Abstract: Behavioral ecologists have devoted considerable effort to identifying the sources of variation in individual reproductive success. Much of this work has focused on the characteristics of individuals, such as their sex and rank. However, many animals live in stable social groups and the fitness of individuals depends at least in part on the outcome of their interactions with other group members. For example, in many primate species, high dominance rank enhances access to resources and reproductive success. The ability to acquire and maintain high rank often depends on the availability and effectiveness of coalitionary support. Allies may be cultivated and coalitions may be reinforced by affiliative interactions such as grooming, food sharing, and tolerance. These findings suggest that if we want to understand the selective pressures that shape the social behavior of primates, it will be profitable to broaden our focus from the characteristics of individuals to the properties of the relationships that they form with others. The goal of this paper is to discuss a set of methods that can be used to quantify the properties of social relationships.

Abstract: It has been hypothesized that group-living mammals engage in reconciliation (post-conflict affiliation between former opponents) to reduce the disruptive costs of aggression and restore opponents' tolerance to baseline levels. Recipients of aggression are sometimes reluctant to tolerate the proximity of a recent opponent, however, in apparent fear that aggression will be renewed. In such cases, reconciliatory behaviour by the aggressor's close kin may substitute for direct reconciliation. We describe a playback experiment with free-ranging baboons (Papio hamadryas ursinus) that examines whether friendly behaviour by the aggressor's kin can substitute for direct reconciliation by the aggressor herself. In the test condition, female subjects who had recently been threatened heard the friendly grunt of one of their aggressor's relatives, mimicking kin-mediated vocal reconciliation. In the control condition, subjects heard the grunt of a dominant female from a different matriline. Subjects responded significantly more strongly in test than in control trials. Moreover, in the next hour they were significantly more likely to tolerate the proximity of both their aggressor and the relative whose grunt they had heard. In contrast, subjects' behaviour towards both control females and other members of their aggressor's matriline was unaffected. We conclude that kin-mediated vocal reconciliation can substitute for direct reconciliation in baboons.

Abstract: Are the cognitive mechanisms underlying language unique, or can similar mechanisms be found in other domains? Recent field experiments demonstrate that baboons' knowledge of their companions' social relationships is based on discrete-valued traits (identity, rank, kinship) that are combined to create a representation of social relations that is hierarchically structured, open-ended, rule-governed, and independent of sensory modality. The mechanisms underlying language might have evolved from the social knowledge of our pre-linguistic primate ancestors.

Abstract: In many of the studies reviewed in this book, eavesdropping takes the
following form: a subject has the opportunity to monitor, or eavesdrop upon, an
interaction between two other animals,Aand B. The subject then uses the information
obtained through these observations to assess A`s and B`s relative dominance
or attractiveness as a mate (e.g. Mennill et al., 2002; Ch. 2). For example, Oliveira
et al. (1998) found that male fighting fish Betta splendens that had witnessed two
other males involved in an aggressive interaction subsequently responded more
strongly to the loser of that interaction than the winner. Subjects-behaviour could
not have been influenced by any inherent differences between the two males, because
subjects responded equally strongly to the winner and the loser of competitive
interactions they had not observed. Similarly, Peake et al. (2001) presented
male great tits Parus major with the opportunity to monitor an apparent competitive
interaction between two strangers by simulating a singing contest using two
loudspeakers. The relative timing of the singing bouts (as measured by the degree
of overlap between the two songs) provided information about each “contestants”
relative status. Following the singing interaction, one of the “contestants” was
introduced into the male`s territory. Males responded significantly less strongly
to singers that had apparently just “lost” the interaction (see also McGregor &
Dabelsteen, 1996; Naguib et al., 1999; Ch. 2).
What information does an individual acquire when it eavesdrops on others?
In theory, an eavesdropper could acquire information of many different sorts:
about A, about B, about the relationship between A and B, or about the place of
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A`s and B`s relationship in a larger social framework. The exact information acquired
will probably reflect the particular species social structure. For example,
songbirds like great tits live in communities in which six or seven neighbours
surround each territory-holding male. Males appear to benefit from the knowledge
that certain individuals occupy specific areas (e.g. Brooks & Falls, 1975), that
competitive interactions between two different neighbours have particular outcomes,
and that these outcomes are stable over time. We would, therefore, expect
an eavesdropping great tit not only to learn that neighbour A was dominant to
neighbour B, for example, but also to form the expectation that A was likely to
defeat B in all future encounters. More speculatively, because the outcome of territorial
interactions are often site specific (reviewed by Bradbury & Vehrencamp,
1998), we would expect eavesdropping tits to learn further that A dominates B
in some areas but B dominates A in others. In contrast, the information gained
from monitoring neighbours interactions would unlikely be sufficient to allow
the eavesdropper to rank all of its neighbours in a linear dominance hierarchy,
because not all neighbouring males would come into contact with one another.
Such information would be difficult if not impossible to acquire; it might also be
of little functional value.
In contrast, species that live in large, permanent social groups have a much
greater opportunity to monitor the social interactions of many different individuals
simultaneously. Monkey species such as baboons Papio cynocephalus, for
example, typically live in groups of 80 or more individuals, which include several
matrilineal families arranged in a stable, linear dominance rank order (Silk et al.,
1999). Offspring assume ranks similar to those of their mothers, and females maintain
close bonds with their matrilineal kin throughout their lives. Cutting across
these stable long-term relationships based on rank and kinship are more transient
bonds: for example, the temporary associations formed between unrelated
females whose infants are of similar ages, and the “friendships” formed between
adult males and lactating females as an apparent adaptation against infanticide
(Palombit et al., 1997, 2001). In order to compete successfully within such groups, it
would seem advantageous for individuals to recognize who outranks whom, who
is closely bonded to whom, and who is likely to be allied to whom (Harcourt, 1988,
1992; Cheney & Seyfarth, 1990; see below). The ability to adopt a third party`s perspective
and discriminate among the social relationships that exist among others
would seem to be of great selective benefit.
In this chapter, we review evidence for eavesdropping in selected primate
species and we consider what sort of information is acquired when one individual
observes or listens in on the interactions of others. We then compare eavesdropping
by primates with eavesdropping in other animal species, focusing on both
potential differences and directions for further research

Abstract: Historically, a dichotomy has been drawn between the semantic communication of human language and the apparently emotional calls of animals. Current research paints a more complicated picture. Just as scientists have identified elements of human speech that reflect a speaker's emotions, field experiments have shown that the calls of many animals provide listeners with information about objects and events in the environment. Like human speech, therefore, animal vocalizations simultaneously provide others with information that is both semantic and emotional. In support of this conclusion, we review the results of field experiments on the natural vocalizations of African vervet monkeys, diana monkeys, baboons, and suricates (a South African mongoose). Vervet and diana monkeys give acoustically distinct alarm calls in response to the presence of leopards, eagles, and snakes. Each alarm call type elicits a different, adaptive response from others nearby. Field experiments demonstrate that listeners compare these vocalizations not just according to their acoustic properties but also according to the information they convey. Like monkeys, suricates give acoustically distinct alarm calls in response to different predators. Within each predator class, the calls also differ acoustically according to the signaler's perception of urgency. Like speech, therefore, suricate alarm calls convey both semantic and emotional information. The vocalizations of baboons, like those of many birds and mammals, are individually distinctive. As a result, when one baboon hears a sequence of calls exchanged between two or more individuals, the listener acquires information about social events in its group. Baboons, moreover, are skilled “eavesdroppers:” their response to different call sequences provides evidence of the sophisticated information they acquire from other individuals' vocalizations. Baboon males give loud “wahoo” calls during competitive displays. Like other vocalizations, these highly emotional calls provide listeners with information about the caller's dominance rank, age, and competitive ability. Although animal vocalizations, like human speech, simultaneously encode both semantic and emotional information, they differ from language in at least one fundamental respect. Although listeners acquire rich information from a caller's vocalization, callers do not, in the human sense, intend to provide it. Listeners acquire information as an inadvertent consequence of signaler behavior.