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, died July 21 in southeastern France. He was 93.

He died at his home in the Camargue wetlands, which is known for flamingos and other birds, said the Tour de Valat, a research center 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, Hans Lukas Hoffmann developed an early fascination with nature. He earned a doctorate in zoology from the University of Basel, the Swiss city where he was born on Jan. 23, 1923. He wrote over 60 books and publications, mostly on birds and their habitats.

Dr. 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, Dr. Hoffmann was honored 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.

His wife, Daria Hoffmann-Razumovsky, died in 2002. Survivors include 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 center four years ago.