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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., Competing Interests: The authors declare that they have no competing interests., (© 2024 Burtner et al.)

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.

Body size and shape fundamentally determine organismal energy requirements by modulating heat and mass exchange with the environment and the costs of locomotion, thermoregulation, and maintenance. Ecologists have long used the physical linkage between morphology and energy balance to explain why the body size and shape of many organisms vary across climatic gradients, e.g., why larger endotherms are more common in colder regions. However, few modeling exercises have aimed at investigating this link from first principles. Body size evolution in bats contrasts with the patterns observed in other endotherms, probably because physical constraints on flight limit morphological adaptations. Here, we develop a biophysical model based on heat transfer and aerodynamic principles to investigate energy constraints on morphological evolution in bats. Our biophysical model predicts that the energy costs of thermoregulation and flight, respectively, impose upper and lower limits on the relationship of wing surface area to body mass (S-MR), giving rise to an optimal S-MR at which both energy costs are minimized. A comparative analysis of 278 species of bats supports the model’s prediction that S-MR evolves toward an optimal shape and that the strength of selection is higher among species experiencing greater energy demands for thermoregulation in cold climates. Our study suggests that energy costs modulate the mode of morphological evolution in bats—hence shedding light on a long-standing debate over bats’ conformity to ecogeographical patterns observed in other mammals—and offers a procedure for investigating complex macroecological patterns from first principles.

Bats possess wings comprised of a flexible membrane and a jointed skeletal structure allowing them to execute complex flight maneuvers such as rapid tight turns. The extent of a bat's tight turning capability can be explored by analyzing a 180-degree U-turn. Prior studies have investigated more subtle flight maneuvers, but the kinematic and aerodynamic mechanisms of a U-turn have not been characterized. In this work, we use 3D optical motion capture and aerodynamic simulations to investigate a U-turn maneuver executed by a great roundleaf bat (Hipposideros armiger: mass = 55 g, span = 51 cm). The bat was observed to decrease its flight velocity and gain approximately 20 cm of altitude entering the U-turn. By lowering its velocity from 2.0 m/s to 0.5 m/s, the centripetal force requirement to execute a tight turn was substantially reduced. Centripetal force was generated by tilting the lift force vector laterally through banking. During the initiation of the U-turn, the bank angle increased from 20 degrees to 40 degrees. During the initiation and persisting throughout the U-turn, the flap amplitude of the right wing (inside of the turn) increased relative to the left wing. In addition, the right wing moved more laterally closer to the centerline of the body during the end of the downstroke and the beginning of the upstroke compared to the left wing. Reorientation of the body into the turn happened prior to a change in the flight path of the bat. Once the bat entered the U-turn and the bank angle increased, the change in flight path of the bat began to change rapidly as the bat negotiated the apex of the turn. During this phase of the turn, the minimum radius of curvature of the bat was 5.5 cm. During the egress of the turn, the bat accelerated and expended stored potential energy by descending. The cycle averaged total power expenditure by the bat during the six wingbeat cycle U-turn maneuver was 0.51 W which was approximately 40% above the power expenditure calculated for a nominally straight flight by the same bat. Future work on the topic of bat maneuverability may investigate a broader array of maneuvering flights characterizing the commonalities and differences across flights. In addition, the interplay between aerodynamic moments and inertial moments are of interest in order to more robustly characterize maneuvering mechanisms., Competing Interests: The authors have declared that no competing interests exist.

The synchronization of flight mechanics with respiration and echolocation call emission by bats, while economizing these behaviors, presumably puts compressive loads on the cartilaginous rings that hold open the respiratory tract. Previous work has shown that during postnatal development of Artibeus jamaicensis (Phyllostomidae), the onset of adult echolocation call emission rate coincides with calcification of the larynx, and the development of flight coincides with tracheal ring calcification. In the present study, I assessed the level of reinforcement of the respiratory system in 13 bat species representing six families that use stereotypical modes of echolocation (i.e. duty cycle % and intensity). Using computed tomography, the degree of mineralization or ossification of the tracheal rings, cricoid, thyroid and arytenoid cartilages were determined for non-echolocators, tongue clicking, low-duty cycle low-intensity, low-duty cycle high-intensity, and high-duty cycle high-intensity echolocating bats. While all bats had evidence of cervical tracheal ring mineralization, about half the species had evidence of thoracic tracheal ring calcification. Larger bats (Phyllostomus hastatus and Pterpodidae sp.) exhibited more extensive tracheal ring mineralization, suggesting an underlying cause independent of laryngeal echolocation. Within most of the laryngeally echolocating species, the degree of mineralization or ossification of the larynx was dependent on the mode of echolocation system used. Low-duty cycle low-intensity bats had extensively mineralized cricoids, and zero to very minor mineralization of the thyroids and arytenoids. Low-duty cycle high-intensity bats had extensively mineralized cricoids, and patches of thyroid and arytenoid mineralization. The high-duty cycle high-intensity rhinolophids and hipposiderid had extensively ossified cricoids, large patches of ossification on the thyroids, and heavily ossified arytenoids. The high-duty cycle high-intensity echolocator, Pteronotus parnellii, had mineralization patterns and laryngeal morphology very similar to the other low-duty cycle high-intensity mormoopid species, perhaps suggesting relatively recent evolution of high-duty cycle echolocation in P. parnellii compared with the Old World high-duty cycle echolocators (Rhinolophidae and Hipposideridae). All laryngeal echolocators exhibited mineralized or ossified lateral expansions of the cricoid for articulation with the inferior horn of the thyroid, these were most prominent in the high-duty cycle high-intensity rhinolophids and hipposiderid, and least prominent in the low-duty cycle low-intensity echolocators. The non-laryngeal echolocators had extensively ossified cricoid and thyroid cartilages, and no evidence of mineralization/ossification of the arytenoids or lateral expansions of the cricoid. While the non-echolocators had extensive ossification of the larynx, it was inconsistent with that seen in the laryngeal echolocators., (© 2020 Anatomical Society.)

Many animals display morphological adaptations of the nose that improve their ability to detect and track odors. Bilateral odor sampling improves an animals' ability to navigate using olfaction and increased separation of the nostrils facilitates olfactory source localization. Many bats use odors to find food and mates and bats display an elaborate diversity of facial features. Prior studies have quantified how variations in facial features correlate with echolocation and feeding ecology, but surprisingly none have asked whether bat noses might be adapted for olfactory tracking in flight. We predicted that bat species that rely upon odor cues while foraging would have greater nostril separation in support of olfactory tropotaxis. Using museum specimens, we measured the external nose and cranial morphology of 40 New World bat species. Diet had a significant effect on external nose morphology, but contrary to our predictions, insectivorous bats had the largest relative separation of nostrils, while nectar feeding species had the narrowest nostril widths. Furthermore, nasal echolocating bats had significantly narrower nostrils than oral emitting bats, reflecting a potential trade-off between sonar pulse emission and stereo-olfaction in those species. To our knowledge, this is the first study to evaluate the evolutionary interactions between olfaction and echolocation in shaping the external morphology of a facial feature using modern phylogenetic comparative methods. Future work pairing olfactory morphology with tracking behavior will provide more insight into how animals such as bats integrate olfactory information while foraging., Competing Interests: The authors have declared that no competing interests exist.

Neomorphic, membrane-associated skeletal rods are found in disparate vertebrate lineages, but their evolution is poorly understood. Here we show that one of these elements-the calcar of bats (Chiroptera)-is a skeletal novelty that has anatomically diversified. Comparisons of evolutionary models of calcar length and corresponding disparity-through-time analyses indicate that the calcar diversified early in the evolutionary history of Chiroptera, as bats phylogenetically diversified after evolving the capacity for flight. This interspecific variation in calcar length and its relative proportion to tibia and forearm length is of functional relevance to flight-related behaviors. We also find that the calcar varies in its tissue composition among bats, which might affect its response to mechanical loading. We confirm the presence of a synovial joint at the articulation between the calcar and the calcaneus in some species, which suggests the calcar has a kinematic functional role. Collectively, this functionally relevant variation suggests that adaptive advantages provided by the calcar led to its anatomical diversification. Our results demonstrate that novel skeletal additions can become integrated into vertebrate body plans and subsequently evolve into a variety of forms, potentially impacting clade diversification by expanding the available morphological space into which organisms can evolve., (© 2019 The Author(s). Evolution © 2019 The Society for the Study of Evolution.)

Bats, with highly articulated wings, are some of the most agile flyers in nature. A novel three-dimensional geometric decomposition framework is developed to reduce the complex kinematics of a bat wing into physical movements used to describe flapping flight: namely flapping, stroke plane deviation and pitching, together with cambering and flexion. The decomposition is combined with aerodynamic simulations to investigate the cumulative effect of each motion on force production, and their primary contribution to the unsteady vortex dynamics. For the nearly straight and level flight of Hipposideros pratti, results show that the flapping motion by itself induced a moderate drag and lift. Stroke plane deviation increased lift, and nullified the drag. With the inclusion of the pitching motion into the kinematics, lift production further increased by a factor of more than 2.5, and exhibited a positive net thrust by virtue of the favorable wing orientation during the upstroke. The primary contribution of cambering, which included a maximum chord line displacement of ≈40% standard mean chord, was the stabilization of the leading edge vortex during the downstroke. This increased mean lift by about 35% at the expense of net thrust. Flexion was perhaps the most complex motion with maximum displacements of 75% standard mean chord. This was instrumental in reducing the negative lift during the upstroke by preventing the formation of strong leading edge vortices. The aerodynamic effective angle of attack emerged as a heuristic parameter to describe lift and net thrust production across movements., Competing Interests: The authors have declared that no competing interests exist.

The use of echolocation allows insectivorous bats to access unique foraging niches by locating obstacles and prey with ultrasounds in complete darkness. To avoid interspecific competition, it is likely that sonar features and wing morphology co-evolved with species vertical distribution, but due to the technical difficulties of studying flight in the vertical dimension, this has never been demonstrated with empirical measurements. The authors equipped 48 wind masts with arrays of two microphones and located the vertical distribution of a community of 19 bat species and two species groups over their annual activity period (>8000 nights). The authors tested the correlation between the proportion of flights at height and the acoustic features of bat calls as well as their wing morphology. The authors found that call peak frequency and bandwidth are good predictors of bat use of the vertical space regardless of their acoustic strategies (i.e., gleaning, hawking, or detecting prey flutter). High wing aspect ratios and high wing loadings were associated with high proportions of time spent at height, confirming hypotheses from the literature.

The striking postcranial anatomy of bats reflects their specialized ecology; they are the only mammals capable of powered flight. Bat postcranial adaptations include a series of membranes that connect highly-modified, or even novel, skeletal elements. While most studies of bat postcranial anatomy have focused on their wings, bat hindlimbs also contain many derived and functionally important, yet less studied, features. In this study, we investigate variation in the membrane and limb musculature associated with the calcar, a neomorphic skeletal structure found in the hindlimbs of most bats. We use diffusible iodine-based contrast-enhanced computed tomography and standard histological techniques to examine the calcars and hindlimb membranes of three bat species that vary ecologically (Myotis californicus, a slow-flying insectivore; Molossus molossus, a fast-flying insectivore; and Artibeus jamaicensis, a slow-flying frugivore). We also assess the level of mineralization of the calcar at muscle attachment sites to better understand how muscle contraction may enable calcar function. We found that the arrangement of the calcar musculature varies among the three bat species, as does the pattern of mineral content within the calcar. M. molossus and M. californicus exhibit more complex calcar and calcar musculature morphologies than A. jamaicensis, and the degree of calcar mineralization decreases toward the tip of the calcar in all species. These results are consistent with the idea that the calcar may have a functional role in flight maneuverability. Anat Rec, 301:441-448, 2018. © 2018 Wiley Periodicals, Inc., (© 2018 Wiley Periodicals, Inc.)

All echolocating bats and whales measured to date emit a directional bio-sonar beam that affords them a number of advantages over an omni-directional beam, i.e. reduced clutter, increased source level and inherent directional information. In this study, we investigated the importance of directional sound emission for navigation through echolocation by measuring the sonar beam of brown long-eared bats, Plecotus auritus Plecotus auritus emits sound through the nostrils but has no external appendages to readily facilitate a directional sound emission as found in most nose emitters. The study shows that P. auritus , despite lacking an external focusing apparatus, emits a directional echolocation beam (directivity index=13 dB) and that the beam is more directional vertically (-6 dB angle at 22 deg) than horizontally (-6 dB angle at 35 deg). Using a simple numerical model, we found that the recorded emission pattern is achievable if P. auritus emits sound through the nostrils as well as the mouth. The study thus supports the hypothesis that a directional echolocation beam is important for perception through echolocation and we propose that animals with similarly non-directional emitter characteristics may facilitate a directional sound emission by emitting sound through both the nostrils and the mouth., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Bats navigate the dark using echolocation. Echolocation is enhanced by external ears, but external ears increase the projected frontal area and reduce the streamlining of the animal. External ears are thus expected to compromise flight efficiency, but research suggests that very large ears may mitigate the cost by producing aerodynamic lift. Here we compare quantitative aerodynamic measures of flight efficiency of two bat species, one large-eared ( Plecotus auritus ) and one small-eared ( Glossophaga soricina ), flying freely in a wind tunnel. We find that the body drag of both species is higher than previously assumed and that the large-eared species has a higher body drag coefficient, but also produces relatively more ear/body lift than the small-eared species, in line with prior studies on model bats. The measured aerodynamic power of P. auritus was higher than predicted from the aerodynamic model, while the small-eared species aligned with predictions. The relatively higher power of the large-eared species results in lower optimal flight speeds and our findings support the notion of a trade-off between the acoustic benefits of large external ears and aerodynamic performance. The result of this trade-off would be the eco-morphological correlation in bat flight, with large-eared bats generally adopting slow-flight feeding strategies., (© 2017 The Author(s).)

Harnessing flight strategies refined by millions of years of evolution can help expedite the design of more efficient, manoeuvrable and robust flying robots. This review synthesizes recent advances and highlights remaining gaps in our understanding of how bird and bat wing adaptations enable effective flight. Included in this discussion is an evaluation of how current robotic analogues measure up to their biological sources of inspiration. Studies of vertebrate wings have revealed skeletal systems well suited for enduring the loads required during flight, but the mechanisms that drive coordinated motions between bones and connected integuments remain ill-described. Similarly, vertebrate flight muscles have adapted to sustain increased wing loading, but a lack of in vivo studies limits our understanding of specific muscular functions. Forelimb adaptations diverge at the integument level, but both bird feathers and bat membranes yield aerodynamic surfaces with a level of robustness unparalleled by engineered wings. These morphological adaptations enable a diverse range of kinematics tuned for different flight speeds and manoeuvres. By integrating vertebrate flight specializations-particularly those that enable greater robustness and adaptability-into the design and control of robotic wings, engineers can begin narrowing the wide margin that currently exists between flying robots and vertebrates. In turn, these robotic wings can help biologists create experiments that would be impossible in vivo ., (© 2017 The Author(s).)

Within the large order of bats, sexual size dimorphism measured by forearm length and body mass is often female-biased. Several studies have explained this through the effects on load carrying during pregnancy, intrasexual competition, as well as the fecundity and thermoregulation advantages of increased female body size. We hypothesized that wing shape should differ along with size and be under variable selection pressure in a species where there are large differences in flight behaviour. We tested whether load carrying, sex differential migration, or reproductive advantages of large females affect size and wing shape dimorphism in the common noctule (Nyctalus noctula), in which females are typically larger than males and only females migrate long distances each year. We tested for univariate and multivariate size and shape dimorphism using data sets derived from wing photos and biometric data collected during pre-migratory spring captures in Switzerland. Females had forearms that are on average 1% longer than males and are 1% heavier than males after emerging from hibernation, but we found no sex differences in other size, shape, or other functional characters in any wing parameters during this pre-migratory period. Female-biased size dimorphism without wing shape differences indicates that reproductive advantages of big mothers are most likely responsible for sexual dimorphism in this species, not load compensation or shape differences favouring aerodynamic efficiency during pregnancy or migration. Despite large behavioural and ecological sex differences, morphology associated with a specialized feeding niche may limit potential dimorphism in narrow-winged bats such as common noctules and the dramatic differences in migratory behaviour may then be accomplished through plasticity in wing kinematics., 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).)

Bat wings, like other mammalian forelimbs, contain many joints within the digits. These joints collectively affect dynamic three-dimensional (3D) wing shape, thereby affecting the amount of aerodynamic force a wing can generate. Bats are a speciose group, and show substantial variation in the number of wing joints. Additionally, some bat species have joints with extensor but no flexor muscles. While several studies have examined the diversity in number of joints and presence of muscles, musculoskeletal variation in the digits has not been interpreted in phylogenetic, functional or ecological contexts. To provide this context, the number of joints and the presence/absence of muscles are quantified for 44 bat species, and are mapped phylogenetically. It is shown that, relative to the ancestral state, joints and muscles were lost multiple times from different digits and in many lineages. It is also shown that joints lacking flexors undergo cyclical flexion and extension, in a manner similar to that observed in joints with both flexors and extensors. Comparison of species with contrasting feeding ecologies demonstrates that species that feed primarily on non-mobile food (e.g. fruit) have fewer fully active joints than species that catch mobile prey (e.g. insects). It is hypothesized that there is a functional trade-off between energetic savings and maneuverability. Having fewer joints and muscles reduces the mass of the wing, thereby reducing the energetic requirements of flapping flight, and having more joints increases the assortment of possible 3D wing shapes, thereby enhancing the range and fine control of aerodynamic force production and thus maneuverability., (© 2016 Anatomical Society.)

The inertial power and inertial force of wings are important factors in evaluating the flight performance of native bats. Based on measurement data of wing size and motions of Eptesicus fuscus, we present a new computational bat wing model with divided fragments of skeletons and membrane. The motions of the model were verified by comparing the joint and tip trajectories with native bats. The influences of flap, sweep, elbow, wrist and digits motions, the effects of different bones and membrane of bat wing, the components on vertical, spanwise and fore-aft directions of the inertial power and force were analyzed. Our results indicate that the flap, sweep, and elbow motions contribute the main inertial power and force; the membrane occupies an important proportion of the inertial power and force; inertial power on flap direction was larger, while variations of inertial forces on different directions were not evident. These methods and results offer insights into flight dynamics in other flying animals and may contribute to the design of future robotic bats.

Adequate postnatal growth is important for young bats to develop skilled sensory and locomotor abilities, which are highly associated with their survival once independent. This study investigated the postnatal growth and development of Scotophilus kuhlii in captivity. An empirical growth curve was established, and the postnatal growth rate was quantified to derive an age-predictive equation. By further controlling the fostering conditions of twins, the differences in the development patterns between pups that received maternal care or were hand-reared were analyzed to determine whether the latter developed in the same manner as their maternally reared counterparts. Our results indicate that both forearm length and body mass increased rapidly and linearly during the first 4 weeks, after which the growth rate gradually decreased to reach a stable level. The first flight occurred at an average age of 39 days with a mean forearm length and body mass of 92.07% and 70.52% of maternal size, respectively. The developmental pattern of hand-reared pups, although similar to that of their maternally reared twin siblings, displayed a slightly faster growth rate in the 4th and 5th weeks. The heavier body mass of hand-reared pups during the pre-fledging period may cause higher wing loading, potentially influencing the flight performance and survival of the bats once independent., (© 2015 Wiley Periodicals, Inc.)

This paper elucidates the aerodynamic role of the dynamically changing wingspan in bat flight. Based on direct numerical simulations of the flow over a slow-flying bat, it is found that the dynamically changing wingspan can significantly enhance the lift. Further, an analysis of flow structures and lift decomposition reveal that the elevated vortex lift associated with the leading-edge vortices intensified by the dynamically changing wingspan considerably contributed to enhancement of the time-averaged lift. The nonlinear interaction between the dynamically changing wing and the vortical structures plays an important role in the lift enhancement of a flying bat in addition to the geometrical effect of changing the lifting-surface area in a flapping cycle. In addition, the dynamically changing wingspan leads to the higher efficiency in terms of generating lift for a given amount of the mechanical energy consumed in flight., (© 2015 The Author(s).)

Recent evidence has shown that the developmental emergence of echolocation calls in young bats follow an independent developmental pathway from other vocalizations and that adult-like echolocation call structure significantly precedes flight ability. These data in combination with new insights into the echolocation ability of some shrews suggest that the evolution of echolocation in bats may involve inheritance of a primitive sonar system that was modified to its current state, rather than the ad hoc evolution of echolocation in the earliest bats. Because the cochlea is crucial in the sensation of echoes returning from sonar pulses, we tracked changes in cochlear morphology during development that included the basilar membrane (BM) and secondary spiral lamina (SSL) along the length of the cochlea in relation to stages of flight ability in young bats. Our data show that the morphological prerequisite for sonar sensitivity of the cochlea significantly precedes the onset of flight in young bats and, in fact, development of this prerequisite is complete before parturition. In addition, there were no discernible changes in cochlear morphology with stages of flight development, demonstrating temporal asymmetry between the development of morphology associated with echo-pulse return sensitivity and volancy. These data further corroborate and support the hypothesis that adaptations for sonar and echolocation evolved before flight in mammals., (© 2015 Anatomical Society.)

Bats evolved the ability of powered flight more than 50 million years ago. The modern bat is an efficient flyer and recent research on bat flight has revealed many intriguing facts. By using particle image velocimetry to visualize wake vortices, both the magnitude and time-history of aerodynamic forces can be estimated. At most speeds the downstroke generates both lift and thrust, whereas the function of the upstroke changes with forward flight speed. At hovering and slow speed bats use a leading edge vortex to enhance the lift beyond that allowed by steady aerodynamics and an inverted wing during the upstroke to further aid weight support. The bat wing and its skeleton exhibit many features and control mechanisms that are presumed to improve flight performance. Whereas bats appear aerodynamically less efficient than birds when it comes to cruising flight, they have the edge over birds when it comes to manoeuvring. There is a direct relationship between kinematics and the aerodynamic performance, but there is still a lack of knowledge about how (and if) the bat controls the movements and shape (planform and camber) of the wing. Considering the relatively few bat species whose aerodynamic tracks have been characterized, there is scope for new discoveries and a need to study species representing more extreme positions in the bat morphospace., (© 2015. Published by The Company of Biologists Ltd.)

Powered flight has evolved three times in the vertebrates: in the birds, the bats and the extinct pterosaurs. The largest bats ever known are at least an order of magnitude smaller than the largest members of the other two groups. Recently, it was argued that different scaling of wingbeat frequencies to body mass in birds and bats can help explain why the largest birds are larger than the largest bats. Here, I extend this argument in two ways. Firstly, I suggest that different respiratory physiologies are key to understanding the restriction on bat maximum size compared with birds. Secondly, I argue that a respiratory physiology similar to birds would have been a prerequisite for the gigantism seen in pterosaurs., (© 2014. Published by The Company of Biologists Ltd.)

Bats are the only mammals capable of true flight. Critical adaptations for flight include a pair of dramatically elongated hands with broad wing membranes. To study the molecular mechanisms of bat wing evolution, we perform genomewide mRNA sequencing and in situ hybridization for embryonic bat limbs. We identify seven key genes that display unique expression patterns in embryonic bat wings and feet, compared with mouse fore- and hindlimbs. The expression of all 5'HoxD genes (Hoxd9-13) and Tbx3, six known crucial transcription factors for limb and digit development, is extremely high and prolonged in the elongating wing area. The expression of Fam5c, a tumour suppressor, in bat limbs is bat-specific and significantly high in all short digit regions (the thumb and foot digits). These results suggest multiple genetic changes occurred independently during the evolution of bat wings to elongate the hand digits, promote membrane growth and keep other digits short. Our findings also indicate that the evolution of limb morphology depends on the complex integration of multiple gene regulatory networks and biological processes that control digit formation and identity, chondrogenesis, and interdigital regression or retention.

Many animals, on air, water, or land, navigate in three-dimensional (3D) environments, yet it remains unclear how brain circuits encode the animal's 3D position. We recorded single neurons in freely flying bats, using a wireless neural-telemetry system, and studied how hippocampal place cells encode 3D volumetric space during flight. Individual place cells were active in confined 3D volumes, and in >90% of the neurons, all three axes were encoded with similar resolution. The 3D place fields from different neurons spanned different locations and collectively represented uniformly the available space in the room. Theta rhythmicity was absent in the firing patterns of 3D place cells. These results suggest that the bat hippocampus represents 3D volumetric space by a uniform and nearly isotropic rate code.

Many birds use a flight mode called undulating or flap-gliding flight, where they alternate between flapping and gliding phases, while only a few bats make use of such a flight mode. Among birds, flap-gliding is commonly used by medium to large species, where it is regarded to have a lower energetic cost than continuously flapping flight. Here, we introduce a novel model for estimating the energetic flight economy of flap-gliding animals, by determining the lift-to-drag ratio for flap-gliding based on empirical lift-to-drag ratio estimates for continuous flapping flight and for continuous gliding flight, respectively. We apply the model to flight performance data of the common swift (Apus apus) and of the lesser long-nosed bat (Leptonycteris yerbabuenae). The common swift is a typical flap-glider while-to the best of our knowledge-the lesser long-nosed bat does not use flap-gliding. The results show that, according to the model, the flap-gliding common swift saves up to 15% energy compared to a continuous flapping swift, and that this is primarily due to the exceptionally high lift-to-drag ratio in gliding flight relative to that in flapping flight for common swifts. The lesser long-nosed bat, on the other hand, seems not to be able to reduce energetic costs by flap-gliding. The difference in relative costs of flap-gliding flight between the common swift and the lesser long-nosed bat can be explained by differences in morphology, flight style and wake dynamics. The model presented here proves to be a valuable tool for estimating energetic flight economy in flap-gliding animals. The results show that flap-gliding flight that is naturally used by common swifts is indeed the most economic one of the two flight modes, while this is not the case for the non-flap-gliding lesser long-nosed bat., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

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.

The ability to fly opens up ecological opportunities but flight mechanics and muscle energetics impose constraints, one of which is that the maximum body size must be kept below a rather low limit. The muscle power available for flight increases in proportion to flight muscle mass and wingbeat frequency. The maximum wingbeat frequency attainable among increasingly large animals decreases faster than the minimum frequency required, so eventually they coincide, thereby defining the maximum body mass at which the available power just matches up to the power required for sustained aerobic flight. Here, we report new wingbeat frequency data for 27 morphologically diverse bat species representing nine families, and additional data from the literature for another 38 species, together spanning a range from 2.0 to 870 g. For these species, wingbeat frequency decreases with increasing body mass as M(b)(-0.26). We filmed 25 of our 27 species in free flight outdoors, and for these the wingbeat frequency varies as M(b)(-0.30). These exponents are strikingly similar to the body mass dependency M(b)(-0.27) among birds, but the wingbeat frequency is higher in birds than in bats for any given body mass. The downstroke muscle mass is also a larger proportion of the body mass in birds. We applied these empirically based scaling functions for wingbeat frequency in bats to biomechanical theories about how the power required for flight and the power available converge as animal size increases. To this end we estimated the muscle mass-specific power required for the largest flying extant bird (12-16 kg) and assumed that the largest potential bat would exert similar muscle mass-specific power. Given the observed scaling of wingbeat frequency and the proportion of the body mass that is made up by flight muscles in birds and bats, we estimated the maximum potential body mass for bats to be 1.1-2.3 kg. The largest bats, extinct or extant, weigh 1.6 kg. This is within the range expected if it is the bat characteristic flight muscle mass and wingbeat frequency that limit the maximum body mass in bats. It is only a tenth the mass of the largest flying extant bird.

Historically, studies concerning bat flight have focused primarily on the wings. By analyzing high-speed video taken on 48 individuals of five species of vespertilionid bats, we show that the capacity to flap the tail-membrane (uropatagium) in order to generate thrust and lift during takeoffs and minimal-speed flight (<1 m (s-1)) was largely underestimated. Indeed, bats flapped the tail-membrane by extensive dorso-ventral fanning motions covering as much as 135 degrees of arc consistent with thrust generation by air displacement. The degree of dorsal extension of the tail-membrane, and thus the potential amount of thrust generated during platform launches, was significantly correlated with body mass (P = 0.02). Adduction of the hind limbs during upstrokes collapsed the tail-membrane thereby reducing its surface area and minimizing negative lift forces. Abduction of the hind limbs during the downstroke fully expanded the tail-membrane as it was swept ventrally. The flapping kinematics of the tail-membrane is thus consistent with expectations for an airfoil. Timing offsets between the wings and tail-membrane during downstrokes was as much as 50%, suggesting that the tail-membrane was providing thrust and perhaps lift when the wings were retracting through the upstoke phase of the wing-beat cycle. The extent to which the tail-membrane was used during takeoffs differed significantly among four vespertilionid species (P = 0.01) and aligned with predictions derived from bat ecomorphology. The extensive fanning motion of the tail membrane by vespertilionid bats has not been reported for other flying vertebrates.

Flying animals exhibit remarkable degrees of both stability and manoeuvrability. Our understanding of these capabilities has recently been improved by the identification of a source of passive damping specific to flapping flight. Examining how this damping effect scales among different species and how it affects active manoeuvres as well as recovery from perturbations provides general insights into the flight of insects, birds and bats. These new damping models offer a means to predict manoeuvrability and stability for a wide variety of flying animals using prior reports of the morphology and flapping motions of these species. Furthermore, the presence of passive damping is likely to have facilitated the evolution of powered flight in animals by providing a stability benefit associated with flapping.

A new study shows that hairs on the flight membranes of bats act as airflow sensors. Neurons in the brain that are sensitive to touch respond to stimulation of the hairs, and removal of the hairs compromises flight performance.

Bats are the only mammals capable of powered flight, and they perform impressive aerial maneuvers like tight turns, hovering, and perching upside down. The bat wing contains five digits, and its specialized membrane is covered with stiff, microscopically small, domed hairs. We provide here unique empirical evidence that the tactile receptors associated with these hairs are involved in sensorimotor flight control by providing aerodynamic feedback. We found that neurons in bat primary somatosensory cortex respond with directional sensitivity to stimulation of the wing hairs with low-speed airflow. Wing hairs mostly preferred reversed airflow, which occurs under flight conditions when the airflow separates and vortices form. This finding suggests that the hairs act as an array of sensors to monitor flight speed and/or airflow conditions that indicate stall. Depilation of different functional regions of the bats' wing membrane altered the flight behavior in obstacle avoidance tasks by reducing aerial maneuverability, as indicated by decreased turning angles and increased flight speed.

The 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.

In this study we compared the wing kinematics of 27 bats representing six pteropodid species ranging more than 40 times in body mass (M(b)=0.0278-1.152 kg), to determine whether wing posture and overall wing kinematics scaled as predicted according to theory. The smallest species flew in a wind tunnel and the other five species in a flight corridor. Seventeen kinematic markers on the midline and left side of the body were tracked in three dimensions. We used phylogenetically informed reduced major axis regression to test for allometry. We found that maximum wingspan (b(max)) and maximum wing area (S(max)) scaled with more positive allometry, and wing loading (Q(s)) with more negative allometry (b(max)∝M(b)(0.423); S(max)∝M(b)(0.768); Q(s)∝M(b)(0.233)) than has been reported in previous studies that were based on measurements from specimens stretched out flat on a horizontal surface. Our results suggest that larger bats open their wings more fully than small bats do in flight, and that for bats, body measurements alone cannot be used to predict the conformation of the wings in flight. Several kinematic variables, including downstroke ratio, wing stroke amplitude, stroke plane angle, wing camber and Strouhal number, did not change significantly with body size, demonstrating that many aspects of wing kinematics are similar across this range of body sizes. Whereas aerodynamic theory suggests that preferred flight speed should increase with mass, we did not observe an increase in preferred flight speed with mass. Instead, larger bats had higher lift coefficients (C(L)) than did small bats (C(L)∝M(b)(0.170)). Also, the slope of the wingbeat period (T) to body mass regression was significantly more shallow than expected under isometry (T∝M(b)(0.180)), and angle of attack (α) increased significantly with body mass [α∝log(M(b))7.738]. 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.

Bat wings are important for thermoregulation, but their role in heat balance during flight is largely unknown. More than 80% of the energy consumed during flight generates heat as a by-product, and thus it is expected that bat wings should dissipate large amounts of heat to prevent hyperthermia. We measured rectal (T(r)) and surface (T(s)) temperatures of Brazilian free-tailed bats (Tadarida brasiliensis) as they emerged from and returned to their daytime roosts and calculated sensible heat transfer for different body regions (head, body, wings, and tail membrane). Bats' T(r) decreased from 36.8°C during emergence flights to 34.4°C during returns, and T(s) scaled positively with ambient temperature (T(a)). Total radiative heat loss from bats was significantly greater for a radiative sink to the night sky than for a sink with temperature equal to T(a). We found that free-ranging Brazilian free-tailed bats, on average, do not dissipate heat from their wings by convection but instead dissipate radiative heat (L) to the cloudless night sky during flight ([Formula: see text] W). However, within the range of T(a) measured in this study, T. brasiliensis experienced net heat loss between evening emergence and return flights. Regional hypothermia reduces heat loss from wings that are exposed to potentially high convective fluxes. Additional research is needed to establish the role of wings in evaporative cooling during flight in 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.

Foraging behaviour of bats is supposedly largely influenced by the high costs of flapping flight. Yet our understanding of flight energetics focuses mostly on continuous horizontal forward flight at intermediate speeds. Many bats, however, perform manoeuvring flights at suboptimal speeds when foraging. For example, members of the genus Rhinolophus hunt insects during short sallying flights from a perch. Such flights include many descents and ascents below minimum power speed and are therefore considered energetically more expensive than flying at intermediate speed. To test this idea, we quantified the energy costs of short manoeuvring flights (<2 min) using the Na-bicarbonate technique in two Rhinolophus species that differ in body mass but have similar wing shapes. First, we hypothesized that, similar to birds, energy costs of short flights should be higher than predicted by an equation derived for bats at intermediate speeds. Second, we predicted that R. mehelyi encounters higher flight costs than R. euryale, because of its higher wing loading. Although wing loading of R. mehelyi was only 20% larger than that of R. euryale, its flight costs (2.61 ± 0.75 W; mean ± 1 SD) exceeded that of R. euryale (1.71 ± 0.37 W) by 50%. Measured flight costs were higher than predicted for R. mehelyi, but not for R. euryale. We conclude that R. mehelyi face elevated energy costs during short manoeuvring flights due to high wing loading and thus may optimize foraging efficiency by energy-conserving perch-hunting.

Body motions (kinematics) of animals can be dimensionally complex, especially when flexible parts of the body interact with a surrounding fluid. In these systems, tracking motion completely can be difficult, and result in a large number of correlated measurements, with unclear contributions of each parameter to performance. Workers typically get around this by deciding a priori which variables are important (wing camber, stroke amplitude, etc.), and focusing only on those variables, but this constrains the ability of a study to uncover variables of influence. Here, we describe an application of proper orthogonal decomposition (POD) for assigning importances to kinematic variables, using dimensional complexity as a metric. We apply this method to bat flight kinematics, addressing three questions: (1) Does dimensional complexity of motion change with speed? (2) What body markers are optimal for capturing dimensional complexity? (3) What variables should a simplified reconstruction of bat flight include in order to maximally reconstruct actual dimensional complexity? We measured the motions of 17 kinematic markers (20 joint angles) on a bat (Cynopterus brachyotis) flying in a wind tunnel at nine speeds. Dimensional complexity did not change with flight speed, despite changes in the kinematics themselves, suggesting that the relative efficacy of a given number of dimensions for reconstructing kinematics is conserved across speeds. By looking at subsets of the full 17-marker set, we found that using more markers improved resolution of kinematic dimensional complexity, but that the benefit of adding markers diminished as the total number of markers increased. Dimensional complexity was highest when the hindlimb and several points along digits III and IV were tracked. Also, we uncovered three groups of joints that move together during flight by using POD to quantify correlations of motion. These groups describe 14/20 joint angles, and provide a framework for models of bat flight for experimental and modeling purposes.

The wake structures of a bat in flight have a number of characteristics not associated with any of the bird species studied to this point. Unique features include discrete vortex rings generating negative lift at the end of the upstroke at medium and high speeds, each wing generating its own vortex loop, and a systematic variation in the circulation of the start and stop vortices along the wingspan, with increasing strength towards the wing tips. Here we analyse in further detail some previously published data from quantitative measurements of the wake behind a small bat species flying at speeds ranging from 1.5 to 7 m s(-1) in a wind tunnel. The data are extended to include both near- and far-wake measurements. The near-/far-wake comparisons show that although the measured peak vorticity of the start and stop vortices decreases with increasing downstream distance from the wing, the total circulation remains approximately constant. As the wake evolves, the diffuse stop vortex shed at the inner wing forms a more concentrated vortex in the far wake. Taken together, the results show that studying the far wake, which has been the standard procedure, nevertheless risks missing details of the wake. Although study of the far wake alone can lead to the misinterpretation of the wake topology, the net, overall circulation of the main wake vortices can be preserved so that approximate momentum balance calculations are not unreasonable within the inevitably large experimental uncertainties.

The evolution of flapping flight in bats from an arboreal gliding ancestor appears on the surface to be a relatively simple transition. However, bat flight is a highly complex functional system from a morphological, physiological, and aerodynamic perspective, and the transition from a gliding precursor may involve functional discontinuities that represent evolutionary hurdles. In this review, I suggest a framework for a comprehensive treatment of the evolution of complex functional systems that emphasizes a mechanistic understanding of the initial state, the final state, and the proposed transitional states. In this case, bats represent the final state and extant mammalian gliders are used as a model for the initial state. To explore possible transitional states, I propose a set of criteria for evaluating hypotheses about the evolution of flight in vertebrates and suggest methods by which we can advance our understanding of the transition from gliding to flapping flight. Although it is impossible ever to know with certainty the sequence of events landing to flapping flight, the field of possibilities can be narrowed to those that maintain the functional continuity of the wing and result in improved aerodynamic performance across this transition. The fundamental differences between gliding and flapping flight should not necessarily be seen as evidence that this transition could not occur; rather, these differences point out compelling aspects of the aerodynamics of animal wings that require further investigation.

Bats (Chiroptera) represent one of the largest and most diverse radiations of mammals, accounting for one-fifth of extant species. Although recent studies unambiguously support bat monophyly and consensus is rapidly emerging about evolutionary relationships among extant lineages, the fossil record of bats extends over 50 million years, and early evolution of the group remains poorly understood. Here we describe a new bat from the Early Eocene Green River Formation of Wyoming, USA, with features that are more primitive than seen in any previously known bat. The evolutionary pathways that led to flapping flight and echolocation in bats have been in dispute, and until now fossils have been of limited use in documenting transitions involved in this marked change in lifestyle. Phylogenetically informed comparisons of the new taxon with other bats and non-flying mammals reveal that critical morphological and functional changes evolved incrementally. Forelimb anatomy indicates that the new bat was capable of powered flight like other Eocene bats, but ear morphology suggests that it lacked their echolocation abilities, supporting a 'flight first' hypothesis for chiropteran evolution. The shape of the wings suggests that an undulating gliding-fluttering flight style may be primitive for bats, and the presence of a long calcar indicates that a broad tail membrane evolved early in Chiroptera, probably functioning as an additional airfoil rather than as a prey-capture device. Limb proportions and retention of claws on all digits indicate that the new bat may have been an agile climber that employed quadrupedal locomotion and under-branch hanging behaviour.

Tetrapod limbs display a great variety of forms and functions. In an attempt to understand factors influencing this diversity, Cretekos and colleagues, in a recent issue of Genes & Development, provide molecular insight into the evolution of wing morphology in bats.

We measured qualitative and quantitative aspects of the head and body pelage of 23 species of Western Australian bats. A functionally appropriate relationship was found with the normal flight speeds and foraging strategy of the bats at three levels of geometric consideration: overall fur texture, individual hair length and cuticular scale attributes (scale type, scale length and diameter, as well as sub-scale detail design). This relationship is best explained by describing the pelage surface as characterised by aerodynamic riblets. For species that utilise high-speed and aerodynamically efficient flight during commuting and foraging, riblets should reduce the skin friction drag of the head and body by up to 10%. The molossids, emballonurids and one pteropid studied have fur that falls within the non-dimensional height range that gives best aerodynamic efficiency, 8

The evolution of sociality is a central theme in evolutionary biology. The vast majority of bats are social, which has been explained in terms of the benefits of communal breeding. However, the causes for segregated male groups remain unknown. In a comparative study, we tested whether diet and morphological adaptations to specific foraging styles, two factors known to influence the occurrence of information transfer, can predict male sociality. Our results suggest that the species most likely to benefit from information transfer--namely, those preying on ephemeral insects and with morphological adaptations to feeding in open habitat--are more likely to form male groups. Our findings also indicate that solitary life was the ancestral state of males and sociality evolved in several lineages. Beyond their significance for explaining the existence of male groups in bats, our findings highlight the importance of information transfer in the evolution of animal sociality.

The flapping flight of animals generates an aerodynamic footprint as a time-varying vortex wake in which the rate of momentum change represents the aerodynamic force. We showed that the wakes of a small bat species differ from those of birds in some important respects. In our bats, each wing generated its own vortex loop. Also, at moderate and high flight speeds, the circulation on the outer (hand) wing and the arm wing differed in sign during the upstroke, resulting in negative lift on the hand wing and positive lift on the arm wing. Our interpretations of the unsteady aerodynamic performance and function of membranous-winged, flapping flight should change modeling strategies for the study of equivalent natural and engineered flying devices.

Three vertebrate groups - birds, bats and pterosaurs - have evolved flapping flight over the past 200 million years. This innovation allowed each clade access to new ecological opportunities, but did the diversification of one of these groups inhibit the evolutionary radiation of any of the others? A related question is whether having the wing attached to the hindlimbs in bats and pterosaurs constrained their morphological diversity relative to birds. Fore- and hindlimb measurements from 894 specimens were used to construct a morphospace to assess morphological overlap and range, a possible indicator of competition, among the three clades. Neither birds nor bats entered pterosaur morphospace across the Cretaceous-Paleogene (Tertiary) extinction. Bats plot in a separate area from birds, and have a significantly smaller morphological range than either birds or pterosaurs. On the basis of these results, competitive exclusion among the three groups is not supported.

Postnatal changes in wing morphology, flight development and aerodynamics were studied in captive free-flying short-nosed fruit bats, Cynopterus sphinx. Pups were reluctant to move until 25 days of age and started fluttering at the mean age of 40 days. The wingspan and wing area increased linearly until 45 days of age by which time the young bats exhibited clumsy flight with gentle turns. At birth, C. sphinx had less-developed handwings compared to armwings; however, the handwing developed faster than the armwing during the postnatal period. Young bats achieved sustained flight at 55 days of age. Wing loading decreased linearly until 35 days of age and thereafter increased to a maximum of 12.82 Nm(-2) at 125 days of age. The logistic equation fitted the postnatal changes in wingspan and wing area better than the Gompertz and von Bertalanffy equations. The predicted minimum power speed (V(mp)) and maximum range speed (V(mr)) decreased until the onset of flight and thereafter the V(mp) and V(mr) increased linearly and approached 96.2% and 96.4%, respectively, of the speed of postpartum females at the age of 125 days. The requirement of minimum flight power (P(mp)) and maximum range power (P(mr)) increased until 85 days of age and thereafter stabilised. The minimum theoretical radius of banked turn (r(min)) decreased until 35 days of age and thereafter increased linearly and attained 86.5% of the r(min) of postpartum females at the age of 125 days.

Experimental measurements and analysis of the flight of bats are presented, including kinematic analysis of high-speed stereo videography of straight and turning flight, and measurements of the wake velocity field behind the bat. The kinematic data reveal that, at relatively slow flight speeds, wing motion is quite complex, including a sharp retraction of the wing during the upstroke and a broad sweep of the partially extended wing during the downstroke. The data also indicate that the flight speed and elevation are not constant, but oscillate in synchrony with both the horizontal and vertical movements of the wing. PIV measurements in the transverse (Trefftz) plane of the wake indicate a complex 'wake vortex' structure dominated by a strong wing tip vortex shed from the wing tip during the downstroke and either the wing tip or a more proximal joint during the upstroke. Data synthesis of several discrete realizations suggests a 'cartoon' of the wake structure during the entire wing beat cycle. Considerable work remains to be done to confirm and amplify these results.