Nicholas B. Davies Nick Davies has spent his life watching birds, and his deep knowledge of their natural history lies at the heart of his outstanding academic career in science. Through his detailed studies of British birds like Pied Wagtails Motacilla alba yarrellii, Dunnocks Prunella modularis and Common Cuckoos Cuculus canorus, Nick played a pivotal role in developing the field of behavioural ecology: the study of adaptation in relation to contemporary ecological and social conditions. In a scientific world enveloped by the fog of big data, his work shines out as a beacon of clarity, where theory is informed by observation and tested incisively through elegant experimentation. Nick’s analyses of adaptation in the natural world place him in a trio of 20th century pioneers who led discoveries in behaviour, ecology and evolution with their field experiments on birds. Nick Davies’ research in the 1970s, 80s and 90s is on a par with David Lack’s work in the 1950s and 60s and Niko Tinbergen’s work in the 1930s and 40s. In common with David Lack, Nick studied natural sciences as an undergraduate at Cambridge (1970–1973). Like Lack, he found the experience underwhelming for the lack of field biology. Even so, Nick (working together with his fellow student Rhys Green – who was also to be awarded the Godman-Salvin Prize) was able to formulate his own research project on the foraging behaviours of warblers at Wicken Fen and presented his work at the annual Edward Grey Institute (EGI) student conference in Oxford in 1973. This secured him a PhD place there, with Euan Dunn. Nick arrived in Oxford just as it became a crucible for research on adaptation. Tinbergen and Lack already had established research groups that studied behaviour from an evolutionary and ecological perspective but now Richard Dawkins was writing The Selfish Gene and John Krebs was championing an economic approach to analyses of decision-making. Nick led the way in test-driving and tinkering with this new thinking in the field, starting with his PhD research on territoriality in Pied Wagtails. A key discovery was that bird behaviour flexibly and adaptively tracks ecological conditions. Pied Wagtails varied the extent of their territory defence adaptively, as prey abundance varied over time. In 1979, Nick returned to the Department of Zoology at Cambridge and he has stayed there ever since (though he retired from his Professorship in 2019). Soon after his return, he remedied the deficiencies in field biology at Cambridge by encouraging original project work on the field course for final year students. He also developed an entirely new set of lectures in behavioural ecology with a unique method of delivery, winning over class after class by playing the part of a student himself and writing his lecture notes effortlessly on an overhead projector as he spoke, for students to transcribe. The clarity and logic of his lectures, and the ease and gentle humour with which he presented them, belied the immense thought and work that Nick put into preparing them. With Nick as a guide, complex ideas were suddenly appealing, difficult experiments seemed straightforward to execute, the tangle of the natural world was unravelled, and students left the lecture theatre feeling a lot cleverer than when they had arrived. From 1981, students worldwide were able to appreciate Nick’s teaching, when the lectures became chapters in his bestselling textbook with John Krebs, An Introduction to Behavioural Ecology (4th edition, 2012). Meanwhile, Nick had begun to study the Dunnocks living in the University’s Botanic Garden. Following in Lack’s footsteps, he marked individual Dunnocks with colour rings and followed them around their everyday lives. He added a modern twist by supplementing his detailed field observations with one of the first genetic analyses of parentage in a wild animal (working with Terry Burke and Mike Bruford). Nick showed that natural selection works in opposing directions on male and female social behaviour: behaviour that promotes female fitness is bad for male fitness, while behaviour that promotes male fitness is bad for female fitness. This, he deduced, is why Dunnocks have such a variable mating system. Selection acting on males favours a polygynous mating system over polyandry, whereas selection acting on females favours polyandry over polygyny. A roll of the ecological dice, through the underlying distribution of food, determines which sex will triumph. Sometimes males win, sometimes females win and sometimes there is a stalemate, with monogamy or polygynandry (multiple males and multiple females on the same territory). With Ben Hatchwell, Nick found that the conflict between the sexes also permeates other aspects of the social lives of Dunnocks. It influences how parents share the work of provisioning offspring and it persists after mating when sperm compete for fertilisations. It pays a female to mate with multiple males because she trades paternity for paternal care, but it pays a male to sire the whole brood and to let the female bear the greater burden of raising young. In a companion study in the Pyrenees on another Prunella species, the Alpine Accentor Prunella collaris, Nick and his group found that the same social rules apply in more rugged terrain. After a decade of Dunnocks, Nick returned to Wicken Fen to start another substantial piece of work - this time focusing on Common Cuckoos. Unable to mark individual Cuckoos for observation, he relied extensively on Tinbergen-style field experiments, in which he played the part of the Cuckoo himself. With Mike Brooke, he made model eggs and ‘laid’ them in hundreds of host nests to watch the owners’ reaction. From these experiments, he unpicked a co-evolutionary arms race of adaptation and counter-adaptation in the Cuckoo and its hosts, showing in detail how the hosts have evolved defences against exploitation by the Cuckoo and how these defences have in turn selected actions in the Cuckoo to circumvent them. He broadcast begging calls at the nest, and manipulated nestling gape colour, to show how the Cuckoo chick fools hosts into feeding it so assiduously, by tapping into the hosts’ usual rules for feeding young. And he discovered how social learning can rapidly activate and spread host defences against Cuckoos through the population. This work is known far beyond academia through Nick’s books (especially Cuckoo: Cheating by Nature), as well as radio and TV programmes. Two threads run through Nick’s lifetime in science. The first is his graciousness, and the personal warmth he exudes for everyone that has the good fortune to meet him. It shows not only in his personal interactions, and his kind and supportive mentoring, but through his empathy. He has spent his life putting himself in the shoes of others, whether it is the students he teaches, the audiences he lectures, the readers he writes for, or the birds he watches so carefully in the field. This is one reason why he is such a superb educator, communicator and naturalist. The second glittering thread is the string of awards and prizes that Nick has accrued in travelling from one scientific discovery to the next. He was elected a Fellow of the Royal Society when he was just 41. His books have received multiple prizes and critical acclaim. He has been awarded the Frink Medal, the ASAB Medal, the Elliot Coues Medal, ISBE Hamilton Prize, and the Croonian Medal and Lecture from the Royal Society. It is entirely fitting that Nick can now add to that list the Godman-Salvin Prize from the British Ornithologists’ Union, for his outstanding and distinguished contribution to ornithology.