Emily L C Shepard

Swansea University | SWAN · Department of Biosciences

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...

Thermal soaring can offer substantial reductions in flight cost, but it is often assumed to be confined to a relatively narrow group of fliers (those with low wing loading relative to their body mass). Using high-frequency movement data, including magnetometry and GPS, we identified thermal soaring in a seabird previously thought to use only flappi...

Predator-prey arms races have led to the evolution of remarkable disguise strategies. While the theoretical benefits of predator camouflage are well established, no study has yet been able to quantify its consequences for hunting success. High-resolution movement data therefore allowed us to study how barn owls (Tyto alba) conceal their approach wh...

Predator-prey arms races have led to the evolution of remarkable disguise strategies. While the theoretical benefits of predator camouflage are well established, no study has yet been able to quantify its consequences for hunting success. High-resolution movement data therefore allowed us to study how barn owls (Tyto alba) conceal their approach wh...

Background Bio-logging devices play a fundamental and indispensable role in movement ecology studies, particularly in the wild. However, researchers are aware of the influence that attaching devices can have on animals, particularly on their behaviour, energy expenditure and survival. The way a device is attached to an animal’s body has also potent...

Albatrosses exploit winds to travel vast distances across the ocean. Their morphology is adapted for low-cost dynamic soaring flight, but these adaptations confer low manoeuvrability, which may be risky when flying over land. This study investigates how wind conditions influence Endangered grey-headed albatross Thalassarche chrysostoma crashes in t...

Predator-prey arms races have led to the evolution of remarkable disguise strategies. While the theoretical benefits of predator camouflage are well established, no study has yet been able to quantify its consequences for hunting success. High-resolution movement data therefore allowed us to study how barn owls (Tyto alba) conceal their approach wh...

Storms can cause widespread seabird stranding and wrecking,1,2,3,4,5 yet little is known about the maximum wind speeds that birds are able to tolerate or the conditions they avoid. We analyzed >300,000 h of tracking data from 18 seabird species, including flapping and soaring fliers, to assess how flight morphology affects wind selectivity, both at...

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, providin...

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 directio...

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 kinemat...

Quantifying stress and energetic responses in animals is a major challenge as existing methods lack temporal resolution and elevate animal stress. We propose “wake respirometry”, a new method of quantifying fine-scale changes in CO2 production in unrestrained animals, using a non-dispersive infrared CO2 sensor positioned downwind of the animal i.e....

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...

Emily Shepard introduces ways flying animals conserve energy inflight.

Flying seabirds are adapted for windy environments 1,2 . Despite this, storms can cause widespread strandings and wrecks, demonstrating that these seabirds are not always able to avoid or compensate for extreme conditions 3,4,5,6,7 . The maximum wind speeds that birds can operate in should vary with morphology and flight style ⁸ , but this has been...

Quantifying activity-specific energy expenditure in free-living animals is a major challenge as current methods require calibration in the lab and animal capture. We propose "wake respirometry", a new method of quantifying fine-scale changes in CO 2 production in unrestrained animals, using a non-dispersive infrared CO 2 sensor positioned downwind...

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 years and find that the response to cyclones varied according to the wind speed and direc...

Remaining cohesive on the move can be beneficial for animal groups. As such, animal groups have evolved coordination mechanisms such as leadership to resolve navigational conflicts of interest. Consistent "leaders" may have an intrinsic advantage over "followers" which compromise on their preferred route to retain cohesion, which highlights the que...

Accelerometers in animal‐attached tags are powerful tools in behavioural ecology, they can be used to determine behaviour and provide proxies for movement‐based energy expenditure. Researchers are collecting and archiving data across systems, seasons and device types. However, using data repositories to draw ecological inference requires a good und...

Habitat-use and distribution models are essential tools of conservation biology. For wide-ranging species, such models may be challenged by the expanse, remoteness and variability of their habitat, these challenges often being compounded by the species’ mobility. In marine environments, direct observations and sampling are usually impractical over...

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 cha...

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 cha...

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. Strai...

Bio-logging devices play a fundamental and indispensable role in movement ecology studies, particularly in the wild. However, researchers are becoming increasingly aware of the potential effects that attaching devices can have on animals, particularly on their behaviour, energy expenditure and survival. The way a device is attached to an animal's b...

Mapping cyclone paths and seabird wintering areas in the North Atlantic reveals hotspots where they overlap. Modelling the energy expenditure of seabirds suggests that an inability to feed in cyclones is likely to be what makes the high wind conditions deadly.

1. Accelerometers in animal-attached tags have proven to be powerful tools in behavioural ecology, being used to determine behaviour and provide proxies for movement-based energy expenditure. Researchers are collecting and archiving data across systems, seasons and device types. However, in order to use data repositories to draw ecological inferenc...

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. Straig...

The power curve provides a basis for predicting adjustments that animals make in flight speed, for example in relation to wind, distance, habitat foraging quality and objective. However, relatively few studies have examined how animals respond to the landscape below them, which could affect speed and power allocation through modifications in climb...

The power curve provides a basis for predicting adjustments that animals make in flight speed, for example in relation to wind, distance, habitat foraging quality and objective. However, relatively few studies have examined how animals respond to the landscape below them, which could affect speed and power allocation through modifications in climb...

The flight speeds that animals should adopt to minimise energy expenditure in different scenarios can be predicted by the curve of power against speed. However, relatively few studies have examined how animals respond to the landscape below them, which could affect speed through modifications in climb rate and perceived predation risk. We equipped...

Wind is a fundamental driver of the distribution and energy expenditure of birds at sea. Wind can also influence mortality at the nest. Yet airflows have never been fully integrated into models of breeding habitat selection. We use computational fluid dynamics to provide the first assessment of whether and how airflows predict the distribution of s...

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%...

Background Global positioning systems (GPS) and altimeters are increasingly used to monitor vertical space use by aerial species, a key aspect of their ecological niche, that we need to know to manage our own use of the airspace, and to protect those species. However, there are various sources of error in flight height data (“height” above ground,...

The evolution of group living transformed the history of animal life on earth, yielding substantial selective benefits. Yet, without overcoming fundamental challenges such as how to coordinate movements with conspecifics, animals cannot maintain cohesion, and coordination is thus a prerequisite for the evolution of sociality in nonstationary animal...

Background Global positioning systems (GPS) and altimeters are increasingly used to monitor vertical space use by aerial species, a key aspect of their niche that we need to know to understand their ecology and conservation needs, and to manage our own use of the airspace. However, there are various sources of error in flight height data (“height”...

Increasing urbanisation is detrimental for some animal species and potentially advantageous for others. Urban-nesting populations of gulls have undergone rapid population increases worldwide, which has resulted in an increase in human-gull conflicts. In order to inform management and conservation decisions in relation to these populations, more inf...

For fast-flying birds, the ability to respond to wind during landing is critical, as errors can lead to injury or even death. Nonetheless, landing ability, and its ecological significance, remain unstudied. We show that for auks, 60% of attempts to land at their cliff nests fail in a strong breeze (80% in near-gale winds). This is most likely becau...

It is fundamentally important for many animal ecologists to quantify the costs of animal activities, although it is not straightforward to do so. The recording of triaxial acceleration by animal‐attached devices has been proposed as a way forward for this, with the specific suggestion that dynamic body acceleration (DBA) be used as a proxy for move...

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 c...

Energy harvesting by animals is important because it provides the power needed for all metabolic processes. Beyond this, efficient food-finding enhances individual fitness [1] and population viability [2], although rates of energy accumulation are affected by environmental- [3] and individual stochasticity [4]. Typically, differences between indivi...

Many large birds rely on thermal soaring flight to travel cross-country. 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, the...

The development of multisensor animal‐attached tags, recording data at high frequencies, has enormous potential in allowing us to define animal behaviour. The high volumes of data, are pushing us towards machine‐learning as a powerful option for distilling out behaviours. However, with increasing parallel lines of data, systems become more likely t...

Background Precision Livestock Farming (PLF) is a promising approach to minimize the conflicts between socio-economic activities and landscape conservation. However, its application on extensive systems of livestock production can be challenging. The main difficulties arise because animals graze on large natural pastures where they are exposed to c...

Behavioural classification methods and results

Dead Reckoning paths methods and example R code

Spatial data and tag descriptions

Raw 3-axial acceleration and magnetometer data recorded at 40 hz

Additional figures of movement paths with associated behaviours

Raw GPS data 1 minute resolution

Animal‐attached technologies can be powerful means to quantify space‐use and behaviour, however, there are also ethical implications associated with capturing and instrumenting animals. Furthermore, tagging approaches are not necessarily well‐suited for examining the movements of multiple individuals within specific, local areas of interest. Here,...

A range of species exploit anthropogenic food resources in behaviour known as 'raiding'. Such behavioural flexibility is considered a central component of a species' ability to cope with human-induced environmental changes. Here, we study the behavioural processes by which raiding male chacma baboons (Papio ursinus) exploit the opportunities and mi...

One contribution of 13 to a theme issue 'Physiological determinants of social behaviour in animals'. Group living has been proposed to yield benefits that enhance fitness above the level that would be achieved through living as solitary individuals. Dominance hierarchies occur commonly in these social assemblages, and result, by definition, in reso...

798594@swansea.ac.uk The attachment of tags to free-ranging birds has greatly enhanced our understanding of avian biology. From the minutia of behaviours to vast transcontinental movements, animal-borne sensors can be used to answer suites of biological questions, relating to the physiology, behaviour and ecology of wild birds in-situ. Tens of th...

Collisions of large soaring raptors with wind turbines and other infrastructures represent a growing conservation concern. We describe a way to leverage knowledge about raptor soaring behaviour to forecast the probability that raptors fly in the rotor‐swept zone. Soaring raptors are theoretically expected to select energy sources (uplift) optimally...

Background: Accelerometers are powerful sensors in many bio-logging devices, and are increasingly allowing researchers to investigate the performance, behaviour, energy expenditure and even state, of free-living animals. Another sensor commonly used in animal-attached loggers is the magnetometer, which has been primarily used in dead-reckoning or i...

Background The use of accelerometers in bio-logging devices has proved to be a powerful tool for the quantification of animal behaviour. While bio-logging techniques are being used on wide range of species, to date they have only been seldom used with non-human primates. This is likely due to three main factors: the long tradition of direct field o...

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...

Birds modulate their flight paths in relation to regional and global airflows in order to reduce their travel costs. Birds should also respond to fine-scale airflows, although the incidence and value of this remains largely unknown. We resolved the three-dimensional trajectories of gulls flying along a built-up coastline, and used computational flu...

Background We are increasingly using recording devices with multiple sensors operating at high frequencies to produce large volumes of data which are problematic to interpret. A particularly challenging example comes from studies on animals and humans where researchers use animal-attached accelerometers on moving subjects to attempt to quantify beh...

External tags fitted to diving birds can affect them in many ways with the most critical effect being an increase in drag. The effects of transmitters can be even more acute due to the presence of a protruding aerial. The study assesses the impact of PTT antenna on the behaviour and energetics of device-equipped guillemots (Uria aalge) in captivity...

External tags fitted to diving birds can affect them in many ways with the most critical effect being an increase in drag. The effects of transmitters can be even more acute due to the presence of a protruding aerial. The study assesses the impact of PTT antenna on the behaviour and energetics of device-equipped guillemots (Uria aalge) in captivity...

This is the program developed as part of the Biotelemetry Event Tree project. For more details please read the associated article and the supplementary information which contain instructions for use.

Background Accelerometry has been used to identify behaviours through the quantification of body posture and motion for a range of species moving in different media. This technique has not been applied to flight behaviours to the same degree, having only been used to distinguish flapping from soaring flight, even though identifying the type of soar...

Background: Smart tags attached to freely-roaming animals recording multiple parameters at infra-second rates are becoming commonplace, and are transforming our understanding of the way wild animals behave. Interpretation of such data is complex and currently limits the ability of biologists to realise the value of their recorded information. Des...

Background Smart tags attached to freely-roaming animals recording multiple parameters at infra-second rates are becoming commonplace, and are transforming our understanding of the way wild animals behave. Interpretation of such data is complex and currently limits the ability of biologists to realise the value of their recorded information. Descr...

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 behav...

How can the ecological consequences of the increasing use of airspace by humans be minimized? Over the past century, humans have increasingly used the airspace for purposes such as transportation, energy generation, and surveillance. Conflict with wildlife may arise from buildings, turbines, power lines, and antennae that project into space and fro...

The use of biotelemetry methods can provide information on animal behaviour, movement ecology and energetics. However, deployment of biotelemetry equipment on free-living animals incurs risk of damage or loss, which can result in high cost and low sample sizes. To facilitate the uptake of these methods, we have recognized the need for a prescribed...

The ability of many animals to access and exploit food is dependent on the ability to move. In the case of scavenging birds, which use soaring flight to locate and exploit ephemeral resources, the cost and speed of movement vary with meteorological factors. These factors are likely to modify the nature of interspecific interactions, as well as indi...

Abstract The metabolic costs of animal movement have been studied extensively under laboratory conditions, although frequently these are a poor approximation of the costs of operating in the natural, heterogeneous environment. Construction of "energy landscapes," which relate animal locality to the cost of transport, can clarify whether, to what ex...

The tortuosity of the track taken by an animal searching for food profoundly affects search efficiency, which should be optimised to maximise net energy gain. Models examining this generally describe movement as a series of straight steps interspaced by turns, and implicitly assume no turn costs. We used both empirical- and modelling-based approach...

Energy management during the breeding season is crucial for central place foragers since parents need to feed themselves and their offspring while being spatially and temporally constrained. In this work, we used overall dynamic body acceleration as a measure of activity and also to allude to the foraging energy expenditure of breeding Imperial cor...

Dynamic body acceleration (DBA) has been used as a proxy for energy expenditure in logger-equipped animals, with researchers summing the acceleration (overall dynamic body acceleration--ODBA) from the three orthogonal axes of devices. The vector of the dynamic body acceleration (VeDBA) may be a better proxy so this study compared ODBA and VeDBA as...

Intersexual differences in the foraging behaviour have been examined in several seabird species, especially those exhibiting sexual size dimorphism. We studied intersex behavioural differences in the Imperial Cormorant (Phalacrocorax atriceps), a size dimorphic seabird. Twenty adults (11 females and 9 males), breeding at Punta León (43°04′S; 64°2′W...

Current understanding of how animals search for and exploit food resources is based on microeconomic models. Although widely used to examine feeding, such constructs should inform other energy-harvesting situations where theoretical assumptions are met. In fact, some animals extract non-food forms of energy from the environment, such as birds that...

The way in which animals use and acquire energy is fundamental to their fitness. Overall dynamic body acceleration (ODBA) has recently been suggested as a new method for the determination of energy expenditure in wild animals. Although the relationship between ODBA and energy expenditure has been calibrated using gas-respirometry, it has only been...

A theoretically valid proxy of energy expenditure is the acceleration of an animal's mass due to the movement of its body parts. Acceleration can be measured by an accelerometer and recorded onto a data logging device. Relevant studies have usually derived a measure of acceleration from the raw data that represents acceleration purely due to moveme...

1. The energetic costs of different behaviours are critical in modulating the behavioural ecology of free-living animals. Despite this, measurement of energy expenditure in the field has proved difficult. 2. A new method with broad application for field studies has been proposed for determining the rate at which animals expend energy, based on meas...

Current guidelines for instrumenting birds state that external devices should not exceed 3–5% of the birds’ body mass; however, the energetic consequences of carrying any given device mass are likely to vary according to the morphology and ecology of the species concerned. We used a freeware program to estimate the mechanical power requirements of...

Colonial seabirds are central place foragers and likely to be subject to substantial competition for resources. Mechanisms proposed for reducing intra-specific competition include differential inter-sex area use mediated by adult choice. We used GPS loggers and dive recorders to study area use and dive depth in a total of 27 male and 26 female impe...

In air-breathing animals, the time and energy costs of each dive are primarily deter- mined by depth-related upthrust, swim speed and temperature. While studies have previously investigated how animals should optimise their behaviour in relation to either upthrust or speed they have rarely been examined in tandem. We took overall dynamic body accel...