A zoo in Wales that made headlines in 2017 after a lynx escaped suffered the deaths of one in five of its animals in 2018, it has emerged.

Borth Wild Animal Kingdom near Aberystwyth reopened last month in defiance of the council after closing temporarily in January following a “directive” to shut its most dangerous animal enclosures since it does not have enough firearms-qualified staff in the event of an animal escaping.

The zoo, which reopened all areas apart from the lion enclosure, is to appeal against the decision in court.

Reports of animals dying in mysterious circumstances began to surface after an escaped lynx was shot dead by a marksman in late 2017 and another was killed accidentally by strangulation while being moved shortly after. A number of other deaths followed.

Some of the reports originated when a volunteer animal handler, Mark Anthony, accused the zoo, which had recently been taken under new ownership by Tracy and Dean Tweedy, of neglect and keeping the animals in inadequate conditions.

The Tweedys have denied the claims. They blame old age and natural causes for a number of the deaths, including those of a wallaby, a squirrel monkey, and a 22-year-old leopard, and said a Burmese python died of a chronic respiratory problem. And they hold Anthony responsible for the deaths of an emu and three bald-headed ibises.

The true scale of the deaths is now clear. A stock list disclosed by Ceredigion council in response to a freedom of information request reveals that 57 of its 305 animals died in 2018.

Five of seven marginated tortoises, seven of 30 primates, five of six parakeets, two of nine meerkats, one of two caiman crocodiles, all seven African giant snails and all five stick insects were among the animals that died, while many others left the zoo.

Anthony is also accused by the Tweedys of selling a number of primates and a crocodile, which reportedly ended up in the pet trade. He has denied all of the claims against him, describing them to the Times as “absolute nonsense” and “downright lies”.

Postmortem documents from Wales Veterinary Science Centre seen by the Guardian note the ibises’ “sudden” deaths. One had ingested a 7cm piece of wood, and another had swallowed a large piece of glass and a long cable tie.

However, postmortems did not take place for every death. It has been claimed that poor shelter and bad management led to the deaths of two monkeys, while Anthony said rats had infested the aviary and animals were kept in cold conditions after the heating broke.

Anthony, who could not be reached for comment, previously worked for a company named Extreme Animal Encounters, which said it supplied animals to the TV and advertising industry. “Virtually any animal you want is available for your production,” an advertisement read. “If we don’t have what you need, we will find it for you at no extra cost.”

The company was dissolved in August 2014, Companies House records show. Its directors, David Crumpton and Samantha Pardoe, have recently begun working at Borth zoo.

Ceredigion council said it was unable to comment on events at the zoo, since it is subject to legal proceedings.

“Borth Wild Animal Kingdom have appealed [against] a direction to close the enclosures of its most dangerous animals – known as category one animals,” a spokesperson said. “The direction was served on 27 January 2020 because of inadequate firearms arrangements in the event of an escape of a dangerous animal.”

In contravention of the council directive, the zoo reopened in February without a full team of firearms-trained staff on site. It has appealed against the order while complaining of delays in the licensing process and a date for the hearing is expected to be named on 26 March.

“Unfortunately, we have not been able to fully resolve the firearms situation due to the powers that be and are opening against the advice of the licensing authority,” a notice on its website said.

“However, we have put in place extra security measures to ensure everyone’s safety and we will be keeping the lion area closed.”

The owners said the zoo needed to remain open to pay for the upkeep of the animals and has reassured visitors that they take their safety extremely seriously.

Council officials said they were already working on contingency plans to rehome category one animals – including a python, two lions, three lynx and five monkeys – if the appeal is unsuccessful.

Campaigners have called for the zoo’s closure. “Borth Wild Animal Kingdom has repeatedly failed to meet its legal requirements to keep wild animals and has put the welfare of animals and the public at risk,” said a spokesperson for the charity Freedom for Animals, which made the FoI request. “It’s time Borth zoo was shut down permanently.”