If any modern scientist can claim to have carried the torch lit by Charles Darwin, it was Edward Osborne Wilson — usually known as "E.O."
Wilson pioneered the study of biological diversity, adding a theoretical dimension to nature conservation — which had previously seemed more like a moral crusade than a scientific endeavor.
He popularized the term "sociobiology," exploring evolutionary explanations of social behavior. And he turned it all into lucid prose, becoming one of the most effective science communicators of his time.
Wilson died on Sunday in Burlington, Massachusetts, according to a statement from the E.O. Wilson Biodiversity Foundation.
Like Darwin, Wilson stirred up a hornet's nest with his evolutionary theories. But in Wilson's case, the most violent attacks against him came not from religious opponents, but from fellow scientists.
After the publication of his 1975 book Sociobiology, Wilson was pilloried by the biologists Richard Lewontin and Stephen Jay Gould, who accused him of justifying eugenics, sexism, and rampant capitalism. But Wilson gave as good as he got, and emerged from the battle with his reputation intact.
Born in Birmingham, Alabama, in 1929, Wilson might have studied birds or mammals if not for an accident at the age of 7.
After piercing his right eye with a fishing hook, he didn't seek treatment and later had to have the lens removed. The accident left Wilson with poor distance vision, as he recounts in his 1994 autobiography, Naturalist. But his left eye remained sharp, leading him to focus on what he called "the little things."
Ants became his life's work — and led to his interest in biodiversity and the evolution of social behavior. As a Harvard fellow in the early 1950s, he traveled to Cuba, Mexico, and the South Pacific, marveling at the diversity of the ants he observed and speculating on how they could have evolved.
Back at Harvard, where he spent his entire career, these thoughts eventually coalesced into The Theory of Island Biogeography, a 1967 book written with the mathematical ecologist Robert MacArthur. Wilson and MacArthur explored how species diverge into new forms on islands, and showed how their risk of extinction is highest if those islands — which can be patches of natural habitat in a human-altered landscape — are small.
This idea still dominates the thinking of conservationists, as they strive to expand and connect fragments of remaining habitat to give wildlife a fighting chance of survival.
Ants are not only incredibly diverse, but they also boast some of the most complex societies in the animal kingdom.
Ant colonies with thousands of individuals work together to raise the young of a single queen. This had long posed a problem for Darwinian theory: In a natural world dominated by the struggle to survive and reproduce, why would female workers in social insect species have evolved to be sterile?
In 1964, the British biologist Bill Hamilton came up with an answer. He argued that evolution is really about the reproduction and survival of genes, rather than individuals. So a gene can spread if it causes behavior that favors copies of itself in an animal's relatives. This "kin selection" theory made sense as an explanation for insect societies, because sister ants and bees — thanks to a quirk of their genetics — share 75% of their genes by descent, rather than the usual 50%.
Hamilton was a great theorist, but a poor communicator. Wilson packaged the idea for wider consumption and united it with other work explaining the evolution of behavior. The final chapter of Sociobiology briefly discussed these ideas in the context of human society — too briefly, Wilson later admitted.
"I should have been more politically careful, by saying this does not imply racism, it does not imply sexism," Wilson told the Harvard Gazette in 2011.
The chapter enraged left-leaning academics led by his Harvard colleagues Gould and Lewontin, who published a blistering assault in the New York Review of Books invoking the horrors of Nazi Germany. Wilson was aghast, but fought back hard, accusing his critics of misrepresenting his positions.
Wilson followed up in 1979 with On Human Nature, which dealt with the evolution of human behavior in more depth, and won the first of his Pulitzer prizes. (The second was for The Ants, published in 1991 with fellow ant guru Bert Hölldobler.)
Wilson's interest in human nature also led him to ponder the centrality of religious belief in human societies.
He lost his Baptist faith in his teens, but in his later years tried to find common ground with evangelical Christians in preserving what they view as divine creation and what he called the "evolutionary epic."
Wilson didn't shy away from conflict. He spent the last few years of his life at odds with many evolutionary biologists after he rejected Hamilton's kin selection theory in a controversial 2010 paper published in Nature. The paper argued that kin selection had little value as a general explanation of sociality, and described insect colonies as "superorganisms."
This echoed an old idea called "group selection," which argued that social behavior can evolve because it benefits a group of animals, improving their survival over rival groups. In the gene-centric era, this had become a discredited theory. For the author of Sociobiology to embrace it, while rejecting kin selection, stunned his peers.
In a subsequent issue of Nature, 137 leading evolutionary biologists signed a letter denouncing Wilson's about-turn. And in a later article in Prospect magazine, Richard Dawkins of Oxford University added: "For Wilson not to acknowledge that he speaks for himself against the great majority of his professional colleagues is — it pains me to say this of a lifelong hero — an act of wanton arrogance."
Arrogance, or supreme intellectual confidence? History, and the relentless struggle between competing scientific ideas, will be the judge.