After delighting in a diet rich in romaine lettuce and cactus fruit for over a century, Gramma, the cherished Galapagos tortoise and the oldest inhabitant of the San Diego Zoo, has passed away.

According to zoo representatives, Gramma, who was born in her natural environment, was estimated to be around 141 years old. She passed away on November 20.

While the exact date of Gramma’s arrival at the San Diego Zoo remains uncertain, officials note that she was transferred from the Bronx Zoo between 1928 and 1931, as part of the initial group of Galapagos tortoises.

The record for the oldest known Galapagos tortoise is held by Harriet, who lived at the Australia Zoo until she reached 175 years. Harriet was collected from the Galapagos Islands in 1835 as a young tortoise, about the size of a dinner plate. This suggests she hatched around 1830 and passed away in 2006.

The Galapagos tortoise species comprises 15 subspecies, of which three are considered extinct. The remainder are classified as vulnerable or critically endangered by the International Union for Conservation of Nature.

Significant strides have been made in breeding these tortoises in captivity over recent decades. Since 1965, over 10,000 young tortoises have been introduced back into the wild, as reported by the Galapagos Conservancy. These efforts have helped some subspecies recover from the verge 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.