Body-mounted accelerometers provide a new prospect for estimating power use in flying birds, as the signal varies with the two major kinematic determinants of aerodynamic power: wingbeat frequency and amplitude. Yet wingbeat frequency is sometimes used as a proxy for power output in isolation. There is therefore a need to understand which kinematic parameter birds vary and whether this is predicted by flight mode (e.g., accelerating, ascending/descending flight), speed or morphology. We investigate this using high-frequency acceleration data from (i) 14 species flying in the wild, (ii) two species flying in controlled conditions in a wind tunnel and (iii) a review of experimental and field studies. While wingbeat frequency and amplitude were positively correlated, R2 values were generally low, supporting the idea that parameters can vary independently. Indeed, birds were more likely to modulate wingbeat amplitude for more energy-demanding flight modes, including climbing and take-off. Nonetheless, the striking variability even within species and flight types, highlights the complexity of describing the kinematic relationships, which appear sensitive to both the biological and physical context. Notwithstanding this acceleration metrics that incorporate both kinematic parameters should be more robust proxies for power than wingbeat frequency alone.

The role of wingbeat frequency and amplitude in flight power

For decades, studies have highlighted links between wind patterns and the behavior, ecology, distribution, energetics and life history of seabirds. However, only relatively recently have advancements in tracking technologies and improvements in the resolution of globally-available wind data allowed wind impacts on seabirds to be quantified across multiple spatiotemporal scales. Here, we review and synthesize current knowledge of the effects of wind on seabirds. We first describe global patterns of wind circulation and relevant atmospheric processes and discuss the relationship between seabird morphology, flight performance and behavior relative to wind. We then develop a conceptual model linking seabird movement strategies to wind, morphology, flight capabilities and central-place constraint. Finally, we examine how wind influences seabird populations via effects on flight efficiency and energetics, and wind impacts associated with climate variability and severe weather. We conclude by highlighting research priorities for advancing our understanding of the effects of wind on seabird ecology and behavior; these include assessing how and to what extent seabirds use ocean waves for efficient flight, understanding how seabirds sense and anticipate wind patterns, and examining how wind has shaped seabird evolution. Future research should also focus on assessing how wind modulates habitat accessibility, and how this knowledge could be incorporated into theory of seabird habitat use. Moreover, approaches that focus on mechanistic links between climate, wind and demography are needed to assess population-level effects, and will be imperative to understanding how seabirds may be impacted by climate-driven changes to wind patterns. 

Effects of wind on the movement, behavior, energetics, and life history of seabirds

The movements of seabirds during the immature period generally remain poorly understood, primarily due to the challenges involved with tracking birds that do not regularly return to a nest. This knowledge gap prevents us from gaining a full understanding of the areas used by seabird populations. Here, we attempted to track the post-fledging movements of Atlantic Puffins Fratercula arctica from Skomer Island (Wales, UK), by deploying geolocators on chicks ready to leave the nest. Despite our very small return rate (just two loggers out of 54, recording 485 and 196 days of data after fledging, respectively), our results provide a first glimpse into the distribution and scale of movements of young Puffins after fledging. The young Puffins undertook movements comparable in scale to those of post-breeding adults, and there were considerable differences between the two individuals. New initiatives to track juvenile seabirds in much larger numbers will hopefully soon provide more insight into seabird post-fledging movements. 

Post-fledging movements of Atlantic Puffins from Skomer Island

Migratory birds travel over impressively long distances. Consequently, they have to adopt flight regimes being both efficient-in order to spare their metabolic resources-and robust to perturbations. This paper investigates the relationship between both aspects, i.e., energetic performance and stability, in flapping flight of migratory birds. Relying on a poly-articulated wing morphing model and a tail-like surface, several families of steady flight regime have been identified and analysed. These families differ by their wing kinematics and tail opening. A systematic parametric search analysis has been carried out, in order to evaluate power consumption and cost of transport. A framework tailored for assessing limit cycles, namely Floquet theory, is used to numerically study flight stability. Our results show that under certain conditions, an inherent passive stability of steady and level flight can be achieved. In particular, we find that progressively opening the tail leads to passively stable flight regimes. Within these passively stable regimes, the tail can produce either upward or downward lift. However, these configurations entail an increase of cost of transport at high velocities penalizing fast forward flight regimes. Our model-based predictions suggest that long range flights require a furled tail configuration, as confirmed by field observations, and consequently need to rely on alternative mechanisms to stabilize the flight. 

/pdf/on-the-role-of-tail-in-stability-and-energetic-cost-of-bird-1t1wx3pq.pdf

On the role of tail in stability and energetic cost of bird flapping flight

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.

Turbulence causes kinematic and behavioural adjustments in a flapping flier

Abstract Animal‐borne telemetry devices provide essential insights into the life‐history strategies of far‐ranging species and allow us to understand how they interact with their environment. Many species in the seabird family Alcidae undergo a synchronous molt of all primary flight feathers during the non‐breeding season, making them flightless and more susceptible to environmental stressors, including severe storms and prey shortages. However, the timing and location of molt remain largely unknown, with most information coming from studies on birds killed by storms or shot by hunters for food. Using light‐level geolocators with saltwater immersion loggers, we develop a method for determining flightless periods in the context of the annual cycle. Four Atlantic puffins (Fratercula arctica) were equipped with geolocator/immersion loggers on each leg to attempt to overcome issues of leg tucking in plumage while sitting on the water, which confounds the interpretation of logger data. Light‐level and saltwater immersion time‐series data were combined to correct for this issue. This approach was adapted and applied to 40 puffins equipped with the standard practice deployments of geolocators on one leg only. Flightless periods consistent with molt were identified in the dual‐equipped birds, whereas molt identification in single‐equipped birds was less effective and definitive and should be treated with caution. Within the dual‐equipped sample, we present evidence for two flightless molt periods per non‐breeding season in two puffins that undertook more extensive migrations (>2000 km) and were flightless for up to 77 days in a single non‐breeding season. A biannual flight feather molt is highly unusual among non‐passerine birds and may be unique to birds that undergo catastrophic molt, i.e., become flightless when molting. Although our conclusions are based on a small sample, we have established a freely available methodological framework for future investigation of the molt patterns of this and other seabird species.

A new biologging approach reveals unique flightless molt strategies of Atlantic puffins

Many vertebrates show lateralized behaviour, or handedness, where an individual preferentially uses one side of the body more than the other. This is generally thought to be caused by brain lateralization and allows functional specializations such as sight, locomotion, and decision-making among other things. We deployed accelerometers on 51 northern gannets, Morus bassanus, to test for behavioural lateralization during plunge dives. When plunge diving, gannets ‘roll’ to one side, and standard indices indicated that 51% of individuals were left-sided, 43% right-sided, and 6% ‘non-lateralized’. Lateralization indices provide no measure of error and do not account for environmental covariance, so we conducted two repeatability analyses on individuals' dive roll direction and angle. Dive side lateralization was highly repeatable among individuals over time at the population level (R = 0.878, p < 0.001). Furthermore, roll angle was also highly repeatable in individuals (R = 0.751, p < 0.001) even after controlling for lateralized state. Gannets show individual specializations in two different parts of the plunge diving process when attempting to catch prey. This is the first demonstration of lateralization during prey capture in a foraging seabird. It is also one of the few demonstrations of behavioural lateralization in a mixed model approach, providing a structure for further exploring behavioural lateralization.

Handedness and individual roll-angle specialism when plunge diving in the northern gannet

Spatial Modelling of Aerial Survey Data Reveals an Important European Storm‐Petrel Hotspot and Its Underlying Drivers Within the North‐East Atlantic

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The role of wingbeat frequency and amplitude in flight power

A new biologging approach reveals unique flightless molt strategies of Atlantic puffins

Handedness and individual roll-angle specialism when plunge diving in the northern gannet

Spatial Modelling of Aerial Survey Data Reveals an Important European Storm‐Petrel Hotspot and Its Underlying Drivers Within the North‐East Atlantic