Foraging behavior and Doppler shift compensation in echolocating hipposiderid bats, I-Iipposideros bicolor and I-Iipposideros speoris

1. Two hipposiderid bats,H. bicolor andH. speoris, were observed in their natural foraging areas in Madurai (South India). Both species hunt close together near the foliage of trees and bushes but they differ in fine structure of preferred hunting space:H. bicolor hunts within the foliage, especially whenH. speoris is active at the same time, whereasH. speoris never flies in dense vegetation but rather in the more open area (Fig. 1, Table 1). 2. Both species emit CF/FM-sounds containing only one harmonic component in almost all echolocation situations. The CF-parts of CF/FM-sounds are species specific within a band of 127–138 kHz forH. speoris and 147–159 kHz forH. bicolor (Tables 2 and 3). 3. H. speoris additionally uses a complex harmonic sound during obstacle avoidance and during laboratory tests for Doppler shift compensation.H. bicolor consistently emits CF/FM-sounds in these same situations (Fig. 2). 4. Both hipposiderid bats respond to Doppler shifts in the returning echoes by lowering the frequency of the emitted sounds (Fig. 3). However, Doppler compensations are incomplete as the emitted frequencies are decreased by only 55% and 56% (mean values) of the full frequency shifts byH. speoris andH, bicolor, respectively. 5. The differences in Doppler shift compensation, echolocating and hunting behavior suggest thatH. speoris is less specialized on echolocation with CF/FM-sounds thanH. bicolor

The use of acoustical cues for prey detection by the Indian false vampire bat, Megaderma lyra

1. The response of the echolocating bat, Megaderma lyra, was tested to different kinds of prey in an outdoor cage. The bats caught larger flying insects (moths, beetles, grasshoppers, and cockroaches) on the wing and also picked up arthropods (solifugid spiders, beetles and cockroaches) and small vertebrates (mice, fishes, frogs and geckoes) from the ground. After touching the prey with the muzzle, the bats were able to differentiate between species. Scorpions and toads were not taken by M. lyra. 2. In lighted and in dark conditions, M. lyra detected and caught prey only when it moved. Dead frogs briskly pulled over the floor were also detected and caught, whereas stationary dead frogs were disregarded by the bats (Table 1). 3. When dead frogs were pulled over the watered surface of a glass plate to eliminate noises by motion, the motion no longer alarmed the bats. From the results of these experiments it was concluded that M. lyra detects prey on the ground by listening to the noise of the moving target only, and not by echolocation (Table 1 C, Fig. 1). Furthermore, M. lyra were not attracted by frog calls. 4. M. lyra differentiated between palatable frogs and non-palatable toads only after touching the prey with the muzzle. 5. Experiments with freshly killed frogs coated with toad secretions or covered with toad skins indicate that M. lyra differentiates between frogs and toads by chemical means. There was no evidence that these prey were differentiated by means of echolocation

Ontogenesis of tonotopy in inferior colliculus of a hipposiderid bat reveals postnatal shift in frequency-place code

The postnatal development of midbrain tonotopy was investigated in the inferior colliculus (IC) of the south Indian CF-FM bat Hipposideros speoris. The developmental progress of the three-dimensional frequency representation was determined by systematic stereotaxic recordings of multiunit clusters from the 1st up to the 7th postnatal week. Additional developmental measures included the tuning characteristics of single units (Figs. 3f; 4f; 5f), the analysis of the vocalised pulse repertoire (Figs. 3e, 4e, 5e), and morphometric reconstructions of the brains of all experimental animals (Fig. 1). The maturation of auditory processing could be divided into two distinct, possibly overlapping developmental periods: First, up to the 5th week, the orderly tonotopy in the IC developed, beginning with the low frequency representation and progressively adding the high frequency representation. With regard to the topology of isofrequency sheets within the IC, maturation progresses from dorsolateral to ventromedial (Figs. 3c, 4c). At the end of this phase the entire IC becomes specialised for narrowly tuned and sensitive frequency processing. This includes the establishment of the 'auditory fovea', i.e. the extensive spatial representation of a narrow band of behaviorally relevant frequencies in the ventromedial part of the IC. In the 5th postnatal week the auditory fovea is concerned with frequencies from 100-118 kHz (Fig. 4c, d). During subsequent development, the frequency tuning of the auditory fovea increases by 20-25 kHz and finally attains the adult range of ca. 125-140 kHz. During this process, neither the bandwidth of the auditory fovea (15-20 kHz) nor the absolute sensitivity of its units (ca. 50 dB SPL) were changed. Further maturation occurred at the single unit level : the sharpness of frequency tuning increased from the 5th to the 7th postnatal weeks (Q-10-dB-values up to 30-60), and upper thresholds emerged (Figs. 4f, 5f). Although in the adult the frequency of the auditory fovea matches that of the vocalised pulses, none of the juvenile bats tested from the 5th to the 7th weeks showed such a frequency match between vocalisation and audition (Figs. 4e, 5e). The results show that postnatal maturation of audition in hipposiderid bats cannot be described by a model based on a single developmental parameter

Movement as a specific stimulus for prey catching behaviour in rhinolophid and hipposiderid bats

1. The echolocating 'long CF/FM-bat'Rhinolophus rouxi and the 'short CF/FM-bats'Hipposideros bicolor andHipposideros speoris were tested for catching responses to moving and non-moving targets. 2. Under our experimental conditions (freshly caught caged bats in a natural environment)Rhinolophus rouxi and Hipposideros speoris only responded to insects of any sort that were beating their wings. The bats showed no reactions whatsoever to nonmoving insects or those walking on the floor or the sides of the cage. 3. Hipposideros bicolor responded in the same way as the above species to wingbeating insects but in addition also attacked walking insects. In 27 presentations 15 walking insects were caught (Fig. 2). 4. Rhinolophus rouxi, Hipposideros speoris and Hipposideros bicolor also detected, approached and seized tethered cockroaches hanging from the ceiling when these were vibrating up and down (Fig. 3). This indicates that any oscillating movement and not specific aspects of wing beating were the key releasers for catching behaviour in all three species. However, a wing beating insect is strongly preferred over a vibrating one in all three species (Fig. 4). 5. Rhinolophus rouxi, Hipposideros speoris and Hipposideros bicolor attacked and seized a dead bait when it was associated with a wing beating device (Fig. 1). All three species responded effectively to beat frequencies as low as 10 beats/s (peak-to-peak amplitude of the wing excursion 20 mm). For lower frequencies the response rates rapidly deteriorated (Fig. 5). 6. Horseshoe bats no longer responded to wing beats of 5 beats/s when the wing beat amplitude was 2 to 1 mm or to wing beats of 2 to 1 beats/s when the amplitude was 3 mm or lower (Fig. 6). This suggests that the speed of the wing is a critical parameter. From these data we infer that the threshold for the catching responses is at a wing speed of about 2 to 1 cm/s. 7. In horseshoe bats (experimental tests) and the two hipposiderid species (behavioural observations) one single wing beat was enough to elicit a catching response (Fig. 8). 8. It is concluded that 'long' and 'short' CF/ FM-bats feature a similar responsiveness to fluttering targets. The sensitivity to oscillating movements is considered as an effective detection mechanism for any sort of potential prey

Hearing Characteristics and Doppler Shift Compensation in South Indian CF-FM Bats

1. Echolocation pulses, Doppler shift compensation behaviour under laboratory conditions and frequency response characteristics of hearing were recorded inRhinolophus rouxi, Hipposideros speoris andHipposideros bicolor. 2. The frequencies of the constant frequency portions of the CF-FM pulses lie at about 82.8 kHz forR. rouxi from Mahabaleshwar, at 85.2 kHz forR. rouxi from Mysore. Hipposiderid bats have considerably higher frequencies at 135 kHz inH. speoris and 154.5 kHz inH. bicolor. The mean sound durations were 50 ms, 6.4 ms and 4.7 ms, respectively. 3. R. rouxi compensates for Doppler shifts in a range up to typically 4 kHz of positive Doppler shifts (Fig. 2). The Doppler shift compensation behaviour is almost identical to that ofR. ferrumequinum. 4. H. speoris andH. bicolor do not compensate for Doppler shifts under laboratory conditions. Doppler shifts in the echoes induce emission frequency changes which are not correlated to the presented Doppler shifts (Fig. 3). 5. The frequency response characteristics of hearing ofR. rouxi show characteristic sensitivity changes near the bat's reference frequency as also found inR. ferrumequinum. The threshold differences between the low threshold at the reference frequency and a few hundred Hz below are 40 to 50 dB in awake bats (Fig. 5). 6. Frequency sensitivity changes near the emitted CF-frequency of the bats are less pronounced inH. speoris or almost absent inH. bicolor

Collicular Responses to the Frequency Modulated Final Part of Echolocation Sounds in Rhinolophusferrum equinum

Collicular evoked potentials in Rhinolophus ferrum equinum show very prominent responses to the final frequency modulated part of a acoustic stimulus, simulating the natural echolocation sound

Audition in vampire bats, Desmodus rotundus

1. Within the tonotopic organization of the inferior colliculus two frequency ranges are well represented: a frequency range within that of the echolocation signals from 50 to 100 kHz, and a frequency band below that of the echolocation sounds, from 10 to 35 kHz. The frequency range between these two bands, from about 40 to 50 kHz is distinctly underrepresented (Fig. 3B). 2. Units with BFs in the lower frequency range (10–25 kHz) were most sensitive with thresholds of -5 to -11 dB SPL, and units with BFs within the frequency range of the echolocation signals had minimal thresholds around 0 dB SPL (Fig. 1). 3. In the medial part of the rostral inferior colliculus units were encountered which preferentially or exclusively responded to noise stimuli. — Seven neurons were found which were only excited by human breathing noises and not by pure tones, frequency modulated signals or various noise bands. These neurons were considered as a subspeciality of the larger sample of noise-sensitive neurons. — The maximal auditory sensitivity in the frequency range below that of echolocation, and the conspicuous existence of noise and breathing-noise sensitive units in the inferior colliculus are discussed in context with the foraging behavior of vampire bats

Environmental correlates of geographic divergence in a phenotypic trait: A case study using bat echolocation

Divergence in phenotypic traits may arise from the interaction of different evolutionary forces, including different kinds of selection (e.g., ecological), genetic drift, and phenotypic plasticity. Sensory systems play an important role in survival and reproduction, and divergent selection on such systems may result in lineage diversification. Such diversification could be largely influenced by selection in different environments as a result of isolation by environment (IbE). We investigated this process using geographic variation in the resting echolocation frequency of the horseshoe bat species, Rhinolophus damarensis, as a test case. Bats were sampled along a latitudinal gradient ranging from 16°S to 32°S in the arid western half of southern Africa. We measured body size and peak resting frequencies (RF) from handheld individual bats. Three hypotheses for the divergence in RF were tested: (1) James’ Rule, (2) IbE, and (3) genetic drift through isolation by distance (IbD) to isolate the effects of body size, local climatic conditions, and geographic distance, respectively, on the resting frequency of R. damarensis. Our results did not support genetic drift because there was no correlation between RF variation and geographic distance. Our results also did not support James’ Rule because there was no significant relationship between (1) geographic distances and RF, (2) body size and RF, or (3) body size and climatic variables. Instead, we found support for IbE in the form of a correlation between RF and both region and annual mean temperature, suggesting that RF variation may be the result of environmental discontinuities. The environmental discontinuities coincided with previously reported genetic divergence. Climatic gradients in conjunction with environmental discontinuities could lead to local adaptation in sensory signals and directed dispersal such that gene flow is restricted, allowing lineages to diverge. However, our study cannot exclude the role of processes like phenotypic plasticity in phenotypic variation