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SUMMARYWe 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. Supplementary material available online at

modes of bats vary widely among families, so we focused on a single family, the Pteropodidae. This family consists of ca. 186 species distributed throughout the paleotropics (Wilson and Reeder, 2005) and is characterized by fruit and nectar-feeding, non-echolocating Accepted 26 None of the bats in our study flew at constant speed, so we used multiple regression to isolate the changes in wing kinematics that correlated with changes in flight speed, horizontal acceleration and vertical acceleration. We uncovered several significant trends that were consistent among species. Our results demonstrate that for medium-to large-sized bats, the ways that bats modulate their wing kinematics to produce thrust and lift over the course of a wingbeat cycle are independent of body size. Supplementary material available online at

SUMMARY Bats (Chiroptera) are generally awkward crawlers, but the common vampire bat (Desmodus rotundus) and the New Zealand short-tailed bat(Mystacina tuberculata) have independently evolved the ability to manoeuvre well on the ground. In this study we describe the kinematics of locomotion in both species, and the kinetics of locomotion in M. tuberculata. We sought to determine whether these bats move terrestrially the way other quadrupeds do, or whether they possess altogether different patterns of movement on the ground than are observed in quadrupeds that do not fly. Using high-speed video analyses of bats moving on a treadmill, we observed that both species possess symmetrical lateral-sequence gaits similar to the kinematically defined walks of a broad range of tetrapods. At high speeds, D. rotundus use an asymmetrical bounding gait that appears to converge on the bounding gaits of small terrestrial mammals, but with the roles of the forelimbs and hindlimbs reversed. This gait was not performed by M. tuberculata. Many animals that possess a single kinematic gait shift with increasing speed from a kinetic walk (where kinetic and potential energy of the centre of mass oscillate out of phase from each other) to a kinetic run (where they oscillate in phase). To determine whether the single kinematic gait of M. tuberculata meets the kinetic definition of a walk, a run, or a gait that functions as a walk at low speed and a run at high speed, we used force plates and high-speed video recordings to characterize the energetics of the centre of mass in that species. Although oscillations in kinetic and potential energy were of similar magnitudes, M. tuberculata did not use pendulum-like exchanges of energy between them to the extent that many other quadrupedal animals do, and did not transition from a kinetic walk to kinetic run with increasing speed. The gait of M. tuberculata is kinematically a walk,but kinetically run-like at all speeds.

SUMMARYThe center of mass (COM) of a flying animal accelerates through space because of aerodynamic and gravitational forces. For vertebrates, changes in the position of a landmark on the body have been widely used to estimate net aerodynamic forces. The flapping of relatively massive wings, however, might induce inertial forces that cause markers on the body to move independently of the COM, thus making them unreliable indicators of aerodynamic force. We used high-speed three-dimensional kinematics from wind tunnel flights of four lesser dog-faced fruit bats, Cynopterus brachyotis, at speeds ranging from 2.4 to 7.8ms -1 to construct a time-varying model of the mass distribution of the bats and to estimate changes in the position of their COM through time. We compared accelerations calculated by markers on the trunk with accelerations calculated from the estimated COM and we found significant inertial effects on both horizontal and vertical accelerations. We discuss the effect of these inertial accelerations on the long-held idea that, during slow flights, bats accelerate their COM forward during 'tip-reversal upstrokes', whereby the distal portion of the wing moves upward and backward with respect to still air. This idea has been supported by the observation that markers placed on the body accelerate forward during tip-reversal upstrokes. As in previously published studies, we observed that markers on the trunk accelerated forward during the tip-reversal upstrokes. When removing inertial effects, however, we found that the COM accelerated forward primarily during the downstroke. These results highlight the crucial importance of the incorporation of inertial effects of wing motion in the analysis of flapping flight. Supplementary material available online at

Flying vertebrates change the shapes of their wings during the upstroke, thereby decreasing wing surface area and bringing the wings closer to the body than during downstroke. These, and other wing deformations, might reduce the inertial cost of the upstroke compared with what it would be if the wings remained fully extended. However, wing deformations themselves entail energetic costs that could exceed any inertial energy savings. Using a model that incorporates detailed three-dimensional wing kinematics, we estimated the inertial cost of flapping flight for six bat species spanning a 40-fold range of body masses. We estimate that folding and unfolding comprises roughly 44 per cent of the inertial cost, but that the total inertial cost is only approximately 65 per cent of what it would be if the wing remained extended and rigid throughout the wingbeat cycle. Folding and unfolding occurred mostly during the upstroke; hence, our model suggests inertial cost of the upstroke is not less than that of downstroke. The cost of accelerating the metacarpals and phalanges accounted for around 44 per cent of inertial costs, although those elements constitute only 12 per cent of wing weight. This highlights the energetic benefit afforded to bats by the decreased mineralization of the distal wing bones.

Gliding is an efficient form of travel found in every major group of terrestrial vertebrates. Gliding is often modelled in equilibrium, where aerodynamic forces exactly balance body weight resulting in constant velocity. Although the equilibrium model is relevant for long-distance gliding, such as soaring by birds, it may not be realistic for shorter distances between trees. To understand the aerodynamics of inter-tree gliding, we used direct observation and mathematical modelling. We used videography (60 -125 fps) to track and reconstruct the three-dimensional trajectories of northern flying squirrels (Glaucomys sabrinus) in nature. From their trajectories, we calculated velocities, aerodynamic forces and force coefficients. We determined that flying squirrels do not glide at equilibrium, and instead demonstrate continuously changing velocities, forces and force coefficients, and generate more lift than needed to balance body weight. We compared observed glide performance with mathematical simulations that use constant force coefficients, a characteristic of equilibrium glides. Simulations with varying force coefficients, such as those of live squirrels, demonstrated better wholeglide performance compared with the theoretical equilibrium state. Using results from both the observed glides and the simulation, we describe the mechanics and execution of inter-tree glides, and then discuss how gliding behaviour may relate to the evolution of flapping flight.

SUMMARYBats 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 forcemeasuring 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 twopoint 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.