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Because it can lead to retaliatory killing, livestock depredation by large carnivores is among the foremost threats to carnivore conservation, and it severely impacts human well-being worldwide. Ongoing climate change can amplify these human-wildlife conflicts, but such issues are largely unexplored, though are becoming increasingly recognized. Here, we assessed how the availability of primary resources and wild prey interact to shape large carnivore selection for livestock rather than wild prey (i.e., via prey switching or apparent competition). Specifically, we combined remotely sensed estimates of primary resources (i.e., water availability and primary productivity), wild prey movement, and 7 years (2015-2021) of reports for livestock depredation by African lions ( Panthera leo ) in the Makgadikgadi Pans ecosystem, Botswana. Although livestock depredation did not vary between wet versus dry seasons, analyses at finer temporal scales revealed higher incidences of livestock depredation when primary production, water availability, and wild prey availability were lower, though the effects of wild prey availability were mediated by water availability. Increased precipitation also amplified livestock depredation events despite having no influence on wild prey availability. Our results suggest that livestock depredation is influenced by the diverse responses of livestock, wild prey, and lions to primary resource availability, a driver that is largely overlooked or oversimplified in studies of human-carnivore conflict. Our findings provide insight into tailoring potential conflict mitigation strategies to fine-scale changes in resource conditions to efficiently reduce conflict and support human livelihoods., Competing Interests: The authors declare no conflicts of interest., (© 2024 The Author(s). Ecology and Evolution published by John Wiley & Sons Ltd.)

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., (© 2024 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

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., Competing Interests: The authors have declared that no competing interests exist.

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.

A simple model is developed to find vertical force profiles and stance durations that minimize either limb mechanical work or peak power demands during bipedal locomotion. The model predicts that work minimization is achieved with a symmetrical vertical force profile, consistent with previous models and observations of adult humans, and data for 487 participants (predominantly 11-18 years old) required to walk at a range of speeds at a Science Fair. Work minimization also predicts the discrete walk-run transition, familiar for adult humans. In contrast, modeled peak limb mechanical power demands are minimized with an early skew in vertical ground reaction force that increases with speed, and stance durations that decrease steadily with speed across the work minimizing walk-run transition speed. The peak power minimization model therefore predicts a continuous walk-run gait transition that is quantitatively consistent with measurements of younger children (1.1-4.7 years) required to locomote at a range of speeds but free to select their own gaits., (Copyright © 2018 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Large mammals that live in arid and/or desert environments can cope with seasonal and local variations in rainfall, food and climate 1 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 transport 2-4 . The terrestrial cost of transport is much higher than for flying (7.5 times) and swimming (20 times) 4 . Terrestrial migrants are usually large 1-3 with anatomical specializations for economical locomotion 5-9 , because the cost of transport reduces with increasing size and limb length 5-7 . Here we used GPS-tracking collars 10 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.

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., (© 2018 The Author(s).)

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.

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., (© 2017. Published by The Company of Biologists Ltd.)

Botswana has the second highest population of cheetah (Acinonyx jubatus) with most living outside protected areas. As a result, many cheetahs are found in farming areas which occasionally results in human-wildlife conflict. This study aimed to look at movement patterns of cheetahs in farming environments to determine whether cheetahs have adapted their movements in these human-dominated landscapes. We fitted high-time resolution GPS collars to cheetahs in the Ghanzi farmlands of Botswana. GPS locations were used to calculate home range sizes as well as number and duration of visits to landscape features using a time-based local convex hull method. Cheetahs had medium-sized home ranges compared to previously studied cheetah in similar farming environments. Results showed that cheetahs actively visited scent marking trees and avoided visiting homesteads. A slight preference for visiting game farms over cattle farms was found, but there was no difference in duration of visits between farm types. We conclude that cheetahs selected for areas that are important for their dietary and social needs and prefer to avoid human-occupied areas. Improved knowledge of how cheetahs use farmlands can allow farmers to make informed decisions when developing management practices and can be an important tool for reducing human-wildlife conflict., Competing Interests: The authors declare no competing or financial interests., (© 2017. Published by The Company of Biologists Ltd.)

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., Competing Interests: The authors have declared that no competing interests exist.

We compare kinematics and wake structure over a range of flight speeds (4.0-8.2 m s(-1)) for two bats that pursue insect prey aerially, Tadarida brasiliensis and Myotis velifer Body mass and wingspan are similar in these species, but M. velifer has broader wings and lower wing loading. By using high-speed videography and particle image velocimetry of steady flight in a wind tunnel, we show that three-dimensional kinematics and wake structure are similar in the two species at the higher speeds studied, but differ at lower speeds. At lower speeds, the two species show significant differences in mean angle of attack, body-wingtip distance and sweep angle. The distinct body vortex seen at low speed in T. brasiliensis and other bats studied to date is considerably weaker or absent in M. velifer We suggest that this could be influenced by morphology: (i) the narrower thorax in this species probably reduces the body-induced discontinuity in circulation between the two wings and (ii) the wing loading is lower, hence the lift coefficient required for weight support is lower. As a result, in M. velifer, there may be a decreased disruption in the lift generation between the body and the wing, and the strength of the characteristic root vortex is greatly diminished, both suggesting increased flight efficiency.This article is part of the themed issue 'Moving in a moving medium: new perspectives on flight'., (© 2016 The Author(s).)

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., (© 2016. Published by The Company of Biologists Ltd.)

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.

African wild dogs (Lycaon pictus) are reported to hunt with energetically costly long chase distances. We used high-resolution GPS and inertial technology to record 1,119 high-speed chases of all members of a pack of six adult African wild dogs in northern Botswana. Dogs performed multiple short, high-speed, mostly unsuccessful chases to capture prey, while cheetahs (Acinonyx jubatus) undertook even shorter, higher-speed hunts. We used an energy balance model to show that the energy return from group hunting and feeding substantially outweighs the cost of multiple short chases, which indicates that African wild dogs are more energetically robust than previously believed. Comparison with cheetah illustrates the trade-off between sheer athleticism and high individual kill rate characteristic of cheetahs, and the energetic robustness of frequent opportunistic group hunting and feeding by African wild dogs.

African wild dogs (Lycaon pictus) are described as highly collaborative endurance pursuit hunters based on observations derived primarily from the grass plains of East Africa. However, the remaining population of this endangered species mainly occupies mixed woodland savannah where hunting strategies appear to differ from those previously described. We used high-resolution GPS and inertial technology to record fine-scale movement of all members of a single pack of six adult African wild dogs in northern Botswana. The dogs used multiple short-distance hunting attempts with a low individual kill rate (15.5%), but high group feeding rate due to the sharing of prey. Use of high-level cooperative chase strategies (coordination and collaboration) was not recorded. In the mixed woodland habitats typical of their current range, simultaneous, opportunistic, short-distance chasing by dogs pursuing multiple prey (rather than long collaborative pursuits of single prey by multiple individuals) could be the key to their relative success in these habitats.

Terrestrial locomotion on legs is energetically expensive. Compared with cycling, or with locomotion in swimming or flying animals, walking and running are highly uneconomical. Legged gaits that minimise mechanical work have previously been identified and broadly match walking and running at appropriate speeds. Furthermore, the 'cost of muscle force' approaches are effective in relating locomotion kinetics to metabolic cost. However, few accounts have been made for why animals deviate from either work-minimising or muscle-force-minimising strategies. Also, there is no current mechanistic account for the scaling of locomotion kinetics with animal size and speed. Here, we report measurements of ground reaction forces in walking children and adult humans, and their stance durations during running. We find that many aspects of gait kinetics and kinematics scale with speed and size in a manner that is consistent with minimising muscle activation required for the more demanding between mechanical work and power: spreading the duration of muscle action reduces activation requirements for power, at the cost of greater work demands. Mechanical work is relatively more demanding for larger bipeds--adult humans--accounting for their symmetrical M-shaped vertical force traces in walking, and relatively brief stance durations in running compared with smaller bipeds--children. The gaits of small children, and the greater deviation of their mechanics from work-minimising strategies, may be understood as appropriate for their scale, not merely as immature, incompletely developed and energetically sub-optimal versions of adult gaits., (© 2015. Published by The Company of Biologists Ltd.)

Many species travel in highly organized groups. The most quoted function of these configurations is to reduce energy expenditure and enhance locomotor performance of individuals in the assemblage. The distinctive V formation of bird flocks has long intrigued researchers and continues to attract both scientific and popular attention. The well-held belief is that such aggregations give an energetic benefit for those birds that are flying behind and to one side of another bird through using the regions of upwash generated by the wings of the preceding bird, although a definitive account of the aerodynamic implications of these formations has remained elusive. Here we show that individuals of northern bald ibises (Geronticus eremita) flying in a V flock position themselves in aerodynamically optimum positions, in that they agree with theoretical aerodynamic predictions. Furthermore, we demonstrate that birds show wingtip path coherence when flying in V positions, flapping spatially in phase and thus enabling upwash capture to be maximized throughout the entire flap cycle. In contrast, when birds fly immediately behind another bird--in a streamwise position--there is no wingtip path coherence; the wing-beats are in spatial anti-phase. This could potentially reduce the adverse effects of downwash for the following bird. These aerodynamic accomplishments were previously not thought possible for birds because of the complex flight dynamics and sensory feedback that would be required to perform such a feat. We conclude that the intricate mechanisms involved in V formation flight indicate awareness of the spatial wake structures of nearby flock-mates, and remarkable ability either to sense or predict it. We suggest that birds in V formation have phasing strategies to cope with the dynamic wakes produced by flapping wings.

There is an ongoing debate about the reasons underlying gait transition in terrestrial locomotion. In bipedal locomotion, the 'compass gait', a reductionist model of inverted pendulum walking, predicts the boundaries of speed and step length within which walking is feasible. The stance of the compass gait is energetically optimal-at walking speeds-owing to the absence of leg compression/extension; completely stiff limbs perform no work during the vaulting phase. Here, we extend theoretical compass gait vaulting to include inclines, and find good agreement with previous observations of changes in walk-run transition speed (approx. 1% per 1% incline). We measured step length and frequency for humans walking either on the level or up a 9.8 per cent incline and report preferred walk-run, walk-compliant-walk and maximum walk-run transition speeds. While the measured 'preferred' walk-run transition speed lies consistently below the predicted maximum walking speeds, and 'actual' maximum walking speeds are clearly above the predicted values, the onset of compliant walking in level as well as incline walking occurs close to the predicted values. These findings support the view that normal human walking is constrained by the physics of vaulting, but preferred absolute walk-run transition speeds may be influenced by additional factors.

Mechanically, the most economical gait for slow bipedal locomotion requires walking as an 'inverted pendulum', with: I, an impulsive, energy-dissipating leg compression at the beginning of stance; II, a stiff-limbed vault; and III, an impulsive, powering push-off at the end of stance. The characteristic 'M'-shaped vertical ground reaction forces of walking in humans reflect this impulse-vault-impulse strategy. Humans achieve this gait by dissipating energy during the heel-to-sole transition in early stance, approximately stiff-limbed, flat-footed vaulting over midstance and ankle plantarflexion (powering the toes down) in late stance. Here, we show that the 'M'-shaped walking ground reaction force profile does not require the plantigrade human foot or heel-sole-toe stance; it is maintained in tip-toe and high-heel walking as well as in ostriches. However, the unusual, stiff, human foot structure--with ground-contacting heel behind ankle and toes in front--enables both mechanically economical inverted pendular walking and physiologically economical muscle loading, by producing extreme changes in mechanical advantage between muscles and ground reaction forces. With a human foot, and heel-sole-toe strategy during stance, the shin muscles that dissipate energy, or calf muscles that power the push-off, need not be loaded at all--largely avoiding the 'cost of muscle force'--during the passive vaulting phase.

Bipedal animals experience ground reaction forces (GRFs) that pass close to the centre of mass (CoM) throughout stance, first decelerating the body, then re-accelerating it during the second half of stance. This results in fluctuations in kinetic energy, requiring mechanical work from the muscles. However, here we show analytically that, in extreme cases (with a very large body pitch moment of inertia), continuous alignment of the GRF through the CoM requires greater mechanical work than a maintained vertical force; we show numerically that GRFs passing between CoM and vertical throughout stance are energetically favourable under realistic conditions; and demonstrate that the magnitude, if not the precise form, of actual CoM-torque profiles in running is broadly consistent with simple mechanical work minimization for humans with appropriate pitch moment of inertia. While the potential energetic savings of CoM-torque support strategies are small (a few per cent) over the range of human running, their importance increases dramatically at high speeds and stance angles. Fast, compliant runners or hoppers would benefit considerably from GRFs more vertical than the zero-CoM-torque strategy, especially with bodies of high pitch moment of inertia--suggesting a novel advantage to kangaroos of their peculiar long-head/long-tail structure.

To date, wake measurements using particle image velocimetry (PIV) of bats in flight have studied only three bat species, all fruit and nectar feeders. In this study, we present the first wake structure analysis for an insectivorous bat. Tadarida brasiliensis, the Brazilian free-tailed bat, is an aerial hunter that annually migrates long distances and also differs strikingly from the previously investigated species morphologically. We compare the aerodynamics of T. brasiliensis with those of other, frugivorous bats and with common swifts, Apus apus, a bird with wing morphology, kinematics and flight ecology similar to that of these bats. The comparison reveals that, for the range of speeds evaluated, the cyclical pattern of aerodynamic forces associated with a wingbeat shows more similarities between T. brasiliensis and A. apus than between T. brasiliensis and other frugivorous bats.

We investigated the detailed kinematics and wake structure of lesser dog-faced fruit bats (Cynopterus brachyotis) flying in a wind tunnel. High speed recordings of the kinematics were conducted to obtain three-dimensional reconstructions of wing movements. Simultaneously, the flow structure in the spanwise plane perpendicular to the flow stream was visualized using time-resolved particle image velocimetry. The flight of four individuals was investigated to reveal patterns in kinematics and wake structure typical for lower and higher speeds. The wake structure identified as typical for both speed categories was a closed-loop ring vortex consisting of the tip vortex and the limited appearance of a counter-rotating vortex near the body, as well as a small distally located vortex system at the end of the upstroke that generated negative lift. We also investigated the degree of consistency within trials and looked at individual variation in flight parameters, and found distinct differences between individuals as well as within individuals.

Over the last decade, interest in animal flight has grown, in part due to the possible use of flapping propulsion for micro air vehicles. The importance of unsteady lift-enhancing mechanisms in insect flight has been recognized, but unsteady effects were generally thought to be absent for the flapping flight of larger animals. Only recently has the existence of LEVs (leading edge vortices) in small vertebrates such as swifts, small bats and hummingbirds been confirmed. To study the relevance of unsteady effects at the scale of large birds [reduced frequency k between 0.05 and 0.3, k=(pifc)/U(infinity); f is wingbeat frequency, U(infinity) is free-stream velocity, and c is the average wing chord], and the consequences of the lack of kinematic and morphological refinements, we have designed a simplified goose-sized flapping model for wind tunnel testing. The 2-D flow patterns along the wing span were quantitatively visualized using particle image velocimetry (PIV), and a three-component balance was used to measure the forces generated by the wings. The flow visualization on the wing showed the appearance of LEVs, which is typically associated with a delayed stall effect, and the transition into flow separation. Also, the influence of the delayed stall and flow separation was clearly visible in measurements of instantaneous net force over the wingbeat cycle. Here, we show that, even at reduced frequencies as low as those of large bird flight, unsteady effects are present and non-negligible and have to be addressed by kinematic and morphological adaptations.

Bats typically roost head-under-heels but they cannot hover in this position, thus, landing on a ceiling presents a biomechanical challenge. To land, a bat must perform an acrobatic flip that brings the claws of the toes in contact with the ceiling and do so gently enough as to avoid injury to its slender hindlimbs. In the present study, we sought to determine how bats land, to seek a link between landing kinematics and ceiling impact forces, and to determine whether landing strategies vary among bat species. To do this, we measured the kinematics and kinetics of landing behaviour in three species of bats as they landed on a force-measuring platform (Cynopterus brachyotis, N=3; Carollia perspicillata, N=5; Glossophaga soricina, N=5). Kinematics were similar for all bats within a species but differed among species. C. brachyotis performed four-point landings, during which body pitch increased until the ventral surface of the body faced the ceiling and the thumbs and hindlimbs simultaneously grasped the surface. Bats of the other two species performed two-point landings, whereby only the hindlimbs made contact with the ceiling. During these two-point landings, the hindlimbs were drawn up the side of the body to come in contact with the ceiling, causing simultaneous changes in body pitch, roll and yaw over the course of the landing sequence. Right-handed and left-handed forms of the two-point landing were observed, with individuals often switching back and forth between them among landing events. The four-point landing of C. brachyotis resulted in larger peak forces (3.7+/-2.4 body weights; median +/- interquartile range) than the two-point landings of C. perspicillata (0.8+/-0.6 body weights) or G. soricina (0.8+/-0.2 body weights). Our results demonstrate that the kinematics and kinetics of landing vary among bat species and that there is a correlation between the way a bat moves its body when it lands and the magnitude of peak impact force it experiences during that landing. We postulate that these interspecific differences in impact force could result because of stronger selective pressure for gentle landing in cave-roosting (C. perspicillata, G. soricina) versus foliage-roosting (C. brachyotis) species.