Edward O. Wilson, a biologist and author who conducted pioneering work on biodiversity, insects and human nature — and won two Pulitzer Prizes along the way — died on Sunday in Burlington, Mass. He was 92.

His death was announced on Monday by the E.O. Wilson Biodiversity Foundation. A cause of death was not given.

Dr. Wilson was survived by his daughter, Catherine. He was preceded in death by his wife, Irene K. Wilson.

“Ed’s holy grail was the sheer delight of the pursuit of knowledge,” Paula J. Ehrlich, chief executive and president of the E.O. Wilson Biodiversity Foundation, and a co-founder of the Half-Earth Project, said in a statement. “A relentless synthesizer of ideas, his courageous scientific focus and poetic voice transformed our way of understanding ourselves and our planet.”

When Dr. Wilson began his career in evolutionary biology in the 1950s, the study of animals and plants seemed to many scientists like a quaint, obsolete hobby. Molecular biologists were getting their first glimpses of DNA, proteins and other invisible foundations of life. Dr. Wilson made it his life’s work to put evolution on an equal footing.

“How could our seemingly old-fashioned subjects achieve new intellectual rigor and originality compared to molecular biology?” Dr. Wilson recalled in 2009. He answered his own question by pioneering new fields of research.

As an expert on insects, Dr. Wilson studied the evolution of behavior, exploring how natural selection and other forces could produce something as extraordinarily complex as an ant colony. He then championed this kind of research as a way of making sense of all behavior — including our own.

As part of his campaign, Dr. Wilson wrote a string of books that influenced his fellow scientists while also gaining a broad public audience. “On Human Nature” won the Pulitzer Prize for general nonfiction in 1979; “The Ants,” which Dr. Wilson wrote with his longtime colleague Bert Hölldobler, won him his second Pulitzer in 1991.

Dr. Wilson also became a pioneer in the study of biological diversity, developing a mathematical approach to questions about why different places have different numbers of species. Later in his career, Dr. Wilson became one of the world’s leading voices for the protection of endangered wildlife.

Dr. Wilson, a professor for 46 years at Harvard, was famous for his shy demeanor and gentle Southern charm, but they hid a fierce determination. By his own admission, he was “roused by the amphetamine of ambition.”

Those ambitions earned him many critics as well. Some condemned what they considered simplistic accounts of human nature. Other evolutionary biologists attacked him for reversing his views on natural selection late in his career.

But while his legacy may be complicated, it remains profound. “He was a visionary on multiple fronts,” Sarah Blaffer Hrdy, a former student of Dr. Wilson’s and a professor emerita at the University of California, Davis, said in a 2019 interview.

Derrick Bryson Taylor contributed reporting.

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