The fate of two killer whales is uncertain after the marine zoo closed on Sunday.
Activists and zoo directors have been locked in a row over what should happen to the orcas, with the French government already blocking a proposal to reintroduce them.
Last month, Marineland Antibes, located near Cannes on the French Riviera, said it would close permanently on January 5 following new animal welfare laws.
The legislation, which bans the use of dolphins and whales in marine zoo shows, was passed in 2021 but goes into effect next year.
Marineland, which describes itself as the largest of its kind in Europe, currently keeps two killer whales – Wiki, 23, and her 11-year-old son Keigo.
Managers say shows featuring killer whales and dolphins attract 90% of Marineland's visitors – and that without them the business would not be viable.
Several destinations have been proposed for the whales but there is disagreement about where they should go and what should happen to them.
Most experts agree that releasing the two whales, specifically Icelandic killer whales, into the wild would be inappropriate because both were born in captivity and would not have the skills needed to survive.
“It's kind of like taking your dog out of the house and sending him into the woods to live freely as a wolf,” Hahn-Strager says.
In 2023, the marine biologist published The Killer Whale Journals, which details her decades-long interest in ocean predators and how they behave.
“These whales, which have spent their entire lives in captivity, have their closest relationship with humans. They are the ones who provide them with food, care, activities and social relationships.
“Killer whales are very social animals, like us humans, and they depend on social bonds. They have established those bonds with their trainers… They depend on humans and that is the only thing they know.”
The deal to send Wiki and Keigo to a marine zoo in Japan, with the support of managers at Marineland, sparked outrage among activists who said they would receive worse treatment.
Last November, the French government blocked the deal, saying that Japan's animal welfare laws were lax compared to those in Europe, and that the 13,000-kilometre (8,000-mile) journey would cause stress on killer whales.
Another option is to send them to the Spanish Marine Zoo in the Canary Islands.
Loro Parque, in Tenerife, adheres to European animal welfare standards, but activists fear Wiki and Kygo will continue to perform there.
There have also been several orca deaths there in the past few years.
A 29-year-old man named Keto died in November, and three other orcas died there between March 2021 and September 2022.
Scientific examinations of those three killer whales by the University of Las Palmas de Gran Canaria show that the deaths were inevitable, Loro Parque says.
Katherine Wise, from the charity World Animal Protection (WAP), told the BBC: “It would be devastating for Wiki and Kijo to end up in another entertainment venue like Loro Parque – from one whale prison to another.”
Wap wants to rehome killer whales in an air-conditioned ocean bay.
“(We) and many others have urged the French government to do everything it can to facilitate the movement of killer whales to a sanctuary off the coast of Nova Scotia.”
“We will close the Gulf to them.”
The organization, which hopes to build the facility in eastern Canada, says it will be able to attract funding if it receives a commitment from the French government to send the two whales there.
The Whale Sanctuary Project (WSP) proposes to enclose a 40-hectare (98-acre) area of seawater with nets.
Wikie and Keijo can then use a large area of water, with human support from veterinarians and social care workers, for the rest of their lives.
The average lifespan of a male killer whale is about 30 years, according to the US National Oceanic and Atmospheric Agency. Females usually live about 50 years.
“Life in the shelter will be as close as possible to what they experienced when growing up in the ocean,” says WSP. “It will be a new life “It will make up for a lot of what happened before.”
This type of project has been done before.
Kiko, the killer whale who starred in the 1993 film Free Willy, was rescued from captivity in 1996 before being transferred to a bay in Iceland in 1998.
Unlike Wikie and Keijo, he was born in the wild and was able to relearn some necessary survival skills while living in the bay for four years.
He eventually left with a group of killer whales that he joined and swam to Norway where he is He died in 2003 after infection.
Strager warns that the proposed refuge may look as foreign to Wikie and Keijo as it does in the open ocean.
“We have this perception that animals have freedom in the same sense that we do, 'Now they're free and they're going to love it.'
“We don't know if they look at freedom the same way…Will they feel afraid because it's so different from what they're used to? I don't know.”
She told the BBC: “I don't think there are any good solutions for animals that have been in captivity all their lives.”
More than 4,000 animals will be relocated from Marineland, which was founded in 1970 by Count Roland de la Poype.
He was a decorated fighter pilot who fought during World War II before establishing himself in the plastics industry and opening Marineland due to his interest in marine life.
Closing down his passion project is the latest step in a campaign targeting marine zoos that has gained momentum over the past 15 years.
Actress Pamela Anderson called for the closure of Marineland in 2017 and held a protest outside its entrance, saying “captivity kills.”
In 2013, the documentary Blackfish detailed how to do just that An orca named Tilikum killed trainer Dawn Brancheau After the show at SeaWorld Orlando in 2010.
He grabbed her and dragged her into the water, where he tore off her arm and drowned her.
The film also shows how Tilikum was also involved in the killing of two other people.
Researchers interviewed in the film argued that killer whales captured from the wild and trained to perform become violent in captivity.
Numbers of visitors and SeaWorld's financial revenues suffered in the wake of the documentary In 2016, they suspended the captive breeding program.
They rejected calls to release the remaining killer whales into the wild, saying they would likely die if left to fend for themselves.
Eighteen months ago, they opened a new marine zoo in the United Arab Emirates, the first of its kind at Sea World outside the United States.
The new $1.2bn (£966m) facility in Abu Dhabi with state-owned entertainment developer Miral includes the world's largest aquarium.
There are no orcas on display here, but to the dismay of activists, dolphins are still present.
Wap helped convince Expedia not to sell more vacations that included captive dolphin shows, and wants other travel companies to follow suit.
“Blackfish was more than just a success — it was a phenomenon,” scientist Naomi Rose wrote in a report for Wap. “I am convinced that this has pushed Western society past the tipping point on the issue of captive cetaceans.”