Flight is a unique adaptation at the core of many behaviours in most bird species, whether it be foraging, migration or breeding. Birds have developed a wide diversity of flight modes (e.g. flapping, gliding, soaring, hovering) which involves very specialized behaviours. A key issue when studying flight behaviours is to understand how they develop through all the ontogenetic stages of birds, from the embryo to the flying adult. This question typically involves classical debates on animal behaviour about the importance of maturation and experience. Here, we review the literature available on the development of flight behaviours in birds. First, we focus on the early period when young birds are not yet capable of flight. We discuss examples and show how endogenous processes (e.g. wing flapping in the nest, flight development timing) and environmental factors (e.g. maternal stress, nutritional stress) can influence the development of flight behaviours. Then, we review several examples showing the different processes involved in the development of flight in flight-capable juveniles (e.g. practice, trial and error learning, social learning). Despite the lack of experimental studies investigating this specific question at different developmental stages, we show that several patterns can be identified, and we anticipate that the development of new tracking techniques will allow us to study this question more thoroughly in more bird species.

Flight is an efficient way of transport over a unit of distance, but it can be very costly over each unit of time, and reducing flight energy expenditures is a major selective pressure in birds. The common swift (Apus apus) is one of the most aerial bird species, performing most behaviours in flight: foraging, sleeping, and also drinking by regularly descending to various waterbodies and skimming over the surface. An energy-saving way to perform such touch-and-go drinking would be to strive to conserve mechanical energy, by transforming potential energy to kinetic energy during the gliding descent, touching water at high speed, and regaining height with minimal muscular work. Using 3D optical tracking, we recorded 163 swift drinking trajectories, over three waterbodies near Rennes, France. Contrarily to the energy conservation hypothesis, we show that swifts approaching a waterbody with a higher mechanical energy (higher height and/or speed 5 s before contact) do not reach water at higher speeds, but do brake, i.e. dissipate mechanical energy to lose both height and speed. Braking seemed to be linked with sharp turns and the use of headwind to some extent, but finer turns and postural adjustments, beyond the resolving power of our tracking data, could also be involved. We hypothesize that this surprisingly costly behaviour results from a trade-off between energy expenditure and safety, because approaching water surface requires fine motor control, and high speed increases the risk of falling into water, which would have serious energetic and survival costs for a swift.

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