuring thyroid hormones in fecal samples of zoo-housed and wild nonhuman primates. Our study aimed to (i) validate the measurement of immunoreactive fecal total triiodothyronine (IF-T3) in wild Assamese macaques (Macaca assamensis) and (ii) to investigate its developmental changes and its response to environmental changes, including stress responses, in immature individuals. Fecal samples and environmental parameters were collected from individuals of three social groups of wild Assamese macaques living at Phu Khieo Wildlife Sanctuary, Northeastern Thailand. Our study confirmed the methodological feasibility and biological validity of measuring IF-T3 in this population. Specifically, the biological validation demonstrated higher IF-T3 levels in immatures compared to adults, and higher levels in females during late gestation compared to the preconception stage. Our analysis of IF-T3 levels in developing immature macaques revealed a significant increase with age. Furthermore, we found a positive association between IF-T3 and immunoreactive fecal glucocorticoid levels, an indicator of the physiological stress response. Neither minimum temperature nor fruit abundance predicted variation in IF-T3 levels in the immatures. Our findings indicate the possibility for differing effects of climatic factors and food availability on thyroid hormone level changes in immature versus adult animals and in wild compared to experimental conditions. Overall, our study provides the basis for further investigations into the role of thyroid hormones in shaping species-specific traits, growth, and overall primate development.

Hormonal responses to mating competition in male Tonkean macaques.

Glucocorticoid and androgen hormones play a prominent role in male reproductive effort. Their production usually increases in non-human primates during mating competition, which may include rivalry for access to receptive females, struggles for high dominance rank, or social pressure on low-ranking individuals. It is generally assumed that glucocorticoids and androgens are associated with mating challenges rather than dominance status, but the involvement of multiple factors makes it difficult to disentangle the two. In this regard, Tonkean macaques provide a suitable model because they are characterized by relaxed dominance and year-round breeding, meaning that there is typically no more than one receptive female in a group, and thus first-ranking males can easily monopolize her. We studied two captive groups of Tonkean macaques over an 80-month period, recording the reproductive status of females, collecting urine from males and sampling behaviors in both sexes. Male urinary hormone concentrations could be affected by increased competition caused by the mating period, the number of males and the degree of female attractiveness. The highest increases in androgens were recorded in males performing female mate-guarding. Despite the importance of dominance status in determining which males can mate, we found no significant effect of male rank on glucocorticoids and only a marginal effect on androgens during mate-guarding. Both types of hormones were more directly involved in the mating effort of males than in their dominance status. Our results show that their function can be understood in light of the particular competitive needs generated by the species-specific social system.

No food left behind: foraging route choices among free-ranging Japanese macaques (Macaca fuscata) in a multi-destination array at the Awajishima Monkey Center, Japan.

Animals must make route choices every day when moving through their habitat while foraging. Choosing an optimal route can be cognitively costly, and primates and other animals have been shown to use simple heuristics, "rules of thumb", to make foraging route choices. We investigated the potential use of heuristics among foraging free-ranging Japanese monkeys (Macaca fuscata) during solitary foraging trials. We also investigated the potential influence of individual variables (age and sex) and social variables (presence in the central group, presence of potential inter- and intraspecific competitors), on the use of heuristics, route length and trial time. We used a multi-destination foraging experiment with 6 platforms in a (4 m × 8 m) Z-array, completed by 29 Japanese macaques in 155 runs at the Awajishima Monkey Center in Japan. Our results showed that the macaques chose routes consistent with heuristics (e.g. nearest neighbour heuristic 19.4%, convex hull heuristic 4.5%) and selected optimal routes (shortest path in 23.9% of the trials). We also identified a potential new heuristic that was used most frequently, that we termed the "sweep heuristic" (27.1% of trials), which we interpreted as a strategy to deal with competitive foraging trade-offs - choosing routes to prioritize not leaving isolated food pieces behind. Age was significantly related to trial time; juvenile macaques were faster than adults and young adults, using speed to gain access to resources. Solitary trials with conspecifics present took significantly longer routes. Our results suggest that contextual factors led to variation in Japanese macaque decision-making, and we suggest that the preferential use of a sweep heuristic may have been a response to high intragroup competition.

Hand preference predicts behavioral responses to threats in Barbary macaques.

The structure and functioning of the brain are lateralized-the right hemisphere processes unexpected stimuli and controls spontaneous behavior, while the left deals with familiar stimuli and routine responses. Hemispheric dominance, the predisposition of an individual using one hemisphere over the other, may lead to behavioral differences; particularly, an individual may be programed to act in a certain way concerning hemispheric dominance. Hand preference is a robust estimator of hemispheric dominance in primates, as each hemisphere controls the opposing side of the body. Studies have found links between hand preference and the exhibition of behaviors in contexts such as exploring and manipulating objects. However, little is known whether hand preference predicts behavioral variations in other ecologically relevant contexts like predation. We investigated the relationship between hand preference and behavioral responses to two types of predator models in captive Barbary macaques (Macaca sylvanus) (n = 22). Besides, a nonpredator novel object was included as control. We found 91% of the macaques to be lateralized with no group-level bias. A higher rate of tension and focus (behavioral response) behavior was found in predator contexts than in the novel object condition. Unlike their right-hand counterparts, individuals with a strong left-hand preference elicited frequent focus and tension behavior toward the predator models. Additionally, the behavioral response varied with predator type. We also found an interaction effect between hand preference and predator type. Our study suggests that hand preference can reliably predict behavioral variations in the context of potential predation. While these results are consistent with lateralized brain function, indicating lateralization a neural mechanism of behavioral variation, the interaction effect between hand preference and predator type elucidates the importance of context-specificity when investigating laterality noninvasively. Future research on other nonhuman primates using the current framework may provide insights into the evolution of laterality and underlying behavioral predispositions.

A somato-cognitive action network alternates with effector regions in motor cortex.

Motor cortex (M1) has been thought to form a continuous somatotopic homunculus extending down the precentral gyrus from foot to face representations1,2, despite evidence for concentric functional zones3 and maps of complex actions4. Here, using precision functional magnetic resonance imaging (fMRI) methods, we find that the classic homunculus is interrupted by regions with distinct connectivity, structure and function, alternating with effector-specific (foot, hand and mouth) areas. These inter-effector regions exhibit decreased cortical thickness and strong functional connectivity to each other, as well as to the cingulo-opercular network (CON), critical for action5 and physiological control6, arousal7, errors8 and pain9. This interdigitation of action control-linked and motor effector regions was verified in the three largest fMRI datasets. Macaque and pediatric (newborn, infant and child) precision fMRI suggested cross-species homologues and developmental precursors of the inter-effector system. A battery of motor and action fMRI tasks documented concentric effector somatotopies, separated by the CON-linked inter-effector regions. The inter-effectors lacked movement specificity and co-activated during action planning (coordination of hands and feet) and axial body movement (such as of the abdomen or eyebrows). These results, together with previous studies demonstrating stimulation-evoked complex actions4 and connectivity to internal organs10 such as the adrenal medulla, suggest that M1 is punctuated by a system for whole-body action planning, the somato-cognitive action network (SCAN). In M1, two parallel systems intertwine, forming an integrate-isolate pattern: effector-specific regions (foot, hand and mouth) for isolating fine motor control and the SCAN for integrating goals, physiology and body movement.

Behavioral responses of solicitors after failure to receive grooming in Macaca fuscata.

Grooming is one of the most common cooperative behaviors among several animal species. However, the tactics used to cope with uncooperative partners in grooming interactions remain unclear. Japanese macaques (Macaca fuscata) solicit grooming from partners through postural behaviors, but may not necessarily receive grooming. This study investigated the behavior of female Japanese macaques after they solicited but did not receive grooming. We predicted that unsuccessful solicitors would engage in grooming interactions with uncooperative partners if they were affiliated. If they were not affiliated, the solicitors would not do so and may seek grooming interactions with other grooming partners. We used a focal-animal sampling method, targeting 17 females at Katsuyama, Okayama Prefecture, Japan. We recognized affiliative relationships by measuring close spatial association. After unsuccessful solicitation, females tended to scratch themselves, suggesting that solicitors may experience anxiety or distress when they do not receive grooming. They also tended to be proximate with affiliated partners after solicitation, regardless of whether the solicitors received grooming from their partners. In contrast, when solicitors failed to receive grooming from unaffiliated partners, their subsequent proximity was lower than when they were groomed. Moreover, unsuccessful solicitors were likely to engage in grooming interactions with affiliated partners who were uncooperative (receivers of unsuccessful solicitations). However, they were less likely to engage in grooming interactions with unaffiliated partners and instead engaged in grooming interactions with other nearby partners. These findings indicate that female Japanese macaques decide whether to engage in grooming interactions with uncooperative partners who have not groomed solicitors based on affiliative relationships and the availability of other grooming partners. It is probable that, when the cost of searching for a grooming partner is low, female Japanese macaques are likely to switch partners, potentially leading to an increase in the benefits obtained from grooming interactions.

RECTAL PROLAPSE IN THE SULAWESI CRESTED BLACK MACAQUE (MACACA NIGRA): MORBIDITY, MORTALITY, AND RISK FACTORS.

Empirical data suggest that rectal prolapse (RP) is common in captive Sulawesi crested black macaques (Macaca nigra) in Europe, resulting in the euthanasia of animals that experience repeat occurrences. However, the prevalence, etiology, and risk factors of RP remain unidentified. The aims of this retrospective study were to assess the morbidity and mortality of RP, to provide an overview of management practices, and to identify risk factors for RP in this species. A questionnaire was sent to all European Ex situ Programme institutions that housed M. nigra between 01 January 2014 and 31 December 2020. Zoological Information Management System medical records and the studbook were used to obtain additional information. The questionnaire had a response rate of 65%, accounting for 204 animals. Of these animals, 25 (12.3%) suffered from at least one RP event during the study period and recurrence was noted in 72%. The majority of prolapses reverted naturally, but 28% of afflicted animals were euthanized for this ailment. Institutions with M. nigra with high frequencies of diarrhea (P= 0.035), those that provided diets of ≥90% vegetables and high-fiber pellet (P < 0.001), and those with more male than female M. nigra (P < 0.001) had increased odds of RP. Institutions that provided fruits daily (P < 0.002) had reduced odds of having RP cases. Although correlation of RP with diet was identified, confounding cannot be excluded, and a detailed dietary analysis needs to take place before altering feeding practices. Acute stressors and detection of protozoa in fecal samples were common findings before an RP event. Demographic analysis indicated that aged females, young males, and subordinate individuals were most affected by this condition. Where tested during an RP intervention, animals had low serum levels of vitamin D. Pedigree analysis hinted at genetic predisposition in this species and requires further investigation.

Using behavior and genital swellings to monitor social dynamics and track reproductive cycling in zoo-housed lion-tailed macaques (Macaca silenus).

Though lion-tailed macaques are managed ex situ as an assurance population, little information is available on the relationship between behavior and reproductive physiology to ensure successful management and evaluate welfare. To facilitate social and reproductive management in a group of one male and three female lion-tailed macaques at Disney's Animal Kingdom®, we observed behaviors associated with dominance and reproduction, rated genital swellings, recorded menses, tracked ovarian cycles as determined by measurement of fecal immunoreactive progesterone (P4) and estradiol (E2), and monitored fecal cortisol (CORT). We analyzed the dominance hierarchy of females, compared hormone patterns of ovarian cycles with genital swellings and copulations relative to menses, and tested for correlations between hormones and behavior. The strength of the dominance hierarchy indicated moderate linearity, but relaxed after the dominant female received anxiolytic medications to reduce intraspecific aggression. Fecal P4 and E2 patterns revealed ovarian cycle lengths averaging 29.7 ± 0.8 days. The timing of single-day E2 peaks varied between and within females' cycles. However, on average, E2 peaks occurred 6-10 days after the first day of menses, with maximum genital swellings and copulations occurring slightly thereafter in uncontracepted females. Female-to-female aggression and sexual behaviors with the male were related to ovarian cycle phase, but correlations between behaviors and hormones varied between females. Initiated aggression was positively correlated with E2 in the dominant and mid-ranked females, while aggression received was positively correlated with P4 or negatively correlated with E2 in the mid-ranked and subordinate females. In one uncontracepted female, sexual behaviors with the male were negatively correlated to P4. CORT was positively correlated to aggression in the dominant and mid-ranked females and negatively correlated to grooming in the subordinate female. Results indicate observations of visual and behavioral cues can be used to track ovarian cycles and evaluate social dynamics and welfare in zoo-housed lion-tailed macaques.

Older Barbary macaques show limited capacity for self-regulation to avoid hazardous social interactions.

According to the Strength-and-Vulnerability-Integration (SAVI) model, older people are more motivated to avoid negative affect and high arousal than younger people. To explore the biological roots of this effect, we investigate communicative interactions and social information processing in Barbary macaques (Macaca sylvanus) living at 'La Forêt des Singes' in Rocamadour, France. The study combines an analysis of the production of (N = 8185 signals, 84 signallers) and responses to communicative signals (N = 3672 events, 84 receivers) with a field experiment (N = 166 trials, 45 subjects). Here we show that older monkeys are not more likely to specifically ignore negative social information or to employ avoidance strategies in stressful situations, although they are overall less sociable. We suggest that the monkeys have only a limited capacity for self-regulation within social interactions and rather rely on general avoidance strategies to decrease the risk of potentially hazardous social interactions.

[Posture Control Associated with Quadrupedal and Bipedal Gait in Japanese Monkeys (Macaca fuscata)].

How the CNS deals with instability of upright posture is the core in the control of bipedal gait. In this review, we summarize our recent findings comparing kinematics and EMG activity during quadrupedal and bipedal gait in Japanese macaques. Trunk/hindlimb muscles showed step cycle-modulated activity, which was more active in bipedal than in quadrupedal gait. For bipedal gait, enhanced activity during longer double support phase was predominantly observed in distal hindlimb muscles. Alternate burst activity in bilateral back muscles cyclically brought back the tilted trunk. In monkeys' quadrupedal gait, hindlimbs formed functional pairs with contralateral forelimbs, unlike in non-primate quadrupeds. These diagonal pairs acted differently on movements of the center of mass (COM). For bipedal gait, the hindlimbs solely carried the COM. Our results suggest that, compared to non-primate quadrupeds, hindlimbs in macaques contribute more critically to weight support and balance control even for quadrupedal gait. Additionally, for more unstable bipedal gait, the monkeys' CNS reinforces such hindlimb roles and actively controls the trunk posture in maintaining dynamic balance, in a manner similar to humans. Studies on Japanese macaques will further our understanding of the neural basis for the control of gait in mammals by bridging non-primate quadrupeds and humans.

Chemical cues of identity and reproductive status in Japanese macaques.

Olfactory communication plays an important role in the regulation of socio-sexual interactions in mammals. There is growing evidence that both human and nonhuman primates rely on odors to inform their mating decisions. Nevertheless, studies of primate chemical ecology remain scarce due to the difficulty of obtaining and analyzing samples. We analyzed 67 urine samples from five captive female Japanese macaques (Macaca fuscata) and 30 vaginal swabs from three of these females using gas chromatography-mass spectrometry and examined the relationship between odor (compounds identified, richness, intensity, and diversity) and female identity as well as cycle phase. We found a total of 36 urine compounds of which we identified 31, and 68 vaginal compounds of which we identified 37. Our results suggest that urine and vaginal odor varied more between individuals than within cycle phases. However, we found that within a female cycle, urine samples from similar phases may cluster more than samples from different phases. Our results suggest that female odor may encode information about identity (vaginal and urine odor) and reproductive status (urine odor). The question of how conspecifics use female urine and vaginal odor remains open and could be tested using bioassays. Our results and their interpretation are constrained by our limited sample size and our study design. Nonetheless, our study provides insight into the potential signaling role of female odor in sexual communication in Japanese macaques and contributes to our understanding of how odors may influence mating strategies in primates.

When males have females on their backs: Male's tolerance, solicitation, and use of female-male mounting in Japanese macaques.

Previous research on Japanese macaques has shown that female-to-male mounting (FMM) is performed by some females as an exaggerated form of sexual solicitation that may occur in the context of high female competition for male mates. This supernormal courtship behavior functions to prompt subsequent male-to-female mounting. In this report, we focused on the male consort partners' responses to FMM. We studied a free-ranging population of Japanese macaques at Arashiyama, Japan, in which FMM is frequent and prevalent. We analyzed 240 consortships involving 31 females and 19 males. We tested three hypotheses regarding male's tolerance, solicitation, and use of FMM. First, we found that FMM was tolerated by male mountees who were no more likely to aggress their female partners during a short time window around a FMM than they were during the rest of the consortship period. Second, we showed that FMM could be triggered by male recipients, via explicit male-to-female sexual solicitations. Third, we found that some males may utilize FMM in a quest for their own sexual stimulation, which sometimes culminated in masturbation by the male during FMM. Our findings indicate that male partners facilitate the expression of FMM both passively (via their tolerance) and actively (via their solicitation). In addition, FMM appears to enhance the sexual arousal of male partners during consortships. We argued that, for females to have expanded their repertoire of sexual solicitations by adopting FMM, male mates must have played a role in the evolutionary origins and maintenance of this nonconceptive but intense and powerful female mating tactic.