LOS ANGELES — Gramma, the cherished Galapagos tortoise and the oldest inhabitant of the San Diego Zoo, has passed away, leaving behind a legacy of over a century filled with her favorite meals of romaine lettuce and cactus fruit.

Zookeepers estimate that Gramma, born in her natural habitat, reached the remarkable age of 141 years before her death on November 20.

While the exact date of Gramma’s arrival at the San Diego Zoo remains uncertain, officials believe she joined from the Bronx Zoo in either 1928 or 1931, becoming part of their pioneering group of Galapagos tortoises.

Throughout the years, as the world evolved, Gramma captivated visitors with her gentle and reserved demeanor. Her lifespan spanned two World Wars and the tenures of 20 U.S. presidents.

Known fondly as “the Queen of the Zoo” by her caregivers, Gramma had been suffering from age-related bone issues that had intensified, leading to the difficult decision of euthanasia, according to zoo representatives.

Following the news of her passing, many shared memories on social media of their first encounters with Gramma during childhood and the joy of introducing her to their own children in later years.

Cristina Park, 69, said one of her earliest memories from her childhood was going to the San Diego Zoo when she was 3 or 4 years old and riding on the back of a tortoise. That’s no longer allowed, but the experience inspired her to keep a small desert tortoise as a pet and learn more about tortoise conservation.

“Just how amazing it is that they managed to live through so much,” Park said. “And yet they’re still there.”

Galapagos tortoises can live for over 100 years in the wild, and close to double that in captivity.

The oldest known Galapagos tortoise was named Harriet, who lived at the Australia Zoo until the age of 175. She was collected from the Galapagos Islands in 1835, when she was just the size of a dinner plate, according to the zoo. This means that she hatched somewhere around 1830, and she died in 2006.

Galapagos tortoises include 15 subspecies of tortoises from the islands, three of which were deemed extinct. The rest are all vulnerable or critically endangered, according to the International Union for Conservation of Nature.

Concerted efforts have been made to breed these tortoises in captivity over the past several decades, with more than 10,000 juveniles released to the wild since 1965, according to the Galapagos Conservancy. Some subspecies have been brought back from the brink of extinction.

In April, four baby Galapagos tortoises were born at the Philadelphia Zoo to first-time parents that were roughly 100 years old, a first in the zoo’s history. In June, Zoo Miami resident and Galapagos tortoise Goliath became a first-time father at the age of 135.