Obituary: Vero Wynne-Edwards

Vero Wynne-Edwards was one of the 20th century's greatest scientific naturalists and original thinkers on population regulation in animals.

The book for which he will be remembered is Animal Dispersion in Relation to Social Behaviour (1962), which was probably the most controversial to appear in biology in the Sixties and Seventies. At 650 pages it was the scholarly result of a lifelong consideration of the processes limiting animal numbers.

In it he proposed that animals collaborate socially for the benefit of the group, that they compete for territory and status rather than for food, with the losers patiently accepting their lot, and that animals are not, as Darwin supposed, always striving to increase their numbers but are instead programmed to regulate them. The mechanisms that prevent animals overexploiting their resources include social displays, territorial behaviour and communal roosting which evolved by group selection.

In Wynne-Edwards's view group selection operates by differential survival of populations. Those populations which showed self-restraint in reproduction and exploitation of resources survived longer than more profligate groups, so that self-regulation of population size developed during the course of evolution. This ran counter to the conventional Darwinian view of natural selection which operates by differential survival of individuals.

These ideas were robustly rejected by adherents of the still widely accepted orthodoxy that natural populations are limited by shortage of food, by predators, parasites and disease or climatic perturbations. More importantly, group selection was considered to be an implausible evolutionary process; conventional selection at the individual level would always override any self-restraint by members of a population.

Nevertheless, the principal contribution of Animal Dispersion was to focus attention on the mechanisms of social evolution, and ideas that have been developed over the past two decades in opposition to Wynne-Edwards's hypothesis remain prominent in much of the current thinking in sociobiology and behavioural ecology. His book was widely read by biologists of all disciplines and a precis of it, published in the Scientific American in 1974, sold 350,000 copies. His contribution to science was recognised in 1970 by election to the Royal Society.

Wynne-Edwards attempted to answer his critics in his second major work, Evolution through Group Selection, published in 1986. He reviewed new evidence which supported his ideas and considered that he had overcome many of the objections to this thesis. The book was sympathetically reviewed at length in Nature and Science although Nature's reviewer concluded, from Wynne-Edwards's failure to confront the tenets of the newly emerging discipline of behavioural ecology, that it was a work of advocacy. And this was the main problem - from a hypothesis, group selection had for Wynne-Edwards become an article of faith.

One colleague, dismissing these ideas to a class in Aberdeen, was chastened when at the end of the lecture, delivered in a room adjacent to an office occupied in retirement by Wynne - as he was known to his academic friends - the door opened and Wynne invited him to listen to a long and very courteous discourse on the error of his ways.

Wynne-Edwards continued to take every opportunity to convince sceptics that group selection works, despite the unfortunate reluctance of some editors to give him journal space. Scientific American commissioned a second article, then declined to publish it, but Wynne-Edwards continued writing, and his last paper on group selection appeared in the Journal of Theoretical Biology in 1993, when he was 87.

Vero Wynne-Edwards's boyhood was spent in the Yorkshire Dales where he was fascinated not only by the local fauna and flora but also by astronomy. He was a meticulous observer of animals and plants who recorded his daily observations in countless notebooks. After Rugby School, he read Zoology at New College, Oxford, in 1924-27, where he acquired a detailed knowledge of the animal kingdom from E.S. Goodrich and was greatly influenced by Julian Huxley and Charles Elton. He gained first class honours and his first appointment was as a "student probationer" at the Marine Biological Association in Plymouth, where he studied fish and crustacea but also published his "extra-curricular" observations on the movement and dispersion of wintering flocks of starlings.

These observations were made during visits from Plymouth to his fiancee Jeannie Morris, one of his Oxford classmates, who lived in Exeter. He would rise early to catch the first bus or train and station himself at a vantage-point before sunrise to watch the first waves of starlings dispersing from roosts. By backtracking he was able to locate all the roosts in Devon. This early enthusiasm for population studies of birds was to last throughout his life.

He was then briefly on the staff of the Zoology Department at Bristol University where, in 1929, he married Jeannie, who was to become his lifelong companion and support. He soon moved to be an Assistant Professor at McGill University, Montreal. On the voyage across the Atlantic, he recorded the locations of seabirds and, as a result of this and several further transatlantic journeys by ship, he concluded that seabirds in the north Atlantic were not randomly distributed but occurred in three zones - coastal, offshore (to the edge of the continental shelf) and pelagic (deep-water). These observations were published by the Boston Society of Natural History in 1935 and resulted in the award of the Walker Prize.

In Canada, Wynne-Edwards developed his interests in montane flora and his explanation of the postglacial distribution of plants in North America earned him fellowship of the Royal Society of Canada and a second Walker Prize.

During the Second World War, he enlisted for training in the Royal Canadian Naval Reserve but retained his post at McGill. He taught electronics to radar mechanics in the Royal Canadian Air Force and was also sent by the Fisheries Research Board of Canada to report on the fisheries resources of the Mackenzie and Yukon rivers and to establish the potential for settlement of men discharged from the armed forces after the war. This was a great adventure and rekindled his interest in fish populations.

Although they shared the responsibilities of head of department, McGill was unable to decide between Wynne-Edwards and N.J. Berrill for the Strathcona Chair of Zoology, and Wynne-Edwards decided to break the deadlock by returning to England. His children were quick to complain that this would deprive them of skiing opportunities and so he was attracted to vacancies further north.

In 1945, he was offered the Regius Chair of Natural History at Aberdeen University, a post which he occupied from 1946 until his retirement in 1974. In 1956 he initiated an important research project on the population ecology and behaviour of red grouse which is still active 40 years later. He established Culterty Field Station as a centre for postgraduate training and research in ecology, and was instrumental in rehousing (and renaming) his Department of Natural History in a new building which was opened in 1970 and which remains one of half a dozen departments of Zoology in the UK. Wynne-Edwards also served as Vice-Principal of Aberdeen University from 1972 to 1974.

A major reshuffle of government-sponsored science took place in 1964-65. A new Natural Environment Research Council (NERC) was formed, incorporating the Nature Conservancy (NC) and the Advisory Committee on Fisheries Research. After serving on the councils of the NC and NERC, Wynne-Edwards took over as NERC Chairman 1968-71. His term of office coincided with a golden age of expansion and prosperity and most of the projects the NERC Council set in motion came to fruition.

However, the NC and the NERC did not integrate well and Wynne-Edwards was unable to reconcile the opposing factions. This led in 1973 to further reorganisation, with the establishment of the Nature Conservancy Council as a quango. The research division of the former Nature Conservancy was retained within the NERC as the Institute of Terrestrial Ecology. There was much ill-feeling about this division and many of his friends saw Wynne-Edwards as one of the casualties. On retirement, he was appointed CBE for his long and distinguished service to the government and science in the UK. Many felt he deserved a greater accolade.

In Aberdeen, Wynne- Edwards was a legend as a cross-country skier and hill walker. At 62, he held the record for the six tops of the Cairngorms, which he covered in 9 hours 34 minutes, a distance of 28 miles with 9,000 feet of climbing. At 76, a few weeks after major abdominal surgery, he was seen on his skis in Glen Tanar, and at 80 he skied down Morven with his daughter.

In the days before university research and teaching required assessment, in order to produce league tables of quality, Vero Wynne-Edwards was the epitome of scholarship and erudition. His authority, delivered with a firm yet gentle touch, commanded respect and inspired affection. As a scientist he lacked personal experience of hypothesis- testing research, but came to appreciate its importance in shedding light on group selection. Although his ideas did not gain wide acceptance, he was widely admired both as natural historian and biological thinker.

He is survived by his wife, with whom he spent his last months in a retirement home looking out over the Dee Valley with his binoculars at the ready.

Vero Copner Wynne-Edwards, natural scientist: born 4 July 1906; Assistant Lecturer in Zoology, Bristol University 1929-30; Assistant Professor of Zoology, McGill University 1930-44, Associate Professor 1944-46; Regius Professor of Natural History, Aberdeen University 1946-74, Vice-Principal 1972-74; FRS 1970; CBE 1973; married 1929 Jeannie Morris (one son, one daughter) died Banchory, Kincardineshire 5 January 1997.