The Columbus Zoo announces the passing of its world-famous matriarch.

Colo, the oldest gorilla on record and the first born in captivity, has died at the Columbus Zoo and Aquarium. She celebrated her 60th birthday on Dec. 22 and had outlived her life expectancy by more than two decades, according to a zoo press release, which announced her death this morning.

She was a phenomenon when she was born in Columbus in 1956, grabbing headlines from around the world as the first western lowland gorilla born into a zoo. Columbus Monthly included a comic strip in our December issue that marked her six decades of life, as well as the September birth of her great-grandson, JJ.

But her story was more sensational than we could cover in just seven frames, going all the way back to her origin, the rogue act of a young zookeeper. Warren Thomas, a second-year veterinary student at the time, put Colo’s parents Mac and Millie together against the zoo director’s orders, according to the Columbus Zoo press release. Thomas also later found the newborn gorilla, still in her amniotic sac, and revived the lifeless baby with mouth-to-mouth resuscitation.

She gained the name Colo (after her hometown of Columbus) in a nationwide “Name the Baby Gorilla” contest. She became the mother of three, the grandmother of 16, the great-grandmother of 12 and the great-great grandmother of three. Colo lived her whole life at the Columbus Zoo and came to be the matriarch of the zoo’s family of gorillas, which included six of her own descendants: daughter Toni; grandson Mac; granddaughter Cassie; great-granddaughters Dotty and Nadami; and JJ.

Colo died overnight in her sleep, about six weeks after veterinary surgeons removed a malignant tumor from underneath her arm on Dec. 3. It’s not yet known if the cancer contributed to her death, according to the zoo’s release, and a necropsy will be performed to determine the official cause. The zoo has designated an area outside the entrance for those who would like to remember the beloved gorilla. Contributions in her honor can be made through the zoo’s website, give.columbuszoo.org/colo.

Her death comes less than a month after her 60th birthday celebration, which was attended by hundreds of fans and garnered national media attention, from People magazine to Men’s Journal. She also received a video birthday greeting from her celebrity admirers: Urban and Shelley Meyer, Betty White (who turned 95 today), Jack and Suzi Hanna, Wayne Newton, Jack and Barbara Nicklaus, Henry Winkler and Carol Burnett. Though they didn’t know it at the time, it was a fitting goodbye to the world’s most famous gorilla.

Check out the video of Colo’s celebrity birthday greeting on People’s website.