GENEVA -- Dr. Luc Hoffmann, a Swiss ornithologist and naturalist with a passion for wetlands who helped create the World Wildlife Fund for Nature and many other conservation groups, has died, wildlife groups said. He was 93.

Hoffmann died Thursday at his home in the Camargue wetlands of southeastern France, which is known for flamingos and other birds, said the Tour de Valat, a research centre he founded there over a half-century ago. No cause of death was given.

A grandson of the founder of the pharmaceutical company Hoffmann-La Roche, now Roche, Lukas "Luc" Hoffmann developed an early fascination with nature. He earned a doctorate in zoology from the University of Basel, where he was born on Jan. 23, 1923. He wrote over 60 books and publications, mostly on birds and their habitats.

Hoffmann was credited with a key role in early moves to protect the Coto Donana wetlands in southern Andalucia, Spain, and was a "driving force" behind the Ramsar Convention, an international treaty on wetlands named for an Iranian city where it was adopted in 1971, WWF said.

Over the years, Hoffmann was honoured in a number of countries, mostly European. In 2003, a Luc Hoffmann chair in field ornithology was established at the Edward Grey Institute at Oxford University.

A private funeral was planned. Hoffmann is survived by four children, eight grandchildren and a great-grandchild, said the MAVA Foundation, another group that he helped found to fund global conservation.

MAVA and WWF set up the Luc Hoffmann Institute research centre four years ago.