Cleo, the stately lioness and a popular mainstay at the Sacramento Zoo, has died. She was 19.

Cleo was “one of the Zoo’s most cherished and iconic animals,” officials said in a statement Tuesday, adding that her “impact on staff, visitors, and the conservation community will never be forgotten. She was adored by many over the years, and the Sacramento Zoo, along with Cleo’s dedicated care team, deeply appreciate all the love and support from our zoo family during this time.”

Zoo officials said the lioness had an assortment of ailments in recent years including a steadily advancing kidney disease as she aged. A recent examination confirmed that Cleo’s kidneys were failing.

“Over the last few months, Cleo began to show signs that her kidney disease was worsening and that it was beginning to impact her comfort, appetite, and routine,” Sacramento Zoo staff veterinarian Dr. Jenessa Gjeltema said in a statement Tuesday announcing Cleo’s death. “After careful consideration of all options, the team made the kind decision to euthanize Cleo.”

The African lioness arrived in Sacramento 17 years ago at just 2 years old. By the end of her life, she was one of the oldest lions in the Association of Zoos and Aquariums African lion Species Survival Plan population.

Cleo and longtime mate, Kamau, who preceded her in death in June 2023, at 16, were perennial favorites at the zoo, no more so than in 2014 when the couple produced a litter of lion cubs. The Sacramento Zoo celebrated the cubs’ first birthday — and the zoo’s 500,000th visitor — the following year.

Like his beloved mate, Kamau, too, was remembered as a “charismatic and iconic presence,” upon his death.

In 2019, the zoo built a larger habitat and enclosure for the pair. Double in size from its earlier environs, its length of glass wall allowed zoo visitors — and Cleo — up close views of each other. More than one image shows the iconic cat, a paw pressed up against the glass, seeking connection with the human on the other side.

Cleo’s caretaker for all of her 17 years in Sacramento, Tom Nakayama, felt a special bond with the African lioness.

“I feel like I had a genuine connection with Cleo,” he said. “She always seemed happy to see me, and I was always happy to see her.”