


Joy was making the move last month to meet a new rule established by the Association of Zoos and Aquariums. The nonprofit that accredits zoos, mostly in the U.S., is requiring facilities with one or two elephants to either add to their pack or get rid of their pachyderms by September 2016, citing research showing that elephants are social animals and belong in a group. The Greenville Zoo said it had to move Joy because it didn't have the money or space for more elephants.

But on the second night of her journey, Joy was found dead in the back of her custom climate-controlled tractor-trailer. She was 44 years old, surpassing the average life expectancy for a female African elephant in a zoo, about 38. Necropsy results are pending, but Joy's age and the stress of the move were likely factors, said Jeff Bullock, Greenville's director. "We had lots of high hopes for her," Mr. Bullock said. "It was like getting punched in the gut." Distraught fans left cards and flowers at her empty exhibit. The zoo plans to put a three-story bird tower in the space.

Malaika, an African elephant born in 1986, does yoga with zookeeper Dale Berryman at the Cheyenne Mountain Zoo in Colorado Springs, Colo. Benjamin Rasmussen for The Wall Street Journal

Greenville is the latest American zoo to bid goodbye to the elephant, long considered the heart of many zoos. Zoos are finding it tough to add to their herds. The U.S. elephant population is aging—more than two-thirds of the African elephants in zoos are over 30. Domestic breeding programs have failed to produce many babies, and ivory poachers have reduced the number of wild elephants available to zoos.

In 1989, more than half of the 147 zoos accredited by the AZA had elephants. Today, only one-third of the 224 accredited zoos have them. Zoos accredited by the group house a total of 304 elephants, though the number on display is lower because some males are segregated for behavioral reasons and some animals are in sanctuaries that are generally not open to the public.

Beyond herd size, the AZA's new requirements also call for more space for the animals. The Greenville Zoo said it would have had to devote one-third of its 14-acre space to elephants to comply.

Animal-rights advocates say they hope zoos phase out elephants altogether because captivity is inherently limiting. Hard surfaces can cause foot disease and arthritis, animal biologists say, and small enclosures can prompt stress-related behaviors such as repetitive rocking and swaying.

A zoo-led campaign called 96 Elephants is raising money to save wild elephants targeted by ivory poachers—as many as 96 African elephants are estimated to be killed every day. The African elephant population has fallen by two-thirds in 40 years to 420,000 elephants, according to 96 Elephants. Fewer than 50,000 Asian elephants remain world-wide, according to the International Union for Conservation of Nature, an advocacy group.

"The statistics are jaw-dropping," said Brendan Wiley, director of the Topeka Zoo in Kansas, which has a 54-year-old Asian elephant and a 44-year-old African elephant. "If we don't do something, unfortunately the future of elephants in zoos will mirror what's going on with elephants in the wild."

Kimba, an African elephant born in 1978, grabs hay from a hanging barrel in the elephant barn at the Cheyenne Mountain Zoo in Colorado Springs. Benjamin Rasmussen for The Wall Street Journal

The Baton Rouge zoo in Louisiana and the Niabi Zoo in Coal Valley, Ill., closed their elephant exhibits last year. Other zoos, including the Bronx Zoo in New York, say they plan to care for the elephants they have for the rest of their lives but have no plans to acquire other elephants. Many smaller zoos are becoming what zookeepers dub a boutique, with a targeted collection of animals, rather than a department store, with two or three of as many animals as possible. Greenville has a relatively large collection of monkeys, reptiles and birds.

Some midsize zoos, including those in San Antonio and Topeka, say they want to stay in the elephant game and plan to raise millions of dollars to build new exhibits with breeding programs. But neither zoo has a construction timeline or animal-acquisition plan. "We are making a big bet, but it's one our community believes in," said Topeka's Mr. Wiley.

Zoos help people learn first-hand how physically majestic elephants are and how threatened they can be in a rapidly changing world. "There is no bigger fan of Disney than me, but 'The Lion King' creates the perception that 'the wild' is still there," Mr. Wiley said. "Our hope is we can increase the population in zoos, and at the same time, have a positive effect elsewhere."

Other zoos are carving out a niche as destinations for aging elephants, including Cheyenne Mountain Zoo in Colorado Springs, where Joy was headed, and the Little Rock Zoo in Arkansas.

Sophie, right, and Babe are elephants at the Little Rock Zoo. Little Rock Zoo

Little Rock is the new home of Babe, 39, and Sophie, 45, who left the Niabi Zoo last October. They now spend their time foraging for acacia vines and alfalfa with 53-year-old Zina. They also get their joints and feet checked daily, and do enrichment activities like searching for treats in a "bagel tree."

"Babe and Sophie fit right in here," said Susan Altrui, Little Rock's marketing director. "We don't mind being a retirement home for elephants." The zoo served as the last home for three female elephants that have died there since 2011.

In Greenville, zookeepers are trying to come up with new ways to keep the pachyderm spirit alive. "We'd like to at least have a bronze elephant statue," Mr. Bullock said. "We want kids to at least get a feel for what it's like to see an elephant."

Lucky and Kimba wander the elephant exhibit in the Cheyenne Mountain Zoo. Benjamin Rasmussen for The Wall Street Journal

Write to Valerie Bauerlein at