Canadian zoologist Anne Innis Dagg at Brookfield Zoo Chicago in 2016. A trip to the zoo with her mother 80 years earlier inspired her trailblazing research into giraffes. Elaisa Vargas/Roundstone Communications

Anne Innis Dagg, a Canadian zoologist who broke new ground in animal research while studying giraffes in the wild and who later campaigned against institutional sexism after she was denied tenure by an all-male committee and told that women belonged in the home instead of the academy, died April 1 at a hospital in Kitchener, Ontario. She was 91.

The cause was complications from pneumonia, said Paul Zimic, the executive producer of “The Woman Who Loves Giraffes,” a 2018 documentary about her life.

An exuberant researcher who seemed as comfortable in the field as she was in the lecture hall, Innis Dagg had a lifelong fascination with giraffes that began when she was 3, when she encountered the long-necked animals for the first time during a visit to the Brookfield Zoo outside Chicago. She later told CBC Radio that when she asked for a book about giraffes, she was told one did not exist.

“So I thought, ‘Well, I’ll learn about giraffes and then I’ll write one.’”

A few years before Jane Goodall began her field studies on chimpanzees in Tanzania, and a decade before Dian Fossey started her research on mountain gorillas in Rwanda, Innis Dagg went to South Africa to study giraffes in the bush near Kruger National Park. She was only 23 when she arrived in 1956 and was considered the first scientist to study giraffes in the wild – and one of only a few researchers at the time to study any animal in its natural habitat.

“If you wanted to know about the species, you would watch it in the zoo, or you’d study it by looking at the bones or looking at museum specimens, trying to figure out the taxonomy,” said Fred Bercovitch, a comparative wildlife biologist on the board of the Anne Innis Dagg Foundation, a conservation and education group. Innis Dagg, he added, was “at the cutting edge” in focusing on animal behavior and ecology, doing research that entered the mainstream only in the 1960s.

For about nine hours a day over an eight-month span, Innis Dagg took notes on the way the world’s tallest land animals moved, ate, fought, socialized and cared for their young. She kept track of about 95 giraffes, using a 16-millimeter camera to film the peculiar way they walked and galloped. When one of the animals was killed, she conducted an autopsy of sorts, drying the intestines and measuring them at 256 feet.

For the most part, she tried to stay out of the way, observing giraffes from inside her car, a rickety Ford Prefect – bought for 200 pounds – that she called Camelo, after camelopardalis, a scientific name for the giraffe.

Beginning in 1956, Anne Innis Dagg spent months studying giraffes in South Africa, often from the confines of her used Ford Prefect. Photos and film footage from the trip were later featured in a 2018 documentary, “The Woman Who Loves Giraffes.” Alexander Matthew/Roundstone Communications

Innis Dagg summarized her findings in a 1958 scientific article, published in the Proceedings of the Zoological Society of London, that laid the groundwork for the book she had long dreamed of writing. Published in 1976 with co-author J. Bristol Foster, “The Giraffe: Its Biology, Behavior, and Ecology” was considered a landmark in the field, pulling together virtually everything that was known about the animals.

“Without her pioneering work, the study of giraffes would not have been as complete,” said Graham Mitchell, a zoologist and physiologist who drew on her research for his 2021 book “How Giraffes Work.” He called Innis Dagg “the doyenne of giraffe researchers,” adding in an email that through her research and writing, she “did much to make the world aware of these remarkable and threatened animals.”

But by the time Innis Dagg published “The Giraffe,” her academic career had been “sidetracked,” as she put it, “by the institutional sexism that was rampant in academia.”

She was working as an assistant zoology professor at the University of Guelph in Ontario, teaching, publishing and conducting research while raising three children with her husband when she was denied tenure in 1971 and told that she would have to leave her job.

The school’s tenure committee said her teaching was “not up to standard” and alleged that her more than 20 peer-reviewed research papers were not of a “desirable scientific sophistication.”

The only committee member to back her, zoology colleague Sandy Middleton, told the Toronto Star much later that he believed Innis Dagg had “ran into the old boys’ network,” which may have sought to punish her because of jealousy over her research.

Innis Dagg unsuccessfully appealed the decision, making headlines in Canadian newspapers after she accused the university of sexism. She noted that, around the same time, two other women who had been briefly employed as zoology professors were denied tenure, in what she believed was a way for the department “to save money, having many large classes taught by academics hired at the lowest rate, then replaced by others also beginning at the salary floor.”

Over the next few years, she tried and failed to secure another academic posting near her family in Toronto. She later said a dean at the University of Waterloo “told me he would never give a married woman tenure because she had a husband to support her.”

When she was passed over for a teaching position at Wilfrid Laurier University in Waterloo and learned that the position went to a man with less experience, she filed a discrimination claim with the Ontario Human Rights Commission. The case didn’t go anywhere, she said, and she turned down part-time positions that she believed the university had offered in an attempt at “conciliation.”

Looking for ways to finance her research, she took a part-time job in 1978 at the University of Waterloo, where she became an academic adviser in the independent studies program. The job helped her continue to work as an independent scholar – she conducted research on homosexuality in animals, the locomotion of camels and the impact of human development on Canadian wildlife – even as she branched into other fields, calling out sexism in books including “Harems and Other Horrors: Sexual Bias in Behavioral Biology” (1983) and “MisEducation: Women & Canadian Universities” (1988).

With few exceptions, like an appearance on the American game show “To Tell the Truth” in 1965, she was largely overlooked by the general public until 2014, when the CBC broadcast an hourlong radio documentary about her work. The feature inspired Alison Reid to make a documentary film, “The Woman Who Loves Giraffes,” which followed Innis Dagg on her first return trip to South Africa in almost 60 years.

Innis Dagg, by then in her 80s, became increasingly in-demand on the academic circuit, attending conferences, accepting honorary doctorates and giving interviews in which she sought to promote science education, especially for women and girls. She also championed conservation efforts for giraffes, which have faced dramatic population declines in recent decades amid habitat loss and poaching.

Anne Innis Dagg in her car in South Africa in 1956. The picture was featured in the documentary “The Woman Who Loves Giraffes.” Alexander Matthew/Roundstone Communications

In 2019, she was appointed a member of the Order of Canada. Earlier that year, she had returned to the University of Guelph, where her academic ambitions had first been thwarted, for a screening of the documentary.

The dean of the university’s College of Biological Science announced that a research scholarship for women had been created in her honor. A letter was also read from the school’s provost and vice president, Charlotte Yates, who wrote that she was extending “an overdue apology for the ways in which you and other women were treated by the institution.”

“Isn’t it weird?” Innis Dagg told the Star, marveling at the crowds that lined up to shake her hand or give her a hug after screenings. “I’ve been ignored my whole life, and just to find out now that I’m actually a person and people really think I’m interesting – it’s pretty amazing.”

The youngest of four children, Anne Christine Innis was born in Toronto on Jan. 25, 1933.

Her mother, Mary Quayle Innis, was an American-born writer and historian who also served as dean of women at the University of Toronto’s University College. Her father, Harold Innis, was a communication theorist who became the head of the political economy department at the University of Toronto; he also helped inspire her love of nature, once making a canoe trip to the Arctic to see beavers and bears.

After graduating from the Bishop Strachan School, a Toronto prep school, Innis Dagg studied at the University of Toronto, receiving a bachelor’s degree in biology in 1955 and a master’s degree in genetics in 1956.

Convinced that the best way to understand giraffes was to study them in the wild, she sent more than a dozen letters to African wildlife officials, looking for someone who might sponsor her research. The replies were not encouraging; some suggested that an unmarried young woman should not be traveling alone, and warned that she might encounter trouble from rhinos and other hazards of the savanna.

After she began signing her letters with a gender-ambiguous name, “A. Innis,” she finally found a sponsor: a rancher, Alexander Matthew, who invited her to stay at his property, Fleur de Lys farm, in apartheid-era South Africa.

By the time she arrived in the country, Matthew had realized from one of her subsequent letters that she was a woman and concluded that it was improper, as a married man with his family out of the country, to host a “girl” at his house. He eventually relented, according to Innis Dagg, after she wrote him letters “every other day for two or three weeks.”

On her way back home to Canada in 1957, she stopped in England and married her fiancé, Canadian scientist Ian Ralph Dagg. He chaired the physics department at the University of Waterloo before his death in 1993. Survivors include their three children, Hugh, Ian and Mary; a brother; and a grandson.

Innis Dagg received a Ph.D. in animal behavior from the University of Waterloo in 1967, using her footage of giraffes for a dissertation on animal gaits. She said that because she was unable to get a tenure-track job and was busy raising children, “there was little opportunity” for a return trip to South Africa.

Still, she was able to conduct fieldwork in the Sahara during the summer of 1973, when she studied camels in triple-digit heat.

“The Jeep I hired broke down in the desert,” she told the Star the next year, recalling her journey home. “I stayed with some nomads until I got a lift in a truck to the railway station. Then our train got derailed in a sandstorm.

“Apart from that, it was quite uneventful.”