Mirror responses in a Japanese macaque troop (Arashiyama West)

Behavior toward two mirrors in the field was observed in the Arashiyama West troop ofMacaca fuscata. Counts of visits to the mirrors, plus scan and focal animal sampling were used. Some animals were marked with fluorescent paint to test informally for self-recognition. A relatively high mean frequency of visits to the reflecting side of both mirrors by all age classes, ranks, and sexes was recorded. There was no age difference in frequency of mirror visits per sample but adults spent more time per visit than subadults who in turn spent more time than juveniles. There was no indication of self-recognition by paint-marked animals. Mirrors appeared to be used to monitor the reflected scene and to look at the self-image. Social behavior in the mirror zone that was not directed toward the mirror was common to all age classes. Species-typical behavior directed toward the mirror was seen in younger animals but very seldom in adults. No threat displays by any animal were observed. We suggest that for adults the mirror image was not seen simply as another monkey.     

Monkeys with mirrors: Some questions for primate psychology

Monkeys do not appear to recognize themselves in mirrors but display social responses to the reflection. This article comprehensively reviews the literature concerning monkeys’ reactions to mirrors, describing the interest shown in reflections, the reinforcing effects of mirror-image stimulation, the social responses of the monkeys, their interpretation of objects reflected in a mirror, and their performance in tests of self-recognition. In all cases limitations of the methods employed restrict conclusions about the monkeys’ behavior. Detailed behavioral observations coupled with comparisons of the psychological properties of mirrors with those of other stimuli promise to reveal more useful information about cognitive mechanisms and abilities of monkeys and other primates.     

Not all chimpanzees (Pan troglodytes) show self-recognition

When a chimpanzee is presented with a mirror it initially responds with social behavior directed toward the reflection. After several hours of exposure to the mirror the social behavior decreases and the mirror is used to guide self-directed responses to previously unobservable parts of the body such as the face. When a distinctively colored mark is unobtrousively applied to the chimpanzee's face and the chimpanzee touches the mark while observing itself in the mirror, this behavior is said to indicate self-recognition. Such self-recognition has been considered to be a robust phenomenon in chimpanzees, with self-directed and mark-directed behaviors both appearing in all socially-housed adult chimpanzees tested. In our study 11 chimpanzees were given mirror exposure and tested with the mark test. Only one of the 11 chimpanzees touched the mark during test, although several showed self-directed behavior using the mirror to guide their movements. Such experimental factors as mirror size, position, or temporal spacing of the mirror exposure, and such subject variables as age, sex, previous social experience, and subspecies were insufficient to explain the difference between the present and previous findings. We suggest that there are individual differences in mirror recognition behavior in chimpanzees, and that further consideration of the factors contributing to this phenomenon, including the development of additional tests for self-recognition, is needed.     

Mirror responses in a group of Miopithecus talapoin

Studies of non-human primate self-recognition in mirrors demonstrate variation both within and between species. This study applied a rigorous methodology that took into account habituation of subjects to the mirror as an object and to the experimental situation. The species observed in our study was Miopithecus talapoin, which has been little studied in the wild or in captivity. Although this species shows several interesting characteristics, including complex social organisation and a high encephalization index, the talapoin monkeys in the study did not pass the mark test; however, they showed a prerequisite for self-recognition, namely comparing their body parts to the image of these in the mirror.     

Social Monitoring Reflects Dominance Relationships in a Small Captive Group of Brown Capuchin Monkeys (Cebus apella)

In several studies of social monitoring in primates, subordinate animals directed more visual attention toward dominant animals than vice versa. This behavior is thought to enable subordinate animals to avoid conflict. We sought to clarify whether visual attention behavior functions in this manner in a small captive group of brown capuchin monkeys, Cebus apella. We tested the hypothesis that social monitoring is related to dominance status. Dominance status was determined based on the directionality of aggressive behavior, and visual attention was quantified by using focal animal sampling. Subordinate animals directed significantly more visual attention toward others than dominant animals. Subordinate animals also looked more frequently at the animals that attacked them and others the most. The results indicate that social monitoring behavior in this captive group was driven by conflict‐avoidance.     

Function of peering behavior among bonobos (Pan paniscus) at Wamba,Zaire

Peering behavior (prolonged gazing within 30 cm by an animal toward another) in wild bonobos (Pan paniscus) at Wamba, Zaire, was studied. A total of 230 peering episodes were observed in various social contexts. Peering behavior was often directed from younger animals toward older ones. In particular, adult females were most frequently involved in peering, with individuals of all age-sex classes. On the other hand, male bonobos seldom took part in peering behavior. Four types of behavior patterns followed the peering behavior: (1) the peerer left; (2) the peeree left; (3) both peerer and peeree stayed but had no further social interaction; and (4) some other social interaction followed. Type (1) was the most frequent. Peering usually led to tolerance by older (dominant) animals of a younger (subordinate) animal’s subsequent actions directed towards the former. Peering was thus concluded to be a unilateral action for initiating affinitive interactions by the peerer.     

A ten-month study of endogenous testosterone levels and behaviour in outdoor-living female rhesus monkeys (Macaca mulatta)

Endogenous testosterone levels were measured in association with sexual, aggressive, and social/affiliative behaviors in 11 outdoor-housed female rhesus monkeys over a ten-month period. Several behaviors (sex directed toward the male, sex received from the male, aggression directed toward the male, submission directed toward the male, submission directed toward the female, and groom another female) were significantly (p<0.05) positively correlated with testosterone in from one to five females. No trends were strong enough across all females to suggest that any of these correlations have species-wide significance. Factor analysis revealed clearcut clusters of behaviors, but elevations in testosterone were not strongly associated with any of these clusters. It is concluded that endogenous testosterone levels have little measurable effect on overt behavior in female rhesus monkeys.     

Mirror,Mirror on the Wall,How Does My Brain Recognize My Image at All?

For decades researchers have used mirrors to study self-recognition. However, attempts to identify neural processes underlying this ability have used photographs instead. Here we used event related potentials (ERPs) to compare self-face recognition in photographs versus mirrors and found distinct neural signatures. Measures of visual self-recognition are therefore not independent of the medium employed.     

Reflections in the rainforest: full-length mirrors facilitate behavioral observations of unhabituated,wild chimpanzees

We describe behaviors of unhabituated wild chimpanzees in Gabon during repeated encounters with large mirrors installed permanently in their home range. Movement in proximity to the mirrors triggered video cameras that recorded the scene. Data are presented for 51 mirror encounters spanning a 3-year period. After initial wariness, mirror-directed aggressive behaviors were common, especially in adult males, but aggression gradually diminished and eventually almost completely ceased. Focusing on the two mirrors that elicited most reactions, the percentage of chimpanzees showing tension or anxiety also decreased across encounters. These mirrors elicited a range of socio-sexual behaviors interpreted as having a reassurance function, especially when group-level tension appeared high. Chimpanzees also occasionally directed these behaviors towards their own reflection. Despite increasing habituation and positive attraction to the mirrors, none of the chimpanzees displayed signs of self-recognition. We conclude that a combination of large mirrors and video traps can provide valuable information about unhabituated, semi-terrestrial primates in their natural habitat, by inducing the primates to stay in one place for longer than they might otherwise do.     

Mirror-image responses in pygmy marmosets (Cebuella pygmaea)

Only a few nonhuman species (chimpanzees and orangutans) have displayed mirror-image recognition of themselves by grooming at a spot that can only be seen with the mirror. Pygmy marmosets have never been observed to self-groom, but they do behave toward mirrors in a manner suggestive of the early stages of mirror-image recognition. They displayed a rapid extinction of social threat responses to their own image and of novelty responses to mirrors, but continued to show mirror-specific responses such as following their own image, playing peek-a-boo, and looking at their image throughout a 28-day period of mirror exposure. The pygmy marmosets used a mirror to locate otherwise unseen conspecifics from other groups and directed threat responses toward the real location of these animals rather than to their mirror-image. Pygmy marmosets displayed the precursor behaviors to mirror-image recognition.     

Mirror-Directed Social Behaviors of Garnett's Greater Bush Baby (Otolemur garnettii)

Many diurnal anthropoid species direct social behaviors toward their own mirror-image as though viewing a conspecific. To determine whether a nocturnal prosimian species would behave similarly, we videotaped social responses of 45 Garnett's greater bush babies (Otolemur garnettii) observing mirror-images for the first time, scored them for frequency and duration, and compared them with the same behaviors directed elsewhere in the test apparatus. Males scentmarked more than females did, principally with the hindfoot, and most when in immediate proximity to the mirror. Bush babies displayed bipedal posture and threat gestures when oriented directly toward a mirror from a near position. Orientation toward the mirror also increased the frequency of arched-back postures; however, this behavior was not contingent on proximity to the mirror or visibility of the mirror-image. The differential expression of specific behaviors toward mirror-images by male and female bush babies supports the view that this nocturnal prosimian, thought to be dependent on olfaction and audition for communication, is capable of specific recognition by visual cues alone.     

Mirror-mediated responses of California scrub jays (Aphelocoma californica) during a caching task and the mark test

Mirror self-recognition, as an index of self-awareness, has been proposed as a precursor for more complex social cognitive abilities, such as prosocial reasoning and cooperative decision-making. Indeed, evidence for mirror self-recognition has been shown for animals possessing complex social cognitive abilities such as great apes, dolphins, elephants and corvids. California scrub jays (Aphelocoma californica) have provided strong evidence that non-human animals are capable of mental state attribution. For instance, scrub jays are reported to use their experience stealing the food of others to infer that other birds may similarly intend to steal from them. If a concept of “self” is required for such complex social cognitive abilities, then scrub jays might be expected to show mirror self-recognition. Thus, we examined whether California scrub jays are capable of mirror self-recognition using two experimental contexts: a caching task and the mark test. During the caching task, we compared the extent to which scrub jays protected their food after caching alone, in the presence of a conspecific and in the presence of a mirror. The birds did not engage in more cache protection behaviours with a mirror present than when caching alone, suggesting scrub jays may have recognized their reflection and so did not expect cache theft. Alternative explanations for this behaviour are also discussed. During the mark test, the scrub jays were surreptitiously marked with a red or plumage-coloured control sticker. The scrub jays showed no evidence of mirror self-recognition during the mark test, as the birds did not preferentially attempt to remove the red mark in the presence of a mirror. Together, the results provide mixed evidence of the mirror self-recognition abilities of California scrub jays. We highlight the need to develop alternative approaches for evaluating mirror self-recognition in non-human animals to better understand its relationship with complex social cognition.     

Male chimpanzee behavior in relation to female ano-genital swelling

This study examined the relationships between male agonistic, affiliative, and sexual behaviors and female estrus condition in captive adolescent and young-adult chimpanzees (Pan troglodytes). Data on agonistic, affiliative, and sexual behaviors of 11 males living in three social groups were collected during daily 45 minute observations over a 5 month period. Female estrus condition was assessed daily using the relative size of the female's ano-genital swelling. It was hypothesized that the presence of maximally tumescent females would generate conflicts between males, so an increase in inter-male agonism was predicted. Males exhibited higher rates of agonism toward other males when at least one female in the group was maximally tumescent. Male affiliative behavior directed toward other males and social play with males were affected by the presence and number of maximally tumescent females. Male sexual behavior increased when maximally tumescent females were present.     

Sex Differential Behavior of Immature Hanuman Langurs (Presbytis entellus) in Ramnagar, South Nepal

We investigated sex differences in the social behavior of immature Hanuman langurs (Presbytis entellus) in the light of sex-specifically different life-courses and Hanuman langur characteristics, such as the individualistic dominance hierarchy and the rarity of intragroup coalitions among adult females. We observed four immature female and four immature male langurs—all members of the same free-ranging multimale multifemale group in Ramnagar, South Nepal—from November 1992 to February 1993 for 288 hr via focal-animal and instantaneous sampling techniques. Immature females spent significantly more time in proximity to other group members than immature males did. They had more physical contact and groomed more. Other immature females were their preferred social partners. Immature males also preferred like-aged females. They restricted their relationships with other immature males to proximity and occasional grooming. Monitoring was directed especially toward adult males. Female behavior can be interpreted as oriented toward integration into the female social network and their age-inverted dominance hierarchy. Males seem to prepare for leaving their natal group and for future strong intrasexual competition.     

False friends are worse than bitter enemies: “Altruistic” punishment of in-group members

One of the most critical features of human society is the pervasiveness of cooperation in social and economic exchanges. Moreover, social scientists have found overwhelming evidence that such cooperative behavior is likely to be directed toward in-group members. We propose that the group-based nature of cooperation includes punishment behavior. Punishment behavior is used to maintain cooperation within systems of social exchange and, thus, is directed towards members of an exchange system. Because social exchanges often take place within groups, we predict that punishment behavior is used to maintain cooperation in the punisher's group. Specifically, punishment behavior is directed toward in-group members who are found to be noncooperators. To examine this, we conducted a gift-giving game experiment with third-party punishment. The results of the experiment (N=90) support the following hypothesis: Participants who are cooperative in a gift-giving game punish noncooperative in-group members more severely than they punish noncooperative out-group members.     

Mirror-Mark Tests Performed on Jackdaws Reveal Potential Methodological Problems in the Use of Stickers in Avian Mark-Test Studies

Some animals are capable of recognizing themselves in a mirror, which is considered to be demonstrated by passing the mark test. Mirror self-recognition capacity has been found in just a few mammals having very large brains and only in one bird, the magpie (Pica pica). The results obtained in magpies have enormous biological and cognitive implications because the fact that magpies were able to pass the mark test meant that this species is at the same cognitive level with great apes, that mirror self-recognition has evolved independently in the magpie and great apes (which diverged 300 million years ago), and that the neocortex (which is not present in the bird''s brains) is not a prerequisite for mirror self-recognition as previously believed. Here, we have replicated the experimental design used on magpies to determine whether jackdaws (Corvus monedula) are also capable of mirror self-recognition by passing the mark test. We found that our nine jackdaws showed a very high interest towards the mirror and exhibited self-contingent behavior as soon as mirrors were introduced. However, jackdaws were not able to pass the mark test: both sticker-directed actions and sticker removal were performed with a similar frequency in both the cardboard (control) and the mirror conditions. We conclude that our jackdaws'' behaviour raises non-trivial questions about the methodology used in the avian mark test. Our study suggests that the use of self-adhesive stickers on sensitive throat feathers may open the way to artefactual results because birds might perceive the stickers tactilely.     

Infant stumptailed macaques reared with mirrors or peers: Social responsiveness,attachment, and adjustment

Comparisons of activity toward mirrors and peers in infant macaques being reared with one of these stimuli as the primary rearing partner revealed markedly greater social responsiveness to a fully accessible cagemate than to one's own reflection. Measures of exploration, aggression, and especially play all revealed the cagemate to be the more potent social stimulus. Mirror-reared infants given additional experience of a live peer behind a transparent partition were less responsive to the mirror than were infants with no social stimulation other than a mirror. In contrast, cagemate-directed behavior of peer-reared infants was not seriously affected by additional exposure to a mirror. A fully accessible peer also elicited more social responding than a peer behind a transparent partition, and infants with experience of both a live cagemate and mirrors were generally more responsive toward the former. Greater agitation in peer-reared than in mirror-reared stumptailed monkeys during separations from their rearing partners suggests that exposure to the physically accessible partner led to stronger attachments. Infants reacted positively to a moderately unfamiliar environment but showed behavioral disruption when placed in a very unfamiliar environment. Disruption was especially evident in peer-reared infants, in which exposure to the unfamiliar environment was compounded with the absence of the attachment figure. Mirror-rearing appeared to reduce the tendency toward ‘isolation syndrome’ behaviors compared to alone-rearing, and these behaviors appear to be less common in stumptailed than in rhesus macaques.     

Responses of capuchin monkeys (Cebus apella) to different conditions of mirror-image stimulation

Group-living brown capuchins were given mirror-image stimulation as follows: (1) mirror 1 m away; (2) mirror attached to the cage-mesh; (3) angled mirrors creating a deflected image; (4) small mirror in the cage; and (5) small, transportable mirrors. The subjects were initially interested in the mirrors in each condition, but they generally habituated to them over the course of repeated presentations. Control (non-reflective) objects were attended to less than mirrors. Facial expressions occurred mostly in Condition 2, lateral and vertical head movements in Condition 3, reaching behind the mirror in Condition 2, looking obliquely into the mirror in Conditions 4 and 5. Despite these diverse conditions of exposure to mirrors, lasting for a total of over three months, no behaviours suggesting self-recognition were seen in the monkeys.     

四川短尾鼩尿液气味在社会等级识别中的作用

本文在中立竞技场中通过两两互作确定四川短尾鼩(Anourosorex squamipes)同性个体间的社会等级,并在此基础上利用其尿液,研究不同社会等级个体的自我或非自我识别能力及模式、尿液气味的行为响应机制,以及社会等级识别能力。结果表明:(1)四川短尾鼩优势个体表现攻击行为较多,从属个体防御行为较多,优势个体的标记行为显著高于从属个体;攻击行为表现为同等级雄性高于同等级雌性,且雌性间的攻击强度低于雄性;(2)从属个体和优势个体分别对自身尿液气味和非自身尿液气味存在明显偏好差异;不同性别、等级个体自我识别模式差异不明显,不同社会等级个体对于自身识别模式和非自身尿液的行为反应模式均不同。不同社会等级个体具有自我识别能力且能力不同;(3)四川短尾鼩能够识别不同社会等级个体的尿液气味,雌性对雄性尿液更感兴趣,雄性对优势雄性尿液选择回避;雄性对其他个体的访问时间与嗅舔频次均显著高于雌性,雌雄个体在识别不同社会等级的尿液气味时存在性二型。     

Watch Your Tone: Social Conditions Modulate Singing Strategies

Bird song is typically depicted as a male singing a long‐distance signal to potentially unknown receivers to (1) deter males and (2) attract females. Nevertheless, many songbirds sing from close distances to a known receiver; males of these species may be under more intense selective pressure to modify their songs depending on the sex of the receiver in order to convey different motivational states (aggression versus courtship) to the different sexes. In a laboratory setting, we examined how receiver sex affected within‐song variation of the close‐range singing behavior in the brown‐headed cowbird (Molothrus ater). Although we know that cowbird song is influenced by flock composition, it is still unclear as to how the cowbird modifies his song based on social context. Using a cross‐correlation analysis of each male's different song types, we found that pairs of songs were significantly more dissimilar if they were directed to females compared with songs directed to males. We subsequently tested whether there were any consistent spectral or temporal patterns in the songs males gave to females versus to males. Our results lend support for the Motivational Structural Rules Hypothesis as songs directed toward males had higher entropy (i.e., harshness) than the same song type directed toward females. Our results suggest that cowbirds may have evolved the ability to alter multiple dimensions of their singing behavior based on receiver sex.