Jane Goodall, the trailblazing naturalist whose intimate observations of chimpanzees in the African wild produced powerful insights that transformed basic conceptions of humankind, has died. She was 91.

A tireless advocate of preserving chimpanzees’ natural habitat, Goodall died on Wednesday morning in California of natural causes, the Jane Goodall Institute announced on its Instagram page.

“Dr. Goodall’s discoveries as an ethologist revolutionized science,” the Jane Goodall Institute said in a statement.

A protege of anthropologist Louis S.B. Leakey, Goodall made history in 1960 when she discovered that chimpanzees, humankind’s closest living relatives, made and used tools, characteristics that scientists had long thought were exclusive to humans.

She also found that chimps hunted prey, ate meat, and were capable of a range of emotions and behaviors similar to those of humans, including filial love, grief and violence bordering on warfare.

In the course of establishing one of the world’s longest-running studies of wild animal behavior at what is now Tanzania’s Gombe Stream National Park, she gave her chimp subjects names instead of numbers, a practice that raised eyebrows in the male-dominated field of primate studies in the 1960s. But within a decade, the trim British scientist with the tidy ponytail was a National Geographic heroine, whose books and films educated a worldwide audience with stories of the apes she called David Graybeard, Mr. McGregor, Gilka and Flo.

“When we read about a woman who gives funny names to chimpanzees and then follows them into the bush, meticulously recording their every grunt and groom, we are reluctant to admit such activity into the big leagues,” the late biologist Stephen Jay Gould wrote of the scientific world’s initial reaction to Goodall.

But Goodall overcame her critics and produced work that Gould later characterized as “one of the Western world’s great scientific achievements.”

Tenacious and keenly observant, Goodall paved the way for other women in primatology, including the late gorilla researcher Dian Fossey and orangutan expert Birutė Galdikas. She was honored in 1995 with the National Geographic Society’s Hubbard Medal, which then had been bestowed only 31 times in the previous 90 years to such eminent figures as North Pole explorer Robert E. Peary and aviator Charles Lindbergh.

In her 80s she continued to travel 300 days a year to speak to schoolchildren and others about the need to fight deforestation, preserve chimpanzees’ natural habitat and promote sustainable development in Africa. She was in California as part of her speaking tour in the U.S. at the time of her death.

Goodall was born April 3, 1934, in London and grew up in the English coastal town of Bournemouth. The daughter of a businessman and a writer who separated when she was a child and later divorced, she was raised in a matriarchal household that included her maternal grandmother, her mother, Vanne, some aunts and her sister, Judy.

She demonstrated an affinity for nature from a young age, filling her bedroom with worms and sea snails that she rushed back to their natural homes after her mother told her they would otherwise die.

When she was about 5, she disappeared for hours to a dark henhouse to see how chickens laid eggs, so absorbed that she was oblivious to her family’s frantic search for her. She did not abandon her study until she observed the wondrous event.

“Suddenly with a plop, the egg landed on the straw. With clucks of pleasure the hen shook her feathers, nudged the egg with her beak, and left,” Goodall wrote almost 60 years later. “It is quite extraordinary how clearly I remember that whole sequence of events.”

When finally she ran out of the henhouse with the exciting news, her mother did not scold her but patiently listened to her daughter’s account of her first scientific observation.

Later, she gave Goodall books about animals and adventure — especially the Doctor Dolittle tales and Tarzan. Her daughter became so enchanted with Tarzan’s world that she insisted on doing her homework in a tree.

“I was madly in love with the Lord of the Jungle, terribly jealous of his Jane,” Goodall wrote in her 1999 memoir, “Reason for Hope: A Spiritual Journey.” “It was daydreaming about life in the forest with Tarzan that led to my determination to go to Africa, to live with animals and write books about them.”

Her opportunity came after she finished high school. A week before Christmas in 1956 she was invited to visit an old school chum’s family farm in Kenya. Goodall saved her earnings from a waitress job until she had enough for a round-trip ticket.

Jane Goodall gives a little kiss to Tess, a 5- or 6-year-old female chimpanzee, in 1997.

(Jean-Marc Bouju / Associated Press)

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She arrived in Kenya in 1957, thrilled to be living in the Africa she had “always felt stirring in my blood.” At a dinner party in Nairobi shortly after her arrival, someone told her that if she was interested in animals, she should meet Leakey, already famous for his discoveries in East Africa of man’s fossil ancestors.

She went to see him at what’s now the National Museum of Kenya, where he was curator. He hired her as a secretary and soon had her helping him and his wife, Mary, dig for fossils at Olduvai Gorge, a famous site in the Serengeti Plains in what is now northern Tanzania.

Leakey spoke to her of his desire to learn more about all the great apes. He said he had heard of a community of chimpanzees on the rugged eastern shore of Lake Tanganyika where an intrepid researcher might make valuable discoveries.

When Goodall told him this was exactly the kind of work she dreamed of doing, Leakey agreed to send her there.

It took Leakey two years to find funding, which gave Goodall time to study primate behavior and anatomy in London. She finally landed in Gombe in the summer of 1960.

On a rocky outcropping she called the Peak, Goodall made her first important observation. Scientists had thought chimps were docile vegetarians, but on this day about three months after her arrival, Goodall spied a group of the apes feasting on something pink. It turned out to be a baby bush pig.

Two weeks later, she made an even more exciting discovery — the one that would establish her reputation. She had begun to recognize individual chimps, and on a rainy October day in 1960, she spotted the one with white hair on his chin. He was sitting beside a mound of red earth, carefully pushing a blade of grass into a hole, then withdrawing it and poking it into his mouth.

When he finally ambled off, Goodall hurried over for a closer look. She picked up the abandoned grass stalk, stuck it into the same hole and pulled it out to find it covered with termites. The chimp she later named David Graybeard had been using the stalk to fish for the bugs.

“It was hard for me to believe what I had seen,” Goodall later wrote. “It had long been thought that we were the only creatures on earth that used and made tools. ‘Man the Toolmaker’ is how we were defined …” What Goodall saw challenged man’s uniqueness.

When she sent her report to Leakey, he responded: “We must now redefine man, redefine tool, or accept chimpanzees as human!”

Goodall’s startling finding, published in Nature in 1964, enabled Leakey to line up funding to extend her stay at Gombe. It also eased Goodall’s admission to Cambridge University to study ethology. In 1965, she became the eighth person in Cambridge history to earn a doctorate without first having a bachelor’s degree.

In the meantime, she had met and in 1964 married Hugo Van Lawick, a gifted filmmaker who had traveled to Gombe to make a documentary about her chimp project. They had a child, Hugo Eric Louis — later nicknamed Grub — in 1967.

Goodall later said that raising Grub, who lived at Gombe until he was 9, gave her insights into the behavior of chimp mothers. Conversely, she had “no doubt that my observation of the chimpanzees helped me to be a better mother.”

She and Van Lawick were married for 10 years, divorcing in 1974. The following year she married Derek Bryceson, director of Tanzania National Parks. He died of colon cancer four years later.

Within a year of arriving at Gombe, Goodall had chimps literally eating out of her hands. Toward the end of her second year there, David Graybeard, who had shown the least fear of her, was the first to allow her physical contact. She touched him lightly and he permitted her to groom him for a full minute before gently pushing her hand away. For an adult male chimpanzee who had grown up in the wild to tolerate physical contact with a human was, she wrote in her 1971 book “In the Shadow of Man,” “a Christmas gift to treasure.”

Jane Goodall plays with Bahati, a 3-year-old female chimpanzee, at the Sweetwaters Chimpanzee Sanctuary, north of Nairobi, on Dec. 6, 1997.

(Jean-Marc Bouju / Associated Press)

Her studies yielded a trove of other observations on behaviors, including etiquette (such as soliciting a pat on the rump to indicate submission) and the sex lives of chimps. She collected some of the most fascinating information on the latter by watching Flo, an older female with a bulbous nose and an amazing retinue of suitors who was bearing children well into her 40s.

Her reports initially caused much skepticism in the scientific community. “I was not taken very seriously by many of the scientists. I was known as a [National] Geographic cover girl,” she recalled in a CBS interview in 2012.

Her unorthodox personalizing of the chimps was particularly controversial. The editor of one of her first published papers insisted on crossing out all references to the creatures as “he” or “she” in favor of “it.” Goodall eventually prevailed.

Her most disturbing studies came in the mid-1970s, when she and her team of field workers began to record a series of savage attacks.

The incidents grew into what Goodall called the four-year war, a period of brutality carried out by a band of male chimpanzees from a region known as the Kasakela Valley. The marauders beat and slashed to death all the males in a neighboring colony and subjugated the breeding females, essentially annihilating an entire community.

It was the first time a scientist had witnessed organized aggression by one group of non-human primates against another. Goodall said this “nightmare time” forever changed her view of ape nature.

“During the first 10 years of the study I had believed … that the Gombe chimpanzees were, for the most part, rather nicer than human beings,” she wrote in “Reason for Hope: A Spiritual Journey,” a 1999 book co-authored with Phillip Berman. “Then suddenly we found that the chimpanzees could be brutal — that they, like us, had a dark side to their nature.”

Critics tried to dismiss the evidence as merely anecdotal. Others thought she was wrong to publicize the violence, fearing that irresponsible scientists would use the information to “prove” that the tendency to war is innate in humans, a legacy from their ape ancestors. Goodall persisted in talking about the attacks, maintaining that her purpose was not to support or debunk theories about human aggression but to “understand a little better” the nature of chimpanzee aggression.

“My question was: How far along our human path, which has led to hatred and evil and full-scale war, have chimpanzees traveled?”

Her observations of chimp violence marked a turning point for primate researchers, who had considered it taboo to talk about chimpanzee behavior in human terms. But by the 1980s, much chimp behavior was being interpreted in ways that would have been labeled anthropomorphism — ascribing human traits to non-human entities — decades earlier. Goodall, in removing the barriers, raised primatology to new heights, opening the way for research on subjects ranging from political coalitions among baboons to the use of deception by an array of primates.

Her concern about protecting chimpanzees in the wild and in captivity led her in 1977 to found the Jane Goodall Institute to advocate for great apes and support research and public education. She also established Roots and Shoots, a program aimed at youths in 130 countries, and TACARE, which involves African villagers in sustainable development.

She became an international ambassador for chimps and conservation in 1986 when she saw a film about the mistreatment of laboratory chimps. The secretly taped footage “was like looking into the Holocaust,” she told interviewer Cathleen Rountree in 1998. From that moment, she became a globe-trotting crusader for animal rights.

In the 2017 documentary “Jane,” the producer pored through 140 hours of footage of Goodall that had been hidden away in the National Geographic archives. The film won a Los Angeles Film Critics Assn. Award, one of many honors it received.

In a ranging 2009 interview with Times columnist Patt Morrison, Goodall mused on topics from traditional zoos — she said most captive environments should be abolished — to climate change, a battle she feared humankind was quickly losing, if not lost already. She also spoke about the power of what one human can accomplish.

“I always say, ‘If you would spend just a little bit of time learning about the consequences of the choices you make each day’ — what you buy, what you eat, what you wear, how you interact with people and animals — and start consciously making choices, that would be beneficial rather than harmful.”

As the years passed, Goodall continued to track Gombe’s chimps, accumulating enough information to draw the arcs of their lives — from birth through sometimes troubled adolescence, maturity, illness and finally death.

She wrote movingly about how she followed Mr. McGregor, an older, somewhat curmudgeonly chimp, through his agonizing death from polio, and how the orphan Gilka survived to lonely adulthood only to have her babies snatched from her by a pair of cannibalistic female chimps.

Jane Goodall in San Diego.

(Sam Hodgson / San Diego Union-Tribune)

Her reaction in 1972 to the death of Flo, a prolific female known as Gombe’s most devoted mother, suggested the depth of feeling that Goodall had for the animals. Knowing that Flo’s faithful son Flint was nearby and grieving, Goodall watched over the body all night to keep marauding bush pigs from violating her remains.

“People say to me, thank you for giving them characters and personalities,” Goodall once told CBS’s “60 Minutes.” “I said I didn’t give them anything. I merely translated them for people.”

Woo is a former Times staff writer.

DAVENPORT, Calif. — On a spit of sand 12 miles north of Santa Cruz, a small, emaciated sea lion lay on its side. The only sign of life was the deep press of its flippers against its belly, relaxing for a few seconds, then squeezing again.

“That’s a classic sign of lepto,” said Giancarlo Rulli, a volunteer and spokesperson with the Marine Mammal Center, pointing to the young animal’s wretched self-embrace. The corkscrew-shaped bacteria, leptospirosis, causes severe abdominal pain in sea lions by damaging their kidneys and inflaming their gastrointestinal tracts. “They hold their stomach just like that. Like a sick child with a bellyache,” he said.

Since the end of June, officials say nearly 400 animals have been reported stranded or sickened along the Central Coast beaches. More than two-thirds of them have died, Rulli said. Hundreds more probably were washed away before anyone spotted them, or died at sea.

The historically large and long bacterial outbreak is adding to an already devastating death toll for the seals, sea lions, dolphins, otters and whales who live in and migrate through the state’s coastal waters.

There are the poisonous algal blooms off the central and southern coasts. There are massive changes in food availability and distribution across the Pacific. And there are growing casualties from ship strikes, record numbers of entanglements in rope and line, and a new heat blob forming in the eastern Pacific.

This year may be remembered as one of the gravest for marine mammals on record. Or, more worryingly, a sign that our ocean environment is changing so drastically that in some places and seasons, it’s becoming uninhabitable for the life it holds.

The network of volunteers who tend to stranded marine life is running ragged, said Rulli, answering dozens of rescue calls a day. “It’s been a brutal year. … It’s been hard on the animals. It’s been traumatic for the volunteers. It’s a lot.”

Whether all of these pressures and changes are related, or are completely separate phenomena happening at the same time in the same place, scientists don’t know.

“We’re trying to build our understanding of how ocean conditions relate to the occurrence of disease. But it’s a work in progress. And the world is changing quickly underneath our feet,” said Jamie Lloyd-Smith, an ecologist and evolutionary biologist at UCLA.

The first outbreak of leptospirosis in sea lions was reported along the West Coast in 1970, said Katie Prager, a disease ecologist at UCLA. By the 1980s, the Marine Mammal Center and others were keeping comprehensive records. They found that the bacterium tended to cause small, annual outbreaks that started in late summer and lasted just a month or two.

Dr. Alissa Deming, left, and veterinarian assistant Malena Berndt give anti-seizure medicine to a California sea lion named Patsy in a recovery room at the Pacific Marine Mammal Center in Laguna Beach after it had seizures from toxic algae blooms in June 2023.

(Allen J. Schaben / Los Angeles Times)

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Every three to five years, however, they’d see a large outbreak in which scores of animals got sick. In 2011 and 2018 during the last two big outbreaks, roughly 300 animals were rescued, Rulli said.

Lloyd-Smith and others say such leptospira-booms are probably driven by typical population dynamics — such as when a large enough cohort of never-exposed young animals get it and pass it around on beaches where the highly social animals congregate.

But this year, the outbreak started more than a month earlier than usual, and the number of sickened animals has surpassed any previously recorded outbreak.

This year seems deadlier, too, Rulli said. Leptospirosis typically kills some two-thirds of the animals it sickens. It’s only an impression at this point, but this year it seems to him like even more.

Looking at the sick pup on the Davenport beach, Rulli shook his head and said the animal was about as sick as he’d ever seen.

The little sea lion was humanely put down soon after it was taken to the Marine Mammal Center’s Castroville clinic, noted on a white board only as “Nameless Carcass.”

Why this year’s outbreak has been so devastating is not clear.

Lloyd-Smith and Prager said the leptospira species that affects sea lions is also found in some terrestrial mammals — such as raccoons, skunks and coyotes. Whether these scavengers are introducing new strains of the bacteria to sea lions on beaches, or the other way around, is not known. Nor is the bacteria’s natural reservoir — an area of research Lloyd-Smith is actively pursuing.

Jeremy Alcantara of the Marine Mammal Center nets an injured sea lion on a dock in Capitola.

(Nic Coury / For The Times)

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On a floating dock below the Capitola wharf, two groups of sea lions were lying down in the unusually sticky, humid air of a recent late-September afternoon. Seven were spooning one another in two small clusters — their flippers outstretched on each other’s bodies, their heads resting on their neighbors’ tummies or backs.

One rested at a distance from the others. It was the one someone had called in about.

For the rescue team, it was the third stop of the day, and it would be another tough one. A quick scan of the eight sea lions showed that another also looked unwell, her hip bones and vertebrae jutted jarringly underneath her blubberless skin.

The rescuers tried to catch the solo sea lion by nabbing her with a large fishing net, but she managed to squirt out of it. Veteran rescuers Jeremy Alcantara and Patrick McDonald regrouped with the others up on the wharf. They decided they’d try for the bony sea lion sunbathing with her friends.

Members of the Marine Mammal Center carry an injured sea lion on the pier in Capitola.

(Nic Coury / For The Times)

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Since April, the state’s stranding network of volunteer rescue crews has been responding daily to calls about sick sea lions, dolphins, whales, sea turtles and birds.

On the Southern California coast, there was a historic domoic acid outbreak that sickened more than 2,100 animals.

In the Bay Area, there was a record-breaking number of dead gray whales.

And from San Diego to Crescent City, they saw an off-the-chart number of whale entanglements — humpbacks and gray whales caught in the ropes and lines of the region’s commercial fisheries.

Now there’s worry that a growing marine heat wave in the Pacific could make things even worse — just as the Trump administration has threatened to pull funding from the National Oceanic and Atmospheric Administration, which provides financial help, research and oceanic data for the beleaguered animal crews.

“Fortunately, these volunteers don’t give up,” Rulli said. “They’re completely dedicated.”

Alcantara and McDonald descended the stairs from the wharf to the floating dock, taking roughly 10 minutes to quietly approach the sunbathing sea lions. The skinny one they were after had her flippers tucked tight against her belly.

A curious gull watched from the water. Tourists and locals gawked from above.

With a swoop of the net they caught her, carried her up the ramp to the wharf, quickly maneuvered her into a crate and then the back of an air-conditioned van that drove her to Castroville, where she was pumped with antibiotics and fluids.

She’s now at the center’s headquarters hospital in Sausalito, said Rulli. But “has not been receptive to offers of sustainable ground herring.”

Woodrow, as she has been named, is stable and the center’s veterinary staff will assess her again.

The explosion and hours-long fire at Chevron’s refinery Thursday night in El Segundo deeply unnerved communities in the South Bay.

The blast sent shock waves throughout the refinery grounds, allegedly injuring at least one worker, and jolting residents as far as a mile away. A 100-foot-tall pillar of fire cast an orange glow over the night sky. And towering plumes of smoke and acrid odors drifted eastward with the onshore winds.

While local regulators are investigating the fire, environmental advocates lament that federal safety agencies likely won’t be joining in the effort to find the cause of Thursday’s explosion — perhaps preventing similar hazardous chemical releases in the future. The incident was one of the most perilous events in the refinery’s 114-year history, adding to a long list of environmental and safety violations, according to public records reviewed by The Times.

Most staff at the Occupational Safety and Health Administration, the federal agency tasked with investigating workplace safety, is not working because of the ongoing federal shutdown. The U.S. Chemical Safety and Hazard Mitigation Board, which determines root causes from dangerous chemical releases, is also furloughed and could lose its funding because of proposed budget cuts by the Trump administration.

“The Trump administration has defunded the Chemical Safety board, and the federal government is shut down right now,” said Joe Lyou, a resident of nearby Hawthorne and president of the Coalition for Clean Air, a statewide nonprofit. “So there is a very good possibility we are never going to know what really caused this, because the experts in figuring this stuff out are no longer there to do that.”

Without clear answers, labor unions are fearful that a similar disaster could endanger thousands of workers at California’s 15 refineries, which are mostly clustered in Southern California and the Bay Area.

“Companies are making billions in profits and still are making it nearly impossible to make sure we’re safe from terrible disasters,” said Joe Uehlein, board president of the Labor Network for Sustainability. “In California, we’ve seen horrific injuries to workers and tens of thousands of residents have had to seek medical attention in refinery accidents. This time, we got lucky.”

The Chemical Safety Board has identified causes of scores of refinery incidents over its history, including the 2015 explosion at the ExxonMobil refinery in Torrance that injured at least two workers.

In that incident, the board’s investigation found multiple safety failures, including a severely eroded safety valve that allowed flammable gases to dangerously seep into unwanted areas. The board also discovered that a large piece of debris almost struck a tank of hydrofluoric acid, which could have resulted in a deadly release of the highly toxic chemical, leading to pressure to cease using the chemical.

But, for the Chevron refinery explosion, there is no guarantee such an investigation will take place. The Trump administration proposed eliminating the budget for the Chemical Safety Board this fiscal year, starting Oct. 1, sunsetting the 27-year-old federal agency. Environmental advocates say that is a mistake.

“They’re undermining our ability to prevent these accidents by taking away the accountability mechanisms in the federal government,” said Lyou. “That’s a huge concern. It’s not politics. Democrats and Republicans live around the Chevron refinery, and they both want to make sure that the refinery is operating safely.”

In the absence of federal regulators, the South Coast Air Quality Management District is investigating potential violations of air quality rules and permit conditions. The refinery will also be required to submit a report analyzing potential causes and equipment breakdowns within 30 days.

So far, the air district has said the fire originated in the refinery’s ISOMAX hydocracking unit, which uses hydrogen to refine oil into jet fuel and diesel. The refinery’s air monitors detected a spike in airborne chemicals after the fire broke out, but air district officials say conditions returned to normal levels after a few hours.

Environmental advocates say the extent of the fallout may not be known until there is a larger examination of air quality monitors.

“I was very surprised that the air district reported they weren’t seeing terribly high levels of pollution,” said Julia May, senior scientist for California-based nonprofit Communities for a Better Environment. “Sometimes in a big refinery fire like this, it goes straight up. But then the smoke comes down in other areas. And that’s a lot of pollution that’s going someplace.”

The Chevron facility had been cited numerous times for environmental and safety violations, according to local and federal records.

The South Coast Air Quality Management District has issued 13 notices of violations over the last 12 months, and 46 in the last five years. Most recently, on Sept. 22, the air district cited the facility for a large chemical leak and failing to keep its equipment in proper working condition.

In August, Chevron representatives had also asked the air district for leniency in assessing compliance with air quality rules while it was working to remove unwanted buildup inside its furnace tubes — conditions that they said risked equipment overheating and potentially failing.

OSHA records show the agency conducted at least 15 inspections at the Chevron refinery in El Segundo over the last decade, identifying 17 violations.

In September 2023, OSHA issued citations related to heat illness prevention requirements, ladderway guardrails and a failure to conduct a thorough hazard analysis — an internal assessment intended to control fires, explosions and chemical releases.

In October 2022, after conducting a planned inspection of the Chevron refinery, OSHA records show the agency identified a “serious” violation of an agency standard requiring employers to “develop, implement and maintain safe work practices to prevent or control hazards,” such as leaks, spills, releases and discharges; and control over entry into hazardous work areas.

During the government shutdown, it’s unclear if OSHA’s pared-down staff will be investigating Thursday’s refinery fire. An OSHA media office phone number went straight to a recorded message stating that the line is not being monitored and “due to a loss of funding, certain government activities have been suspended and I’m unable to respond to your message at this time.”

For some environmentalists, the Chevron refinery fire has underscored why it’s necessary to transition away from fossil fuels altogether.

“They [the refineries] have great workers and great fire departments to respond, but this is an inherently dangerous operation that handles hundreds of thousands of barrels per day of flammable explosive materials under high temperature and high pressure,” said May, the senior scientist for Communities for a Better Environment.

“When something goes wrong, you can have a runaway fire. They did a great job at getting it under control. But do we really want antiquated dirty energy in our communities?”