Personality heterophily and friendship as drivers for successful cooperation.

Cooperation is widespread and arguably a pivotal evolutionary force in maintaining animal societies. Yet, proximately, what underlying motivators drive individuals to cooperate remains relatively unclear. Since 'free-riders' can exploit the benefits by cheating, selecting the right partner is paramount. Such decision rules need not be based on complex calculations and can be driven by cognitively less-demanding mechanisms, like social relationships (e.g. kinship, non-kin friendships, dyadic tolerance), social status (e.g. dominance hierarchies) and personalities (social and non-social traits); however, holistic evidence related to those mechanisms is scarce. Using the classical 'loose-string paradigm', we tested cooperative tendencies of a hierarchical primate, the long-tailed macaque (Macaca fascicularis). We studied three groups (n = 21) in their social settings, allowing partner choice. We supplemented cooperation with observational and experimental data on social relationships, dominance hierarchies and personality. Friendship and dissimilarities in non-social 'exploration' and 'activity-sociability' personality traits predicted the likelihood of cooperative dyad formation. Furthermore, the magnitude of cooperative success was positively associated with friendship, low rank-distance and dissimilarity in the activity-sociability trait. Kinship did not affect cooperation. While some findings align with prior studies, the evidence of (non-social) personality heterophily promoting cooperation may deepen our understanding of the proximate mechanisms and, broadly, the evolution of cooperation.

The ability to inhibit impulses is related to social behavior in long-tailed macaques.

Performance in cognitive tasks has been linked to differences in species' social organization, yet to understand its function its relationship to within-species variation in behavior should also be explored. One important cognitive capacity, the ability to inhibit impulses, is typically better in egalitarian than despotic primate species and in primate species with strong fission-fusion dynamics. A different line of research indicates that a high ability to inhibit impulses is related to less aggressive behavior and more socio-positive behavior. However, within species the relationship between performance on cognitive inhibition tasks and variation in social behavior remains to be explored. Here we investigate how performance in a typical inhibition task in cognitive research is related to aggressive and socio-positive behavior in despotic long-tailed macaques. Twenty individuals living in two naturalistic mixed-sex groups were tested with the Plexiglass Hole Task. Aggressive behavior and three types of socio-positive behavior (neutral/friendly approaches, socio-positive signaling, and grooming others) among group members were measured. Individuals differed in their ability to inhibit impulses. Individuals that were not good at inhibiting impulses showed higher rates of aggressive behavior, but also more socio-positive signals, whereas inhibition was not related to neutral/friendly approaches and grooming. These results confirm the positive link between impulsiveness and aggression. In addition, the results indicate that some social-positive behavior may be enhanced when inhibition is limited. In this species, benefits potentially derived from aggression and socio-positive signals match a low ability to inhibit impulses, suggesting that a low ability to inhibit impulses may actually be advantageous. To understand differences between species in cognitive skills, understanding the benefits of variation in a cognitive capacity within a species is crucial.

External environmental conditions impact nocturnal activity levels in proboscis monkeys (Nasalis larvatus) living in Sabah, Malaysia.

Recently, several diurnal nonhuman anthropoids have been identified displaying varying degrees of nocturnal activity, which can be influenced by activity "masking effects"-external events or conditions that suppress or trigger activity, temporarily altering normal activity patterns. Environmental masking characteristics include nocturnal temperature, rainfall, cloud cover, and moon brightness. Similarly, other ecological characteristics, including proximity to humans and predators and daytime activity, may also trigger or suppress nocturnal activity. Understanding the effects of external conditions on activity patterns is pertinent to effective species conservation. We investigated the presence of nocturnal activity and the influence of masking effects on the level of nocturnal activity displayed by wild proboscis monkeys (Nasalis larvatus) in Sabah, Malaysian Borneo. Dual-axis accelerometers were attached by collar to six male proboscis monkeys from different one-male, multi-female groups to record activity continuously (165-401 days each). We measured the monkeys' nocturnal and diurnal activity levels and investigated the effects of seven potential masking effects. Nocturnal activity was much lower than diurnal activity. Still, proboscis monkeys did display varying levels of nocturnal activity. Generalized linear mixed models identified higher nocturnal activity in the study individuals during nights with cooler temperatures, higher rainfall, and after higher diurnal activity. These three masking effects affected nocturnal activity levels during the observation period that informed our model, although they did not predict nocturnal activity outside of this period. While the generalizability of these results remains uncertain, this study highlights the utility of accelerometers in identifying activity patterns and masking effects that create variability in these patterns.

Tolerant and intolerant macaques show different levels of structural complexity in their vocal communication.

We tested the social complexity hypothesis which posits that animals living in complex social environments should use complex communication systems. We focused on two components of vocal complexity: diversity (number of categories of calls) and flexibility (degree of gradation between categories of calls). We compared the acoustic structure of vocal signals in groups of macaques belonging to four species with varying levels of uncertainty (i.e. complexity) in social tolerance (the higher the degree of tolerance, the higher the degree of uncertainty): two intolerant species, Japanese and rhesus macaques, and two tolerant species, Tonkean and crested macaques. We recorded the vocalizations emitted by adult females in affiliative, agonistic and neutral contexts. We analysed several acoustic variables: call duration, entropy, time and frequency energy quantiles. The results showed that tolerant macaques displayed higher levels of vocal diversity and flexibility than intolerant macaques in situations with a greater number of options and consequences, i.e. in agonistic and affiliative contexts. We found no significant differences between tolerant and intolerant macaques in the neutral context where individuals are not directly involved in social interaction. This shows that species experiencing more uncertain social interactions displayed greater vocal diversity and flexibility, which supports the social complexity hypothesis.

Low relationship quality predicts scratch contagion during tense situations in orangutans (Pongo pygmaeus).

Primates show various forms of behavioral contagion that are stronger between kin and friends. As a result, behavioral contagion is thought to promote group coordination, social cohesion, and possibly state matching. Aside from contagious yawning, little is known about the contagious effect of other behaviors. Scratching is commonly observed during arousal and as such may play a role within group dynamics. While the Bornean orangutan (Pongo pygmaeus) is commonly considered the least social great ape, orangutans do engage in social interactions. Therefore, their social organization makes them a suitable case for studying the social function of behavioral contagion. Through behavioral observations of captive orangutans, we recorded all yawn and scratch events together with the corresponding behavior of all bystander group-members. As yawning was rarely observed, no conclusions could be drawn regarding this behavior. Scratching was contagious and occurred within 90 s after the triggering scratch. Specifically, orangutans showed increased scratch contagion when they had seen a weakly bonded individual scratch during tense contexts. When the orangutan had not seen the triggering scratch, the contagiousness of scratching was not affected by context or relationship quality. Our results indicate that behavioral contagion is not simply higher between individuals with stronger social relationships, but that the contagiousness of behaviors may vary based on the context and on social factors. We discuss these findings in light of an adaptive function that may reduce aggression.

Determining overweight and underweight with a new weight-for-height index in captive group-housed macaques.

Housing primates in naturalistic groups provides social benefits relative to solitary housing. However, food intake may vary across individuals, possibly resulting in overweight and underweight individuals. Information on relative adiposity (the amount of fat tissue relative to body weight) is needed to monitor overweight and underweight of group-housed individuals. However, the upper and lower relative adiposity boundaries are currently only known for macaques living solitarily in small cages. We determined the best measure of relative adiposity and explored the boundaries of overweight and underweight to investigate their incidence in group-housed adult male and female rhesus macaques and long-tailed macaques living in spacious enclosures at the Biomedical Primate Research Centre (BPRC), the Netherlands. During yearly health checks different relative adiposity measures were obtained. For long-tailed macaques, comparable data on founder and wild animals were also available. Weight-for-height indices (WHI) with height to the power of 3.0 (WHI3.0) for rhesus macaques and 2.7 (WHI2.7) for long-tailed macaques were optimally independent of height and were highly correlated with other relative adiposity measures. The boundary for overweight was similar in group-housed and solitary-housed macaques. A lower boundary for underweight, based on 2% body fat similar to wild primates, gave a better estimate for underweight in group-housed macaques. We propose that for captive group-housed rhesus macaques relative adiposity should range between 42 and 67 (WHI3.0) and for long-tailed macaques between 39 and 62 (WHI2.7). The majority of group-housed macaques in this facility have a normal relative adiposity, a considerable proportion (17-23%) is overweight, and a few (0-3%) are underweight.

Is Inspiring Group Members an Effective Predictor of Social Dominance in Early Adolescence? Direct and Moderated Effects of Behavioral Strategies, Social Skills, and Gender on Resource Control and Popularity.

Dominance in the peer group is important for adolescents. Resource Control Theory posits that both coercive and prosocial (positively assertive) strategies are associated with dominance. Combining Resource Control Theory with Socioanalytic Theory on personality, we hypothesized that inspiring group members would be an additional effective strategy. This study examined whether the three behavioral strategies and two types of social skills (social competence and manipulation) predicted dominance (resource control and popularity). Participants were 619 Dutch adolescents (Mage = 13.1; 47% female) in the first grade of secondary school. They completed peer reports (behavioral strategies and dominance) and self-reports (social skills). Only inspirational and coercive strategies substantially predicted dominance. Main effects of social skills emerged. Moderation between strategies and social skills was only observed for girls (e.g., coercive strategy use was associated with more popularity for girls with higher levels of social manipulation skills). This study furthered our understanding of the predictors of dominance in adolescence by including inspirational behavior and examining prosocial and antisocial skills.

Comparing physical and social cognitive skills in macaque species with different degrees of social tolerance.

Contemporary evolutionary theories propose that living in groups drives the selection of enhanced cognitive skills to face competition and facilitate cooperation between individuals. Being able to coordinate both in space and time with others and make strategic decisions are essential skills for cooperating within groups. Social tolerance and an egalitarian social structure have been proposed as one specific driver of cooperation. Therefore, social tolerance is predicted to be associated with enhanced cognitive skills that underpin communication and coordination. Social tolerance should also be associated with enhanced inhibition, which is crucial for suppressing automatic responses and permitting delayed gratification in cooperative contexts. We tested the performance of four closely related non-human primate species (genus Macaca) characterized by different degrees of social tolerance on a large battery of cognitive tasks covering physical and social cognition, and on an inhibitory control task. All species performed at a comparable level on the physical cognition tasks but the more tolerant species outperformed the less tolerant species at a social cognition task relevant to cooperation and in the inhibitory control task. These findings support the hypothesis that social tolerance is associated with the evolution of sophisticated cognitive skills relevant for cooperative social living.

Male long-tailed macaques (Macaca fascicularis) understand the target of facial threat.

The cognitive demands of group living have resulted in the development of social competences in a wide range of animal species. Primates are well aware of the complex social structure within their group and infer information about social status by observing interactions of others. A capacity used to infer this information, Visual Perspective Taking (VPT), is present in apes and in monkeys. However, it is unclear whether monkeys really understand that another individual is looking at a specific target. We investigated whether monkeys understand the target of attention of conspecifics using a new paradigm, based on expectancy violation. Subjects were exposed to pictures of scenes involving group members. These pictures either represented congruent (agonistic signals consistent with the dominance hierarchy) or incongruent (signals contradict the dominance hierarchy) social situations. The only difference between scenes concerned the looking direction, that is, the target of attention, and facial expression of the central monkey in the picture. Female subjects did not differ in their looking times to incongruent and congruent scenes, but results may be confounded by their longer looking times at scenes involving kin than non-kin. Male subjects looked significantly longer at incongruent than congruent scenes, suggesting that they understand the target of attention of other individuals. Alternative explanations involving simpler cognitive capacities were excluded. This implies that monkey species share social cognitive capacities underlying VPT with apes and humans. Am. J. Primatol. 78:720-730, 2016. © 2016 Wiley Periodicals, Inc.

Inferences about food location in three cercopithecine species: an insight into the socioecological cognition of primates.

Many animal species use a variety of cognitive strategies to locate food resources. One strategy is to make inferences by exclusion, i.e., perceiving the absence of reward as a cue that another location should be investigated. The use of such advanced cognitive strategies may be more prominent in species that are known to frequently solve social challenges, and inferential reasoning has mainly been investigated in social species such as corvids, dogs, dolphins and non-human primates. In this paper, we investigate how far social intricacy may explain the disparity of reasoning performances observed in three cercopithecine species that differ in the density of their social network and the diversity of their social partners. We used standard reasoning tasks, testing the volume concept and inference by exclusion using visual and auditory modalities. We showed that Old World monkeys can infer the location of invisible food by exclusion. In addition, Tonkean macaques and olive baboons had greater performances in most tasks compared to rhesus macaques. These responses are consistent with the social complexity displayed by these three species. We suggest that the cognitive strategies required to navigate through a demanding social world are involved in the understanding of the physical domain.

The behavioral context of visual displays in common marmosets (Callithrix jacchus).

Communication is important in social species, and may occur with the use of visual, olfactory or auditory signals. However, visual communication may be hampered in species that are arboreal have elaborate facial coloring and live in small groups. The common marmoset fits these criteria and may have limited visual communication. Nonetheless, some (contradictive) propositions concerning visual displays in the common marmoset have been made, yet quantitative data are lacking. The aim of this study was to assign a behavioral context to different visual displays using pre-post-event-analyses. Focal observations were conducted on 16 captive adult and sub-adult marmosets in three different family groups. Based on behavioral elements with an unambiguous meaning, four different behavioral contexts were distinguished: aggression, fear, affiliation, and play behavior. Visual displays concerned behavior that included facial expressions, body postures, and pilo-erection of the fur. Visual displays related to aggression, fear, and play/affiliation were consistent with the literature. We propose that the visual display "pilo-erection tip of tail" is related to fear. Individuals receiving these fear signals showed a higher rate of affiliative behavior. This study indicates that several visual displays may provide cues or signals of particular social contexts. Since the three displays of fear elicited an affiliative response, they may communicate a request of anxiety reduction or signal an external referent. Concluding, common marmosets, despite being arboreal and living in small groups, use several visual displays to communicate with conspecifics and their facial coloration may not hamper, but actually promote the visibility of visual displays.

Orangutan (Pongo spp.) whistling and implications for the emergence of an open-ended call repertoire: a replication and extension.

One of the most apparent discontinuities between non-human primate (primate) call communication and human speech concerns repertoire size. The former is essentially fixed to a limited number of innate calls, while the latter essentially consists of numerous learned components. Consequently, primates are thought to lack laryngeal control required to produce learned voiced calls. However, whether they may produce learned voiceless calls awaits investigation. Here, a case of voiceless call learning in primates is investigated--orangutan (Pongo spp.) whistling. In this study, all known whistling orangutans are inventoried, whistling-matching tests (previously conducted with one individual) are replicated with another individual using original test paradigms, and articulatory and acoustic whistle characteristics are compared between three orangutans. Results show that whistling has been reported for ten captive orangutans. The test orangutan correctly matched human whistles with significantly high levels of performance. Whistle variation between individuals indicated voluntary control over the upper lip, lower lip, and respiratory musculature, allowing individuals to produce learned voiceless calls. Results are consistent with inter- and intra-specific social transmission in whistling orangutans. Voiceless call learning in orangutans implies that some important components of human speech learning and control were in place before the homininae-ponginae evolutionary split.

Does tolerance allow bonobos to outperform chimpanzees on a cooperative task? A conceptual replication of Hare et al., 2007.

Across various taxa, social tolerance is thought to facilitate cooperation, and many species are treated as having species-specific patterns of social tolerance. Yet studies that assess wild and captive bonobos and chimpanzees result in contrasting findings. By replicating a cornerstone experimental study on tolerance and cooperation in bonobos and chimpanzees (Hare et al. 2007 Cur. Biol. 17, 619-623 (doi:10.1016/j.cub.2007.02.040)), we aim to further our understanding of current discrepant findings. We tested bonobos and chimpanzees housed at the same facility in a co-feeding and cooperation task. Food was placed on dishes located on both ends or in the middle of a platform. In the co-feeding task, the tray was simply made available to the ape duos, whereas in the cooperation task the apes had to simultaneously pull at both ends of a rope attached to the platform to retrieve the food. By contrast to the published findings, bonobos and chimpanzees co-fed to a similar degree, indicating a similar level of tolerance. However, bonobos cooperated more than chimpanzees when the food was monopolizable, which replicates the original study. Our findings call into question the interpretation that at the species level bonobos cooperate to a higher degree because they are inherently more tolerant.

Food sharing with friends and acquaintances: A study in preschool boys and girls.

Introduction: The current study examined whether preschoolers in a (semi-)natural situation shared more food with friends or acquaintances, and whether this was different between boys and girls, older and younger children, and for preferred and non- preferred food. In order to do so, we replicated and extended the classical work of Birch and Billman in a Dutch sample. Methods: Participants included 91 children aged between 3 to 6 years (52.7% boys, 93.4% Western European) from a middle- to upper-middle-class neighborhood in the Netherlands. Results: The results revealed that children shared more non-preferred than preferred food with others. Girls gave more non-preferred food to acquaintances than to friends, whereas boys gave more to friends than to acquaintances. No effect of relationship was found for preferred food. Older children shared more food than younger children. Compared to acquaintances, friends made more active attempts to get food. Moreover, children who were not shared with were just as likely to share food as children who were shared with. Discussion: Overall, only a small degree of agreement with the original study was found: Some significant findings could not be replicated, and some unconfirmed hypotheses of the original study were supported. The results underscore both the need for replications and studying the effect of social-contextual factors in natural settings.

Inequity aversion in relation to effort and relationship quality in long-tailed Macaques (Macaca fascicularis).

Social animals may employ evolved implicit rules to maintain a balance between cooperation and competition. Inequity aversion (IA), the aversive reaction to an unequal distribution of resources, is considered such a rule to avoid exploitation between cooperating individuals. Recent studies have revealed the presence of IA in several nonhuman species. In addition, it has been shown that an effort is crucial for this behavior to occur in animals. Moreover, IA may well depend on the partner's identity. Although dominant individuals typically monopolize food, subordinate individuals obtain less preferred food and usually do not protest. Furthermore, "friends" may pay less attention to equity than "nonfriends." We tested whether long-tailed macaques show IA with different cost-benefit ratios. In addition, we determined whether IA depends on relationship quality (RQ). Dominant subjects expressed IA only when a small effort was required. At a very large effort, however, long-tailed macaques did not show IA, possibly owing to bottom effects on the number of rewards they aim to receive. Moreover, and contrary to our predictions, an individual's inequity response was similar when tested with a "friend" or a "nonfriend." Therefore, we conclude that long-tailed macaques show IA only in conditions of moderate effort, yet that IA seems independent of RQ. Furthermore, IA may not be domain specific. Altogether, IA may be a trait present in all species that habitually cooperate, independent of their social organization.

The Older the Better: Infanticide Is Age-Related for Both Victims and Perpetrators in Captive Long-Tailed Macaques.

In wild primates, infanticide is a risk that is especially prevalent when a new male takes over the alpha position. Insight into risk factors related to infanticide may decrease the incidence of infanticide in captivity during male introductions. We investigated several risk factors of infanticide derived from hypotheses explaining infanticide in the wild and tested this in captive long-tailed macaques (Macaca fascicularis) using demographic data spanning a 25.5-year period. Factors that are related to infanticide in the wild explained a large proportion, but not all incidences, of infanticide in captivity. Consistent with the wild data, infants young enough to decrease the interbirth interval (<215 days) were at risk of being killed. In contrast to studies from the wild, infanticidal males were more than 2.5 years younger than non-infanticidal males. This indicates that captive settings can lead to new risks since relatively young males may gain the alpha position, promoting infanticide. Therefore, we propose the adolescent male risk hypothesis as a captive risk factor in which subadult males pose a risk of infanticide. In conclusion, the ages of both males and infants are related to infanticide in captivity and have to be taken into account during male introductions.

Individual behavioral correlates of tail biting in pre-finishing piglets.

Introduction: Tail biting is a widespread problem in pig production systems and has a negative impact on both animal welfare and farm income. This explorative study aims to validate how tail biting is related to general behaviors at the individual level and explore whether these behaviors are related to a particular type of tail biting: two-stage, sudden-forceful, obsessive, or epidemic. Methods: This research was conducted in a standard commercial setting where 89 tail-docked pre-finishing piglets divided into 8 groups were observed 4 days per week from 5 to 8 weeks of age. Each piglet was observed for a total of 160 min using continuous focal sampling. Ten individual behaviors were recorded based on the general behaviors expected to be linked to giving tail biting (PCA1), receiving tail biting (PCA2), and tail biting damage (PCA3). These PCAs were assembled and related to tail biting given, tail biting received, and tail biting lesions. Results: Tail biting did not lead to major damage on the piglets' tail at 8 weeks of age but was observed 420 times, where most of the individuals (72%) were categorized as "biters and victims." When relating PCA1 with tail biting given, piglets that gave more tail biting showed more "active exploration." When relating PCA2 with tail biting received, piglets receiving more tail biting were more "explored while active" and "attacked and explored." When relating PCA2 with tail biting lesions, piglets presenting lesions showed more "agonism." Surprisingly, tail biting lesions were not significantly related to PCA3. The relationship between explorative behaviors and tail biting indicates that the pre-damage stage of two-stage tail biting was the predominant tail biting type, while the damaging stage was likely incipient. The relationship between tail biting and aggression, as well as the minor tail lesions observed suggest that sudden-forceful tail biting was probably present even though it was rarely seen. Obsessive and epidemic tail biting were not observed. Discussion: This study demonstrates that studying tail biting at the individual level helps to identify the type of tail biting present. This gives directions to farmers for applying appropriate measures to prevent the development of tail biting behavior in piglets.

Multigenerational Social Housing and Group-Rearing Enhance Female Reproductive Success in Captive Rhesus Macaques (Macaca mulatta).

To optimize costs and reproductive success, rhesus macaques in biomedical primate research facilities are often peer-reared. Older, dependent infants are typically removed from their natal group to enhance female reproduction. The minimal husbandry age-norm of infant removal is ten months. These practices deviate from species-specific behavior and may reduce welfare, suggesting a trade-off between female reproduction and welfare. However, the effect of breeding group type and rearing history on female reproductive success (i.e., birth rate; inter-birth interval (IBI); offspring survival) is unclear. This retrospective study investigated whether group type (i.e., peer groups versus multigenerational groups) and rearing history (i.e., peer- or hand-reared; group-reared with peer- or hand-reared mother; group-reared) affected female reproductive success in captive rhesus macaques. Data on female reproduction between 1996 and 2019 were collected at the Biomedical Primate Research Centre, Rijswijk. Birth rates were higher in multigenerational breeding groups than in peer groups. Moreover, group-reared females had higher offspring survival than peer- or hand-reared females. IBI was not affected by breeding group type or female rearing history. However, females in both peer and multigenerational breeding groups typically conceived earlier after giving birth than the husbandry infant removal age-norm of ten months. Thus, infant removal at an age of ten months does not enhance a female's reproduction. Altogether, female reproduction and non-human primate welfare can simultaneously be optimized through multigenerational breeding groups and group-rearing.

Overweight Management through Mild Caloric Restriction in Multigenerational Long-Tailed Macaque Breeding Groups.

Caloric restriction (CR) is an effective method to reduce overweight in captive non-human primates (NHPs). CR has been applied to individually- and pair-housed NHPs, but whether applying CR can be effective and safe in group-housed NHPs has not yet been assessed. This study investigates the effect of mild (20%) CR on adult overweight and biochemical parameters, immature growth, veterinary consultations, and reproductive success in multigenerational long-tailed macaque (Macaca fascicularis) breeding groups. Data were derived from anthropometric measurements and blood samples during yearly health checks, complemented with retrospective data on veterinary consultations and reproductive success. Adult body measures decreased after CR, with heavier individuals and females losing more weight compared to leaner individuals and males. CR lowered cholesterol levels in adults but had no overall effect on other biochemical parameters. Yet, biochemical parameters of individuals with high baseline values were reduced more compared to individuals with low baseline values. Immature growth, veterinary consultations and reproductive success were not influenced by CR. Thus, CR targeted the right individuals, i.e., overweight adults, and had no adverse effects on the variables examined in this study. This implies that mild CR can be a valuable overweight management strategy in group-housed NHPs.

Emotional bookkeeping and differentiated affiliative relationships: Exploring the role of dynamics and speed in updating relationship quality in the EMO-model.

Emotional bookkeeping is the process by which primates integrate the emotional effects of social interactions to form internal representations of their affiliative relationships. The dynamics and speed of this process, which comprises the formation, maintenance and fading out of affiliative relationships, are not clear. Empirical data suggest that affiliative relationships are slowly formed and do not easily fade out. The EMO-model, an agent-based model designed to simulate the social life of primates capable of emotional bookkeeping, was used to explore the effects of different types of internal relationship dynamics and speeds of increase and decrease of relationship strength. In the original EMO-model the internal dynamics involves a fast built-up of a relationship independent of its current quality, alongside a relatively fast fading out of relationship quality. Here we explore the effect of this original dynamics and an alternative dynamics more in line with empirical data, in combination with different speeds of internal relationship quality increase and decrease, on the differentiation and stability of affiliative relationships. The alternative dynamics leads to more differentiated and stable affiliative relationships than the original dynamics, especially when the speed with which internal relationship quality increases is low and the speed with which it decreases is intermediate. Consequently, individuals can groom different group members with varying frequency and support a rich social life with stable preferred partners and attention to several others. In conclusion, differentiated and stable affiliative relationships are especially formed when friends are not made too quickly and not forgotten too easily.