Echolocation in the large molossid bats Eumops glaucinus and Nyctinomops macrotis

Eumops glaucinus and Nyctinomops macrotis, the largest molossid bats in Cuba, were investigated. Both species of bats share the same guild in the island and are similar in size, which allow the prediction of overlapping echolocation inventories following both the vocal plasticity hypothesis and the scaling hypothesis. In addition, large body size predicts the emission of low frequency calls in the human audible range. Calls recorded during hunting show that the bats' echolocation repertoires are very similar and of low frequency, with most differences in search calls. Matches were found in the calls' design, duration, slope, bandwidth, and spectral parameters. Statistical differences between search calls are consistent with the predictions from the scaling hypothesis, considering that E. glaucinus is only slightly larger than N. macrotis. The echolocation calls emitted by both species are in the frequency range below 20-25 kHz, which identifies both species as the only ones with echolocation in the human audible range in Cuba.     

Discrimination of wingbeat motion by bats,correlated with echolocation sound pattern

Summary Bats of the species Rhinolophus rouxi, Hipposideros lankadiva and Eptesicus fuscus were trained to discriminate between two simultaneously presented artificial insect wingbeat targets moving at different wingbeat rates. During the discrimination trials, R. rouxi, H. lankadiva and E. fuscus emitted long-CF/FM, short-CF/FM and FM echolocation sounds respectively. R. rouxi, H. lankadiva and E. fuscus were able to discriminate a difference in wingbeat rate of 2.7 Hz, 9.2 Hz and 15.8 Hz, respectively, between two simultaneously presented targets at an absolute wingbeat rate of 60 Hz, using a criterion of 75% correct responses.The performance of the different bat species is correlated with the echolocation signal design used by each species, particularly with the presence and relative duration of a narrowband component preceding a broadband FM component. These results provide behavioral evidence supporting the hypothesis that bats that use CF/FM echolocation sounds have adaptations for the perception of insect wingbeat motion and that long-CF/FM species are more specialized for this task than short-CF/FM species.Abbreviations CF constant frequency - FM frequency modulation     

Echolocation calls in Central American emballonurid bats: signal design and call frequency alternation

In southern Central America, 10 species of emballonurid bats occur, which are all aerial insectivores: some hunt flying insects preferably away from vegetation in open space, others hunt in edge space near vegetation and one species forages mainly over water. We present a search call design of each species and link signal structure to foraging habitat. All emballonurid bats use a similar type of echolocation call that consists of a central, narrowband component and one or two short, frequency-modulated sweeps. All calls are multi-harmonic, generally with most energy concentrated in the second harmonic. The design of search calls is closely related to habitat type, in particular to distance of clutter. Emballonurid bats foraging in edge space near vegetation and over water used higher frequencies, shorter call durations and shorter pulse intervals compared with species mostly hunting in open, uncluttered habitats. Peak frequency correlated negatively with body size. Regular frequency alternation between subsequent calls was typical in the search sequences of four out of 10 species. We discuss several hypotheses regarding the possible role of this frequency alternation, including species identification and partitioning of acoustic channels. Furthermore, we propose a model of how frequency alternation could increase the maximum detection distance of obstacles by marking search calls with different frequencies.     

Echolocation behavior of rufous horseshoe bats hunting for insects in the flycatcher-style

Summary The echolocation behavior ofRhinolophus rouxi when waiting for insects on a perch and when pursuing them in short hunting flights is described. It reveals that flycatcher-style hunting is one foraging strategy utilized by this species. It also suggests that bats use fluttering target information to classify their prey.     

Echolocation, olfaction and fruit display: how bats find fruit of flagellichorous cucurbits

1. The relationship between plant morphology and the senses used by dispersal agents to find fruit was examined. 'Flagellichory' (fruit borne on pendulous structures), a costly morphology associated with dispersal by bats, is focused on.
2. Using Gurania spinulosa , a flagellichorous vine, and its major dispersal agent, Phyllostomus hastatus , the hypothesis was tested that flagellichory increases the conspicuousness of fruit to bats that use echolocation to find fruit.
3. The responses of wild-caught P. hastatus to various fruiting branch morphologies and fruit odour were recorded. Phyllostomus hastatus used echolocation rather than olfaction to detect fruit, and consistently chose fruit displayed on pendulous leafless branches, ignoring fruit held among leaves on horizontal branches.
4. By comparing echolocation signals with the distance between fruiting branches of G. spinulosa and surrounding vegetation, it was shown that pendulous fruiting branches present clear, clutter-free targets that can be detected by echolocating bats. This is the first demonstration of neotropical frugivorous bats using echolocation to find fruit.     

Managing anthropogenic driven range expansion behaviourally: Mediterranean bats in desert ecosystems

Bat abundance and activity in deserts is affected by the distribution of water sources. Kuhl’s pipistrelle (Pipistrellus kuhlii) is one of several bat species that penetrated the Negev desert from Mediterranean habitats following anthropogenic development, and potentially competes with desert dwelling bats of the same guild. Pipistrellus kuhlii is an obligate drinker requiring a clear ‘swoop zone’ to drink. Thus, to reduce the competitive load that P. kuhlii has on desert dwelling species, we tested the effectiveness of obstructions above water surfaces, which we hypothesised would reduce the drinking ability of P. kuhlii, thus providing desert species a competitive edge. We obstructed the water surface of a swimming pool and successfully prevented P. kuhlii from drinking. Next, we manipulated natural water sources used by bats for foraging. This was done on two spatial and temporal scales: on 11 adjacent ephemeral water pools in Nahal Zin where each treatment was applied for several nights and on eight isolated natural pools in the Negev where treatments were alternated every 20 min. Manipulation of adjacent pools had no effect on activity levels of bats, but in isolated pools, the obstruction reduced the proportion of P. kuhlii activity by up to 15%. Although this tool cannot be used for management without further examination of its long-term effects on other bats and wildlife, this study does demonstrate its feasibility. Conservation schemes aiming to mitigate effects of undesirable species can be designed by identifying ecological differences between competing species, and manipulating the environment accordingly.     

Echolocation versus echo suppression in humans

Several studies have shown that blind humans can gather spatial information through echolocation. However, when localizing sound sources, the precedence effect suppresses spatial information of echoes, and thereby conflicts with effective echolocation. This study investigates the interaction of echolocation and echo suppression in terms of discrimination suppression in virtual acoustic space. In the ‘Listening’ experiment, sighted subjects discriminated between positions of a single sound source, the leading or the lagging of two sources, respectively. In the ‘Echolocation’ experiment, the sources were replaced by reflectors. Here, the same subjects evaluated echoes generated in real time from self-produced vocalizations and thereby discriminated between positions of a single reflector, the leading or the lagging of two reflectors, respectively. Two key results were observed. First, sighted subjects can learn to discriminate positions of reflective surfaces echo-acoustically with accuracy comparable to sound source discrimination. Second, in the Listening experiment, the presence of the leading source affected discrimination of lagging sources much more than vice versa. In the Echolocation experiment, however, the presence of both the lead and the lag strongly affected discrimination. These data show that the classically described asymmetry in the perception of leading and lagging sounds is strongly diminished in an echolocation task. Additional control experiments showed that the effect is owing to both the direct sound of the vocalization that precedes the echoes and owing to the fact that the subjects actively vocalize in the echolocation task.     

Sex and segregation in temperate bats

Many temperate insectivorous bats show marked sexual segregation during the summer, but in spectacular, pre-hibernation swarming, gather at caves to mate. In many species, sexual segregation is probably due to a gradient in aerial insect availability that confines females to lower elevations, where high reproductive costs are met by an abundant and reliable food supply. In the hawking and trawling Myotis daubentonii, we show that alongside inter-sexual segregation, there is intra-male segregation and suggest that this results from the exclusion of most males from high-quality habitat. These apparently excluded males suffer reduced foraging efficiency and mating success relative to males that roost with the females in summer. Changes in resources and behaviour at the end of the summer lead to a change in strategy that gives all males a chance to mate during swarming, but this does not overcome the paternity advantage to males that spend the summer with the females.     

Non-kin cooperation in bats

Many bats are extremely social. In some cases, individuals remain together for years or even decades and engage in mutually beneficial behaviours among non-related individuals. Here, we summarize ways in which unrelated bats cooperate while roosting, foraging, feeding or caring for offspring. For each situation, we ask if cooperation involves an investment, and if so, what mechanisms might ensure a return. While some cooperative outcomes are likely a by-product of selfish behaviour as they are in many other vertebrates, we explain how cooperative investments can occur in several situations and are particularly evident in food sharing among common vampire bats (Desmodus rotundus) and alloparental care by greater spear-nosed bats (Phyllostomus hastatus). Fieldwork and experiments on vampire bats indicate that sharing blood with non-kin expands the number of possible donors beyond kin and promotes reciprocal help by strengthening long-term social bonds. Similarly, more than 25 years of recapture data and field observations of greater spear-nosed bats reveal multiple cooperative investments occurring within stable groups of non-kin. These studies illustrate how bats can serve as models for understanding how cooperation is regulated in social vertebrates.     

Moult, flight performance and wingbeat kinematics during take-off in European starlings Sturnus vulgaris

The effects of natural moult on avian flight performance have received relatively little attention, yet moult is an important part of the annual cycle. Quantification of flight costs will help to explain the diversity of moult patterns observed in avian taxa. Take‐off from the ground requires a high power output from the flight muscles compared to other modes of flight, and is an important feature of foraging and predation escape. The present study was designed to quantify the effect of natural moult and new plumage on the take‐off strategy, kinematics, and flight performance of European starlings Sturnus vulgaris. A high‐speed (185 Hz) cine camera was used to film seven European starlings on three occasions: session 1, two weeks prior to the onset of moult; session 2, during mid‐moult; and session 3, two weeks after full plumage had re‐grown. From subsequent film analysis, we assessed take‐off speed and angle, the energy gained per wingbeat, and wingtip kinematics. Take‐off strategy (measured by angle and speed) altered through the course of the three experimental sessions, i.e. ascent angle decreased and take‐off speed increased. Energy gained per wingbeat did not vary, suggesting there was no significant decrease in flight performance due to moult, but there was a significant improvement in take‐off performance due to renewal of flight plumage. Wingbeat amplitude increased in association with moult and after flight plumage had been completely renewed. The European starlings incurred relatively minor flight costs due to moult, when comparing before‐moult with during‐moult take‐off performance. The apparent absence of additional flight costs associated with moult may reflect a decreased mechanical performance of year‐old feathers (which are replaced during the moult) and may also help to explain the relatively long duration of the moult in this species. This study also provides evidence of the benefits of plumage renewal, as take‐off performance is improved after moult has been completed.     

Different Auditory Feedback Control for Echolocation and Communication in Horseshoe Bats

Auditory feedback from the animal''s own voice is essential during bat echolocation: to optimize signal detection, bats continuously adjust various call parameters in response to changing echo signals. Auditory feedback seems also necessary for controlling many bat communication calls, although it remains unclear how auditory feedback control differs in echolocation and communication. We tackled this question by analyzing echolocation and communication in greater horseshoe bats, whose echolocation pulses are dominated by a constant frequency component that matches the frequency range they hear best. To maintain echoes within this “auditory fovea”, horseshoe bats constantly adjust their echolocation call frequency depending on the frequency of the returning echo signal. This Doppler-shift compensation (DSC) behavior represents one of the most precise forms of sensory-motor feedback known. We examined the variability of echolocation pulses emitted at rest (resting frequencies, RFs) and one type of communication signal which resembles an echolocation pulse but is much shorter (short constant frequency communication calls, SCFs) and produced only during social interactions. We found that while RFs varied from day to day, corroborating earlier studies in other constant frequency bats, SCF-frequencies remained unchanged. In addition, RFs overlapped for some bats whereas SCF-frequencies were always distinctly different. This indicates that auditory feedback during echolocation changed with varying RFs but remained constant or may have been absent during emission of SCF calls for communication. This fundamentally different feedback mechanism for echolocation and communication may have enabled these bats to use SCF calls for individual recognition whereas they adjusted RF calls to accommodate the daily shifts of their auditory fovea.     

高颅鼠耳蝠回声定位声波特征与分析

在自建的野外实验事内,录制雌雄成体高颅鼠耳蝠Myotis siligorensis不同行为状态(飞行、爬行、手持)回声定位声波,利用单因素方差分析(One-Way ANOVA)对不同状态及不同性别的回声定位声波参数进行均值多重比较和显著性差异分析.结果 表明,高颅鼠耳蝠回声定位声波为长的宽频带调频(FM)声波,有1~3个谐波,但能量主要集中在第一谐波.不同行为状态下,高颅鼠耳蝠声脉冲持续时间、声脉冲间隔和能率环均存在显著性差异(P<0.05),第一谐波起始频率和终止频率、主频率、带宽差异不显著;第一谐波终止频率、带宽、声脉冲持续时间和能率环的性别差异显著(P<0.05),具有性别二态性.高颅鼠耳蝠回声定位声波特征体现了其在捕食策略和捕食生境方面的生态适应.     

Arctiid moth clicks can degrade the accuracy of range difference discrimination in echolocating big brown bats,Eptesicus fuscus

Summary Four big brown bats (Eptesicus fuscus) born and raised in captivity were trained using the Yes/No psychophysical method to report whether a virtual sonar target was at a standard distance or not. At threshold bats were able to detect a minimum range difference of 6 mm (a t of 36 s).Following threshold determinations, a click burst 1.8 ms long containing 5 pulses from the ruby tiger moth, Phragmatobia fuliginosa (Arctiidae), was presented randomly after each phantom echo. The sound energy of the click burst was -4 dB relative to that of the phantom echo. Clicks presented for the very first time could startle naive bats to different degrees depending on the individual.The bats' performance deteriorated by as much as 4000% when the click burst started within a window of about 1.5 ms before the phantom echo (Fig. 4). Even when one of ten phantom echoes was preceded by a click burst, the range difference discrimination worsened by 200% (Fig. 9). Hence, clicks falling within the 1.5 ms time window seem to interfere with the bat's neural timing mechanism.The clicks of arctiid moths appear to serve 3 functions: they can startle naive bats, interfere with range difference determinations, or they can signal the moth's distastefulness, as shown in earlier studies.Abbreviations peSPL peak equivalent sound pressure level - sd standard deviation - FM frequency modulation - CF constant frequency - EPROM erasable programmable read only memory     

Neural mechanisms of target ranging in FM bats: physiological evidence from bats and frogs

Echolocating bats assess target range by the delay in echo relative to the emitted sonar pulse. Earlier studies in FM bats showed that a population of neurons in auditory centers above the inferior colliculus (IC) is tuned to echo delay, with different neurons tuned to different echo delays. A building block for delay-tuned responses is paradoxical latency shift (PLS), featuring longer response latencies to more intense sounds. PLS is first created in the IC, where neurons exhibit unit-specific quantum increase in response latency with increasing sound level. Other IC neurons display oscillatory discharges whose period is unit-specific and level tolerant, indicating that this is attributable to cell’s intrinsic properties. High-threshold inhibition of oscillatory discharge produces PLS, indicating that oscillatory discharge is a building block for PLS. To investigate the cellular basis of oscillatory discharges, we performed whole-cell patch-clamp recordings from IC neurons in leopard frogs (which also exhibit oscillatory discharges and PLS). These recordings show that IC neurons are heterogeneous displaying diverse biophysical phenotypes; each phenotype (and cell) has its own membrane time constant, input resistance, and strengths of I _h, I _kir, I _kv—these intrinsic properties give rise to cell-specific resonance which can be observed through current and afferent stimulations.