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Animals use diverse solutions to land on vertical surfaces. Here we show the unique landing of the gliding gecko, Hemidactylus platyurus. Our high-speed video footage in the Southeast Asian rainforest capturing the first recorded, subcritical, short-range glides revealed that geckos did not markedly decrease velocity prior to impact. Unlike specialized gliders, geckos crashed head-first with the tree trunk at 6.0 ± 0.9 m/s (~140 body lengths per second) followed by an enormous pitchback of their head and torso 103 ± 34° away from the tree trunk anchored by only their hind limbs and tail. A dynamic mathematical model pointed to the utility of tails for the fall arresting response (FAR) upon landing. We tested predictions by measuring foot forces during landing of a soft, robotic physical model with an active tail reflex triggered by forefoot contact. As in wild animals, greater landing success was found for tailed robots. Experiments showed that longer tails with an active tail reflex resulted in the lower adhesive foot forces necessary for stabilizing successful landings, with a tail shortened to 25% requiring over twice the adhesive foot force., Siddall, Byrnes, Full, and Jusufi observe the function of the Asian flat-tailed gecko tail in gliding and landing on tree trunks in the field, complemented with mathematical and robotic models made of soft active materials. Altogether, their models show how geckos use an active tail reflex to brace the impact and reduce the risk of falling from landings on tree trunks.

Trade-offs in maneuverability and stability are essential in ecologically relevant situations with respect to robustness of locomotion, with multiple strategies apparent in animal model systems depending on their habitat and ecology. Free appendages such as tails and ungrounded limbs may assist in navigating this trade-off by assisting with balance, thereby increasing the acceleration that can be achieved without destabilizing the body. This comparative analysis explores the inertial mechanisms and, in some cases, fluid dynamic mechanisms by which appendages contribute to the stabilization of gait and perturbation response behaviors in a wide variety of animals. Following a broad review of examples from nature and bio-inspired robotics that illustrate the importance of appendages to the control of body orientation, two specific cases are examined through preliminary experiments: the role of arm motion in bipedal gait termination is explored using trajectory optimization, and the role of the cheetah’s tail during a deceleration maneuver is analyzed based on motion capture data. In both these examples, forward rotation of the appendage in question is found to counteract the unwanted forward pitch caused by the braking forces. It is theorized that this stabilizing action may facilitate more rapid deceleration by allowing larger or longer-acting braking forces to be applied safely.

Arboreal mammals navigate a highly three dimensional and discontinuous habitat. Among arboreal mammals, squirrels demonstrate impressive agility. In a recent “viral” YouTube video, unsuspecting squirrels were mechanically catapulted off of a track, inducing an initially uncontrolled rotation of the body. Interestingly, they skillfully stabilized themselves using tail motion, which ultimately allowed the squirrels to land successfully. Here we analyze the mechanism by which the squirrels recover from large body angular rates. We analyzed from the video that squirrels first use their tail to help stabilizing their head to visually fix a landing site. Then the tail starts to rotate to help stabilizing the body, preparing themselves for landing. To analyze further the mechanism of this tail use during mid-air, we built a multibody squirrel model and showed the righting strategy based on body inertia moment changes and active angular momentum transfer between axes. To validate the hypothesized strategy, we made a squirrel-like robot and demonstrated a fall-stabilizing experiment. Our results demonstrate that a squirrel’s long tail, despite comprising just 3% of body mass, can inertially stabilize a rapidly rotating body. This research contributes to better understanding the importance of long tails for righting mechanisms in animals living in complex environments such as trees.

Locomotion in unstructured and irregular environments is an enduring challenge in robotics. This is particularly true at the small scale, where relative obstacle size increases, often to the point ...

Due to the difficulty of manipulating muscle activation in live, freely swimming fish, a thorough examination of the body kinematics, propulsive performance, and muscle activity patterns in fish during undulatory swimming motion has not been conducted. We propose to use soft robotic model animals as experimental platforms to address biomechanics questions and acquire understanding into subcarangiform fish swimming behavior. We extend previous research on a bio-inspired soft robotic fish equipped with two pneumatic actuators and soft strain sensors to investigate swimming performance in undulation frequencies between 0.3 and 0.7 Hz and flow rates ranging from 0 to 20 cms in a recirculating flow tank. We demonstrate the potential of eutectic gallium–indium (eGaIn) sensors to measure the lateral deflection of a robotic fish in real time, a controller that is able to keep a constant undulatory amplitude in varying flow conditions, as well as using Particle Image Velocimetry (PIV) to characterizing swimming performance across a range of flow speeds and give a qualitative measurement of thrust force exerted by the physical platform without the need of externally attached force sensors. A detailed wake structure was then analyzed with Dynamic Mode Decomposition (DMD) to highlight different wave modes present in the robot’s swimming motion and provide insights into the efficiency of the robotic swimmer. In the future, we anticipate 3D-PIV with DMD serving as a global framework for comparing the performance of diverse bio-inspired swimming robots against a variety of swimming animals.

Arboreal animals face numerous challenges when negotiating complex three-dimensional terrain. Directed aerial descent or gliding flight allows for rapid traversal of arboreal environments, but presents control challenges. Some animals, such as birds or gliding squirrels, have specialized structures to modulate aerodynamic forces while airborne. However, many arboreal animals do not possess these specializations but still control posture and orientation in mid-air. One of the largest inertial segments in lizards is their tail. Inertial reorientation can be used to attain postures appropriate for controlled aerial descent. Here, we discuss the role of tail inertia in a range of mid-air reorientation behaviors using experimental data from geckos in combination with mathematical and robotic models. Geckos can self-right in mid-air by tail rotation alone. Equilibrium glide behavior of geckos in a vertical wind tunnel show that they can steer toward a visual stimulus by using rapid, circular tail rotations to control pitch and yaw. Multiple coordinated tail responses appear to be required for the most effective terminal velocity gliding. A mathematical model allows us to explore the relationship between morphology and the capacity for inertial reorientation by conducting sensitivity analyses, and testing control approaches. Robotic models further define the limits of performance and generate new control hypotheses. Such comparative analysis allows predictions about the diversity of performance across lizard morphologies, relative limb proportions, and provides insights into the evolution of aerial behaviors.

Synopsis Tails are a defining characteristic of chordates and show enormous diversity in function and shape. Although chordate tails share a common evolutionary and genetic-developmental origin, tails are extremely versatile in morphology and function. For example, tails can be short or long, thin or thick, and feathered or spiked, and they can be used for propulsion, communication, or balancing, and they mediate in predator–prey outcomes. Depending on the species of animal the tail is attached to, it can have extraordinarily multi-functional purposes. Despite its morphological diversity and broad functional roles, tails have not received similar scientific attention as, for example, the paired appendages such as legs or fins. This forward-looking review article is a first step toward interdisciplinary scientific synthesis in tail research. We discuss the importance of tail research in relation to five topics: (1) evolution and development, (2) regeneration, (3) functional morphology, (4) sensorimotor control, and (5) computational and physical models. Within each of these areas, we highlight areas of research and combinations of long-standing and new experimental approaches to move the field of tail research forward. To best advance a holistic understanding of tail evolution and function, it is imperative to embrace an interdisciplinary approach, re-integrating traditionally siloed fields around discussions on tail-related research.

We propose the use of bio-inspired robotics equipped with soft sensor technologies to gain a better understanding of the mechanics and control of animal movement. Soft robotic systems can be used to generate new hypotheses and uncover fundamental principles underlying animal locomotion and sensory capabilities, which could subsequently be validated using living organisms. Physical models increasingly include lateral body movements, notably back and tail bending, which are necessary for horizontal plane undulation in model systems ranging from fish to amphibians and reptiles. We present a comparative study of the use of physical modeling in conjunction with soft robotics and integrated soft and hyperelastic sensors to monitor local pressures, enabling local feedback control, and discuss issues related to understanding the mechanics and control of undulatory locomotion. A parallel approach combining live animal data with biorobotic physical modeling promises to be beneficial for gaining a better understanding of systems in motion.

Summary Acrobatic geckos can sprint at high speeds over challenging terrain [ 1 ], scamper up the smoothest surfaces [ 2 ], rapidly swing underneath leaves [ 3 ], and right themselves in midair by swinging only their tails [ 4 , 5 ]. From our field observations, we can add racing on the water’s surface to the gecko’s list of agile feats. Locomotion at the air-water interface evolved in over a thousand species, including insects, fish, reptiles, and mammals [ 6 ]. To support their weight, some larger-legged vertebrates use forces generated by vigorous slapping of the fluid’s surface followed by a stroke of their appendage [ 7 , 8 , 9 , 10 , 11 , 12 ], whereas smaller animals, like arthropods, rely on surface tension to walk on water [ 6 , 13 ]. Intermediate-sized geckos (Hemidactylus platyurus) fall squarely between these two regimes. Here, we report the unique ability of geckos to exceed the speed limits of conventional surface swimming. Several mechanisms likely contribute in this intermediate regime. In contrast to bipedal basilisk lizards [ 7 , 8 , 9 , 10 ], geckos used a stereotypic trotting gait with all four limbs, creating air cavities during slapping to raise their head and anterior trunk above water. Adding surfactant to the water decreased velocity by half, confirming surface tension’s role. The superhydrophobic skin could reduce drag during semi-planing. Geckos laterally undulated their bodies, including their submerged posterior trunk and tail, generating thrust for forward propulsion, much like water dragons [ 14 ] and alligators [ 15 ]. Geckos again remind us of the advantages of multi-functional morphologies providing the opportunity for multiple mechanisms for motion.

Undulatory motion of the body is the dominant mode of locomotion in fishes, and numerous studies of body kinematics and muscle activity patterns have provided insights into the mechanics of swimming. However, it has not been possible to investigate how key parameters such as the extent of bilateral muscle activation affect propulsive performance due to the inability to manipulate muscle activation in live, freely swimming fishes. In this article we extend previous work on passive flexible mechanical models of undulatory propulsion by using actively controlled pneumatic actuators attached to a flexible foil to gain insight into undulatory locomotion and mechanisms for body stiffness control. Two soft actuators were attached on each side of a flexible panel with stiffness comparable to that of a fish body. To study how bilateral contraction can be used to modify axial body stiffness during swimming, we ran a parameter sweep of actuator contraction phasing and frequency. Thrust production by the soft pneumatic actuators was tested at cyclic undulation frequencies ranging from 0.3 to 1.2 Hz in a recirculating flow tank at flow speeds up to 28 cm/s. Overall, this system generated more thrust at higher tail beat frequencies, with a plateau in thrust above 0.8 Hz. Self-propelled speed was found to be 0.8 foil lengths per second or ∼13 cm/s when actuated at 0.55 Hz. This active pneumatic model is capable of producing substantial trailing edge amplitudes with a maximum excursion equivalent to 1.4 foil lengths, and of generating considerable thrust. Altering the extent of bilateral co-contraction in a range from -22% to 17% of the cycle period showed that thrust was maximized with some amount of simultaneous left-right actuation of ∼3% to 6% of the cycle period. When the system is exposed to water flow, thrust was substantially reduced for conditions of greatest antagonistic overlap in left-right actuation, and also for the largest latencies introduced. This experimental platform provides a soft robotic testbed for studying aquatic propulsion with active control of undulatory kinematics.

Understanding the biomechanics of rapid running locomotion plays an important role in comparative biomechanics and bio-inspired engineering and is an integral part of animal welfare. However, this is not easily achieved using conventional methods of gait analysis: measuring ground reaction forces using a force plate, mainly on irregular granular terrain i.e. greyhounds in racing conditions or in animal’s natural habitats i.e. cheetahs in natural terrain. An alternative to measuring forces externally via force platforms embedded in track ways, we can attach inertial measurement units to agile quadrupeds to measure the effects of rapid running and turning. Here we deployed an IMU equipped with a tri-axial accelerometer on sprinting greyhounds to analyze rapid locomotion behaviors like dynamic banking and turning in conditions equivalent to racing. High speed videography and paw print analysis of the entire race were used for calibration. The results are beneficial in locomotion analysis and welfare of greyhounds.

Synopsis Animals that fall upside down typically engage in an aerial righting response so as to reorient dorsoventrally. This behavior can be preparatory to gliding or other controlled aerial behaviors and is ultimately necessary for a successful landing. Aerial righting reflexes have been described historically in various mammals such as cats, guinea pigs, rabbits, rats, and primates. The mechanisms whereby such righting can be accomplished depend on the size of the animal and on anatomical features associated with motion of the limbs and body. Here we apply a comparative approach to the study of aerial righting to explore the diverse strategies used for reorientation in midair. We discuss data for two species of lizards, the gecko Hemidactylus platyurus and the anole Anolis carolinensis, as well as for the first instar of the stick insect Extatosoma tiaratum, to illustrate size-dependence of this phenomenon and its relevance to subsequent aerial performance in parachuting and gliding animals. Geckos can use rotation of their large tails to reorient their bodies via conservation of angular momentum. Lizards with tails well exceeding snout-vent length, and correspondingly large tail inertia to body inertia ratios, are more effective at creating midair reorientation maneuvers. Moreover, experiments with stick insects, weighing an order of magnitude less than the lizards, suggest that aerodynamic torques acting on the limbs and body may play a dominant role in the righting process for small invertebrates. Both inertial and aerodynamic effects, therefore, can play a role in the control of aerial righting. We propose that aerial righting reflexes are widespread among arboreal vertebrates and arthropods and that they represent an important initial adaptation in the evolution of controlled aerial behavior.

Escaping from predators often demands that animals rapidly negotiate complex environments. The smallest animals attain relatively fast speeds with high frequency leg cycling, wing flapping or body undulations, but absolute speeds are slow compared to larger animals. Instead, small animals benefit from the advantages of enhanced maneuverability in part due to scaling. Here, we report a novel behavior in small, legged runners that may facilitate their escape by disappearance from predators. We video recorded cockroaches and geckos rapidly running up an incline toward a ledge, digitized their motion and created a simple model to generalize the behavior. Both species ran rapidly at 12–15 body lengths-per-second toward the ledge without braking, dove off the ledge, attached their feet by claws like a grappling hook, and used a pendulum-like motion that can exceed one meter-per-second to swing around to an inverted position under the ledge, out of sight. We discovered geckos in Southeast Asia can execute this escape behavior in the field. Quantification of these acrobatic behaviors provides biological inspiration toward the design of small, highly mobile search-and-rescue robots that can assist us during natural and human-made disasters. We report the first steps toward this new capability in a small, hexapedal robot.

In 1969, a palaeontologist proposed that theropod dinosaurs used their tails as dynamic stabilizers during rapid or irregular movements, contributing to their depiction as active and agile predators. Since then the inertia of swinging appendages has been implicated in stabilizing human walking, aiding acrobatic manoeuvres by primates and rodents, and enabling cats to balance on branches. Recent studies on geckos suggest that active tail stabilization occurs during climbing, righting and gliding. By contrast, studies on the effect of lizard tail loss show evidence of a decrease, an increase or no change in performance. Application of a control-theoretic framework could advance our general understanding of inertial appendage use in locomotion. Here we report that lizards control the swing of their tails in a measured manner to redirect angular momentum from their bodies to their tails, stabilizing body attitude in the sagittal plane. We video-recorded Red-Headed Agama lizards (Agama agama) leaping towards a vertical surface by first vaulting onto an obstacle with variable traction to induce a range of perturbations in body angular momentum. To examine a known controlled tail response, we built a lizard-sized robot with an active tail that used sensory feedback to stabilize pitch as it drove off a ramp. Our dynamics model revealed that a body swinging its tail experienced less rotation than a body with a rigid tail, a passively compliant tail or no tail. To compare a range of tails, we calculated tail effectiveness as the amount of tailless body rotation a tail could stabilize. A model Velociraptor mongoliensis supported the initial tail stabilization hypothesis, showing as it did a greater tail effectiveness than the Agama lizards. Leaping lizards show that inertial control of body attitude can advance our understanding of appendage evolution and provide biological inspiration for the next generation of manoeuvrable search-and-rescue robots.

Geckos are nature's elite climbers. Their remarkable climbing feats have been attributed to specialized feet with hairy toes that uncurl and peel in milliseconds. Here, we report that the secret to the gecko's arboreal acrobatics includes an active tail. We examine the tail's role during rapid climbing, aerial descent, and gliding. We show that a gecko's tail functions as an emergency fifth leg to prevent falling during rapid climbing. A response initiated by slipping causes the tail tip to push against the vertical surface, thereby preventing pitch-back of the head and upper body. When pitch-back cannot be prevented, geckos avoid falling by placing their tail in a posture similar to a bicycle's kickstand. Should a gecko fall with its back to the ground, a swing of its tail induces the most rapid, zero-angular momentum air-righting response yet measured. Once righted to a sprawled gliding posture, circular tail movements control yaw and pitch as the gecko descends. Our results suggest that large, active tails can function as effective control appendages. These results have provided biological inspiration for the design of an active tail on a climbing robot, and we anticipate their use in small, unmanned gliding vehicles and multisegment spacecraft.

Unlike the falling cat, lizards can right themselves in mid-air by a swing of their large tails in one direction causing the body to rotate in the other. Here, we developed a new three-dimensional analytical model to investigate the effectiveness of tails as inertial appendages that change body orientation. We anchored our model using the morphological parameters of the flat-tailed house gecko Hemidactylus platyurus. The degree of roll in air righting and the amount of yaw in mid-air turning directly measured in house geckos matched the model’s results. Our model predicted an increase in body roll and turning as tails increase in length relative to the body. Tails that swung from a near orthogonal plane relative to the body (i.e. 0‐30 ◦ from vertical) were the most effective at generating body roll, whereas tails operating at steeper angles (i.e. 45‐60 ◦ ) produced only half the rotation. To further test our analytical model’s predictions, we built a bio-inspired robot prototype. The robot reinforced how effective attitude control can be attained with simple movements of an inertial appendage.