Ceredigion council said Borth Wild Animal Kingdom was told in April and May 2017, months before one escaped.

Lillith was shot after weeks on the run and a source close to the zoo said she died due to the owners' incompetence.

But Tracy Tweedy, the zoo's co-owner, said they had cut the foliage back as far as they thought they needed to.

Despite the warnings from a vet and a zoo licensing officer, the trees were not cut back until after she jumped over the perimeter fence.

The source, who has decades of experience with zoos and wildlife parks, said: "They lost their own lynx - I knew it was able to climb a tree and that tree put it over a height of the night den, it could jump out of the enclosure.

"I told them the trees inside the enclosure were too high."

The council said a vet warned the zoo operators in April last year of the situation, and a licensing officer had advised them at first in person and then in an email on 4 May 2017.

Mrs Tweedy said that when she and her husband took over the zoo on 17 May 2017, they "cut back vegetation in all of the enclosures, including the lynx enclosure, and this was done on several occasions over the summer".

She said: "Obviously, in hindsight, we did not cut it back enough but I would like to point out that these were conditions from the previous owner."

Mrs Tweedy added that the council's request to cut back vegetation had been "a general statement about the entire zoo".

"We have worked continuously since taking over last year to improve enclosures and public and staff safety and we will continue to do this," she said.

The source also revealed that a second lynx Nilly died in what owners called a handling error three days before Lilleth was shot - something the zoo did not make public for six days.

In a Facebook post on 13 November, the zoo simply said Nilly died "last week".

The council said it understood Nilly died on 7 November and this was disclosed to officers the following day.

Mrs Tweedy said it was not announced sooner as "the focus had to be kept on finding and returning Lillith".

The Eurasian lynx escaped between 24 and 29 October 2017 before being found at a nearby caravan site where she was shot dead by a marksman on 10 November.

The council blamed the zoo for its failure to catch Lilleth before she posed a risk to the public and Dyfed-Powys Police is investigating threats made against the marksman.

The zoo's ban on keeping certain dangerous "category one" animals was lifted in July on the understanding it employed "a suitably qualified, competent and experienced person to manage the collection within six months".

Nilly's post-mortem examination by an independent vet was inconclusive, so no action could be taken as there was "no prospect of a successful prosecution".

The source said: "100% Lilleth would be alive if it had been properly managed.

"I wish them every success if they run a petting zoo for the rest of their lives, but these people have lions."