Gramma, the beloved Galápagos tortoise and the San Diego Zoo’s oldest resident, has died at an estimated age of 141.
Known for her sweet, shy demeanour and a diet of romaine lettuce and cactus fruit, she passed away on 20 November.
Born in her native habitat, Gramma arrived at the San Diego Zoo from the Bronx Zoo in either 1928 or 1931, as part of their inaugural group of Galápagos tortoises.
Throughout her extraordinary life, she witnessed two World Wars and the terms of 20 US presidents, captivating generations of visitors.
Her dedicated care specialists affectionately dubbed her "the Queen of the Zoo."
Sadly, Gramma had been suffering from age-related bone conditions that had recently worsened, leading to the difficult decision to euthanise her.
Many visitors have since shared memories on social media, recalling their first encounters with Gramma as children and later returning with their own families.

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.
Galápagos 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 Galápagos Conservancy. Some subspecies have been brought back from the brink of extinction.
In April, four baby Galápagos 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 Galápagos tortoise Goliath became a first-time father at the age of 135.
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