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Ellen Paul

Colin Pennycuick 1933 - 2019

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Colin Pennycuick, who has died aged 86, was the pre-eminent researcher in animal flight over the last century. He focused on the flight of bats and birds (and their possible ancestors), and asked the question: how do they work? To answer this deceptively simple question he brought to bear a mix of sharp logic and original and practical invention.

Though he sought to ground his work in the rigorous application of physics and mathematics, he was not satisfied with abstract results and conclusions by themselves, but always sought to democratise his findings, first to the biological sciences community and then to the huge population of lay people fascinated with birds and their flight escapades.

Pennycuick was an expert glider pilot, and gained some notoriety by piloting his craft in and around flocks of vultures, storks and eagles in Africa, and condors in Peru.

The son of Brig James Pennycuick and his wife, Marjorie, Pennycuick was born in Windsor, Berkshire. His family followed his father’s army postings, which in 1938 took them to Singapore, which they left in 1941 shortly before the Japanese invasion. Pennycuick was later sent as a boarder to Wellington college, Berkshire, studied zoology as an undergraduate at Merton College, Oxford, and worked on his PhD at Peterhouse, Cambridge. There he studied muscle mitochondria, whose task of converting oxygen and nutrients into energy he viewed as the basic engine of flight.

During two years’ national service with the RAF, he flew Provosts and Vampires, early jet-powered aircraft. He subsequently worked as a postdoctoral fellow at the Animal Behaviour Laboratory in Madingley, Cambridge, and in 1964 began a long association with the zoology department at Bristol University as a lecturer.

Colin Pennycuick at work in Iceland in 1995. His career took him as far afield as Nairobi, Peru and the South Georgia Islands. Photograph: Sverrir Thorstensen

He used the first computer at the university to design a tiltable wind tunnel, which he built from scratch and hung in a stairwell. He developed and adapted aeronautical ideas from helicopter theory to bird flight and tested their application based on meticulous observations of the free-flying pigeons which he kept in a loft on the roof of the building.

In 1968 he traveled to Nairobi, which he made his base for three years, installing his tilting wind tunnel between two acacia trees to study bat flight in the same manner as he had previously done with pigeons. He then spent another two years in the Serengeti national park as deputy director of the research station there. He learned how to fly his powered glider alongside pelicans, storks and vultures, documenting for the first time their extraordinary and essential abilities to travel economically over large distances by exploiting thermals.

From here on, his career was not so much a list of academic positions and research topics as a restless migration (frequently aerial, frequently self-piloted) of his own. He flew back to Bristol in 1973 via Addis Ababa, Cairo and Crete, in and around the Shetlands, France and Sweden, and down to Bird Island in South Georgia, Antarctica. There he first used his “ornithodolite”, an instrument he designed for measuring birds’ flight paths and speed, to track in detail the soaring flight of albatrosses. He found that the standard explanation – that they could power their flight by following a specific trajectory through a wind shear profile – was only partly responsible for their ability to fly continuously, without flapping for very long times, and that instead they used the wind in several different ways.

In 1983, he left for Miami University, which became a handy launch point for expeditions to the Everglades, Tennessee, Pennsylvania and Idaho, and further afield in Puerto Rico, the Bahamas and Peru. In 1992 he left Miami, via Greenland, Iceland and Sweden. He began a continuing association with the animal ecology group at Lund University in Sweden, tracking migratory birds by radar, and in 1994 the bird flight wind tunnel was inaugurated there by the king of Sweden.

In the late 1990s he collaborated with the Wildlife and Wetlands Trust at Slimbridge, in Gloucestershire, in tracking whooper swans, which as the largest flapping bird can provide a stringent test of aerodynamic theory at relatively large extremes of scale. He appeared in the 2003 BBC radio series Swan Migration Live, which tracked six Bewick’s swans and a whooper swan from Arctic Russia to the UK, with updates on their progress on the Today programme each morning.

In 2008 Pennycuick took part in an even bigger and more ambitious Radio 4 project, World on the Move: Great Animal Migrations, which tracked brent and white-fronted geese from the UK to Canada. With the aid of very accurate meteorological data, combined with measurements of wing beat frequency and wing shape, he modelled a gauge that could estimate the fuel consumed while these geese were migrating: this would give audiences, and the scientific community, some idea of the effort involved.

Pennycuick’s primary goal was to provide and test a physically reasonable theory of vertebrate flight, which could then be used to predict and understand how and why birds and bats do what they do. Many of his inventions, in techniques, procedures and instrumentation, were absolutely novel because he thought his own thoughts and proceeded by himself, according to the rigorous rules of logic and scientific inquiry.

A rich and exuberant publication history burst from his activities, starting with the first practical flight theory papers in 1968 and going on to include the books Animal Flight (1972), Bird Flight Performance (1989) and Modelling the Flying Bird (2008). In later years he increasingly focused his efforts on his flight software package, which grew from a small custom Basic program to a rather versatile application with graphical interface. As well as biologists, engineers wanting to know how birds manage to achieve the things they do with apparent economy of effort and energy expenditure used the program, and both groups learned from it, which gave Pennycuick particular pleasure.

He was appointed research professor in zoology at the University of Bristol in 1993, and senior research fellow in 1997. He was elected fellow of the Royal Society in 1990, and was made honorary companion of the Royal Aeronautical Society in 1994. In 1996 he was awarded an honorary doctorate by Lund University.

In 1992 he married Sandy Winterson. She and his son, Adam, survive him.

Colin Pennycuick at work in Iceland in 1995. His career took him as far afield as Nairobi, Peru and the South Georgia Islands. Photograph: Sverrir Thorstensen

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