Proactive cursorial and ambush predation risk avoidance in four African herbivore species.

Most herbivores must balance demands to meet nutritional requirements, maintain stable thermoregulation and avoid predation. Species-specific predator and prey characteristics determine the ability of prey to avoid predation and the ability of predators to maximize hunting success. Using GPS collar data from African wild dogs, lions, impala, tsessebes, wildebeest and zebra in the Okavango Delta, Botswana, we studied proactive predation risk avoidance by herbivores. We considered predator activity level in relation to prey movement, predator and prey habitat selection, and preferential use of areas by prey. We compared herbivore behaviour to lion and wild dog activity patterns and determined the effect of seasonal resource availability and prey body mass on anti-predator behaviour. Herbivore movement patterns were more strongly correlated with lion than wild dog activity. Habitat selection by predators was not activity level dependent and, while prey and predators differed to some extent in their habitat selection, there were also overlaps, probably caused by predators seeking habitats with high prey abundance. Areas favoured by lions were used by herbivores more when lions were less active, whereas wild dog activity level was not correlated with prey use. Prey body mass was not a strong predictor of the strength of proactive predation avoidance behaviour. Herbivores showed stronger anti-predator behaviours during the rainy season when resources were abundant. Reducing movement when top predators are most active and avoiding areas with a high likelihood of predator use during the same periods appear to be common strategies to minimize predation risk. Such valuable insights into predator-prey dynamics are only possible when using similar data from multiple sympatric species of predator and prey, an approach that should become more prevalent given the ongoing integration of technological methods into ecological studies.

Markerless 3D kinematics and force estimation in cheetahs.

The complex dynamics of animal manoeuvrability in the wild is extremely challenging to study. The cheetah (Acinonyx jubatus) is a perfect example: despite great interest in its unmatched speed and manoeuvrability, obtaining complete whole-body motion data from these animals remains an unsolved problem. This is especially difficult in wild cheetahs, where it is essential that the methods used are remote and do not constrain the animal's motion. In this work, we use data obtained from cheetahs in the wild to present a trajectory optimisation approach for estimating the 3D kinematics and joint torques of subjects remotely. We call this approach kinetic full trajectory estimation (K-FTE). We validate the method on a dataset comprising synchronised video and force plate data. We are able to reconstruct the 3D kinematics with an average reprojection error of 17.69 pixels (62.94% PCK using the nose-to-eye(s) length segment as a threshold), while the estimates produce an average root-mean-square error of 171.3N ( ≈ 17.16% of peak force during stride) for the estimated ground reaction force when compared against the force plate data. While the joint torques cannot be directly validated against ground truth data, as no such data is available for cheetahs, the estimated torques agree with previous studies of quadrupeds in controlled settings. These results will enable deeper insight into the study of animal locomotion in a more natural environment for both biologists and roboticists.

There and back again - a zebra's tale.

Animals need to navigate between resources such as water, food and shelter, and how they achieve this is likely to vary with species. Here, using high-accuracy GPS data, we studied repeated journeys made by wild plains zebra (Equus quagga) through a naturally vegetated environment to explore whether they consistently follow the same route through the area or whether they use a range of routes to reach their goal. We used a model to distinguish and quantify these two possibilities and show that our observations are consistent with the use of multiple routes. Our model performs better than assuming a uniform angular distribution of trajectories. The typical separation of the routes was found to be small (1.96 m), while the scale at which neighbouring trajectories are informative to direction of travel was found to be large (with a confidence interval of 1.19-26.4 m). Our observations are consistent with the hypothesis that zebra are able to navigate without having to return to previously used routes, instead using numerous different routes of similar trajectories.

Artificial mass loading disrupts stable social order in pigeon dominance hierarchies.

Dominance hierarchies confer benefits to group members by decreasing the incidences of physical conflict, but may result in certain lower ranked individuals consistently missing out on access to resources. Here, we report a linear dominance hierarchy remaining stable over time in a closed population of birds. We show that this stability can be disrupted, however, by the artificial mass loading of birds that typically comprise the bottom 50% of the hierarchy. Mass loading causes these low-ranked birds to immediately become more aggressive and rise-up the dominance hierarchy; however, this effect was only evident in males and was absent in females. Removal of the artificial mass causes the hierarchy to return to its previous structure. This interruption of a stable hierarchy implies a strong direct link between body mass and social behaviour and suggests that an individual's personality can be altered by the artificial manipulation of body mass.

There and back again - a zebra's tale.

Animals need to navigate between resources such as water, food and shelter and how they achieve this is likely to vary with species. Here, using high accuracy GPS data, we study repeated journeys made by wild zebra (Equus quagga) through a naturally vegetated environment to explore whether they consistently follow the same route through the area or whether they use a range of routes to reach their goal. We use a model to distinguish and quantify these two possibilities and show that our observations are consistent with the use of multiple routes. Our model performs better than assuming uniform angular distribution of trajectories. The typical separation of the routes was found to be small (1.96 m), while the scale at which neighboring trajectories are informative to direction of travel was found to be large (with a confidence interval of (1.19, 26.4) m). Our observations are consistent with the hypothesis that zebra are able to navigate without having to return to previously used routes, instead using numerous different routes of similar trajectories.

Ground reaction forces of overground galloping in ridden Thoroughbred racehorses.

The horse has evolved to gallop economically at high speed. Limb force increases with speed but direct measures of limb ground reaction forces (GRFs) at gallop are sparse. This study reports GRFs for multiple limbs, using force plates, across seven Thoroughbred racehorses during ridden galloping. The results show peak vertical GRF values of 13.6 N kg-1 (non-lead hindlimb), 12.3 N kg-1 (lead hindlimb), 14.0 N kg-1 (non-lead forelimb) and 13.6 N kg-1 (lead forelimb) at 11.4 m s-1 and recorded values are consistent with those predicted from duty factor. The distribution of body weight between the forelimbs and hindlimbs is approximated to 50:50, and is variable with speed, unlike the 60:40 commonly stated for cursorial quadrupeds in the literature. An even distribution of load on all limbs may help minimise accumulation of fatigue and assist in injury avoidance. Cranio-caudal force data concur with the observation that horses apply a net accelerative impulse with the hindlimbs and a net decelerative impulse with the forelimbs. Capturing GRFs enhances our knowledge on the mechanics of galloping in fast-moving species and provides insight into injury risk and factors limiting athletic performance.

Energy turnover in mammalian skeletal muscle in contractions mimicking locomotion: effects of stimulus pattern on work, impulse and energetic cost and efficiency.

Active muscle performs various mechanical functions during locomotion: work output during shortening, work absorption when resisting (but not preventing) lengthening, and impulse (force-time integral) whenever there is active force. The energetic costs of these functions are important components in the energy budget during locomotion. We investigated how the pattern of stimulation and movement affects the mechanics and energetics of muscle fibre bundles isolated from wild rabbits (Oryctolagus cuniculus). The fibres were from muscles consisting of mainly fast-twitch, type 2 fibres. Fibre length was held constant (isometric) or a sinusoidal pattern of movement was imposed at a frequency similar to the stride frequency of running wild rabbits. Duty cycle (stimulation duration×movement frequency) and phase (timing of stimulation relative to movement) were varied. Work and impulse were measured as well as energy produced as heat. The sum of net work (work output-work input) and heat was taken as a measure of energetic cost. Maximum work output was produced with a long duty cycle and stimulation starting slightly before shortening, and was produced quite efficiently. However, efficiency was even higher with other stimulation patterns that produced less work. The highest impulse (considerably higher than isometric impulse) was produced when stimulation started while the muscle fibres were being lengthened. High impulse was produced very economically because of the low cost of producing force during lengthening. Thus, locomotion demanding high work, high impulse or economical work output or impulse requires a distinct pattern of stimulation and movement.

Parsimonious test of dynamic interaction.

In recent years, there have been significant advances in the technology used to collect data on the movement and activity patterns of humans and animals. GPS units, which form the primary source of location data, have become cheaper, more accurate, lighter and less power-hungry, and their accuracy has been further improved with the addition of inertial measurement units. The consequence is a glut of geospatial time series data, recorded at rates that range from one position fix every several hours (to maximize system lifetime) to ten fixes per second (in high dynamic situations). Since data of this quality and volume have only recently become available, the analytical methods to extract behavioral information from raw position data are at an early stage of development. An instance of this lies in the analysis of animal movement patterns. When investigating solitary animals, the timing and location of instances of avoidance and association are important behavioral markers. In this paper, a novel analytical method to detect avoidance and association between individuals is proposed; unlike existing methods, assumptions about the shape of the territories or the nature of individual movement are not needed. Simulations demonstrate that false positives (type I error) are rare (1%-3%), which means that the test rarely suggests that there is an association if there is none.

Possible causes of divergent population trends in sympatric African herbivores.

Sympatric herbivores experience similar environmental conditions but can vary in their population trends. Identifying factors causing these differences could assist conservation efforts aimed at maintaining fully functional ecosystems. From 1996-2013, tsessebe and wildebeest populations in the Okavango Delta, Botswana, declined by 73% and 90%, respectively, whereas zebra populations remained stable. These sympatric, medium sized herbivores are exposed to similar natural and anthropogenic pressures, but apparently differ in their responses to those pressures. To identify factors that could cause these differences, we fitted GPS-enabled collars to six zebra, eight tsessebe and seven wildebeest in the Moremi Game Reserve, Botswana. We calculated utilisation distributions (UDs) from GPS data, and used 95% isopleths to compare seasonal home range size between species. We calculated utilisation intensity (UI) from the UDs and generated spatial layers representing resources and disturbances, and then used model averaging to identify factors affecting UI for each species. We calculated second and third order habitat selection ratios to determine whether species were habitat specialists or generalists. Zebra occupied larger home ranges than tsessebe and wildebeest, showed weaker responses to spatial variables and displayed no third order habitat selection; zebra social systems are also more fluid, allowing for information exchange between stable harems. Herbivore species that are sedentary, occupy small home ranges, are habitat specialists and exist in relatively isolated groups are likely to be less resistant and resilient to the rapid pace of environmental change forecast by climate change scenarios. Resources contained within existing protected areas are unlikely to maintain populations of such species at sufficiently high levels, potentially leading to functional extinction. Special precautions may be needed to ensure that such species can persist in the wild, such as buffer zones around existing protected areas, which would allow greater potential for adaptive movement should current environmental conditions change.

The running kinematics of free-roaming giraffes, measured using a low cost unmanned aerial vehicle (UAV).

The study of animal locomotion can be logistically challenging, especially in the case of large or unhandleable animals in uncontrolled environments. Here we demonstrate the utility of a low cost unmanned aerial vehicle (UAV) in measuring two-dimensional running kinematics from free-roaming giraffes (Giraffa camelopardalis giraffa) in the Free State Province, South Africa. We collected 120 Hz video of running giraffes, and calibrated each video frame using metatarsal length as a constant object of scale. We tested a number of methods to measure metatarsal length. The method with the least variation used close range photography and a trigonometric equation to spatially calibrate the still image, and derive metatarsal length. In the absence of this option, a spatially calibrated surface model of the study terrain was used to estimate topographical dimensions in video footage of interest. Data for the terrain models were collected using the same equipment, during the same study period. We subsequently validated the accuracy of the UAV method by comparing similar speed measurements of a human subject running on a treadmill, with treadmill speed. At 8 m focal distance we observed an error of 8% between the two measures of speed. This error was greater at a shorter focal distance, and when the subject was not in the central field of view. We recommend that future users maximise the camera focal distance, and keep the subject in the central field of view. The studied giraffes used a grounded rotary gallop with a speed range of 3.4-6.9 ms-1 (never cantering, trotting or pacing), and lower duty factors when compared with other cursorial quadrupeds. As this pattern might result in adverse increases in peak vertical limb forces with speed, it was notable to find that contralateral limbs became more in-phase with speed. Considering the latter pattern and the modest maximal speed of giraffes, we speculate that tissue safety factors are maintained within tolerable bounds this way. Furthermore, the angular kinematics of the neck were frequently isolated from the pitching of the body during running; this may be a result of the large mass of the head and neck. Further field experiments and biomechanical models are needed to robustly test these speculations.

Terrestrial mammalian wildlife responses to Unmanned Aerial Systems approaches.

Unmanned Aerial Systems (UAS) are increasingly being used recreationally, commercially and for wildlife research, but very few studies have quantified terrestrial mammalian reactions to UAS approaches. We used two Vertical Take-off and Landing (VTOL) UAS to approach seven herbivore species in the Moremi Game Reserve, Botswana, after securing the relevant permissions. We recorded responses to 103 vertical and 120 horizontal approaches, the latter from three altitudes above ground level (AGL). We ran mixed logistic regressions to identify factors triggering (i) any response and (ii) an evasive response. We included effects of activity, altitude, direction of approach, distance, habitat, herd type, herd size, other species, target species, time, VTOL type and wind strength. Response triggers were linked to altitude, distance, habitat and target species. Elephant (Loxodonta africana), giraffe (Giraffa camelopardalis), wildebeest (Connochaetes taurinus) and zebra (Equus quagga) were most affected by VTOL approach, impala (Aepyceros melampus) and lechwe (Kobus leche) were least responsive, and tsessebe (Damaliscus lunatus) displayed intermediate sensitivity. VTOLs flown lower than 60 m AGL and closer than 100 m horizontal distance from target animals triggered behavioural responses in most species. Enforced regulations on recreational UAS use in wildlife areas are necessary to minimise disturbance to terrestrial mammals.

The locomotor kinematics and ground reaction forces of walking giraffes.

Giraffes (Giraffa camelopardalis) possess specialised anatomy. Their disproportionately elongate limbs and neck confer recognised feeding advantages, but little is known about how their morphology affects locomotor function. In this study, we examined the stride parameters and ground reaction forces from three adult giraffes in a zoological park, across a range of walking speeds. The patterns of GRFs during walking indicate that giraffes, similar to other mammalian quadrupeds, maintain a forelimb-biased weight distribution. The angular excursion of the neck has functional links with locomotor dynamics in giraffes, and was exaggerated at faster speeds. The horizontal accelerations of the neck and trunk were out of phase compared with the vertical accelerations, which were intermediate between in and out of phase. Despite possessing specialised morphology, giraffes' stride parameters were broadly predicted from dynamic similarity, facilitating the use of other quadrupedal locomotion models to generate testable hypotheses in giraffes.

Remarkable muscles, remarkable locomotion in desert-dwelling wildebeest.

Large mammals that live in arid and/or desert environments can cope with seasonal and local variations in rainfall, food and climate1 by moving long distances, often without reliable water or food en route. The capacity of an animal for this long-distance travel is substantially dependent on the rate of energy utilization and thus heat production during locomotion-the cost of transport2-4. The terrestrial cost of transport is much higher than for flying (7.5 times) and swimming (20 times)4. Terrestrial migrants are usually large1-3 with anatomical specializations for economical locomotion5-9, because the cost of transport reduces with increasing size and limb length5-7. Here we used GPS-tracking collars10 with movement and environmental sensors to show that blue wildebeest (Connochaetes taurinus, 220 kg) that live in a hot arid environment in Northern Botswana walked up to 80 km over five days without drinking. They predominantly travelled during the day and locomotion appeared to be unaffected by temperature and humidity, although some behavioural thermoregulation was apparent. We measured power and efficiency of work production (mechanical work and heat production) during cyclic contractions of intact muscle biopsies from the forelimb flexor carpi ulnaris of wildebeest and domestic cows (Bos taurus, 760 kg), a comparable but relatively sedentary ruminant. The energetic costs of isometric contraction (activation and force generation) in wildebeest and cows were similar to published values for smaller mammals. Wildebeest muscle was substantially more efficient (62.6%) than the same muscle from much larger cows (41.8%) and comparable measurements that were obtained from smaller mammals (mouse (34%)11 and rabbit (27%)). We used the direct energetic measurements on intact muscle fibres to model the contribution of high working efficiency of wildebeest muscle to minimizing thermoregulatory challenges during their long migrations under hot arid conditions.

Movement patterns and athletic performance of leopards in the Okavango Delta.

Although leopards are the most widespread of all the big cats and are known for their adaptability, they are elusive and little is known in detail about their movement and hunting energetics. We used high-resolution GPS/IMU (inertial measurement unit) collars to record position, activity and the first high-speed movement data on four male leopards in the Okavango Delta, an area with high habitat diversity and habitat fragmentation. Leopards in this study were generally active and conducted more runs during the night, with peaks in activity and number of runs in the morning and evening twilight. Runs were generally short (less than 100 m) and relatively slow (maximum speed 5.3 m s-1, mean of individual medians) compared to other large predators. Average daily travel distance was 11 km and maximum daily travel distance was 29 km. No direct correlation was found between average daily temperature and travel distance or between season and travel distance. Total daily energy requirements based on locomotor cost and basal metabolic rate varied little between individuals and over time. This study provides novel insights into movement patterns and athletic performance of leopards through quantitative high-resolution measurement of the locomotor, energetic, spatial and temporal movement characteristics. The results are unbiased by methodological and observational limitations characteristic of previous studies and demonstrate the utility of applying new technologies to field studies of elusive nocturnal species.

Biomechanics of predator-prey arms race in lion, zebra, cheetah and impala.

The fastest and most manoeuvrable terrestrial animals are found in savannah habitats, where predators chase and capture running prey. Hunt outcome and success rate are critical to survival, so both predator and prey should evolve to be faster and/or more manoeuvrable. Here we compare locomotor characteristics in two pursuit predator-prey pairs, lion-zebra and cheetah-impala, in their natural savannah habitat in Botswana. We show that although cheetahs and impalas were universally more athletic than lions and zebras in terms of speed, acceleration and turning, within each predator-prey pair, the predators had 20% higher muscle fibre power than prey, 37% greater acceleration and 72% greater deceleration capacity than their prey. We simulated hunt dynamics with these data and showed that hunts at lower speeds enable prey to use their maximum manoeuvring capacity and favour prey survival, and that the predator needs to be more athletic than its prey to sustain a viable success rate.

Suite of simple metrics reveals common movement syndromes across vertebrate taxa.

BACKGROUND: Because empirical studies of animal movement are most-often site- and species-specific, we lack understanding of the level of consistency in movement patterns across diverse taxa, as well as a framework for quantitatively classifying movement patterns. We aim to address this gap by determining the extent to which statistical signatures of animal movement patterns recur across ecological systems. We assessed a suite of movement metrics derived from GPS trajectories of thirteen marine and terrestrial vertebrate species spanning three taxonomic classes, orders of magnitude in body size, and modes of movement (swimming, flying, walking). Using these metrics, we performed a principal components analysis and cluster analysis to determine if individuals organized into statistically distinct clusters. Finally, to identify and interpret commonalities within clusters, we compared them to computer-simulated idealized movement syndromes representing suites of correlated movement traits observed across taxa (migration, nomadism, territoriality, and central place foraging). RESULTS: Two principal components explained 70% of the variance among the movement metrics we evaluated across the thirteen species, and were used for the cluster analysis. The resulting analysis revealed four statistically distinct clusters. All simulated individuals of each idealized movement syndrome organized into separate clusters, suggesting that the four clusters are explained by common movement syndrome. CONCLUSIONS: Our results offer early indication of widespread recurrent patterns in movement ecology that have consistent statistical signatures, regardless of taxon, body size, mode of movement, or environment. We further show that a simple set of metrics can be used to classify broad-scale movement patterns in disparate vertebrate taxa. Our comparative approach provides a general framework for quantifying and classifying animal movements, and facilitates new inquiries into relationships between movement syndromes and other ecological processes.

An exploratory clustering approach for extracting stride parameters from tracking collars on free-ranging wild animals.

Changes in stride frequency and length with speed are key parameters in animal locomotion research. They are commonly measured in a laboratory on a treadmill or by filming trained captive animals. Here, we show that a clustering approach can be used to extract these variables from data collected by a tracking collar containing a GPS module and tri-axis accelerometers and gyroscopes. The method enables stride parameters to be measured during free-ranging locomotion in natural habitats. As it does not require labelled data, it is particularly suitable for use with difficult to observe animals. The method was tested on large data sets collected from collars on free-ranging lions and African wild dogs and validated using a domestic dog.

Cheetah Reunion - The Challenge of Finding Your Friends Again.

Animals navigate their environment using a variety of senses and strategies. This multiplicity enables them to respond to different navigational requirements resulting from habitat, scale and purpose. One of the challenges social animals face is how to reunite after periods of separation. We explore a variety of possible mechanisms used to reunite the members of a cheetah coalition dispersed within a large area after prolonged separation. Using GPS data from three cheetahs reuniting after weeks of separation, we determined that 1) the likelihood of purely coincidental reunion is miniscule 2) the reunion occurred in an area not normally frequented 3) with very little time spent in the region in advance of the reunion. We therefore propose that timely encounter of scent markings where paths cross is the most likely mechanism used to aid the reunion.

Improving the accuracy of estimates of animal path and travel distance using GPS drift-corrected dead reckoning.

Route taken and distance travelled are important parameters for studies of animal locomotion. They are often measured using a collar equipped with GPS. Collar weight restrictions limit battery size, which leads to a compromise between collar operating life and GPS fix rate. In studies that rely on linear interpolation between intermittent GPS fixes, path tortuosity will often lead to inaccurate path and distance travelled estimates. Here, we investigate whether GPS-corrected dead reckoning can improve the accuracy of localization and distance travelled estimates while maximizing collar operating life. Custom-built tracking collars were deployed on nine freely exercising domestic dogs to collect high fix rate GPS data. Simulations were carried out to measure the extent to which combining accelerometer-based speed and magnetometer heading estimates (dead reckoning) with low fix rate GPS drift correction could improve the accuracy of path and distance travelled estimates. In our study, median 2-dimensional root-mean-squared (2D-RMS) position error was between 158 and 463 m (median path length 16.43 km) and distance travelled was underestimated by between 30% and 64% when a GPS position fix was taken every 5 min. Dead reckoning with GPS drift correction (1 GPS fix every 5 min) reduced 2D-RMS position error to between 15 and 38 m and distance travelled to between an underestimation of 2% and an overestimation of 5%. Achieving this accuracy from GPS alone would require approximately 12 fixes every minute and result in a battery life of approximately 11 days; dead reckoning reduces the number of fixes required, enabling a collar life of approximately 10 months. Our results are generally applicable to GPS-based tracking studies of quadrupedal animals and could be applied to studies of energetics, behavioral ecology, and locomotion. This low-cost approach overcomes the limitation of low fix rate GPS and enables the long-term deployment of lightweight GPS collars.

Determining position, velocity and acceleration of free-ranging animals with a low-cost unmanned aerial system.

Unmanned aerial systems (UASs), frequently referred to as 'drones', have become more common and affordable and are a promising tool for collecting data on free-ranging wild animals. We used a Phantom-2 UAS equipped with a gimbal-mounted camera to estimate position, velocity and acceleration of a subject on the ground moving through a grid of GPS surveyed ground control points (area ∼1200 m(2)). We validated the accuracy of the system against a dual frequency survey grade GPS system attached to the subject. When compared with GPS survey data, the estimations of position, velocity and acceleration had a root mean square error of 0.13 m, 0.11 m s(-1) and 2.31 m s(-2), respectively. The system can be used to collect locomotion and localisation data on multiple free-ranging animals simultaneously. It does not require specialist skills to operate, is easily transported to field locations, and is rapidly and easily deployed. It is therefore a useful addition to the range of methods available for field data collection on free-ranging animal locomotion.