Oto, Japan’s oldest lion, died Monday of natural causes at the Toyohashi Zoo & Botanical Park in Aichi Prefecture, according to the zoo.

She was 25½ years old — equivalent to about 100 human years.

Born on Aug. 26, 1999, at Asahiyama Zoo in the city of Asahikawa, Hokkaido, Oto had been at the Toyohashi zoo since July 2001.

Despite her age, she retained a glossy coat and had no chronic illnesses.

However, by late February, she started to sleep more and eat less. The zoo made adjustments to her diet and environment to keep her as comfortable as possible in her final days.

“Beloved by lion enthusiasts across the country, Oto brought joy and lasting memories to countless visitors. We would like to express our deep gratitude to her for gracing us with her presence. May she rest in peace,” the zoo said in a statement on its official website.

The zoo will set up a floral tribute stand at the lion enclosure from March 11 to 20.