It only took moments for Naruto, a male crested black macaque, to snap photos of his own face - amber-colored eyes looking straight at the camera, mouth open in a half-grin - using a camera belonging to wildlife photographer (and human) David J. Slater.
The legal battle over who owned the ensuing "monkey selfies," on the other hand, stretched for nearly two years.
In 2015, People for the Ethical Treatment of Animals filed a lawsuit on behalf of the primate, claiming that Naruto owned copyrights to the photos, which had gone viral.
On Monday, Slater and the animal rights group reached an agreement out of court. As part of the settlement, Slater will donate 25 percent of proceeds from sales or usage of the "monkey selfies" to charities in Indonesia that protect crested macaques, considered a critically endangered species.
"PETA and David Slater agree that this case raises important, cutting-edge issues about expanding legal rights for nonhuman animals, a goal that they both support, and they will continue their respective work to achieve this goal," the group said in a joint statement with Slater. "As we learn more about Naruto, his community of macaques, and all other animals, we must recognize appropriate fundamental legal rights for them as our fellow global occupants and members of their own nations who want only to live their lives and be with their families."
The agreement put an end to what PETA called a "groundbreaking lawsuit" that sought to "extend fundamental rights to animals for their own sake - not in relation to the ways in which they can be exploited by humans." Over the past two years, Slater has described the legal battle with PETA (and, separately, with Wikimedia Commons) in less glowing terms, comparing it to, well, a monkey on his back.
"Photographers are under enough pressure these days to make a living," Slater wrote last month in response to a Facebook user who sided with the photographer. "We [don't need] ridiculous interpretations of what copyright is from ignorant people like PETA and Wikipedia."
Slater did not immediately respond to an interview request Tuesday. On his Facebook page, the photographer struck a more conciliatory tone regarding his settlement with PETA, writing that their shared goal is "far more important than battles over copyright between me and a monkey I want to help."
Attorneys on both sides asked the 9th U.S. Circuit Court of Appeals to dismiss the case and throw out a lower-court decision that said animals cannot own copyrights, according to the Associated Press. Andrew J. Dhuey, an attorney for Slater, did not specify to the AP how much money the photographer has made from the selfies or if he would keep the remaining 75 percent of future revenue.
Slater has long argued that his 2011 trip to the Tangkoko-Batuangus Nature Reserve on the Indonesian island of Sulawesi was intended to bring attention to help the endangered primates - and that the photos would not have been possible had he not set up the equipment beforehand.
"Promotion and conservation of the crested black macaque, an extremely endangered relative of ours, was my original intention when I visited Sulawesi. I believed that comparing their personality to ours, through a photograph, could only gain our respect and our love for them," Slater wrote. "I see animals with personalities in exactly the same way as I see my fellow humans. The way in which I regard wildlife makes them so special to me and I want to share this. I hope that more and more people will try to see animals in the way I do. If so, then the conservation of our planet and the animals we have evolved with, will become a top priority within our society."
On his photography blog, Slater described how the selfies were made possible after he spent three days shadowing a troupe of macaques in the jungles of Sulawesi, trying to earn their trust. It was on the second day that a group of monkeys began grabbing his camera, prompting Slater to mount it on a tripod to see if they would play with it. They did for about 30 minutes, he wrote:
So I put my camera on a tripod with a very-wide-angle lens, settings configured such as predictive autofocus, motorwind, even a flashgun, to give me a chance of a facial close-up if they were to approach again for a play. I duly moved away and bingo, they moved in, fingering the toy, pressing the buttons and fingering the lens. I was then to witness one of the funniest things ever as they grinned, grimaced and bared teeth at themselves in the reflection of the large glassy lens. . . . They played with the camera until, of course, some images were inevitably taken! . . . It was like the joy of seeing your new baby learn about something new and becoming enlightened with a new toy. They loved the shutter noise, but most of all they loved their own faces, "chimping" away in what seemed to me to be total fun for them. . . ."
The ensuing "monkey selfies" quickly went viral and were featured in a 2014 photography book by Slater called "Wildlife Personalities." Notably, he specified in the book that "the shutter was pressed by the monkey."
PETA then sued for damages, alleging that Slater had infringed on Naruto's copyright in a complaint filed in U.S. District Court in Northern California.
"The U.S. copyright law does not make any distinction as to who can be the author of a copyrightable work based upon the species," Jeffrey Kerr, general counsel to PETA, told The Washington Post in 2015. "If a human being had made this or similar selfie photographs, they would be the copyright owner of the photograph. Naruto is no different since he is a macaque."
A federal judge disagreed, ruling last January that the copyright act did not apply to animals. However, PETA appealed, drawing the legal battle out even further - and leaving open the door for another year's worth of pun-filled headlines about the case. (None of this changed with news of the agreement: "Who Owns a Monkey Selfie? Settlement Should Leave Him Smiling," the New York Times declared. "Monkey see. Monkey sue. Monkey settle," the AP tweeted.)
In announcing the settlement, PETA cast it as a victory and said it would continue fighting for legal rights for animals.
"Naruto and the famous 'monkey selfie' photographs that he undeniably took clearly demonstrate that he and his fellow macaques - like so many other animals - are highly intelligent, thinking, sophisticated beings worthy of having legal ownership of their own intellectual property and holding other rights as members of the legal community," PETA said.
Slater, for his part, thanked people for donating and encouraging him throughout the legal process, and said he is eager to return to the field.
"This has seriously helped me to rekindle my passions, and once again go out and continue my work as a wildlife photographer and conservationist," Slater said.
He did not specify whether he would be encouraging animals to take their own photographs again.