Lemurs have lost one of their greatest conservation proponents, Alison Jolly, the famed primatologist whose research in the 1960s proved for the first time that not all primates live in male-dominated societies. Jolly passed away on Feb. 6 at the age of 76, just a few days before a new paper predicted a lemur extinction crisis in Madagascar.

Jolly first traveled to Madagascar in 1963, where her studies resulted in several important papers and popular books. Chief among her observations were that right-tailed lemurs and about a dozen other species lived in female-dominated societies. All scientific research prior to that point indicated that primates—including humans—lived in exclusively male-dominated groups. "All females, whether dominant or subordinate in the female hierarchy, are dominant over males," Jolly wrote in her 1966 book, "Lemur Behavior: A Madagascar Field Study."

The discovery was greeted with resistance by the scientific community of the day. "This was a real surprise to people in the '60s,” primatologist Patricia Wright told the New York Times. "Female leaders were still so rare. And here comes a woman presenting a model of primates where the females are leaders — effective leaders."

Her work also detailed her theory that intelligence evolved not through ecological factors but through social behavior.

Jolly's studies also revealed an important aspect of conservation: preserving species also requires meeting the needs of the people who live in the same regions. This, too, was a controversial topic decades ago. Jolly and her husband Richard, a noted economics, wrote a paper for an international conference about conservation in Madagascar in 1970. They argued that the needs of people needed to be balanced with that of wildlife. The paper was so controversial that it was never officially published. It circulated unofficially for many years. "The paper circulated informally instead," her friend Alison Richard told Yale's alumni magazine. The paper helped "to establish an approach to conservation that included the needs and aspirations of people as well as the island's unique and endangered natural communities."

In addition to her research and writing, Jolly was known for mentoring young conservation scientists in Madagascar as well as for writing books to teach the Malagasy children about lemurs. She also served five years as president of the International Primatology Society and was, at the time of her death, a visiting scientist at the University of Sussex.

Jolly's legacy will live on. Her final book, "Saving Madagascar: Conservation Diaries of Alison Jolly," will be published posthumously. A restored mining forest was named for her just last month. And a species of mouse lemur, Microcebus jollyae, was named after her in 2006.

Jolly is survived by her husband, four children and four great-grandchildren.

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