Thomas Hobbes and Jean-Jacques Rousseau set the stage in the eighteenth century for a protracted and polemicized debate about human political nature. If we continue to look for simplified answers that lead strongly in one direction, the discourse promises to be lively—but interminable. A more sophisticated set of questions can be fashioned. However, to generate them we must understand our complex and multifaceted human nature in a relatively full evolutionary context, one that spans the last seven million years.
I suggest that the views of Rousseau and Hobbes may reflect human nature quite accurately—but only if we combine their contradictory viewpoints, rather than allowing them to compete. Humans do seem to enjoy autonomy and serenity. At the same time, they seem to have a competitive penchant for domination that leads to conflict and creates a need for governance. Natural selection is the agency responsible for both facets, so we must look to the evolutionary basis of our political nature if we are to understand these opposing tendencies.
Modeling Our Precursors
Let us begin by considering preadaptations that could have led to the emergence of egalitarianism. To this end I shall create a comprehensive political portrait of the Common Ancestor of humans and the three African great apes. The four-species cladistic methodology I utilize is based on conservative behavioral features that all four species share today (Wrangham 1987). If we compare this kind of modeling to that done with just a single species, the former is far more reliable; it is highly unlikely that four different lineages would have accidentally converged on a behavioral feature lacking in the Common Ancestor.
The shared features I identify on the basis of extant behaviors will be projected back about seven million years, as measured by “molecular time” (Wolpoff 1996), to the Common Ancestor from which the four extant hominid lineages are descended. At that juncture the gorilla lineage went one way and the lineage ancestral to chimpanzees, bonobos, and humans went another. Perhaps five million years ago another split took place, this time with the Pan and human lineages diverging. Eventually, between one and a half and three million years ago, bonobos and chimpanzees diverged to become two separate species.
This four-species cladistic model has scientifically conservative properties that are highly desirable, in that any behavior determined to have been present in the Common Ancestor surely will have been present in all species within the human line—from hominids through to Anatomically Modern Humans. Because I shall also use a narrower, chimpanzee-based referential model, I must address the fact that referential models have been criticized (see Tooby and DeVore 1987; Potts 1987; Stanford and Allen 1991; Moore 1996; Stanford 1998b), and that there are in fact two chimpanzee models.
If we consider the two Pan species, it would appear that the chimpanzee (Pan troglodytes) may have been rather conservative in its evolution (Wrangham 1987; Wrangham and Peterson 1996) in comparison with the bonobo (Pan paniscus). Thus, when the human and Pan lineages diverged, their mutual ancestor could have been more similar to a chimpanzee than to a bonobo. This assumption is based on the fact that bonobos exploit a narrower range of environments than chimpanzees (Wrangham and Peterson 1996), and that such environments are likely to be different from those of their mutual ancestor.
Chimpanzees may be considerably more “generalist” in their behavioral ecology than bonobos, but if we compare chimpanzees with humans the disparity is still greater. As the ultimate generalists (see Potts 1996), we humans have developed huge brains, almost hairless bodies, and language along with symbolic culture. We can exist in virtually any earthly environment as long as it is not too frozen, utterly desiccated, under water, or at too high an altitude. Chimpanzees are lesser generalists: they continue to adapt to a rather wide spectrum of tropical forest and savanna environments similar to those of their ancestors five million years ago (Wrangham and Peterson 1996).
The Original Cladistic Model
In arriving at an innovative experimental model of the hominoid African Common Ancestor as of seven million years ago, Wrangham (1987:55) says:
The behavioral variables selected for analysis are those which theory suggests are primary determinants of other aspects of social organization … They are adult grouping patterns, relationships among mothers, relationships among breeding males, sexual relationships and intergroup relationships. These are considered to be fundamental aspects of social organization because they reflect rather directly the reproductive strategies of breeding adults, which are themselves influenced strongly by ecological variables.
With respect to groups, Wrangham determined that the ancestor of gorillas, the two Pan species, and humans lived in semiclosed, stable social networks with some lone males, and that relations between groups were hostile. It was uncertain whether males formed political alliances within the group, even though males stalked and attacked other males. Wrangham judged that females usually left their natal group to breed and rarely engaged in alliancing with their own kind. With respect to sexual relationships, extant African apes and humans are so variable that nothing could be assumed about promiscuity versus polygyny, or about the duration of relationships. Intergroup relations were hostile on the part of males, but territorial defense could not be deemed universal because gorilla harem leaders seem not to defend the natural resources they exploit.
There is a fundamental question that Wrangham (1987) did not address, even though it probably is just as basic to individual reproductive competition and behavioral ecology as the behaviors enumerated in the quotation above. What about individual competition by means of dominance and submission, and its effect on the internal structure of these semiclosed groups?
In 1987 it would have been easy enough to determine that pronounced social dominance hierarchies existed in gorillas, chimpanzees, and bonobos. Humans, in contrast, posed something of a problem. In spite of interpretations by Lee (1979), Woodburn (1982), and Boehm (1982b, 1984a, 1994b), many anthropologists were still tending to take hunter-gatherer egalitarianism at face value. As we have seen, a few were implying that such societies were “anarchic,” while others seemed to feel that they were naturally “without hierarchy.” With humans in chiefdoms and nations being characterized as hierarchical, but the foragers whose genes we inherited being viewed as having insignificant hierarchy, the call was perhaps a difficult one. However, the ethologists Tiger and Fox (1971) had already countered Fried (1967), taking the position that humans were by nature hierarchical, and Eibl-Eibesfeldt (1974) had attacked the myth of peaceful, nonhierarchical hunter-gatherers.
Another behavioral feature not on Wrangham’s list, one I feel to be potentially important from the standpoint of behavioral ecology, is group leadership. Again, mixed messages were coming from the hunter-gatherer literature. Some bands had named leaders, while others merely had an array of individuals who served as functional leaders when the context was appropriate.
Here, with somewhat different interests, I shall try to develop a Common Ancestral cladistic model more politically complete than previously. The issues of social dominance hierarchy and group leadership will be treated in some detail, along with the question of coalitions among both males and females. This updated, politically oriented model will be based on a better understanding of human egalitarianism, and also on information about captive apes. Because my interest is in the Common Ancestor’s preadaptive potential, captive data are relevant. As we try to understand the political evolution of a species as behaviorally labile as our own, large groups of apes in zoos provide useful information about how labile the Common Ancestor is likely to have been, and about its behavioral potential.
A Political Portrait of the Common Ancestor
Defense of Reproductively Relevant Resources
A political behavior that Wrangham (1987) considered but deemed questionable in the Common Ancestor is territorial defense of natural resources. He found this characteristic to be variable in humans and chimpanzees, doubtful in bonobos, and absent in gorillas. He also suggested that open-country chimpanzees might be utilizing ranges too large to defend, but methodological difficulties have blocked identification of patterns of grouping and intergroup relations where this species inhabits arid lands (McGrew, Baldwin, and Tutin 1981; Moore 1996).
Human territorial defense is variable because some peoples, particularly the simpler foragers who are thought to best approximate our prehistoric forebears (Woodburn 1982; Knauft 1991), appear sometimes to have very little active intergroup conflict and less than highly proprietary attitudes about the land they use (Kelly 1995). However, in view of extant behaviors involving hostile social boundary defense and active perimeter defense as defined by Cashdan (1983) and discussed by Kelly with respect to foragers on several continents, we do well to heed Service’s warning that hunter-gatherers probably had a greater degree of group conflict in the past.
With respect to bonobos, we have seen that the adult males do appear to be antagonistic or at least hold aloof when groups meet, even as females move between groups with friendly interactions. In this case, new information has become available since 1987. Mountain gorillas remain a problem, but their ranges overlap, as do those of lowland gorillas. We do have reports of lowland gorilla groups engaging in aggressive encounters over fruiting trees and stands of bamboo, and also over acquisition or defense of females (Yamigawa et al. 1996:94; Tutin 1996:63–64), a behavior seen also in mountain gorillas. There is some uncertainty about how distinct western lowland gorillas are from mountain gorillas; but if one considers them together, the Common Ancestor did defend natural resources. If one considers them separately, mountain gorillas show no definitive evidence of territorial behavior—even though the potential may be present.
This point merits further discussion. My interest here is more in politics than in behavioral ecology per se. Wrangham’s (1987) definition of territorial defense was based on models of animal territoriality, which classically involve highly predictable, active defense of precisely bounded pieces of real estate. If we change the definition to “defense of any and all reproductively relevant resources in a group context,” we have seen that mountain gorilla silverbacks defend their harems vigorously against interlopers, and they sometimes have their sons as allies in doing so (Fossey 1983).
On this basis, it can be said that the Common Ancestor had a definite tendency to defend reproductively relevant resources in a way that involved semiclosed groups. Giving further credence to a “defense of resources” assumption for the Common Ancestor is the fact that studies of great apes and human foragers are usually made during times of relative plenty, whereas times of serious scarcity are more likely to stimulate direct competition between groups.
The Critical Question of Hierarchy
Was the Common Ancestor hierarchical or not? The question hinges solely on humans. We have just seen that if their egalitarianism is taken to indicate a true lack of hierarchy, no determination can be made about the Common Ancestor. The problem disappears if egalitarian society is seen as involving a mere reversal in the direction of social dominance. I have already mentioned the debate in Current Anthropology (Boehm 1993) and the subsequent critiques of my reverse dominance hierarchy interpretation (see especially Knauft 1993, 1994b; Erdal and Whiten 1994, 1996; Barclay 1993; Boehm 1994b, 1997a; de Waal 1996). A minority of researchers still prefer to see hunter-gatherers as having very little hierarchy, rather than emphasizing the dominance of the rank and file.
My own position can be summarized as follows: all humans are innately prepared to engage in dominance-and-submission behavior (for example, Blurton-Jones 1972), either in orthodox hierarchies or in reverse hierarchies that are operated decisively by the rank and file. If all humans live hierarchically in one way or another, this pattern removes the only obstacle with respect to determining the nature of ancestral social organization. The exact nature of the Common Ancestor’s hierarchy cannot be specified, but in all probability it was of the orthodox variety that produces one or more dominant individuals at the top. This top ancestral ape could have been male or female; to judge from role assignments in our four extant species, it was more likely to have been a male.
Leadership
Decisive individual leadership is prominent in gorillas, noteworthy among chimpanzees, and at least present in moderate form in bonobos and egalitarian foragers. It is very prominent in some human societies. Thus, at least a moderate degree of leadership must have been present in the Common Ancestor. While leaders may influence the group in setting a direction for travel, another important function (shared by all four species) is the pacification of conflicts within the group.
From the standpoint of power and politics, this issue is important. Silverback gorillas use pig grunts to stop fights (Harcourt 1979; Fossey 1983; see also Watts 1996:22), and dominant chimpanzees routinely pacify the quarrels of subordinates (Boehm 1994a; see also de Waal 1982; Goodall 1986). Bonobos are less well studied, but they have some of this behavior as well (Kano 1992; Parish, personal communication), and extant humans exhibit it strongly (Boehm 1986). Egalitarian foragers keep their leaders so weak that their interventions tend to be ineffective, but they do try (see von Fürer-Haimendorf 1967; Knauft 1991; Kelly 1995). The Common Ancestor had group leaders, and they helped to pacify conflicts.
Small Political Coalitions
I have saved for last a type of political behavior that will figure prominently in the interpretation of human political evolution. This is coalition behavior, by which individuals form short-term or longer-term alliances. I shall treat political coalitions of entire groups separately from the smaller, within-group coalitions discussed here.
In 1987 Wrangham judged female small alliances with other females to be rare in gorillas, chimpanzees, and humans, and basically unknown in bonobos. Similar male alliances were very rare in gorillas and bonobos, but common in humans and chimpanzees. Wrangham (1987:59) rated the Common Ancestor as having rare female alliancing and questionable male alliancing. A decade later Wrangham and Peterson (1996), using new information published by Kano (1992), suggested that female coalitions are prominent in bonobos, whereas males do not form alliances to compete within the group. These findings do not appear to have changed Wrangham’s original assessments for the Common Ancestor.
In chimpanzees and bonobos, we have seen that coalitions work strongly to determine the rank positions of individuals. In bonobos, it is well-bonded females who act together physically to advance their own political fortunes and those of their sons. In chimpanzees, it is primarily the males whose extreme status rivalry leads them to form political coalitions, although wild females help one another to resist the domination attempts of adolescent males and to combine forces in other contexts, and mothers at least support their sons in contests with other males by vocalizing (Goodall 1986). Female small coalitions mainly have to do with self-protection and with defense of resources. Male small coalitions aim at increasing rank, and, if possible, unseating the alpha.
With respect to female chimpanzees, Goodall’s (1986) assessment that female hierarchies are relatively muted has been augmented by Pusey, Williams, and Goodall’s (1997) finding that female rank correlates with reproductive success. That makes it less surprising that in captivity a group of chimpanzees was dominated by an alpha female in the absence of any adult males. In taking over the Arnhem Zoo group, this alpha female was helped by a female coalition partner who also ranked high (de Waal 1982). At Mahale, Nishida and Hosaka (1996) report that wild female chimpanzees can act in pairs, or slightly larger units, to support favored males who engage in contests with other males. In two reported instances the females actively supported the alpha male against the same rival, and in one case more than two females were involved (see also Nishida et al. 1995).
Thus, small coalitions play a meaningful role in the political lives of chimpanzees and bonobos, even though the sexes engage in very different ways in the wild. The linking of male bonobos with their mothers (and their mothers’ allies) provides them with highly stable coalitional backing. By contrast, male chimpanzees’ partnerships with other males tend to be expedient and therefore less durable, while females have their own coalitions separately from males.
Gorillas present something of an empirical problem with respect to small coalitions. Lowland gorillas are just coming under study, and their ecology is somewhat different from that of mountain gorillas. Wild mountain gorillas are well studied, though, and even if coalition behavior is rare I can set out some relevant details.
We know that harems are dominated by enormous silverbacks, so female coalitions have relatively little play. However, if resident blackbacks who are sons of the silverback harem leader are not driven from the group, they may act as allies of their father when the silverback’s control is challenged by an interloper. Fossey (1983:75–76) reported a son’s receiving profuse wounds while supporting his father against a usurper who tried to take over the harem, and I judge this to be an important form of coalition behavior. Fossey (1983:66–67) also reported a meeting of two harems during which younger females from one wanted to play with females in the other, their natal group. The more dominant male eventually moved into the other male’s group to retrieve his own females, and at one point several females of that group rushed him, screaming, even though their intimidated harem master stayed to one side. On this basis it can be said that small male and female coalitions are found, if rarely, among wild mountain gorillas.
Among lowland gorillas, Watts (1996:22) stated that males within a group are prone to compete strongly for breeding privileges, with adolescents often leaving their natal groups; but also silverbacks are more tolerant of very close kin than of other males growing up in their harems. This tolerance provides them with allies in defending their harems against outsiders. In this respect lowland gorillas seem similar to mountain gorillas: small coalitions, usually fathers and sons, are operative for males.
Probably the most widespread type of small political coalition among humans is the linkage between close male kinsmen. When one is killed, the others often seek to retaliate, a type of behavior frequently found among hunter-gatherers. It is prevalent as well among tribesmen, who sometimes retaliate as entire clans. In addition, in most human groups families or extended families act together in other political contexts, as when factions form. Small coalitions are widespread.
Let us now triangulate to the Common Ancestor. It would seem that small coalitions of some kind existed, but the question arises whether they involved males or females. Small male political coalitions are formed by humans, chimpanzees, and mountain and lowland gorillas, but not by bonobos; small female political coalitions appear to occur very frequently in bonobos, moderately in chimpanzees, weakly in humans, and rarely in mountain gorillas. Furthermore, among bonobos coalitions are formed by mothers and sons. What can be said conservatively about the Common Ancestor is that small coalitions were present in one sex or the other. And it seems likely that if one sex engaged in such behavior, the potential would have existed for the other sex to do so as well.
Political Macrocoalitions
In the next chapter I trace the evolutionary development of egalitarianism. By my definition, egalitarian society is the product of a large, well-united coalition of subordinates who assertively deny political power to the would-be alphas in their group. For this reason, we will be particularly interested in the capacity of the Common Ancestor to act cooperatively in large coalitions that unify the group—as opposed to small ones that basically divide the group. Here I broaden the definition of political alliancing to include what I have called macrocoalitions (Boehm 1992). These units are formed in the course of intergroup hostilities or collective defense against predators, but also when all or most of a group gangs up on a persona non grata within the group.
Defense of Resources
We have seen that well-studied chimpanzee communities in predominantly forested environments regularly patrol their perimeters, with all the males acting as a single coalition much as humans do when conducting major raids. At Gombe the chimpanzees also engage in “excursions” (Goodall 1986): essentially, to acquire food, an entire community moves into territory normally used by neighbors on an exclusive basis. In one such case, a determined coalition of defending males (from the Mitumba community) discovered a larger group of intruders who were not on their own turf, and by bluffing furiously drove them away (Jane Goodall, personal communication).
Adult male bonobos stick together and appear to be antagonistic when different groups meet—even though systematic patrolling and gang attacks on individuals thus far have not been reported. This too can be taken as macrocoalition behavior. Mountain and lowland gorillas exhibit individual and sometimes coalitional defense of harems, with the few adult males all participating, and lowland gorilla groups engage in contests over concentrated food sources. Humans show a variable but widespread pattern of raiding, feuding, and warfare that often involves smaller groups of males, but sometimes entails all the males of one group actively attacking another group. Although gorillas may be a weak link in this particular argument because their coalitions are so small and infrequent, at the very least the Common Ancestor had some potential to form large coalitions against conspecific interlopers. If harems were absent, these coalitions are likely to have been large.
Defense against Predators
Certain birds engage in mobbing as a type of group bluffing, as do many primates, and mobbing often involves actual physical contact. The basic principle is that a determined group of “weaklings” can scare away or at least seriously confuse a single predator who is far more powerful—and also discourage it from stalking them because their predictable mobbing behavior will spoil its hunting activities.
Pythons are not the only dangerous predators that chimpanzees dare to mob. At Mahale, observers heard a group of chimpanzees vocalizing aggressively, and when they arrived the chimpanzees had driven a female leopard into a sheltered place. One of the chimpanzees entered her refuge and emerged with a leopard cub, which was killed, eaten at, played with, carried, and groomed (Byrne and Byrne 1988). Such mobbing behavior serves to reduce the reproductive success of predators. In Tai Forest in West Africa, chimpanzees were subject to serious predation by leopards, but were able to defend themselves if they were not caught alone (Boesch 1991). In all these cases, coalition behavior enabled the prey species to cope with an individually more powerful predator.
Human hunters often work in groups to hunt large game, just as chimpanzees do, and we may assume that they also resist predation by staying in groups. Bonobos do not seem to have many predators, and data are unavailable about possible mobbing even though such behavior seems possible. Gorillas appear to be a definite exception. Their best defense against predation apparently is sexual dimorphism in body size, and it seems possible that silverbacks protect their harems on a solo basis. The only evidence is Schaller’s (1963) report of finding a silverback gorilla and a leopard both dead, apparently from mutually inflicted wounds. Given the lack of data for bonobos and gorillas, mobbing of predators by the Common Ancestor must be deemed questionable.
Internal Macrocoalitions
Very large coalitions that direct their power against members of their own group are particularly interesting as preadaptations for human egalitarianism. Let us begin with hunter-gatherer moral communities. These may be viewed as large subordinate coalitions formed to neutralize the power of aggressive deviants. Deviance is culturally defined, and outside of the family for an individual “egalitarian” to seek or attain dominant political power is always deviant. When serious inclinations in this direction become evident, the entire group mobilizes into a large political coalition. Had the band been unable to mobilize in this way, it seems highly unlikely that our human type of egalitarianism could have arisen; instead, prehistoric hunter-gatherers would have been dominated by alpha individuals in a position to use coercive force and work their will on inferiors.
It is vital, at this point, to assess whether collective subordinate rebellion is present in the other three African hominoids. Such a potential definitely is present in chimpanzees. In the wild, just once in thirty-five years of observation, the Gombe chimpanzees worked as a large coalition to prevent a long-established alpha male from making a comeback—this, after his displacement by a much younger male (Goodall 1992).
Wilkie’s challenge to Goblin had taken place in the context of mating competition. The female in question was Candy, and Goblin suffered severe wounds to his abdomen and scrotum. Without intervention by a veterinarian who was visiting the park, this defeated alpha male probably would have died of the abscesses that developed. But Goblin recovered and, being a political animal noted for his persistence (see Goodall 1986), he attempted to stage a comeback. When Goblin tried to enter a large group led by Wilkie, the younger males united to drive him away. He was peripheralized for a time before he worked his way back into the hierarchy at a much lower position. In effect, Goblin was ostracized until he gave up his intention to regain the top position.
It would be anthropmorphizing to use the moralistic term “political deviance” in this connection, but it is evident that as alpha candidate Goblin had become a persona non grata. The parallel with human egalitarians is significant, because most of the chimpanzee males united as a single large coalition and collectively dominated their former leader when his intention was to dominate them again. Normally, chimpanzees coalesce into macrocoalitions only to patrol, to hunt, or to mob predators. But they obviously have the potential to mob a member of their own group, and use it occasionally.
A similar report comes from Mahale (Nishida and Hosaka 1996:130): “In 1991, Kalunde, then alpha, chased Ntologi, who was roaming alone as an exiled ex–alpha male. At that time some adult females, together with some adult males, cooperated with Kalunde against Nitologe.” This episode, further described in Nishida (1994), is quite similar to what happened at Gombe. It seems that if a wild chimpanzee group is followed for several decades, eventually a group-internal macrocoalition will be observed in action. Unfortunately, we do not yet have a comparable depth of study in West Africa.
This natural behavior is paralleled by the Arnhem Zoo females, and also by females at Yerkes Regional Primate Research Center in Atlanta. At Arnhem, after adult males were introduced to the group, females no longer occupied the alpha position. But as a large coalition they actively selected one alpha male and limited the functions of a successor (de Waal 1982). At Yerkes, females in the large study group collectively wounded several adult males who were introduced, and finally accepted Jimoh, a small male that two of the females had known previously at another captive location (de Waal 1996:91–92, 131). They also controlled Jimoh, subsequently their alpha, when he attacked a lesser male (in an episode to be described later), so their macrocoalition was powerful indeed. This behavior is directly paralleled by captive lowland gorilla females who in effect voted for a resident young blackback and rejected a new silverback. Bonobos show no sign of forming very large coalitions within their groups, but wild females regularly operate in coalitions larger than pairs, as they easily maintain political parity with males who are larger and more muscular.
Because of bonobos’ being less studied than chimpanzees, I can only offer a “probable” opinion with respect to the likelihood of the Common Ancestor’s forming group-wide macrocoalitions aimed at dominating otherwise dominant individuals within the group. However, in all four species individuals who otherwise would be decisively subordinated combine forces, in groups larger than two, to neutralize the power of their superiors.
Political Effects of Subordinate Coalitions
By examining three types of large-coalition behavior, I have concluded that seven million years ago in its groups the Common Ancestor had the potential to cope collectively with conspecific interlopers, and also to gang up on powerful individuals within the group. By “gang up” I mean that there was the potential to undermine alpha power by excluding certain politically potent individuals from a dominant, alpha-type role. While today it is human egalitarians who carry this pattern to an extreme (doing so moralistically as unified groups of both males and females), the preadaptive potential for subordinate-coalition behavior would seem to have been considerable in the Common Ancestor.
I believe it to have been greater still in the mutual ancestor of chimpanzees and humans—assuming that bonobos are more derived than chimpanzees and can be set aside for purposes of reconstruction. If we move to five million years ago to consider this mutual ancestor, it is likely that all the males of a group hunted together, mobbed predators together, and protected natural resources together, and within the group very large coalitions of males and females had the potential to work effectively to control power at the top.
The Psychology of Subordinate Intransigence
Simple Dominance-and-Submission Models
The mission of behavioral ecologists is to describe species-typical behaviors accurately, and then to understand their ecological and evolutionary ramifications. My interest in this volume is in assessing the ancestral preadaptive potential with respect to a rather unusual type of political behavior, by which a rank and file’s collective insubordination acts to significantly neutralize a social dominance hierarchy. Because I have been dealing in behavioral potential, I have given far more weight to behaviors taking place in captivity than would normally be the case. Another liberty I now take is to consider the political motivations and intentions of subordinates when they join forces to reduce the power of those above them.
Ever since the work of Tinbergen and Lorenz, it has been customary to identify dispositions to “dominate” and “submit” as coevolved behavioral building blocks that produce social dominance hierarchies in a wide variety of social species. In small-brained species such as birds (Schelderup-Ebbe 1922; Tinbergen 1961), or fish (Lorenz 1963), analyses of their dominance-submission interactions imply that they are behaving more or less like automatons—even though they are quite efficient as “learning machines.” By this I mean that they appear to be equipped with on-off switches that make them either dominate or submit on what amounts to a binary basis. Once they have assessed an adversary’s fighting ability, in effect the switch is thrown one way or the other. More recently, Pulliam and Dunford (1980) and Lumsden and Wilson (1981) have emphasized the ethological importance of thinking in terms of decision-making; decision modeling has become rather popular in behavioral ecology, and in human behavioral ecology in particular. I extend this approach, to consider motivations and choices that attend political life within a primate social dominance hierarchy.
If one discards this implicit on-off model, ambivalence becomes a prominent element in decision behavior (Boehm 1989, 1996). I hypothesize that in a hierarchically organized, socially labile species that exhibits a substantial degree of status rivalry are both strong preferences to dominate in situations of competition and inclinations to submit or flee. In the absence of a something like a binary switch, these two orientations work against each other constantly as individuals make their political decisions. The inclination to dominate is mitigated by fear of getting into the wrong fight. If fear overwhelms political ambition or desire for a specific prize, submission is the result. Such fear is useful to individual reproductive success, for the loser of a fight can be wounded or killed.
When we tease apart the act of submission, it appears to be composed of several competing motivational elements. One is the underlying desire to dominate, which is assumed to be continuing even as submission takes place. Another is the aforementioned fear, which results in a posture or gesture of appeasement, a submissive vocalization or facial expression, or possibly flight. Being subordinated also may be complicated by positive orientations: for example, the submitter may anticipate becoming less tense, for often appeasement signals lead to immediate cessation of the threat or even to friendly intercourse. A useful long-term relationship may be involved, one that involves protection against predators, help against political rivals, sharing of food, or routinized and satisfying social contact.
The critical hypothesis here is that submitters often would prefer to dominate, therefore they are likely to be ambivalent when they submit. Negative feelings about being subordinated stem from the basic competitive dispositions that make for status rivalry in the first place: a would-be winner hates to lose—or at least is ambivalent about losing, even though submission has its obvious advantages. The primatological literature is replete with accounts of ambitious subordinates who bide their time until the situation is ripe for rank reversal or a takeover of the top position. Frequently they engage in minor challenges along the way—even though they are obliged to submit (for instance, de Waal 1982; Goodall 1986).
A striking example of this ambivalence is a protracted dominance instability, in which two individuals are competing for rank and cannot arrive at the clear-cut dominance-and-submission arrangement that would both simplify their lives and eliminate the risk of being wounded. Such instabilities are reported for many despotic primate species, but probably the best data come from Arnhem Zoo. For this large group, with its ease of continuous observation, de Waal (1982) has described pairs of males in the throes of protracted instability. Their political ambivalence is obvious from body language and facial expression, as each tries to bluff down the other. Both are feeling aggressive and fearful at the same time, wanting to display or attack but simultaneously exhibiting fear-grins. In one case in which two males wanted to be dominant but were indecisive and exhibiting fear-grins, one of them actually covered his mouth with his hand—apparently to deprive his rival of information that could be of strategic use.
The Chimpanzee Waa as Ambivalent Vocalization
Humans have the gift of speech, a trait that makes it much easier to identify political ambivalences. We have examined statements and philosophies of egalitarians who obviously are ambivalent about status differences: they praise and respect competent warriors and hunters, yet deny them power. While chimpanzees do not have speech, they are a particularly expressive species. Their behavioral repertoire is well suited for observers to make inferences about politically-relevant emotions and intentions in the context of dominance interactions. One chimpanzee vocalization in particular lends itself nicely to the case for ambivalence—the waa-bark (Goodall 1986).
Let me detail a typical group-intimidation scene. When Goblin erects his hair as he readies himself for a display, subordinates are already moving toward trees or holding them in position to climb. When the display begins, a chorus of submissive pant-grunts and pant-barks erupts, but also a great deal of apparently terrified screaming as trees are climbed rapidly, in fear. Once the subordinates are well up in their trees, some of the screaming vocalizations change in the direction of waa-barks. This acoustical transition requires some explanation.
Chimpanzees have a graded call system (Marler 1969), which means that certain calls can and do mutate into other calls along an acoustical continuum. For example, a scream can grade, gradually or quickly, into a waa. When a chimpanzee screams, air is expelled from the vocal cavity as the animal’s lips are drawn back to expose the teeth; this amounts to a fear-grin, which is hard wired. The vocalization has a high, thin, bleating quality, which tends to be continuous except when air intake is necessary. (Orthographically, this vocalic production can be roughly represented by ee, as in “see.”) By contrast, a waa involves expelling air while the lips are pursed, and an “ah-like” vowel is voiced.
The waa vocalization has a variety of acoustical forms, none of which really sounds like barking. Sometimes as the call begins, it sounds like the English semivowel w. This is because the chimpanzee’s pursed lips are being gradually opened. As the vocalization continues, with a definite ah quality to the vocalic production, the lips remain somewhat pursed, rounded but more open. Like the scream, this classic waa can be prolonged until it is interrupted by the need to breathe. A possible variant might be called the wow, because the lips are moved toward a closed position again at the end, to produce a terminal semivowel effect. Another variant that has struck my ear (but may well have no special meaning to chimpanzees) is the eeyow, which exhibits the semivowel effect only at the end of the call and before that vocalically resembles our English ee sound more than our ah. As will presently be seen, a similar variant has been noted in captivity by de Waal (1996).
These are merely personal impressions of wild calls I have heard. I have not used spectrographic analysis to categorize them, but rather an ear that is attuned to human speech sounds and to what is known about articulation of sounds in human language. That same ear also has been exposed to thousands of chimpanzee vocalizations. In the course of sixteen months of fieldwork and many hours of examining videotaped vocalizations, I have been able to find no statistical correlation between the three phonetic variations I seem to hear and the behavioral contexts in which they occur.
For this reason, I take the three waas to be in free variation. Until it can be proved otherwise, there is only a single waa call; statistically, the pure waa seems to be more frequent than the wow or the eeyow variant. When a scream grades into a classic waa, my impression is that the quality of the vowel-sound is what changes first: the screaming ee sound moves, without necessarily an abrupt transition, toward the ah sound of the classic waa. Once the ah is established, when intake of air becomes necessary the semivowel sound appears at the initial position and a waa results; the fear-grin has been relaxing and the lips can now be pursed.
According to Goodall’s (1986) behavioral analysis of chimpanzee calls, what is taking place at the level of the emotions is a transition from fear to hostile defiance. If we look at the typical behavioral context in which the waa is emitted, the facts support this inference: when the displaying alpha male is at close range, screams of pure fear are emitted, and they tend to continue as trees are being climbed. Once the threat is reduced by the high predictability of the alpha male’s staying on the ground to continue his display, often the screams grade into waas.
Goodall (1986) has called the waa a vocalization of defiance, and de Waal (1996) refers to it as “indignant.” The social context suggests a hostile resistance to power or authority, and this is consistent with the ambivalent psychological state described a few pages back. However, the call is emitted in other behavioral contexts as well. After examining several hundred hours of videotape in the laboratory at the Jane Goodall Research Center at the University of Southern California, I came to the conclusion that many (probably most) waas at Gombe are given by subordinates expressing hostility to their superiors from a safe distance. A less usual context occurred when a mother vocally supported her late-adolescent son as he challenged a male ranking above him. This too can be categorized as defiance by a subordinate, for any female ranks lower than an adult male, and in this anecdote the son was challenging an adult male. Waas may be given simultaneously by individuals whose rank is close and who are engaged in a conflict of some type, a behavioral context that suggests mutual defiance. In one videotaped incident, two mothers quickly retrieve their respective offspring after the infants’ rough play has resulted in screaming; then the two of them issue waas at each other for a moment before things calm down. The mothers were females of relatively high rank.
Waas also are used in a very different political context. In one videotaped sequence, two juveniles are playing and the play becomes quite rough. The smaller juvenile screams, disengages to solicit aid from nearby adults who ignore him, goes back to play, again is hurt, and screams loudly. A large adult female, Gigi, is resting five yards away. She issues a waa at that point but does nothing more. It has the effect of temporarily damping the agonistic behavior, but play resumes and again the larger juvenile hurts the smaller. This time Gigi not only waas but erects her hair and gets to her feet, taking a step toward the two. They completely disengage, and the play session is terminated as Gigi lies down again. In this instance the waa was used not in a subordinate capacity, but in a control role. Gigi was on the verge of physically pacifying the conflict by use of her vastly superior strength, but she accomplished the same purpose from a distance: the waa was a warning by a dominant that the two subordinates were about to experience an active intervention.
In another instance of waa use in a context of dominant control, rather than insubordination, Goblin and Evered are eating some meat. Goblin’s little brother Gimble is begging so persistently that both of the adult males repeatedly issue waas at him. As he waas, Evered advances toward Gimble and directs arm threats at him.
Waas also are interspersed with other calls when chimpanzee communities are reacting to “foreign entities” of various types. They are mixed with hunting barks when chimpanzees go after colobus monkeys or bushpigs. Waas were directed at the python in the episode detailed earlier, along with hostile, aggressive wraaas. When two patrols meet and begin to vocalize, again waas are mixed with wraaas and also a variety of hoots (Goodall 1986). These are situations of contest, fraught with hostility and danger, and the basic orientation appears to be both aggressive and defensive. The defensive side is true even in hunting, where a similar mixture of calls is heard. Bushpigs are formidable killing machines and colobus males bite at the testicles of their larger simian predators (Stanford 1998a).
The vast majority of waas appear to be used by defiant subordinates, but obviously their frequency is affected by the fact that alpha males put on their intimidation displays on a daily basis. Clearly, this call (or set of calls) is not dedicated just to the expression of subordinate defiance. What does seem to hold constant, however, is that waas invariably express hostility—and that if fear is involved, it is suborned to the hostility. When an alpha male begins to display and a subordinate goes screaming up a tree, we may interpret this as a submissive act of fear; but when that same subordinate begins to waa as the display continues, it is an open, hostile expression of insubordination. I believe that a similar emotional and behavioral orientation underlies the human moral community’s labeling of domination behaviors by stronger individuals as deviant.
As suggested by de Waal (1996), chimpanzees too have some capacity to “rule from below” in a way that is stable over time. For some years Frans de Waal has been observing a large chimpanzee group in a spacious enclosure at the Yerkes Regional Primate Research Center, and he provides fascinating observations of the use of the waa vocalization by females there. As at Arnhem Zoo, the Yerkes females act as a power coalition and have been able to fiercely reject a number of candidates for alpha male. After they accepted Jimoh, this small male eventually dominated even the alpha female. As alpha male he worked zealously on the community’s behalf to break up fights between other chimpanzees. In the following anecdote, however, Jimoh is pursuing his own self-interest and it is the females subordinate to him who collectively change the course of the conflict.
Jimoh … once detected a secret mating between Socko, an adolescent male, and one of Jimoh’s favorite females. Socko and the female had wisely disappeared from view, but Jimoh had gone looking for them. Normally, the old male would merely chase off the culprit, but for some reason—perhaps because the female had repeatedly refused to mate with Jimoh himself that day—he this time went full speed after Socko and did not give up. He chased him all around the enclosure—Socko screaming and defecating in fear, Jimoh intent on catching him.
Before he could accomplish his aim, several females close to the scene began to “woaow” bark. This indignant sound is used in protest against aggressors and intruders. At first the callers looked around to see how the rest of the group was reacting; but when others joined in, particularly the top-ranking female, the intensity of their calls quickly increased until literally everyone’s voice was part of a deafening chorus. The scattered beginning almost gave the impression that the group was taking a vote. Once the protest had swelled to a chorus, Jimoh broke off his attack with a nervous grin on his face; he got the message. Had he failed to respond, there would no doubt have been concerted female action to end the disturbance. (de Waal 1996:91–92)
This display of collectivized subordinate power can be compared with two of the waa-behaviors cited above for wild chimpanzees. Individually, the females were all subordinate to Jimoh, so their waas can be compared with those of Fifi at Gombe, who waaed in support of her adolescent son Freud when he attacked the adult male Atlas to enter the male dominance hierarchy. Atlas was dominant to Fifi. However, the chorus of Yerkes waa vocalizations can be taken as a collectively dominant warning that grew out of scattered subordinate protests. Contextually, these manipulative waas are directly comparable to those of Gigi, who had dominant power on her side when she similarly stopped a conflict at a distance, without having to intervene physically.
In a sense, the waa vocalizations of large chimpanzee groups can be considered an expression of “public opinion,” as we would put it in human terms. Indeed, this description by de Waal of an emerging female consensus can be compared with Silberbauer’s description, cited earlier, of G/wi foragers in the Kalahari as they gradually arrive at a decision of the entire band about where to migrate next. Human decisions to sanction a bully are likely to develop on a similar basis.
A handful of scattered subordinate protests up in trees can be ignored by a superior as he displays, but an entire group waaing in a context that suggests imminent physical intervention will get his attention. In this sense, waa-barks provide a signal by which individuals in various roles can read the political dynamics that are taking place in their group. The subordinates, if they sense enough support, may be emboldened to rebel in deed, rather than by voice alone. The dominant can react to such collectivized threats and submit to the subordinate coalition, as Jimoh did, or he can try for intimidation, as Goblin did against Wilkie, and find himself in exile.
By definition, ambivalence involves mixed emotions. Indirect but persuasive evidence suggests that chimpanzees can experience serious and protracted ambivalence in situations in which they must submit, or in which they are not sure what to do. Offered as further evidence are the two competitive contexts I earlier described, in which chimpanzee subordinate males set aside their usual response (appeasement and submission or flight) to threaten or actively attack a superior. Both involved rivalry over prized commodities. With meat and with mates, prior possession can embolden a subordinate to act on the aggressive side of his ambivalence: instead of merely resenting domination, the intransigent subordinate is prepared to fight in spite of his adversary’s established dominance.
Egalitarian Ambivalence
What about political ambivalence in humans? In societies that are egalitarian, subordinate intransigence is pervasive and results in vigilant suppression of alpha-type behaviors. In human societies that are hierarchical, subordinates appear to buy into the notion of social and political hierarchy. Nevertheless, they subject their dominators to a certain kind of cost-benefit accounting. If their leaders are fair-minded and attentive to the needs of their people, subordinates remain appreciative—and docile. If they feel the leaders are abusing their powers, however these may be defined locally in terms of political legitimacy, they become ambivalent, hostile, potentially rebellious, and disposed to act forcefully.
My suggestion is that any behaviorally flexible animal that lands on the despotic side of Vehrencamp’s political scale will, in a subordinate role, react with partial hostility when subjected to intimidation that breeds fear and stress. Possible situations include those in which its freedom of action is curtailed, or when it loses specific prizes in food or mating competition. Such motivational ambivalence is most likely to be expressed when the animal senses a possible opportunity to gain rank and is testing the resolve of a superior. The assumption of political ambivalence would seem to include our African Common Ancestor. Like any despotic primate, this ape was prone to become ambivalent about being dominated whenever a subordinate role was necessary and some major prize was at issue. Presumably, the mutual ancestor of humans and chimpanzees was the same, but was more prone to act collectively in an “insubordinate” capacity.
In human nature terms, our discussion is relevant to Fried’s (1967) passing speculation that there may be a universal drive to parity. The disposition in question is not one that orients us specifically to equality, but one that makes us resentful of being unduly subordinated, however that happens to be defined individually—or culturally. Today’s human egalitarians define inappropriate domination culturally, and do so on a hair-trigger basis. Their ingenious invention is to define the ideal society in a way such that no main political actor gets to dominate another. Then they see to it, as a group, that anyone who tries to infringe seriously on this rule is himself dominated. In the next chapter I develop a scenario for how this curious and wonderful state of political affairs originated.