Characteristics of the acoustic field of interfering reflections and the echolocation hearing of the dolphin

A model of the acoustic field of interfering reflections from steel cylinders was developed. Analysis of the model showed the availability of great potential resources for a decrease of the influence of unwanted echoes and hence for increasing the signal-to-clatter ratio. The conformity of the available models of the echolocation hearing of the dolphin to the acoustic field of the clatter was considered. The participation of mandidle mental foramens in conducting the echo to the cochlea was considered. In this case the hearing aperture is determined by the dimensions of mental foramens, while the hearing base is determined by the distance between the mental foramens of the left and right mandible halves. There are good reasons to believe that the optimal dimensions of the aperture and the base of echolocation hearing of Odontoceti essentially increase the effectiveness of defense of their sonar from reverberation.     

Mechanisms of sound reception and conduction in the dolphin

Morphology of the dolphin’s lower jaw, model and behavioral experiments are discussed with the aim of exploring the mechanisms of sound reception and conduction to the lower jaw canals, taking into account the known concepts of acoustics and the theory of grouped antennas. It is shown that the left and right rows of mental foramens with the respective mandibular canal and tissues of the canals are forming the new external ear and the new external auditory duct whereby sound (in the frequency band of 0.1–160 kHz) is transmitted into the middle ear, in contrast to the dolphin’s nonfunctional outer ear. This new external ear is created by nature as a receiving array of traveling-wave antennas located in the throat of an acoustic horn (the respective mandibular canal). The results give reason to assume the existence of a similar new external ear in Odontoceti.     

Why do shrews twitter? Communication or simple echo-based orientation

Shrews are very vocal animals. We tested behaviourally whether the high-pitched laryngeal ‘twittering’ calls of as-yet unclear function serve for communication or echo-based orientation. We used a representative species from each of the two largest phylogenetic groups of shrews. In both species, experimental manipulation of substrate density, but not of the likelihood of conspecific presence, affected the shrews'' call rate when exploring an unknown environment. This adaptation of call rate to the degree of habitat clutter parallels bat echolocation and suggests that shrews may use the echoes and reverberations of their calls for identifying routes through their habitat or for probing habitat type. To assess the acoustic feasibility of shrew echo orientation, we ensonified shrew habitats in the field with an ‘artificial shrew’ (small speaker mounted close to a sensitive microphone). The data showed that shrew-like calls can indeed yield echo scenes useful for habitat assessment at close range, but beyond the range of the shrews'' vibrissae.     

A comprehensive computational model of animal biosonar signal processing

Computational models of animal biosonar seek to identify critical aspects of echo processing responsible for the superior, real-time performance of echolocating bats and dolphins in target tracking and clutter rejection. The Spectrogram Correlation and Transformation (SCAT) model replicates aspects of biosonar imaging in both species by processing wideband biosonar sounds and echoes with auditory mechanisms identified from experiments with bats. The model acquires broadband biosonar broadcasts and echoes, represents them as time-frequency spectrograms using parallel bandpass filters, translates the filtered signals into ten parallel amplitude threshold levels, and then operates on the resulting time-of-occurrence values at each frequency to estimate overall echo range delay. It uses the structure of the echo spectrum by depicting it as a series of local frequency nulls arranged regularly along the frequency axis of the spectrograms after dechirping them relative to the broadcast. Computations take place entirely on the timing of threshold-crossing events for each echo relative to threshold-events for the broadcast. Threshold-crossing times take into account amplitude-latency trading, a physiological feature absent from conventional digital signal processing. Amplitude-latency trading transposes the profile of amplitudes across frequencies into a profile of time-registrations across frequencies. Target shape is extracted from the spacing of the object’s individual acoustic reflecting points, or glints, using the mutual interference pattern of peaks and nulls in the echo spectrum. These are merged with the overall range-delay estimate to produce a delay-based reconstruction of the object’s distance as well as its glints. Clutter echoes indiscriminately activate multiple parts in the null-detecting system, which then produces the equivalent glint-delay spacings in images, thus blurring the overall echo-delay estimates by adding spurious glint delays to the image. Blurring acts as an anticorrelation process that rejects clutter intrusion into perceptions.     

Echolocation constraints of Daubenton's Bat foraging over water

1. Daubenton's Bats ( Myotis daubentonii ) foraging over a stream concentrated their activity over calm surfaces, avoiding an adjacent area with small ripples (< 3 cm high). Aerial insects were most abundant over the ripples, so insect distribution could not explain why the bats avoided this area.
2. The bats flew low over water and always ( N = 22) directed the head forwards, presumably emitting the echolocation beam parallel to the surface, thus minimizing clutter. At an angle of incidence of 30° there was significantly more clutter from the rippled water.
3. The ripples produced ultrasonic noises in the form of transient pulses at an average rate of 6·2 per second. In the present case, such pulses were common enough potentially to interfere with target detection by the bats. Transient noises and echo clutter from moving ripples may be the principal reason why bats generally avoid foraging low over turbulent water.
4. The target strength of a potential insect prey at the water surface and the source levels of the bats' searching signals were measured to use in estimating the echo level at the bat when it detects the prey. The echo level at detection (+ 38 dB sound pressure level) was about the same as the clutter level extrapolated to the detection distance. This suggests that Daubenton's Bat operates at very low signal-to-noise ratios when foraging for insects near the water surface.     

门齿孔位置在中国古人类化石与现代人群的表现及其演化意义

魏敦瑞在研究周口店北京直立人化石时发现,位于上颌骨硬腭表面的门齿孔位置在周口店标本靠后,而在现代人靠近齿槽。此后,门齿孔位置作为具有演化意义的形态特征被用于古人类学研究。迄今,对门齿孔位置在中国古人类化石表现的专门研究仅有周口店一件标本,而在现代中国人的数据尚属空白。鉴于此,本文对门齿孔位置在中国古人类化石以及现代中国人标本进行了观察、测量和数据统计。在此基础上,结合世界其他地区古人类数据资料,本文对门齿孔位置在中国古人类化石的表现特点及其演化意义进行了探讨。本研究发现,从更新世早期到更新世晚期,门齿孔位置在中国古人类呈现由后向前的总体变化趋势。更新世早期和中期直立人（郧县曲远河口、周口店）的门齿孔位置都比较靠后;中更新世晚期的部分中国古人类（大荔、长阳、华龙洞）的门齿孔位置前移,并与现代人接近,而金牛山和巢县门齿孔位置比较靠后,位于直立人范围;在更新世晚期,所有中国古人类都比较靠前,位于现代人变异范围。本文对现代人标本的观测显示,门齿孔位置在现代中国人比较靠前。现代人门齿孔大小及形态存在较大变异,这种表现特点在一定程度上影响对门齿孔位置及演化意义的判定。几乎全部现代人标本门齿孔前缘呈开放状态,门齿管从开口处的门齿孔向后上方呈不同程度倾斜走向。这一发现与魏敦瑞提出的现代人门齿管垂直走向的观点不同。综合本文采集的中国古人类化石、现代中国人标本门齿孔位置数据,以及世界其他地区古人类门齿孔位置数据资料,我们认为门齿孔位置在人类演化过程中呈现较规律的变化,门齿孔位置靠后应该是一项相对原始的特征。     

The detection of phantom targets in noise by serotine bats; negative evidence for the coherent receiver

Summary Using a target simulator three serotine bats,Eptesicus serotinus, were trained to judge whether a phantom target was present or absent. The echolocation sounds emitted by the bats during the detection were intercepted by a microphone, amplified and returned by a loudspeaker as an artificial echo, with a delay of 3.2 ms and a sound level determined by the overall gain and cry amplitude. The cry level of each pulse was measured and the echo level received by the bat was calculated. The target was presented in 50% of the trials and the gain adjusted using conventional up/down procedures. Under these conditions between 40 and 48 dB peSPL were required for 50% detection (Figs. 2, 3).In a subsequent experiment the phantom target was masked with white noise (N_o) with a spectrum level of –113 dB re. 1 Pa·Hz^–1/2. The thresholds were increased by 7–14 dB. Energy density (S) of a single pulse was measured and used to estimate S/N_o, which ranged from 36–49 dB at threshold. Theoretically the coherent receiver model predicts the ratio between hits and false alarms observed for the bats at a S/N_o of ca. 1–2 dB. Since the bats require 40–50 dB higher S/N_o (Fig. 3), this is taken as negative evidence for coherent reception (cross correlation).Furthermore, a strong sensitivity to clutter was found since there seemed to exist a fixed relationship between thresholds and clutter level.Abbreviations C clutter - Nbw noise in a specified bandwidth - No noise in i Hz bandwidth - peSPL peak equivalent sound pressure level - S signal energy - SD standard deviation - Y/N Yes/No psychometry - 2AFC two alternative forced choice psychometry     

BEHAVIORAL EVIDENCE FOR HEARING THROUGH THE LOWER JAW BY AN ECHOLOCATING DOLPHIN (TURSIOPS TRUNCATUS)

On the basis of disputed physiological evidence the fat-filled lower jaw of odontocete cetaceans has previously been hypothesized as the primary pathway to the inner ear for acoustic signals. To gain behavioral evidence, a dolphin was trained to perform an echolocation task while wearing suction cups over its eyes and either of two neoprene robber hoods over its lower jaw. One hood allowed returning acoustic signals to pass. The other substantially attenuated such signals. The dolphin's performance was significantly hindered while wearing the attenuating hood ( P <. 001, ψ²) as would be expected if the lower jaw was critically important in the reception of high frequency signals.     

Discrimination of jittered sonar echoes by the echolocating bat,Eptesicus fuscus: The shape of target images in echolocation

1. Behavioral experiments with jittering echoes examined acoustic images of sonar targets in the echolocating bat, Eptesicus fuscus, along the echo delay or target range axis. Echo phase, amplitude, bandwidth, and signal-to-noise ratio were manipulated to assess the underlying auditory processes for image formation. 2. Fine delay acuity is about 10 ns. Calibration and control procedures indicate that this represents temporal acuity rather than spectral discrimination. Jitter discrimination curves change in phase when the phase of one jittering echo is shifted by 180 degrees relative to the other, showing that echo phase is involved in delay estimation. At an echo detectability index of about 36 dB, fine acuity is 40 ns, which is approximately as predicted for the delay accuracy of an ideal receiver. 3. Compound performance curves for 0 degrees and 180 degrees phase conditions match the crosscorrelation function of the echoes. The locations of both 0 degrees and 180 degrees phase peaks in the performance curves shift along the time axis by an amount that matches neural amplitude-latency trading in Eptesicus, confirming a temporal basis for jitter discrimination.     

Novel Acoustic Technology for Studying Free-Ranging Shark Social Behaviour by Recording Individuals' Interactions

Group behaviours are widespread among fish but comparatively little is known about the interactions between free-ranging individuals and how these might change across different spatio-temporal scales. This is largely due to the difficulty of observing wild fish groups directly underwater over long enough time periods to quantify group structure and individual associations. Here we describe the use of a novel technology, an animal-borne acoustic proximity receiver that records close-spatial associations between free-ranging fish by detection of acoustic signals emitted from transmitters on other individuals. Validation trials, held within enclosures in the natural environment, on juvenile lemon sharks Negaprion brevirostris fitted with external receivers and transmitters, showed receivers logged interactions between individuals regularly when sharks were within 4 m (∼4 body lengths) of each other, but rarely when at 10 m distance. A field trial lasting 17 days with 5 juvenile lemon sharks implanted with proximity receivers showed one receiver successfully recorded association data, demonstrating this shark associated with 9 other juvenile lemon sharks on 128 occasions. This study describes the use of acoustic underwater proximity receivers to quantify interactions among wild sharks, setting the scene for new advances in understanding the social behaviours of marine animals.     

Can you hear me now? Range‐testing a submerged passive acoustic receiver array in a Caribbean coral reef habitat

Submerged passive acoustic technology allows researchers to investigate spatial and temporal movement patterns of many marine and freshwater species. The technology uses receivers to detect and record acoustic transmissions emitted from tags attached to an individual. Acoustic signal strength naturally attenuates over distance, but numerous environmental variables also affect the probability a tag is detected. Knowledge of receiver range is crucial for designing acoustic arrays and analyzing telemetry data. Here, we present a method for testing a relatively large‐scale receiver array in a dynamic Caribbean coastal environment intended for long‐term monitoring of multiple species. The U.S. Geological Survey and several academic institutions in collaboration with resource management at Buck Island Reef National Monument (BIRNM), off the coast of St. Croix, recently deployed a 52 passive acoustic receiver array. We targeted 19 array‐representative receivers for range‐testing by submersing fixed delay interval range‐testing tags at various distance intervals in each cardinal direction from a receiver for a minimum of an hour. Using a generalized linear mixed model (GLMM), we estimated the probability of detection across the array and assessed the effect of water depth, habitat, wind, temperature, and time of day on the probability of detection. The predicted probability of detection across the entire array at 100 m distance from a receiver was 58.2% (95% CI: 44.0–73.0%) and dropped to 26.0% (95% CI: 11.4–39.3%) 200 m from a receiver indicating a somewhat constrained effective detection range. Detection probability varied across habitat classes with the greatest effective detection range occurring in homogenous sand substrate and the smallest in high rugosity reef. Predicted probability of detection across BIRNM highlights potential gaps in coverage using the current array as well as limitations of passive acoustic technology within a complex coral reef environment.     

Interpulse interval modulation by echolocating big brown bats (<Emphasis Type="Italic">Eptesicus fuscus</Emphasis>) in different densities of obstacle clutter

Big brown bats (Eptesicus fuscus) use biosonar to find insect prey in open areas, but they also find prey near vegetation and even fly through vegetation when in transit from roosts to feeding sites. To evaluate their reactions to dense, distributed clutter, bats were tested in an obstacle array consisting of rows of vertically hanging chains. Chains were removed from the array to create a curved corridor of three clutter densities (high, medium, low). Bats flew along this path to receive a food reward after landing on the far wall. Interpulse intervals (IPIs) varied across clutter densities to reflect different compromises between using short IPIs for gathering echoes rapidly enough to maneuver past the nearest chains and using longer IPIs so that all echoes from one sound can be received before the next sound is emitted. In high-clutter density, IPIs were uniformly shorter (20–65 ms) than in medium and low densities (40–100 ms) and arranged in “strobe groups,” with some overlap of echo streams from different broadcasts, causing pulse-echo ambiguity. As previously proposed, alternating short and long IPIs in strobe groups may allow bats to focus on large-scale pathfinding tasks as well as close-in obstacle avoidance.     

The use of pulse‐echo acoustic microscopy to non‐invasively determine the sex of living larval sea lampreys

A rapid, accurate and non‐invasive method to determine the sex of larval sea lampreys Petromyzon marinus , using wide‐field pulse‐scanning acoustic microscopy, is described. Cross‐sectional pulse‐echo scans were made with a high‐resolution acoustic microscope in 48 larvae (110–130 mm total length, L _T), and the acoustic images generated showed such internal body structures as the gonad, intestine, kidneys, cardinal veins, notochord and musculature. Females were identified by the presence of a relatively large (1–1·5 mm diameter) ovary, which was considerably less reflective to the 15 to 25 MHz acoustic signals than the surrounding kidney tissue. Males were recognized by the lack of the large non‐reflective ovary and, in some cases, the appearance of a small (0·2–0·3 mm) testis with slightly stronger reflective properties than the kidney. Identification of sex was confirmed by optical microscopy following dissection, and in a blind test on an additional 10 specimens (121–168 mm L _T), the acoustic method reliably identified sex in 100% of the larvae. These results indicate that acoustic microscopy can determine the sex of live sea lamprey larvae in c . 30 s per animal, a process which until now required dissection or invasive surgery.     

Molecular diet analysis finds an insectivorous desert bat community dominated by resource sharing despite diverse echolocation and foraging strategies

Interspecific differences in traits can alter the relative niche use of species within the same environment. Bats provide an excellent model to study niche use because they use a wide variety of behavioral, acoustic, and morphological traits that may lead to multi‐species, functional groups. Predatory bats have been classified by their foraging location (edge, clutter, open space), ability to use aerial hawking or substrate gleaning and echolocation call design and flexibility, all of which may dictate their prey use. For example, high frequency, broadband calls do not travel far but offer high object resolution while high intensity, low frequency calls travel further but provide lower resolution. Because these behaviors can be flexible, four behavioral categories have been proposed: (a) gleaning, (b) behaviorally flexible (gleaning and hawking), (c) clutter‐tolerant hawking, and (d) open space hawking. Many recent studies of diet in bats use molecular tools to identify prey but mainly focus on one or two species in isolation; few studies provide evidence for substantial differences in prey use despite the many behavioral, acoustic, and morphological differences. Here, we analyze the diet of 17 sympatric species in the Chihuahuan desert and test the hypothesis that peak echolocation frequency and behavioral categories are linked to differences in diet. We find no significant correlation between dietary richness and echolocation peak frequency though it spanned close to 100 kHz across species. Our data, however, suggest that bats which use both gleaning and hawking strategies have the broadest diets and are most differentiated from clutter‐tolerant aerial hawking species.     

Spatial processing within the mustache bat echolocation system: possible mechanisms for optimization

1. The directionality of an echolocation system is determined by the acoustic properties of both the emitter and receiver, i.e., by the radiation pattern of the emitted pulse and the directionally of the external ears. We measured the directionality of the echolocation system of the greater mustache bat (Pteronotus parnellii) at the 30 kHz, 60 kHz and 90 kHz harmonics of its echolocation pulse by summing, at points throughout the frontal sound field, the echo attenuation due to the spread of pulse energy and the attenuation due to the spread of pulse energy and the attenuation due to the directionality of its external ears. The pulse radiation pattern at the 3 harmonics was measured by comparing the output of a microphone moved throughout the frontal sound field against a second reference microphone at the center of the field. External ear directionality at the 3. harmonics was measured by presenting free-field sounds throughout the frontal sound field, and recording the intensity thresholds of cochlear microphonic potentials, and the intensity thresholds of monaural neurons in the inferior colliculus tuned to one of the 3 harmonics. 2. When compared with ear directionality, the echolocation system was found to be more directional for the center of the sound field in several respects. At all harmonics, attenuation of sounds originating in the peripheral part of the field was increased by 10 to 13 dB. Areas of maximum sound intensity contracted toward the center of the field. Also, the isointensity contours of the echolocation system were more radially symmetrical about the center of the field. 3. At 60 kHz, sound intensity along the azimuth within the echolocation system was nearly constant 26 degrees to either side of the center of the field. This suggests that the radiation pattern of the echolocation pulse and the directionality of the external ears complement one another to produce an acoustic environment at the center of the sound field in which stimulus intensity is stabilized to allow more effective analysis of various aspects of the echolocation target. In particular, we suggest that this intensity stabilization may allow the bat to more effectively resolve the interaural intensity differences it uses to localize prey. 4. Predictions of the azimuthal spatial tuning of binaurally sensitive neurons in the inferior colliculus within the echolocation system were compared with their spatial tuning when only ear directionality is considered.(ABSTRACT TRUNCATED AT 400 WORDS)     

Measuring fish length and assessing behaviour in a high‐biodiversity reach of the Upper Yangtze River using an acoustic camera and echo sounder

In the past decade improved acoustic hard‐ and software have enabled estimations of abundance and distribution patterns of aquatic organism, including non‐intrusive monitoring of fish migrations and behaviour. In this study, a high frequency acoustic camera (DIDSON‐LR, 1.2 MHz, 0.7 MHz) and a portable split‐beam scientific echo sounder (Simrad EY60, 200 kHz) collected acoustic data on 192 and 157 individuals within 24 hr (19–20 April 2011) in the Mituo reach of the Yangtze River, China. Mean fish length estimated from the acoustic camera data was 18.7 ± 5.6 cm, with an average swimming speed of 0.19 ± 0.13 m s^?1. The mean fish target strength (TS) produced by the echo sounder was ?43.8 ± 4.4 dB, which corresponded to 5.7–119.9 cm fish length when converted by three different TS‐length equations. Average swimming speed was 0.11 ± 0.06 m s^?1 from the echo sounder. Compared with the actual fish catch by the three layers of drift gill net in the survey area, the target length indicated by DIDSON was more accurate than the EY60 results, which were highly affected by the choice of TS length equations. It was determined that the two devices used synchronously could estimate fish length effectively to investigate their behaviour and distribution.     

A review of detection range testing in aquatic passive acoustic telemetry studies

Passive acoustic telemetry provides an important tool to study the spatial ecology and behaviour of organisms in marine and freshwater systems, but understanding the detection range of acoustic receivers is critical for interpreting acoustic data and establishing receiver spacing to maximize study efficiency. This study presents a comprehensive review of how acoustic detection range has been considered and assessed to date, summarizes important variables to monitor when determining the detection range of a receiver array, and provides recommendations to account for detection range during experimental design, analysis and data interpretation. A total of 378 passive acoustic telemetry studies (1986–2012) were scored against a set of pre-defined criteria to provide a standardized assessment of how well detection range was accounted for, from a maximum possible score of 45. Scores ranged from 0 to 39 (11.1 ± 0.4; mean ± 1 SE). Over the past decade mean scores have been consistently between 6.7 and 12.9 which indicates that detection range has not been adequately considered in most contemporary acoustic telemetry studies. Given the highly variable nature of detection range over space and time, it is necessary to create a culture of detection range testing among the scientific community. For robust telemetry studies it is recommended that consideration of detection range should be given a greater focus within study design, execution and data analysis. To aid array design in new systems, short-term detection range tests should be conducted in the most representative area of the study system prior to deployment. As well, fixed distance sentinel tags should ideally be deployed at a representative receiver site within the array to provide a continuous assessment of detection range and influential environmental parameters should be monitored to facilitate modeling of detection range variability over time. When warranted, data analysis should incorporate modeled variation in detection ranges.     

Performance of remote acoustic receivers within a coral reef habitat: implications for array design

Remote monitoring technologies are increasingly being implemented in the marine environment to better understand the movement patterns of taxa. Coral reefs are no exception. However, there is a paucity of information relating to the performance of acoustic receivers on coral reefs. Our results suggest that the detection performance of acoustic receivers may be significantly impacted by the unique nature of the reef environment. This study assessed the performance of passive acoustic receivers on a typical inner-shelf fringing reef, Orpheus Island, on the Great Barrier Reef, Australia. The detection range and diel performance variability of acoustic receivers was assessed using two parallel lines of 5 VR2W receivers spanning 125?m, deployed on the reef base and reef crest. Two 9-mm acoustic transmitters were moored at opposite ends of each receiver line. The working detection range for receivers was found to be approximately 90?m for the transmitter moored on the reef base and just 60?m for the transmitter moored on the reef crest. However, the detection range on the reef crest increased to 90?m when just the reef crest receivers were considered, highlighting importance of optimal receiver deployment. No diel patterns in receiver performance or detection capacities were detected, suggesting that no corrections are required when interpreting nocturnal versus diurnal activity patterns. We suggest that studies aiming for complete coverage of a site within a reef environment will require receivers in close (<100?m) proximity, and that the placement depth of receivers must be a major consideration, with shallow receivers exhibiting a greater detection range than those on the reef slope. Our results highlight the challenges imposed by coral reefs for acoustic telemetry and the importance of receiver placement for studies conducted within these habitats.     

A combined radio and acoustic transmitter for fixing direction and range of freshwater fish (RAFIX)

A mobile system incorporating a novel transmitter for tracking the position of freshwater fish is described. The transmitter emits both acoustic and radio pulses simultaneously to allow a range estimate from the delay between signals arriving at a receiver station. Results of field trials are presented and the usefulness of the system is discussed in relation to other tracking methods.     

Bats' avoidance of real and virtual objects: implications for the sonar coding of object size

Fast movement in complex environments requires the controlled evasion of obstacles. Sonar-based obstacle evasion involves analysing the acoustic features of object-echoes (e.g., echo amplitude) that correlate with this object's physical features (e.g., object size). Here, we investigated sonar-based obstacle evasion in bats emerging in groups from their day roost. Using video-recordings, we first show that the bats evaded a small real object (ultrasonic loudspeaker) despite the familiar flight situation. Secondly, we studied the sonar coding of object size by adding a larger virtual object. The virtual object echo was generated by real-time convolution of the bats’ calls with the acoustic impulse response of a large spherical disc and played from the loudspeaker. Contrary to the real object, the virtual object did not elicit evasive flight, despite the spectro-temporal similarity of real and virtual object echoes. Yet, their spatial echo features differ: virtual object echoes lack the spread of angles of incidence from which the echoes of large objects arrive at a bat's ears (sonar aperture). We hypothesise that this mismatch of spectro-temporal and spatial echo features caused the lack of virtual object evasion and suggest that the sonar aperture of object echoscapes contributes to the sonar coding of object size.