THE Scottish zoologist who was studying Cecil the lion has told of his “outrage” at the killing of the animal - but said the global outcry it has triggered will help secure conservation efforts in the future.

Professor David Macdonald, founder and director of the Wildlife Conservation Research Unit (WildCRU) at Oxford University, said he was considering having to close an anti-poaching project in Zimbabwe only three weeks ago because of a lack of funding.

Now the hundreds of thousands of pounds donated to the research unit in the wake of the death of Zimbabwe’s most famous lion will be used to continue the anti-poaching patrols and expand other conservation work.

Last week it emerged Cecil, a 13-year-old lion who had been monitored since 2008, was lured outside protected national park land by the scent of bait before being shot with a bow and arrow, tracked for two days, then skinned and beheaded. The hunter was later named as American dentist Walter Palmer.

Macdonald also revealed that in one of his last encounters with Cecil - whose movements were tracked through a GPS collar - he witnessed the lion have a close brush with hunters.

In an interview with the Sunday Herald he said: “It was only a few months ago I was watching him in the field and rejoicing in his magnificence and beauty.

“My response to his killing is on two levels: one as a human being who has dedicated his life to wildlife and therefore feels strongly about these issues and the other is as a scientist.

“As a human of course I was shocked and saddened, and when I learned this particular death was allegedly due to illegal activity, I was not only saddened I was outraged.

“On the other hand, I look for good to come out of this - there is more data for us to understand about the pattern of interaction between people and wildlife, about the behaviour of the surviving lion and how can we better improve our advice regarding conservation in this context.”

Macdonald, who is originally from Glasgow, has been interested in wildlife from an early age when he would make casts of fox footprints he found in golf course bunkers - and has studied animals ranging and wood mice in the UK to meerkats in the Kalahari and proboscis monkeys in Borneo.

He founded the WildCRU 20 years ago with colleague Dr Andrew Loveridge, which runs various research projects including understanding the factors threatening lion conservation throughout Africa.

A few years ago it was thought there could be as few as 30,000 lions left in Africa – but Macdonald said more recent evidence has suggested that number has dropped even further. Cecil was one of a number of lions being studied through satellite tracking which enabled his movements to be followed from hour to hour through the vast and often impenetrable landscape of Zimbabwe’s Hwange National Park.

Macdonald said he and Loveridge had previously witnessed the lion having a close encounter with hunters after straying near the border of the national park.

“We could see there were hunters on the other side - on that occasion he turned back and we were relieved and rejoiced,” he said.

“But of course this time it didn’t work out. So I am saddened by that, but I seek to learn from it.”

Palmer has faced a storm of hate messages on social media in the wake of the incident, with police said to be investigating threats made against him.

The dentist, from Minneapolis, has claimed he believed the hunt was legal and that he had no idea the lion was a local favourite, or being studied by scientists. On Friday Zimbabwe’s environment minister said extradition is being sought for Palmer and the government wanted him tried in the country because he “violated our laws”.

A petition urging Palmer to be sent to Zimbabwe to face justice is also being reviewed by the White House after gathering more than 140,000 signatures.

Two Zimbabweans - a professional hunter and a farm owner – appeared in court last week in connection with the killing of the lion and were subsequently bailed.

When asked about what should happen to those involved in the killing of Cecil, Macdonald said they were likely to face a heavy penalty under the law, but added he had no idea about the complicity of Palmer.

He said: “He may have been complicit or he may have been unaware of what was going on.

“I could not comment – I take the view that people are innocent until proven guilty.

“But insofar as this was illegal it is reprehensible and should be punished accordingly.”

While the death of Cecil the lion – which Palmer reportedly paid around £32,000 to hunt – has triggered huge debate about trophy hunting, Macdonald said outlawing big game hunting was not necessarily the answer.

He said: “I think modern conservation is extremely complicated and needs to integrate various different factors and needs that involve people with wildlife.

“What I do deplore is bad management. So while I would take an impartial position on things that are legal and simply seek to inform about the facts and consequences of those activities, I take a very dismal view of bad management.”

But he said the “remarkable” response from millions of people around the globe would help in future conservation efforts run by WildCRU.

A host of celebrities have also voiced their anger at the killing, including comedian Ricky Gervais, TV show host Sharon Osbourne, actress Joanna Lumley and former Spice Girl Geri Halliwell.

Nearly £250,000 was raised in around 48 hours after American chat show host Jimmy Kimmel launched an appeal on Tuesday night and US billionaire philanthropists Tom Kaplan and Daphne Recanati Kaplan have pledged to match new donations to the unit up to the value of £64,000.

It is hoped that the Cecil appeal will eventually reach £500,000 with an appeal site launched at everydayhero.co.uk/event/Wildcru-Cecil-the-lion.

Macdonald said: “There has been a remarkable response around the world with millions of people showing their interest.

“Although they are inspired to do so by the case of Cecil - one lion that happens to have had this misfortune - I take their reaction as a sort of metaphor that there is a world population of people who deeply value wildlife and nature and the environment and this is being expressed at the moment through their concern for Cecil.”

Macdonald said the funds raised so far would guarantee the continuation of the anti-poaching force and potentially expand his team’s research work on lions to other countries including Botswana and Zambia.

“Only three weeks ago we were talking about having to close down our anti-poaching force which goes around very courageous work day-to-day cutting snares in the national park,” he said.

“For example – the male lion which is the co-member of the coalition with Cecil had been snared a couple of years ago. The anti-poaching force found him, rescued him, patched him up and he is fine now.”

A vital part of the unit’s work is also bringing Zimbabwean students to Oxford to study conservation, Macdonald said.

He added: “We already train some young Zimbabwean conservation biologists and have given scholarships to the WildCRU Centre in Oxford.

“If as a result of these donations we can train more young Zimbabweans to be custodians of their own wildlife, it will be a truly wonderful memorial to Cecil.”