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Woodland Park Zoo’s African elephant, Watoto, on display for more than four decades, was euthanized Friday morning after Seattle zookeepers found her collapsed and unable to rise in an outdoor exhibit yard.

Watoto, 45, was discovered in the south yard of the Elephant Forest exhibit about 7 a.m. after zookeepers arrived to work. Attempts to lift her with cloth straps and heavy machinery failed.

Zookeepers and medical staff made the “difficult decision” to euthanize Watoto because her health continued to deteriorate, zoo officials said. A necropsy will be conducted to determine a possible cause of death. Results will be publicly released.

Watoto, which is Swahili for “children,” was plucked from the African wild at age 2 and transported to Seattle through an international animal exchange, which was a common practice at the time. Today, in most cases, U.S. zoos are prohibited from importing elephants from protected wildlife areas.

Two other female elephants, both Asian, remain at the zoo: Chai, 35, and Bamboo, 47. They are housed in a building from 1989 that is divided into four sections of confinement, the largest measuring about 23 by 38 feet, with about an acre of shared outdoor space.

Friday’s unexpected death elicited widespread public condolences. It also underscored the continuing debate in Seattle and nationally over the humaneness of elephant captivity.

As the world’s largest land mammal, elephants are also considered the most humanlike. They live as families — mothers and daughters bond for life — and possess memory as well as a wide range of emotions, including grief and love, research has shown.

Captive elephants typically have shorter life spans, according to research conducted at the Amboseli National Park in Kenya. Wild African female elephants have lived up to 70 years old, with 56 as the median age.

Seattle Mayor Ed Murray called the death “very sad news” in a statement Friday, but added, “At the same time, I do believe that today’s news should reopen a dialogue in this city about the proper habitat for elephants.”

Watoto’s death marks a continuing decline in the number of elephants remaining inside U.S. zoos. Breeding programs have encountered widespread failure, often from injury and disease. About 284 elephants remain in accredited U.S. zoos.

A Seattle Times investigation in 2012 found that for every elephant born in captivity, on average, two others die. The zoo industry has publicly maintained that elephants are “thriving” in captivity.

The Times found that Woodland Park’s youngest female, Chai, endured 112 unsuccessful attempts at artificial insemination. She gave birth in November 2000 after she was bred at the Dickerson Park Zoo in Missouri.

But the calf, Hansa, died at age 6 from an infectious herpes virus known as EEHV. Zoo officials remain uncertain how the deadly disease was transmitted.

Nationally, at least 26 zoos have closed or plan to phase out their elephant exhibits, including those in San Francisco, Detroit, Chicago and New York.

Nonetheless, Woodland Park Zoo announced in March a $3 million expansion plan to add more elephants, revamp an aging, landlocked exhibit and reactivate a breeding program.

At the time, Deborah Jensen, zoo president and CEO, said the zoo hoped to relocate Watoto to another accredited facility.

Jensen and other nationally prominent zoo officials have steadfastly defended elephant captivity as a way to raise public awareness and donations that help to preserve elephants in their natural habitats.

But dozens of Seattle activists and national animal-welfare organizations have pushed Woodland Park to retire its elephants to one of two nonprofit sanctuaries, in California or Tennessee, which provide lifelong care for the animals on open acres of rolling land.

Watoto had no known major medical problems, though she had lost a tusk years ago.

Seattle activist Alyne Fortgang, who has monitored Woodland’s program for nearly a decade, said, “Watoto’s suffering is over. She’s free at last.”

Michael J. Berens: 206-464-2288 or Twitter @MJBerens1