Darwin: Religion & Ethics ─ Reading #2
The Importance of Groups in Human Evolution

Human Morality Begins with a “We”
Michael Tomasello

Center for Humans and Nature


[1] Chimpanzees are in many ways highly cooperative. Chimpanzee “friends” support one another in fights and groom one another to remove lice and other vermin. In appropriate circumstances individuals will help their groupmates fetch out-of-reach objects, and sometimes even share food with them.  

[2] But chimpanzees are not moral in the human sense of individuals self-regulating their behavior by a sense of the right and wrong ways to treat others. Thus, despite our moral concern for them as living creatures, we do not allow them to roam freely in our midst for fear they will attack our children, steal our food, destroy our property, and generally wreak havoc without regard for anyone else. And if they did all of these terrible things, no one would blame them or hold them responsible. Why not? Why are they not moral in the human sense? 

[3] The answer is that morality is a uniquely human form of cooperation that evolved under particular circumstances. At some point after the split with other great apes, humans were forced by some ecological change— perhaps a population explosion of monkeys who outcompeted them for their normal fruits and vegetation—into a new feeding niche: they began to specialize in foods that could be acquired only, or more easily, via cooperation (e.g., large game). The new way of thinking was: “we” are foraging together interdependently and so “we” should share the spoils together “fairly.”  

[4] It is in this context that human morality arose. Of course humans retained many self-serving motives; natural selection requires that individuals not cooperate themselves out of existence. But this new way of life created a new set of cooperative motives that competed with self-serving motives. Of special importance were:

you >me, in which I am motivated to sacrifice for your welfare (to a point)

you = me, in which I see you as an equally deserving partner

we >me, in which I subordinate myself to our agreed-upon norms of conduct 

[5] The sacrificing for others, you >me motive became especially strong in early humans because they were especially interdependent with one another. If the only way I can get food is by teaming up with you, then if you are hurt or ill I had best do everything I can to help. And in a marketplace of partner choice, with many potential partners, I had better make sure that others perceive me as a cooperative partner as well— otherwise I will starve. We care for each other— and about what others think of us— because we are interdependent. 

[6] The equality with others, you = me motive is even more clearly moral as it is not about ensuring my own long-term survival. The point is that as early human individuals collaborated with a partner toward a common goal, they understood that each of them was equally causally responsible for the resources produced—neither could have done it alone. Moreover, as they engaged in a particular collaborative foraging activity repeatedly, for example, hunting for antelope, they understood that for their mutual success each had to play their roles adequately. Each role thus had mutually known standards that applied to anyone in that role; they were impartial. In this context, early humans developed a sense of collaborative partners as equally deserving of the spoils: we who collaborate together treat one another (as opposed to free riders) fairly. 

[7] The subordination of self to the collective, we >me motive reflects the Rousseauian insight that in interdependent collaboration “we” judge each of us. When we agree (make a joint commitment) to hunt antelope together, we each agree at the same time to play our role the way we both know it should be played. If one of us slacks off, the other will call him out on it. The slacker cannot just blow off the criticism because she herself believes the criticism is legitimate; we agreed at the time of the joint commitment that this is how we should do things for our mutual success (and she will be shunned in the future for not keeping to the commitment). When I do break my commitment, I feel guilty precisely because I believe the joint commitment is legitimate and that my equal partner does not deserve to be treated this way.  

[8] In A Natural History of Human Morality, I argue that in the process of moving to a more cooperative, interdependent lifestyle humans developed these three non-selfish motives. And then came the move to cultural groups in which the interdependent “we” was everyone in my tribe, which did not include others in the out-group. Humans did not become moral angels during this process, but there did come into existence a moral option— what is best for us, however defined— that might be chosen. But it is not always clear what is the moral option; moral dilemmas are all too real. Moral dilemmas exist because human morality is not a homogeneous set of rules that automatically apply everywhere, but rather a motley of social propensities that humans have accumulated over hundreds of thousands of years aimed at promoting “us,” as defined in various ways.  

[9] If we now ask whether morality is serving us well in the contemporary world, with all of its cultural and political complexities, the only evolutionary answer is: we are still here. Whether we will be here much longer depends, it would appear, on whether all of us earthlings choose to conceive of ourselves as a “we,” working together for mutual benefit.

D. S. Wilson

Essay XV: Group Selection in the Wild September 18, 2009



[1] I have argued that prejudice against group selection is impervious to evidence from laboratory experiments. It is also impervious to evidence from the wild. I will focus on one of many examples that can be provided. In 1995, Robert Heinsohn and Craig Packer published an important paper on territorial defense in lions in the journal Science. As good experimental field biologists, they had played recordings of lions from neighboring territories to observe how females of the focal territory responded. They discovered that the same individuals consistently arrived first at the scene while others consistently lagged behind. There seemed to be bravehearts and cowardly lions within the same pride.


[2] Heinsohn and Packer looked for an advantage to counteract the cost of territorial defense for the bravehearts within their own pride and couldn't find it. The bravehearts weren't socially dominant, they didn't have more offspring, and they didn't punish the cowardly lions, who simply seemed to be cheating and getting away with it. The bravehearts were providing a public good at their own expense, an animal version of the tragedy of the commons made famous by Garrett Hardin in the 1960's. Here is how Heinsohn and Packer described the situation to the best of their knowledge:

[3] Female lions share a common resource, the territory; but only a proportion of females pay the full costs of territorial defense. If too few females accept the responsibilities of leadership, the territory will be lost. If enough females cooperate to defend the range, their territory is maintained, but their collective effort is vulnerable to abuse by their companions. Leaders do not gain "additional benefits" from leading, but they do provide an opportunity for laggards to gain a free ride.

[4] Let me pair this passage with the canonical passage by Darwin in The Descent of Man, Ch.3:

It must not be forgotten that although a high standard of morality gives but a slight or no advantage to each individual man and his children over other men of the same tribe, yet that an increase in the number of well-endowed men and advancement in the standard of morality will certainly give an immense advantage to one tribe over another. There can be no doubt that a tribe including many members who, from possessing in a high degree the spirit of patriotism, fidelity, obedience, courage, and sympathy, were always ready to aid one another, and to sacrifice themselves for the common good, would be victorious over most other tribes; and this would be natural selection.

[5] I hope you can see the similarity between these two passages. Darwin was identifying what I call the original problem; traits that benefit the whole group are often disadvantageous within the group. The counterbalance for cheating does not reside within the group; it resides in the process of more cooperative groups outcompeting less cooperative groups. Darwin's example was hypothetical. Heinsohn and Packer seem to be providing a real-world example with territorial defense in lions. They even stress the importance of between-group competition as the primary influence on the evolution of lion sociality.


[6] Before continuing, let me issue two caveats. First, I have the highest respect for Heinsohn and Packer. They are topnotch scientists who can only be admired, not only for conducting such arduous research but also for attempting controlled experiments in the wild. Second, the last word has not been written on lion social behavior. Perhaps they or someone else will find a within-group advantage for bravehearts in the future. I'm interested in how they interpret their current data. They don't interpret it as a provisional example of group selection. They don't even mention group selection as a possibility. I doubt that it even occurred to them to regard group selection as a viable possibility, even though a description comparable to Darwin's flowed from their own pen!

[7] For those who feel impelled to shout "kin selection!" because lion prides are composed of related females, my reply is the same: Genetic relatedness explains why bravehearts are clustered in some prides and cowards in others. Cowards still have the advantage within each pride, the observation that Heinsohn and Packer find so puzzling. If they had a better understanding of kin selection seen through the lens of multilevel selection theory (see T&R XIII), they wouldn't be so mystified by their own data.

 [8] This example illustrates a problem that pervades the evolutionary literature. Group selection became such a pariah concept that most people don't look for it. They haven't looked for it for so long that they forgot what it looks like and can't even recognize it when it bites them in the butt! Even when they do recognize it, they are tempted to describe it in a way that doesn't use the G word to make it more palatable to their peers. In this fashion, our students continue to learn that the rejection of group selection was a triumphant advance for evolutionary theory while the evidence for group selection lies all around us. 

[9] Science is no better than political revisionism if this situation is allowed to persist. One reason that I am writing this series of blogs is because I am an idealist about science. I regard it as the best cultural system we have for holding people accountable for what they say. Scientists have a responsibility to keep track of the history of their ideas and to acknowledge mistakes from the past, no matter how large. Unfortunately, like religion, science as practiced often falls short of science as idealized. The group selection controversy is an embarrassment for science and the sooner its shortcomings are corrected, the better.

Debates About Morality | Evolution & Morality

Jeffrey Schloss

Center for Humans & Nature


[1] As the new year arrives, the holiday-season movie It’s a Wonderful Life has played in many households. Evolutionary biologist Stephen J. Gould employed this film as a metaphor for the contingent a-directionality of evolution— how a single event (in this case one human life) can randomly but profoundly alter historical outcome. But the film offers another theme, involving the tension between the pursuit of individual gain and the investment in collective welfare. Mr. Potter, the mercenary villain, decries “starry-eyed dreamers” who work on behalf of “a discontented, lazy rabble.” The protagonist George Bailey laudably— though with considerable struggle— relinquishes his own dreams in order to support friends and neighbors in his community. And in so doing he ends up recognizing the richness of a life embedded in the enriching of others.   

[2] While the moral message may seem attractive, when first released, some viewed it as gravely immoral: the FBI deemed its communitarian views were politically and morally subversive. Today’s civic climate is brooding with similar ethical tensions. The fascinating thing is that these tensions reflect a variety of long-standing debates over the relationship of evolutionary theory— and the very character of nature itself— to morality. A number of these questions are ably and graciously explored by contributors to this series.  

[3] One issue concerns whether, and in what way, evolution relates to morality at all. Herbert Spencer, who coined the phrase “survival of the fittest,” invested his life in developing an evolutionary morality, rooted in (and strongly affirming of) unrestrained competitive struggle between individuals. Darwin had a comparably biological but more benign view of morality, proposing that social sentiments may facilitate personal sacrifice that helps the group compete with other groups. On the other hand, Alfred Wallace, co-discoverer of natural selection, concluded that selection cannot explain morality, because humans value right and wrong beyond benefits to individual or group. Thus, a “Higher Intelligence” guiding our origins is required. And finally we have T. H. Huxley, “Darwin’s bulldog,” who might be viewed as a Mercutio invoking “a plague on both your houses”— biological and supernatural. He acknowledged the “paradox that ethical nature, while born of cosmic nature, is necessarily at enmity with its parent.” For Huxley, like Wallace, moral virtue does not arise from but is actually opposed to success in the struggle for existence; yet, the capacity to restrain biological dispositions toward selfishness and value the welfare of others is not supernatural. Rather, it is rooted in human law and custom.  

[4] These basic views have animated discussion since (and actually long before) Darwin. In particular, the “cultural virtue versus biological selfishness” perspective was subsequently emphasized by Freud and has been prominently promoted by manifold evolutionary biologists who, like Richard Dawkins, exhort, “Let us try to teach generosity and altruism, because we are born selfish.”   

[5] Although that view has been criticized as cynical— primatologist Frans de Waal refers to it as “veneer” morality, or a thin coat of moral goodness imposed on an averse biological substrate— it is also just too simplistic. For one thing, characterizing biology as uniformly “selfish” wrongly equates the self-replicating efficacy of genes with strictly self-serving behaviors of the creatures that carry genes. But we know that genes can be transmitted successfully via all sorts of behaviors that benefit another at cost to the actor. Think of bees giving their lives defending a hive.  

[6] Moreover, when it comes to human relationships, the “born selfish” claim conflates self-fulfilling consequences with self-serving motives in our interactions with others. As George Bailey discovers when he receives unanticipated support from those he helped, genuinely caring about others can have positive but unsought personal outcomes: George was so gratefully rewarded precisely because he did not seek reward. Indeed, most of us recognize both giving and receiving, without vigilant demands of reciprocity, as wonderful hallmarks of human friendship.  

[7] Finally, the “cultural virtue versus biological selfishness” claim attributes an utterly and idealistically disembodied goodness and autonomy to culture. Even if culture altogether transcended biology, that would not in itself elevate us to moral structures that value the welfare of others. We construct cultures from the Machiavellian to the merciful. But culture is not autonomous. Variability notwithstanding, there are widely shared central tendencies across human cultures— like the incest taboo— that plausibly reflect interaction between social-mediation and biological endowment.  

[8] So our evolved biology is not unilaterally inclined toward selfishness, nor is culture a transcendent source of goodness. And failing to consider how culture and biology interact makes us more vulnerable to their ambivalent influence. Pascal observes that “humans are neither angels nor beasts, and unfortunately… those who would act the angel, act the beast”: moral wisdom recognizes our unique capacities to envision the good but also our shared frailties in willing it into action.  

[9] This involves another issue of ongoing debate: understanding how our biology “votes.” More nuanced and more vigorous than Darwin’s and Spencer’s differences, there continues to be disagreement over the extent to which natural selection can establish traits via benefits to groups in addition to— or even over— those of unrelated individuals. Such “group selection” might better facilitate the development of morality as a direct, and sometimes self-relinquishing, adaptation for social function. Two caveats are important. First, driven by competition, neither individual nor group level selection readily gives rise to sacrificing for those outside the group, or “love your enemy.” But second, what constitutes a “group” is not fixed. Here culture comes in, because social practices and values can influence— and indeed enlarge— how we attribute group identity and how we construe the domain of moral concern.  

[10] Indeed, the very idea of a distinctly “biological vote” in our behavior may be naïve, since all traits— especially higher cognition— are shaped by social environment. And all the various evolutionary accounts of morality recognize that it has profoundly enlarged our fundamental capacities for sociality compared to other primates. Moreover, since morality’s origin, modulation over cultural history has progressively expanded the scale of social organization. “Love your enemy” may be intrinsically harder than “love your neighbor,” but we recognize more folks as neighbors now than we did in the Paleolithic.  

[11] That brings us, then, to a third debate and back to Gould’s use of Wonderful Life: is evolution truly a-directional? While evolution itself is amoral, a widely recognized series of major evolutionary transitions reflect a profound directionality relevant to the emergence of morality. Living organisms have evolved from simple (prokaryotic) to complex (eukaryotic) cells, from single to multicellular organisms, from asexual to sexual reproducers, from solitary individuals to eusocial communities, and from primate to human societies. Each of these transitions has entailed the emergence of new levels of collective function through obligate interdependence. And with the emergence of human sociality, the scale of cooperation has continued to increase, facilitated by moral solutions to cooperative dilemmas and (more controversially) perhaps belief in moral deities.   

[12] However, the emergence of morality is not the climax of a romance novel. Across scales of biotic organization—from genes to cells to social relations— the possibility of defecting on mutually beneficial interaction is ongoing. And the capacity for suffering due to social exclusion or betrayal emerges, then intensifies, over the course of evolution. Moreover, none of us plays a benignant role in the moral drama by rote recitation. Recall that George Bailey was torn by perceptions of tension between his own fulfillment and obligations to others. And George lived in the face-to-face community of Bedford Falls, not the teeming metropolis of New York City!  

[13] In writing this essay and considering the voluminous literature on these topics, I’m mindful that the good is considerably easier to talk about than to do. Thus, I appreciate the confession from Herbert Spencer, who, although he gave abundant license to pursuing self-interest, nevertheless acknowledged his own “daily” moral struggle, due “more from defect of will than of knowledge.” In seeking a beneficent society, it’s important to contest significant differences in moral understanding. But it may be no less important to charitably acknowledge our shared defects of will. The most insidiously divisive out-group attributions consist in deeming others not just to be errant in moral knowledge, but— unlike “us”— to be corrupt in moral will. Being mindful of this, we might even find that some enemies become neighbors.