Atheists often claim that evolution provides a rational basis for objective ethical standards. Don't count on it. It was a common line of thinking and claims about evolution for "the good of the group" were common. The downfall was when V.C. Wynne Edwards tried to make this intuition rigorous in his book Animal Dispersion in Relation to Social Behavior. By putting rigorous arguments on the table Wynne-Edwards made it easy to falsify the theory.
That is precisely what George Williams did. He pointed out that evolution for the good of the group created what economists call the free rider problem. Suppose a group of animals is facing a particular tough environment. The moral thing to do is for all the animals to reduce their food intake. That will make the group stronger and the members along with it. But suppose a free rider lacked this moral gene. It kept eating at its normal food intake. At the end of the winter (or dry season, or whatever) it would be stronger and healthier than the moral animals. This would result in increased reproductive success. The immoral free riders would have more offspring than the moral animals. Soon the moral gene would go extinct.
This does not deny the existence of kin selection (behaving altruistically towards family members who share your genes) and reciprocal altruism (behaving altruistically in exchange for a favor in return and/or social status). They clearly give reasons to act morally. But only in a fairly limited sphere. Only the most naive species of neoclassical economist would argue that all human interaction can be modeled with the assumption of perfect information. In the real world asymmetrical information is common and people can't know if you've behaved honestly or not. Hence the expression: character is doing the right thing even when nobody is watching.
One of the most striking cases where animals can further their self-interest at the expense of the group is through non-monogamous reproduction. Polygamy is very common in the animal kingdom in species such as gorillas, elephant seals, and many others. It results in one alpha male defeating weaker males and acquiring a harem of females. The first thing the conquering alpha does is kill the infants of the deposed alpha. That brings the females into heat sooner. The alpha male can procreate faster by killing the infants. Martin Daly and Margo Wilson wrote the book homicide to test this theory on humans. They found that stepfathers were 70 to 100 times more likely to kill their children than biological offspring. As D.S. Wilson points out, that number is probably too low. Many of the married "biological fathers" were likely cuckolded by their wives - they were killing another man's offspring.
Species with helpless offspring are not typically polygamous because helpless offspring need fathers. But males can procreate on the sly and cuckold other males. That allows them to pass on more of their genes at the expense of other males. Adultery is rational for monogamous females too - it allows them to "trade up" to higher quality genes. Not every woman can marry an alpha, but she can certainly copulate with one. Males develop various defense mechanisms like jealousy, mate-guarding, domestic violence, and cooperative cuckolding (if you can't beat 'em, join 'em). (See The Myth of Monogamy by evolutionary biologist David Barash for a good summary of the rational side of immoral behavior.)
David Sloan Wilson deserves enormous credit for braving the ridicule that used to await any biologist with the nerve to champion group selection. He created the theory of multilevel selection. It puts to rest the old debates over the proper unit of selection in biology: genes, individuals, or groups? The answer is 'yes.' Evolution can happen at all levels. Consider the case of hunter-gatherers. Contrary to secular progressive opinion, they are very warlike. About 25% of adult males die in warfare (the defining book is Lawrence Keeley's War Before Civilization, but I prefer Steven LeBlanc's Constant Battles.). In each tribe immoral free riders will prosper and pass on more of their genes than their moral peers. Selection is happening at the level of individuals (and genes). But tribes with a lot of free riders will not be able to cooperate as effectively as moral tribes. They will take heavy losses in battle. Thus at the level of groups selection is also happening and it favors moral behavior.
The lesson: strong hostilities between groups are essential to make group selection work. The price of group selection is a small circle of moral consciousness and a harsh distinction between those who are Like Us and The Other.
Sometimes evolution solves the free rider problem. Then selection at the lower level is effectively eliminated. This is what happens in our bodies. It is in the self-interest of each cell to reproduce like crazy even if it hurts the bodies overall health but this does not happen (except in dysfunctional cases like cancer). Some species have solved the free rider problem at the group level rather than the cellular level. These are the eusocial species and they are only insects such as ants and bees. There is only one eusocial animal species, the naked mole rat. E.O Wilson colorfully describes eusocial species as a superorganism.
Are eusocial species moral? D.S. Wilson says they are and defines morality "acting in ways that are good for the group." Conversely, immoral behavior is acting in ways that are bad for the group. I think that definition fails. For one thing, don't think of superorganisms as a utopia. Instead think of them as a totalitarian state in which the dictator found a way to gain utter and complete power over the masses, to the point at which the masses had no choice but to work for the group. Consider how our bodies solve the free rider problem. Cells with mutations are immediately killed. Cell lines that get too old are killed. You don't want to risk a cell deciding to do its own thing so strict conformity is demanded.
In fact, the essential feature of a superorganism is the formation of a reproductive caste which is allowed to reproduce. The non-reproductive castes are not allowed to reproduce. Sometimes female worker ants try to lay their own eggs but they are promptly eaten by other workers. When naked mole rat queens die there is a bloodfest as other females fight to become the new queen. The winner achieves her position over the dead bodies of her rivals. The losers who survive are of course forbidden from reproducing. Their only way to pass on their genes is indirectly - by sacrificing for the rest of the group. And given their denial of reproductive opportunity, sacrifice is their best strategy. It makes superorganisms enormously successful. Ants are the kings of the insect kingdom. Insect species that are not eusocial like ants are forced to inhabit minor niches.
It is a moot point because humans are obviously not a superorganism. We are not divided into reproductive and sterile castes. But the fact is that group selection is not morality. It suffers from two fatal flaws. The first is that it does not solve the problem of warfare with other groups. Ants are an extraordinarily warlike species. The second is that superorganisms are totalitarian. In fact, the only way that true morality is possible is if we have the power to do evil, but freely choose the good. That is also a rough definition of 'natural right' but I'll save that for another post.
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