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Abstract Insect wings must be flexible, light, and strong to allow dynamic behaviors such as flying, mating, and feeding. When winged insects eclose into adults, their wings unfold, actuated hydraulically by hemolymph. Flowing hemolymph in the wing is necessary for functioning and healthy wings, both as the wing forms and as an adult. Because this process recruits the circulatory system, we asked, how much hemolymph is pumped into wings, and what happens to the hemolymph afterwards? Using Brood X cicadas ( Magicicada septendecim ), we collected 200 cicada nymphs, observing wing transformation over 2 h. Using dissection, weighing, and imaging of wings at set time intervals, we found that within 40 min after emergence, wing pads morphed into adult wings and total wing mass increased to ~ 16% of body mass. Thus, a significant amount of hemolymph is diverted from body to wings to effectuate expansion. After full expansion, in the ~ 80 min after, the mass of the wings decreased precipitously. In fact, the final adult wing is lighter than the initial folded wing pad, a surprising result. These results demonstrate that cicadas not only pump hemolymph into the wings, they then pump it out, producing a strong yet lightweight wing. 

Abstract Recent observations of wingless animals, including jumping nematodes, springtails, insects, and wingless vertebrates like geckos, snakes, and salamanders, have shown that their adaptations and body morphing are essential for rapid self-righting and controlled landing. These skills can reduce the risk of physical damage during collision, minimize recoil during landing, and allow for a quick escape response to minimize predation risk. The size, mass distribution, and speed of an animal determine its self-righting method, with larger animals depending on the conservation of angular momentum and smaller animals primarily using aerodynamic forces. Many animals falling through the air, from nematodes to salamanders, adopt a skydiving posture while descending. Similarly, plant seeds such as dandelions and samaras are able to turn upright in mid-air using aerodynamic forces and produce high decelerations. These aerial capabilities allow for a wide dispersal range, low-impact collisions, and effective landing and settling. Recently, small robots that can right themselves for controlled landings have been designed based on principles of aerial maneuvering in animals. Further research into the effects of unsteady flows on self-righting and landing in small arthropods, particularly those exhibiting explosive catapulting, could reveal how morphological features, flow dynamics, and physical mechanisms contribute to effective mid-air control. More broadly, studying apterygote (wingless insects) landing could also provide insight into the origin of insect flight. These research efforts have the potential to lead to the bio-inspired design of aerial micro-vehicles, sports projectiles, parachutes, and impulsive robots that can land upright in unsteady flow conditions. 

Although most binaural organisms locate sound sources using neurological structures to amplify the sounds they hear, some animals use mechanically coupled hearing organs instead. One of these animals, the parasitoid flyOrmia ochracea(O. ochracea), has astoundingly accurate sound localization abilities. It can locate objects in the azimuthal plane with a precision of 2°, equal to that of humans, despite an intertympanal distance of only 0.5 mm, which is less than$1/100$th of the wavelength of the sound emitted by the crickets that it parasitizes.O. ochraceaaccomplishes this feat via mechanically coupled tympana that interact with incoming acoustic pressure waves to amplify differences in the signals received at the two ears. In 1995, Mileset aldeveloped a model of hearing mechanics inO. ochraceathat represents the tympana as flat, front-facing prosternal membranes, though they lie on a convex surface at an angle from the flies’ frontal and transverse planes. The model works well for incoming sound angles less than$\pm {30}^{\circ }$but suffers from reduced accuracy (up to 60% error) at higher angles compared to response data acquired fromO. ochraceaspecimens. Despite this limitation, it has been the basis for bio-inspired microphone designs for decades. Here, we present critical improvements to this classic hearing model based on information from three-dimensional reconstructions ofO. ochracea’s tympanal organ. We identified the orientation of the tympana with respect to a frontal plane and the azimuthal angle segment between the tympana as morphological features essential to the flies’ auditory acuity, and hypothesized a differentiated mechanical response to incoming sound on the ipsi- and contralateral sides that depend on these features. We incorporated spatially-varying model coefficients representing this asymmetric response, making a new quasi-two-dimensional (q2D) model. The q2D model has high accuracy (average errors of under 10%) for all incoming sound angles. This improved biomechanical model may inform the design of new microscale directional microphones and other small-scale acoustic sensor systems.

 

This paper numerically studies the flow dynamics of aerial undulation of a snake-like model, which is adapted from the kinematics of the flying snake (Chrysopelea) undergoing a gliding process. The model applies aerial undulation periodically in a horizontal plane where a range of angle of attack (AOA) is assigned to model the real gliding motion. The flow is simulated using an immersed-boundary-method-based incompressible flow solver. Local mesh refinement mesh blocks are implemented to ensure the grid resolutions around the moving body. Results show that the undulating body produces the maximum lift at 45° of AOA. Vortex dynamics analysis has revealed a series of vortex structures including leading-edge vortices (LEV), trailing-edge vortices, and tip vortices around the body. Changes in other key parameters including the undulation frequency and Reynolds number are also found to affect the aerodynamics of the studied snake-like model, where increasing of undulation frequency enhances vortex steadiness and increasing of Reynolds number enhances lift production due to the strengthened LEVs. This study represents the first study of both the aerodynamics of the whole body of the snake as well as its undulatory motion, providing a new basis for investigating the mechanics of elongated flexible flyers.

 

An insect’s living systems—circulation, respiration, and a branching nervous system—extend from the body into the wing. Wing hemolymph circulation is critical for hydrating tissues and supplying nutrients to living systems such as sensory organs across the wing. Despite the critical role of hemolymph circulation in maintaining healthy wing function, wings are often considered “lifeless” cuticle, and flows remain largely unquantified. High-speed fluorescent microscopy and particle tracking of hemolymph in the wings and body of the grasshopperSchistocerca americanarevealed dynamic flow in every vein of the fore- and hindwings. The global system forms a circuit, but local flow behavior is complex, exhibiting three distinct types: pulsatile, aperiodic, and “leaky” flow. Thoracic wing hearts pull hemolymph from the wing at slower frequencies than the dorsal vessel; however, the velocity of returning hemolymph (in the hindwing) is faster than in that of the dorsal vessel. To characterize the wing’s internal flow mechanics, we mapped dimensionless flow parameters across the wings, revealing viscous flow regimes. Wings sustain ecologically important insect behaviors such as pollination and migration. Analysis of the wing circulatory system provides a template for future studies investigating the critical hemodynamics necessary to sustaining wing health and insect flight.

 

In snakes, the skin serves for protection, camouflage, visual signaling, locomotion, and its ability to stretch facilitates large prey ingestion. The flying snakes of the genusChrysopeleaare capable of jumping and gliding through the air, requiring additional functional demands: its skin must accommodate stretch in multiple directions during gliding and, perhaps more importantly, during high‐speed, direct‐impact landing. Is the skin of flying snakes specialized for gliding? Here, we characterized the material properties of the skin ofChrysopelea ornataand compared them with two nongliding species of colubrid snakes,Thamnophis sirtalisandPantherophis guttatus, as well as with previously published values. The skin was examined using uniaxial tensile testing to measure stresses, and digital image correlation methods to determine strains, yielding metrics of strength, elastic modulus, strain energy, and extensibility. To test for loading orientation effects, specimens were tested from three orientations relative to the snake's long axis: lateral, circumferential, and ventral. Specimens were taken from two regions of the body, pre‐ and pos‐tpyloric, to test for regional effects related to the ingestion of large prey. In comparison withT. sirtalisandP. guttatus,C. ornataexhibited higher post‐pyloric and lower pre‐pyloric extensibility in circumferential specimens. However, overall there were few differences in skin material properties ofC. ornatacompared to other species, both within and across studies, suggesting that the skin of flying snakes is not specialized for gliding locomotion. Surprisingly, circumferential specimens demonstrated lower strength and extensibility in pre‐pyloric skin, suggesting less regional specialization related to large prey.

 

ABSTRACT Flying snakes flatten their body to form a roughly triangular cross-sectional shape, enabling lift production and horizontal acceleration. While gliding, they also assume an S-shaped posture, which could promote aerodynamic interactions between the fore and the aft body. Such interactions have been studied experimentally; however, very coarse models of the snake's cross-sectional shape were used, and the effects were measured only for the downstream model. In this study, the aerodynamic interactions resulting from the snake's posture were approximated using two-dimensional anatomically accurate airfoils positioned in tandem to mimic the snake's geometry during flight. Load cells were used to measure the lift and drag forces, and flow field data were obtained using digital particle image velocimetry (DPIV). The results showed a strong dependence of the aerodynamic performance on the tandem arrangement, with the lift coefficients being generally more influenced than the drag coefficients. Flow field data revealed that the tandem arrangement modified the separated flow and the wake size, and enhanced the lift in cases in which the wake vortices formed closer to the models, producing suction on the dorsal surface. The downforce created by the flow separation from the ventral surface of the models at 0 deg angle of attack was another significant factor contributing to lift production. A number of cases showing large variations of aerodynamic performance included configurations close to the most probable posture of airborne flying snakes, suggesting that small postural variations could be used to control the glide trajectory. 

The insect circulatory system contains an open hemocoel, in which the mechanism of hemolymph flow control is ambiguous. As a continuous fluidic structure, this cavity should exhibit pressure changes that propagate quickly. Narrow-waisted insects create sustained pressure differences across segments, but their constricted waist provides an evident mechanism for compartmentalization. Insects with no obvious constrictions between segments may be capable of functionally compartmentalizing the body, which could explain complex hemolymph flows. Here, we test the hypothesis of functional compartmentalization by measuring pressures in a beetle and recording abdominal movements. We found that the pressure is indeed uniform within the abdomen and thorax, congruent with the predicted behavior of an open system. However, during some abdominal movements, pressures were on average 62% higher in the abdomen than in the thorax, suggesting that functional compartmentalization creates a gradient within the hemocoel. Synchrotron tomography and dissection show that the arthrodial membrane and thoracic muscles may contribute to this dynamic pressurization. Analysis of volume change suggests that the gut may play an important role in regulating pressure by translating between body segments. Overall, this study suggests that functional compartmentalization may provide an explanation for how fluid flows are managed in an open circulatory system.