Echolocation in the notch-eared bat,Myotis emarginatus

Summary In Myotis emarginatus, the patterns of echolocation sounds vary with different foraging habitats: In commuting flights the echolocation sounds are linearly frequency modulated sweeps that start at about 100 kHz, terminate at 40 kHz, and have a duration of 1–3 ms. They consist of a loud first harmonic. The second and third harmonics are at least 15 dB fainter than the first one and often undetectable. A distinctly different type of sound is emitted when the bats search for flying insects in open spaces. The sounds are reduced in bandwidth and elongated by a constant frequency component that follows the initial frequency modulated part. Typically, sounds start at about 94 kHz and terminate in a constant frequency component at about 40–45 kHz. The average duration of the constant frequency tail is 2.8 ms; this approximately doubles the length of the pulse, with the longest recorded sound lasting 7.2 ms. When bats are foraging near and within foliage, and gleaning prey from foliage, echolocation sounds are brief (average 1 ms) frequency modulated pulses with a broad bandwidth. The pulses start at about 105 kHz and sweep down to 25 kHz. During gleaning within a building, the frequency range of the sounds is shifted to higher frequencies and extends from 124 to 52 kHz. When the bats forage for aireal insects in a confined area that creates echo-clutter, they emit sounds similar to those used during gleaning within buildings except that sound durations are extended to about 1.8 ms. In each foraging area, the echolocation sounds emitted during the search for and approach to prey are similar in structure. Sound and pause durations are reduced in the approach phase. Irrespective of foraging style and habitat, immediately before capture the bat emits a rapid and stereotyped sequence of 2-10 echolocation pulses (final buzz). These pulses are brief (0.2–0.5 ms), frequency modulated sounds with a reduced bandwidth. The sounds start at 45 kHz and sweep down to 35–20 kHz. The repetition rate is increased up to 200 pulses/s. Offprint requests to: G. Neuweiler     

The echolocation and hunting behavior of Daubenton's bat,Myotis daubentoni

Summary The echolocation and hunting behavior of Daubenton's bat (Myotis daubentoni) were studied in the field under completely natural conditions using a multiflash photographic system synchronized with high-speed tape recordings. The hunting behavior of M. daubentoni is separated into four stages. In the search flight stage Daubenton's bat flies with an average speed of 3.4±0.6 m/s SD usually within 30 cm over water surfaces searching for insects. After the detection of potential prey, the approach flight stage occurs, during which the bat approaches the target in a goal-directed flight. The stage tail down indicates that M. daubentoni is close to the potential prey (approximately 10–22 cm) and is preparing for the catch. The insects are caught with the interfemoral membrane, the feet, and sometimes with the additional aid of a wing. In the stage head down, the bat seizes the prey during flight. Immediately afterwards, Daubenton's bat returns to search flight. M. daubentoni shows the typical echolocation behavior of a vespertilionid bat, emitting frequency-modulated (FM) echolocation signals. The three behavioral stages search, approach, and terminal phase (Griffin et al. 1960) are used to describe the pulse pattern of foraging M. daubentoni in the field. The terminal phase (or buzz) of Daubenton's bat is separated into two parts: buzz I and buzz II. Buzz II is distinguished from buzz I by the following characteristics: a sharp drop in terminal frequency, a distinct reduction in the bandwidth of the first harmonic, a continuous high repetition rate throughout the phase in the range 155–210 Hz, very short pulses (0,25–0.3 ms) and interpulse intervals (4.5–5.0 ms) at the end of the phase, and a distinct decrease in duty cycle. A pause in echolocation separates the end of the terminal phase from the ongoing search phase. The reduction in sound duration after the detection of a target and during pursuits with successfull or attempted catches is discussed in relation to the actual distance of the bat to the target at each stage. It is likely that Daubenton's bat reduces sound duration during approach and terminal phase in order to prevent an overlap of an outgoing pulse with the returning echo from the target. It is argued that the minimum detection distance can be estimated from the sound duration during search flight. Estimates of detection and reaction distances of M. daubentoni based upon synchronized photos and echolocation sequences are given to corroborate this hypothesis. An average detection distance of 128 cm and an average reaction distance of 112 cm were determined. Each behavioral stage of foraging M. daubentoni is characterized by a distinct pattern of echolocation signals and a distinct stage in hunting behavior. The approach flight in hunting behavior coincides with the approach phase and with buzz I in echolocation behavior. The stage tail down corresponds to buzz II. The stage head down is correlated with a pause in echolocation. Immediately afterwards, the bat returns into search flight and into the search phase, emitting search signals.     

Unusual echolocation behavior in a small molossid bat, <Emphasis Type="Italic">Molossops temminckii,</Emphasis> that forages near background clutter

When searching for flying insects, Molossops temminckii uses unusual echolocation calls characterized by upward modulation of frequency vs time (UFM). Call frequency increases asymptotically in the relatively long (∼8 ms) pulses from a starting frequency of ∼40 kHz to a long narrowband tail at ∼50 kHz. When approaching a prey, the bat progressively increases the duration of calls and intersperses in the sequence broadband downwardly frequency-modulated signals with a terminal frequency of about 53 kHz, which totally replaces the UFM signals at the end of the approach phase. The sequence progresses to a capture buzz resembling those from other molossid and vespertilionid bats. The M. temminckii wing morphology is characterized by an average aspect ratio and a high wing loading, suggesting that it is more maneuverable than the typical Molossidae but less than typical Vespertilionidae. M. temminckii regularly forages near clutter, where it needs to pay attention to the background and might face forward and backward masking of signals. We hypothesize that the UFM echolocation signals of M. temminckii represent an adaptation to foraging near background clutter in a not very maneuverable bat needing a broad attention window. The broadband component of the signal might serve for the perception of the background and the narrowband tail for detection and perhaps classification of prey. Bats may solve the signal masking problems by separating emission and echoes in the frequency domain. The echolocation behavior of M. temminckii may shed light on the evolution of the narrowband frequency analysis echolocation systems adopted by some bats foraging within clutter.     

Foraging behavior and echolocation of wild horseshoe bats Rhinolophus ferrumequinum and R. hipposideros (Chiroptera,Rhinolophidae)

Summary 1. Echolocation and foraging behavior of the horseshoe bats Rhinolophus ferrumequinum and R. hipposideros feeding under natural conditions are described. 2. The calls of both species consisted predominantly of a long CF segment, often initiated and terminated by brief FM sweeps of substantial bandwidth. 3. R. hipposideros typically flew close to vegetation, and fed by aerial hawking, gleaning and by pouncing on prey close to the ground. R. hipposideros called with a CF segment close to 112 kHz which is the second harmonic of the vocalization; its calls included low intensity primary harmonics, and had prominent initial and terminal FM sweeps of considerable bandwidth. When searching for prey on the wing it had longer interpulse intervals than R. ferrumequinum, but emitted shorter pulses at a higher repetition rate; overall it had a similar duty cycle to R. ferrumequinum. 4. R. ferrumequinum, calling with a CF segment close to 83 kHz, also used harmonics other than the dominant secondary in its calls, and modified its echolocation according to ecological conditions. This species showed certain parallels with R. rouxi of Asia. It was observed feeding by aerial hawking and by flycatching. When scanning for prey from a perch (perch hunting), calls were of shorter duration, and interpulse intervals were on average longer, than when bats were flying. Mean duty cycle was longer in flight, and the bandwidths and frequency sweep rates of the FM segments in the calls increased in comparison with perched bats. 5. FM information may facilitate determination of target range and the location and nature of obstacles; it may also be involved in the interpretation of echoes and the detection of moving targets among clutter. The rising FM sweep initiating the call in both species when flying (and to a lesser extent perch hunting) in the wild must have a significant adaptive role, and should be considered an essential component of the call; rhinolophids should be termed FM/CF/FM bats.Abbreviations CF constant frequency - FM frequency modulated - FM₁ initial rising frequency sweep - FM₂ terminal falling frequency sweep - PRR pulse repetition rate - SD standard deviation - SNR signal-to-noise ratio     

Fishing and echolocation behavior of the greater bulldog bat,Noctilio leporinus,in the field

When hunting for fish Noctilio leporinus uses several strategies. In high search flight it flies within 20–50 cm of the water surface and emits groups of two to four echolocation signals, always containing at least one pure constant frequency (CF) pulse and one mixed CF-FM pulse consisting of a CF component which is followed by a frequency-modulated (FM) component. The pure CF signals are the longest, with an average duration of 13.3 ms and a maximum of 17 ms. The CF component of the CF-FM signals averages 8.9 ms, the FM sweeps 3.9 ms. The CF components have frequencies of 52.8–56.2 kHz and the FM components have an average bandwidth of 25.9 kHz. A bat in high search flight reacts to jumping fish with pointed dips at the spot where a fish has broken the surface. As it descends to the water surface the bat shows the typical approach pattern of all bats with decreasing pulse duration and pulse interval. A jumping fish reveals itself by a typical pattern of temporary echo glints, reflected back to the bat from its body and from the water disturbance. In low search flight N. leporinus drops to a height of only 4–10 cm, with body parallel to the water, legs extended straight back and turned slightly downward, and feet cocked somewhat above the line of the legs and poised within 2–4 cm of the water surface. In this situation N. leporinus emits long series of short CF-FM pulses with an average duration of 5.6 ms (CF 3.1 and FM 2.6) and an average pulse interval of 20 ms, indicating that it is looking for targets within a short range. N. leporinus also makes pointed dips during low search flight by rapidly snapping the feet into the water at the spot where it has localized a jumping fish or disturbance. In the random rake mode, N. leporinus drops to the water surface, lowers its feet and drags its claws through the water in relatively straight lines for up to 10m. The echolocation behavior is similar to that of high search flight. This indicates that in this hunting mode N. leporinus is not pursuing specific targets, and that raking is a random or statistical search for surface fishes. When raking, the bat uses two strategies. In directed random rake it rakes through patches of water where fish jumping activity is high. Our interpretation is that the bat detects this activity by echolocation but prefers not to concentrate on a single jumping fish. In the absence of jumping fish, after flying for several minutes without any dips, N. leporinus starts to make very long rakes in areas where it has hunted successfully before (memory-directed random rake). Hunting bats caught a fish approximately once in every 50–200 passes through the hunting area.     

Echolocation and foraging behavior of the lesser bulldog bat, Noctilio albiventris : preadaptations for piscivory?

We studied variability in foraging behavior of Noctilio albiventris (Chiroptera: Noctilionidae) in Costa Rica and Panamá and related it to properties of its echolocation behavior. N. albiventris searches for prey in high (>20 cm) or low (<20 cm) search flight, mostly over water. It captures insects in mid-air (aerial captures) and from the water surface (pointed dip). We once observed an individual dragging its feet through the water (directed random rake). In search flight, N. albiventris emits groups of echolocation signals (duration 10–11 ms) containing mixed signals with constant-frequency (CF) and frequency-modulated (FM) components, or pure CF signals. Sometimes, mostly over land, it produces long FM signals (duration 15–21 ms). When N. albiventris approaches prey in a pointed dip or in aerial captures, pulse duration and pulse interval are reduced, the CF component is eliminated, and a terminal phase with short FM signals (duration 2 ms) at high repetition rates (150–170 Hz) is emitted. Except for the last pulses in the terminal phase N. albiventris avoids overlap between emitted signals and echoes returning from prey. During rakes, echolocation behavior is similar to that in high search flight. We compare N. albiventris with its larger congener, N. leporinus, and discuss behavioral and morphological specializations that can be interpreted as preadaptations favoring the evolution of piscivory as seen in N. leporinus. Prominent among these specializations are the CF components of the echolocation signals which allow detection and evaluation of fluttering prey amidst clutter-echoes, high variability in foraging strategy and the associated echolocation behavior, as well as morphological specializations such as enlarged feet for capturing prey from the water surface. Received: 21 April 1997 / Accepted after revision: 12 January 1998     

Echolocation call design and limits on prey size: a case study using the aerial-hawking bat Nyctalus leisleri

The echolocation calls used by Nyctalus leisleri during search phase in open air space are between 9 and 14 ms long, with the peak energy between 24 and 28 kHz. The pulses are shallowly frequency-modulated with or without an initial steep frequency-modulated component. The diet consists primarily of small flies (Diptera), including many chironomids (wingspan 9–12 mm) and yellow dung flies (Scatophaga; wingspan 24 mm), but also of some larger insects such as dung beetles (Coleoptera; Scarabaeoidea), caddis-flies (Trichoptera) and moths (Lepidoptera). The echo target strength of some prey items was measured. Contrary to models based on standard targets such as spheres or disks, the echo strength of real insects was found to be virtually independent of the emitted frequency within the 20–100 kHz frequency range. A model was used to calculate probable detection distances of the prey by the bat. Using narrow-band calls of 13.7 ± 2.7 ms duration, a bat would detect the two smallest size classes of insect at greatest range using calls of 20 kHz. The results may therefore explain why many species of large and medium sized aerial-hawking bats use low-frequency calls and still eat mostly relatively small insects. The data and model challenges the assumption that small prey are unavailable to bats using low-frequency calls.     

Foraging habitat and echolocation behaviour of Schneider's leafnosed bat, Hipposideros speoris, in a vegetation mosaic in Sri Lanka

The Hipposideridae and Rhinolophidae are closely related families of bats that have similar echolocation (long-duration pure-tone signal, high duty cycle) and auditory systems (Doppler-shift compensation, auditory fovea). Rhinolophid bats are known to forage in highly cluttered areas where they capture fluttering insects, whereas the foraging habitat of hipposiderid bats is not well understood. Compared to rhinolophids, hipposiderid calls are shorter in duration, have lower duty cycles, and they exhibit only partial Doppler-shift compensation. These differences suggest that the foraging habitat of the two families may also differ. We tested this hypothesis by studying foraging and echolocation of Hipposideros speoris at a site with a range of vegetation types. Bats foraged only while in flight and used all available closed and edge habitats, including areas adjacent to open space. Levels of clutter were high in forest and moderate in other foraging areas. Prey capture (n=42) occurred in edge vegetation where it bordered open space. Echolocation signals of H. speoris lacked an initial upward frequency-modulated sweep and were of moderate duration (5.1-8.7 ms). Sequences had high duty cycles (23-41%) and very high pulse repetition rates (22.8-60.6 Hz). Variation in signal parameters during search phase flight across foraging habitats was low. H. speoris showed a greater flexibility in its use of foraging habitat than is known for any rhinolophid species. Our study confirmed that there are differences in habitat use between hipposiderid and rhinolophid bats and we suggest that this divergence is a consequence of differences in their echolocation and auditory systems.     

Echolocation,development, and vocal communication in the lesser bulldog bat,Noctilio albiventris

Summary Foraging and echolocation behavior and its ontogeny in the lesser bulldog bat, Noctilio albiventris, were studied in Panama under field and captive conditions. The vocalizations utilized for echolocation and communication were monitored. Adult N. albiventris captured insect prey from the water surface employing various combinations of CF/FM (constant frequency and frequency modulated) signals. The proportions of CF/FM and the repetition rate were a function of the bat's activity. Most adults exhibited post-sunset and pre-dawn foraging activity, although several telemetered lactating females foraged for only the half hour after dusk, spending the rest of the night with their babies in the roost. When the juveniles began to leave the roost at the age of two months, they appeared to accompany their mothers on initial flights.Captive infant Noctilio developed slowly, and did not fly until about 5–6 weeks postnatally. They continued to nurse for almost 3 months, even though they were capable of eating solid food at about 6 weeks. Previous to weaning, mothers fed their infants with masticated food from their cheekpouches.At birth, Noctilio emit a combination of long FM isolation calls and shorter CF/FM pulses. Mothers nurse only their own babies which they appear to recognize by a vocal signature contained in the infants' isolation calls. The individual isolation calls, as well as the mother's communication sounds, appear to be variations of an FM sinusoidal wave. The periodicity and amplitude change, and different portions including harmonics are added or deleted. The short CF/FM signals of the infant evolve into the adult orientation type signals as the CF component increases in frequency and the repetition rate increases. These sounds appear to serve a dual function in communication and echolocation. Mother-young pairs were observed to call antiphonally, utilizing CF/short FM signals in retrieval situations. This duetting was also observed in bats flying over the Chagras River after the time the juveniles began to fly, and may function to maintain vocal contact during initial foraging flights.Deceased     

High altitude echolocation of insects by bats

Summary The orientation sounds of many bats, almost certainly belonging to the genus Tadarida, were recorded at altitudes of 100 to 300 m above the ground by means of an ultrasonic radio microphone. Both in North Queensland, Australia, and in southern Utah and Nevada, USA, bats were often more numerous at 200 to 300 m than near the ground. Rapid increases in pulse repetition rate often indicated that these bats were actively hunting flying insects. The absence of clutter at high altitudes may significantly facilitate the detection and capture of insect prey.     

Foraging behaviour and echolocation in the rufous horseshoe bat (Rhinolophus rouxi) of Sri Lanka

Summary In October 1984 foraging areas and foraging behaviour of the rufous horseshoe bat, Rhinolophus rouxi, were studied around a nursery colony on the hill slopes of Sri Lanka. The bats only foraged in dense forest and were not found in open woodlands (Fig. 1). This strongly supports the hypothesis that detection of fluttering prey is by pure tone echolocation within or close to echo-cluttering foliage. During a first activity period after sunset for about 30–60 min, the bats mainly caught insects on the wing. This was followed by a period of inactivity for another 60–120 min. Thereafter the bats resumed foraging throughout the night. They mainly alighted on specific twigs and foraged in flycatcher style. Individual bats maintained individual foraging areas of about 20x20 m. They stayed in this area throughout the night and returned to the same area on subsequent nights. Within this area the bats generally alighted on twigs at the same spots. Foraging areas were not defended against intruders. The bats echolocated throughout the night at an average repetition rate of 9.6±1.4 sounds/s. While hanging on twigs they scanned the surrounding area for flying prey by turning their bodies continuously around their legs. On average they performed one brief catching flight every 2 min and immediately returned to one of their favourite vantage points. Echolocation sounds may consist of up to three parts, a brief initial frequency-modulated (FM) component, a long constant frequency (CF) part lasting for about 40–50 ms, and a final FM part again (Fig. 4b, c). Adult males and females emitted pure tone frequencies in separate bands, the males from 73.5–77 kHz and the females from 76.5–79 kHz (Fig. 5). During scanning for prey from vantage points, the bats mostly emitted pure tones without any FM component (Fig. 4a). The last few pure tones emitted before take-off were prolonged to about 60 ms duration. The final FM part was therefore not an obligatory component of the echolocation signals in horseshoe bats. During flight and especially during emergence from the cave, most sounds consisted of a pure tone and loud initial and final FM sweeps. We therefore suggest that the initial FM part might also be relevant for echolocation. From our observations we conclude that the FM components are especially important during obstacle avoidance. In most sounds emitted in the field a fainter first harmonic was present. It was usually up to 30 dB fainter than the second harmonic, but in some instances it was as loud or even distinctly louder than the second one (Fig. 6a). Even within one sound the intensity relationship between the two harmonics may be reversed. We therefore suggest that the first harmonic is an integral part of the signal and relevant for information analysis in echolocation.     

Echolocation beam shape in emballonurid bats, Saccopteryx bilineata and Cormura brevirostris

The shape of the sonar beam plays a crucial role in how echolocating bats perceive their surroundings. Signal design may thus be adapted to optimize beam shape to a given context. Studies suggest that this is indeed true for vespertilionid bats, but little is known from the remaining 16 families of echolocating bats. We investigated the echolocation beam shape of two species of emballonurid bats, Cormura brevirostris and Saccopteryx bilineata, while they navigated a large outdoor flight cage on Barro Colorado Island, Panama. C. brevirostris emitted more directional signals than did S. bilineata. The difference in directionality was due to a markedly different energy distribution in the calls. C. brevirostris emitted two call types, a multiharmonic shallowly frequency-modulated call and a multiharmonic sweep, both with most energy in the fifth harmonic around 68?kHz. S. bilineata emitted only one call type, multiharmonic shallowly frequency-modulated calls with most energy in the second harmonic (~46?kHz). When comparing same harmonic number, the directionality of the calls of the two bat species was nearly identical. However, the difference in energy distribution in the calls made the signals emitted by C. brevirostris more directional overall than those emitted by S. bilineata. We hypothesize that the upward shift in frequency exhibited by C. brevirostris serves to increase directionality, in order to generate a less cluttered auditory scene. The study indicates that emballonurid bats are forced to adjust their relative harmonic energy instead of adjusting the fundamental frequency, as the vespertilionids do, presumably due to a less flexible sound production.     

Flexible bat echolocation: the influence of individual,habitat and conspecifics on sonar signal design

Acoustic signals which are used in animal communication must carry a variety of information and are therefore highly flexible. Echolocation has probably such functions and could prove as flexible. Measurable variabitlity can indicate flexibility in a behaviour. To quantify variability in bat sonar and relate to behavioural and environmental factors, I recorded echolocation calls of Euderma maculatum, Eptesicus fuscus, Lasiurus borealis and L. cinereus while the bats hunted in their natural habitat. I analysed 3390 search phase calls emitted by 16 known and 16 unknown individuals foraging in different environmental and behvioural situations. All four species used mainly multiharmonic signals that showed considerable intra- and inter-individual variability in the five signal variables I analysed (call duration, call interval, highest and lowest frequency and frequency with maximum energy) and also in the shape of the sonagram. A nested multivariate analysis of variance identified the influences of individual, hunting site, close conspecifics and of each observation on the frequency with maximum energy in the calls, and on other variables measured. Individual bats differed in multiple comparisons, most often in the main call frequency and least often in call interval. In a discriminant function analysis with resubstitution, 56–76% of a species' calls were assigned to the correct individual. Distinct individual call patterns were recorded in special situations in all species and the size of foraging areas in forested areas influenced temporal and spectral call structure. Echolocation behaviour was influenced by the presence of conspecifics. When bats were hunting together, call duration decreased and call interval increased in all species, but spectral effects were less pronounced. The role of morphometric differences as the source of individually distinct vocalizations is discussed. I also examined signal adaptations to long range echolocation and the influence of obstacle distance on echolocation call design. My results allow to discuss the problems of echo recognition and jamming avoidance in vespertilionid bats.     

Fruit detection and discrimination by small fruit-eating bats (Phyllostomidae): echolocation call design and olfaction

We studied the role of echolocation and other sensory cues in two small frugivorous New World leaf-nosed bats (Phyllostomidae: Artibeus watsoni and Vampyressa pusilla) feeding on different types of fig fruit. To test which cues the bats need to find these fruit, we conducted behavioral experiments in a flight cage with ripe and similar-sized figs where we selectively excluded vision, olfaction, and echolocation cues from the bats. In another series of experiments, we tested the discrimination abilities of the bats and presented sets of fruits that differed in ripeness (ripe, unripe), size (small, large), and quality (intact(infested with caterpillars). We monitored the bats' foraging and echolocation behavior simultaneously. In flight, both bat species continuously emitted short (<2 ms), multi-harmonic, and steep frequency-modulated (FM) calls of high frequencies, large bandwidth, and very low amplitude. Foraging behavior of bats was composed of two distinct stages: search or orienting flight followed by approach behavior consisting of exploration flights, multiple approaches of a selected fruit, and final acquisition of ripe figs in flight or in a brief landing. Both bat species continuously emitted echolocation calls. Structure and pattern of signals changed predictably when the bats switched from search or orienting calls to approach calls. We did not record a terminal phase before final acquisition of a fruit, as it is typical for aerial insectivorous bats prior to capture. Both bat species selected ripe over unripe fruit and non-infested over infested fruit. Artibeus watsoni preferred larger over smaller fruit. We conclude from our experiments, that the bats used a combination of odor-guided detection together with echolocation for localization in order to find ripe fruit and to discriminate among them.     

The use of Doppler-shifted echoes as a flutter detection and clutter rejection system: the echolocation and feeding behavior of Hipposideros ruber (Chiroptera: Hipposideridae)

Summary Hipposideros ruber use CF/FM echolocation calls to detect the wing flutter of their insect prey. Fluttering prey were detected whether the insects were flying or sitting on a surface, and prey in either situation were captured with equal success (approximately 40% of capture attempts). Stationary prey were ignored. The bats did not use visual cues or the sounds of wing flutter to locate their prey. Wing flutter detection suggests that H. ruber exploit the Doppler-shifted information in echoes of their echolocation calls. These bats fed primarily upon moths, usually those of between 10 and 25 mm wingchord, although moths of less than 5 mm and greater than 40 mm wingchord were also attacked and captured. They showed no evidence of selecting moths on the basis of species or other taxonomic distinction, and occasionaly captured other insects.     

Natterer’s bat (Myotis nattereri Kuhl, 1818) hawks for prey close to vegetation using echolocation signals of very broad bandwidth

We present a hitherto unknown prey perception strategy in bats: Myotis nattereri (Vespertilionidae, Chiroptera) is able to perceive prey by echolocation within a few centimeters of echo-cluttering vegetation, by using frequency-modulated search signals of very large bandwidth (up to 135 kHz). We describe the species’ search behavior and echolocation repertoire from the field and from experiments in a flight tent. In the field, bats varied signal parameters in relation to their distance from vegetation and usually flew close to vegetation. In the flight tent, M. nattereri detected and localized prey by echolocation alone as close as 5 cm from vegetation. Apparently, the bats were able to tolerate some overlap between prey and clutter echoes. Passive prey cues (vision, olfaction, prey-generated sounds) were not used in prey perception. The bats selected prey by size. The animals performed aerial catches and produced approach sequences typical for aerial hawking bats, but were able to do so within a few centimeters of the substrate. M. nattereri thus has access to silent, suspended prey very close to vegetation (e.g., spiders, and caterpillars on threads). Received: 29 September 1999 / Received in revised form: 12 February 2000 / Accepted: 12 February 2000     

Plasticity in echolocation signals of European pipistrelle bats in search flight: implications for habitat use and prey detection

We studied the echolocation and hunting behavior of three aerial insectivorous species of bats (Vespertilionidae: Pipistrellus) in the field in order to characterize the signals used by the bats and to determine how call structure varies in relation to habitat structure (uncluttered versus cluttered space). We documented free-flying, naturally foraging wild pipistrelles in various habitats using multiflash stereophotography combined with simultaneous sound recordings. Then we reconstructed the bat's flight position in three-dimensional space and correlated it with the corresponding echolocation sequences. In all three species of pipistrelles, signal structure varied substantially. In echolocation sequences of the search phase we found a consistent association of signal types with habitat types. In uncluttered habitats (obstacles more than 5 m from the bat) pipistrelles emitted almost exclusively narrowband signals with bandwidths less than 15 kHz. In cluttered habitats (obstacles less than 5 m from the bat) they switched to signals with bandwidths of more than 15 kHz. Wideband signals were also used when the bats were turning in cluttered and uncluttered spaces and for an instant after turning away from obstacles. Prey detection occured only when the outgoing signal did not overlap with the returning echo from potential prey. The bats also avoided overlap of echoes from potential prey and obstacles. Based on the results of this study, we propose an overlap-free window within which pipistrelles may detect potential prey and which allows predictions of minimum distances to prey and clutter-producing objects. Correspondence to: E.K.V. Kalko     

The role of synchronized calling,ambient light,and ambient noise,in anti-bat-predator behavior of a treefrog

Summary Male treefrogs, Smilisca sila (Hylidae), produce calls of varying complexity and demonstrate a remarkable ability to synchronize their calls with those of neighbors. The bat Trachops cirrhosus eats frogs and uses the frogs' advertisement calls as locational cues. The bats are less likely to respond to synchronous calls than to asynchronous calls, and when given a choice prefer complex calls to simple calls.Experiments with bat models indicate that, like other frogs, S. sila probably uses visual cues to detect hunting bats. In response to bat models the frogs decreased both the number and the complexity of their calls. The calling behavior of the frogs was sampled in the field during periods with and without artificial illumination. The frogs produced fewer and less complex calls, and they tended to call from more concealed sites, during the period without illumination, when presumably it would have been more difficult for the frogs to detect hunting bats. S. sila tended to call from sites with higher ambient noise level, the noise primarily originating from waterfalls. The frequencies of the dominant energies in the waterfall sounds completely overlapped the frequency range of the S. sila call; thus waterfalls might mask the frog calls. When given a choice between calls produced near and away from waterfall sounds, bats preferred the latter.     

Ecological niche and phylogeny: the highly complex echolocation behavior of the trawling long-legged bat, Macrophyllum macrophyllum

Bats produce echolocation signals that reflect the sensory tasks they perform. In open air or over water, bats encounter few or no background echoes (clutter). Echolocation of such bats is the primary cue for prey perception and varies with the stage of approach to prey, typically comprising search, approach, and terminal group calls. In contrast, bats that glean stationary food from rough surfaces emit more uniform calls without a distinct terminal group. They use echolocation primarily for orientation in space and mostly need additional sensory cues for finding food because clutter echoes overlap strongly with food echoes. Macrophyllum macrophyllum is the only Neotropical leaf-nosed bat (Phyllostomidae) that hunts in clutter-poor habitat over water. As such, we hypothesized that, unlike all other members of its family, but similar to other trawling and aerial insectivorous bats, M. macrophyllum can hunt successfully by using only echolocation for prey perception. In controlled behavioral experiments on Barro Colorado Island, Panamá, we confirmed that echolocation alone is sufficient for finding prey in M. macrophyllum. Furthermore, we showed that pattern and structure of echolocation signals in M. macrophyllum are more similar to aerial and other trawling insectivorous bats than to close phylogenetic relatives. Particularly unique among phyllostomid bats, we found distinct search, approach, and terminal group calls in foraging M. macrophyllum. Call structure, however, consisting of short, multiharmonic, and steep frequency-modulated signals, closely resembled those of other phyllostomid bats. Thus, echolocation behavior in M. macrophyllum is shaped by ecological niche as well as by phylogeny.     

Hunting in unfamiliar space: echolocation in the Indian false vampire bat, Megaderma lyra, when gleaning prey

The literature suggests that in familiar laboratory settings, Indian false vampire bats (Megaderma lyra, family Megadermatidae) locate terrestrial prey with and without emitting echolocation calls in the dark and cease echolocating when simulated moonlit conditions presumably allow the use of vision. More recent laboratory-based research suggests that M. lyra uses echolocation throughout attacks but at emission rates much lower than those of other gleaning bats. We present data from wild-caught bats hunting for and capturing prey in unfamiliar conditions mimicking natural situations. By varying light level and substrate complexity we demonstrated that hunting M. lyra always emit echolocation calls and that emission patterns are the same regardless of light/substrate condition and similar to those of other wild-caught gleaning bats. Therefore, echoic information appears necessary for this species when hunting in unfamiliar situations, while, in the context of past research, echolocation may be supplanted by vision, spatial memory or both in familiar spaces.Communicated by T. Czeschlik