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The development of increasingly affordable ultrasonic detectors and automatic classifiers has increasingly boosted the use of acoustic recording of echolocation calls to survey bats all over the world. Echolocation call keys are crucial to reliably classify acoustic recordings, but those are not available for many regions, such as China. In the present study, we conducted a systematic review of bat echolocation studies across continental China and developed an acoustic identification key. Based on 130 studies, published from 1999 to 2020, we obtained echolocation parameters from 64 bat species (47.4% of the total echolocating species known from the country). This review highlights the lack of echolocation references from continental China for 71 species, from which, the echolocation of 21 has never been described. Additionally, we developed the Bat Knowledge Index, an indicator that allows the identification of key priority areas for future research on bat acoustics and geographic biases in the existing studies, further highlighting the lack of studies in the northwestern regions of the country. The compiled identification key provides easy-to-follow identification steps to classify the echolocation calls of 114 species (84.5% of all Chinese echolocating species) to either species (11.4%) or phonic group level (88.6%). This key represents a useful baseline tool and a further step to increase bioacoustic studies’ suitability in China.

Artificial lights may be altering interactions between bats and moth prey. According to the allotonic frequency hypothesis (AFH), eared moths are generally unavailable as prey for syntonic bats (i.e., bats that use echolocation frequencies between 20 and 50 kHz within the hearing range of eared moths) due to the moths' ability to detect syntonic bat echolocation. Syntonic bats therefore feed mainly on beetles, flies, true bugs, and non-eared moths. The AFH is expected to be violated around lights where eared moths are susceptible to exploitation by syntonic bats because moths' evasive strategies become less effective. The hypothesis has been tested to date almost exclusively in areas with permanent lighting, where the effects of lights on bat diets are confounded with other aspects of human habitat alteration. We undertook diet analysis in areas with short-term, localized artificial lighting to isolate the effects of artificial lighting and determine if syntonic and allotonic bats (i.e., bats that use echolocation frequencies outside the hearing range of eared moths) consumed more moths under conditions of artificial lights than in natural darkness. We found that syntonic bats increased their consumption of moth prey under experimentally lit conditions, likely owing to a reduction in the ability of eared moths to evade the bats. Eared moths may increase in diets of generalist syntonic bats foraging around artificial light sources, as opposed to allotonic species and syntonic species with a more specialized diet.

Understanding the evolutionary origins of communication signals requires careful study of multiple species within a known phylogenetic framework. Most cricket species produce low-frequency calls for mate attraction, whereas they startle to high-frequency sounds similar to bat echolocation. Male crickets in the tribe Lebinthini produce high-frequency calls, to which females reply with vibrational signals. This novel communication system likely evolved by male sensory exploitation of acoustic startle to high-frequency sounds in females. This behavior was previously described for the Lebinthini from Asia. Here we demonstrate that this novel communication system is found in a Neotropical species, Ponca hebardi, and is therefore likely shared by the whole tribe Lebinthini, dating the origin of this behavior to coincide with the origin of echolocation in bats. Furthermore, we document male duets involving both acoustic and vibratory signals not previously described in crickets, and we tentatively interpret it as competitive masking between males.

Molossops temminckii is the only species of the Molossidae capable of exploring background-cluttered space. We tested the hypothesis that M. temminckii uses the clutter rejection strategy in cluttered environments and determined whether the echolocation calls of this species show large-scale geographic variation. We recorded echolocation calls of 10 individuals of M. temminckii using 4 treatments with different degrees of clutter. We compared individuals recorded in Venezuela with individuals recorded in Brazil to verify geographic variation in calls. In the cluttered environments, M. temminckii emitted short downward frequency-modulated calls with short pulse intervals. In the uncluttered environments, M. temminckii emitted long upward frequency-modulated calls with longer intervals and shorter bandwidth. The decreased pulse duration and interval in cluttered environments support the hypothesis that M. temminckii uses the clutter rejection strategy. Additionally, we found that in Brazil, M. temminckii emits shorter calls with higher minimum frequency compared with bats recorded in Venezuela, indicating large-scale geographic variation, probably because of environmental factors.Molossops temminckii e a unica especie de Molossidae com capacidade de explorar areas com obstaculos de fundo. Nesse trabalho testamos a hipotese do uso da estrategia de rejeicao de obstaculo em ambientes fechados, e determinamos se ha variacao geografica de larga escala nos chamados de ecolocalizacao de M. temminckii. Gravamos 10 individuos, em 4 tratamentos, com diferentes graus de obstaculos. Comparamos os chamados de ecolocalizacao gravados na Venezuela e Brasil para verificar a variacao geografica. Em ambientes com obstaculos, M. temminckii emite chamados curtos, de frequencia modulada descendente, com intervalos curtos entre os chamados. Nos ambientes abertos, emite chamados longos, de frequencia modulada ascendente, com intervalos mais longos e menor largura de banda. A menor duracao e o menor intervalo em ambientes fechados suportam a hipotese que M. temminckii utiliza a estrategia de rejeicao de obstaculo. No Brasil, M. temminckii emite chamados mais curtos, com frequencia minima mais alta, em comparacao com os morcegos gravados na Venezuela. Esse fato indica variacao geografica de larga escala e fatores ambientais provavelmente contribuem para essa variacao.

Recent technological advances have permitted collection of immense data sets through automated recordings that are primarily aimed at capturing bat echolocation. Analyses of echolocation calls are used to identify species, relative abundance, and some aspects of behaviour, such as foraging or commuting. Here we propose that social calls recorded in flight are also valuable tools for understanding bat ecology and behaviour. First, we examine how and why the acoustic structure of social calls differ from echolocation. Differences in form make social calls often, but not always, easy to identify. We then use a case study on in-flight song in Brazilian free-tailed bat (Tadarida brasiliensis (I. Geoffroy, 1824)) to show that what may appear as echolocation may instead be predominantly used for social communication. Next, we review three basic functions of in-flight social calls, including examples of each, and develop a framework for testing these alternative functions using automated recordings. In a second case study, we use automated recordings of the endangered Florida bonneted bat (Eumops floridanus (G.M. Allen, 1932)) to illustrate how behavioural information can be gleaned by examining patterns of social call production. Finally, we discuss why and how social calls provide novel information that can be crucial for conservation and management efforts.

Automated recording units are frequently used for passive acoustic monitoring of taxa, including bats. Detection and species-level identification of bat echolocation calls using manual techniques is a common practice, often supplementing automated analysis by software. However, few standardized protocols exist for manual analysis, which is challenging for novices and impedes comparisons among research groups. In this two-part review, I first summarize and synthesize current approaches to manual call analysis. Three observations about the processes used to conduct manual call identification emerge: (1) there are significant knowledge gaps and few comparisons of interoperator variability; (2) they are individual- and location-specific, with no standardized underlying framework; and (3) they are often not well-described in the peer-reviewed literature. In response to these observations, I then conduct a comparative analysis of the fields of clinical reasoning (the study of medical decision-making) and the identification of bat echolocation calls. Clinical reasoning is a mature area of research and findings from this field may inform practices and instructional strategies for manually identifying echolocation calls. I demonstrate similarities between clinical reasoning and call identification processes and then make recommendations on how to apply findings from the clinical reasoning literature to call identification practices and training.

Bioacoustic research has made several advancements in developing systems to record extensive acoustic data and classify bat echolocation calls to species level using automated classifiers. ...

Echolocating bats are regularly studied to investigate auditory‐guided behaviors and as important bioindicators. Bioacoustic monitoring methods based on echolocation calls are increasingly used for risk assessment and to ultimately inform conservation strategies for bats. As echolocation calls transmit through the air at the speed of sound, they undergo changes due to atmospheric and geometric attenuation. Both the speed of sound and atmospheric attenuation, however, are variable and determined by weather conditions, particularly temperature and relative humidity. Changing weather conditions thus cause variation in analyzed call parameters, limiting our ability to detect, and correctly analyze bat calls. Here, I use real‐world weather data to exemplify the effect of varying weather conditions on the acoustic properties of air. I then present atmospheric attenuation and speed of sound for the global range of weather conditions and bat call frequencies to show their relative effects. Atmospheric attenuation is a nonlinear function of call frequency, temperature, relative humidity, and atmospheric pressure. While atmospheric attenuation is strongly positively correlated with call frequency, it is also significantly influenced by temperature and relative humidity in a complex nonlinear fashion. Variable weather conditions thus result in variable and unknown effects on the recorded call, affecting estimates of call frequency and intensity, particularly for high frequencies. Weather‐induced variation in speed of sound reaches up to about ±3%, but is generally much smaller and only relevant for acoustic localization methods of bats. The frequency‐ and weather‐dependent variation in atmospheric attenuation has a threefold effect on bioacoustic monitoring of bats: It limits our capability (1) to monitor bats equally across time, space, and species, (2) to correctly measure frequency parameters of bat echolocation calls, particularly for high frequencies, and (3) to correctly identify bat species in species‐rich assemblies or for sympatric species with similar call designs.

Predation on swarms of prey, especially using visual information, has drawn much interest in studies of collective movement. Surprisingly, in the field of biosonar this aspect of prey detection, which is probably very common, has received little to no attention. Here, we combine computer simulations and actual echo measurements to accurately estimate the echo sound pressure of insect swarms of different size and density. We show that swarm echo sound pressure increases with 3dB for every doubling of insect number, irrespective of swarm density. Thus swarms will be much easier to detect than single insects. Many of the insects bats eat are so small that they are only detectable by echolocation at very short distances. By focusing on detection of swarms of insects, a bat may increase its operating range and diversify its diet. Interestingly, interference between the sound waves reflected from a swarm of insects can sometimes result in echoes that are much weaker than echoes from single insects. We show that bats can reduce this problem by increasing the bandwidth of their echolocation calls. Specifically, a bandwidth of 3–8 kHz would guarantee receiving loud echoes from any angle relative to the swarm. Indeed, many bat species, and specifically bats hunting in open spaces, where swarms are abundant, use echolocation signals with a bandwidth of several kHz. Our results might also explain how the first echolocating bats that probably had limited echolocation abilities, could detect insects through swarm hunting., Author summary When bats hunt, they often encounter insects that fly in swarms. Echolocating bats emit sonar signals to search for prey and it is currently unknown what such swarms look like to a bat. Unlike vision, sonar senses the delay or distance to objects directly. We show that when bats hunt for insects in the sky, the echoes from the insects in a swarm will most of the time sum up and therefore become much louder than the echo of a single insect. Every time an insect swarm would double in number, a bat would hear an echo that is 3dB stronger. This could enable a bat to detect prey from longer distances and some bats might thus profit from swarm hunting. However, the echoes reflected from the many insects in the swarm also create acoustic interference so that sometimes the summed echo is actually weak at a certain frequency. We show how bats could deal with this drawback. It is known that most bats do not use sonar signals with a single tone but that they modulate their tones. Our analysis shows that this modulation can solve the problem of spectral interference ensuring that the swarm-echo is always loud.

Bat acoustic libraries are important tools that assemble echolocation calls to allow the comparison and discrimination to confirm species identifications. The Sonozotz project represents the first nation‐wide library of bat echolocation calls for a megadiverse country. It was assembled following a standardized recording protocol that aimed to cover different recording habitats, recording techniques, and call variation inherent to individuals. The Sonozotz project included 69 species of echolocating bats, a high species richness that represents 50% of bat species found in the country. We include recommendations on how the database can be used and how the sampling methods can be potentially replicated in countries with similar environmental and geographic conditions. To our knowledge, this represents the most exhaustive effort to date to document and compile the diversity of bat echolocation calls for a megadiverse country. This database will be useful to address a range of ecological questions including the effects of anthropogenic activities on bat communities through the analysis of bat sound., The Sonozotz project represents the first nation‐wide library of bat echolocation calls for a megadiverse country.

Bats are a species-rich order of mammals providing key ecosystem services. Because bats are threatened by human action and also serve as important bioindicators, monitoring their populations is of utmost importance. However, surveying bats is difficult because of their nocturnal habits, elusiveness and sensitivity to disturbance. Bat detectors allow echolocating bats to be surveyed non-invasively and record species that would otherwise be difficult to observe by capture or roost inspection. Unfortunately, several bat species cannot be identified confidently from their calls so acoustic classification remains ambiguous or impossible in some cases. The popularity of automated classifiers of bat echolocation calls has escalated rapidly, including that of several packages available on purchase. Such products have filled a vacant niche on the market mostly in relation to the expanding monitoring efforts related to the development of wind energy production worldwide. We highlight that no classifier has yet proven capable of providing correct classifications in 100% of cases or getting close enough to this ideal performance. Besides, from the literature available and our own experience we argue that such tools have not yet been tested sufficiently in the field. Visual inspection of calls whose automated classification is judged suspicious is often recommended, but human intervention a posteriori represents a circular argument and requires noticeable experience. We are concerned that neophytes – including consultants with little experience with bats but specialized into other taxonomical groups – will accept passively automated responses of tools still awaiting sufficient validation. We remark that bat call identification is a serious practical issue because biases in the assessment of bat distribution or habitat preferences may lead to wrong management decisions with serious conservation consequences. Automated classifiers may crucially aid bat research and certainly merit further investigations but the boost in commercially available software may have come too early. Thorough field tests need to be carried out to assess limitations and strengths of these tools.

This study was supported by the Carlsberg Foundation via a Semper Ardens grant, ONR, N00014-17-1- 2736; AFOSR FA9550-14-1-0398, and NSF NCS-FO:1734744 and a Human Frontiers Science Program Long-Term Fellowship to AS. These experiments were approved by The Danish Council for Experiments on Animals under permit number: 2016-15-0201-00989 and by the Johns Hopkins University Animal Care and Use Committee under protocol number BA17A107. We thank Uwe Firzlaff and Lutz Wiegrebe for their help. 1. To study sensorimotor behaviour in wild animals, it is necessary to synchronously record the sensory inputs available to the animal, and its movements. To do this, we have developed a biologging device that can record the primary sensory information and the associated movements during foraging and navigating in echolocating bats. 2. This 2.6 -gram tag records the sonar calls and echoes from an ultrasonic microphone, while simultaneously sampling fine-scale movement in three dimensions from wideband accelerometers and magnetometers. In this study, we tested the tag on an European noctula (Nyctalus noctula) during target approaches and on four big brown bats (Eptesicus fuscus) during prey interception in a flight room. 3. We show that the tag records both the outgoing calls and echoes returning from objects at biologically relevant distances. Inertial sensor data enables the detection of behavioural events such as flying, turning, and resting. In addition, individual wing-beats can be tracked and synchronized to the bat's sound emissions to study the coordination of different motor events. 4. By recording the primary acoustic flow of bats concomitant with associated behaviours on a very fine time-scale, this type of biologging method will foster a deeper understanding of how sensory inputs guide feeding behaviours in the wild. Postprint

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.

Echolocation calls produced by bats in their larynges allow these flying, nocturnal mammals to orient and find food at night. The acoustic signals are not like bird song, and even individual bats exhibit great flexibility in call design and between-species overlap is common. As a result, identifying bats to species by their echolocation calls even in communities with few bat species can be difficult. Unfortunately, the situation is worse still. As a result of several factors — some to do with microphones, some with environment, some with bats, and the calls themselves — acoustic information transmitted to and transduced by microphones can be dramatically different from the actual signal produced by the bat and as would be recorded on axis, close to its mouth using ideal microphones under ideal conditions. We outline some of these pitfalls and discuss ways to make the best of a bad situation. Overall, however, we stress that many of these factors cannot be ignored and do impact our recordings.

Active echolocation enables bats to orient and hunt the night sky for insects. As a counter-measure against the severe predation pressure many nocturnal insects have evolved ears sensitive to ultrasonic bat calls. In moths bat-detection was the principal purpose of hearing, as evidenced by comparable hearing physiology with best sensitivity in the bat echolocation range, 20–60 kHz, across moths in spite of diverse ear morphology. Some eared moths subsequently developed sound-producing organs to warn/startle/jam attacking bats and/or to communicate intraspecifically with sound. Not only the sounds for interaction with bats, but also mating signals are within the frequency range where bats echolocate, indicating that sound communication developed after hearing by “sensory exploitation”. Recent findings on moth sound communication reveal that close-range (~ a few cm) communication with low-intensity ultrasounds “whispered” by males during courtship is not uncommon, contrary to the general notion of moths predominantly being silent. Sexual sound communication in moths may apply to many eared moths, perhaps even a majority. The low intensities and high frequencies explain that this was overlooked, revealing a bias towards what humans can sense, when studying (acoustic) communication in animals.

Quantitative methods for species identification are commonly used in acoustic surveys for animals. While various identification models have been studied extensively, there has been little study of methods for selecting calls prior to modeling or methods for validating results after modeling. We obtained two call libraries with a combined 1556 pulse sequences from 11 North American bat species. We used four acoustic filters to automatically select and quantify bat calls from the combined library. For each filter, we trained a species identification model (a quadratic discriminant function analysis) and compared the classification ability of the models. In a separate analysis, we trained a classification model using just one call library. We then compared a conventional model assessment that used the training library against an alternative approach that used the second library. We found that filters differed in the share of known pulse sequences that were selected (68 to 96%), the share of non-bat noises that were excluded (37 to 100%), their measurement of various pulse parameters, and their overall correct classification rate (41% to 85%). Although the top two filters did not differ significantly in overall correct classification rate (85% and 83%), rates differed significantly for some bat species. In our assessment of call libraries, overall correct classification rates were significantly lower (15% to 23% lower) when tested on the second call library instead of the training library. Well-designed filters obviated the need for subjective and time-consuming manual selection of pulses. Accordingly, researchers should carefully design and test filters and include adequate descriptions in publications. Our results also indicate that it may not be possible to extend inferences about model accuracy beyond the training library. If so, the accuracy of acoustic-only surveys may be lower than commonly reported, which could affect ecological understanding or management decisions based on acoustic surveys.

In their 1960 paper about bats using echolocation to find and track flying insects, Donald R. Griffin, Fredric A. Webster and Charles R. Michael (Animal Behaviour, 8, 141–154) changed the face of research on this behaviour. They moved the field of echolocation from documenting that this animal or that one could echolocate to demonstrating an adaptive value of echolocation. They used experiments with captive bats, fruit flies, mosquitoes and crane flies to illustrate how bats used a ‘feeding buzz’ as they closed with their prey. The topic remains current today, and one of the first papers in Nature in 2013 (Jacobsen et al., 493, 93–96) presented more information about feeding buzzes building on the platform that Griffin et al. had established. In the intervening period, literally thousands of papers have been published about echolocation, demonstrating how curious minds, technological advances and basic information about natural history can result in diversification of a field of research. We have learned that bats can use echolocation to recognize water surfaces and to find insect prey on spider webs. The continuum between orientation and social functions of echolocation means that this behaviour not only influences foraging and negotiating obstacle paths, but is also a cue that brings individuals together. Acoustic wars between bats and potential insect prey have further enriched the discipline by identifying acoustic measures and countermeasures used by the players. Parallel studies with toothed whales have provided further examples of the enrichment that echolocation brings to the lives of animals and those who study them.

We examined bat collision mortality, activity and species composition at an 89-turbine wind resource area in farmland of north-central Iowa from mid-Apr. to mid-Dec., 2003 and mid-Mar. to mid-Dec., 2004. We found 30 bats beneath turbines on cleared ground and gravel access areas in 2003 and 45 bats in 2004. After adjusting for search probability, search efficiency and scavenging rate, we estimated total bat mortality at 396 ± 72 (95% ci) in 2003 and 636 ± 112 (95% ci) in 2004. Although carcasses were mostly migratory tree bats, we found a considerable proportion of little brown bats (Myotis lucifugus). We recorded 1465 bat echolocation call files at turbine sites ( = 34.88 call files/detector-night) and 1536 bat call files at adjacent non-turbine sites ( = 36.57 call files/detector-night). Bat activity did not differ significantly between turbine and non-turbine sites. A large proportion of recorded call files were made by Myotis sp. but this may be because we detected activity at ground leve...

Ultrasonic detectors are powerful tools for the study of bat ecology. Many options are available for deploying acoustic detectors including various weatherproofing designs and microphone orientations, but the impacts of these options on the quantity and quality of the bat calls that are recorded are unknown. We compared the impacts of three microphone orientations (horizontal, 45°, and vertical) and two weatherproofing designs (polyvinyl chloride tubes and the BatHat) on the number of calls detected, call quality, and species detected by the Anabat II bat detector system at 17 sites in central Kentucky in May and June 2008. Detectors with BatHat weatherproofing recorded significantly fewer call sequences, pulses per file, species per site, and lower quality calls. Detectors in the horizontal position also tended to record fewer files, fewer species, and lower quality calls. These results illustrate potential impacts of deployment method on quality and quantity of data obtained. Because weatherproofing and orientation impacted the quality and quantity of data recorded, comparison of results using different methodologies should be made with caution.

Ecological constraints often shape the echolocation pulses emitted by bat species. Consequently some (but not all) bats emit species-specific echolocation pulses. Because echolocation pulses are often intense and emitted at high rates, they are potential targets for eavesdropping by other bats. Echolocation pulses can also vary within species according to sex, body size, age, social group and geographic location. Whether these features can be recognised by other bats can only be determined reliably by playback experiments, which have shown that echolocation pulses do provide sufficient information for the identification of sex and individual in one species. Playbacks also show that bats can locate conspecifics and heterospecifics at foraging and roost sites by eavesdropping on echolocation pulses. Guilds of echolocating bat species often partition their use of pulse frequencies. Ecology, allometric scaling and phylogeny play roles here, but are not sufficient to explain this partitioning. Evidence is accumulating to support the hypothesis that frequency partitioning evolved to facilitate intraspecific communication. Acoustic character displacement occurs in at least one instance. Future research can relate genetic population structure to regional variation in echolocation pulse features and elucidate those acoustic features that most contribute to discrimination of individuals.

Automated remote ultrasound detectors allow large amounts of data on bat presence and activity to be collected. Processing of such data involves identifying bat species from their echolocation calls. Automated species identification has the potential to provide more consistent, predictable, and potentially higher levels of accuracy than identification by hu- mans. In contrast, identification by humans permits flexibility and intelligence in identification, as well as the incorpora- tion of features and patterns that may be difficult to quantify. We compared humans with artificial neural networks (ANNs) in their ability to classify short recordings of bat echolocation calls of variable signal to noise ratios; these sequen- ces are typical of those obtained from remote automated recording systems that are often used in large-scale ecological studies. We presented 45 recordings (1-4 calls) produced by known species of bats to ANNs and to 26 human participants with 1 month to 23 years of experience in acoustic identification of bats. Humans correctly classified 86% of recordings to genus and 56% to species; ANNs correctly identified 92% and 62%, respectively. There was no significant difference between the performance of ANNs and that of humans, but ANNs performed better than about 75% of humans. There was little relationship between the experience of the human participants and their classification rate. However, humans with

SUMMARYThrough staged free-flight encounters between echolocating bats and praying mantids, we examined the effectiveness of two potential predator-evasion behaviors mediated by different sensory modalities: (1) power dive responses triggered by bat echolocation detected by the mantis ultrasound-sensitive auditory system, and (2) `last-ditch' maneuvers triggered by bat-generated wind detected by the mantis cercal system. Hearing mantids escaped more often than deafened mantids (76% vs 34%, respectively; hearing conveyed 42%advantage). Hearing mantis escape rates decreased when bat attack sequences contained very rapid increases in pulse repetition rates (escape rates 16 p.p.s. 10 ms–1; escape rates>60% for transition slopes 16 p.p.s. 10 ms–1) could circumvent mantis/insect auditory defenses. However, echolocation attack sequences containing such transitions occurred in only 15% of the trials. Since mantis ultrasound-mediated responses are not 100% effective, cercal-mediated evasive behaviors triggered by bat-generated wind could be beneficial as a backup/secondary system. Although deafened mantids with functioning cerci did not escape more often than deafened mantids with deactivated cerci (35%vs 32%, respectively), bats dropped mantids with functioning cerci twice as frequently as mantids with deactivated cerci. This latter result was not statistically reliable due to small sample sizes, since this study was not designed to fully evaluate this result. It is an interesting observation that warrants further investigation, however, especially since these dropped mantids always survived the encounter.

Big brown bats (Eptesicus) emit multiple-harmonic FM sounds (FM1, FM2) and exploit the relative weakening of higher harmonics in lowpass echoes from the surrounding scene to suppress clutter by defocusing of wideband images. Only echoes from a frontally located targets arrive as unfiltered, focused images. Experiments using electronically generated echoes show that lowpass filtering of masking echoes causes clutter masking to disappear. Lowpass filtering induces amplitude-latency trading, which retards response times at higher frequencies in clutter echoes relative to lower frequencies. Introducing countervailing changes in presentation-times of higher frequencies in electronically generated clutter echoes restores masking. In the big brown bat's inferior colliculus, FM sounds mimicking broadcasts and echoes evoke ~1 spike per sound at each neuron's best frequency; however, amplitude tuning is very broad. To exploit their high acuity for detecting coherence or non-coherence of echo responses, bats work in the latency domain instead, removing background objects through deliberate imposition of response de-synchronization and concomitant inattention on undesired clutter. Overall, the results indicate that big brown bats use neuronal response timing for virtually all auditory computations of echo delay, including clutter rejection. This use of active perceptual processes in biosonar instead of conventional sonar processes opens a new view toward biomimetic design.

Bats use echolocation calls that are suited for foraging in particular habitats. Many generalist bats alter their call design patterns as they move between open and spatially complex areas. This plasticity probably allows them to respond to the perceptual challenges posed by different structural environments. I tested the hypothesis that bats learn to produce these habitat-specific calls. I recorded the echolocation calls and foraging success of 14 juvenile little brown bats, Myotis lucifugus, as they foraged repeatedly in a large flight arena in two habitat treatments: open space and cluttered space. Calls were plastic across habitats. Furthermore, as bats gained experience, their foraging performance improved significantly in both open and cluttered habitat treatments. These improvements were related to changes in echolocation behaviour within the open habitat, which is consistent with the hypothesis that bats learn to produce more efficient calls in the open. Although the bats changed some aspects of their calls in the cluttered treatment, their use of space in the arena was the best predictor of foraging success. Bats that foraged in the centre of the arena were more successful. When bats switched habitat treatments, performance in the open was not affected by previous experience in the cluttered treatment. Bats that had previously experienced the open treatment, however, tended to use a more ‘open-area’ call when foraging in cluttered space. This result is discussed in light of habitat selection in young M. lucifugus.

A comparison of bat activity levels in the Coastal Plain of South Carolina among 5 habitat types: forested riparian areas, clearcuts, young pine plantations, mature pine plantations and pine savannas, using time expansion radio-microphones and integrated detectors to simultaneously monitor bat activity at three heights in each habitat type.

Ultrasonic hearing in moths evolved to detect bat echolocation cries and has been co-opted into the mating system of some species. Some moths evade or approach ultrasonic stimuli that overlap in their spectral and temporal features. The present study addressed the evaluation of ultrasonic stimuli by lesser wax- moths, a species in which ultrasonic hearing functions in bat evasion and pair formation. Playback experiments show that the role of different stimulus features varies with behavioural context. Flying moths have a single evasive response, which is performed mainly on the basis of pulse length and stimulus power: they dive upon hearing loud pulses � 1 ms in length, and respond more strongly to high-power stimuli above that pulse length. Female moths walking on the ground respond with positive phonotaxis mainly on the basis of pulse rate by orienting toward pulses delivered at rates � 30 pulses s � 1 . Consequently, moths on the

This review is yet another attempt to explain how echolocation in bats or bat-like mammals came into existence. Attention is focused on neuronal specializations in the ascending auditory pathway of echolocating bats. Three different mechanisms are considered that may create a specific auditory sensitivity to echos: (1) time-windows of enhanced echo-processing opened by a corollary discharge of neuronal vocalization commands; (2) differentiation and expansion of ensembles of combination-sensitive neurons in the midbrain; and (3) corticofugal top-down modulations. The second part of the review interprets three different types of echolocation as adaptations to ecological niches, and presents the sophisticated cochlear specializations in constant-frequency/frequency-modulated bats as a case study of finely tuned differentiation. It is briefly discussed how a resonant mechanism in the inner ear of constant-frequency/frequency-modulated bats may have evolved in common mammalian cochlea.

The responses of female Tettigonia viridissima to simulated bat echolocation calls were examined during tethered flight. The insects responded with three distinct behaviours, which occurred at graded stimulus intensities. At low intensities (threshold 54 dB SPL), T. viridissima responded by steering away from the sound source (negative phonotaxis). At intensities approximately 10 dB higher, beating of the hindwing was interrupted, although the insect remained in the flight posture. A diving response (cessation of the wingbeat, closure of the forewings and alignment of the legs against the body) occurred with a threshold of 76 dB SPL. Considering these thresholds, we estimate that the diving response occurs at approximately the sound amplitude at which many aerial-hawking bats first receive echoes from the insect. The other behaviours probably occur before the bat detects the insect and should therefore be interpreted as early avoidance behaviours. The repertoire of startle responses in T. viridissima, with directional and non-directional components, is similar to those of crickets and moths, but quite different from those described for another bushcricket (Neoconocephalus ensiger), which shows only a non-directional response. This supports the conclusion that bat-evasive behaviours are not conserved within the Tettigoniidae, but instead are shaped by the ecological constraints of the insects.

The tuning and pure-tone physiology of the T-cell prothoracic auditory interneuron were investigated in the nocturnal katydid Neoconocephalus ensiger. The T-cell is extremely sensitive and broadly tuned, particularly to high-frequency ultrasound (?:20 kHz). Adult thresholds were lowest and showed their least variability for frequencies ranging from 25 to 80 kHz. The average best threshold of the T-cell in N. ensiger ranged from 28 to 38 dB SPL and the best frequency from 20 to 27 kHz. In females, the T-cell is slightly more sensitive to the range of frequencies encompassing the spectrum of male song. Tuning of the T-cell in non-volant nymphs was comparable with that of adults, and this precocious ultrasound sensitivity supports the view that it has a role in the detection of terrestrial sources of predaceous ultrasound. In adults, T-cell tuning is narrower than that of the whole auditory (tympanic) organ, but only at audio frequencies. Superthreshold physiological experiments revealed that T-cell responses were ultrasound-biased, with intensity/response functions steeper and spike latencies shorter at 20, 30 and 40 kHz than at 5, 10 and 15 kHz. The same was also true for T-cell stimulation at 90 ° compared with stimulation at 0 ° within a frequency, which supports early T-cell research showing that excitation of the contralateral ear inhibits ipsilateral T-cell responses. In a temporal summation experiment, the integration time of the T-cell at 40 kHz (integration time constant τ=6.1 ms) was less than half that measured at 15 kHz (τ=15.0 ms). Moreover, T-cell spiking in response to short-duration pure-tone trains mimicking calling conspecifics (15 kHz) and bat echolocation hunting sequences (40 kHz) revealed that temporal pattern-copying was superior for ultrasonic stimulation. Apparently, T-cell responses are reduced or inhibited by stimulation with audio frequencies, which leads to the prediction that the T-cell will encode conspecific song less well than bat-like frequency-modulated sweeps during acoustic playback. The fact that the T-cell is one of the most sensitive ultrasound neurons in tympanate insects is most consistent with it serving an alarm, warning or escape function in both volant and non-volant katydids (nymphs and adults).

This study examines the contribution of GABAergic inhibition to the discharge pattern and pulse duration tuning characteristics of 101 bat inferior collicular neurons by means of bicuculline application to their recording sites. When stimulated with single pulses, 56 (55%) neurons discharged 1 or 2 impulses (phasic responders), 42 (42%) discharged 3–10 impulses (phasic bursters) and 3 (3%) discharged impulses throughout the stimulus duration (tonic responders). Bicuculline application increased the number of impulses and changed the discharge patterns of 66 neurons. Using 50% difference between maximal and minimal responses as a criterion, the duration tuning characteristics of these neurons can be described as band-pass (20, 20%), long-pass (17, 17%), short-pass (33, 32%), and all-pass (31, 31%). Each band-pass neuron discharged maximally to a specific duration (the best duration) which was at least 50% larger than the neuron's responses to a long-duration pulse and a short-duration pulse. In contrast, each long- or short-pass neuron discharged maximally to a range of long or short duration pulses. Bicuculline application changed the duration tuning characteristics of 65 neurons. Possible mechanisms underlying duration tuning characteristics and the behavioral relevance to bat echolocation are discussed.

One of the most efficient mechanisms to optimize signal-to-noise ratios is the Lombard effect - an involuntary rise in call amplitude due to ambient noise. It is often accompanied by changes in the spectro-temporal composition of calls. We examined the effects of broadband-filtered noise on the spectro-temporal composition of horseshoe bat echolocation calls, which consist of a constant-frequency component and an initial and terminal frequency-modulated portion. We find that the frequency-modulated components became larger for almost all noise conditions, whereas the bandwidth of the constant-frequency component increased only when broadband-filtered noise was centered on or above the calls' dominant or fundamental frequency. This indicates that ambient noise modifies the associated acoustic parameters of the Lombard-effect, such as spectro-temporal features, independently and could significantly affect the bat's ability to detect and locate targets. Our findings may be of significance in evaluating the impact of environmental noise on echolocation behavior in bats.

Bat detectors are commonly used to monitor bat behaviour. Earlier research has suggested that there may be systematic differences in the response of different detectors to bat calls. Such differences would have important implications for the comparability of quantitative surveys conducted with bat detectors. The present study examines variability within and between brands of bat detector in accuracy of tuning, directionality and sensitivity to different types of bat echolocation call in bat detectors from three manufacturers. The consistency of results from a field survey incorporating the three brands in a standardised methodology are also examined. Significant differences were found within and between brands in directionality and sensitivity which would lead to bias in bat surveys. The implications of these findings for bat surveys are discussed, as are the design features of importance for species identification.

Previous studies have shown that inferior collicular neurons of the big brown bat, Eptesicus fuscus, serve as short-, band-, long- and all-pass filters for sound durations. Neurons with band-, short- and long-pass filtering characteristics discharged maximally to a specific sound duration or a range of sound durations. In contrast, neurons with all-pass filtering characteristics do not have duration selectivity. To determine if duration-tuning characteristics of collicular neurons were tolerant to changes in sound intensity, we examined the duration-tuning characteristics of collicular neurons using a wide range of sound intensities. Duration-tuning characteristics examined included the type, bandwidth and slope of duration-tuning curves. Sound intensity delivered within 20 dB of minimum threshold did not affect duration-tuning characteristics of all collicular neurons studied. Sound intensities at still higher levels did not affect the tuning characteristics of two-thirds of collicular neurons but decreased the duration selectivity and changed the duration-tuning curves of the remaining one-third of neurons from one type to another. However, these two groups of duration-tuning collicular neurons were not separately organized inside the inferior colliculus. The biological relevance of these findings to bat echolocation is discussed.

This study examines the role of neural inhibition in auditory spatial selectivity of inferior collicular neurons of the big brown bat, Eptesicus fuscus, using a two-tone inhibition paradigm. Two-tone inhibition decreases auditory spatial response areas but increases the slopes of directional sensitivity curves of inferior collicular neurons. Inferior collicular neurons have either directionally-selective or hemifield directional sensitivity curves. A directionally-selective curve always has a peak which is at least 50% larger than the minimum. A hemifield directional sensitivity curve rises from an ipsilateral angle by more than 50% and either reaches a plateau or declines by less than 50% over a range of contralateral angles. Two-tone inhibition does not change directionally-selective curves but changes most hemifield directional sensitivity curves into directionally-selective curves. Auditory spatial selectivity determined both with and without two-tone inhibition increases with increasing best-excitatory frequency. Sharpening of auditory spatial selectivity by two-tone inhibition is larger for neurons with smaller differences between excitatory and inhibitory best frequencies. The effect of two-tone inhibition on auditory spatial selectivity increases with increasing inhibitory tone intensity but decreases with increasing intertone interval. The implications of these findings in bat echolocation are discussed.

Clicks emitted by arctiid moths interfere with the ranging ability of echolocating bats. To identify possible neural correlates of this interference, we recorded responses of single units in the nuclei of the lateral lemniscus to combinations of a broad-band click and a test signal (pure tones or frequency-modulated sweeps). In 77% of 87 units tested, clicks interfered with neural responses to the test stimuli. The interference fell into two categories: latency ambiguity and suppression. Units showing latency ambiguity responded to both the click and the test signal. However, when the click occurred within a window of approximately 3 ms before the onset of the test signal, the latency of the response to the test signal was affected. Units that were suppressed did not respond to clicks. Nevertheless, when a click was presented immediately before or simultaneously with a test signal, the response to the test signal was eliminated. Both types of units were found throughout the lateral lemniscus except for the columnar division of the ventral nucleus, where all units tested exhibited latency ambiguity. There is a close match between the single unit data and previous studies of range difference discrimination in the presence of clicks.

The ability of the noctuid A1 cell acoustic receptor to encode biologically relevant information from bat echolocation calls is examined. Short-duration stimuli (less than approximately 6 ms) reduce the dynamic resolution of the receptor, making intensity, and hence range, estimates of foraging bats unreliable. This low dynamic range is further reduced by inaccurate encoding of stimulus intensity, reducing the real dynamic range of the A1 cell to 1 bit at stimulus durations below 3.1 ms. Interspike interval is also an unreliable measure of stimulus intensity at low stimulus levels and/or for short-duration stimuli. The quantity of information encoded per stimulus is reduced as the presentation rate of stimuli is increased. The spontaneous generation of A1 cell action potentials may reduce the ability of the moth to discriminate bat from non-bat signals. Even with a recognition criterion of three A1 cell spikes per call, the moth would regularly make wrong decisions about a bat being present in the immediate environment. Removing this noise would necessitate a considerable loss of information through filtering at the interneurone level. It is proposed that, for bats using short-duration calls, the moth would only be able to recognise an approaching bat from the repetitious nature of the incoming signal.

Bat surveys are frequently undertaken using ultrasonic detectors to determine the species present in an area on the basis of the identity of echolocation calls. We compared three techniques for using the Anabat II detector: the detector pointed along tracks (flyways) versus the detector pointed across tracks (non-flyways); recording output to audio cassette (analogue) versus direct recording to computer (digital); and active hand-held recording versus static automatic recording. In addition, we derived a species-accumulation curve from all-night Anabat recordings in the wet–dry tropics of the Northern Territory. We found no significant difference between flyway and non-flyway recordings; significantly more calls were identified from digital recordings; and significantly more species were detected using hand-held than static recordings. Species-accumulation analysis suggests that the minimum time required to achieve a satisfactory (80%) inventory of bat species at a site is during the three-hour period immediately after sunset. We use our findings to make recommendations for the design of bat surveys using the Anabat II detector.

Stationary and walking tympanate moths (Arctiidae, Geometridae, Noctuidae, and Notodontidae) exhibited discrete behavioural responses to electronically simulated bat echolocation calls. The presence of a stationary response in a given species was strongly correlated with the presence of a walking response in the same species. Some inter- and intra-specific variability in the presence of this behaviour was observed and the evolutionary implications of this variability are discussed. Such behavioural responses by nonvolant moths to ultrasound may protect them from predation by gleaning bats.

The tympanate, arctiid moth,Cycnia tenera responds to pulsed, 30 kHz acoustic stimuli resembling bat echolocation signals by emitting trains of clicks. This phonoresponse was used to determine that this moth is maximally sensitive to stimulus pulse repetition rates of 30–50 pulses/s, rates typically emitted by bats shortly before they close with their targets. At rates both above and below this optimum moths exhibit higher thresholds and reduced responsiveness. These data suggest thatC. tenera is capable of using the repetition rate emitted by an approaching bat as a cue in determining the relative proximity of the bat. The use of repetition rate information should allow this moth both an unambiguous indication of a bat at very close range as well as the ability to distinguish sources of nocturnal, high-frequency sounds not emitted by predators.

1. Many arctiid and ctenuchid moths produce clicking sounds in response to the ultrasonic cries of bats. Clicks were recorded from the two arctiid moth speciesArctia caja, the garden tiger, andPhragmatobia fuliginosa, the ruby tiger. The threshold for eliciting clicks was around 60 to 75 dB pe SPL in both species.A. caja produced single clicks, andP. fuliginosa bursts of clicks. The maximum intensity of the clicks was 90 to 94 dB pe SPL at 5 cm forA. caja and 85 dB pe SPL at 5 cm forP. fuliginosa. The clicks contain most energy in the frequency range from 40 to 80 kHz (Figs. 2, 3). 2. Pipistrelle bats (Pipistrellus pipistrellus) were trained to sit on a platform and discriminate the difference in range,Δd, to two targets. The minimum Δd the bats could discriminate with more than 75% success rate was 1.5 cm. 3. The targets had built-in electrostatic loudspeakers through which different sounds could be played back to the bat. Playback of arctiid moth clicks from both targets did not disturb the bat's discrimination accuracy. The success rate did not decrease at anyΔd, and the minimumΔ d in the presence of clicks was 1 cm. 4. The clicks played from both loudspeakers did not influence the acoustic behavior or discrimination behavior of the bats in any obvious way. In all trials the bats went through a period with long (3 ms) slowly repeated (12–15 pulses/s) cries, a period with shorter cries and increased PRR (20 pulses/s) in which the decision seemed to be made, and finally a period with very short cries (0.5 ms) repeated at rates of up to 150 pulses/s (Figs. 4 and 5). The cries were FM sweeps from 120 kHz to 55 kHz with a second harmonic, which was strongest in the short cries. 5. The bats' response to the playback of different sounds, such as noise and recorded bat cries, from either the left or right loudspeaker, suggested that the bats reacted to clicks as if they were noise. The playback of sounds from only one speaker at a time decreased the bats' success rate, since the bats were attracted to the sounds (Figs. 6 and 7). 6. A secretion from the cervical glands ofA. caja, which contains choline ester, was given to a bat if it crawled towards a clicking target. Both bats tested in this way learned to associate the clicks with a noxious reward and avoided the clicks after just one or two trials (Fig. 8). 7. These results suggest that the function of the garden tiger and ruby tiger clicks in nature is to warn the bat of the moth's distastefulness, and not to ‘jam’ the bat's sonar system.

The dogbane tiger moth (Cycnia tenera Hubner; Arctiidae) responds to ultrasonic, artificial bat echolocation signals by emitting stereotyped trains of high-frequency, rapidly repeated clicks. By comparing this response in intact and headless moths, the role of protocerebral auditory inter-neurones suggested by other studies was examined. Individual moths were tested intact and decapitated, and their response differences analysed. Response latency and threshold (dB) did not alter with the removal of the head but response duration and responsiveness to stimulus trains were significantly reduced in headless moths. These data are interpreted as suggesting the existence of a reflex arc connecting the moth's tympanic organ (ear) with its sound-producing structure (tymbal), and as providing preliminary evidence that the role of higher-order interneurones is primarily that of response reinforcement.

Analysis of the clicks produced by some arctiid moths shows that their acoustic characteristics (power spectra and frequency–time structures) are remarkably similar to those of frequency-modulated echolocation calls produced by many bats as they close with their prey. We suspect that the clicks are initially processed as echoes by the auditory systems of these bats and function by interfering with information processing by the bat.