ALTRUISM AND THE HYPOTHESIS OF
META-SELECTION IN HUMAN EVOLUTION

Ferdinand Knobloch, M.D.*

*Professor Emeritus of Psychiatry, University of British Columbia, Vancouver, Canada; Chairman and Training Analyst, Canadian Society for Integrated Psychotherapy and Psychoanalysis (a member of the International Federation of Psychoanalytic Societies); Past President, Psychotherapy Section, World Psychiatric Association.

The problem of altruism in human evolution, although discussed from the very beginning of Darwinism, has not reached a generally recognized solution until today. I will propose the meta-selction hypothesis as a possible partial answer. Heuristically, my study of transference and resistance contributed to its formulation.

THE DARWINISTS PUZZLE OVER ALTRUISM
Since the very beginning, evolutionary theory has been concerned with the problems of selfishness and altruism. Charles Darwin (1871) concluded that selfish individuals are favored by evolution. At the same time, he had no doubt "that tribe including many members who, from possessing in high degree the spirit of patriotism and fidelity, obedience, courage, and sympathy, were always ready to give aid to each other and sacrifice themselves for the common good, would be victorious over most other tribes; and this would be natural selection" (p. 166). Wilson & Sober (1994) summarized the issue laconically: "Altruism involves a conflict between levels of selection. Groups of altruists beat groups of nonaltruists, but nonaltruists also beat altruists within groups" (p. 599).
But Darwin's comments are puzzling and he lets us down by failing to elaborate, complains Cronin (1991) in her book prefaced by J. Maynard Smith. She suspects group selectionism which she rejects, scorning it as "greater-goodism." Cronin joins Hamilton in concluding that Darwin dealt with human altruism, saw the problem, discussed it, but left it unsolved (p. 327).

ARE WE BORN SELFISH?
Dawkins (1976, 1989), who represents the prevailing opinion of contemporary sociobiology, says: "Be warned tthat if you wish, as I do, to build
a society in which individuals cooperate generously and unselfishly towards a common good, you can expect little help from biological nature. I am not advocating morality based on evolution. Let us try to teach generosity and altruism, because we are born selfish (p. 3)." Dawkins talks here about altruism in the usual sense of the word, psychological or motivational altruism, and not about evolutionary altruism and egoism, the "selfish gene," which is broadly accepted and will not be discussed here; see Sober and Wilson, 1998).
Of many authors who have similar views, only one, Badcock (1986), will be mentioned here because of his attempt at psychoanalytic explanation. Our motives are selfish, he claims, but they are repressed and so we believe we are altruistic. According to de Waal (1996), what Badcock asserts is that we are sophisticated hypocrites (p. 238).
The problem is not only theoretical and the educators are worried. They ask: What shall we tell our children? (Thiessen, 1996). And Hrdy (1996), a prominent student of evolution, comes to a surprising and controversial conclusion:
I question whether sociobiology should be taught at the high school level, or even the undergraduate level, because it's very threatening to students still in the process of shaping their own priorities… The whole message of sociobiology is oriented toward the success of the individual. It's Machiavellian, and unless a student has a moral framework already in place, we could be producing social monsters by teaching this. It really fits in very nicely with the yuppie "me first" ethos, which I prefer not to encourage. So, I think it's important to have a certain level of maturity, with values already settled, before this kind of stuff is taught. I would hate to see sociobiology become a substitute for morality. It's a nasty way to live ( p. 19).

VARIETIES OF ALATRUISM
When talking about "altruism" we have to distinguish between psychological (motivational) altruism, that is "altruism" as we understand in ordinary language, and evolutionary altruism, describing behavior which increases the fitness of others at the expense of the fitness of the altruist. In that regard, the present-day sociobiology distinguishes sharply between kin and non-kin members of the group. Altruism among kin members, kin altruism, presents no theoretical problems because, even if the individual's altruistic behavior leads to decreased fitness, it increases, if all goes well, the fitness of one's children or other kin members, so the same genes are preserved (inclusive fitness). In contrast, nothing else can exist among non-kin members than reciprocal altruism, that is, an exchange of equivalent values - something for something else - or the lineage of altruists would die out.
But do people always bargain for reciprocal compensation? Are not people sometimes inclined to help others, without thinking on reciprocal compensation, and accepting gratefulness as reward? That is supported by such empirical studies as of Caporael and al. (1989) and Batson and Shaw (1991). However, the discussion is clouded by traditional philosophical struggles among hedonism, egoism and altruism, as described by Sober and Wilson (1998) who, however, did not quite escape from it, in my opinion. Though I accept their tentative conclusion that people are sometimes genuinely altruistic, I do not use the tern "altruism" as they do, which does not allow for any kind of reward, agreeing with Krebs (1961) and Mook (1981) that egoism and altruism is a false dichotomy.
. The standard argument of sociobiologists against any altruism among non-kin members beyond reciprocal altruism is that individuals, being handicapped by having altruistic genes, would die out. But Sober & Wilson (1998) argue that it is not necessarily so. The authors are representatives of the resurrected group selection theories, which since the sixties was declared as outdated-- not theoretically impossible, but so improbable that it does not need to be considered . They accept that in each group, the number of altruists gradually diminishes. However, the total number of altruists in all related groups grows. This counter-intuitive result is based on Simpson's paradox (p. 25). That this, however, can take place, is conditioned by1. there must be a population of groups; 2. the groups differ in the proportion of altruists; 3. there is a positive correlation between number of altruists and the success of the groups (as Darwin envisaged); and 4. the groups interact in certain limited ways (p. 23).
Although the argument of Sober and Wilson seems persuasive to me, and although it seems to me -- as the erratic path toward truth is often zigzagging -- that the naive group selectionists of the past were replaced by fanatics of individualism (scornful of "greater-goodism "of group selectionists), I will leave the final solution of this controversy to the expert biologists (see the reviews of Sober and Wilson's recent book, e.g., by Maynard-Smith, 1998; Trivers, 1999ab;
and the answer by Wilson and Sober, 1999).
Luckily, the hypothesis of meta-selection, based on the idea of breeding by higher-order breeders, namely groups-as-a-whole, is immune to the outcome of this controversy; for this reason, the controversy does not need to be described here in detail.
HE PATH OF META-SELECTION
One of the stimuli to the meta-election hypothesis came from re-reading the first chapter, on domestication, in Darwin's (1859) Origin of Species. He writes: "The key is man's power of accumulative selection: nature gives successive variations; man adds them up in certain directions useful to him. In this sense he may be said to have made for himself useful breeds…" (p. 47).
Let us imagine that the breeders who made for themselves "useful breeds" were supernatural breeders of humans, Biblical tribal gods - Yahweh, Baal,
or Dagon. These tribal gods competed among themselves for the success of their tribes much more fiercely than the breeders of racehorses do. We are told that with the assistance of Yahweh, in the city of Jericho, all citizens, including children, were killed (Joshua, 6:21). However, they prescribed altruism to the members of their tribes ("love thy neighbor"), submission to authorities, and other rules enhancing the tribal cohesion and fitness. By dispensing rewards and punishments extended to future generations (viz. the Yahweh of the Old Testament).
These supernatural breeders do not need to exist to be effective, if they are prescriptive symbols for the tribes functioning as higher-order breeders. In fierce competition with other groups, they are driven to create selectively cohesive groups. In order to be cohesive, the group demands the bonding of members with altruism beyond reciprocity to non-kin members - and this is what higher-order breeders achieved. This is a more flexible and rich alternative solution to that of the mega-communities of ants and bees, characterized by extreme altruism, but governed by rigid rules, in their undertakings as huge "family businesses."
When using the metaphors of Biblical gods, we must not forget that these complex heavenly figures are of recent origin, not more than ten thousand years old, whereas the EEA (era of evolutionary adaptedness) during which human traits developed, stretched for hundred of thousands of years. We have to assume that these gods had primitive predecessors or equivalent ways to drive the group to be more and more cohesive and function better as units.
Good organization requires some hierarchy. Now, who does the organizing and why? The organizers (roughly synonymous: leaders, power coalition, power elite, ruling class, governing class, and economic and cultural brokers) are those who have formal or informal power to influence planning, make decisions and coordinate the activities of the members, resulting in the distribution of benefits and costs. These leaders care about the welfare of the group as a whole, at least to the degree they are sufficiently rewarded and can retain their power in the group. The success of the group brings rewards to its members (though often grossly uneven), and with it increased inclusive fitness.
Who pays for the organizing efforts? In clubs, there is a club membership fee, in political states, the government collects revenues. In the process of evolution, with the help of the organizers as executors influencing the distribution of benefits and costs, new rules of meta-selection, different from the rules of natural and sexual selection, emerged. These rules created a disposition to pay the "membership fee" willingly, even with pleasure. To act with concern for the welfare of the group and to enjoy receiving and showing altruism beyond reciprocity, to do good to others and receive their appreciation and gratitude, is often regarded as one of the life's greatest pleasures. Indeed, it even affects positively one's physical health (Selye, 1956).
The leadership enjoyed special rewards and privileges, though the leaders of EEA would blush if they would see the privileges of kings of the first civilizations with their thousands of concubines and slaves.

It is further assumed that meta-selection is a continuation of a process which started long before human development. Even in chimpanzees, and presumably still more among hominids, there are signs pointing in the direction of meta-selection. As de Waal (1996) observes about chimpanzees:

"Undoubtedly, prescriptive rules and a sense of order derive from a hierarchical organization, one in which the subordinate pays close attention to the dominant. Not that every social rule is necessarily established through coercion and dominance, but prototype rule enforcement comes from above" (p. 92).

In a simplified form, most of the basic features of human social systems can be seen in chimpanzee groups as described by de Waal (1982, 1989, 1996). For example, vis-a-vis the development from dominance to leadership (1996):
I learned from this development not only that the control role can be performed by the second-in-command but also that the group has a say in who performs it and how. It appears to be one of those social contracts in which one party is constrained by the expectations of the others. As a result, the control role may be regarded as an umbrella shielding the weak against the strong.... It is as if the group looks for the most effective arbitrator in its midst, then throws its weight behind this individual to give him a broad base of support for guaranteeing peace and order. In this sense, then, there is a payoff to the control role: instead of coming from powerful friends, the return favor comes from those who at first glance appear powerless. Although high-ranking individuals have disproportionate privileges and influence, dominance also depends to some degree on acceptance from below. (p. 130).
Though the rewards and coercion from above is essential for the development of meta-selection, it would be erroneous to interpret the hypothesis of meta-selection as ascribing the exclusive shaping power to the group organizers. Even in chimpanzees, the pressures from other roles - peers, sexual partners and even from the subordinates - have shaping effect. As de Waal (1996), comparing humans and chimpanzees, says:
Subordinates have a way of joining forces: alliance formation, a characteristic of the primate order, blunts absolute power. Originally, alliances chiefly served the acquisition of dominance.... The same instrument, however, permits lower echelons to rise up against and even overthrow rulers. Such use is already visible in the chimpanzee...(p. 127).
In short, the dismantling of despotic hierarchies in the course of hominoid evolution brought an emphasis on leadership rather than dominance, and made the privileges of high status contingent upon services to the community (such as effective conflict management) (p. 132).
The hypothesis of meta-selection assumes that there was never a time in human development without hierarchical structure, and that is also the opinion of de Waal (1996) about our past. This is what he says about the so-called egalitarian groups of hunter-gatherers: " Status differences are never completely abandoned, however. Instead of having no hierarchy at all, egalitarian societies occupy one end on a spectrum of dominance styles running from tolerant to despotic"(p. 125).
A group competing for its survival has to be hierarchically organized and that is propelled by the general desire to have power over others, and by differential abilities to fulfill the task and maintenance functions of the group. A remarkable tightening of the hierarchy took place after the hunters-gatherers era, but the last centuries have witnessed an opposite trend, though an undulating one, in which there has been a flattening of the pyramid of power with an increasing sensitivity to the needs of every individual.
What are the pressures of the leadership on the group members, through dispensing rewards and costs, including punishments? 1. It requires de-emphasizing the difference between kin and non-kin.(NOTE 1) 2. It establishes the exchange rates in social exchange, and that is the basis of the accepted rules of fairness, justice and morale behavior which infiltrate the group schema (see NOTE 2) of individuals. 3. It reinforces the hierarchical structure. 4. It creates new benefits, both material and symbolic, for those who protect the group, improve its life and bring sacrifices, and show altruism.
The outcome of these pressures is the development of altruism beyond reciprocity, the development of sympathy (responsiveness to the needs of others in self-other relationship) and empathy (responsiveness by identification), and feeling rewarded by helping others, particularly when they express their gratitude. Meta-selection creates an elaborate group schema (NOTE 3), which is a way for the group to control the individual 24 hours a day. The harmonious relationship with one's group and with one's group schema protects one from the fear of ostracism, brings a feeling of security, and contributes to physiological balance.
The present hypothesis (Knobloch, 1996), attempting to explain altruism beyond reciprocity toward non-kin members of one's group, proposes meta-selection as a third kind of human selection, in addition to natural and sexual selection. It is assumed to be an intra-specific selection, similar to sexual selection, but whereas sexual selection is individual selection, meta-selection is thought to be a special kind of group selection, a process of self-domestication
or self-socialization.
Tendencies based on meta-selection will be often in conflict with tendencies based on natural and sexual selection. This is nothing new: body shapes, ornaments, color and behavior created by sexual selection sometimes interfere with the safety dictated by natural selection - a peacock's tail is an example (summarized by Cronin, 1991, p. 227).
Similarly, tendencies shaped by natural selection may be in conflict with those shaped by meta-selection, e.g., a conflict between self-interest and altruism which pushes hard to encourage sacrifice for the good of the group. It seems that the meta-selection hypothesis can best explain such extreme variants as self-sacrifice, kamikaze, mass suicides of the members of religious sects, and possibly also self-restraint, asceticism, and self-purification (Lopreato, 1984). Altruism toward the group is also often in conflict with nepotism, something which is not news for politicians who often struggle and fail in these conflicts, longing nostalgically for the privileges of kings.
An example of the conflict between sex selection and meta-selection is the conflict between a daughter of high social standing, attracted to a man low in the hierarchy, and her parents, staunchly defending the stability of the social system.
The dilemmas of harmonizing the tendencies flowing from natural, sex and meta-selection are existential dilemmas of human destiny, which we face every day. Different circumstances and cultures call for different solutions. This may be one of the causes why evolution promoted a mix of behavioral phenotypes (Wilson, 1996).
The existence of pro-social behavior and altruism has often been regarded as a challenge to evolutionary theory. What selection pressures could possibly mold human nature to act contrary to a radical self-interest? The argument is that meta-selection creates new conditions of social life in which the right mix of altruism and self-interest leads to higher inclusive fitness than self-interest alone. Selfish persons not liked or ostracized by their group do not do well (Frank, 1988). Throughout the ages and perhaps more than ever in this century, millions of people risked their lives in war, partly because to avoid fighting was even more dangerous, partly to avoid ostracism.

Because of the high value of altruism, pretending to be an altruistic person or pretending to have altruistic motives may be the most frequent kind of cheating. And the deceit of others may be combined with self-deceit, such as a rationalization leading to a belief that one's motives are noble. Such self-deceit leads to behavior which makes the ostensible intentions more convincing.
So the Darwin's idea quoted earlier: "Groups of altruists beat groups of nonaltruists, but nonaltruists also beat altruists within groups" needs a slight modification : The nonaltruists who beat altruists within groups must be successful cheaters pretending altruism- and such talent is rare!
If the hypothesis of meta-selection is correct, we are born not only selfish, as Dawkins claims, but also altruistic- and volatile. It seems that meta-selection took great deal care of generosity and altruism which Dawkins was worried about. Without it, his program of education would be doomed …
DISCUSSION.
Does the postulate of meta-selection meet the conditions which have been set as necessary for group selection? First of all, there should be a population of groups with variable qualities of cohesion and inclusive fitness. There certainly was such a population of hunter-gatherers in groups with variable ability to deal with the task functions and maintenance functions of the group. Clearly, in the context of fierce competition, groups which were better organized and cooperative, as Darwin noted, survived and had higher inclusive fitness.
Second, was there a high rate of extinction? This can be answered positively. Even today, the rate of extinction is high among hunter-gatherers, between 2 and 30 percent every 25 years, as Soltis, Boyd and Richerson (1995) found studying clans in New Guinea.
Thirdly, is there small migration between groups? This can be answered less satisfactorily. It can be assumed that in raids the men were killed, but many non-pregnant women were taken as booty and through their children slowed the genetic changes in the process of meta-selection. Nonetheless, even so, this form of selection was powerful enough - the more so, that it was not random, involving, as in breeding animals, an element of human goal-directedness ("the commands of the tribal gods") which speeded the process. Further, it was intensified by the developing mechanisms of group schema, development of language and cultural conformism (Boyd and Richerson, 1985, 1990).
It should be obvious that group selection by meta-selection as described here is different from earlier versions, the most important version being the one defended most systematically and eloquently by Sober and Wilson (1998).
The hypothesis of meta-selection was discussed on several forums (e.g., Knobloch ,1996, 1997) and in print (Knobloch, 1996; Barkow & Knobloch, 1996; Wilson & Knobloch, 1997). For exmaple, Dr. Barkow agreed with me that, psychologically, a small social group is a model of all social systems:
…intrapsychically, there are only small groups - it does not matter if the reality is that we are dealing nation-states or with groups of evil spirits, our psychology evolved in a small-group context and that is how we conceptualize and react emotionally, in terms of small groups.
However, Dr. Barkow was cautious about the meta-selection hypothesis and pressed for empirical data:
The most important problem is an empirical one - how to demonstrate that leaders do enforce the values you say they do, and how to demonstrate that their followers enhance their genetic fitness by accepting the influence of the leaders. One empirical approach might be that of Betzig's despotism work - she shows that the men at the top have immense reproductive success; you would have to show that the middlemen/loyal followers have greater fitness than the rebels.

It was rewarding that in the discussions, no serious flaws were found in the meta-selection hypothesis. However, the general attitude was cautious -it would surprising if it would be otherwise. The hypothesis will require huge work both conceptually, and in collecting empirical data, to be accepted or rejected. After all, it took one hundred years, before his idea of sexual selection, which Darwin himself added to natural selection, was generally accepted (Cronin, 1991, 123-243).
SUMMARY
According to the meta-selection hypothesis which posstulates a speckal kind of group selection, organized and cohesive human groups became higher-level breeders. These higher-level breeders had selective power to manipulate their members, by a rewards-costs system of social exchange, in such a way that altruism beyond reciprocity, which originally was a handicap, became a potential fitness advantage for those who were able to strike a balance between altruistic attitudes resulting from meta-selection and self-interest resulting from natural selection. The relationship among non-kin members within a group became more similar to that among kin members vis-a-vis altruism beyond reciprocity, social exchange became flexible in its exchange rates, following cultural norms, and group schema developed as an instrument for the fine-tuning of social adjustment and the tightening of group control over private life.
A fragment of a discussion with Barkow gives an idea how complex and arduous the testing of the meta-selection hypothesis will be. But whatever the results of testing will show, it is hoped that the theme of meta-selection opens new alternatives for thinking about the old puzzles of altruism, self-sacrifice and self-restraint which will stimulate both evolutionary psychologists and psychoanalysts striving as they strive for a deeper and unified understanding of human nature.
NOTES.
NOTE 1. TRANSFERENCE AND EVOLUTION. The importance of transference for evolutionary theory is that transference means the shift of relationship from a kin member, usually a member of one's nuclear family, to a non-kin person. But Freud not only drew attention to the fact of transference, he designed an ingenious, yet deceptively simple technique, which can be easily converted into an experimental technique, and which amplifies transference and makes it manifest: psychoanalytic technique itself. (The patient is relaxed on a couch, free-associating, with the therapist out of sight and reacting minimally, depriving the patient of a significant degree of social feedback. The restriction of feedback forces the patient to substitute it by an assumptive feedback and - most significantly - ascribe to the analyst the characteristics and reactions of significant persons from childhood, in particular, parents.) This quasi-experimental technique reveals how surprisingly easy it is for human beings to shift attitudes and feelings from a kin member to a non-kin person.
In some ways, the object fixation of non-human animals is more rigid. Imprinting in birds determines the life-long recognition of conspecifics and even later the recognition of sexual objects, which may lead to the bizarre choices of humans or human-related objects present during hatching (e.g., a bird may court the boots of an experimenter). Some breeds of dogs, attached to a person in a sensitive period, are bound for life; if the human disappears, they become permanently demoralized (Lorenz, 1961).
Although there is a rich body of literature on the relationship of psychoanalysis and evolutionary theory (e.g., Bacciagaluppi, 1989, 1994, 1998; Badcock, 1986; Glantz & Pears, 1989; Slavin & Kriegman, 1992; and Wenegrat, 1984), the significance of ttransferencee for evolutionary theory has not been to my knowledge discussed as possibly one of the most important contributions of psychoanalysis to evoluytionary theory.
NOTE 2. SOCIAL EXCHANGE
The empirical basis of Freud's psycho-economics was his observation about the strength and direction of motivational pressures, pressures similar to vectors which support or conflict with each other. If we accept, as is suggested here, that psychodynamics are basically private sociodynamics, it takes us naturally to social exchange theories, inspired, as in Freud's case, by economic thinking. As might be expected, the range of values people exchange is much broader than those dealt with in economics, and entails goods, services, status, love, information and money (Foa and Foa, 1976). As mentioned above, group schema represents a "parallel market of imaginary values," resulting in rewards and costs - for example, a good or bad conscience.
Cosmides (1989) has examined social exchange in evolution. She means by it "cooperation between two or more individuals for mutual benefit" (p. 52). She argues that as part of the development of social intelligence, and as an important instrument of differential survival in humans, a special module developed in evolution which assesses the cost-benefit exchange of values and detects people who cheat on social contracts - she labels it a "Darwinian algorithm."
Social exchange is regarded here as an important instrument of meta-selection, motivating to pursue equitable, fair and just transactions. But whereas the basis of social exchange is, according to the hypothesis of Cosmides, an Darwinian algorithm, the exchange rates are flexible and determined by culture.
Note 3: GROUP SCHEMA In our past, we lived 99 per cent as hunters-gatherers, in small groups of some 30 persons. Individual and inclusive fitness has always correlated with the capacity of social adaptation. So not surprisingly, " individual psychology is at the same time social psychology"(Freud, 1921, p.18). And we live in a small social group 24 hours a day, since our social reality is freely copied in our "virtual reality," our dreams and day dreams. Our schematic group we live in I named group schema (Knobloch, 1963; also this Journal, Knobloch & Knobloch, 1979a). It is akin to later developed concept (as noticed by Bacciagaluppi, 1994) of Bowlby's "internal working models, 1979, p. 469. It is composed of role schemas: male-female authorities, male-female peers, male-female subordinates and male-female intimate/sexual partners. (Figure 1.) The group schema has three functions. First, it is a cognitive map instructing how to behave toward persons in different roles; it is mostly beneficial, but can be harmful, as in self-defeating behavior based on self-fulfilling prophecies. Second, it is a playground for fantasy and trial exploration in problem solving (Freud's probeweises Handeln, trial acting). It is most vivid in dreams and daydreams, but operates in the background of our consciousness all the time. And, finally, it is a parallel market of social exchange, providing imaginary benefits and costs. The reasons for this formulation, avoiding the intrapsychic-interpersonal fallacy, were explained also in this Journal (Knobloch & Knobloch, 1979a).
Group schema is regarded here as an important instrument of meta-selection. Through the group schema, the society controls the mental life of a person 24 hours a day, it is a way of "self-domestication or "self-socialization".
An important assumption here is that role schemas and their interactions are programmed in evolution, not only such obvious ones as mother-baby or sexually attractive bodies, but also others as well developed in hominoids- such as the ritualized interactions of subordinate chimpanzees with the alpha male, reminiscent of ceremonies at a despotic emperor's court. The innate elements of role schemas are what the ethologists Lorenz and Tinbergen called innate releasing mechanisms or social releasers (originally called innate schemas), responded to by fixed action patterns (Lorenz, 1981). The group schema has similarity with Harlow's (1971, 1983) affectional systems described in rhesus monkeys, but in principle the same according to him in all primates. The secure attachment between a baby monkey and the mother is necessary for the next step of socialization, a healthy relation with peers, which in turn makes possible the relationship with sexual partners. A disturbed bond between a baby and its mother in early childhood leads to disturbed relationship with peers and later with sexual partners, often resulting in unsuccessful mating and failure in mothering. The role schemas are akin to Sullivan's personifications and they can be called, with Jung, archetypes. Though basically programmed by evolution, they are highly individualized in ontogenesis, and, as Freud showed, molded by early relationships, in a process reminiscent of imprinting in other animals.

Figure 1. Group schema
Group schema reflect the structure of the small social group and is composed of role-schemas of Self, Authorites, Peers and Intimate/Sexual Partners. Programmed in evolution, highly individualized ontogenesis.

Acknowledgements. I thank Richard Marcuse for editorial assistance in the preparation of this paper, a short version of which was presented at the Ninth Annual Meeting of the Human Behavior and Evolution Society, Tucson, Arizona, June 4-8, 1997.
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