Classifications Of Behavioral Interactions

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How, in practice, can we impartially assign positive or negative implications to behavioral interactions such as proximity or grooming that are ambivalent, without recourse to the very theories we seek to support? One solution lies in sequence analysis, using unequivocal behavior patterns as anchors; high-intensity attacks are certainly detrimental to the receiver's fitness, whereas allowing mating is usually beneficial to fitness. If transition probabilities revealed grooming as a predictor of attack, then it might usefully be classed as aggressive, but if grooming often precedes acceptance of mating, it is probably amicable. Firm grounding of a behavior in context may justify an interpretative classification. Often, however, it is impossible to establish a uniform link between ambivalent behaviors (e.g., grooming) and anchors (e.g., coalitionary aid or attack) until the whole social network of interactions within the group has been described. Descriptions based on ultimate function should be eschewed until such links are established.

A very long list of behavior patterns or relational types could be described and analyzed at each level of the structural hierarchy of social dynamics. Here, we consider just one example at each of three levels: grooming as an interaction, dominance as a relationship, and the concept of "social group" as a structure.

Allogrooming can be abundant, observable, and clearly bidirectional or unidirectional, a combination of attributes that have made it perhaps the most frequently studied social interaction. Despite the common assumption that it is amicable or beneficent, it may also cue either dominance or submission in a ritualized context. The style of grooming, be it simultaneous, alternating, or routinely one-sided, is often largely species typical and even a small sample may give important clues to social relations in a wider context. Grooming has been the focus of many primatological studies (reviewed by Goosen 1987), among which it appears to have more to do with social bonding than with hygiene. For example, there is a significant correlation between time spent grooming and group size but not body size (Dunbar 1988). The idea that primates compete for grooming access in a social setting has also been very influential (Seyfarth 1983).

To unravel more complex intricacies requires more probing analytical techniques, of which an especially rigorous example is Hemelrijk and Ek's

(1991) study of reciprocity and interchange of grooming and agonistic "support" during conflict in captive chimpanzees. Reciprocity was defined as one act being exchanged for the same thing (grooming for grooming), whereas interchange was defined as involving two different kinds of acts being bartered (i.e., grooming for support). They created an actor matrix (who initiates to whom) and a receiver matrix (who is recipient from whom) and used a correlation procedure, the Kr test, to examine links between the two matrices while taking account of individual variation in tendency to direct acts. They also attempted to retain stationarity in the data by distinguishing between periods when an alpha male was clearly established or when alpha status was in dispute. Hemelrijk (1990) had shown that grooming and support were both independently correlated with dominance rank, so whenever reciprocity or interchange was part of a triangle of significant correlations, they partialed out the third variable to see whether the remaining two retained significance. Between the females, for example, a significant interchange between grooming and support received did not depend on dominance rank. Females groomed individuals that had supported them regardless of rank, indicating a social bond. However, the reverse was not true in that support was not given in return for grooming, so it was not possible to "buy" support with grooming. Patterns of relationships were different between the sexes.

Cheney (1992) used the distribution of grooming among individuals within a group to examine whether adversity, in the form of rivalry between groups, had a uniting effect within the group; the expectation of this hypothesis was a more egalitarian spread of grooming within the group when competition with neighbors was intense. This expectation was not upheld, perhaps because neglected individuals would suffer a greater loss of fitness by shirking in their support of their own group than by attempting to trade this support for better treatment, even assuming a lack of punishment for failure to cooperate (Clutton-Brock and Parker 1995). Even when intergroup competition was strong, marked hierarchies in grooming favoritism characterized intra-group relations. This study used the distribution of grooming to generate data with which to infer intragroup cohesion and equality. The ultimate functional explanations given depend directly on the validity of that assumption. Our acceptance of such hypotheses may be influenced more by our intuitive anthropomorphic bias for their seeming correctness than by our purely objective assessment of the supporting evidence. This is the dilemma of being a social mammal attempting to describe the societies of other mammals.

The site to which grooming is directed may have social implications. Among long-tailed macaques (Macaca fascicularis), for example, low-ranking females expose the chest, face, and belly less often to higher-ranking females than vice versa. Male—male dyads almost never groomed the face, chest, and belly. Similarly the face, chest, and belly were underrepresented in grooming between individuals that groomed each other less frequently. One interpretation is that recipients of grooming expose less vulnerable parts of their body to, and avoid eye contact with, individuals they perceive as potentially dangerous (Moser et al. 1991; Borries 1992). Another analysis of the form, distribution, and context of social grooming was undertaken in two groups of Tonkean macaques (Macaca tonkeana) by Thierry et al. (1990). They found that kinship and dominance role had no effect on the form or distribution of social grooming among adult females, the most common class of individuals to be observed grooming.

The caveat that no one method of disentangling social relationships is universally appropriate has been stressed by Dunbar (1976). For example, in a study of grooming in three species of macaques, Schino et al. (1988) found that whereas frequency and total duration of grooming were highly correlated, there was low correlation between its total duration and mean duration, and frequency and mean duration were not correlated. A closer inspection of the data also revealed that even this general equivalence between measures of frequency and total duration did not always hold within specific grooming relationships. In general, frequency is a measure of initiation and mean duration is a measure of continuation. In Schino et al.'s case, both measures were needed to interpret adequately the macaques' society.

Not surprisingly, one important stimulus for grooming in rodents is soiled fur (Geyer and Kornet 1982). However, because grooming can be undertaken alone or mutually, it has social implications transcending cleanliness. Stopka and Macdonald (in press) used a similar approach to Hemelrijk and Ek's (1991) analysis of grooming in captive chimpanzees to examine patterns of reciprocity in grooming among wood mice (Apodemus sylvaticus). A powerful method for investigating this involved row-wise matrix correlation and the Mantel test (de Vries et al. 1993). Allogrooming was less common than auto-grooming and was most commonly directed by male wood mice to females. Having described a social network in grooming, Markov chain analysis then revealed that the termination rate of female contact behavior (expressed as her tendency to flee the male's attentions) depended largely on the male's tendency to terminate allogrooming. The pattern of transition rates from one behavior to another revealed that the most common transition was from allogrooming of the female by the male to male nasoanal contact with the female. This indicates the motivation underlying the male's tendency to allogroom females:

males groom a female in order to gain the opportunity to sniff her nasoanal region and, ultimately, to decrease her tendency to flee, thereby allowing them the opportunity to mate. Females are most likely to mate with males that groom them the most. This constitutes an unusual example of the market effect (Noe et al. 1991) in a short-lived mammal, with females essentially selling sex in exchange for grooming.

Allogrooming is the clearest and most common form of cooperation among badgers. It appears to be a valuable supplement to self-grooming because it is focused on the parts of a badger (back of neck, shoulders, and upper rump) that are most difficult for an individual to reach for itself (Stewart 1997; Stewart and Macdonald, unpublished data). The proportion of grooming directed to each segment of the body differs, but is complementary between self-grooming and allogrooming (figure 10.2).

Allogrooming cooperation is maintained in this case by the simple expedient of ensuring that as much grooming is given as received. Adult badgers groom one another simultaneously using a responsive rule set (Axelrod and Hamilton 1981) that dictates that grooming can be initiated generously, but rapidly withdrawn (at a mean of just 1.2 seconds) if the recipient does not respond in kind. If the recipient reciprocates, then the two badgers groom each

Figure 10.2 Silhouettes represent the proportion of time, per square centimeter, spent grooming different portions of the body surface. It shows that although self-grooming or allogrooming alone is strongly biased toward particular body regions, a combination of self-grooming and allogrooming produces a fairly even body coverage. This supports a generally utilitarian function for badger grooming. The slight overrepresentation of the shoulder is the result of that being the most common site of grooming initiation during greeting, indicating an additional social function or a necessary social etiquette during such cooperative relations. R = rump, S = side, H = head, T = tail, B = belly, C = chest.

Self-grooming

Figure 10.2 Silhouettes represent the proportion of time, per square centimeter, spent grooming different portions of the body surface. It shows that although self-grooming or allogrooming alone is strongly biased toward particular body regions, a combination of self-grooming and allogrooming produces a fairly even body coverage. This supports a generally utilitarian function for badger grooming. The slight overrepresentation of the shoulder is the result of that being the most common site of grooming initiation during greeting, indicating an additional social function or a necessary social etiquette during such cooperative relations. R = rump, S = side, H = head, T = tail, B = belly, C = chest.

other simultaneously until one individual defects. The other will then retaliate to this defection, typically in less than half a second, by terminating allo-grooming. No elaborate scorekeeping or partner recognition is required to guard against cheating in this cooperative system. (see figure 10.3a).

We hypothesize that the bartering of allogrooming is so direct because in the putatively primitive society of the badger, individuals spend most of their waking hours foraging separately from one another (Kruuk 1978). As a consequence, there are few opportunities to pay back beneficence in other currencies such as coalitionary aid. Perhaps more importantly, there are also few opportunities for badgers to exert manipulative pressure and extort grooming from other individuals by threat of negative reciprocity. An exception may help prove the rule: During the breeding season, when larger badgers can despotically control breeding opportunities around the sett, smaller individuals sometimes abandoned the "You scratch my back, I'll scratch yours" strategy and groomed larger individuals even without reciprocation (figure 10.3c). The only other exception to the direct reciprocation rule was found when mothers groomed cubs that were too young to reciprocate grooming, a behavior that doubtless accrues fitness benefits in terms of cub survival for the mother (figure 10.3b).

Dominance

Terms such as dominance are used to describe predictable aspects of repeated dyadic interactions (i.e., relationships). The term dominance is a valuable concept made controversial largely by a proliferation of definitions; Drews (1993), having reviewed 13 definitions, concluded that dominance is characterized by a consistent outcome in favor of the same individual within a dyad, within which the opponent invariably yields rather than escalating the interaction. This excludes many uses of dominance and, in particular, does not allow the winner of a single agonistic interaction to be defined as dominant. A dominance hierarchy may be produced by ordering the dominance relationships between dyads. There may be a problem in determining what constitutes yielding as an outcome. If a limiting commodity is involved, then yielding may be allowing the "winner" deferential access to the commodity. Where no commodity is involved, a working definition may be used. For example, Chase (1982) considered an overall yielding response to occur if a hen chicken delivered any combination of three strong aggressive contact acts when there was more than a 30-minute interval after the third action during which the receiver of the actions did not attack the initiator. A certain degree of subjective intu-

Figure 10.3 Barplots of badger allogrooming behavior: (a) Typical pattern of simultaneously reciprocal allogrooming bouts for one individual (A) with other individuals (B, C, D, and E) and all combined (All) in a group-grooming huddle. (The grooming interactions between B, C, D, and E, are not shown. (b) Allogrooming interactions of a mother and her cub, showing the characteristically oneway beneficence. (c) Grooming of one adult male to a large adult female, one of the small minority of observations in which allogrooming between adults is not simultaneously reciprocal.

Figure 10.3 Barplots of badger allogrooming behavior: (a) Typical pattern of simultaneously reciprocal allogrooming bouts for one individual (A) with other individuals (B, C, D, and E) and all combined (All) in a group-grooming huddle. (The grooming interactions between B, C, D, and E, are not shown. (b) Allogrooming interactions of a mother and her cub, showing the characteristically oneway beneficence. (c) Grooming of one adult male to a large adult female, one of the small minority of observations in which allogrooming between adults is not simultaneously reciprocal.

ition may go into such definitions; for example, fox cubs may launch quite ferocious attacks on their mother when food is under dispute, but the lack of reaction and effective supplanting of the mother at the food would not be interpreted by many researchers as a sign that the cubs are meaningfully dominant to the mother.

Dominance is often assumed to depend on body size, as a proxy for combative prowess, and this may generally be true (Lindstedt et al. 1986; Hansson 1992). However, this may sometimes be too simple an interpretation (Bar-bault 1988). For example, Berdoy et al. (1994) showed that among male Norway rats (Rattus norvegicus) age (itself broadly correlated with weight) was a better predictor of dominance than was weight. Age of male is also an important predictor of mate choice by female spotted hyenas (Crocuta crocuta; Hofer and East 1993). Advanced age, obviously a necessary corollary of good survival, may also be a better measure of fitness than current musclepower.

There is a tendency to assume too readily that all societies are arranged in a straightforward dominance hierarchy. Our own work with badgers revealed some of the complexities of elucidating and interpreting the dominance concept. We were interested in feeding interactions because the pattern of food availability and its manner of exploitation are believed to be central to badger social organization (Woodroffe and Macdonald 1995b). We investigated feeding dominance by establishing artificial food stations in the field (Stewart 1997). This experimental approach was deemed necessary because although dominance interactions may be key components of a social system, they may also be too rarely expressed to investigate with statistical rigor in a purely natural context. If a longer period is spent accumulating data, the requirement of stationarity may be breached and dominance relations may change during the observation period. Artificial provisioning experiments allow wild social groups with settled relationships to serve as subjects while improving the number, quality, and rate of observations. There are clear perils to the approach, however; ethically we had to ensure that injurious levels of aggression were not provoked (Cuthill 1991), and scientifically the general relevance of the experimental protocol had to be verified in an unmanipulated context (Wrangham 1974; Dunbar 1988). For this reason we maintained surveillance and control over the experiments using a live infrared video link and pursued further related observations in different social contexts.

We found that when badgers were presented with a single food source requiring contest competition for access, there was little direct evidence of default yielding to certain challengers and hence strict dominance relations: a feeding badger generally escalated aggression to some degree against any chal lenger. However, the study revealed predictors of contest success, so that we could predict relative supremacy in a dyad. These predictors of supremacy included length and weight and, to a lesser degree, age. These attributes alone were not sufficient to predict the direction of supremacy in a dyad, however, because the role of challenged feeding individual also proved to be associated with contest success. And even when an individual occupied the same role against the same adversary, we still could not fully predict the contest winner; subsequent rechallenges often had a different outcome, indicating that some motivational factor such as hunger can also play a decisive role. Hence, in the badger's case it did not prove helpful to attempt construction of a dominance hierarchy, linear or otherwise, from the provisioned patch data. A measure of relative supremacy based on an individual's characteristics and role in a contest proved to be more heuristically valuable.

To further complicate matters, it became clear that if more food sources were provisioned at one site, so that contest competition was no longer necessary for feeding access, scramble (Milinski and Parker 1991) became the preferred method of competition. This cast doubt on the relevance of our predictors of relative supremacy derived from contest competition because most natural badger foods are presented in a way that does allow such scramble competition. Interestingly, however, the behavioral patterns of contest we observed during feeding competition (e.g., side-to-side flank barging) were also seen in male—male competition for estrous females. Analysis of those data revealed that weight, length, and age as well as prior ownership were also predictors of copulation access, leading us to believe that these assets may indeed have general relevance in predicting the outcome of competitive dyadic interactions among similarly motivated badgers.

Social Groups

In terms of their constituent individuals, a group may be transient, as in herds of wildebeest, or nearly permanent, as in packs of wolves. Some may even move daily between these extremes; in the dry season, stable groups of capy-baras may coalesce at water holes into transient herds (Macdonald 1981; Herrera and Macdonald 1989). Many groups can be defined spatially, in terms of the proximity that members maintain to each other. Within territorial species, cohabiting occupants constitute a spatial group, and may be brought together by limited or variably available resources or by sociological advantage (Macdonald and Carr 1989). Membership of spatial groups can be assigned using spatial criteria, within which indices of association can be derived from dyadic proximity interactions; Macdonald et al. (1987J report systematic patterns in the frequency with which members of a farm cat colony sit adjacent to other individuals. Among these cats, and among the aforementioned capybaras, the social status of some low-status individuals is defined by their spatial peripher-alization. As Martin and Bateson (1993) noted, extra meaning can be given to such criteria if it can be shown that there is some aspect of synchrony or complementarity between the actions of individuals in each defined group or dyad relative to outgroup individuals. Clutton-Brock et al. (1982) used a behavior similarity index to quantify such complementarity. At a broader scale there may even be physiological synchrony within groups or between clusters of groups, as within packs of dwarf mongooses (Creel et al. 1991) or between packs of Ethiopian wolves (Sillero-Zubiri et al. 1998).

Spatially coherent groups can include a variety of types. Brown's (1975) schema, adapted by Lehner (1996), is useful in that context. It identified five broad categories of group: kin groups (families and extended families), mating groups (pairs, harems, leks, and spawning groups), colonial groups (e.g., nesting sea birds), survival groups (groups drawn to each other for reasons of foraging, herding, or huddling) and aggregation groups (formed by physical factors acting on animals to force them into one geographical location).

Although membership of a spatial group is a frugal expression of nonran-dom associations, a social group should exhibit some persistent convergence of behavior or unified purpose that results directly from the association and interaction of the constituent network of individuals. That is not to say there will be no competition, antagonism, or discord within the group. It may be endemic. But to differentiate a social group (e.g., a pride of lions) from an aggregation (brown bears at a salmon run may be an example), there must be some element of cooperation (positive interaction) to offset the disharmony. Some populations of badgers are organized into groups that cohabit within the same territory, develop social relationships, and include close kin, yet for a long time there was scant evidence that they accrued any sociological benefit from their association. However as more has been learned of the social system of badgers, more potential advantages of social living have been found. These include winter huddling to conserve energy (Roper 1992), sporadic alloparental behavior (Woodroffe 1993), and occasional examples of coalitionary aid (Stewart 1997). We have even observed young individuals apparently using the main sett as an information center and trailing older adults to new feeding grounds after weaning. The important point to emerge was that none of these advantages appeared sufficient to explain social grouping, particularly because at the same time disadvantages of group living have been identified (Rogers et al. 1997; Woodroffe and Macdonald 1995b). Rather, these social advantages appear to be secondary benefits derived from, but not fully explaining, their group living habit. On one hand, this spatial congruity is distinct from the aggregation of jackals or bears at a carcass or salmon leap, and perhaps even distinct from the community of antagonistic monogamous pairs of maras (Dolichotis patagonum) at a communal breeding den (Taber and Macdonald 1992). On the other hand, the badger spatial group is equally distinct from the social group of African wild dogs, whose cooperative hunting or collective defense of prey leads sociologically to per capita advantage (Creel 1997; see also Packer and Caro 1997). Furthermore, the jackals meeting at a carcass may come from many different territories, but they may nonetheless meet frequently and have well-established social relationships. Indeed, as Macdonald and Courtenay (1996) showed, neighboring territorial canids may even be bonded by familial ties (see also Evans et al. 1989 for genetic evidence of links between adjoining groups of badgers). It is therefore difficult to formulate a precise definition, and social group risks becoming a nebulous concept. A social group constitutes a network of variously close affiliations: closely connected subclusters of individuals comprising subgroups, and groups connected by primary links between core individuals defined as supergroups. Clusters preferentially and mutually exchanging services, support, or aid may be said to display friendships (Smuts 1985), and where clusters act in concert against others we may define them as coalitions. Of course, the benefits an individual can accrue through associations are not always mutually beneficial, and Kraft et al. (1994) designed a diving-for-food experiment to illustrate theft in rat society. Rats were trained to dive into water in order to obtain food from another compartment. All individuals learned to dive; however, those that were able to steal effectively from their diving companions often opted for this less arduous means of securing food.

Macdonald et al. (1987) distinguished a hierarchy of questions to be tackled in quantifying cat sociality, and these could be adapted to a simple description of any society. First, on the basis of their individual comings and goings, what are the probabilities of a given dyad of cats being available to each other for interaction? Second, when they are simultaneously present, what proximity do they maintain? Third, what is the overall frequency of interactions between them, and how does that translate to a rate per unit time in association? Fourth, what are the qualities of interaction? Fifth, what are the directions of flow from initiator to recipient for each type of interaction?

First, indices of association raise the question of what level of proximity constitutes being together, and the answer may vary between species from physical to visual or olfactory contact (Martin and Bateson 1993). Clutton-Brock et al. (1982) classified red deer within 50 m of each other as in the same party, whereas farm cats generally position themselves within 1 m of the nearest neighbor (figure 10.4).

Indices of association can be calculated in which x is the number of observation periods during which A and B are observed together, ya the number of observation periods during which only A is observed, yb is the number of observation periods during which only B is observed, andyab is the number of observation periods during which both A and B are observed but they are not together (see figure 10.5). In some studies, as Ginsberg and Young (1992) pointed out, the proportion of individual A's time spent with B [x/(x + ya + yab)] and the proportion of individual B's time spent with A [x/(x + yb + yab)] may differ only because of a viewing bias concerning ya or yb (see also Cairns and Schwager 1987). However, in the case of the farm cats the value x + ya + yab differs from x + yb + yab because individuals were genuinely present at the resource center at different frequencies (figure 10.5).

This leads to a problem with the simplest association index [x/(x + ya + yb + yab)] because A's association with B is very strong, whereas B's association

Figure 10.4 What constitutes proximity between individuals differs between species and may take account of physical contact, sight, sound, and smell. Farm cats within a matrilineal group may cluster closely, but with individuals positioned in different annuli with respect to the most central females, whereas two red deer stags might be interacting intimately over a distance much wider than that separating entire societies of cats.

Figure 10.4 What constitutes proximity between individuals differs between species and may take account of physical contact, sight, sound, and smell. Farm cats within a matrilineal group may cluster closely, but with individuals positioned in different annuli with respect to the most central females, whereas two red deer stags might be interacting intimately over a distance much wider than that separating entire societies of cats.

Figure 10.5 Considerations in scoring indices of association are illustrated by the comings and going of farm cats at a resource center. Observations may be biased by the observer's success in seeing them or by genuine differences in individuals' attendance records, but the time for which the two cats are simultaneously present defines the opportunity for interacting in the calculation of the example association index.

X / (X+Ya+Yab) = 0.2: 20% X / (X+Yb+Yab) = 0.8: 80% X / (X+Ya+Yb+Yab) = 0.2

Figure 10.5 Considerations in scoring indices of association are illustrated by the comings and going of farm cats at a resource center. Observations may be biased by the observer's success in seeing them or by genuine differences in individuals' attendance records, but the time for which the two cats are simultaneously present defines the opportunity for interacting in the calculation of the example association index.

with A is not necessarily so, and vice versa. For example, if cat A is at the resource center 40 times out of 40 scan censuses and cat B is there 10 times out of the same 40 scan censuses, and they actually sit together during 8 scans, then according to the simple association index, their association index is [8/(8 + 30 + 0 + 2)] = 8/40 = 0.2. However, looking at this another way, it is also true that while at the resource center cat A sits with cat B only 20 percent of its time, whereas cat B spends 80 percent of its time sitting with cat A. This asymmetry is accommodated if we calculate an association index for each cat rather than for each pair of cats. Thus cat A's association with cat B is [x/(x + ya + yab)] = 0.2, whereas cat B's association with cat A is [x/(x + yb + yab)] = 0.8.

Kerby and Macdonald (1988) and Macdonald et al. (in press) took scan samples of presence and proximity of cats at 30-min intervals at three colonies (small, medium, and large memberships), and made ad libitum records of the social interactions of focal cats. Data were summarized for each cat each month, and statistical analyses used these individual summary scores to avoid problems associated with pooling data (Machlis et al. 1985; Leger and Didrichsons 1994). Some generalizations spanned all colonies: the sexes did not differ sig nificantly in their average monthly rate of presence, whereas age classes did; juvenile cats had the highest average monthly rate of presence, and adult males the lowest. Furthermore, each sex—age class displayed a favorite nearest-neighbor distance in terms of its tendency to maintain a given level of proximity to other cats (figure 10.6).

At the large colony, when kittens were within 1 m of an adult female, this was their mother on 71.1 percent and 58.3 percent of occasions for male and female kittens, respectively, whereas adult males tended to be within 1 m of male juveniles most frequently and of male kittens least frequently. Despite such generalizations, there were also significant differences in the pattern of proximity between the colonies. For example, at the large colony, adult males were significantly more frequently within 1 m of adult females, but at the medium-sized colony no such tendency emerged. A further corollary of spatial structure was that certain females occupied spatially peripheral positions, whereas others lived centrally, and the latter had significantly higher reproductive success (figure 10.7).

Taking into account indices of association, it is possible to explore the frequency, rate, and quality at which cats interact. Take three cats, cat A spending twice as much time in the company of cat B as of cat C (figure 10.8). If A rubs on B and C 40 and 8 times respectively, and attacks them 10 and 2 times respectively, then A rubs on and attacks B at only 2.5 times the rate at which it rubs on or attacks C, although A rubs and attacks B five times as often as C. The quality of A's relationship with B and C is the same in that 80 percent of its interactions with both are rubbing and 20 percent are attacking.

In reality, at Kerby and Macdonald's (1988) large colony, individual central males averaged 602 (±254, SE) initiations relative to 502 (±142) for peripheral males, but the latter were present so seldom that their hourly rate of interaction when present averaged 6.9 (±1.5), in comparison to the central males at 2.3 (±0.7). The situation was opposite for females: Peripheral females interacted at low frequency, and when their presence was taken into account, they interacted at an even lower rate. Furthermore, in the context of grooming, Macdonald et al. (1987) showed that whereas the flow of licking differed significantly between pairs of interacting cats, it was invariably symmetric. In marked contrast, the flow of rubbing, though also differing between dyads, correlated with aggression but not grooming and was generally highly asymmetric. They concluded that in the society of female cats, rubbing as a measure of status flowed centripetally toward central dominant individuals; grooming, which was exchanged reciprocally, was a measure of social (and often genetic) alliances, but unrelated to status (figure 10.9).

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