LOS ANGELES — After more than 100 years of indulging in her favorite treats of romaine lettuce and cactus fruit, Gramma, the cherished Galapagos tortoise and the oldest inhabitant of the San Diego Zoo, has passed away.

Zoo officials noted that Gramma was believed to be about 141 years old, having been born in her natural environment. She passed away on November 20.

Although the exact date of Gramma’s arrival at the San Diego Zoo is uncertain, records suggest she was transferred from the Bronx Zoo sometime between 1928 and 1931 as part of the zoo’s inaugural group of Galapagos tortoises.

Throughout her long life, Gramma charmed zoo visitors with her gentle and reserved nature, witnessing a century of change including two World Wars and the terms of 20 U.S. presidents.

Revered as “the Queen of the Zoo” by her caretakers, Gramma had been battling age-related bone conditions, which worsened recently, leading to the decision to euthanize her, according to the zoo.

Many visitors commented on social media about getting to first visit Gramma when they were young, and being able to come back years later with their kids.

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.