The Rwandan and national parks office said the giant old silverback "succumbed to old age" on Monday after falling ill in the past week.

"He has been sick. He's been weakening. It's in the last week that he started going down," Rosette Rugamba, head of the tourism and national parks office told AFP.

The service said the 200 kilo (450 pound) primate "has been observed closely by researchers his entire life," most notably by legendary American primatologist Dian Fossey who studied his group in the 1970s.

The link with Fossey may yet continue as Titus could be buried in the gorilla graveyard next to the remains of Fossey's mountain cabin in Karisoke, Rwanda.

Rugamba said burial arrangements have yet to be made.

Of more immediate concern is the removal of Titus' body from among his adoring group in the Virunga mountains in northwestern Rwanda.

"The other gorillas are mourning. They are cleaning him," she said.

"You have to be very careful. You can't just remove the body."

"Our vet's on standby to do an autopsy," Rugamba said, adding that this is standard procedure for any mountain gorilla in Rwanda.

Fossey is credited with bringing the primates' plight to the world's attention and most likely saving the species from extinction.

Until she was hacked to death in 1985, Fossey lived on the bamboo-clad slopes of Rwanda's Virunga mountains, surrounded by her beloved gorillas.

Her isolated life in the mountains of Rwanda, where she would single-handedly fight poachers and cattle herders who encroached on the gorillas' habitat, was immortalised in the 1988 Hollywood movie "Gorillas in the Mist" which featured Titus.

In 2008, the BBC filmed a documentary entitled "Titus: the Gorilla King".

"Titus was really one of our success stories -- every year we celebrate that this species is still alive," Rugamba said, adding: "Gorillas are more valued, more protected and the numbers are growing".

Mountain gorillas are found only on the slopes of the Virungas on the borders of Rwanda, Uganda and the Democratic Republic of Congo and fewer than 700 individuals are left, according to the Dian Fossey Gorilla Fund International.

Once hunted for their meat, and for the ornamental value of their skulls and paws, the gorillas have more recently had to survive warfare encroaching on their habitat in one of Africa's most conflict-scarred regions.

Gorillas have also been vulnerable to traps set for other animals, and females have been killed so that their babies could be sold as pets.

Both Rwanda and Uganda have turned gorilla tracking into a major eco-tourism industry and a big foreign-currency earner, while war had stalled similar development in eastern DR Congo.

In Rwanda gorilla tourism has not only helped the economy but has done much to boost the image of a country that used to make the headlines only for its 1994 genocide.

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