PROFESSOR Vero Copner Wynne-Edwards, former Regius Professor of Natural History and vice-principal, University of Aberdeen; born July 4, 1906; died Banchory, January 5, 1997

PROFESSOR Vero Copner Wynne-Edwards was an internationally renowned ecologist and naturalist whose major work caused considerable controversy and led to research, debate, and discussion which continues today.

His book on animal behaviour, Animal Dispersion in Relation to Social Behaviour, in 1962 will always be remembered by biologists. It was the result of a lifetime of observation and thought and dealt with the processes limiting animal numbers.

He suggested that species imposed their own checks through social processes to prevent their numbers increasing beyond sustainable levels. The ideas, which he stood by throughout his life, are unacceptable in modern biology but they stimulated fruitful debate and investigation.

He was invited to publish a precis of his book in Scientific American, the reprint of which sold more than any other.

On his retirement in 1974, in typical fashion, he warned that material lust could destroy the earth as a habitat for man. Addressing his final graduation ceremony he said the lust could leave our descendants with ``nothing but a sucked orange, a wretched plundered world in which the flower of human life could only wither and fall''.

There had to be a political and social bridge built across the gulf between a generation devoted to the unlimited growth of material wealth and a new generation content with an economy in balance with the carrying capacity of crops and other vegetation on which human survival and all other life depended.

He continued to refine his ideas in retirement and in 1986 published a second major work, Evolution through group selection, in which he attempted to meet the objections of his critics to his 1962 work. He continued to write until his eyesight failed and his last paper was published in 1993.

Professor Wynne-Edwards was educated at Leeds Grammar and Rugby School and graduated from Oxford with first class honours in zoology in 1927. He was Senior Scholar at New College, Oxford, in 1927-29. He lectured at Bristol and Leeds Universities before becoming assistant professor of zoology at McGill University, Montreal, in 1930 and remained there until 1946 when he took up his Aberdeen post.

He was involved in numerous important research expeditions including the MacMillan Baffin Island expedition in 1937, the Canadian Fisheries Research Board expeditions to Mackenzie River and Yukon Territory in 1944 and 1945, and to Arctic Canada in 1950 and 1953.

He was visiting Professor of Conservation at the University of Louisville, Kentucky, in 1959. He served on a number of national committees including the Red Deer Commission, the Royal Commission on Environmental Pollution, the Nature Conservancy and the Natural Environment Research Council of which he was chairman from 1968-71. He was elected a Fellow of the Royal Society in 1970 and became a Commander of the Order of the British Empire in 1973.

He was awarded the Godman Salvin Medal of the British Trust for Ornithology in 1977, the Neill Prize of the Royal Society of Edinburgh in 1977, and the Frink Medal of the Zoological Society of London in 1980.

Professor Paul Racey, a colleague and friend, said he was much loved and held in great affection by everyone who knew him. He had continued to visit the university regularly after his retirement.

He said he was a keen hill walker who always outpaced his younger colleagues and in 1968, when he was 62, walked the six tops of the Cairngorms - 28 miles - in what was one of the fastest times recorded for the challenge. He continued hill walking and skiing late into life.

He lived in Torphins until moving into a Banchory retirement home where he died.

He is survived by his wife, Jeannie, and a son and daughter, Hugh and Janet, who both live in Canada. His grand-daughter, Catherine Wynne-Edwards, is distinguished in the same field.