Edward O. Wilson, the sometimes-controversial US biologist, professor and author known as "Darwin's natural heir," who studied insects and human behavior, has died at the age of 92, his foundation said Monday.

The Harvard University professor emeritus wrote hundreds of scientific papers and more than 30 books, focusing on ants, global conservation, and the link between human behavior and genes. Time magazine once wrote that Wilson had had "one of the great careers in 20th-century science." 

Condolences pour in 

On Monday, the E.O. Wilson Biodiversity Foundation announced his death Sunday in Massachusetts.

"Ed's holy grail was the sheer delight of the pursuit of knowledge," said Paula Ehrlich, president of the Biodiversity Foundation. "A relentless synthesizer of ideas, his courageous scientific focus and poetic voice transformed our way of understanding ourselves and our planet."

Steven Pinker, a cognitive scientist at Harvard, called him "a great scientist" on Twitter, complementing "his generosity and willingness to engage."

"It would be hard to understate Ed's scientific achievements, but his impact extends to every facet of society. He was a true visionary with a unique ability to inspire and galvanize. He articulated, perhaps better than anyone, what it means to be human," David J. Prend, chairman of the board at his foundation, said in a statement.

What did he study throughout his career?

Wilson first became known as "the ant man" for his work documenting how insects communicate through pheromones.

This work got more widespread attention with his 1975 book, "Sociobiology: The New Synthesis," which breathed fresh controversy into the "nature-versus-nurture" debate.

In the book, Wilson focused primarily on his theories of the genetic impact on animal behavior, but the last chapter proposed that there were genetic components to human behavior, as well. He cited areas like a predisposition for the division of labor based on gender, tribalism, male dominance and parental-child bonding. 

Critics were quick to denounce Wilson's theory, accusing him of providing excuses for social injustice, discrimination against women, or even violence or Nazism. Here, too, his nickname as "Darwin's heir" rang true, given how Darwin's theories on evolution were hijacked by typically racist political movements in the late 19th and early 20th centuries. 

Wilson would later say when challenged that he never meant to posit that genes determine all of human behavior, but "in rough terms ... maybe 10% of it," also saying that the scope of the backlash almost prompted him to quit his work. 

In 1991, he returned to his work on insects, winning his second nonfiction Pulitzer Prize for the book he co-wrote with with Harvard colleague Bert Holldobler, "The Ants."

In his 2006 book, "The Creation," he wrote that the fields of science and religion, "the most powerful social forces on Earth," should work together for the preservation of nature and the fight against climate change.

In recent years, he called for the protection of diverse species and ecosystems.

"Biodiversity [is] being eroded at an accelerating rate by human activity," he said in a 2014 lecture at North Carolina's Duke University. "And the loss is going to inflict a heavy price in wealth, security and spirit, unless we staunch it."

jc/msh (AP, AFP, Reuters)