Bat Flight Research Articles

Bats and Mines: Evaluating Townsend's Big-Eared Bat (Corynorhinus townsendii) Maternity Colony Behavioral Response to Gating

With the loss and modification of natural roosting habitat afforded by caves, abandoned mines have assumed increased importance as alternative roosting sites for Townsend's big-eared bats (Corynorhinus townsendii). However, increased human safety concerns have led to accelerated abandoned mine closure programs. To protect roosting sites in mines with significant bat activity, bat-compatible gates are installed that allow continued access to mine workings. Aside from ensuring public safety, these gates provide protection from disturbance to roosting bats. We evaluated the effects of gating on bat flight behavior at maternity colonies in 2 previously gated (control) and 2 ungated (treatment) mines that were gated during this study. We used an infrared video camera to record bat flight behavior at the entrances to each of the 4 study mines for 2 consecutive mornings and a single night each month during the warm season. Entrance (03:00–06:00) and emergence (21:00–24:00) surveys comprised 3 consecutive hours. Overall circling activity increased more than 6-fold at openings of treatment mines following gating (P < 0.001). Crowding during emergence was significantly higher (P = 0.023) in newly gated mines than in previously gated mines. Gates affect subadults during the initial-volancy periods, as detected through collisions with the gates. Increased activity of bats and collisions with the gate, which result in bats falling to the ground at mine openings, may amplify vulnerability to predators and increase energetic demands.

Aerodynamic mechanism of forces generated by twisting model-wing in bat flapping flight

The aerodynamic mechanism of the bat wing membrane along the lateral border of its body is studied. The twist-morphing that alters the angle of attack (AOA) along the span-wise direction is observed widely during bat flapping flight. An assumption is made that the linearly distributed AOA is along the span-wise direction. The plate with the aspect ratio of 3 is used to model a bat wing. A three-dimensional (3D) unsteady panel method is used to predict the aerodynamic forces generated by the flapping plate with leading edge separation. It is found that, relative to the rigid wing flapping, twisting motion can increase the averaged lift by as much as 25% and produce thrust instead of drag. Furthermore, the aerodynamic forces (lift/drag) generated by a twisting plate-wing are similar to those of a pitching rigid-wing, meaning that the twisting in bat flight has the same function as the supination/pronation motion in insect flight.

Sugar flux through the flight muscles of hovering vertebrate nectarivores: a review

In most vertebrates, uptake and oxidation of circulating sugars by locomotor muscles rises with increasing exercise intensity. However, uptake rate by muscle plateaus at moderate aerobic exercise intensities and intracellular fuels dominate at oxygen consumption rates of 50% of maximum or more. Further, uptake and oxidation of circulating fructose by muscle is negligible. In contrast, hummingbirds and nectar bats are capable of fueling expensive hovering flight exclusively, or nearly completely, with dietary sugar. In addition, hummingbirds and nectar bats appear capable of fueling hovering flight completely with fructose. Three crucial steps are believed to be rate limiting to muscle uptake of circulating glucose or fructose in vertebrates: (1) delivery to muscle; (2) transport into muscle through glucose transporter proteins (GLUTs); and (3) phosphorylation of glucose by hexokinase (HK) within the muscle. In this review, we summarize what is known about the functional upregulation of exogenous sugar flux at each of these steps in hummingbirds and nectar bats. High cardiac output, capillary density, and blood sugar levels in hummingbirds and bats enhance sugar delivery to muscles (step 1). Hummingbird and nectar bat flight muscle fibers have relatively small cross-sectional areas and thus relatively high surface areas across which transport can occur (step 2). Maximum HK activities in each species are enough for carbohydrate flux through glycolysis to satisfy 100 % of hovering oxidative demand (step 3). However, qualitative patterns of GLUT expression in the muscle (step 2) raise more questions than they answer regarding sugar transport in hummingbirds and suggest major differences in the regulation of sugar flux compared to nectar bats. Behavioral and physiological similarities among hummingbirds, nectar bats, and other vertebrates suggest enhanced capacities for exogenous fuel use during exercise may be more wide spread than previously appreciated. Further, how the capacity for uptake and phosphorylation of circulating fructose is enhanced remains a tantalizing unknown.

Lift enhancement by dynamically changing wingspan in forward flapping flight

Dynamically stretching and retracting wingspan has been widely observed in the flight of birds and bats, and its effects on the aerodynamic performance particularly lift generation are intriguing. The rectangular flat-plate flapping wing with a sinusoidally stretching and retracting wingspan is proposed as a simple model for biologically inspired dynamic morphing wings. Numerical simulations of the low-Reynolds-number flows around the flapping morphing wing are conducted in a parametric space by using the immersed boundary method. It is found that the instantaneous and time-averaged lift coefficients of the wing can be significantly enhanced by dynamically changing wingspan in a flapping cycle. The lift enhancement is caused by both changing the lifting surface area and manipulating the flow structures responsible to the vortex lift generation. The physical mechanisms behind the lift enhancement are explored by examining the three-dimensional flow structures around the flapping wing.

Hindlimb motion during steady flight of the lesser dog-faced fruit bat, Cynopterus brachyotis.

In bats, the wing membrane is anchored not only to the body and forelimb, but also to the hindlimb. This attachment configuration gives bats the potential to modulate wing shape by moving the hindlimb, such as by joint movement at the hip or knee. Such movements could modulate lift, drag, or the pitching moment. In this study we address: 1) how the ankle translates through space during the wingbeat cycle; 2) whether amplitude of ankle motion is dependent upon flight speed; 3) how tension in the wing membrane pulls the ankle; and 4) whether wing membrane tension is responsible for driving ankle motion. We flew five individuals of the lesser dog-faced fruit bat, Cynopterus brachyotis (Family: Pteropodidae), in a wind tunnel and documented kinematics of the forelimb, hip, ankle, and trailing edge of the wing membrane. Based on kinematic analysis of hindlimb and forelimb movements, we found that: 1) during downstroke, the ankle moved ventrally and during upstroke the ankle moved dorsally; 2) there was considerable variation in amplitude of ankle motion, but amplitude did not correlate significantly with flight speed; 3) during downstroke, tension generated by the wing membrane acted to pull the ankle dorsally, and during upstroke, the wing membrane pulled laterally when taut and dorsally when relatively slack; and 4) wing membrane tension generally opposed dorsoventral ankle motion. We conclude that during forward flight in C. brachyotis, wing membrane tension does not power hindlimb motion; instead, we propose that hindlimb movements arise from muscle activity and/or inertial effects.

Bat flight and zoonotic viruses.

Bats are sources of high viral diversity and high-profile zoonotic viruses worldwide. Although apparently not pathogenic in their reservoir hosts, some viruses from bats severely affect other mammals, including humans. Examples include severe acute respiratory syndrome coronaviruses, Ebola and Marburg viruses, and Nipah and Hendra viruses. Factors underlying high viral diversity in bats are the subject of speculation. We hypothesize that flight, a factor common to all bats but to no other mammals, provides an intensive selective force for coexistence with viral parasites through a daily cycle that elevates metabolism and body temperature analogous to the febrile response in other mammals. On an evolutionary scale, this host-virus interaction might have resulted in the large diversity of zoonotic viruses in bats, possibly through bat viruses adapting to be more tolerant of the fever response and less virulent to their natural hosts.

Cool sperm: why some placental mammals have a scrotum.

Throughout the Cenozoic, the fitness benefits of the scrotum in placental mammals presumably outweighed the fitness costs through damage, yet a definitive hypothesis for its evolution remains elusive. Here, I present an hypothesis (Endothermic Pulses Hypothesis) which argues that the evolution of the scrotum was driven by Cenozoic pulses in endothermy, that is, increases in normothermic body temperature, which occurred in Boreotheria (rodents, primates, lagomorphs, carnivores, bats, lipotyphylans and ungulates) in response to factors such as cursoriality and climate adaptation. The model argues that stabilizing selection maintained an optimum temperature for spermatogenesis and sperm storage throughout the Cenozoic at the lower plesiomorphic levels of body temperature that prevailed in ancestral mammals for at least 163million years. Evolutionary stasis may have been driven by reduced rates of germ-cell mutations at lower body temperatures. Following the extinction of the dinosaurs at the Cretaceous-Palaeogene boundary 65.5mya, immediate pulses in endothermy occurred associated with the dramatic radiation of the modern placental mammal orders. The fitness advantages of an optimum temperature of spermatogenesis outweighed the potential costs of testes externalization and paved the way for the evolution of the scrotum. The scrotum evolved within several hundred thousand years of the K-Pg extinction, probably associated initially with the evolution of cursoriality, and arguably facilitated mid- and late Cenozoic metabolic adaptations to factors such as climate, flight in bats and sociality in primates.

A modelling approach to infer the effects of wind farms on landscape connectivity for bats

Little is known about the potentially disrupting effects of wind farms on the habitat connectivity of flying vertebrates at the landscape scale. We developed a regional-scale model to assess the wind farm impact on bat migration and commuting routes. The model was implemented for the bat Nyctalus leisleri in a region of central Italy currently undergoing considerable wind farm development. A Species Distribution Model (SDM) for N. leisleri was generated using the MaxEnt algorithm based on 47 presence records (reduced to 19 after the autocorrelation procedure) and 10 environmental variables derived from topographic and land cover maps. We used the SDM to create a map of connectivity using the software UNICOR to identify potential commuting corridors (PCCs). The incidence of each wind farm on bat flight corridors was assessed by overlaying the existing (380) and planned (195) turbine locations onto the PCCs. The SDM was statistically robust (AUC > 0.8). Most of the corridors were concentrated in the western part of the region, which hosts the largest suitable areas for the species; most of the existing (54 %) and planned (72 %) wind farms interfered with important corridors connecting the western and the eastern parts of the region. Our results provide key information on the impact of the wind farm industry on biodiversity on a regional scale. The novel approach adopted, based on SDM and connectivity analysis, could be easily extended to other flying vertebrates and landscapes and constitutes a promising planning tool necessary for harmonizing the development of renewable energy infrastructures with issues of biodiversity conservation.

Bats (Tadarida brasiliensis) jam conspecifics in food competition

We here describe field experiments testing whether bats adaptively produce sounds to interfere with (“jam”) the echolocation of other bats. Visual observations, low-light videography, and ultrasound recording with microphone arrays allowing reconstruction of bat flight paths were used to document interactions between Mexican free-tailed bats (Tadarida brasiliensis) at two foraging locations in southern Arizona and New Mexico. We tested three sets of predictions based on the jamming hypothesis and two competing hypotheses—cooperative foraging and food patch defense. Bats produced putative jamming calls (termed sinFM calls) that overlapped temporally and spectrally with “feeding buzz” calls made by conspecifics when attacking insects. Bat capture success decreased by 400% when sinFM calls were present compared to when they were absent. Behavioral sequences consisted of two or more bats sequentially making feeding buzzes within a restricted area while another bats made sinFM calls. After making sinFM calls bats frequently turned toward where the other bat made its feeding buzz and then made a buzz of its own. Together, the results support the hypothesis that bats jam conspecifics in extended bouts of food competition. This is the first known case of echolocating animals adaptively jamming conspecifics.

Straight-line climbing flight aerodynamics of a fruit bat

From flight data obtained on a fruit bat, Cynopterus brachyotis, a kinematic model for straight-line flapping motion is extracted and analyzed in a computational fluid dynamics (CFD) framework to gain insight into the complexity of bat flight. The intricate functional mechanics and architecture of the bat wings set it apart from other vertebrate flight. The extracted kinematic model is simulated for a range of Reynolds numbers, to observe the effect these phenomena have on the unsteady transient mechanisms of the flow produced by the flapping wings. The Strouhal number calculated from the data is high indicating that the oscillatory motion dominates the flow physics. From the obtained data, the bat exhibits fine control of its mechanics by actively varying wing camber, wing area, torsional rotation of the wing, forward and backward translational sweep of the wing, and wing conformation to dictate the fluid dynamics. As is common in flapping flight, the primary force generation is through the attached unsteady vortices on the wing surface. The bat through varying the wing camber and the wing area modulates this force output. The power requirement for the kinematics is analyzed and correlated with the aerodynamic performance.

A protocol and calibration method for accurate multi-camera field videography

Stereo videography is a powerful technique for quantifying the kinematics and behavior of animals, but it can be challenging to use in an outdoor field setting. We here present a workflow and associated software for performing calibration of cameras placed in a field setting and estimating the accuracy of the resulting stereoscopic reconstructions. We demonstrate the workflow through example stereoscopic reconstructions of bat and bird flight. We provide software tools for planning experiments and processing the resulting calibrations that other researchers may use to calibrate their own cameras. Our field protocol can be deployed in a single afternoon, requiring only short video clips of light, portable calibration objects.

Fine-tuned echolocation and capture-flight ofMyotis capacciniiwhen facing different sized insects and fish prey

Formerly thought to be a strictly insectivorous trawling bat, recent studies have shown that Myotis capaccinii also preys on fish. To determine whether differences exist in bat flight behaviour, prey handling and echolocation characteristics when catching fish and insects of different size, we conducted a field experiment focused on the last stage of prey capture. We used synchronized video and ultrasound recordings to measure several flight and dip features as well as echolocation characteristics, focusing on terminal buzz phase I, characterized by a call rate exceeding 100 Hz, and buzz phase II, characterized by a drop in the fundamental well below 20 kHz and a repetition rate exceeding 150 Hz. When capturing insects, bats used both parts of the terminal phase to the same extent, and performed short and superficial drags on the water surface. In contrast, when preying on fish, buzz I was longer and buzz II shorter, and the bats made longer and deeper dips. These variations suggest that lengthening buzz I and shortening buzz II when fishing is beneficial, probably because buzz I gives better discrimination ability and the broader sonar beam provided by buzz II is useless when no evasive flight of the prey is expected. Additionally, bats continued emitting calls beyond the theoretical signal-overlap zone, suggesting that they might obtain information even when they have surpassed that threshold, at least initially. This study shows that M. capaccinii can regulate the temporal components of its feeding buzzes and modify prey capture technique according to the target.

Perception and Steering Control in Paired Bat Flight

Animals within groups need to coordinate their reactions to perceived environmental features and to each other in order to safely move from one point to another. This paper extends our previously published work on the flight patterns of Myotis velifer that have been observed in a habitat near Johnson City, Texas. Each evening, these bats emerge from a cave in sequences of small groups that typically contain no more than three or four individuals, and they thus provide ideal subjects for studying leader-follower behaviors. By analyzing the flight paths of a group of M. velifer, the data show that the flight behavior of a follower bat is influenced by the flight behavior of a leader bat in a way that is not well explained by existing pursuit laws, such as classical pursuit, constant bearing and motion camouflage. Thus we propose an alternative steering law based on virtual loom, a concept we introduce to capture the geometrical configuration of the leader-follower pair. It is shown that this law may be integrated with our previously proposed vision-enabled steering laws to synthesize trajectories, the statistics of which fit with those of the bats in our data set. The results suggest that bats use perceived information of both the environment and their neighbors for navigation.

Detailed Analysis of Variables Affecting Wing Kinematics of Bat Flight

Detailed Analysis of Variables Affecting Wing Kinematics of Bat Flight

Morphing Models of a Bat Wing in Flapping Flight

Kinematic models, which contain flapping and deformations, of a bat flapping wing when it flies at medium speeds are presented in the study. The deformations include cambering - morphing in the chordwise direction, bending - morphing in the spanwise direction and twisting respecting angles of attack (AOA) varying along the length of the wing. In the study, we assumed a circular-arc-shaped deformation when cambering or bending occurs, and introduced controlling parameters for each morphing. Then we showed the expressions of the controlling parameters varying with time at the bat flight speed of 3m/s according to the data provided by a literature, and verified the reasonability of the assumptions at last.

Investigation of the diving reflex using an ice bag – Comparison of free breathing and no breathing

Bats are unique among extant flying animals, as they have compliant wings and an echolocation sensory system that distinguish them from birds and insects. Flying in the dark, guided by echolocation, has influenced the aerodynamics of bat flight perhaps more than previously realized and resulted in a characteristic flight that is now being revealed.

Interpretation of body-mounted accelerometry in flying animals and estimation of biomechanical power

An idealized energy fluctuation model of a bird's body undergoing horizontal flapping flight is developed, focusing on the biomechanical power discernible to a body-mounted accelerometer. Expressions for flight body power constructed from root mean square dynamic body accelerations and wingstroke frequency are derived from first principles and presented in dimensionally appropriate units. As wingstroke frequency increases, the model generally predicts a gradual transition in power from a linear to an asymptotically cubic relationship. However, the onset of this transition and the degree to which this occurs depends upon whether and how forward vibrations are exploited for temporary energy storage and retrieval. While this may vary considerably between species and individual birds, it is found that a quadrature phase arrangement is generally advantageous during level flight. Gravity-aligned vertical acceleration always enters into the calculation of body power, but, whenever forward acceleration becomes relevant, its contribution is subtractive. Several novel kinematic measures descriptive of flapping flight are postulated, offering fresh insights into the processes involved in airborne locomotion. The limitations of the model are briefly discussed, and departures from its predictions during ascending and descending flight evaluated. These findings highlight how body-mounted accelerometers can offer a valuable, insightful and non-invasive technique for investigating the flight of free-ranging birds and bats.

Study of bat flight behavior by combining thermal image analysis with a LiDAR forest reconstruction

The nature of forest structure plays an important role in the study of foraging behaviors of bats. In this study, we demonstrate a new combined methodology that uses both thermal imaging technology and a ground-based LiDAR system to record and reconstruct Eptesicus fuscus (big brown bats) flight trajectories in three-dimensional (3-D) space. The combination of the two 3-D datasets provided a fine-scale reconstruction of the flight characteristics adjacent to and within the forests. A 3-D forest reconstruction, assembled from nine Echidna Validation Instrument LiDAR scans over the 1 ha site area, provided the essential environmental variables for the study of bat foraging behaviors, such as the canopy height, terrain, location of the obstacles, and canopy openness at a bat roosting and maternity site in Petersham, Massachusetts. Flight trajectories of 24 bats were recorded over the 25 m × 37.5 m region within the LiDAR forest reconstruction area. The trajectories were reconstructed using imaging data from multiple FLIR ThermoVision SC8000 cameras and were co-registered to the 3-D forest reconstruction. Twenty-four of these flight trajectories were categorized into four different behavior groups according to velocity and altitude analysis of the flight trajectories. Initial results showed that although all bats were guided by echolocation and avoided hitting a tree that was in all of their flight paths, different bats chose different flight routes. This study is an initial demonstration of the power of coupling thermal image analysis and LiDAR forest reconstructions. Our goal was to break ground for future ecological studies, where more extensive flight trajectories of bats can be coupled with the canopy reconstructions to better establish responses of bats to different habitat characteristics and clutter, which includes both static (trees) and dynamic (other bats) obstacles.

Bat flight with bad wings: is flight metabolism affected by damaged wing membranes?

Infection of North American bats with the keratin-digesting fungus Geomyces destructans often results in holes and ruptures of wing membranes, yet it is unknown whether flight performance and metabolism of bats are altered by such injuries. I conducted flight experiments in a circular flight arena with Myotis albescens and M. nigricans individuals with an intact or ruptured trailing edge of one of the plagiopatagial membranes. In both species, individuals with damaged wings were lighter, had a higher aspect ratio (squared wing span divided by wing area) and an increased wing loading (weight divided by wing area) than conspecifics with intact wings. Bats with an asymmetric reduction of the wing area flew at similar speeds to conspecifics with intact wings but performed fewer flight manoeuvres. Individuals with damaged wings showed lower metabolic rates during flight than conspecifics with intact wings, even when controlling for body mass differences; the difference in mass-specific metabolic rate may be attributable to the lower number of flight manoeuvres (U-turns) by bats with damaged wings compared with conspecifics with intact wings. Possibly, bats compensated for an asymmetric reduction in wing area by lowering their body mass and avoiding flight manoeuvres. In conclusion, it may be that bats suffer from moderate wing damage not directly, by experiencing increased metabolic rate, but indirectly, by a reduced manoeuvrability and foraging success. This could impede a bat's ability to gain sufficient body mass before hibernation.

Myological adaptations to fast enduring flight in European free-tailed bats, Tadarida teniotis (Molossidae, Chiroptera)

Molossid bats are considered as morphologically specialized for fast enduring flights within obstacle-free environments. The only European species of the family, the European free-tailed bat Tadarida teniotis, exhibits a similar flight behavior and is thus expected to demonstrate analogous morphological adaptations. The present report investigates external and muscular forelimb features that could be associated with open air high-speed flight patterns. For the scope of the study, we calculated standard external ecomorphological indices and carried out gross anatomical dissections on six adult specimens of T. teniotis from northern Greece. In this way, we qualitatively and quantitatively examined patterns of relative mass and strength in selected muscles. In terms of external morphology, T. teniotis possessed long and narrow wings with slightly pointed tips. Regarding forelimb muscular anatomy, the species is characterized by powerful wing adductors (e.g. pectorales), an enlargement of the coracoid head of biceps brachii, and equal mass of elbow joint flexors and extensors. These characteristics, partly shared by other molossids, may correlate to a powerful, controlled downstroke and a semi-extended elbow for fast enduring non-maneuverable flight.