Zoo employees and workers with the Wild Animal Sanctuary organization prepared the big cats and the other animals for the long journey.
Activists have long held the U.S. territory’s government responsible for the deaths and ill health of animals reported for more than a decade at the zoo in the western town of Mayaguez.
Wild Animal Sanctuary is overseeing the removal of the animals and other creatures at the zoo. They will be transferred initially to a refuge in Colorado until a permanent home is found for them.
Some of the smaller specimens like reptiles and some monkeys were already transferred, but the bigger mammals like the lions, the bear and the camel required special measures for traveling.
A government-appointed committee said that in the past decade two pumas died at the zoo. It also raised concerns about an underweight chimpanzee as well as a rhinoceros named Felipe that was limping and said there was a lack of shelter for animals.
In January, an American black bear named Nina who had stopped eating died from a heart attack at more than 20 years old. Black bears can live up to 35 years in captivity.
In March, federal authorities said they were dropping all investigations into the zoo after reaching an agreement with local officials to transfer the animals to sanctuaries on the U.S. mainland. The announcement angered many activists.
The zoo opened originally in 1954, but it has been closed since hurricanes Irma and Maria battered Puerto Rico in September 2017.