E.O. Wilson, a Harvard biologist who pioneered the field of sociobiology, fiercely fought to protect species, and taught generations to look at the natural world and human behavior in novel and thought-provoking ways, died Sunday in Burlington, according to the E.O. Wilson Biodiversity Foundation. He was 92.
Dr. Wilson was a humanist and optimist at heart — and, on occasion, a pugilist — as he burrowed deeply into the insect world, particularly ants, and connected what he saw to more universal concerns about human behavior and humanity itself.
A scientist-author who appealed to non-academic readers with books such as “On Human Nature” and “The Ants,” both Pulitzer Prize winners, Dr. Wilson exhibited a native Southerner’s gift for storytelling, combined with a seemingly inexhaustible supply of energy and subjects to explore. He belonged to a select group of scientists — such figures as Stephen Hawking, Richard Feynman, and Stephen Jay Gould — whose influence and stature extended far beyond their classrooms, laboratories, and journal articles.

A mentor to countless scientists, he continued working and publishing well into his 80s and beyond. In 2017 alone, Dr. Wilson published two books, “Half-Earth: Our Planet’s Fight for Life” and “Origins of Creativity,” that reflected his wide-ranging intellectual interests and deepening concerns for the fate of the Earth.
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“The ideal scientist thinks like a poet and only later works like a bookkeeper,” he wrote in “Letters to a Young Scientist,” one of more than 30 books he wrote, co-wrote, or edited, both nonfiction and fiction.
He further urged aspiring scientists to “put passion ahead of training” and immerse themselves in underexplored fields of study, then prepare to work relentlessly.
“Real scientists do not take vacations,” he maintained; they take field trips.
According to Harvard entomologist Brian Farrell, who took over Dr. Wilson’s former university lab, his most lasting contributions include his work on island biogeography, a theory co-developed with ecologist Robert MacArthur. It proposes that the number of species found in an island environment reflects a balance between new colonizing species and ones consigned to extinction.
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Their theory was originally tested by eradicating all insect species from an uninhabited Florida island, then observing how new ones established themselves. Painstaking observation combined with bold leaps of imagination became trademarks of Dr. Wilson as he continued to explore the natural world and publish his findings.
More broadly, Farrell said, Dr. Wilson became a great synthesizer of fields as disparate as literature, the arts, and hard science — and someone unafraid when stirring up a hornet’s nest of controversy.
“What mattered to Ed is not whether you’re right but that you have this compass that drives you forward,” Farrell said. “Fearless is probably the single best word that captures his spirit.”

Edward Osbourne Wilson was born in Birmingham, Ala., on June 10, 1929. An only child, Sonny (as he was then called) was moved around a great deal in his youth, mostly growing up in Mobile, Washington, D.C., and Florida. His parents divorced when he was young, and he wound up attending 16 schools in 12 years.
At age 7, Sonny was fishing off a dock when a pinfish spine pierced his right eye. His vision was permanently impaired, adding to a hearing problem he already had. The accident changed a youngster already deeply interested in the natural world.
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“The attention of my surviving eye turned to the ground,” he recalled. “I would thereafter celebrate the little things of the world.”
A lifelong love affair with nature — ant species in particular — soon began.
In Washington, D.C., where his father worked for the Rural Electrification Administration, Sonny spent hours collecting bug and butterfly specimens in a nearby park.
“Adults . . . are prone to undervalue the mental growth that occurs during daydreaming and aimless wandering,” he later wrote. He began corresponding with Marion Smith of Washington’s National Museum of Natural History, who encouraged his study of ants, a field known as myrmecology.
After attending from Gulf Coast Military Academy, Dr. Wilson entered the University of Alabama, picking up two degrees before going to Harvard University, where he earned a PhD in 1955.
That same year, he married Irene Kelley, a poet. They settled in Lexington and had one child, Catherine, a professor of philosophy. Irene died in August. Catherine is Dr. Wilson’s only immediate survivor, according to his foundation.
During the 1940s and 1950s, he traveled throughout the Caribbean and South Pacific, studying and collecting numerous ant species.
His office at Harvard’s Museum of Comparative Zoology would grow to contain nearly 14,000 species of ants, reportedly the world’s largest collection. Among the first field scientists to discover how ant species and others communicated through chemical excretion, Dr. Wilson was credited with having identified and described more than 450 new species of ants.
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During the 1950s and ’60s, Dr. Wilson forged collaborations with several prominent scientists, among them mathematician William Bossert and MacArthur, the ecologist, with whom he published his first book, “The Theory of Island Biogeography,” in 1967.
In 1975, Dr. Wilson published “Sociobiology: The New Synthesis,” a groundbreaking work exploring the genetic and evolutionary roots of animal behavior. The book expanded upon his observations of insect behavior to consider more socially advanced species, including humans. In arguing for the existence of a gene-based human nature — thereby suggesting limits on how much humans are capable of remaking themselves — Dr. Wilson ignited a firestorm of protest from many left-leaning social scientists.
Branded a genetic determinist, he faced calls for his removal from the Harvard faculty, culminating in one protester dumping a jug of water on his head at a scientific conference. Among his most vocal opponents were Harvard colleagues Gould and Richard Lewontin, who accused Dr. Wilson of providing “genetic justification” for racism, sexism, and opposition to pro-gay policies.
Harvard molecular biologist James Watson, who co-discovered the molecular structure of DNA, became another high-profile adversary. Watson once described naturalists such as Dr. Wilson as “stamp collectors.” (Dr. Wilson returned the favor, calling Watson and his ilk “test tube jockeys.”)
Their feud was more than personal. It underscored the tension between competing approaches to biological research, one centered on laboratory observation, the other on field study. The two men later patched up their differences.
In his 1994 memoir, “Naturalist,” Dr. Wilson admitted he could be stubborn, even combative, when backed into a corner. As a child, “I never picked a fight,” he wrote. “But once started, I never quit, even when losing.”
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Other widely read books of Dr. Wilson’s include “Consilience: The Unity of Knowledge,” “The Diversity of Life,’’ which in 1993 predicted a wave of species extinctions, and “Anthill: A Novel.”
More recently, in “Half-Earth” and the global project it spawned, Dr. Wilson proposed setting aside half the Earth’s territory (land and water) for species conservation. He and his allies, among them the musician Paul Simon and economist Jeffrey Sachs, calculate that without a concerted effort, half the planet’s species could disappear by 2100.
During 2019-20, he published two more books, “Tales from the Ant World” and “Genesis: The Deep Origin of Societies.” He also is the subject of the 2021 biography, “Scientist: E.O. Wilson: A Life in Nature” by Pulitzer Prize-winning author Richard Rhodes.
A 2002 Boston Globe profile described him as “America’s foremost public defender of the natural world.”
“Ed is the most embracing, magical, positive, funny person I know,” said John “Ike” Williams, his longtime lawyer and book agent. “He’s the modern Darwin, a scientist who added a broader humanist element” to his work, much as Albert Einstein did in crusading against the use of nuclear weapons.
With books like “Consilience” — the term refers to the synthesis of knowledge gleaned from anthropology, sociology, psychology, and other fields — Dr. Wilson achieved two milestones, according to Williams. He reached an audience “not necessarily knowledgeable about science, but without dumbing it down,” he said. And he placed human beings in two important contexts: as the most destructive of species and as potential saviors of all other life forms.
At Harvard, Dr. Wilson held the title of Pellegrino University research professor emeritus and honorary curator in entomology. He also taught at Columbia and Drake universities and the University of California Berkeley, retiring from full-time teaching duties in 1996. He later helped establish the E.O. Wilson Biodiversity Foundation and Encyclopedia of Life, an online, open-source database.
Over his lengthy career, Dr. Wilson won numerous awards and honors, among them a National Medal of Science, Audubon Medal, Explorers Club Medal, Leidy Award, Carl Sagan Award for Public Understanding of Science, International Prize for Biology, TED Prize, Nierenberg Prize, Kistler Prize, Lewis Thomas Award for science writing, International Cosmos Prize, and Humanist of the Year Award.
In 1995, Time magazine named him one of America’s 25 most influential people. More than a quarter-century later, his influence as a thinker, writer, scientific observer, and planetary advocate remained strong. As did his unshakable optimism.
“To strive against odds on behalf of all of life,” he wrote in “Half-Earth,” “would be humanity at its most noble.”
Joseph Kahn can be reached at josephpkahn@gmail.com.
CORRECTION: In a previous version of this story, a quote was erroneously attributed to Harvard molecular biologist James Watson. The Globe regrets the error.