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.     

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     

Echolocation behavior of the bat <Emphasis Type="Italic">Vespertilio murinus</Emphasis> reveals the border between the habitat types “edge” and “open space”

We test the hypothesis that echolocation behavior can be used to find the border between bat habitats. Assuming that bats react to background targets in “edge space” but not in “open space”, we determined the border between these two habitat types for commuting individuals of the parti-colored bat Vespertilio murinus. We recorded sequences of bats’ echolocation signals while they flew parallel to the walls of large buildings and to the ground and determined the signals’ average bandwidth, duration, and pulse interval. These parameters varied systematically with the estimated horizontal and vertical distances between the bats and the background. A distinct effect of horizontal distance to the background on echolocation behavior was found for horizontal distances of less than 6 m, thus indicating the border between edge and open space. Only a few bats flew at vertical distances below 5 m. However, enough passages at vertical distances of 5 m and above indicated that the vertical border is somewhere below a distance of 5 m. Within edge space, V. murinus reacted to the background by reducing signal duration, increasing bandwidth at closer distances, and often emitting one signal per wing beat. In open space, signal parameters did not vary as a function of distance to the background. There, V. murinus emitted the longest signals with the narrowest bandwidth and often made one or two wing beats without emitting a pulse. With our data we support with statistical methods the hypothesis that echolocation behavior reveals the border between the habitat types “edge” and “open space”.     

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.     

Dynamic adjustment of biosonar intensity to habitat clutter in the bat Macrophyllum macrophyllum (Phyllostomidae)

Echolocating bats adjust the time–frequency structure such as sweep rate and pulse interval of their sonar calls when they move from open space to vegetation-dense environments. Emitted call intensity is equally important for echolocation, but adjustment of signal intensity to different habitats has never been systematically studied in any bat species. To address this question, we recorded sonar calls of the Neotropical trawling insectivorous bat Macrophyllum macrophyllum (Phyllostomidae) at three sites with different obstacle densities (clutter). We found a clear correlation between emitted intensity and degree of clutter, with intensity proportional to decreasing clutter. In highly cluttered, semicluttered, and open spaces, M. macrophyllum emitted calls with mean source levels (sound pressure level (SPL) 10 cm from the bat’s mouth) of 100, 105, and 111 dB SPL root mean square (rms), respectively. To our knowledge, this is the first documentation of dynamic intensity adjustments in bats. Phyllostomid bats were previously considered silent, but the 111-dB SPL rms emitted by free-ranging M. macrophyllum in open space is comparable to output in aerial insectivorous bats from other families. Our results suggest that the acoustic constraints of habitats are better predictors of call intensity than phylogeny and therefore likely to be major drivers shaping the sonar system of bats in the course of evolution.     

Echolocation behavior and signal plasticity in the Neotropical bat Myotis nigricans (Schinz, 1821) (Vespertilionidae): a convergent case with European species of Pipistrellus?

We used both field and flight cage observations to investigate the echolocation and foraging behavior of the seldom studied, small, aerial insectivorous bat Myotis nigricans (Vespertilionidae) in Panama. In contrast to its temperate congeners, M. nigricans foraged extensively in open space and showed an echolocation behavior well adapted to this foraging habitat. It broadcast narrowband echolocation signals of 7 ms duration that enhance the chance of prey detection in open space. Because of rhythmical alternations of signal amplitude from signal to signal in our sound recordings of search signals in open space, we conclude that the bats scanned their environment with head movements, thereby enlarging their search volume. In edge-and-gap situations, and in the flight cage, M. nigricans introduced an initial broadband component to its search calls. In the field and in the flight cage, M. nigricans hawked for prey in aerial catches; gleaning was never observed. M. nigricans demonstrates call structures, such as narrow bandwidth and rather long signals adapted to foraging predominantly in open space. Moreover, call structure is highly plastic, allowing M. nigricans to forage in edge-and-gap situations also. These adaptations in call structure and plasticity have evolved convergently at least twice within the genus Myotis. Finally, M. nigricans echolocation and foraging behavior parallels that of the small, aerial, insectivorous pipistrelle bats (Vespertilionidae), which are not closely related to M. nigricans but forage in similar habitats.     

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.     

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.     

The effectiveness of katydid (<Emphasis Type="Italic">Neoconocephalus ensiger</Emphasis>) song cessation as antipredator defence against the gleaning bat <Emphasis Type="Italic">Myotis septentrionalis</Emphasis>

Many nocturnal katydids (Orthoptera: Tettigoniidae) produce intense calling songs, and some bat species use these songs to detect and locate prey. One Nearctic katydid species, Neoconocephalus ensiger, ceases or pauses singing in response to bat echolocation calls. We tested the hypothesis that song cessation is an effective defence against gleaning bats (i.e., bats that take prey from surfaces). We observed Myotis septentrionalis, a sympatric bat species that uses prey-generated sounds when gleaning, attack and feed on singing N. ensiger in an outdoor flight room. These bats demonstrated a preference for the calling song of N. ensiger over a novel cricket calling song when they were broadcast from a speaker in the flight room. Bats attacked speakers broadcasting N. ensiger calling song as long as the song was continuous and aborted their attack if the sound stopped as they approached, regardless of whether a katydid was present as a physical target on the speaker. Echolocation calls were recorded during attacks and no significant differences were found between continuous and interrupted song approaches for four call parameters, suggesting that M. septentrionalis may not use echolocation to locate silent prey. Therefore, song cessation by katydids in response to ultrasound is an effective defence against gleaning bats.     

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     

Prey detection in trawling insectivorous bats: duckweed affects hunting behaviour in Daubenton's bat, Myotis daubentonii

Daubenton's bat, a trawling vespertilionid bat species, hunts for insects that fly close to, or rest on, the water surface. During summer, many ponds at which Daubenton's bats hunt become gradually covered with duckweed. The purpose of this study was to investigate the effects of duckweed cover on the hunting behaviour of Daubenton's bats and on the ultrasound-reflecting properties of the water surface. Our study revealed the following. (1) Daubenton's bat avoids water surfaces covered with duckweed. (2) Prey abundance was related to the number of foraging Daubenton's bats but was independent of duckweed cover. (3) When mealworms were presented among standardized amounts of duckweed to naturally foraging Daubenton's bats, they caught significantly less mealworms when the duckweed cover was increased. (4) Measurements with ultrasonic signals show that a water surface covered with duckweed returns a much stronger background echo at small angles (i.e. parallel to the water surface) compared to an uncovered water surface. It seems likely that a cover of duckweed on the water surface interferes with prey detection by masking the echoes returning from prey. (5) It was relatively difficult for the bats to discriminate small patches of duckweed from mealworms. The proposed discrimination mechanism for this trawling bat species suggests that single duckweed patches can also be mistaken for natural prey by Daubenton's bats. Received: 4 January 1998 / Accepted after revision: 19 July 1998     

Ground gleaning in horseshoe bats: comparative evidence from<Emphasis Type="Italic"> Rhinolophus blasii</Emphasis>,<Emphasis Type="Italic"> R. euryale</Emphasis> and<Emphasis Type="Italic"> R. mehelyi</Emphasis>

The 71 species of horseshoe bat (genus Rhinolophus) use echolocation calls with long constant-frequency (CF) components to detect and localize fluttering insects which they seize in aerial captures or glean from foliage. Here we describe ground-gleaning as an additional prey-capture strategy for horseshoe bats. This study presents the first record and experimental evidence for ground-gleaning in the little-studied Blasius horseshoe bat (Rhinolophus blasii). The gleaning bouts in a flight tent included landing, quadrupedal walking and take-off from the ground. The bats emitted echolocation calls continuously during all phases of prey capture. Both spontaneously and in a choice experiment, all six individuals attacked only fluttering insects and never motionless prey. These data suggest that R. blasii performs ground-gleaning largely by relying on the same prey-detection strategy and echolocation behaviour that it and other horseshoe bats use for aerial hawking.We also studied the Mediterranean horseshoe bat (R. euryale) in the flight tent. All four individuals never gleaned prey from the ground, though they appeared to be well able to detect fluttering moths on the ground. It is not known yet whether ground-gleaning plays a role in Mehelys horseshoe bat (R. mehelyi). In a performance test, we measured the ability of these three European species of middle-sized horseshoe bats (R. euryale, R. mehelyi and R. blasii) to take-off from the ground. All were able to take flight even in a confined space; i.e. the willingness to ground-glean in R. blasii is not related to a superior take-off performance. In contrast to ground-gleaning bats of other phylogenetic lineages, R. blasii appears not to be a specialist, but rather shows a remarkable behavioural flexibility in prey-capture strategies and abilities. We suggest that the key innovation of CF echolocation paired with behavioural flexibility in foraging strategies might explain the evolutionary success of Rhinolophus as the second largest genus of bat.Communicated by T. Czeschlik     

Echolocation in two very small bats from Thailand Craseonycteris thonglongyai and Myotis siligorensis

Summary The echolocation and hunting behavior of two very small bats, Craseonycteris thonglongyai (Hill) and Myotis siligorensis (Horsfield), from Thailand, were investigated using multiflash photographs, video, and high-speed tape recordings with a microphone array that allowed determination of distance and direction to the bats. C. thonglongyai is the world's smallest mammal and M. siligorensis is only slightly larger. Both bats hunted insects in open areas. The search signals of C. thonglongyai were 3.5 ms long multiharmonic constant frequency (CF) signals with a prominent second harmonic at 73 kHz repeated at around 22 Hz. The band width (BW) of the short terminal frequency modulated (FM) sweep increased during the very short approach phase. In the final buzz the CF component disappeared, the duration decreased to 0.2 ms, and the repetition rate increased to 215 Hz (Figs. 2, 3, 4). There was no drop in frequency in the buzz. The video recordings of C. thonglongyai indicated that it seizes insects directly with the mouth (Fig. 1). M. siligorensis produced 5.4 ms long CF search signals at 66 kHz. The repetition rate was around 13 Hz. In the approach phase an initial broad band FM sweep was added. The buzz consisted of two phases, buzz I and buzz II. Buzz 11 was characterized by short cry durations (around 0.3 ms), a constant high repetition rate (185 Hz), a distinct drop in frequency, and a prominent second harmonic (Figs. 5, 6, 7). The drop in frequency, apparently typical of vespertilionid bats, has been explained by physiological limitations in sound production. However, C. thonglongyai produced very short signals at very high repetition rates without any frequency drop. The drop may be of adaptive value since it enables M. siligorensis to produce very short signals with high sweep rates. The drop moves the pronounced second harmonic into the frequency range of most interest to the bat (Fig. 7D). The sweep rate in this frequency range may now increase to twice the maximum rate that the vocal cords can produce directly. C. thonglongyai and M. siligorensis belong to different superfamilies, Emballonuroidea and Vespertilionoidea, respectively. In spite of their phylogenetic distance they produce strikingly similar search signals of narrow BW around 70 kHz with high source levels (100–115 dB peSPL peak equivalent sound pressure level). We argue that the signal resemblance is due to the similarity in size and hunting behavior of the two bats both hunting insects in open areas. High frequencies are heavily attenuated in air, but because of their small size the bats are restricted to hunting small insects which only reflect echoes at high frequencies. Thus, the emitted frequency is probably the lowest possible given the prey size. Hence, the two bats can only maximize the range of their sonar by decreasing the BW and emitting high intensities. Correspondence to: A. Surlykke     

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.     

The roles of echolocation and olfaction in two Neotropical fruit-eating bats, Carollia perspicillata and C. castanea, feeding on Piper

We studied the echolocation and foraging behavior of two Neotropical frugivorous leaf-nosed bats (Carollia perspicillata, C. castanea: Phyllostomidae) in a flight cage. To test which cues Carollia uses to detect, identify, and localize ripe Piper fruit, their preferred natural food, we conducted experiments under semi-natural conditions with ripe, unripe, and artifical fruits. We first offered the bats ripe fruits and documented their foraging behavior using multiflash stereophotography combined with simultaneous sound recordings. Both species showed a similar, stereotyped foraging pattern. In searchflight, the bats circled through the flight cage in search of a branch with ripe fruit. After finding such a branch, the bats switched to approach behavior, consisting of multiple exploration flights and the final approach when the bats picked up the fruit at its tip and tore it off in flight. Our behavioral experiments revealed that odor plays an important role in enabling Carollia to find ripe fruit. While foraging, Carollia always echolocated and produced multiharmonic, frequency-modulated (FM) signals of broad bandwidth, high frequency, short duration, and low intensity. We discriminated an orientation phase (mostly a single pulse per wingbeat) and an approach phase (groups of two to six pulses per wing beat). We conclude from the bats' behavioral reaction to real and artificial fruit as well as from characteristic patterns in their echolocation behavior that during exploration flights, Carollia changes from primarily odor-oriented detection and initial localization of ripe fruit to a primarily echo-oriented final localization of the position of the fruit. Received: 27 March 1997 / Accepted after revision: 28 February 1998     

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     

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.     

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 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.