Turkey vultures tune their airspeed to changing air density.

Animals must tune their physical performance to changing environmental conditions, and the breadth of environmental tolerance may contribute to delineating the geographic range of a species. A common environmental challenge that flying animals face is the reduction of air density at high elevation and the reduction in the effectiveness of lift production that accompanies it. As a species, turkey vultures (Cathartes aura) inhabit a >3000 m elevation range, and fly considerably higher, necessitating that they accommodate for a 27% change in air density (0.890 to 1.227 kg m-3) through behavior, physiology or biomechanics. We predicted that birds flying at high elevation would maintain aerodynamic lift performance behaviorally via higher flight speeds, rather than increases in power output or local phenotypic adaptation. We used three-dimensional videography to track turkey vultures flying at three elevations, and data supported the hypothesized negative relationship between median airspeed and air density. Additionally, neither the ratio of horizontal speed to sinking speed nor flapping behavior varied with air density.

Olfactory sampling volume for pheromone capture by wing fanning of silkworm moth: a simulation-based study.

Odours used by insects for foraging and mating are carried by the air. Insects induce airflows around them by flapping their wings, and the distribution of these airflows may strongly influence odour source localisation. The flightless silkworm moth, Bombyx mori, has been a prominent insect model for olfactory research. However, although there have been numerous studies on antenna morphology and its fluid dynamics, neurophysiology, and localisation algorithms, the airflow manipulation of the B. mori by fanning has not been thoroughly investigated. In this study, we performed computational fluid dynamics (CFD) analyses of flapping B. mori to analyse this mechanism in depth. A three-dimensional simulation using reconstructed wing kinematics was used to investigate the effects of B. mori fanning on locomotion and pheromone capture. The fanning of the B. mori was found to generate an aerodynamic force on the scale of its weight through an aerodynamic mechanism similar to that of flying insects. Our simulations further indicate that the B. mori guides particles from its anterior direction within the ~ 60° horizontally by wing fanning. Hence, if it detects pheromones during fanning, the pheromone can be concluded to originate from the direction the head is pointing. The anisotropy in the sampling volume enables the B. mori to orient to the pheromone plume direction. These results provide new insights into insect behaviour and offer design guidelines for robots for odour source localisation.

Measurement of wing motion, deformation, and inertial forces of a biomimetic butterfly.

This paper introduces a method for measuring wing motion, deformation, and inertial forces in bio-inspired aircraft research using a camera motion capture system. The method involves placing markers on the wing surface and fitting rigid planes to determine the wing's spatial axis. This allows for describing the wing's rigid motion and obtaining deformation characteristics, such as deflection, twist angle, and gap distance of the forewing and hindwing. An image-based method is proposed for determining wing mass distribution, mass blocks, and mass points for inertial force measurement. The study addresses wing motion, deformation, and inertial force measurement in a real butterfly-like flapping wing vehicle and demonstrates the effectiveness of the approach. The results reveal that inertial forces play a negligible role in the generation of lift peaks and contribute minimal lift during the entire flapping cycle. Furthermore, a transitional phase between downstroke and upstroke is found in flexible wing motion, which has high lift production. This measurement approach offers a rapid and effective solution to experimental challenges in bio-inspired aircraft design and optimization.

Intraspecific variation in the pterosaur Rhamphorhynchus muensteri-implications for flight and socio-sexual signaling.

Pterosaurs were the first powered flying vertebrates, with a fossil record that stretches back to about 230 million years before present. Most species are only known from one to three specimens, which are most often fragmentary. However, Rhamphorhynchus muensteri is known from numerous excellent specimens, including multiple specimens with soft tissue preservation. As such, Rhamphorhynchus muensteri is one of the only pterosaurs amenable to analysis for intraspecific variation. It has been previously predicted that elements directly involved in the flight apparatus, such as those of the forelimb, will be more highly constrained in their proportions than other parts of the skeleton. We investigated the degree of variation seen in elements and body parts of Rhamphorhynchus, which represents the best model system among pterosaurs for testing these expectations of intraspecific variation. We recover evidence for high levels of constraint throughout the appendicular and axial elements (head, neck, torso, tail, forelimbs, hindlimbs), suggesting that all were important for flight. We further find that tail variation increases among the largest specimens, suggesting reduced constraint and/or stronger sexual selection on the tail in more mature individuals.

Generating controlled gust perturbations using vortex rings.

To understand the locomotory mechanisms of flying and swimming animals, it is often necessary to develop assays that enable us to measure their responses to external gust perturbations. Typically, such measurements have been carried out using a variety of gusts which are difficult to control or characterize owing to their inherently turbulent nature. Here, we present a method of generating discrete gusts under controlled laboratory conditions in the form of a vortex rings which are well-characterized and highly controllable. We also provide the theoretical guidelines underlying the design of gust generators for specific applications. As a case study, we tested the efficacy of this method to study the flight response of freely-flying soldier flies Hermetia illucens. The vortex ring based method can be used to generate controlled gusts to study diverse phenomena ranging from a natural flight in insects to the artificial flight of insect-sized drones and micro-aerial vehicles.

Exploration-exploitation model of moth-inspired olfactory navigation.

Navigation of male moths towards females during the mating search offers a unique perspective on the exploration-exploitation (EE) model in decision-making. This study uses the EE model to explain male moth pheromone-driven flight paths. Wind tunnel measurements and three-dimensional tracking using infrared cameras have been leveraged to gain insights into male moth behaviour. During the experiments in the wind tunnel, disturbance to the airflow has been added and the effect of increased fluctuations on moth flights has been analysed, in the context of the proposed EE model. The exploration and exploitation phases are separated using a genetic algorithm to the experimentally obtained dataset of moth three-dimensional trajectories. First, the exploration-to-exploitation rate (EER) increases with distance from the source of the female pheromone is demonstrated, which can be explained in the context of the EE model. Furthermore, our findings reveal a compelling relationship between EER and increased flow fluctuations near the pheromone source. Using an olfactory navigation simulation and our moth-inspired navigation model, the phenomenon where male moths exhibit an enhanced EER as turbulence levels increase is explained. This research extends our understanding of optimal navigation strategies based on general biological EE models and supports the development of bioinspired navigation algorithms.

Contrasting seasonal responses to wind in migrating songbirds on a globally important flyway.

During spring migration, nocturnal migrants attempt to minimize their travel time to reach their breeding grounds early. However, how they behave and respond to unfavourable conditions during their springtime travels is much less understood. In this study, we reveal the effects of atmospheric factors on nocturnal bird migration under adverse conditions during spring and autumn, based on one of the most detailed bird migration studies globally, using radar data from 13 deployments over a period of seven years (2014-2020) in the Levant region. Using ERA5 reanalysis data, we found that migratory birds maintain similar ground speeds in both autumn and spring migrations, but during spring, when encountering unfavourable winds, they put more effort into maintaining their travel speed by increasing self-powered airspeed by 18%. Moreover, we report for the first time that spring migrants showed less selectivity to wind conditions and migrated even under unfavourable headwind and crosswind conditions. Interestingly, we discovered that temperature was the most important weather parameter, such that warm weather substantially increased migration intensities in both seasons. Our results enhance our understanding of bird migration over the Levant region, one of the world's largest and most important migration flyways, and the factors controlling it. This information is essential for predicting bird migration, which-especially under the ongoing anthropogenic changes-is of high importance.

Wing deformation improves aerodynamic performance of forward flight of bluebottle flies flying in a flight mill.

Insect wings are flexible structures that exhibit deformations of complex spatiotemporal patterns. Existing studies on wing deformation underscore the indispensable role of wing deformation in enhancing aerodynamic performance. Here, we investigated forward flight in bluebottle flies, flying semi-freely in a magnetic flight mill; we quantified wing surface deformation using high-speed videography and marker-less surface reconstruction and studied the effects on aerodynamic forces, power and efficiency using computational fluid dynamics. The results showed that flies' wings exhibited substantial camber near the wing root and twisted along the wingspan, as they were coupled effects of deflection primarily about the claval flexion line. Such deflection was more substantial for supination during the upstroke when most thrust was produced. Compared with deformed wings, the undeformed wings generated 59-98% of thrust and 54-87% of thrust efficiency (i.e. ratio of thrust and power). Wing twist moved the aerodynamic centre of pressure proximally and posteriorly, likely improving aerodynamic efficiency.

Leading-edge curvature effect on aerodynamic performance of flapping wings in hover and forward flight.

This study investigates the role of leading-edge (LE) curvature in flapping wing aerodynamics considering hovering and forward flight conditions. A scaled-up robotic model is towed along its longitudinal axis by a rack gear carriage system. The forward velocity of the robotic model is changed by varying the advance ratioJfrom 0 (hovering) to 1.0. The study reveals that the LE curvature has insignificant influence on the cycle-average aerodynamic lift and drag. However, the time-history lift coefficient shows that the curvature can enhance the lift around the middle of downstroke. This enhanced lift is reduced from 5% to 1.2% asJchanged from 0 to 1.0. Further flow examinations reveal that the LE curvature is beneficial by enhancing circulation only at the outboard wing sections. The enhanced outboard circulation is found to emanate from the less stretched leading-edge vortices (LEVs), weakened trailing-edge vortices (TEVs), and the coherent merging of the tip vortices (TVs) with the minor LEVs as observed from the phase-lock planar digital particle image velocimetry measurements. The far-wake observation shows that the LE curvature enhances the vorticity within the TV, helping to reduce the overall flow fluctuations in the far field. These findings can be extended to explain the predominantly straight LE wing shape with a small amount of curvature only observed near the wing tip for flapping fliers with Re from 103to 104.

Age dominates flight distance and duration, while body size shapes flight speed in Bombus terrestris L. (Hymenoptera: Apidae).

Flight plays a crucial role in the fitness of insect pollinators, such as bumblebees. Despite their relatively large body size compared with their wings, bumblebees can fly under difficult ambient conditions, such as cooler temperatures. While their body size is often positively linked to their foraging range and flight ability, the influence of age remains less explored. Here, we studied the flight performance (distance, duration and speed) of ageing bumblebee workers using tethered flight mills. Additionally, we measured their intertegular distance and dry mass as proxies for their body size. We found that the flight distance and duration were predominantly influenced by age, challenging assumptions that age does not play a key role in foraging and task allocation. From the age of 7 to 14 days, flight distance and duration increased sixfold and fivefold, respectively. Conversely, the body size primarily impacted the maximum and average flight speed of workers. Our findings indicate that age substantially influences the flight distance and duration in bumblebee workers, affecting foraging performance and potentially altering task allocation strategies. This underscores the importance of considering individual age and physiological changes alongside body size/mass in experiments involving bumblebee workers.

Earliest evidence of avian primary feather moult.

Feather moulting is a crucial process in the avian life cycle, which evolved to maintain plumage functionality. However, moulting involves both energetic and functional costs. During moulting, plumage function temporarily decreases between the shedding of old feathers and the full growth of new ones. In flying taxa, a gradual and sequential replacement of flight feathers evolved to maintain aerodynamic capabilities during the moulting period. Little is known about the moult strategies of non-avian pennaraptoran dinosaurs and stem birds, before the emergence of crown lineage. Here, we report on two Early Cretaceous pygostylian birds from the Yixian Formation (125 mya), probably referable to Confuciusornithiformes, exhibiting morphological characteristics that suggest a gradual and sequential moult of wing flight feathers. Short primary feathers interpreted as immature are symmetrically present on both wings, as is typical among extant flying birds. Our survey of the enormous collection of the Tianyu Museum confirms previous findings that evidence of active moult in non-neornithine pennaraptorans is rare and likely indicates a moult cycle greater than one year. Documenting moult in Mesozoic feathered dinosaurs is critical for understanding their ecology, locomotor ability and the evolution of this important life-history process in birds.

A chemo-mechanical constitutive model for muscle activation in bat wing skins.

Birds, bats and insects have evolved unique wing structures to achieve a wide range of flight capabilities. Insects have relatively stiff and passive wings, birds have a complex and hierarchical feathered structure and bats have an articulated skeletal system integrated with a highly stretchable skin. The compliant skin of the wing distinguishes bats from all other flying animals and contributes to bats' remarkable, highly manoeuvrable flight performance and high energetic efficiency. The structural and functional complexity of the bat wing skin is one of the least understood although important elements of the bat flight anatomy. The wing skin has two unusual features: a discrete array of very soft elastin fibres and a discrete array of skeletal muscle fibres. The latter is intriguing because skeletal muscle is typically attached to bone, so the arrangement of intramembranous muscle in soft skin raises questions about its role in flight. In this paper, we develop a multi-scale chemo-mechanical constitutive model for bat wing skin. The chemo-mechanical model links cross-bridge cycling to a structure-based continuum model that describes the active viscoelastic behaviour of the soft anisotropic skin tissue. Continuum models at the tissue length-scale are valuable as they are easily implemented in commercial finite element codes to solve problems involving complex geometries, loading and boundary conditions. The constitutive model presented in this paper will be used in detailed finite element simulations to improve our understanding of the mechanics of bat flight in the context of wing kinematics and aerodynamic performance.

Visual lateralization in the sky: Geese manifest visual lateralization when flying with pair mates.

The brain's sensory lateralization involves the processing of information from the sensory organs primarily in one hemisphere. This can improve brain efficiency by reducing interference and duplication of neural circuits. For species that rely on successful interaction among family partners, such as geese, lateralization can be advantageous. However, at the group level, one-sided biases in sensory lateralization can make individuals predictable to competitors and predators. We investigated lateral preferences in the positioning of pair mates of Greater white-fronted geese Anser albifrons albifrons. Using GPS-GSM trackers, we monitored individual geese in flight throughout the year. Our findings indicate that geese exhibit individual lateral biases when viewing their mate in flight, but the direction of these biases varies among individuals. We suggest that these patterns of visual lateralization could be an adaptive trait for the species with long-term social monogamy, high levels of interspecies communication and competition, and high levels of predator and hunting pressure.

Gliding toward an understanding of the origin of flight in bats.

Bats are the only mammals capable of powered flight and have correspondingly specialized body plans, particularly in their limb morphology. The origin of bat flight is still not fully understood due to an uninformative fossil record but, from the perspective of a functional transition, it is widely hypothesized that bats evolved from gliding ancestors. Here, we test predictions of the gliding-to-flying hypothesis of the origin of bat flight by using phylogenetic comparative methods to model the evolution of forelimb and hindlimb traits on a dataset spanning four extinct bats and 231 extant mammals with diverse locomotor modes. Our results reveal that gliders exhibit adaptive trait optima (1) toward relatively elongate forelimbs that are intermediate between those of bats and non-gliding arborealists, and (2) toward relatively narrower but not longer hindlimbs that are intermediate between those of non-gliders and bats. We propose an adaptive landscape based on limb length and width optimal trends derived from our modeling analyses. Our results support a hypothetical evolutionary pathway wherein glider-like postcranial morphology precedes a bat-like morphology adapted to powered-flight, setting a foundation for future developmental, biomechanical, and evolutionary research to test this idea.

Insect ecology: Tracking migratory butterflies through pollen.

Painted ladies are well-known migratory butterflies, but confirmation of the details of their swarming flights through Europe has evaded scientists until now. It was their role as pollinators, carrying pollen grains on their flights, that helped unlock the secrets of their migrations.

How touch helps hummingbirds hover.

Georgina Mills discusses new research looking at how hummingbirds use somatosensation to adjust their flight.

Universal wing- and fin-beat frequency scaling.

We derive an equation that applies for the wing-beat frequency of flying animals and to the fin-stroke frequency of diving animals like penguins and whales. The equation states that the wing/fin-beat frequency is proportional to the square root of the animal's mass divided by the wing area. Data for birds, insects, bats, and even a robotic bird-supplemented by data for whales and penguins that must swim to stay submerged-show that the constant of proportionality is to a good approximation the same across all species; thus the equation is universal. The wing/fin-beat frequency equation is derived by dimensional analysis, which is a standard method of reasoning in physics. We finally demonstrate that a mathematically even simpler expression without the animal mass does not apply.

Perception of motion salience shapes the emergence of collective motions.

Despite the profound implications of self-organization in animal groups for collective behaviors, understanding the fundamental principles and applying them to swarm robotics remains incomplete. Here we propose a heuristic measure of perception of motion salience (MS) to quantify relative motion changes of neighbors from first-person view. Leveraging three large bird-flocking datasets, we explore how this perception of MS relates to the structure of leader-follower (LF) relations, and further perform an individual-level correlation analysis between past perception of MS and future change rate of velocity consensus. We observe prevalence of the positive correlations in real flocks, which demonstrates that individuals will accelerate the convergence of velocity with neighbors who have higher MS. This empirical finding motivates us to introduce the concept of adaptive MS-based (AMS) interaction in swarm model. Finally, we implement AMS in a swarm of ~102 miniature robots. Swarm experiments show the significant advantage of AMS in enhancing self-organization of the swarm for smooth evacuations from confined environments.

Asynchronous haltere input drives specific wing and head movements in Drosophila.

Halteres are multifunctional mechanosensory organs unique to the true flies (Diptera). A set of reduced hindwings, the halteres beat at the same frequency as the lift-generating forewings and sense inertial forces via mechanosensory campaniform sensilla. Though haltere ablation makes stable flight impossible, the specific role of wing-synchronous input has not been established. Using small iron filings attached to the halteres of tethered flies and an alternating electromagnetic field, we experimentally decoupled the wings and halteres of flying Drosophila and observed the resulting changes in wingbeat amplitude and head orientation. We find that asynchronous haltere input results in fast amplitude changes in the wing (hitches), but does not appreciably move the head. In multi-modal experiments, we find that wing and gaze optomotor responses are disrupted differently by asynchronous input. These effects of wing-asynchronous haltere input suggest that specific sensory information is necessary for maintaining wing amplitude stability and adaptive gaze control.