Frans de Waal, primatologist who argued that apes also have culture, shame and gender roles – obituary

Dr Frans de Waal at a primate research centre in Atlanta, Georgia, 2007
Dr Frans de Waal at a primate research centre in Atlanta, Georgia, 2007 - JACK KEARSE

Frans de Waal, who has died of cancer aged 75, was a distinguished Dutch-American primatologist whose research not only changed our understanding of the animal world but raised important questions about the human condition.

To de Waal, as to other evolutionary biologists post-Darwin, chimps, bonobos, macaques and human beings are simply different types of ape but, according to de Waal, ones who share more characteristics than commonly supposed.

In a series of books de Waal argued that while animal primates, like humans, can be violent and aggressive, they are also capable of empathy and altruism and have a concept of fairness that lies at the foundation of the human moral compass. While the stereotypical aggressive alpha male does sometimes rise to the top in a chimpanzee society, his reign is usually short and ends with his murder or exile. The most common and successful alphas “are typically not necessarily the biggest, strongest, meanest ones around ... Most alphas protect the underdog, keep the peace and reassure those who are distressed.”

Chimps, he found, are even capable of guilt and shame, emotions that had been thought exclusively human. “The standard notion of humanity as the only form of life to have made the step from the natural to the cultural realm,” de Waal maintained, “is in urgent need of correction.”

Frans de Waal in 2021
Frans de Waal in 2021 - Catherine Marin

Early on these were provocative views, not so much because they offended the religious idea that humans alone bear the image of God, but because they were at odds with progressive theories that human beings are born blank slates, unfettered by biological determinism and free to design their own behaviour – an assumption regarded by some as indispensable for any hope of curing society’s ills. De Waal was also sometimes accused of anthropomorphism – attributing human characteristics to animals on scant evidence. He responded that the problem was human exceptionalism.

De Waal never shied away from controversy. His 2013 book The Bonobo and the Atheist asked whether religious belief was an essential component of human morality. He concluded that morality comes from within, and is part of human nature; the role of religion is secondary.

Then, in his last book Different: What Apes Can Teach Us About Gender (2022), he marched fearlessly into one of the most fraught debates of our age.


His central premise was that males and females behave differently and that those differences have a basis in biology. “When scientists have studied how monkeys respond to toys, it has turned out that their choices are not what you would call sexually neutral,” he wrote. “It turns out that monkeys mimicked the sex-linked preferences of human children. Cars, trucks and balls were chosen more by males; females preferred dolls.”

If gender roles have a biological basis, then the skills implied by those roles still have to be acquired by learning and imitation. Female ape orphans in zoos often have no idea what to do with newborns as adults: a female bonobo who grew up in a human home was mystified by the males with obvious erections she encountered on meeting her own kind.

“Gender fluidity”, meanwhile, is common in both animal and human societies. Bonobos, who are as close to humans genetically as chimpanzees, have sex frequently, much of it homosexual, and even where a particular trait is selected for in one sex it will also be present in members of the opposite sex to a lesser but still important degree; in all primate species there are males with more feminine characteristics and female tomboys. The flexibility to alter social roles enhances survival.

Just as in human societies, too, there are “outliers” who do not conform to gender stereotypes: a female chimpanzee named Donna would raise her hairy coat like a male, enjoyed wrestle-play with alpha males and showed no interest in mating with males; a capuchin monkey called Lonnie had sexual relationships that were exclusively gay. De Waal reckoned that between five and 10 per cent of chimpanzee populations are “gender non-conforming apes”.

Unlike in human society, however, chimpanzees and other animal primates are fully accepting of this diversity. In animals, de Waal said, “I don’t find the kind of intolerance we have in human societies.”

Are We Smart Enough to Know How Smart Animals Are?
Are We Smart Enough to Know How Smart Animals Are?

One of seven boys, Franciscus Bernardus Maria “Frans” de Waal was born on October 29 1948 in s’Hertogenbosch, in the Netherlands, where he trained as a zoologist and ethologist at the universities of Nijmegen, Groningen and in Utrecht where he took a PhD under the biologist Jan van Hooff, with a dissertation entitled “Agonistic interactions and relations among Java-monkeys”, concerning aggressive behaviour and alliance formation in macaques.

Subsequent research with the world’s largest captive colony of chimpanzees, at Arnhem Zoo, became the basis for his first book, Chimpanzee Politics: Power and Sex Among Apes (1982), which compared the behaviour of chimps involved in power struggles with that of human politicians and offered the first description of primate behaviour in terms of planned social strategies, reconciliation and coalition-building. Chimps, he revealed, often come together after fights and kiss and make up. Newt Gingrich would included de Waal’s book on the reading list he gave new Republican arrivals at the US House of Representatives in 1994.

In 1981 de Waal moved to the United States to join the Wisconsin National Primate Research Center, from where he moved in 1991 to Emory University in Atlanta, Georgia, where he became Professor of Primate Behavior in the Department of Psychology and director of the Living Links Center at the Yerkes National Primate Research Center.

Over four decades, in scientific papers and in a series of popular books, de Waal proceded to shatter long-held ideas about what it means to be an animal and a human. In 2007 Time magazine included him in its list of the 100 most influential people in the world.

Mama's Last Hug
Mama's Last Hug

Much of the popular appeal of his books lay in the anecdotes. He wrote about how two grizzled male chimps, normally sworn enemies, slung their arms around each other’s shoulders to form a barrage between a newborn and a threatening young alpha male, and about a bonobo named Kuni who picked up an injured starling, climbed a tree, spread the bird’s wings and then released it.

Perhaps the most affecting story, told in Mama’s Last Hug (2019), was about the final encounter between an ageing chimpanzee matriarch at Arnhem Zoo and Jan van Hooff, who had studied her over 40 years. Though on her deathbed, Mama was potentially dangerous so van Hooff approached cautiously.

In de Waal’s account, the chimp sensed his trepidation and hugged him, hooting softly in his ear and tapping with her fingertips on his neck and the back of his head, exactly as she would reassure a frightened chimpanzee infant. She died soon afterwards. Footage of their farewell has been viewed more than 10 million times online.

Among numerous honours, de Waal was a Knight of the Order of the Netherlands Lion and elected to the American Academy of Arts and Sciences and the US National Academy of Sciences.

In 1980 de Waal married Catherine Marin, who survives him.

Frans de Waal, born October 29 1948, died March 14 2024