Buckling-induced sound production in the aeroelastic tymbals of Yponomeuta.

The loss of elastic stability (buckling) can lead to catastrophic failure in the context of traditional engineering structures. Conversely, in nature, buckling often serves a desirable function, such as in the prey-trapping mechanism of the Venus fly trap (Dionaea muscipula). This paper investigates the buckling-enabled sound production in the wingbeat-powered (aeroelastic) tymbals of Yponomeuta moths. The hindwings of Yponomeuta possess a striated band of ridges that snap through sequentially during the up- and downstroke of the wingbeat cycle-a process reminiscent of cellular buckling in compressed slender shells. As a result, bursts of ultrasonic clicks are produced that deter predators (i.e. bats). Using various biological and mechanical characterization techniques, we show that wing camber changes during the wingbeat cycle act as the single actuation mechanism that causes buckling to propagate sequentially through each stria on the tymbal. The snap-through of each stria excites a bald patch of the wing's membrane, thereby amplifying sound pressure levels and radiating sound at the resonant frequencies of the patch. In addition, the interaction of phased tymbal clicks from the two wings enhances the directivity of the acoustic signal strength, suggesting an improvement in acoustic protection. These findings unveil the acousto-mechanics of Yponomeuta tymbals and uncover their buckling-driven evolutionary origin. We anticipate that through bioinspiration, aeroelastic tymbals will encourage novel developments in the context of multi-stable morphing structures, acoustic structural monitoring, and soft robotics.

Effects of forest fragmentation on the dietary ecology and activity of a nocturnal lemur community in North West Madagascar.

Deforestation and habitat fragmentation is the primary threat to primate populations. The primates that live within degraded and anthropogenically disturbed habitats typical of fragmented landscapes have to cope with lower availability of resources in comparison to primates in continuous, undisturbed forests. While some species are sensitive to forest fragmentation, some evidence exists to suggest that primates can alter their behavior and adapt to such changes, which enables their survival in suboptimal habitat. In this study, we assessed how forest fragmentation and its associated edge-effects impact the feeding ecology and activity levels of a nocturnal primate community in the Sahamalaza-Iles Radama National Park, North West Madagascar. From March 06, 2019 to May 17, 2022, we collected data on tree and invertebrate phenology at our study site, and feeding ecology and activity for 159 lemur individuals from four species. Fruit and flower availability varied significantly between continuous and fragmented forest, and between forest core and edge areas, with continuous forest exhibiting higher continuous fruit and flower availability. Lemur feeding ecology varied significantly too, as the feeding niches of all four species were significantly different between continuous and fragmented forest and between core and edge areas. However, lemur activity levels were mostly consistent among all forest areas. The results of this study suggest that nocturnal lemurs are able to adapt their dietary ecology in response to the available food sources within their habitat. Due to this flexible ecology and dietary plasticity, the lemurs do not need to significantly alter their behavior in different environments to fulfill their dietary needs. While nocturnal lemurs demonstrate adaptability and flexibility to degraded habitat, it is unclear how far this plasticity will stretch considering that Madagascar's forests are still being cleared at an alarming rate. Urgent conservation action is therefore needed to ensure the future of lemur habitat.

Moth wings are acoustic metamaterials.

Metamaterials assemble multiple subwavelength elements to create structures with extraordinary physical properties (1-4). Optical metamaterials are rare in nature and no natural acoustic metamaterials are known. Here, we reveal that the intricate scale layer on moth wings forms a metamaterial ultrasound absorber (peak absorption = 72% of sound intensity at 78 kHz) that is 111 times thinner than the longest absorbed wavelength. Individual scales act as resonant (5) unit cells that are linked via a shared wing membrane to form this metamaterial, and collectively they generate hard-to-attain broadband deep-subwavelength absorption. Their collective absorption exceeds the sum of their individual contributions. This sound absorber provides moth wings with acoustic camouflage (6) against echolocating bats. It combines broadband absorption of all frequencies used by bats with light and ultrathin structures that meet aerodynamic constraints on wing weight and thickness. The morphological implementation seen in this evolved acoustic metamaterial reveals enticing ways to design high-performance noise mitigation devices.

Models for resonant acoustic metasurfaces with application to moth wing ultrasound absorption.

Taking as bioinspiration the remarkable acoustic absorption properties of moth wings, we develop a simple analytical model that describes the interaction between acoustic pressure fields, and thin elastic plates incorporating resonant sub-structures. The moth wing is an exemplar of a natural acoustic metamaterial; the wings are deeply subwavelength in thickness at the frequencies of interest, the absorption is broadband and the tiny scales resonate on the moth wing acting in concert. The simplified model incorporates only the essential physics and the scales are idealized to flat rigid rectangular plates coupled via a spring to an elastic plate that forms the wing; all the components are deep-subwavelength at desired frequencies. Based on Fourier analysis, complemented by phenomenological modelling, our theory shows excellent agreement with simulation mimicking the moth-wing structure. Moth wings operate as broadband sound absorbers employing a range of scale sizes. We demonstrate that a random distribution of scale sizes generates a broadband absorption spectrum. To further illustrate the potential of the model, we design a deeply sub-wavelength acoustic counterpart of electromagnetically induced reflectance. This article is part of the theme issue 'Wave generation and transmission in multi-scale complex media and structured metamaterials (part 2)'.

Acoustic deterrents influence foraging activity, flight and echolocation behaviour of free-flying bats.

Acoustic deterrents have shown potential as a viable mitigation measure to reduce human impacts on bats; however, the mechanisms underpinning acoustic deterrence of bats have yet to be explored. Bats avoid ambient ultrasound in their environment and alter their echolocation calls in response to masking noise. Using stereo thermal videogrammetry and acoustic methods, we tested predictions that: (i) bats would avoid acoustic deterrents and forage and social call less in a 'treated airspace'; (ii) deterrents would cause bats to fly with more direct flight paths akin to commuting behaviour and in line with a reduction in foraging activity, resulting in increased flight speed and decreased flight tortuosity; and (iii) bats would alter their echolocation call structure in response to the masking deterrent sound. As predicted, overall bat activity was reduced by 30% and we recorded a significant reduction in counts of Pipistrellus pygmaeus (27%), Myotis spp. (probably M. daubentonii) (26%), and Nyctalus spp. and Eptesicus spp. (68%) passes. Pipistrellus pygmaeus feeding buzzes were also reduced by the deterrent in relation to general activity (by 38%); however, social calls were not (only 23% reduction). Bats also increased their flight speed and reduced the tortuosity of their flight paths, and P. pygmaeus reduced echolocation call bandwidth and start frequency of calls in response to deterrent playback, probably owing to the masking effect of the sound. Deterrence could therefore be used to remove bats from areas where they forage, for example wind turbines and roads, where they may be under threat from direct mortality.

Biomechanics of a moth scale at ultrasonic frequencies.

The wings of moths and butterflies are densely covered in scales that exhibit intricate shapes and sculptured nanostructures. While certain butterfly scales create nanoscale photonic effects, moth scales show different nanostructures suggesting different functionality. Here we investigate moth-scale vibrodynamics to understand their role in creating acoustic camouflage against bat echolocation, where scales on wings provide ultrasound absorber functionality. For this, individual scales can be considered as building blocks with adapted biomechanical properties at ultrasonic frequencies. The 3D nanostructure of a full Bunaea alcinoe moth forewing scale was characterized using confocal microscopy. Structurally, this scale is double layered and endowed with different perforation rates on the upper and lower laminae, which are interconnected by trabeculae pillars. From these observations a parameterized model of the scale's nanostructure was formed and its effective elastic stiffness matrix extracted. Macroscale numerical modeling of scale vibrodynamics showed close qualitative and quantitative agreement with scanning laser Doppler vibrometry measurement of this scale's oscillations, suggesting that the governing biomechanics have been captured accurately. Importantly, this scale of B. alcinoe exhibits its first three resonances in the typical echolocation frequency range of bats, suggesting it has evolved as a resonant absorber. Damping coefficients of the moth-scale resonator and ultrasonic absorption of a scaled wing were estimated using numerical modeling. The calculated absorption coefficient of 0.50 agrees with the published maximum acoustic effect of wing scaling. Understanding scale vibroacoustic behavior helps create macroscopic structures with the capacity for broadband acoustic camouflage.

Neural representation of bat predation risk and evasive flight in moths: A modelling approach.

Most animals are at risk from multiple predators and can vary anti-predator behaviour based on the level of threat posed by each predator. Animals use sensory systems to detect predator cues, but the relationship between the tuning of sensory systems and the sensory cues related to predator threat are not well-studied at the community level. Noctuid moths have ultrasound-sensitive ears to detect the echolocation calls of predatory bats. Here, combining empirical data and mathematical modelling, we show that moth hearing is adapted to provide information about the threat posed by different sympatric bat species. First, we found that multiple characteristics related to the threat posed by bats to moths correlate with bat echolocation call frequency. Second, the frequency tuning of the most sensitive auditory receptor in noctuid moth ears provides information allowing moths to escape detection by all sympatric bats with similar safety margin distances. Third, the least sensitive auditory receptor usually responds to bat echolocation calls at a similar distance across all moth species for a given bat species. If this neuron triggers last-ditch evasive flight, it suggests that there is an ideal reaction distance for each bat species, regardless of moth size. This study shows that even a very simple sensory system can adapt to deliver information suitable for triggering appropriate defensive reactions to each predator in a multiple predator community.

Nathusius' bats optimize long-distance migration by flying at maximum range speed.

Aerial migration is the fastest, yet most energetically demanding way of seasonal movement between habitats. However, for many taxa, and bats in particular, we lack a clear understanding of the energy requirements for migration. Here, we examined the energetic cost and flight speed of the long-distance migratory Nathusius' bat (Pipistrellus nathusii). We measured flight metabolism in relation to airspeed in a wind tunnel, inferred the optimal traveling speed over long distances, i.e. maximum range speed, and compared this value with flight speed measured in wild conspecifics. Body mass and wing morphologies were similar in captive and wild bats, indicating that the body condition of captive bats was similar to that of migratory bats. Nine out of the 12 captive bats exhibited a U-shaped relationship between flight metabolic power and airspeed when flying in the wind tunnel. The flight metabolic rate across all airspeeds averaged 0.98±0.28 W, which corresponds well to established allometric relationships between flight metabolic rate and body mass for bats. During summer migration, P. nathusii traveled at an average speed of 6.9±0.7 m s-1, which was significantly higher than the minimum power speed (5.8±1.0 m s-1), yet within the range of expected maximum range speed inferred from wind tunnel experiments. This suggests that P. nathusii may migrate at an energetically optimal speed and that aerial refueling does not substantially lower migratory speed in P. nathusii.

Spatiotemporal distribution of individuals as an indicator for the social system of Lepilemur sahamalaza.

Primate social systems are highly diverse, complicating the classification of particularly elusive species that are difficult to observe. The spatial distribution of individuals over time is a critica lindicator for the social organization and long-term studies are important to establish patterns of social interactions. In recent years, species of the cryptic, nocturnal sportive lemurs of the genus Lepilemur were found to live in pairs in which a single male and a single female share and defend a mutual home range. The present study aimed to forward research into this underrepresented genus by determining the social organization and structure of the Sahamalaza sportive lemur, L. sahamalaza. We collected 773.15 hr of behavioral and GPS data during a period of 10 months (between 2015 and 2016) on 14 individuals: eight females and six males. There was no evidence of pair-specific home range use as intra- and intersexual home range overlap was high. No pattern of social interactions between focal individuals could be distinguished despite high range overlap. Individuals met and interacted infrequently, resulting in an interaction rate of 0.32 interactions/hr. Sleeping associations between adult individuals were never observed. While both sexes had access to multiple potential mating partners, range sizes or ranging distances did not increase in mating periods. Overall, the social system of Sahamalaza sportive lemurs exhibits aspects of a solitary social organization and structure with potential for the polygamous mating system. These findings underline the importance of detailed social ecology studies that can provide the basis for understanding potential environmental influences on social system variability of closely related species.

Sahamalaza Sportive Lemur, Lepilemur sahamalaza, Vocal Communication: Call Use, Context and Gradation.

Stereotypical vocalisations can facilitate long-distance communication in dense and, thus, sound-degrading forest habitats. Despite this, primate vocal repertoires often also include gradations between different call types that are used in social interactions. Because many nocturnal primates show a solitary social structure, it has been difficult to assess the role vocalisations play in mediating their social encounters. Here we aim to expand on the call types of Sahamalaza sportive lemurs, Lepilemur sahamalaza, as well as investigate their variability and use in social contexts. Through long-term behavioural observations with concurrent recording of vocalisations of known and unknown individuals between 2015 and 2016, we described three previously unrecorded calls and expanded the vocal repertoire to 12 different call types that were used in both targeted social and broadcast solitary contexts. Linear discriminant function and cluster analysis supported initial observations that vocalisations within call types were variable and that at least two call types can be classified as "graded," contrary to expectations. Despite variations in call-context connections, no clear patterns of call use could be established. However, measurable differences in vocal behaviour between the seasons and the sexes indicate that calling is used in a reproductive context, similar to other nocturnal and diurnal primates.

Ultrasound avoidance by flying antlions (Myrmeleontidae).

The acoustic arms race between insectivorous bats and their invertebrate prey has led to the convergent evolution of ultrasound hearing in seven orders of nocturnal insects. Upon hearing the echolocation calls of an approaching bat, such insects take defensive action. Here, we document a previously unknown sense of ultrasound hearing and phonotactic flight behaviour in the neuropteran family Myrmeleontidae (antlions). The antlion Myrmeleon hyalinus was presented with sound pulses at ultrasonic frequencies used by echolocating bats and its response thresholds in tethered flight determined. Behaviours included abdominal twitches, wing flicks, brief pauses in flight and flight cessation. Such behaviours create erratic evasive flight manoeuvres in other eared insects, particularly mantids and lacewings. Antlions responded best to ultrasound between 60 and 80 kHz (75 dB peSPL at 80 kHz), showing response thresholds similar to those of the related lacewings (Neuroptera, Chrysopidae). Yet, at lower ultrasonic frequencies (20-50 kHz), antlions were far less sensitive than lacewings. Based on calculated response distances, we conclude that antlions respond only after having been detected by bats rather than using early evasive flights. We argue that the high response threshold for low-frequency ultrasound is adaptive for an insect that is mainly active close to and within vegetation, because a behavioural response to the lower ultrasonic frequencies used by high-flying bats would result in evasive action in the absence of actual predation risk.

Sensorimotor Model of Obstacle Avoidance in Echolocating Bats.

Bat echolocation is an ability consisting of many subtasks such as navigation, prey detection and object recognition. Understanding the echolocation capabilities of bats comes down to isolating the minimal set of acoustic cues needed to complete each task. For some tasks, the minimal cues have already been identified. However, while a number of possible cues have been suggested, little is known about the minimal cues supporting obstacle avoidance in echolocating bats. In this paper, we propose that the Interaural Intensity Difference (IID) and travel time of the first millisecond of the echo train are sufficient cues for obstacle avoidance. We describe a simple control algorithm based on the use of these cues in combination with alternating ear positions modeled after the constant frequency bat Rhinolophus rouxii. Using spatial simulations (2D and 3D), we show that simple phonotaxis can steer a bat clear from obstacles without performing a reconstruction of the 3D layout of the scene. As such, this paper presents the first computationally explicit explanation for obstacle avoidance validated in complex simulated environments. Based on additional simulations modelling the FM bat Phyllostomus discolor, we conjecture that the proposed cues can be exploited by constant frequency (CF) bats and frequency modulated (FM) bats alike. We hypothesize that using a low level yet robust cue for obstacle avoidance allows bats to comply with the hard real-time constraints of this basic behaviour.

Delayed response and biosonar perception explain movement coordination in trawling bats.

Animal coordinated movement interactions are commonly explained by assuming unspecified social forces of attraction, repulsion and alignment with parameters drawn from observed movement data. Here we propose and test a biologically realistic and quantifiable biosonar movement interaction mechanism for echolocating bats based on spatial perceptual bias, i.e. actual sound field, a reaction delay, and observed motor constraints in speed and acceleration. We found that foraging pairs of bats flying over a water surface swapped leader-follower roles and performed chases or coordinated manoeuvres by copying the heading a nearby individual has had up to 500 ms earlier. Our proposed mechanism based on the interplay between sensory-motor constraints and delayed alignment was able to recreate the observed spatial actor-reactor patterns. Remarkably, when we varied model parameters (response delay, hearing threshold and echolocation directionality) beyond those observed in nature, the spatio-temporal interaction patterns created by the model only recreated the observed interactions, i.e. chases, and best matched the observed spatial patterns for just those response delays, hearing thresholds and echolocation directionalities found to be used by bats. This supports the validity of our sensory ecology approach of movement coordination, where interacting bats localise each other by active echolocation rather than eavesdropping.

A whispering bat that screams: bimodal switch of foraging guild from gleaning to aerial hawking in the desert long-eared bat.

Echolocating bats have historically been classified as either loud aerial hawkers or whispering gleaners. Some bat species can forage in multiple ways and others have demonstrated limited flexibility in the amplitude of their echolocation calls. The desert long-eared bat, Otonycteris hemprichii, has been said to be a passive gleaning whispering bat preying on terrestrial arthropods such as scorpions. Using an acoustic tracking system, we recorded individuals flying at foraging and drinking sites and compared their flight height, flight speed, call duration, pulse interval and source levels with those of gleaning individuals previously recorded using the same setup. We found differences in all variables with the strongest difference in source levels, where bats called at a mean of 119 dB peSPL (compared with 75 dB peSPL when gleaning). Bat faecal analysis indicated that their diet differed from previous studies and that prey species were capable of flight. We conclude that the bats switched from passive gleaning to capturing airborne insects (aerial hawking). Although whispering bats have been known to opportunistically catch insects on the wing, in the present study we show a full bimodal switch between foraging guilds with the respective changes in source level to those typical of a true aerial hawker.

Forest fragmentation and edge effects impact body condition, fur condition and ectoparasite prevalence in a nocturnal lemur community.

Forest fragmentation and edge effects are two major threats to primate populations. Primates inhabiting fragmented landscapes must survive in a more degraded environment, often with lower food availability compared to continuous forests. Such conditions can have deleterious effects on animal physiological health, yet some primates thrive in these habitats. Here, we assessed how forest fragmentation and associated edge effects impact three different components of physiological health in a nocturnal primate community in the Sahamalaza-Iles Radama National Park, northwest Madagascar. Over two periods, 6 March 2019-30 October 2019 and 10 January 2022-17 May 2022, we collected data on body condition, fur condition scores and ectoparasite prevalence for 125 Mirza zaza, 51 Lepilemur sahamalaza, 27 Cheirogaleus medius and 22 Microcebus sambiranensis individuals, and we compared these metrics between core and edge areas of continuous forest and fragmented forest. Body condition scores for all species varied between areas, with a positive response to fragmentation and edge effects observed for M. zaza and L. sahamalaza and a negative response for C. medius and M. sambiranensis. Fur condition scores and ectoparasite prevalence were less variable, although M. zaza and L. sahamalaza had a significantly negative response to fragmentation and edge effects for these two variables. Interestingly, the impacts of fragmentation and edge effects on physiological health were variable-specific. Our results suggest that lemur physiological responses to fragmentation and edge effects are species-specific, and body condition, fur condition and ectoparasite prevalence are impacted in different ways between species. As other ecological factors, including food availability and inter/intraspecific competition, likely also influence physiological health, additional work is required to determine why certain aspects of lemur physiology are affected by environmental stressors while others remain unaffected. Although many nocturnal lemurs demonstrate resilience to fragmented and degraded habitats, urgent conservation action is needed to safeguard the survival of their forest habitats.

The simple ears of noctuoid moths are tuned to the calls of their sympatric bat community.

Insects with bat-detecting ears are ideal animals for investigating sensory system adaptations to predator cues. Noctuid moths have two auditory receptors (A1 and A2) sensitive to the ultrasonic echolocation calls of insectivorous bats. Larger moths are detected at greater distances by bats than smaller moths. Larger moths also have lower A1 best thresholds, allowing them to detect bats at greater distances and possibly compensating for their increased conspicuousness. Interestingly, the sound frequency at the lowest threshold is lower in larger than in smaller moths, suggesting that the relationship between threshold and size might vary across frequencies used by different bat species. Here, we demonstrate that the relationships between threshold and size in moths were only significant at some frequencies, and these frequencies differed between three locations (UK, Canada and Denmark). The relationships were more likely to be significant at call frequencies used by proportionately more bat species in the moths' specific bat community, suggesting an association between the tuning of moth ears and the cues provided by sympatric predators. Additionally, we found that the best threshold and best frequency of the less sensitive A2 receptor are also related to size, and that these relationships hold when controlling for evolutionary relationships. The slopes of best threshold versus size differ, however, such that the difference in threshold between A1 and A2 is greater for larger than for smaller moths. The shorter time from A1 to A2 excitation in smaller than in larger moths could potentially compensate for shorter absolute detection distances in smaller moths.

A test of the senses: fish select novel habitats by responding to multiple cues.

Habitat-specific cues play an important role in orientation for animals that move through a mosaic of habitats. Environmental cues can be imprinted upon during early life stages to guide later return to adult habitats, yet many species must orient toward suitable habitats without previous experience of the habitat. It is hypothesized that multiple sensory cues may enable animals to differentiate between habitats in a sequential order relevant to the spatial scales over which the different types of information are conveyed, but previous research, especially for marine organisms, has mainly focused on the use of single cues in isolation. In this study, we investigated novel habitat selection through the use of three different sensory modalities (hearing, vision, and olfaction). Our model species, the French grunt, Haemulon flavolineatum, is a mangrove/seagrass-associated reef fish species that makes several habitat transitions during early life. Using several in situ and ex situ experiments, we tested the response of fish toward auditory, olfactory, and visual cues from four different habitats (seagrass beds, mangroves, rubble, and coral reef). We identified receptivity to multiple sensory cues during the same life phase, and found that different cues induced different reactions toward the same habitat. For example, early-juvenile fish only responded to sound from coral reefs and to chemical cues from mangroves/seagrass beds, while visual cues of conspecifics overruled olfactory cues from mangrove/seagrass water. Mapping these preferences to the ecology of ontogenetic movements, our results suggest sequential cue use would indeed aid successful orientation to novel key habitats in early life.

Detection distances in desert dwelling, high duty cycle echolocators: A test of the foraging habitat hypothesis.

High Duty Cycle (HDC) echolocating bats use high frequency echolocation pulses that are clutter resistant, but their high frequencies give them limited range. Despite their unique ability to reject background clutter while simultaneously detecting fluttering prey, the frequency of their echolocation pulses has a strong correlation with level of environmental clutter, lower frequency pulses of HDC bats being associated with more open environments. The Foraging Habitat Hypothesis (FHH) proposes that the ecological significance of these lower frequency pulses in HDC bats in open environments is that they allow longer prey detection distances. To test the FHH, we compared the frequencies, Source Levels (SLs) and detection distances of Rhinolophus capensis, a HDC bat that has been shown to vary its call frequency in relation to habitat structure. As a further test of the FHH we investigated the SLs and detection distances of Rhinolophus damarensis (a heterospecific species that occurs in the same open desert environment as R. capensis but echolocates at a higher dominant pulse frequency). In the open desert, R. capensis emitted both lower frequency and higher SL pulses giving them longer detection distances than R. capensis in the cluttered fynbos. SL contributed more to differences in detection distances in both R. capensis and R. damarensis than frequency. In a few instances, R. damarensis achieved similar detection distances to desert-inhabiting R. capensis by emitting much higher SLs despite their average SLs being lower. These results suggest that lower frequency echolocation pulses are not a prerequisite for open desert living but may increase detection distance while avoiding energetic costs required for high SLs.

Moth wings as sound absorber metasurface.

In noise control applications, a perfect metasurface absorber would have the desirable traits of not only mitigating unwanted sound, but also being much thinner than the wavelengths of interest. Such deep-subwavelength performance is difficult to achieve technologically, yet moth wings, as natural metamaterials, offer functionality as efficient sound absorbers through the action of the numerous resonant scales that decorate their wing membrane. Here, we quantify the potential for moth wings to act as a sound-absorbing metasurface coating for acoustically reflective substrates. Moth wings were found to be efficient sound absorbers, reducing reflection from an acoustically hard surface by up to 87% at the lowest frequency tested (20 kHz), despite a thickness to wavelength ratio of up to 1/50. Remarkably, after the removal of the scales from the dorsal surface the wing's orientation on the surface changed its absorptive performance: absorption remains high when the bald wing membrane faces the sound but breaks down almost completely in the reverse orientation. Numerical simulations confirm the strong influence of the air gap below the wing membrane but only when it is adorned with scales. The finding that moth wings act as deep-subwavelength sound-absorbing metasurfaces opens the door to bioinspired, high-performance sound mitigation solutions.

Hemprich's long-eared bat (Otonycteris hemprichii) as a predator of scorpions: whispering echolocation, passive gleaning and prey selection.

Over 70% of the droppings of the gleaning bat Otonycteris hemprichii can contain scorpion fragments. Yet, some scorpions found in its desert habitat possess venom of the highest known toxicity, rendering them a very dangerous prey. In this study, we describe how O. hemprichii catches and handles scorpions, quantify its flight and echolocation behaviour in the field, investigate what sensory modality it uses to detect scorpions, and test whether it selects scorpions according to their size or toxicity. We confirmed that O. hemprichi is a whispering bat (approx. 80 dB peSPL) with short, multi-harmonic calls. In a flight room we also confirmed that O. hemprichii detects scorpions by their walking noises. Amplitudes of such noises were measured and they reach the flying bat at or below the level of echoes of the loess substrate. Bats dropped straight onto moving scorpions and were stung frequently even straight in their face. Stings did not change the bats' behaviour and caused no signs of poisoning. Scorpions were eaten including poison gland and stinger. Bats showed no preference neither for any of the scorpion species nor their size suggesting they are generalist predators with regard to scorpions.