SAN DIEGO—Gramma, the Galápagos tortoise who became the San Diego Zoo’s living emblem of endurance, was euthanized last week at an estimated age of 141 years, marking the end of an era for one of the world’s oldest zoo residents. Her death, attributed to progressive bone deterioration, closes a chapter that spanned the Roaring Twenties, the Great Depression, two world wars, and the digital age. Zoo officials described her as the ‘Queen of the Zoo,’ a title earned through nearly a century of quiet majesty amid Balboa Park’s enclosures.

Arriving between 1928 and 1931 as part of the zoo’s inaugural shipment of Galápagos tortoises, Gramma witnessed the institution’s evolution from a modest collection to a global conservation powerhouse. The Los Angeles Times reported that she was among the first of her kind brought to the Balboa Park campus, her lifespan bridging more than 20 U.S. presidents from Herbert Hoover to Donald Trump. Her passing, confirmed by the zoo on November 21, underscores the tortoise’s iconic status in reptile husbandry and captive longevity studies.

A Relic of Early 20th-Century Conservation

The San Diego Zoo, founded in 1916, acquired Gramma during a period when Galápagos tortoises—Chelonoidis niger—were being imported en masse for zoos and private collections, often with scant regard for sustainability. According to the zoo’s own records, several tortoises from that 1928 cohort remain, estimated at over 100 years old. Gramma’s hatch year is approximated at 1884, placing her birth amid Ecuador’s turbulent politics and the Galápagos Islands’ exploitation by whalers and settlers.

Her transport to California reflected the era’s fascination with exotic megafauna. The San Diego Union-Tribune detailed how she endured nearly a century on display, outliving fellow imports and becoming a fixture for generations of visitors. Zoo veterinarians monitored her closely in recent years as mobility waned, opting for humane euthanasia when quality of life diminished.

Longevity Lessons from a Giant

Galápagos tortoises are renowned for lifespans exceeding 100 years in the wild and up to 177 in captivity, as with Harriet, who died at 175 in Australia’s Great Barrier Reef Wonderland. Gramma’s case bolsters data on ectotherm resilience, with weights up to 417 kg and adaptations like flared shells for island-specific foraging. Conservationists note her generation’s role in breeding programs that have repatriated subspecies, such as the Española giant tortoise.

Posts on X from accounts like SanDiegoVille mourned her as a ‘devastating loss’ amid a tough year for zoo animals, highlighting her 141-year tenure as unmatched. Industry insiders point to her as a benchmark for enclosure design—spacious, vegetated habitats mimicking volcanic islands—which has informed standards at facilities worldwide.

Economic and Institutional Impact

For the San Diego Zoo Wildlife Alliance, Gramma represented billions in cumulative visitor revenue over decades, her presence a draw in an industry generating $20 billion annually in the U.S. alone, per the Association of Zoos and Aquariums. Her story amplified fundraising for Galápagos programs, including genetic rescue efforts for once-extinct lineages rediscovered on Fernandina Island.

The zoo’s tortoise program, bolstered by Gramma’s lineage, has produced hatchlings now bolstering wild populations. She spanned history, from world wars to economic booms. As zoos pivot toward ex-situ conservation amid climate threats, her legacy informs protocols for aging megafauna.

Behind-the-Scenes Husbandry Challenges

Veterinary teams managed Gramma’s decline with laser therapy, supplements, and custom ramps, per zoo statements. Bone deterioration, common in geriatric chelonians, stems from decades of load-bearing on massive shells. Experts compare her to wild counterparts, where predation absence and low-metabolism diets enable centenarian lifespans.

The Times of San Diego quoted zookeepers calling her the ‘original resident,’ a sentinel through expansions like the 2024 Elephant Odyssey. Her euthanasia reflects ethical shifts in zoo medicine, prioritizing welfare over spectacle.

Broader Conservation Ripple Effects

Gramma’s death arrives as Galápagos efforts intensify: 86 Española tortoises repatriated in 2023 (Galápagos Conservancy). Hybrids and purebreds from zoo stock, including San Diego descendants, restore ecosystems by controlling invasive plants. X discussions underscore public sentiment, with users noting her as a ‘190-year-old’ peer in viral tortoise lore, though Gramma’s verified age stands at 141.

The species’ 15 subspecies—12 extant—face habitat loss, yet captive assurance populations thrive. San Diego’s remaining ancients, per zoo estimates, continue Gramma’s watch, guiding research into reptile senescence and biotech parallels for human gerontology.

Legacy in a Changing Zoo Landscape

As animal welfare scrutiny rises—evidenced by recent U.S. legislation—Gramma exemplifies successful long-term care. Her era predates modern accreditations, yet her vitality validated early practices. SanDiegoVille framed 2025 as a ‘difficult year,’ linking her loss to broader menagerie challenges.

Industry leaders view her as a pivot point: from menagerie curiosities to ark-like repositories. With Galápagos tourism booming—$500 million yearly—zoos like San Diego bridge public engagement and field science, ensuring tortoises like Gramma endure beyond individual lifespans.