Turbulence causes kinematic and behavioural adjustments in a flapping flier.

Turbulence is a widespread phenomenon in the natural world, but its influence on flapping fliers remains little studied. We assessed how freestream turbulence affected the kinematics, flight effort and track properties of homing pigeons (Columba livia), using the fine-scale variations in flight height as a proxy for turbulence levels. Birds showed a small increase in their wingbeat amplitude with increasing turbulence (similar to laboratory studies), but this was accompanied by a reduction in mean wingbeat frequency, such that their flapping wing speed remained the same. Mean kinematic responses to turbulence may therefore enable birds to increase their stability without a reduction in propulsive efficiency. Nonetheless, the most marked response to turbulence was an increase in the variability of wingbeat frequency and amplitude. These stroke-to-stroke changes in kinematics provide instantaneous compensation for turbulence. They will also increase flight costs. Yet pigeons only made small adjustments to their flight altitude, likely resulting in little change in exposure to strong convective turbulence. Responses to turbulence were therefore distinct from responses to wind, with the costs of high turbulence being levied through an increase in the variability of their kinematics and airspeed. This highlights the value of investigating the variability in flight parameters in free-living animals.

Estimating fine-scale changes in turbulence using the movements of a flapping flier.

All animals that operate within the atmospheric boundary layer need to respond to aerial turbulence. Yet little is known about how flying animals do this because evaluating turbulence at fine scales (tens to approx. 300 m) is exceedingly difficult. Recently, data from animal-borne sensors have been used to assess wind and updraft strength, providing a new possibility for sensing the physical environment. We tested whether highly resolved changes in altitude and body acceleration measured onboard solo-flying pigeons (as model flapping fliers) can be used as qualitative proxies for turbulence. A range of pressure and acceleration proxies performed well when tested against independent turbulence measurements from a tri-axial anemometer mounted onboard an ultralight flying the same route, with stronger turbulence causing increasing vertical displacement. The best proxy for turbulence also varied with estimates of both convective velocity and wind shear. The approximately linear relationship between most proxies and turbulence levels suggests this approach should be widely applicable, providing insight into how turbulence changes in space and time. Furthermore, pigeons were able to fly in levels of turbulence that were unsafe for the ultralight, paving the way for the study of how freestream turbulence affects the costs and kinematics of animal flight.

Pelagic seabirds reduce risk by flying into the eye of the storm.

Cyclones can cause mass mortality of seabirds, sometimes wrecking thousands of individuals. The few studies to track pelagic seabirds during cyclones show they tend to circumnavigate the strongest winds. We tracked adult shearwaters in the Sea of Japan over 11 y and found that the response to cyclones varied according to the wind speed and direction. In strong winds, birds that were sandwiched between the storm and mainland Japan flew away from land and toward the eye of the storm, flying within ≤30 km of the eye and tracking it for up to 8 h. This exposed shearwaters to some of the highest wind speeds near the eye wall (≤21 m s-1) but enabled them to avoid strong onshore winds in the storm's wake. Extreme winds may therefore become a threat when an inability to compensate for drift could lead to forced landings and collisions. Birds may need to know where land is in order to avoid it. This provides additional selective pressure for a map sense and could explain why juvenile shearwaters, which lack a map sense, instead navigating using a compass heading, are susceptible to being wrecked. We suggest that the ability to respond to storms is influenced by both flight and navigational capacities. This may become increasingly pertinent due to changes in extreme weather patterns.

Airflow modelling predicts seabird breeding habitat across islands.

Wind is fundamentally related to shelter and flight performance: two factors that are critical for birds at their nest sites. Despite this, airflows have never been fully integrated into models of breeding habitat selection, even for well-studied seabirds. Here, we use computational fluid dynamics to provide the first assessment of whether flow characteristics (including wind speed and turbulence) predict the distribution of seabird colonies, taking common guillemots Uria aalge breeding on Skomer Island as our study system. This demonstrates that occupancy is driven by the need to shelter from both wind and rain/wave action, rather than airflow characteristics alone. Models of airflows and cliff orientation both performed well in predicting high-quality habitat in our study site, identifying 80% of colonies and 93% of avoided sites, as well as 73% of the largest colonies on a neighbouring island. This suggests generality in the mechanisms driving breeding distributions and provides an approach for identifying habitat for seabird reintroductions considering current and projected wind speeds and directions.

Projected increases in surface melt and ice loss for the Northern and Southern Patagonian Icefields.

The Northern Patagonian Icefield (NPI) and the Southern Patagonian Icefield (SPI) have increased their ice mass loss in recent decades. In view of the impacts of glacier shrinkage in Patagonia, an assessment of the potential future surface mass balance (SMB) of the icefields is critical. We seek to provide this assessment by modelling the SMB between 1976 and 2050 for both icefields, using regional climate model data (RegCM4.6) and a range of emission scenarios. For the NPI, reductions between 1.5 m w.e. (RCP2.6) and 1.9 m w.e. (RCP8.5) were estimated in the mean SMB during the period 2005-2050 compared to the historical period (1976-2005). For the SPI, the estimated reductions were between 1.1 m w.e. (RCP2.6) and 1.5 m w.e. (RCP8.5). Recently frontal ablation estimates suggest that mean SMB in the SPI is positively biased by 1.5 m w.e., probably due to accumulation overestimation. If it is assumed that frontal ablation rates of the recent past will continue, ice loss and sea-level rise contribution will increase. The trend towards lower SMB is mostly explained by an increase in surface melt. Positive ice loss feedbacks linked to increasing in meltwater availability are expected for calving glaciers.

Moving in a moving medium: new perspectives on flight.

One of the defining features of the aerial environment is its variability; air is almost never still. This has profound consequences for flying animals, affecting their flight stability, speed selection, energy expenditure and choice of flight path. All these factors have important implications for the ecology of flying animals, and the ecosystems they interact with, as well as providing bio-inspiration for the development of unmanned aerial vehicles. In this introduction, we touch on the factors that drive the variability in airflows, the scales of variability and the degree to which given airflows may be predictable. We then summarize how papers in this volume advance our understanding of the sensory, biomechanical, physiological and behavioural responses of animals to air flows. Overall, this provides insight into how flying animals can be so successful in this most fickle of environments.This article is part of the themed issue 'Moving in a moving medium: new perspectives on flight'.