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Bird migrations are impressive behavioral phenomena, representing complex spatiotemporal strategies to balance costs of living while maximizing fitness. The field of bird migration research has made great strides over the past decades, yet fundamental gaps remain. Technologies have sparked a transformation in the study of bird migration research by revealing remarkable insights into the underlying behavioral, cognitive, physiological and evolutionary mechanisms of these diverse journeys. Here, we aim to encourage broad discussions and promote future studies by highlighting research fields that are characterized by major knowledge gaps or conflicting evidence, namely the fields of navigation, social learning, individual development, energetics and conservation. We approach each topic by summarizing the current state of knowledge and provide a future outlook of ideas and state-of-the-art methods to further advance the field. Integrating knowledge across these disciplines will allow us to understand the adaptive abilities of different species and to develop effective conservation strategies in a rapidly changing world.

Animal behavior is elicited, in part, in response to external conditions, but understanding how animals perceive the environment and make the decisions that bring about these behavioral responses is challenging.Animal heads often move during specific behaviors and, additionally, typically have sensory systems (notably vision, smell, and hearing) sampling in defined arcs (normally to the front of their heads). As such, head‐mounted electronic sensors consisting of accelerometers and magnetometers, which can be used to determine the movement and directionality of animal heads (where head “movement” is defined here as changes in heading [azimuth] and/or pitch [elevation angle]), can potentially provide information both on behaviors in general and also clarify which parts of the environment the animals might be prioritizing (“environmental framing”).We propose a new approach to visualize the data of such head‐mounted tags that combines the instantaneous outputs of head heading and pitch in a single intuitive spherical plot. This sphere has magnetic heading denoted by “longitude” position and head pitch by “latitude” on this “orientation sphere” (O‐sphere).We construct the O‐sphere for the head rotations of a number of vertebrates with contrasting body shape and ecology (oryx, sheep, tortoises, and turtles), illustrating various behaviors, including foraging, walking, and environmental scanning. We also propose correcting head orientations for body orientations to highlight specific heading‐independent head rotation, and propose the derivation of O‐sphere‐metrics, such as angular speed across the sphere. This should help identify the functions of various head behaviors.Visualizations of the O‐sphere provide an intuitive representation of animal behavior manifest via head orientation and rotation. This has ramifications for quantifying and understanding behaviors ranging from navigation through vigilance to feeding and, when used in tandem with body movement, should provide an important link between perception of the environment and response to it in free‐ranging animals., We provide a new visualization of animal body and head orientation that shows in an intuitive manner how the animal frames the environment. Metrics derived from this should help us understand how animals make decisions to adopt particular behaviours.

Background Understanding what animals do in time and space is important for a range of ecological questions, however accurate estimates of how animals use space is challenging. Within the use of animal-attached tags, radio telemetry (including the Global Positioning System, ‘GPS’) is typically used to verify an animal’s location periodically. Straight lines are typically drawn between these ‘Verified Positions’ (‘VPs’) so the interpolation of space-use is limited by the temporal and spatial resolution of the system’s measurement. As such, parameters such as route-taken and distance travelled can be poorly represented when using VP systems alone. Dead-reckoning has been suggested as a technique to improve the accuracy and resolution of reconstructed movement paths, whilst maximising battery life of VP systems. This typically involves deriving travel vectors from motion sensor systems and periodically correcting path dimensions for drift with simultaneously deployed VP systems. How often paths should be corrected for drift, however, has remained unclear. Methods and results Here, we review the utility of dead-reckoning across four contrasting model species using different forms of locomotion (the African lion Panthera leo, the red-tailed tropicbird Phaethon rubricauda, the Magellanic penguin Spheniscus magellanicus, and the imperial cormorant Leucocarbo atriceps). Simulations were performed to examine the extent of dead-reckoning error, relative to VPs, as a function of Verified Position correction (VP correction) rate and the effect of this on estimates of distance moved. Dead-reckoning error was greatest for animals travelling within air and water. We demonstrate how sources of measurement error can arise within VP-corrected dead-reckoned tracks and propose advancements to this procedure to maximise dead-reckoning accuracy. Conclusions We review the utility of VP-corrected dead-reckoning according to movement type and consider a range of ecological questions that would benefit from dead-reckoning, primarily concerning animal–barrier interactions and foraging strategies.

Background Fine-scale data on animal position are increasingly enabling us to understand the details of animal movement ecology and dead-reckoning, a technique integrating motion sensor-derived information on heading and speed, can be used to reconstruct fine-scale movement paths at sub-second resolution, irrespective of the environment. On its own however, the dead-reckoning process is prone to cumulative errors, so that position estimates quickly become uncoupled from true location. Periodic ground-truthing with aligned location data (e.g., from global positioning technology) can correct for this drift between Verified Positions (VPs). We present step-by-step instructions for implementing Verified Position Correction (VPC) dead-reckoning in R using the tilt-compensated compass method, accompanied by the mathematical protocols underlying the code and improvements and extensions of this technique to reduce the trade-off between VPC rate and dead-reckoning accuracy. These protocols are all built into a user-friendly, fully annotated VPC dead-reckoning R function; Gundog.Tracks, with multi-functionality to reconstruct animal movement paths across terrestrial, aquatic, and aerial systems, provided within the Additional file 4 as well as online (GitHub). Results The Gundog.Tracks function is demonstrated on three contrasting model species (the African lion Panthera leo, the Magellanic penguin Spheniscus magellanicus, and the Imperial cormorant Leucocarbo atriceps) moving on land, in water and in air. We show the effect of uncorrected errors in speed estimations, heading inaccuracies and infrequent VPC rate and demonstrate how these issues can be addressed. Conclusions The function provided will allow anyone familiar with R to dead-reckon animal tracks readily and accurately, as the key complex issues are dealt with by Gundog.Tracks. This will help the community to consider and implement a valuable, but often overlooked method of reconstructing high-resolution animal movement paths across diverse species and systems without requiring a bespoke application.

Quantum simulation using synthetic systems is a promising route to solve outstanding quantum many-body problems in regimes where other approaches, including numerical ones, fail

Significance Flapping flight is extremely costly for large birds, yet little is known about the conditions that force them to flap. We attached custom-made “flight recorders” to Andean condors, the world’s heaviest soaring birds, documenting every single wingbeat and when and how individuals gained altitude. Remarkably, condors flapped for only 1% of their flight time, specifically during takeoff and when close to the ground. This is particularly striking as the birds were immature. Thus, our results demonstrate that even inexperienced birds can cover vast distances over land without flapping. Overall, this can help explain how extinct birds with twice the wingspan of condors could have flown., Flight costs are predicted to vary with environmental conditions, and this should ultimately determine the movement capacity and distributions of large soaring birds. Despite this, little is known about how flight effort varies with environmental parameters. We deployed bio-logging devices on the world’s heaviest soaring bird, the Andean condor (Vultur gryphus), to assess the extent to which these birds can operate without resorting to powered flight. Our records of individual wingbeats in >216 h of flight show that condors can sustain soaring across a wide range of wind and thermal conditions, flapping for only 1% of their flight time. This is among the very lowest estimated movement costs in vertebrates. One bird even flew for >5 h without flapping, covering ∼172 km. Overall, > 75% of flapping flight was associated with takeoffs. Movement between weak thermal updrafts at the start of the day also imposed a metabolic cost, with birds flapping toward the end of glides to reach ephemeral thermal updrafts. Nonetheless, the investment required was still remarkably low, and even in winter conditions with weak thermals, condors are only predicted to flap for ∼2 s per kilometer. Therefore, the overall flight effort in the largest soaring birds appears to be constrained by the requirements for takeoff.

Thus far, ecophysiology research has predominantly been conducted within controlled laboratory-based environments, owing to a mismatch between the recording technologies available for physiological monitoring in wild animals and the suite of behaviours and environments they need to withstand, without unduly affecting subjects. While it is possible to record some physiological variables for free-living animals using animal-attached logging devices, including inertial-measurement, heart-rate and temperature loggers, the field is still in its infancy. In this opinion piece, we review the most important future research directions for advancing the field of ‘physiologging’ in wild animals, including the technological development that we anticipate will be required, and the fiscal and ethical challenges that must be overcome. Non-invasive, multi-sensor miniature devices are ubiquitous in the world of human health and fitness monitoring, creating invaluable opportunities for animal and human physiologging to drive synergistic advances. We argue that by capitalizing on the research efforts and advancements made in the development of human wearables, it will be possible to design the non-invasive loggers needed by ecophysiologists to collect accurate physiological data from free-ranging animals ethically and with an absolute minimum of impact. In turn, findings have the capacity to foster transformative advances in human health monitoring. Thus, we invite biomedical engineers and researchers to collaborate with the animal-tagging community to drive forward the advancements necessary to realize the full potential of both fields. This article is part of the theme issue ‘Measuring physiology in free-living animals (Part II)’.

International audience; Many large birds rely on thermal soaring flight to travel crosscountry. As such, they are under selective pressure to minimise the time spent gaining altitude in thermal updrafts. Birds should be able to maximise their climb rates by maintaining a position close to the thermal core through careful selection of bank angle and airspeed; however, there have been few direct measurements of either parameter. Here, we apply a novel methodology to quantify the bank angles selected by soaring birds using on-board magnetometers. We couple these data with airspeed measurements to parameterise the soaring envelope of two species of Gyps vulture, from which it is possible to predict 'optimal' bank angles. Our results show that these large birds respond to the challenges of gaining altitude in the initial phase of the climb, where thermal updrafts are weak and narrow, by adopting relatively high, and conserved, bank angles (25-35 deg). The bank angle decreased with increasing altitude, in a manner that was broadly consistent with a strategy of maximising the rate of climb. However, the lift coefficients estimated in our study were lower than those predicted by theoretical models and wind-tunnel studies. Overall, our results highlight how the relevant currency for soaring performance changes within individual climbs: when thermal radius is limiting, birds vary bank angle and maintain a constant airspeed, but speed increases later in the climb in order to respond to decreasing air density.

Biologists studying animals in their natural environment are increasingly using sensors such as accelerometers in animal-attached `smart' tags because it is widely acknowledged that this approach can enhance the understanding of ecological and behavioural processes. The potential of such tags is tempered by the difficulty of extracting animal behaviour from the sensors which is currently primarily dependent on the manual inspection of multiple time series graphs. This is time consuming and error-prone for the domain expert and is now the limiting factor for realising the value of tags in this area. We introduce TimeClassifier, a visual analytic system for the classification of time series data for movement ecologists. We deploy our system with biologists and report two real-world case studies of its use.

Vultures are thought to form networks in the sky, with individuals monitoring the movements of others to gain up-to-date information on resource availability. While it is recognized that social information facilitates the search for carrion, how this facilitates the search for updrafts, another critical resource, remains unknown. In theory, birds could use information on updraft availability to modulate their flight speed, increasing their airspeed when informed on updraft location. In addition, the stylized circling behaviour associated with thermal soaring is likely to provide social cues on updraft availability for any bird operating in the surrounding area. We equipped fiveGypsvultures with GPS and airspeed loggers to quantify the movements of birds flying in the same airspace. Birds that were socially informed on updraft availability immediately adopted higher airspeeds on entering the inter-thermal glide; a strategy that would be risky if birds were relying on personal information alone. This was embedded within a broader pattern of a reduction in airspeed (approx. 3 m s−1) through the glide, likely reflecting the need for low speed to sense and turn into the next thermal. Overall, this demonstrates (i) the complexity of factors affecting speed selection over fine temporal scales and (ii) thatGypsvultures respond to social information on the occurrence of energy in the aerial environment, which may reduce uncertainty in their movement decisions.

Background Whether, and how, animals move requires them to assess their environment to determine the most appropriate action and trajectory, although the precise way the environment is scanned has been little studied. We hypothesized that head attitude, which effectively frames the environment for the eyes, and the way it changes over time, would be modulated by the environment. Method To test this, we used a head-mounted device (Human-Interfaced Personal Observation platform - HIPOP) on people moving through three different environments; a botanical garden (‘green’ space), a reef (‘blue’ space), and a featureless corridor, to examine if head movement in the vertical axis differed between environments. Template matching was used to identify and quantify distinct behaviours. Conclusions The data on head pitch from all subjects and environments over time showed essentially continuous clear waveforms with varying amplitude and wavelength. There were three stylised behaviours consisting of smooth, regular peaks and troughs in head pitch angle and variable length fixations during which the head pitch remained constant. These three behaviours accounted for ca. 40 % of the total time, with irregular head pitch changes accounting for the rest. There were differences in rates of manifestation of behaviour according to environment as well as environmentally different head pitch values of peaks, troughs and fixations. Finally, although there was considerable variation in head pitch angles, the peak and trough values bounded most of the variation in the fixation pitch values. It is suggested that the constant waveforms in head pitch serve to inform people about their environment, providing a scanning mechanism. Particular emphasis to certain sectors is manifest within the peak and trough limits and these appear modulated by the distribution of the points where fixation, interpreted as being due to objects of interest, occurs. This behaviour explains how animals allocate processing resources to the environment and shows promise for movement studies attempting to elucidate which parts of the environment affect movement trajectories.