Primates and Philosophers
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Primates and Philosophers

How Morality Evolved

Frans de Waal, Stephen Macedo, Josiah Ober, Stephen Macedo, Josiah Ober

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Primates and Philosophers

How Morality Evolved

Frans de Waal, Stephen Macedo, Josiah Ober, Stephen Macedo, Josiah Ober

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Can virtuous behavior be explained by nature, and not by human rational choice? "It's the animal in us, " we often hear when we've been bad. But why not when we're good? Primates and Philosophers tackles this question by exploring the biological foundations of one of humanity's most valued traits: morality.
In this provocative book, renowned primatologist Frans de Waal argues that modern-day evolutionary biology takes far too dim a view of the natural world, emphasizing our "selfish" genes and reinforcing our habit of labeling ethical behavior as humane and the less civilized as animalistic. Seeking the origin of human morality not in evolution but in human culture, science insists that we are moral by choice, not by nature.
Citing remarkable evidence based on his extensive research of primate behavior, de Waal attacks "Veneer Theory, " which posits morality as a thin overlay on an otherwise nasty nature. He explains how we evolved from a long line of animals that care for the weak and build cooperation with reciprocal transactions. Drawing on Darwin, recent scientific advances, and his extensive research of primate behavior, de Waal demonstrates a strong continuity between human and animal behavior. He probes issues such as anthropomorphism and human responsibilities toward animals. His compelling account of how human morality evolved out of mammalian society will fascinate anyone who has ever wondered about the origins and reach of human goodness.
Based on the Tanner Lectures de Waal delivered at Princeton University's Center for Human Values in 2004, Primates and Philosophers includes responses by the philosophers Peter Singer, Christine M. Korsgaard, and Philip Kitcher and the science writer Robert Wright. They press de Waal to clarify the differences between humans and other animals, yielding a lively debate that will fascinate all those who wonder about the origins and reach of human goodness.

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Frans de Waal

We approve and we disapprove because we cannot do otherwise. Can we help feeling pain when the fire burns us? Can we help sympathizing with our friends?
—Edward Westermarck (1912 [1908]: 19)

Why should our nastiness be the baggage of an apish past and our kindness uniquely human? Why should we not seek continuity with other animals for our “noble” traits as well?
—Stephen Jay Gould (1980: 261)

Homo homini lupus—“man is wolf to man”—is an ancient Roman proverb popularized by Thomas Hobbes. Even though its basic tenet permeates large parts of law, economics, and political science, the proverb contains two major flaws. First, it fails to do justice to canids, which are among the most gregarious and cooperative animals on the planet (Schleidt and Shalter 2003). But even worse, the saying denies the inherently social nature of our own species.
Social contract theory, and Western civilization with it, seems saturated with the assumption that we are asocial, even nasty creatures rather than the zoon politikon that Aristotle saw in us. Hobbes explicitly rejected the Aristotelian view by proposing that our ancestors started out autonomous and combative, establishing community life only when the cost of strife became unbearable. According to Hobbes, social life never came naturally to us. He saw it as a step we took reluctantly and “by covenant only, which is artificial” (Hobbes 1991 [1651]: 120). More recently, Rawls (1972) proposed a milder version of the same view, adding that humanity’s move toward sociality hinged on conditions of fairness, that is, the prospect of mutually advantageous cooperation among equals.
These ideas about the origin of the well-ordered society remain popular even though the underlying assumption of a rational decision by inherently asocial creatures is untenable in light of what we know about the evolution of our species. Hobbes and Rawls create the illusion of human society as a voluntary arrangement with self-imposed rules assented to by free and equal agents. Yet, there never was a point at which we became social: descended from highly social ancestors—a long line of monkeys and apes—we have been group-living forever. Free and equal people never existed. Humans started out—if a starting point is discernible at all—as interdependent, bonded, and unequal. We come from a long lineage of hierarchical animals for which life in groups is not an option but a survival strategy. Any zoologist would classify our species as obligatorily gregarious.
Having companions offers immense advantages in locating food and avoiding predators (Wrangham 1980; van Schaik 1983). Inasmuch as group-oriented individuals leave more offspring than those less socially inclined (e.g., Silk et al. 2003), sociality has become ever more deeply ingrained in primate biology and psychology. If any decision to establish societies was made, therefore, credit should go to Mother Nature rather than to ourselves.
This is not to dismiss the heuristic value of Rawls’s “original position” as a way of getting us to reflect on what kind of society we would like to live in. His original position refers to a “purely hypothetical situation characterized so as to lead to certain conceptions of justice” (Rawls 1972: 12). But even if we do not take the original position literally, hence adopt it only for the sake of argument, it still distracts from the more pertinent argument that we ought to be pursuing, which is how we actually came to be what we are today. Which parts of human nature have led us down this path, and how have these parts been shaped by evolution? Addressing a real rather than hypothetical past, such questions are bound to bring us closer to the truth, which is that we are social to the core.
A good illustration of the thoroughly social nature of our species is that, second to the death penalty, solitary confinement is the most extreme punishment we can think of. It works this way only, of course, because we are not born as loners. Our bodies and minds are not designed for life in the absence of others. We become hopelessly depressed without social support: our health deteriorates. In one recent experiment, healthy volunteers deliberately exposed to cold and flu viruses got sick more easily if they had fewer friends and family around (Cohen et al. 1997). While the primacy of connectedness is naturally understood by women—perhaps because mammalian females with caring tendencies have outreproduced those without for 180 million years—it applies equally to men. In modern society, there is no more effective way for men to expand their age horizon than to get and stay married: it increases their chance of living past the age of sixty-five from 65 to 90 percent (Taylor 2002).
Our social makeup is so obvious that there would be no need to belabor this point were it not for its conspicuous absence from origin stories within the disciplines of law, economics, and political science. A tendency in the West to see emotions as soft and social attachments as messy has made theoreticians turn to cognition as the preferred guide of human behavior. We celebrate rationality. This is so despite the fact that psychological research suggests the primacy of affect: that is, that human behavior derives above all from fast, automated emotional judgments, and only secondarily from slower conscious processes (e.g., Zajonc 1980, 1984; Bargh and Chartrand 1999).
Unfortunately, the emphasis on individual autonomy and rationality and a corresponding neglect of emotions and attachment are not restricted to the humanities and social sciences. Within evolutionary biology, too, some have embraced the notion that we are a self-invented species. A parallel debate pitting reason against emotion has been raging regarding the origin of morality, a hallmark of human society. One school views morality as a cultural innovation achieved by our species alone. This school does not see moral tendencies as part and parcel of human nature. Our ancestors, it claims, became moral by choice. The second school, in contrast, views morality as a direct outgrowth of the social instincts that we share with other animals. In the latter view, morality is neither unique to us nor a conscious decision taken at a specific point in time: it is the product of social evolution.
The first standpoint assumes that deep down we are not truly moral. It views morality as a cultural overlay, a thin veneer hiding an otherwise selfish and brutish nature. Until recently, this was the dominant approach to morality within evolutionary biology as well as among science writers popularizing this field. I will use the term “Veneer Theory” to denote these ideas, tracing their origin to Thomas Henry Huxley (although they obviously go back much further in Western philosophy and religion, all the way to the concept of original sin). After treating these ideas, I review Charles Darwin’s quite different standpoint of an evolved morality, which was inspired by the Scottish Enlightenment. I further discuss the views of Mencius and Westermarck, which agree with those of Darwin.
Given these contrasting opinions about continuity versus discontinuity with other animals, I then build upon an earlier treatise (de Waal 1996) in paying special attention to the behavior of nonhuman primates in order to explain why I think the building blocks of morality are evolutionarily ancient.


In 1893, for a large audience in Oxford, England, Huxley publicly reconciled his dim view of the natural world with the kindness occasionally encountered in human society. Huxley realized that the laws of the physical world are unalterable. He felt, however, that their impact on human existence could be softened and modified if people kept nature under control. Thus, Huxley compared humanity with a gardener who has a hard time keeping weeds out of his garden. He saw human ethics as a victory over an unruly and nasty evolutionary process (Huxley 1989 [1894]).
This was an astounding position for two reasons. First, it deliberately curbed the explanatory power of evolution. Since many consider morality the essence of humanity, Huxley was in effect saying that what makes us human could not be handled by evolutionary theory. We can become moral only by opposing our own nature. This was an inexplicable retreat by someone who had gained a reputation as “Darwin’s Bulldog” owing to his fierce advocacy of evolution. Second, Huxley gave no hint whatsoever where humanity might have unearthed the will and strength to defeat the forces of its own nature. If we are indeed born competitors, who don’t care about the feelings of others, how did we decide to transform ourselves into model citizens? Can people for generations maintain behavior that is out of character, like a shoal of piranhas that decides to turn vegetarian? How deep does such a change go? Would not this make us wolves in sheep’s clothing: nice on the outside, nasty on the inside?
This was the only time Huxley broke with Darwin. As Huxley’s biographer, Adrian Desmond (1994: 599), put it: “Huxley was forcing his ethical Ark against the Darwinian current which had brought him so far.” Two decades earlier, in The Descent of Man, Darwin (1982 [1871]) had unequivocally included morality in human nature. The reason for Huxley’s departure has been sought in his suffering at the cruel hand of nature, which had taken the life of his beloved daughter, as well as his need to make the ruthlessness of the Darwinian cosmos palatable to the general public. He had depicted nature as so thoroughly “red in tooth and claw” that he could maintain this position only by dislodging human ethics, presenting it as a separate innovation (Desmond 1994). In short, Huxley had talked himself into a corner.
Huxley’s curious dualism, which pits morality against nature and humanity against other animals, was to receive a respectability boost from Sigmund Freud’s writings, which throve on contrasts between the conscious and subconscious, the ego and superego, Love and Death, and so on. As with Huxley’s gardener and garden, Freud was not just dividing the world into symmetrical halves: he saw struggle everywhere. He explained the incest taboo and other moral restrictions as the result of a violent break with the freewheeling sexual life of the primal horde, culminating in the collective slaughter of an overbearing father by his sons (Freud 1962 [1913]). He let civilization arise out of the renunciation of instinct, the gaining of control over the forces of nature, and the building of a cultural superego (Freud 1961 [1930]).
Humanity’s heroic combat against forces that try to drag him down remains a dominant theme within biology today, as illustrated by quotes from outspoken Huxleyans. Declaring ethics a radical break with biology, Williams wrote about the wretchedness of nature, culminating in his claim that human morality is a mere by-product of the evolutionary process: “I account for morality as an accidental capability produced, in its boundless stupidity, by a biological process that is normally opposed to the expression of such a capability” (Williams 1988:438).
Having explained at length that our genes know what is best for us, programming every little wheel of the human survival machine, Dawkins waited until the very last sentence of The Selfish Gene to reassure us that, in fact, we are welcome to chuck all of those genes out the window: “We, alone on earth, can rebel against the tyranny of the selfish replicators” (Dawkins 1976: 215). The break with nature is obvious in this statement, as is the uniqueness of our species. More recently, Dawkins (1996) has declared us “nicer than is good for our selfish genes,” and explicitly endorsed Huxley: “What I am saying, along with many other people, among them T. H. Huxley, is that in our political and social life we are entitled to throw out Darwinism, to say we don’t want to live in a Darwinian world” (Roes, 1997: 3; also Dawkins 2003).
Darwin must be turning in his grave, because the implied “Darwinian world” is miles removed from what he himself envisioned (see below). What is lacking in these statements is any indication of how we can possibly negate our genes, which the same authors at other times have depicted as all-powerful. Like the views of Hobbes, Huxley, and Freud, the thinking is thoroughly dualistic: we are part nature, part culture, rather than a well-integrated whole. Human morality is presented as a thin crust underneath of which boil antisocial, amoral, and egoistic passions. This view of morality as a veneer was best summarized by Ghiselin’s famous quip: “Scratch an ‘altruist,’ and watch a “hypocrite’ bleed” (Ghiselin 1974: 247; figure 1).
Figure 1 The popular view of morality among biologists during the past quarter of a century was summarized by Ghiselin (1974: 247): “Scratch an “altruist,’ and watch a “hypocrite’ bleed.” Humans were considered thoroughly selfish and competitive, with morality being no more than an afterthought. Summarized as “Veneer Theory,” this idea goes back to Darwin’s contemporary, Thomas Henry Huxley. It is visualized here tongue-in-cheek as human nature bad to its core.
Veneer Theory has since been popularized by countless science writers, such as Wright (1994), who went so far as to claim that virtue is absent from people’s hearts and souls, and that our species is potentially but not naturally moral. One might ask: “But what about the people who occasionally experience in themselves and others a degree of sympathy, goodness, and generosity?” Echoing Ghiselin, Wright replies that the “moral animal” is essentially a hypocrite:
[T]he pretense of selflessness is about as much part of human nature as is its frequent absence. We dress ourselves up in tony moral language, denying base motives and stressing our at least minimal consideration for the greater good; and we fiercely and self-righteously decry selfishness in others. (Wright 1994: 344)
To explain how we manage to live with ourselves despite this travesty, theorists have called upon self-deception. If people think they are at times unselfish, so the argument goes, they must be hiding their true motives from themselves (e.g., Badcock 1986). In the ultimate twist of irony, anyone who fails to believe that we are fooling ourselves, and feels that genuine kindness actually exists in the world, is considered a wishful thinker, hence accused of fooling him-or herself.
Some scientists have objected, however:
It is frequently said that people endorse such hypotheses [about human altruism] because they want the world to be a friendly and hospitable place. The defenders of egoism and individualism who advance this criticism thereby pay themselves a compliment; they pat themselves on the back for staring reality squarely in the face. Egoists and individualists are objective, they suggest, whereas proponents of altruism and group selection are trapped by a comforting illusion. (Sober and Wilson 1998: 8–9)
These back-and-forth arguments about how to reconcile everyday human kindness with evolutionary theory seem an unfortunate legacy of Huxley, who had a poor understanding of the theory that he so effectively defended against its detractors. In the words of Mayr (1997: 250): “Huxley, who believed in final causes, rejected natural selection and did not represent genuine Darwinian thought in any way It is unfortunate, considering how confused Huxley was, that his essay [on ethics] is often referred to even today as if it were authoritative.”
It should be pointed out, though, that in Huxley’s time there was already fierce opposition to his ideas (Desmond 1994), some of which came from Russian biologists, such as Petr Kropotkin. Given the harsh climate of Siberia, Russian scientists traditionally were far more impressed by the battle of animals against the elements than against each other, resulting in an emphasis on cooperation and solidarity that contrasted with Huxley’s dog-eat-dog perspective (Todes 1989). Kropotkin’s (1972 [1902]) Mutual Aid was an attack on Huxley, but written with great deference for Darwin.
Although Kropotkin never formulated his theory with the precision and evolutionary logic available to Trivers (1971) in his seminal paper on reciprocal altruism, both pondered the origins of a cooperative, and ultimately moral, society without invoking false pretense, Freudian denial schemes, or cultural indoctrination. In this they proved the true followers of Darwin.


Evolution favors animals that assist each other if by doing so they achieve long-term benefits of greater value than the benefits derived from going it alone and competing with others. Unlike cooperation resting on simultaneous benefits to all parties involved (known as mutualism), reciprocity involves exchanged acts that, while beneficial to the recipient, are costly to the performer (Dugatkin 1997). This cost, which is generated because there is a time lag between giving and receiving, is eliminated as soon as a favor of equal value is returned to the performer (for treatments of this issue since Trivers 1971, see Axelrod and Hamilton 1981; Rothstein and Pierotti 1988; Taylor and McGuire 1988). It is in these theories that we find the germ of an evolutionary explanation of morality that escaped Huxley.
It is important to clarify that these theories do not conflict by any means with popular ideas about the role of selfishness in evolution. It is only recently that the concept of “selfishness” has been plucked from the English language, robbed of its vernacular meaning, and applied outside of the psychological domain. Even though the term is seen by some as synonymous with self-serving, English does have different terms for a reason. Selfishness implies the intention to serve oneself, hence knowledge of ...

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