WASHINGTON (Reuters) - Last year, after a Minnesota dentist sparked an uproar by killing a popular lion named Cecil while on safari in Zimbabwe, the U.S. Fish & Wildlife Service placed similar African lions on the endangered species list, making it illegal to import them as trophies to the United States.

The tiger Arjuna is pictured with Moksha Bybee at Myrtle Beach Safari in Myrtle Beach, South Carolina, U.S, in this handout photo provided by MyrtleBeachSafari.com. MyrtleBeachSafari.com/Handout via Reuters

But for African lions and other threatened and endangered species, there’s an exception to this rule: Hunters, circuses, zoos, breeders and theme parks can get permits to import, export or sell endangered animals if they can demonstrate that the transactions will “enhance the survival” of the species.

Often, records show, this requirement is met in part by making a cash contribution to charity - usually a few thousand dollars. The practice has angered both animal-rights activists who say it exploits wildlife and exhibitors who describe the process as unfair and arbitrary.

In the last five years, the vast majority of the estimated 1,375 endangered species permits granted by the Fish & Wildlife Service involved financial pledges to charity, according to agency documents reviewed by Reuters.

For a $2,000 pledge, the Fish & Wildlife Service permitted two threatened leopard cubs to be sent from a roadside zoo to a small animal park. After a $5,000 pledge, the agency approved the transfer of 10 endangered South African penguins to a Florida theme park.

An application now under final consideration would permit a South Carolina safari park operator to send 18 endangered tigers to Mexico to participate in a multimillion-dollar movie – for a $10,000 donation to charity.

Craig Hoover, a senior Fish & Wildlife Service official, said his agency considers many factors before granting an endangered species permit – among them, a species’ biological needs, threats and population size. Charitable contributions to conservation programs are just one factor in granting permit evaluations, and not a requirement, he said.

“It’s not necessarily all that is considered,” said Hoover. “There may have been an education component, an outreach component, a captive breeding component.”

“INDIRECT BENEFITS” TO WILDLIFE

Under the Endangered Species Act, exception permits may be granted only “for scientific purposes or to enhance the propagation or survival of the affected species.”

According to a recent Fish & Wildlife Service document reviewed by Reuters: “Very few of the Endangered Species Act permits that we issue have direct benefits to the species in the wild. Most applicants provide an indirect benefit, such as monetary support, to meet the enhancement requirement.”

Late Friday, U.S. Representative Brendan Boyle, a Pennsylvania Democrat who serves on the House Foreign Affairs and Oversight Committees, asked the agency to halt the practice.

Boyle said exemptions to the endangered species law are intended for humanitarian or environmental purposes, such as providing medical attention to a wounded animal, not commercial uses. He said the charity pledges are “unreliable at best and amount to an empty promise in exchange for an exemption to our bedrock species conservation law.”

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The agency usually does not try to independently confirm that donations are actually made or that the charities, often located overseas, are worthy, an agency document says. “Typically, we rely on the applicant,” the document notes. Hoover said applicants supply this information through annual reports and agency grant programs.

“PAY TO PLAY” FOR ELEPHANTS

Last year, People for the Ethical Treatment of Animals sued the Fish & Wildlife Service over a 2014 endangered species permit issued to Tarzan Zerbini Circus of Webb City, Missouri. The permit allowed Tarzan Zerbini to take two elephants, Shelly and Marie, on a Canadian circus tour - on the condition that it pledge $15,000 annually to an elephant charity and raise another $50,000 annually from patrons.

“We call it ‘pay-to-play’ because that’s what exactly what’s going on, allowing these people to promise money in exchange for being able to harm endangered animals,” said PETA general counsel Jeff Kerr. “The Fish & Wildlife Service is actively conspiring and cooperating with people to violate the Endangered Species Act through this program.”

The agency defended itself against PETA’s claim that the process is illegal, but the lawsuit apparently triggered a government investigation of Tarzan Zerbini’s financial pledge. Records show the service determined that the circus had only contributed half the amount promised and had raised little, if anything, from patrons. On April 21, the permit was suspended. Last week, PETA withdrew the lawsuit.

Larry Solheim, a Tarzan Zerbini consultant who served as general manager for 26 years, said the circus made good-faith efforts to comply with its pledges. He said honest mistakes and misunderstandings caused the other half of the money to be contributed late and said technical issues hampered efforts to raise the $50,000 from patrons.

Solheim said the concept of requiring conservation efforts is a good idea. But he described the permit process as too focused on foreign donations. He called it “a game” that can resemble “political extortion.”

“You’re just essentially buying a permit if you pay this conservation fee,” he said. “It’s just totally subjective – if they want to have this kind of requirement, they need to have clear guidelines.”

John Cuneo, whose Hawthorn Corp leases endangered animals to circuses and is often criticized by PETA, said he has lost business for failing to promise to make the charitable payments.

“It makes me so mad,” Cuneo said. “It feels like a scam.”

Hoover, the agency official, said PETA and the animal exhibitors are wrong.

“TIGER ISLAND”

“We would deny any form of ‘pay to play policy’ is in place, formally or informally,” Hoover said. He added: “We would deny that we tell people they must” make charitable contributions, “but if they are engaging in activity where the import or export isn’t contributing to conservation, then there must be some other means by which they must be contributing conservation.”

The permit application to send 18 tigers to Mexico for a Hollywood movie was filed by Bhagavan Antle, who operates the Myrtle Beach Safari in South Carolina. Antle declined to name the people behind the movie, which is tentatively titled “Tiger Island.” The plot revolves around tigers living on an abandoned island, and a group of children who end up shipwrecked there.

The permit is still pending, but records show that Fish & Wildlife officials directed that Antle confirm a pledge of $10,000 to charity and a promise that the movie will have a conservation theme. He has agreed to do so, and said he thinks the agency’s process is good because it helps endangered animals. Antle said $10,000 is a fair contribution for the right to use 18 tigers on a multimillion-dollar motion picture.

“The movie company thinks it’s a hardship – to spend $10,000 for what used to be free,” Antle said. But he added, “If it becomes a big hit movie, that will change more hearts and minds than a $10 million contribution to conservation.”

PENGUINS TO MIAMI

Last year, the Fish & Wildlife Service approved the sale of 10 African penguins from a California theme park to the Miami Seaquarium.

“We are thrilled that our guests will be able to observe these fascinating creatures and at the same time learn about this endangered species and what we can do to help preserve our feathered friends,” Andrew Hertz, the Seaquarium’s general manager, said in a press release in February. A spokeswoman said Hertz wasn’t available for an interview.

The sea park built a new exhibit for the birds called “Penguin Isle,” a spectacle that includes a 9,000-gallon pool with an acrylic underwater swimming tunnel, allowing visitors to come “face to face” with the penguins, the release says. The Seaquarium, which averages about 600,000 customers a year, charges $99 for a family of four.

As part of the endangered species permit approval, the seller agreed to make an annual contribution for five years to a South African charity that rescues penguins soiled by oil spills.

The annual pledge: $1,000.