Genetic divergence and echolocation call frequency in cryptic species of Hipposideros larvatus s.l. (Chiroptera: Hipposideridae) from the Indo-Malayan region

The intermediate leaf-nosed bat ( Hipposideros larvatus ) is a medium-sized bat distributed throughout the Indo-Malay region. In north-east India, bats identified as H. larvatus captured at a single cave emitted echolocation calls with a bimodal distribution of peak frequencies, around either 85 kHz or 98 kHz. Individuals echolocating at 85 kHz had larger ears and longer forearms than those echolocating at 98 kHz, although no differences were detected in either wing morphology or diet, suggesting limited resource partitioning. A comparison of mitochondrial control region haplotypes of the two phonic types with individuals sampled from across the Indo-Malay range supports the hypothesis that, in India, two cryptic species are present. The Indian 98-kHz phonic bats formed a monophyletic clade with bats from all other regional populations sampled, to the exclusion of the Indian 85-kHz bats. In India, the two forms showed 12–13% sequence divergence and we propose that the name Hipposideros khasiana for bats of the 85-kHz phonic type. Bats of the 98-kHz phonic type formed a monophyletic group with bats from Myanmar, and corresponded to Hipposideros grandis , which is suggested to be a species distinct from Hipposideros larvatus . Differences in echolocation call frequency among populations did not reflect phylogenetic relationships, indicating that call frequency is a poor indicator of evolutionary history. Instead, divergence in call frequency probably occurs in allopatry, possibly augmented by character displacement on secondary contact to facilitate intraspecific communication.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 119–130.     

Molecular phylogeny of hipposiderid bats from Southeast Asia and evidence of cryptic diversity

Old World leaf-nosed bats (Hipposideridae) are among the most widespread and ecologically diverse groups of insectivorous bats in the Old World tropics. However, phylogenetic relationships in Hipposideridae are poorly resolved at both the generic and species levels, and deep genetic divergence within several Southeast Asian species suggests that current taxonomy underestimates hipposiderid diversity in this region. We used mitochondrial and nuclear sequence data to conduct the first extensive molecular phylogenetic analysis of Southeast Asian hipposiderid bats. Inclusion of multiple samples per taxon allowed testing for evidence of evolutionarily distinct lineages within taxa currently defined as single species. In contrast to earlier phylogenies based on morphometrics, molecular data support monophyly of Hipposideros, but are ambiguous regarding the monophyly of Hipposideridae. With a few exceptions, molecular data also support currently recognized species groups classified by qualitative morphological characters. Widespread paraphyly and polyphyly within many currently recognized species of Hipposideros indicates that evolutionary diversity in the genus is underrepresented by current nomenclature. Comparison of available morphological and echolocation data suggest that both geographic isolation and ecological selection have contributed to the diversification of Southeast Asian hipposiderid bats.     

Recent loss of vitamin C biosynthesis ability in bats

The traditional assumption that bats cannot synthesize vitamin C (Vc) has been challenged recently. We have previously shown that two Old World bat species (Rousettus leschenaultii and Hipposideros armiger) have functional L-gulonolactone oxidase (GULO), an enzyme that catalyzes the last step of Vc biosynthesis de novo. Given the uncertainties surrounding when and how bats lost GULO function, exploration of gene evolutionary patterns is needed. We therefore sequenced GULO genes from 16 bat species in 5 families, aiming to establish their evolutionary histories. In five cases we identified pseudogenes for the first time, including two cases in the genus Pteropus (P. pumilus and P. conspicillatus) and three in family Hipposideridae (Coelops frithi, Hipposideros speoris, and H. bicolor). Evolutionary analysis shows that the Pteropus clade has the highest ω ratio and has been subjected to relaxed selection for less than 3 million years. Purifying selection acting on the pseudogenized GULO genes of roundleaf bats (family Hipposideridae) suggests they have lost the ability to synthesize Vc recently. Limited mutations in the reconstructed GULO sequence of the ancestor of all bats contrasts with the many mutations in the ancestral sequence of recently emerged Pteropus bats. We identified at least five mutational steps that were then related to clade origination times. Together, our results suggest that bats lost the ability to biosynthesize vitamin C recently by exhibiting stepwise mutation patterns during GULO evolution that can ultimately lead to pseudogenization.     

Standard energetics of leaf-nosed bats (Hipposideridae): its relationship to intermittent- and protracted-foraging tactics in bats and birds

Basal rates of metabolism within the insectivorous genera Hipposideros and Ascelliscus, Old World leaf-nosed bats (Hipposideridae), ranged from 58% to 77% of the mammalian standard. The larger species, Hipposideros diadema and Hipposideros maggietaylori, effectively thermoregulated at ambient temperatures down to 9 degrees C, whereas two smaller species, Hipposideros galeritus and Hipposideros cervinus, occasionally permitted body temperatures to fall below 32 degrees C. The low basal rates of metabolism in hipposiderids correlated with a predatory life-style characterized by intermittent flight from a perch to capture insects, a correlation similar to that found in nonpasserine birds. Intermittent-foraging bats and nonpasserines collectively had basal rates of metabolism that averaged 75% of those that pursue insects during protracted flight. However, no difference in basal rate was found between protracted- and intermittent-foraging passerines, which had basal rates 1.8- and 2.4-times those of protracted-foraging and intermittent-foraging bats and nonpasserines, respectively. Bats, swifts, and caprimulgids that enter torpor have basal rates that are 85% of those of similar species that do not enter torpor. Body mass, order affiliation, foraging mode, and propensity to enter into torpor collectively account for 97% of the variation in basal rate of metabolism in insectivorous bats and birds. Foraging style therefore appears to be a factor contributing to the diversity in endotherm energetics. Minimal thermal conductance in the genus Hipposideros ranged from 75% to 102% of the mammalian standard. Birds have minimal thermal conductances that are 75% of mammals and intermittent foragers have minimal conductances that are 78% of protracted foragers.     

‘Compromise’ in Echolocation Calls between Different Colonies of the Intermediate Leaf-Nosed Bat (Hipposideros larvatus)

Each animal population has its own acoustic signature which facilitates identification, communication and reproduction. The sonar signals of bats can convey social information, such as species identity and contextual information. The goal of this study was to determine whether bats adjust their echolocation call structures to mutually recognize and communicate when they encounter the bats from different colonies. We used the intermediate leaf-nosed bats (Hipposideros larvatus) as a case study to investigate the variations of echolocation calls when bats from one colony were introduced singly into the home cage of a new colony or two bats from different colonies were cohabitated together for one month. Our experiments showed that the single bat individual altered its peak frequency of echolocation calls to approach the call of new colony members and two bats from different colonies adjusted their call frequencies toward each other to a similar frequency after being chronically cohabitated. These results indicate that the ‘compromise’ in echolocation calls might be used to ensure effective mutual communication among bats.     

六种共栖蝙蝠的回声定位信号及翼型特征的比较

本研究于2006 年5 ～ 8 月在桂林市七星公园七星岩洞进行,比较分析了共栖2 科(蹄蝠科和蝙蝠科)6 种共75 只蝙蝠的回声定位信号和翼型特征。普氏蹄蝠的回声定位叫声为短而多谐波的CF/ FM 型,主频率为61.2±0.8 kHz, 具有高翼载、低翼展比和中等翼尖指数; 大蹄蝠的回声定位叫声为单CF/ FM 型,主频率为68. 6 ±0.7 kHz,具有高翼载、低翼展比和中等翼尖指数;中蹄蝠的回声定位叫声为单CF / FM 型,主频率为85.2 ±0.5 kHz,具有中等翼载、低翼展比和中等翼尖指数;高颅鼠耳蝠的回声定位叫声为长带宽的FM 型,主频率为50.7 ±3.8 kHz,具有低翼载、中等翼展比和低翼尖指数;大足鼠耳蝠回声定位叫声为FM 型,主频率为39.9 ±3.2 kHz,具有中等翼载、低翼展比和高翼尖指数;绒山蝠回声定位叫声为短而多谐波的FM 型,主频率为49.0± 0. 4 kHz,具有高翼载、中等翼展比和低翼尖指数。经单因素方差分析表明,6 种蝙蝠之间绝大部分的形态和声音参数差异显著(One-way ANOVA,P < 0. 05)。以上结果说明,6 种同地共栖蝙蝠种属特异的回声定位叫声
和形态结构体现出了相互之间的生态位分离,从而降低了种间竞争压力,使得6 种蝙蝠能够同地共存。     

Comparative collicular tonotopy in two bat species adapted to movement detection,Hipposideros speoris andMegaderma lyra

Summary The tonotopic organization of the inferior colliculus (IC) in two echolocating bats,Hipposideros speoris andMegaderma lyra, was studied by multiunit recordings.InHipposideros speoris frequencies below the range of the echolocation signals (i.e. below 120 kHz) are compressed into a dorsolateral cap about 400–600 m thick. Within this region, neuronal sheets of about 4–5 m thickness represent a 1 kHz-band.In contrast, the frequencies of the echolocation signals (120–140 kHz) are overrepresented and occupy the central and ventral parts of the IC (Fig. 3). In this region, neuronal sheets of about 80 m thickness represent a 1 kHz-band. The largest 1 kHz-slabs (400–600 m) represent frequencies of the pure tone components of the echolocation signals (130–140 kHz).The frequency of the pure tone echolocation component is specific for any given individual and always part of the overrepresented frequency range but did not necessarily coincide with the BF of the thickest isofrequency slab. Thus hipposiderid bats have an auditory fovea (Fig. 10).In the IC ofMegaderma lyra the complete range of audible frequencies, from a few kHz to 110 kHz, is represented in fairly equal proportions (Fig. 7). On the average, a neuronal sheet of 30 m thickness is dedicated to a 1 kHz-band, however, frequencies below 20 kHz, i.e. below the range of the echolocation signals, are overrepresented.Audiograms based on thresholds determined from multiunit recordings demonstrate the specific sensitivities of the two bat species. InHipposideros speoris the audiogram shows two sensitivity peaks, one in the nonecholocating frequency range (10–60 kHz) and one within the auditory fovea for echolocation (130–140 kHz).Megaderma lyra has extreme sensitivity between 15–20 kHz, with thresholds as low as –24 dB SPL, and a second sensitivity peak at 50 kHz (Fig. 8).InMegaderma lyra, as in common laboratory mammals, Q_10dB-values of single units do not exceed 30, whereas inHipposideros speoris units with BFs within the auditory fovea reach Q_10dB-values of up to 130.InMegaderma lyra, many single units and multiunit clusters with BFs below 30 kHz show upper thresholds of 40–50 dB SPL and respond most vigorously to sound intensities below 30 dB SPL (Fig. 9). Many of these units respond preferentially or exclusively to noise. These features are interpreted as adaptations to detection of prey-generated noises.The two different tonotopic arrangements (compare Figs. 3 and 7) in the ICs of the two species are correlated with their different foraging behaviours. It is suggested that pure tone echolocation and auditory foveae are primarily adaptations to echo clutter rejection for species foraging on the wing close to vegetation.Abbreviations BF Best frequency - CF constant frequency - FM frequency modulated - IC inferior colliculus - HS Hipposideros speoris     

Electrical responses of the superior olivary complex in Vespertilionidae and Rhinolophidae to ultrasonic stimuli with different fill frequency

Global and single unit responses of the superior olivary complex were investigated during ultrasonic stimulation at different frequencies in two species of bats from the Vespertilionidae, which emit frequency-modulated signals and the Rhinolophidae, which utilize almost monochromatic (80 ± 1 kHz) echolocation cries. Maximal sensitivity to ultrasound in the Vespertilionidae was found at frequencies of 10–40 kHz, and in the Rhinolophidae also within the range 10–40 kHz but with a second increase in sensitivity in the region 82–86 kHz. Sharply tuned neurons were more numerous in the Rhinolophidae than in the Vespertilionidae. Neurons whose response in the echolocation frequency band changed in character depending on the fill frequency of the stimulus were found in Rhinolophidae: a phasic discharge occurs over a wide range of frequencies and a tonic discharge at the characteristic frequency; the latter was also observed over a limited range of intensities.A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 5, No. 1, pp. 33–39, January–February, 1973.     

Variation in the Sonar of an Aerial-hawking Bat (Eptesicus nilssonii)

Echolocation calls from 10 individually marked female northern bats (Eptesicus nilssonii) were recorded as the bats foraged at three distinct feeding sites (territories) near their maternity roost in southern Sweden (57° N). In addition, recordings of unmarked bats were made in northern Sweden (65° N). The frequency at maximum amplitude of “search phase” echolocation pulses was bimodally distributed, with peaks around 29–30 kHz and 31–32 kHz and was negatively correlated with pulse duration. The frequency at maximum amplitude was related to flight altitude (bats used higher frequencies when they flew near the ground) and also differed among the feeding sites. Hence, much of the variation, probably including regional differences, was behavioural and is interpreted as short term (in the order of s or min) adaptation to current foraging situations. Variation among individual bats, caused by age and size, seemed to be of less importance. Individuals did not use exclusive frequency bands.     

Characteristics of unit responses of the cochlear nuclei of bats (Rhinolophidae) to single and paired ultrasonic stimuli

Characteristics of single unit responses of the cochlear nuclei of greater horseshoe bats to ultrasonic stimuli with a filling frequency within the echolocation range were investigated. In most neurons three unconnected regions of responses were found, with characteristic frequencies equal to 1/2 and 1/3 of the basic frequency, within the range 80–90 kHz. The response regions had inhibitory zones, one of which was higher than the basic characteristic frequency whereas the other two were overlapped by the complementary response regions. Selectivity of the neurons to the frequency of stimulation increased with a change in the characteristic frequency from 90 to 80 kHz; it was maximal in the band 80–80.5 kHz.     

毛细属线虫中国新记录种的记述(嘴刺目:鞭虫科)

毛细属Capillaria中国新记录种——越南毛细线虫C.vietnamensis Meszaros,1973,采自广东省惠州市小蹄蝠Hipposideros pomona、大黄蝠Scotophilus heathi,龙门县长翅蝠Miniopterus schreibersi,广州市扁颅蝠Tylonycteris pa...     

The cochlear frequency map of the mustache bat,Pteronotus parnellii

The frequency-place map of the cochlea of mustache bats was constructed by the analysis of HRP-transport patterns in spiral ganglion cells following iontophoretic tracer injections into cochlear nucleus regions responsive to different frequencies. The cochlea consists of 5 half turns (total length 14.3 mm) and the representation of certain frequency bands can be assigned to specific cochlear regions: The broad high frequency range between 70 and 111 kHz is represented in the most basal half turn within only 3.2 mm. This region is terminated apically by a distinct narrowing of the scala vestibuli that coincides with a pronounced increase in basilar membrane (BM) thickness. The narrow intermediate frequency range between 54 and 70 kHz is expanded onto 50% of cochlear length between 4.0 and 11.1 mm distance from apex. The frequency range around 60 kHz, where the tuning characteristics of the auditory system are exceptionally sharp, is located in the center of this expanded BM-region in the second half turn within a maximum of innervation density. These data can account for the vast overrepresentation of neurons sharply tuned to about 60 kHz at central stations of the auditory pathway. In the cochlear region just basal to the innervation maximum, where label from injections at 66 and 70 kHz was found, a number of morphological specializations coincide: the BM is maximally thickened, innervation density is low, the spiral ligament is locally enlarged, and the 'thick lining', a dense covering of the scala tympani throughout the basal halfturn, suddenly disappears. Low frequencies up to 54 kHz are represented within the apical half turns over a 4 mm span of the basilar membrane. The data are compared to the cochlea of horseshoe bats and the possible functional role of the morphological discontinuities for sharp tuning and the generation of otoacoustic emissions is discussed.     

基于核基因RAG1部分序列探讨菊头蝠科和蹄蝠科的系统发育关系

应用核基因重组激活基因1(Recombination activating gene,RAG1)部分序列对贵州9种菊头蝠和5种蹄蝠的系统发育关系进行了研究,运用贝叶斯法(Bayesian inference,BI)、最大似然法(Maximum likelihood,ML)和邻接法(Neighbor-joining,NJ)构建了系统进化树。研究结果表明:1.菊头蝠科和蹄蝠科形成两个平行的分支,支持它们是两个平行科的结论。2.马铁菊头蝠是菊头蝠科中第一个独立出来的亚分支,与其余8种菊头蝠的亲缘关系最远;Rhinolophus sp.与大菊头蝠聚合为一支,推测Rhinolophus sp.可能是大菊头蝠,或者是大菊头蝠的近缘种,它们是继马铁菊头蝠之后分离出来的第二亚分支;中华菊头蝠、栗黄菊头蝠、中菊头蝠聚合为第三亚分支,贵州菊头蝠、大耳菊头蝠、菲菊头蝠聚合为第四亚分支,此两分支形成姊妹群。3.在蹄蝠科的亚分支中,三叶蹄蝠、小蹄蝠、普氏蹄蝠依次分离出来,最后分出的是大蹄蝠和中蹄蝠。     

Convergence of reference frequencies by multiple CF–FM bats (Rhinolophus ferrumequinum nippon) during paired flights evaluated with onboard microphones

The constant frequency component of the second harmonic (CF(2)) of echolocation sounds in Rhinolophus ferrumequinum nippon were measured using onboard telemetry microphones while the bats exhibited Doppler-shift compensation during flights with conspecifics. (1) The CF(2) frequency of pulses emitted by individual bats at rest (F (rest)) showed a long-term gradual decline by 0.22?kHz on average over a period of 3?months. The mean neighboring F (rest) (interindividual differences in F (rest) between neighboring bats when the bats were arranged in ascending order according to F (rest)) ranged from 0.08 to 0.11?kHz among 18 bats in a laboratory colony. (2) The standard deviation of observed echo CF(2) (reference frequency) for bats during paired flights ranged from 50 to 90?Hz, which was not significantly different from that during single flights. This finding suggests that during paired flights, bats exhibit Doppler-shift compensation with the same accuracy as when they fly alone. (3) In 60?% (n?=?29) of the cases, the difference in the reference frequency between two bats during paired flights significantly decreased compared to when the bats flew alone. However, only 15?% of the cases (n?=?7) showed a significant increase during paired flights. The difference in frequency between two bats did not increase even when the reference frequencies of the individuals were not statistically different during single flights.     

Rudnicula goffi n. sp. (Acariformes: Trombiculidae) from the diadem leaf-nosed bat Hipposideros diadema (Geoffroy) (Chiroptera: Hipposideridae) on Bali,Indonesia

Systematic Parasitology - A new species of chigger mite, Rudnicula goffi n. sp., is described from the diadem leaf-nosed bat Hipposideros diadema (Geoffroy) (Hipposideridae) caught in the Goa...     

Frequency discrimination threshold at search call frequencies in the echolocating bat, Eptesicus fuscus

While searching for prey in open spaces, Epteisicus fuscus emits long-duration, downward frequency-modulated calls which cover a frequency band of about 28-22 kHz. In the ascending auditory pathways of E. fuscus, neurons tuned to these search call frequencies are characterised by a remarkably high frequency selectivity and very sensitive absolute thresholds. We investigated whether this narrow tuning is reflected in an exceptional psychoacoustic frequency discrimination ability. The average frequency difference limen of E. fuscus at search call frequencies determined in a two-alternative, forced-choice experiment amounted to about 420 Hz, corresponding to a Weber ratio of 0.017. This value is similar to those found in non-echolocating mammals, and an order of magnitude larger than the frequency difference limens of bats emitting constant-frequency call components. We discuss these differences in frequency difference limen, and relate them to different echolocation strategies.     

Molecular phylogeny of hipposiderid bats (Chiroptera: Hipposideridae) from Solomon Islands and Cape York Peninsula,Australia

Throughout the Old World Tropics, examples of cryptic diversity have been demonstrated in leaf‐nosed bats from the Family Hipposideridae. In addition to cryptic diversity, the widespread distributions and graded morphologies of some species render the taxonomy of this family poorly resolved. We used sequences of the mitochondrial gene ND2 and the nuclear gene RAG1 to reconstruct molecular phylogenetic relationships for hipposiderid species. Specimens from Solomon Island and Cape York Peninsula, Australia, were sequenced and combined with published sequences from Southeast Asia. Our results suggest the presence of undescribed diversity in northern Australia. Sequences of ND2 and RAG1 for samples of Cape York Hipposideros ater were distinct from those of Southeast Asia, with corrected sequence divergence estimates for the mitochondrial gene exceeding 23%. In comparison, there was no cryptic diversity detected in Solomon Island specimens when compared to conspecifics from Southeast Asia. On the contrary, three species members of the diadema morphological group from the region exhibit variability in the mitochondrial gene ND2, far below that of other species included in the analyses.     

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

Summary The postnatal development of midbrain tonotopy was investigated in the inferior colliculus (IC) of the south Indian CF-FM batHipposideros 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.Abbreviations BF best frequency - CF constant frequency - Cer cerebellum - CN cochlear nucleus - CO auditory cortex - CUF cuneiform nucleus - DAB days after birth - FAL forearm length - FM frequency modulation - IC inferior colliculus - NLL nucleus of the lateral lemniscus - PAG periaqueductal gray - SC superior colliculus     

Modelling the prey detection performance of Rhinonicteris aurantia (Chiroptera: Hipposideridae) in different atmospheric conditions discounts the notional role of relative humidity in adaptive evolution

We examined a recent notion that differences in echolocation call frequency amongst geographic groups of constant frequency (CF)-emitting bats is the result of a trade-off between maximising prey detection range at lower frequencies and enhancing small-prey resolution at higher frequencies in different atmospheric (relative humidity; RH) environments. Isolated populations of the endemic Australian orange leaf-nosed bat Rhinonicteris aurantia were used as an example since geographic isolation in different environments has been a precursor to differences in their characteristic echolocation call frequencies (mean difference c. 6 kHz; means of 114.64 and 120.99 kHz). The influence of both atmospheric temperature and RH on maximum prey detection range was explored through mathematical modelling. This revealed that temperature was of similar importance to relative humidity and that under certain circumstances, each could reduce the effect of the other on ultrasound attenuation rates. The newly developed models contain significant conceptual improvements in method compared to other recent approaches, and can be applied to the situation of any other species of bat. For a given set of atmospheric conditions, the prey detection range of R. aurantia was reduced slightly when call frequency increased by 6 kHz, but an increase in RH, temperature or both reduced detection range significantly. A similar trend was also evident in prey detection volume ratios calculated for the same conditions. Spatial volume ratios were applied to assess the impact of changed atmospheric conditions and prey size on foraging ecology. Reductions in detection range associated with increases in RH and/or temperature also varied in relation to the size (cross sectional area) of insect prey. Modelling demonstrated that small (6 kHz) movements in call frequency could not compensate for the changes in prey detection range and spatial detection volumes that result from significant changes in atmospheric temperature or RH. The notion that differences in RH are the primary cause leading to adaptive evolution and speciation in CF-emitting bats by precipitating intraspecific differences in the mean call frequency of geographically isolated bat populations was not supported by the results of this case study.     

Ontogenesis of the echolocation system in the rufous horseshoe bat,Rhinolophus rouxi (Audition and vocalization in early postnatal development)

1. The development of vocalization and hearing was studied in Sri Lankan horseshoe bats (Rhinolophus rouxi) during the first postnatal month. The young bats were caught in a nursing colony of rhinolophids in which birth took place within a two week period. 2. The new-born bats emitted isolation calls through the mouth. At the beginning these calls consisted of pure tones with frequencies below 10 kHz (Fig. 1). During the first postnatal week the call frequency increased to about 15 kHz, and the fundamental was augmented by two to four harmonics. No evoked potentials to pure tone stimuli could be elicited in the inferior colliculus of this age group, i.e., auditory processing at the midbrain level was not demonstrable. 3. Evoked potentials were first recorded in the second week, broadly tuned to 15-45 kHz, with a maximum sensitivity between 15-25 kHz. In the course of the second week, however, higher frequencies up to 60 kHz became progressively incorporated into the audiogram (Fig. 3). The fundamental frequency of the multiharmonic isolation calls, emitted strictly through the mouth, increased to about 20 kHz. 4. In the bats' third postnatal week an increased hearing sensitivity (auditory filter) emerged, sharply tuned at frequencies between 57 and 60 kHz (Fig. 4e). The same individuals were also the first to emit long constant frequency echolocation calls through the nostrils (Fig. 4c). The energy of the calls was arranged in harmonic frequency bands with the second harmonic exactly tuned to the auditory filter. These young bats continued to emit isolation calls through the mouth, which were, however, not harmonically related to the echolocation calls (Fig. 4b, d). 5. During the fourth week, both the auditory filter and the matched echolocation pulses (the second harmonic) shifted towards higher frequencies (Fig. 5). During the fifth week the fundamental frequency of the calls was progressively attenuated, and both the second harmonic of the pulses and the auditory filter reached the frequency range typical for adult bats of 73-78 kHz (Fig. 6). 6. The development of audition and vocalization is discussed with regard to possible interactions of both subsystems, and their incorporation into the active orientation system of echolocation.