Darwin on the evolution of morality

Darwin argued for the biological basis of morality in his Descent of Man (1871). Beginning with the thesis of the continuity of man and animals, he tried to explain the origin of the moral sense, or conscience, as understood as an ability to discern right and wrong, and to feel guilty if one realizes to have done wrong. His argument is that, in any animal with social instincts and sufficient intellectual powers, a moral sense would be developed. Although Darwin's argument had some missing links, I try to show that his argument can be consistently reconstructed, in view of the recent development of evolutionary biology and behavioral ecology. As I understand, Darwin's basic tenet is reductionism via evolutionary processes (natural selection, in particular): morality can be reduced to a combination of non-moral factors, each of which can be shared with other animals; you do not have to assume that morality is sui generis.
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