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Through the 1860s natural selection was already being applied to the growth of society. A.R. Wallace saw cooperation strengthening the moral bonds within primitive tribes. Advocates of social Darwinism, in contrast, complained that modern civilization was protecting the “unfit” from natural selection. Francis Galton argued that particular character traits—even drunkenness and genius—were inherited and that “eugenics,” as it would come to be called, would stop the genetic drain. The trend to explain the evolution of human races, morality, and civilization was capped by Darwin in his two-volume The Descent of Man, and Selection in Relation to Sex (1871). The book was authoritative, annotated, and heavily anecdotal in places. The two volumes were discrete, the first discussing the evolution of civilization and human origins among the Old World monkeys. (Darwin’s depiction of a hairy human ancestor with pointed ears led to a spate of caricatures.) The second volume responded to critics like Argyll, who doubted that the iridescent hummingbird’s plumage had any function—or any Darwinian explanation. Darwin argued that female birds were choosing mates for their gaudy plumage. Darwin as usual tapped his huge correspondence network of breeders, naturalists, and travelers worldwide to produce evidence for that. Such “sexual selection” happened among humans too. With primitive societies accepting diverse notions of beauty, aesthetic preferences, he believed, could account for the origin of the human races.

Darwin’s explanation was also aimed partly at Wallace. Like so many disillusioned socialists, Wallace had become engaged in spiritualism. He argued that an overdeveloped human brain had been provided by the spirit forces to move humanity toward millennial perfection. Darwin had no time for that. Even though he eventually attended a séance with Galton and the novelist George Eliot (Marian Evans) at his brother’s house in 1874, he was appalled at “such rubbish,” and in 1876 he sent £10 toward the costs of the prosecution of the medium Henry Slade.

Darwin finished another long-standing line of work. Since studying the moody orangutans at London Zoo in 1838, through the births of his 10 children (whose facial contortions he duly noted), Darwin had been fascinated by expression. As a student he had heard the attacks on the idea that peoples’ facial muscles were designed by God to express their unique thoughts. Now his photographically illustrated The Expression of the Emotions in Man and Animals (1872) expanded the subject to include the rages and grimaces of asylum inmates, all to show the continuity of emotions and expressions between humans and animals.

The gentle Darwin elicited tremendous devotion. A protective circle formed around him, locked tight by Huxley and Hooker. It was they who ostracized detractors, particularly the Roman Catholic zoologist Saint George Jackson Mivart. Nor did Darwin forget it: he helped raise £2,100 to send a fatigued Huxley on holiday in 1873, and his pestering resulted in the impecunious Wallace’s being added to the Civil List in 1881. Darwin was held in awe by many, the more so because he was rarely seen. And when he was seen—for example, by the Harvard philosopher John Fiske, a privileged visitor to Down House in 1873—he was found to be “the dearest, sweetest, loveliest old grandpa that ever was.”

Darwin wrote his autobiography between 1876 and 1881. It was composed for his grandchildren, rather than for publication, and it was particularly candid on his dislike of Christian myths of eternal torment. To people who inquired about his religious beliefs, however, he would only say that he was an agnostic (a word coined by Huxley in 1869).

The treadmill of experiment and writing gave so much meaning to his life. But as he wrapped up his final, long-term interest, publishing The Formation of Vegetable Mould, Through the Action of Worms (1881), the future looked bleak. Such an earthy subject was typical Darwin: just as he had shown that today’s ecosystems were built by infinitesimal degrees and the mighty Andes by tiny uplifts, so he ended on the monumental transformation of landscapes by Earth’s humblest denizens.

Suffering from angina, he looked forward to joining the worms, contemplating “Down graveyard as the sweetest place on earth.” He had a seizure in March 1882 and died of a heart attack on April 19. Influential groups wanted a grander commemoration than a funeral in Downe, something better for the gentleman naturalist who had delivered the “new Nature” into the new professionals’ hands. Galton had the Royal Society request the family’s permission for a state burial. Huxley, who by taking over the public debate had preserved Darwin’s reputation of “sweet and gentle nature blossomed into perfection,” as a newspaper put it, convinced the canon of Westminster Abbey to bury the diffident agnostic there. And so Darwin was laid to rest with full ecclesiastical pomp on April 26, 1882, attended by the new nobility of science and the state.