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With her toothy grin, large brown eyes and friendly expression, this preening female looks like a natural in front of the camera.

But it seems when the crested black macaque pushed the camera button and took her first ever 'selfie' - a picture which would be viewed millions of times on the internet - she may have been frightened.

'That's not a big smile,' explains award-winning British wildlife photographer David Slater, whose camera the macaque commandeered to take her picture.

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This is me in the jungle: The iconic 'selfie' of a female crested black macaque that captured the hearts of millions on the internet

'She saw herself reflected in the lens and primates will bare their teeth when they see another they don't recognise. It may signal that she is not sure what to do.'

The picture was snapped when David, 49, followed a troupe of around 20 primates on an expedition in the tropical forests on the Indonesian island of Sulawesi.

Yet it has now become the subject of a copyright battle, with David threatening to sue a website that is allowing the image to be downloaded for free - claiming it is the work of the animal, not David.

The pictures were taken in 2011, when David, of Coleford, Glos, rose at dawn every day to try to capture the macaques in their local habitat. But it turned out to be a case of monkey-see, monkey-do, as the primates, which boast blazing reddish eyes and spiky hairdos, quickly took David's job in hand.

At the end of the second day, the animals got so used to David, they started coming up to groom him, looking for insects in his hair and inspecting his camera equipment.

Monkey gets snap happy with photographers camera

Why the long face? Ol' Red Eyeys looks a bit down in the mouth for this perfect selfie

David, a professional photographer for 15 years, says: 'These lovely little primates are particularly inquisitive and intelligent. They have amazing human-like characteristics, not least their "punk" hairdo.

'But their red eyes seal the fascination for me; they are unnerving when they make eye contact with you - a memory of our ancient ancestors, perhaps? They started to get quite mischievous, jumping all over my equipment. At first the click of the lens scared them away, but they soon came back.'

He anchored the camera to a tripod, and then lay on the floor holding it so they could not carry it off. For the next half an hour, he let the monkeys fool around in front of the lens for the ultimate selfie session.

David says: 'Sure enough they put their muzzles right into the lens and starting snapping away. It was like the joy of seeing your new baby learn to play with a toy. It was totally fun.'

Me and my primates: David Slater and a macaque hold hands for the photographer's own selfie

Time for lunch: I know these taste good - but how do you open it?

Just as in the human world, most of the selfie-snappers were females - macaques live in groups of up to 70, dominated by one alpha male. The photoshoot came to an abrupt end when the male head of the clan decided his harem had monkeyed around enough.

'He jumped on my back and gave me a thump,' says David. 'They have formidably large teeth, so I decided to call it a day.' When he looked through the pictures, David knew he had something special.

Although hundreds of shots were blurry, four were perfectly in focus. When he released the pictures, he immediately got emails from people saying they had made them smile.

But it was only when the selfie craze took off about 18 months ago, that the pictures became iconic as a reflection of how similar to us animals can be.

David says: 'They show these monkeys are as fascinated with their own image as we are.'

The main threat to the crested black macaque is man hunting them for food. They are trapped as black-market meat and served as a delicacy.

Girls just wanna have fun: The inquisitive monkeys father around the camera is Sulawesi

There are estimated to be between 4,000 and 6,000 on the island, a fall of 80 per cent in the past 40 years. Indeed, while David was in Sulawesi he came across a close relative of the crested black macaque, a booted macaque, whose hand was stuck in a painful trap.

'I got him released for a 20p packet of cigarettes as a bribe. The value of these monkeys' lives is astonishingly low,' he says.

As well as the one-in-a-million shot that every wildlife photographer dreams of, David had hoped the pictures would raise awareness of the how endangered these animals are.

But the selfies have instead turned into an international legal row. Because monkeys can't own copyright, the images have been added to Wikimedia's bank of free images - and both the macaques and David miss out on royalties of tens of thousands of pounds.

David says the income should come to him - with a share going to a conservation project to protect these endangered creatures. But despite the fight, David says: 'One day I would like to meet them again. I owe them all a big thank-you.'