Before the king's burial, his subjects take their time to clean his body. It is not a ritual that should be lightly interfered with and, as the mighty, fallen monarch receives his last ablutions, medical personnel are keeping a respectuful distance.

Titus, the most studied and photographed ape in history, has died. The king was a gorilla called Titus. Although he had been deposed by his son, death seemed to have restored him to his full glory. The mighty silverback was once the dominant head of a tight-knit group of the great apes whose kingdom was on the eastern slopes of the Karisoke volcano in Rwanda's border lands, and a vital figure in the battle for the survival of the species.

Rosette Rugamba, head of tourism and national parks in Rwanda, said: "The other gorillas are mourning. They are cleaning him. You have to be very careful. You can't just remove the body." The Rwanda national parks office said the 450lb, 35-year-old gorilla had "succumbed to old age" after a short illness.

Officials described Titus as "possibly the most remarkable gorilla ever known", referring to his rise to dominance of he largest known group of gorillas in the world. And Sir David Attenborough, the naturalist who was famously upstaged by Titus when the gorilla, then five, climbed on his back, said yesterday he was sorry that Titus should have died when he was still "quite young". He added: "He was a charming little animal. Certainly, it was very memorable and I haven't been allowed to forget it."

Titus was given his name by the great US primatologist Dian Fossey, who came to the Mikeno chain of volcanoes in the 1970s to conduct a census of the great apes. Over two decades in Rwanda, she followed a group led by a male she had named "Uncle Bert" after a relative.

In 1974, Fossey gave one of Bert's offspring the name Titus. Ten years later, the primatologist was murdered in her Rwandan forest cabin, possibly because of her work opposing the exploitation of the animals. She was buried at the Karisoke Research Centre, which continues that work. In the years that followed, Titus became possibly the most photographed and studied gorilla in history.

One of the first of a generation of apes whose lives have been documented since birth, Titus left a remarkable record that has demonstrated how far human understanding of gorilla behaviour has come. For Titus, fame first came with his part in the film of Fossey's life, Gorillas In The Mist and latterly as the star of a recent BBC documentary The Gorilla King.

Ian Redmond, who worked with Fossey and later with Sigourney Weaver, who played her in the biopic, remembered his first meeting with the ape when the latter film was released last year. "Titus is special to me because he's the first gorilla I ever saw," he said. "As I approached this thicket, I saw this little black furry thing which was Titus being chased by his brother. It was like joining a family picnic - it was quite something."

But upon his birth in August 1974, Titus was initially thought to be an underdeveloped baby with little chance of survival. The environment gave little cause for optimism. His uncle Digit, who was Fossey's favourite gorilla, was killed by poachers in 1978, and had his head and hands hacked off; gorillas were often killed for the ornamental value of their skulls and paws. The same poachers later returned to slaughter Titus's father, Bert.

When Titus met Attenborough at the age of five, he was effectively an orphan, in a group with no silverback. But he survived. And later, after several years in a bachelor group, Titus eventually ascended to lead a mixed clan known as the "Beetsme" group, named for a former dominant male. Titus is thought to have sired more offspring than any other gorilla recorded. He is also credited with guiding his group out of danger during Rwanda's 1994 genocide.

Titus led the Beetsme group for 15 years. But gradually - in a tale of filial ambition worthy of Shakespeare - Titus's son, Kuryama, usurped him at the head of the group. Eventually, Titus accepted it. But he retained his affectionate entourage, who continued to groom him as he followed in his son's wake.

Now tourists will pay $500 for permits to see the great apes, Kuryama's leadership is likely to be a different era. Before Fossey's arrival in the region, though, the gorilla population was on course for extinction. The population of mountain gorillas is still endangered, but it has now stabilised. It was the battle for their survival that may have eventually claimed Fossey's life. Her murder has never been solved, but nor has her legacy expired, a legacy Titus lived out, and which, it is hoped, his tenuously surviving clan will continue to prove.

Although officials have refused to confirm the arrangements, it is likely that Titus will also be laid to rest at Karisoke, near the woman who gave him his name.