Captive dolphins,Tursiops truncatus,develop signature whistles that match acoustic features of human-made model sounds

This paper presents a cross-sectional study testing whether dolphins that are born in aquarium pools where they hear trainers' whistles develop whistles that are less frequency modulated than those of wild dolphins. Ten pairs of captive and wild dolphins were matched for age and sex. Twenty whistles were sampled from each dolphin. Several traditional acoustic features (total duration, duration minus any silent periods, etc.) were measured for each whistle, in addition to newly defined flatness parameters: total flatness ratio (percentage of whistle scored as unmodulated), and contiguous flatness ratio (duration of longest flat segment divided by total duration). The durations of wild dolphin whistles were found to be significantly longer, and the captive dolphins had whistles that were less frequency modulated and more like the trainers' whistles. Using a standard t-test, the captive dolphin had a significantly higher total flatness ratio in 9/10 matched pairs, and in 8/10 pairs the captive dolphin had significantly higher contiguous flatness ratios. These results suggest that captive-born dolphins can incorporate features of artificial acoustic models made by humans into their signature whistles.     

Who is whistling? Localizing and identifying phonating dolphins in captivity

Acoustic communication through whistles is well developed in dolphins. However, little is known on how dolphins are using whistles because localizing the sound source is not an easy task. In the present study, the hyperbola method was used to localize the sound source using a two-hydrophone array. A combined visual and acoustic method was used to determine the identity of the whistling dolphin. In an aquarium in Mexico City where two adult bottlenose dolphins were housed we recorded 946 whistles during 22 days. We found that a dolphin was located along the calculated hyperbola for 72.9% of the whistles, but only for 60.3% of the whistles could we determine the identity of the whistling dolphin. However, sometimes it was possible to use other cues to identify the whistling dolphin. It could be the animal that performed a behavior named “observation” at the time whistling occurred or, when a whistle was only recorded on one channel, the whistling dolphin could be the animal located closest to the hydrophone that captured the whistle. Using these cues, 15.4% of the whistles were further ascribed to either dolphin to obtain an overall identification efficiency of 75.7%. Our results show that a very simple and inexpensive acoustic setup can lead to a reasonable number of identifications of the captive whistling dolphin: this is the first study to report such a high rate of whistles identified to the free swimming, captive dolphin that produced them. Therefore, we have a data set with which we can investigate how dolphins are using whistles. This method can be applied in other aquaria where a small number of dolphins is housed; though, the actual efficiency of this method will depend on how often dolphins spend time next to each other and on the reverberation conditions of the pool.     

Echolocation signals of wild dolphins

Most of our understanding of dolphin echolocation has come from studies of captive dolphins performing various echolocation tasks. Recently, measurements of echolocation signals in the wild have expanded our understanding of the characteristics of these signals in a natural setting. Measuring undistorted dolphin echolocation signals with free swimming dolphins in the field can be a challenging task. A four hydrophone array arranged in a symmetrical star pattern was used to measure the echolocation signals of four species of dolphins in the wild. Echolocation signals of the following dolphins have been measured with the symmetrical star array: white-beaked dolphins in Iceland, Atlantic spotted dolphins in the Bahamas, killer whales in British Columbia, and dusky dolphins in New Zealand. There are many common features in the echolocation signals of the different species. Most of the signals had spectra that were bimodal: two peaks, one at low frequencies and another about an octave higher in frequency. The source level of the sonar transmission varies as a function of 20logR, suggesting a form of time-varying gain but on the transmitting end of the sonar process rather than the receiving end. The results of the field work call into question the issue of whether the signals used by captive dolphins may be shaped by the task they are required to perform rather than what they would do more naturally.     

圈养印太瓶鼻海豚(Tursiops aduncus)正弦型哨叫声与其行为及水环境关系分析

为分析圈养印太瓶鼻海豚与所处水环境间的关系,观测和记录了两只圈养印太瓶鼻海豚一年内的发声、行为及水环境温度、盐度及酸碱度。通过时频滤波和分析,筛选出一年中17:00到08:00内海豚发出的正弦型哨叫声。经统计、比对和相关性分析,得到:正弦型哨叫声发生量与海豚不良情绪行为发生量线性正相关(拟合优度R2=93.89%),8月—10月发生频次最多,此时海豚不良情绪行为也最多;该类型信号发生量占比与所处水环境月平均温度和盐度有关,并拟合出它们的关系式(拟合优度R2=68.61%),但受月平均酸碱度影响不大;水环境月平均温度和盐度在一定范围内时,海豚不良情绪行为发生量占比可以控制在12%以下。研究结果为今后利用海豚哨叫声判断海豚生物行为、健康状况、圈养环境舒适度等提供一定的科学参考。      

圈养瓶鼻海豚对20 kHz连续声信号的行为响应研究

将20 kHz连续声信号作为刺激信号,测试了厦门某海湾圈养的两只瓶鼻海豚对该信号的行为变化。通过对比信号发射期与间歇期海豚相对声源的水面距离、露出水面的次数以及水下发出的click定位声信号的数目等变化,判断发射信号对海豚行为的影响。给出了瓶鼻海豚对测试信号产生躲避行为的声压级门限(154±2 dB re 1μPa,rms),并与鼠海豚的躲避声压门限级进行了对比。结果表明:信号发射期,瓶鼻海豚移离了声源位置,增加了露出水面的次数,水下发出click声信号的次数明显减少。因此,瓶鼻海豚对20kHz连续信号产生了行为改变。      

Behavioral responses of two captive bottlenose dolphins (Tursiops truncatus) to a continuous 50 kHz tone

At present, the fundamental frequencies of signals of most commercially available acoustic alarms to deter small cetaceans are below 20 kHz, but it is not well ascertained whether higher frequencies have a deterrent effect on bottlenose dolphins (Tursiops truncatus). Two captive bottlenose dolphins housed in a floating pen were subjected to a continuous pure tone at 50 kHz with a source level of 160 ± 2 dB (re 1 μPa, rms). The behavioral responses of dolphins were judged by comparing surfacing distance relative to the sound source, number of surfacings, and number of echolocation clicks produced, during forty 15 min baseline periods with forty 15 min test periods (four sessions per day, 40 sessions in total). On all 10 study days, surfacing distance and the number of surfacings increased while click production decreased during broadcasts of test sound. The avoidance threshold sound pressure level for a continuous 50 kHz tone for the bottlenose dolphins, in the context of this study, was estimated to be 144 ± 2 dB (re 1 μPa, rms). The results indicated that a continuous 50 kHz tonal signal can deter bottlenose dolphins from an area.     

An optical telemetry device to identify which dolphin produces a sound

A small telemetry device, called a "vocalight," was designed for attachment to a dolphin's head using a suction cup. The vocalight lights up a variable number of light-emitting diodes depending upon the loudness of sounds received at a hydrophone within the suction cup. If vocalights matched for sensitivity are put on each dolphin within a captive group, observers can identify which dolphin produces a vocalization. Use of vocalights indicates that source levels of whistles from captive bottlenosed dolphins, Tursiops truncatus, range from approximately 125 to over 140 dB re: 1 microPa at 1 m.     

Variation in the hearing sensitivity of a dolphin population determined through the use of evoked potential audiometry

A portable electrophysiological data collection system was used to assess hearing in a captive population of bottlenose dolphins by recording auditory evoked potentials (AEPs). The AEP system used a transducer embedded in a suction cup to deliver amplitude modulated tones to the dolphin through the lower jaw. Evoked potentials were recorded noninvasively using surface electrodes. Adaptive procedures allowed hearing thresholds to be estimated from 10 to 150 kHz in a single ear in about 45 min. Hearing thresholds were measured in 42 bottlenose dolphins (28 male, 14 female), ranging in age from 4 to 47 years. Variations in hearing sensitivity with age and sex followed patterns seen in humans and terrestrial mammals: generally, within the population there was a progressive loss of high frequency hearing with age and an earlier onset of hearing loss in males than in females. Hearing loss generally occurred between the ages of 20 and 30, and all animals over the age of 27 had some degree of hearing loss. Two dolphins with profound hearing loss were found within the population. Aberrant hearing patterns were observed in related dolphins suggesting genetic links to hearing ability may exist.     

Echolocation range of captive and free-ranging baiji (Lipotes vexillifer), finless porpoise (Neophocaena phocaenoides), and bottlenose dolphin (Tursiops truncatus).

The interclick intervals of captive dolphins are known to be longer than the two-way transit time between the dolphin and a target. In the present study, the interclick intervals of free-ranging baiji, finless porpoises, and bottlenose dolphins in the wild and in captivity were compared. The click intervals in open waters ranged up to 100-200 ms, whereas the click intervals in captivity were in the order of 4-28 ms. Echolocation of free-ranging dolphins appears to adapt to various distance in navigation or ranging, sometimes up to 140 m. Additionally, the difference of waveform characteristics of clicks between species was recognized in the frequency of maximum energy and the click duration.     

印太瓶鼻海豚(Tursiops aduncus)通讯声信号分类及特征参数的环境差异性分析

通过长期记录室内水池环境下两只印太瓶鼻海豚通讯信号,并与海湾自然环境下同样的两只海豚所发出的通讯信号进行比较分析,从信号类型、声谱特征等方面研究生活环境变化下瓶鼻海豚通讯信号的差异性。结果表明,生活环境的差异,会改变瓶鼻海豚通讯信号。海湾自然环境下,瓶鼻海豚通讯信号以正弦型信号为主;而室内水池环境下,上扫型信号比例明显增多,而正弦型信号减少。两种环境下,瓶鼻海豚通讯信号在持续时间、拐点数、起始频率、结束频率、最小频率、最大频率等存在显著性差异(p<0.05),但信号的频率变化量相近(p=0.29)。结果为提高海豚通讯信号认知和增强海豚生物行为研究提供一定的科学参考,同时也为仿生隐蔽通信提供技术支撑。      

圈养宽吻海豚自由游动状态和训练期间通讯信号比较研究

根据频率特性对圈养宽吻海豚(Tursiops truncatus)在自由游动和训练两种实验条件下的声通讯信号进行分类,并利用双尾t检验统计分析方法对两种条件下的信号声谱参数进行统计比较。结果显示,宽吻海豚在自由状态下通讯信号的种类多并以正弦型为主,而训练期间的通讯信号则大多数为上扫频类。此外,统计分析表明起始频率不能反映这两种状态的不同(p=0.22)。结束频率、最小频率、最大频率、频率变化量、拐点数、环形数、阶数、波形数和周期等则显示了两种状态显著的差异性(p＜0.05)。结果为今后海豚声行为研究提供一定的科学参考和基础。      

Acoustic features of objects matched by an echolocating bottlenose dolphin

The focus of this study was to investigate how dolphins use acoustic features in returning echolocation signals to discriminate among objects. An echolocating dolphin performed a match-to-sample task with objects that varied in size, shape, material, and texture. After the task was completed, the features of the object echoes were measured (e.g., target strength, peak frequency). The dolphin's error patterns were examined in conjunction with the between-object variation in acoustic features to identify the acoustic features that the dolphin used to discriminate among the objects. The present study explored two hypotheses regarding the way dolphins use acoustic information in echoes: (1) use of a single feature, or (2) use of a linear combination of multiple features. The results suggested that dolphins do not use a single feature across all object sets or a linear combination of six echo features. Five features appeared to be important to the dolphin on four or more sets: the echo spectrum shape, the pattern of changes in target strength and number of highlights as a function of object orientation, and peak and center frequency. These data suggest that dolphins use multiple features and integrate information across echoes from a range of object orientations.     

Source parameters of echolocation clicks from wild bottlenose dolphins (Tursiops aduncus and Tursiops truncatus)

The Indian Ocean and Atlantic bottlenose dolphins (Tursiops aduncus and Tursiops truncatus) are among the best studied echolocating toothed whales. However, almost all echolocation studies on bottlenose dolphins have been made with captive animals, and the echolocation signals of free-ranging animals have not been quantified. Here, biosonar source parameters from wild T. aduncus and T. truncatus were measured with linear three- and four-hydrophone arrays in four geographic locations. The two species had similar source parameters, with source levels of 177-228 dB re 1 μPa peak to peak, click durations of 8-72 μs, centroid frequencies of 33-109 kHz and rms bandwidths between 23 and 54 kHz. T. aduncus clicks had a higher frequency emphasis than T. truncatus. The transmission directionality index was up to 3 dB higher for T. aduncus (29 dB) as compared to T. truncatus (26 dB). The high directionality of T. aduncus does not appear to be only a physical consequence of a higher frequency emphasis in clicks, but may also be caused by differences in the internal properties of the sound production system.     

Classification of echolocation clicks from odontocetes in the Southern California Bight

This study presents a system for classifying echolocation clicks of six species of odontocetes in the Southern California Bight: Visually confirmed bottlenose dolphins, short- and long-beaked common dolphins, Pacific white-sided dolphins, Risso's dolphins, and presumed Cuvier's beaked whales. Echolocation clicks are represented by cepstral feature vectors that are classified by Gaussian mixture models. A randomized cross-validation experiment is designed to provide conditions similar to those found in a field-deployed system. To prevent matched conditions from inappropriately lowering the error rate, echolocation clicks associated with a single sighting are never split across the training and test data. Sightings are randomly permuted before assignment to folds in the experiment. This allows different combinations of the training and test data to be used while keeping data from each sighting entirely in the training or test set. The system achieves a mean error rate of 22% across 100 randomized three-fold cross-validation experiments. Four of the six species had mean error rates lower than the overall mean, with the presumed Cuvier's beaked whale clicks showing the best performance (<2% error rate). Long-beaked common and bottlenose dolphins proved the most difficult to classify, with mean error rates of 53% and 68%, respectively.     

圈养瓶鼻海豚通讯信号分析及融合分类方法

针对圈养条件下瓶鼻海豚通讯信号（whistle）分类时混叠大量回声定位信号（click）导致分类正确率降低的问题,提出了一种基于机器学习的融合分类方法。分别提取whistle信号的时频分布特征训练随机森林分类器,梅尔时频图特征训练卷积神经网络分类器,在此基础上设计融合判决器对混叠whistle信号进行分类识别。对圈养海豚声信号采集实验数据的分类识别结果表明,融合分类方法具有更好的分类性能,对混叠whistle信号分类正确率大于94%,优于时频分布特征分类器和梅尔时频图特征分类器,能够提高混叠信号的分类能力。      

Linking the sounds of dolphins to their locations and behavior using video and multichannel acoustic recordings

It is difficult to attribute underwater animal sounds to the individuals producing them. This paper presents a system developed to solve this problem for dolphins by linking acoustic locations of the sounds of captive bottlenose dolphins with an overhead video image. A time-delay beamforming algorithm localized dolphin sounds obtained from an array of hydrophones dispersed around a lagoon. The localized positions of vocalizing dolphins were projected onto video images. The performance of the system was measured for artificial calibration signals as well as for dolphin sounds. The performance of the system for calibration signals was analyzed in terms of acoustic localization error, video projection error, and combined acoustic localization and video error. The 95% confidence bounds for these were 1.5, 2.1, and 2.1 m, respectively. Performance of the system was analyzed for three types of dolphin sounds: echolocation clicks, whistles, and burst-pulsed sounds. The mean errors for these were 0.8, 1.3, and 1.3 m, respectively. The 95% confidence bound for all vocalizations was 2.8 m, roughly the length of an adult bottlenose dolphin. This system represents a significant advance for studying the function of vocalizations of marine animals in relation to their context, as the sounds can be identified to the vocalizing dolphin and linked to its concurrent behavior.     

Echolocation behavior of franciscana dolphins (Pontoporia blainvillei) in the wild

Franciscana dolphins are small odontocetes hard to study in the field. In particular, little is known on their echolocation behavior in the wild. In this study we recorded 357 min and analyzed 1019 echolocation signals in the Rio Negro Estuary, Argentina. The clicks had a peak frequency at 139 kHz, and a bandwidth of 19 kHz, ranging from 130 to 149 kHz. This is the first study describing echolocation signals of franciscana dolphins in the wild, showing the presence of narrow-band high frequency signals in these dolphins. Whether they use other vocalizations to communicate or not remains uncertain.     

Echo features used by human listeners to discriminate among objects that vary in material or wall thickness: implications for echolocating dolphins

Echolocating dolphins extract object feature information from the acoustic parameters of echoes. To gain insight into which acoustic parameters are important for object discrimination, human listeners were presented with echoes from objects used in two discrimination tasks performed by dolphins: Hollow cylinders with varying wall thicknesses (+/-0.2, 0.3, 0.4, and 0.8 mm), and spheres made of different materials (steel, aluminum, brass, nylon, and glass). The human listeners performed as well or better than the dolphins at the task of discriminating between the standard object and the comparison objects on both the cylinders (humans=97.1%; dolphin=82.3%) and the spheres (humans= 86.6%; dolphin= 88.7%). The human listeners reported using primarily pitch and duration to discriminate among the cylinders, and pitch and timbre to discriminate among the spheres. Dolphins may use some of the same echo features as the humans to discriminate among objects varying in material or structure. Human listening studies can be used to quickly identify salient combinations of echo features that permit object discrimination, which can then be used to generate hypotheses that can be tested using dolphins as subjects.     

Classification of Risso's and Pacific white-sided dolphins using spectral properties of echolocation clicks

The spectral and temporal properties of echolocation clicks and the use of clicks for species classification are investigated for five species of free-ranging dolphins found offshore of southern California: short-beaked common (Delphinus delphis), long-beaked common (D. capensis), Risso's (Grampus griseus), Pacific white-sided (Lagenorhynchus obliquidens), and bottlenose (Tursiops truncatus) dolphins. Spectral properties are compared among the five species and unique spectral peak and notch patterns are described for two species. The spectral peak mean values from Pacific white-sided dolphin clicks are 22.2, 26.6, 33.7, and 37.3 kHz and from Risso's dolphins are 22.4, 25.5, 30.5, and 38.8 kHz. The spectral notch mean values from Pacific white-sided dolphin clicks are 19.0, 24.5, and 29.7 kHz and from Risso's dolphins are 19.6, 27.7, and 35.9 kHz. Analysis of variance analyses indicate that spectral peaks and notches within the frequency band 24-35 kHz are distinct between the two species and exhibit low variation within each species. Post hoc tests divide Pacific white-sided dolphin recordings into two distinct subsets containing different click types, which are hypothesized to represent the different populations that occur within the region. Bottlenose and common dolphin clicks do not show consistent patterns of spectral peaks or notches within the frequency band examined (1-100 kHz).