Daytime spatial relationships in a family group of black rhinoceros Diceros bicornis at the Hiroshima City Asa Zoological Park, Japan.

Although black rhinoceros Diceros bicornis are mostly solitary in the wild, the Hiroshima City Asa Zoological Park (Asa Zoo) has kept a family group together during the daytime, with good reproductive performance over five decades. Management procedures at the zoo include temporary single housing of the mother before and after giving birth, which facilitates maintenance of a compatible family group. We recorded intra-group spatial relationships for 4 years and 4 months, during which time an adult female reared two consecutive calves. During daytime she remained in an enclosure with her new calf, one or two older offspring, and an adult male, the sire of all her offspring. Proximity (within two adult body-lengths) scores between the mother and her two calves were especially high during the first year after birth, and only slightly lower for her older offspring. The adult male had the lowest proximity scores. The spatial relationships were visualized by applying multidimensional scaling (MDS) to the proximity scores. Mother and calves were plotted close to each other, with older offspring slightly farther apart on the two-dimensional MDS representation; the adult male was more distant from the other group members. These findings indicate clear follower-type characteristics in the mother-calf pair and also older immature offspring, albeit to a lesser degree. Although black rhinoceros are generally solitary in the wild, our results duplicate observations of some wild black rhinoceros groups containing an adult female, her calf, and an older immature, with adult males being largely solitary.

Factors affecting captive female giraffe stress response: Male presence, small enclosure, and low temperature.

To improve animal welfare based on suitable social housing conditions, it is important to understand the factors that trigger high-stress responses. Wild giraffes live in a fission-fusion society and males and females are rarely in the same herd for a long period. The captive condition of belonging to a herd with the same individuals for months or years is uncommon in nature. To understand the effect of male presence on female stress levels, fecal glucocorticoid metabolite (fGCM) levels and social interactions in two captive female giraffes were investigated. Additionally, the effect of enclosure size and temperature on fGCM level and social interactions were examined. The results showed no significant difference in the fGCM levels of females based on male presence. The frequency of agonistic behavior by the dominant female toward the subordinate female was significantly increased when a male was present. The subordinate female was significantly less likely to approach the dominant female and showed decreased affiliative and agonistic interactions toward the dominant female when a male was present. The frequencies of agonistic interactions between females were higher in the small enclosure regardless of male presence. Low temperature triggered higher fGCM levels and increased agonistic interaction in an aged female. The findings of this study suggest that these multiple factors should be considered individually to promote the welfare of captive giraffes.

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.

Male care in a free-ranging group of Japanese macaques (Macaca fuscata).

Among the Macaca species, adult male Japanese macaques are the least likely to perform male care (i.e., affiliative interactions between adult males and immatures, including holding, carrying, and grooming); however, they perform male care for infants, albeit infrequently. We examined 17 cases of male care observed for the first time when the immature was younger than 1 year of age. Eleven of the 31 adult males who remained as central males during the 30-year observation period performed male care. Their age and dominance rank did not influence the occurrence of male care. Most cases were first recorded between the last part of the mating season and the first part of the birth season (January-March), whereas male care was rarely observed during the mating season (October-December). In 12 of the 17 cases, male care ceased within 6 months after the first observation, whereas in the remaining cases, it continued for at least 1 year. In 15 of the 17 cases, males tended to perform male care for matrilineally unrelated female infants of low-ranking mothers. In some cases, the male and infant mother showed grooming interactions for 6 months both before and after the start of male care, whereas such grooming interactions were never recorded either before or after the start of male care in other cases. We also examined some hypotheses on male-immature associations and the probable benefits that males and infants might acquire through male care.

Professor Masao Kawai, a pioneer and leading scholar in primatology and writer of animal stories for children.

This Editorial is dedicated to Professor Masao Kawai (Fig. 1), who passed away on May 14th, 2021. He served as the sixth Editor-in-Chief of Primates for 15 years (1981-1995). The following is a collection of memories from 20 scholars. Fig. 1 Professor Masao Kawai, a pioneer and leading scholar in primatology (photo taken in January 2020; Courtesy of Ryoko Kawai).

Triadic grooming among adult females in a free-ranging group of Japanese macaques.

This study aimed to quantitatively describe triadic grooming bouts in a free-ranging group of Japanese macaques (Macaca fuscata). Two types of triadic grooming bouts occurred less frequently and for a shorter duration than dyadic grooming bouts: straight-line type (SL), where individual A grooms B, who then grooms C; and two-to-one type (TO), where individuals X1 and X2 groom Y. TO-type grooming was recorded more often than SL-type grooming. As in the dyadic grooming bouts, two females who had direct grooming interactions in both SL-type (i.e., between A and B or between B and C) and TO-type groomings (i.e., between X1 and Y and between X2 and Y) and two groomers who did not have direct grooming interactions in TO-type grooming were closely related to each other in more than the half of the pairs recorded. Groomers were more likely to be subordinate to groomees in triadic grooming. Almost all of the observed pairs in SL- and TO-type groomings were also recorded in dyadic grooming bouts. These findings indicate that like dyadic grooming, triadic grooming bouts are largely influenced by blood relatedness through maternal lines and dominance relationships between participants. Based on affiliative relationships maintained through dyadic grooming, triadic grooming could be influenced by such affiliative relationships and then function to strengthen bonds between participants. Possible relationships between triadic grooming and the level of social tolerance among individuals are discussed from the viewpoints of interspecies differences among macaque species and regional differences in Japanese macaques.

High but not low tolerance populations of Japanese macaques solve a novel cooperative task.

Currently, it has been revealed that high levels of tolerance facilitate the occurrence of cooperative behavior in animals. This predicts that Japanese macaques (Macaca fuscata) would fail to solve cooperative food-retrieval tasks because of their low level of social tolerance. However, Japanese macaques exhibit regional differences in their levels of tolerance. In this study, we showed how inter-group difference in social tolerance could affect cooperation, by presenting a well-established cooperative rope-pulling task with two free-ranging groups of Japanese macaques that exhibit different levels of social tolerance. We used the task that required two macaques to pull both ends of a single rope simultaneously to obtain food rewards. We found that some macaques from the more tolerant group successfully solved the task, and one of them learned to wait for a partner when a partner was absent. In contrast, however, those of the less tolerant group almost never succeeded in the task. These results indicate that Japanese macaques possess the abilities to cooperate with conspecifics, but such abilities may be constrained in their typically despotic society.

Temporal adjustment of short calls according to a partner during vocal turn-taking in Japanese macaques.

Turn-taking is a common feature in human speech, and is also seen in the communication of other primate species. However, evidence of turn-taking in vocal exchanges within a short time frame is still scarce in nonhuman primates. This study investigated whether dynamic adjustment during turn-taking in short calls exists in Japanese macaques Macaca fuscata. We observed exchanges of short calls such as grunts, girneys, and short, low coos during social interactions in a free-ranging group of Japanese macaques. We found that the median gap between the turns of two callers was 250 ms. Call intervals varied among individuals, suggesting that call intervals were not fixed among individuals. Solo call intervals were shorter than call intervals interrupted by responses from partners (i.e., exchanges) and longer than those between the partner's reply and the reply to that call, indicating that the monkeys did not just repeat calls at certain intervals irrespective of the social situation. The differences in call intervals during exchanged and solo call sequences were explained by the response interval of the partner, suggesting an adjustment of call timing according to the tempo of the partner's call utterance. These findings suggest that monkeys display dynamic temporal adjustment in a short time window, which is comparable with turn-taking in human speech.

Vocalizations during post-conflict affiliations from victims toward aggressors based on uncertainty in Japanese macaques.

We investigated the use of vocalizations called "grunts," "girneys," and "coos" accompanied by post-conflict affiliative interaction between former opponents (reconciliation) in Japanese macaques (Macaca fuscata). Although reconciliation functions to repair bonds, such interactions sometimes entail risks of receiving further aggression. Vocalizations can be used at a distance from the former opponent; thus, we predict that vocalizations are used particularly by victims of a conflict, and are frequently used in situations of uncertainty when it is difficult for them to estimate whether the former opponent will resume aggression. In addition, we predict that vocalizations are effective in preventing further aggression. To test these hypotheses, we conducted observations of post-conflict and matched-control situations in female Japanese macaques living in a free-ranging group. We found that former opponents tended to be attracted to each other within the first minute following a conflict, thus demonstrating reconciliation behavior. Vocalizations were more frequently used by the victims in post-conflict interactions than under control situations; however, this tendency was not found in aggressors. When affiliation with the former opponent occurred, victims were more likely to use vocalizations towards less familiar opponents. These findings suggest that Japanese macaques used vocalizations more often when interacting with less predictable former opponents. Victims were more likely to receive aggression from former aggressors when engaged in affiliations with them than under no such affiliations. No significant differences were found in the probability of the victims receiving aggression, regardless of whether they used vocalizations; thus, whether the victim benefits from using vocalizations in these contexts remains unclear. Japanese macaques form despotic societies and therefore, further aggression was inevitable, to some degree, after a conflict. The use of vocalizations by a victim was found to depend on the nature of their relationship with the aggressor; however, the effectiveness of this behavior requires further investigation.

Japanese macaque (Macaca fuscata) mothers huddle with their young offspring instead of adult females for thermoregulation.

It is unclear whom animals select to huddle with for thermoregulation. In this study, we investigated whom Japanese macaque (Macaca fuscata) mothers huddled with-their young offspring or other adult group members-when there is need for thermoregulation. We used a focal-animal sampling method, targeting 17 females at Katsuyama, Okayama Prefecture, Japan. A majority of huddling among adult females was recorded during winter season (December, January, and February). Females who had young (0- or 1-year-old) offspring huddled less frequently with other adult females compared to females who did not have young offspring in winter. However, including young offspring, the frequency of huddling with any other individuals did not differ by whether females had young offspring. Moreover, the females who did not have young offspring huddled with other adult females more often in cloudy than in sunny weather during winter season. In contrast, females who had young offspring increased huddling with their young offspring in cloudy than in sunny weather, but did not do so with other adult females. This study indicates that Japanese macaque mothers huddle with their young offspring instead of other adult females when there is need for thermoregulation.