DIEL VARIATION IN ECHOLOCATION BEHAVIOR OF WILD HARBOR PORPOISES

The echolocation rate and behavior of wild harbor porpoises were studied using a harbor porpoise click detector (POD) deployed close to the bottom at 40 m depth in Scottish waters, UK, April—June 2001. Echolocation variables were compared among four diel phases; morning, day, evening, and night. The echolocation encounter rate, the minimum interclick interval per train, and the proportion of echolocation click trains with a minimum interclick interval below 10 msec were all significantly higher at night than during the day. The variation in echolocation rate implies that porpoises increased their echolocation rate and/or visited the depth of the POD more often at night than during the day. Further, the changes in minimum interclick interval per train suggest that they used their echolocation for foraging or investigating objects at a close range to a higher extent, and acoustically explored the environment at greater distances at night than during the day.     

Noise affects porpoise click detections – the magnitude of the effect depends on logger type and detection filter settings

Automatic click detectors and full-bandwidth sound recorders are widely used in passive acoustic monitoring of small cetaceans. Detection of these signals depends on a variety of factors, including signal to noise ratio. Passive acoustic monitoring is often used to study impact of underwater noise on small cetaceans, but as detection probability is affected by changes in signal to noise ratio, variable noise levels may affect conclusions drawn from these experiments. Therefore, we examine how different detectors and filters perform in varying ocean noise conditions. C-PODs and full-bandwidth recorders (Wildlife Acoustics, SM2M+) were deployed at two stations in an environment with fluctuating ambient noise for 42 days. Noise level and harbour porpoise (Phocoena phocoena) click trains simultaneously recorded on both loggers were compared. Overall, we found that porpoise click detections by the algorithm used to analyse full-band recorder data (Pamguard) paralleled detections by the C-POD. However, Pamguard detected significantly more clicks than the C-POD. A decrease in detections was seen for both loggers with increasing noise in the band 20 –160 kHz, in particular for levels above 100 dB re 1μPa rms. We also found that the Pamguard detection function changed the least over varying noise conditions when compared to the C-POD detectors. This study sheds light on the fact that inference of animal presence/absence or density that are based on echolocation cues (here, Porpoise Positive Minutes) shall account for the acoustic environments where probability of detecting signals may be affected by variability in ambient noise levels.     

The Diel Rhythms of Biosonar Behavior in the Yangtze Finless Porpoise (Neophocaena asiaeorientalis asiaeorientalis) in the Port of the Yangtze River: The Correlation between Prey Availability and Boat Traffic

Information on the habitat use of the critically endangered Yangtze finless porpoise (Neophocaena asiaeorientalis asiaeorientalis) is critical for its conservation. The diel biosonar behavior of the porpoise in the port areas of the Yangtze River was examined along with simultaneous observations of fish density and boat traffic. Biosonar pulses from the porpoises were detected for 1233 min (5.77%) over a 21,380 min duration of effective observations. In total, 190 (5.63%) buzzes (an indication of prey capture attempts) were recorded among the 3372 identified click trains. Of the 168 echolocation encounters (bouts of click trains less than eight min apart), 150 (89.3%) involved single animals, indicating that solitary porpoises were frequently present and feeding in the port areas. Significant diel patterns were evident involving the biosonar behavior of the porpoises (including click trains and buzzes), fish density and boat traffic. The frequencies of the click trains and buzzes were significantly lower during the day than in the evening and at night, which suggests that porpoises in this region are primarily engaged in crepuscular and nocturnal foraging. The lack of a significant diel pattern in the echolocation encounters indicates the importance of the port in porpoise conservation. A forced feeding schedule may be associated with the lack of a significant correlation between porpoise acoustics and boat traffic. Overall, prey availability appears to be the primary factor that attracts porpoises. Additionally, porpoises tend to migrate or remain downstream in the morning and migrate or remain upstream in the evening, most likely to follow their prey. The findings of this study can be used to improve the conservation of the Yangtze finless porpoise.     

THE CLICK-SOUNDS OF NARWHALS (MONODON MONOCEROS) IN INGLEFIELD BAY, NORTHWEST GREENLAND

We studied the sounds of narwhals ( Monodon monoceros ) foraging in the open waters in Northwest Greenland. We used a linear, vertical array of three hydrophones (depth 10 m, 30 m, 100 m) with a fourth hydrophone (depth 30 m) about 20 m from the vertical array. A smaller fifth hydrophone (depth 2 m) allowed for registering frequencies up to 125 kHz (± 2 dB) when signals were recorded at 762 mm/set on an instrumentation tape recorder. Clicks were the prevalent signals, but we heard whistles occasionally. We separated the clicks into two classes: click trains that had rates of 3-10 clicks/sec and click bursts having rates of 110-150 clicks/sec. The spectra of train clicks had maximum amplitudes at 48 ± 10 kHz and a duration of 29 ± 6 psec. The spectra of burst clicks had maximum amplitudes at 19 ± 1 kHz and a duration of 40 ± 3 psec. By analogy with other dolphin species, narwhals presumably use the clicks for echolocation during orientation and for locating prey. The narwhal click patterns resemble those of insectivorous bats. Click trains might correspond to bat searching signals and click bursts to the bat's terminal "buzz", emitted just before prey capture.     

Determinants of echolocation click frequency characteristics in small toothed whales: recent advances from anatomical information

1. The pulse-like clicking sounds made by odontocetes for echolocation (biosonar) can be roughly classified by their frequency characteristics into narrow-band high-frequency (NBHF) clicks with a sharp peak at around 130 kHz and wide-band (WB) clicks with a moderate peak at 30–100 kHz. Structural differences in the sound-producing organs between NBHF species and WB species have not been comprehensively discussed, nor has the formation of NBHF and WB clicks.
2. A review of the sound-producing organs, including the latest findings, could lead to a new hypothesis about the sound production mechanisms. In the current review, data on echolocation click characteristics and on the anatomical structure of the sound-producing organs were compared in 33 species (14 NBHF species and 19 WB species).
3. We review interspecific information on the characteristics of click frequencies and data from computed tomography scans and morphology of the sound-producing organs, accumulated in conventional studies. The morphology of several characteristic structures, such as the melon, the dense connective tissue over the melon (the ‘porpoise capsule’), and the vestibular sacs, was compared interspecifically.
4. Interspecific comparisons suggest that the presence or absence of the porpoise capsule is unlikely to affect echolocation frequency. Folded structures in the vestibular sacs, features that have been overlooked until now, are present in most species with NBHF sound production and not in WB species; the vestibular sacs are therefore likely to be important in determining echolocation click frequency characteristics. The acoustical properties of the shape of the melon and vestibular sacs are important topics for future investigations about the relationship between anatomical structure and sound-producing mechanisms for echolocation clicks.

     

CHANGES IN CLICK SOURCE LEVELS WITH DISTANCE TO TARGETS: STUDIES OF FREE-RANGING WHITE-BEAKED DOLPHINS LAGENORHYNCHUS ALBIROSTRIS AND CAPTIVE HARBOUR PORPOISES PHOCOENA PHOCOENA

ABSTRACT

Probably all odontocetes use echolocation for spatial orientation and detection of prey. We used a four hydrophone “Y” array to record the high frequency clicks from free-ranging White-beaked Dolphins Lagenorhynchus albirostris and captive Harbour Porpoises Phocoena phocoena. From the recordings we calculated distances to the animals and source levels of the clicks. The recordings from White-beaked Dolphins were made in Iceland and those from Harbour Porpoises at Fjord & Baelt, Kerteminde, Denmark during prey capture. We used stringent criteria to determine which clicks could be defined as being on the acoustic axis. Two dolphin and nine porpoise click series could be used to track individual animals, which presumably focused on the array hydrophones or a fish right in front of the array. The apparent source levels of clicks in the individual tracks increased with range. One individual White-beaked Dolphin and three Harbour Porpoises regulate their output signal level to nearly compensate for one-way transmission loss while approaching a target. The other dolphin regulated the output differently. For most of the recordings the sound level at the target remains nearly constant and the echo level at the animal increases as it closes on the target.     

The beluga whale produces two pulses to form its sonar signal

Odontocete cetaceans use biosonar clicks to acoustically probe their aquatic environment with an aptitude unmatched by man-made sonar. A cornerstone of this ability is their use of short, broadband pulses produced in the region of the upper nasal passages. Here we provide empirical evidence that a beluga whale (Delphinapterus leucas) uses two signal generators simultaneously when echolocating. We show that the pulses of the two generators are combined as they are transmitted through the melon to produce a single echolocation click emitted from the front of the animal. Generating two pulses probably offers the beluga the ability to control the energy and frequency distribution of the emitted click and may allow it to acoustically steer its echolocation beam.     

Field assessment of C‐POD performance in detecting echolocation click trains of bottlenose dolphins (Tursiops truncatus)

We evaluated the performance of dolphin echolocation detectors (C‐PODs) in the New River, North Carolina, by ground‐truthing echolocation detections with digital acoustic recordings. We deployed C‐PODs at three sites for a total of 204 monitoring hours. We also performed detection range trials at two sites where water depths ranged from 1.0 to 4.5 m. We used Detection Positive Minutes (DPMs), minutes of C‐POD recordings that contained at least one echolocation click train, to indicate the presence of at least one dolphin. The C‐PODs performed well in detecting dolphin click trains, although all units performed conservatively by failing to detect some echolocation events and therefore underestimated the true occurrence of dolphins. C‐PODs reported only a small number of false detections, as indicated by low false positive rates ranging between 1% and 4% for individual units. Overall, C‐PODs performed with a high accuracy (72%–91%) and detected echolocation at a distance of at least 933 m. We conclude that C‐PODs hold considerable promise in future monitoring studies of this species, but recommend a careful study design especially in complex, coastal environments.     

BOOK REVIEW

Echolocation calls of four species of insectivorous bats of central Chile were recorded and characterized to determine vocal signatures that allow their identification in the field. Pulses of Tadarida brasiliensis were characterized by the highest duration and the lowest values for all frequencies, which do not overlap those of the remaining species. Tadarida emits narrowband, shallow frequency-modulated search calls. All three vespertilionid species studied (Histiotus montanus, Lasiurus varius and Myotis chiloensis) showed similar echolocation design to one another, consisting of a downward frequency modulation at the beginning of the signal followed by a narrowband quasi-constant frequency component; however, their calls differ by their spectral characteristics. Discriminant function analysis of six acoustic parameters (duration, initial frequency, slope frequency modulation, peak frequency, minimal and maximal frequencies) gave an overall classification of 87.4%, suggesting species could be correctly classified based on echolocation calls. Call duration and minimal frequency were the variables most important for species identification.     

Can pipistrelles, Pipistrellus pipistrellus (Schreber, 1774) and Pipistrellus pygmaeus (Leach, 1825), foraging in a group, change parameters of their signals?

Echolocation behaviour and the structure of calls of Pipistrellus pygmaeus and Pipistrellus pipistrellus were studied by using a time expansion bat detector. Echolocation signals were recorded in the field in south-eastern Moravia and northern Bohemia (Czech Republic) and in an ad hoc experimental laboratory. For each of the species, multivariate analysis of variance (MANOVA) indicated significant differences in calls produced inside the experimental room and in the open. Paired t -tests and MANOVA were also used to reveal influences of interindividual contacts in each of the cryptic species on the spectral patterns of call variables. Differences were found in the spectral variables of echolocation calls of an individual flying in the room alone and in a group of conspecifics. The possibility that bats use their flexibility to avoid mutual disturbances of echolocation calls was tested. We found that bats flying in a group modify the parameters of their echolocation signals according to the presence of other individuals of the same species. These differences can indicate jamming avoidance and recognition of own echoes. However, they did not change the parameters if individuals of another species were present. Social calls are more numerous when bats fly in a mixed-species group than in a monospecific group.     

Echolocation calls produced by Kuhl's pipistrelles in different flight situations

Flexibility in the echolocation call structure of bats can improve their performances, because, in some situations, some signal designs are better than others. Hence, at least some bats should adjust their echolocation calls according to the setting in which they are operating but also to the specific task at hand, that is their behavioral intention. We studied variation in the echolocation calls of Pipistrellus kuhlii emitted during four flight situations that were similar in setting but differed in behavioral context: emergence from a roost, commuting to and from foraging sites, foraging and returning to a roost. Echolocation calls produced by P. kuhlii differed significantly according to the flight situation. Call types differed most distinctly between foraging and commuting. We also found a high variance in the emergence calls we recorded, perhaps reflecting pre- and post-takeoff calls. Discriminant function analysis on calls emitted while foraging, commuting or returning to the roost classified the calls to the correct group 73.3% of the time. The differences between bats' echolocation calls in different flight situations might indicate an intrinsic change in the bat's behavior. Recognizing these differences could be crucial when using call variables to identify bat species.     

TARGET DISCRIMINATION BY AN ECHOLOCATING FINLESS PORPOISE, NEOPHOCAENA PHOCAENOIDES

The ability of a finless porpoise (Neophocaena phocaenoides) to discriminate the material and size of a target by echolocation was investigated. The porpoise was required to choose a standard target of a 15-mm diameter solid steel cylinder from two stimuli, a standard and a comparison target, presented simultaneously. The porpoise could distinguish a standard cylinder from acrylic resin and brass targets, but had difficulties distinguishing it from an aluminum cylinder. In size discrimination, the porpoise could distinguish the standard from 12-, 18-, and 20-mm diameter cylinders, but had difficulties distinguishing it from a 14-mm diameter cylinder. Echo measurements suggest that the porpoise is able to detect: (1) time difference between two echo highlights to within approximately 1 μS, (2) frequency shifts of approximately 7 kHz in a broadband echo having a peak frequency of nearly 140 kHz, (3) time-separation pitch of approximately 30 kHz, and (4) target strength differences of approximately 1 dB.     

Things that go bump in the night: evolutionary interactions between cephalopods and cetaceans in the tertiary

Echolocation has evolved independently in several vertebrate groups, and hypotheses about the origin of echolocation in these groups often invoke abiotic mechanisms driving morphological evolution. In bats, for example, the ecological setting associated with the origin of echolocation has been linked to global warming during the Palaeocene–Eocene; similarly, the origin of toothed whales (odontocetes) has been broadly correlated with the establishment of the circum-Antarctic current. These scenarios, and the adaptational hypotheses for the evolution of echolocation with which they are associated, neglect a consideration of possible biotic mechanisms. Here we propose that the origin of echolocation in odontocetes was initially an adaptation for nocturnal epipelagic feeding – primarily on diel migrating cephalopods. We test this hypothesis using data on the temporal, geographical, and water column distributions of odontocetes and cephalopods, and other global events from their respective tertiary histories. From this analysis, we suggest that echolocation in early odontocetes aided nocturnal feeding on cephalopods and other prey items, and that this early system was exapted for deep diving and hunting at depths below the photic zone where abundant cephalopod resources were available 24 h a day. This scenario extends to the evolution of other cephalopod feeding (teuthophagous) marine vertebrates such as pinnipeds and Mesozoic marine reptiles.     

Synthetic communication signals influence wild harbour porpoise (Phocoena phocoena) behaviour

We used our novel and programmable Porpoise Alarm (PAL, patd.) to synthesize life-like, electronic harbour porpoise communication signals based on those described for captive animals. In the Little Belt, Denmark, we employed PAL (source level 158 ± 1 dB p–p re 1 μPa@1 m; centroid frequency 133 ± 8.5 kHz) to synthesize three aggressive click train types termed “A”, “F3” and “M1” to naive, free-living harbour porpoises. Via theodolite tracking (372 h of total visual effort spread over 10 expeditions) we found that, depending on signal type, porpoises either avoid or become attracted to PAL: Signal types “A” and “F3” are slight deterrents, porpoises increasing minimum range (+23 to 32 m, respectively), whereas “M1” attracts porpoises, reducing range (by ? 29 m). As determined via archival acoustic detectors (AADs), both signals “F3” and “M1” led the animals to significantly intensify their click rate (by +10% and 68%, respectively) while signal “A” led to a significant reduction ( ? 59%). We propose that equipping fishing gear with PAL emitting signal “F3” could potentially reduce porpoise by-catch by increasing (1) awareness through enhanced echolocation and (2) distance to the nets. Detection probability and radius of PAL/AAD tandems could be improved by emitting signal “M1” to focus porpoise echolocation signals on the AAD. The signal may also be useful in luring animals away from hazards, which may be helpful for conservation measures prior to the onset of harmful acoustic activities such as pile-driving, seismic exploration or ammunition clearance.     

The modulation rate transfer function of a harbour porpoise (Phocoena phocoena)

During echolocation, toothed whales produce ultrasonic clicks at extremely rapid rates and listen for the returning echoes. The auditory brainstem response (ABR) duration was evaluated in terms of latency between single peaks: 5.5 ms (from peak I to VII), 3.4 ms (I–VI), and 1.4 ms (II–IV). In comparison to the killer whale and the bottlenose dolphin, the ABR of the harbour porpoise has shorter intervals between the peaks and consequently a shorter ABR duration. This indicates that the ABR duration and peak latencies are possibly related to the relative size of the auditory structures of the central nervous system and thus to the animal’s size. The ABR to a sinusoidal amplitude modulated stimulus at 125 kHz (sensitivity threshold 63 dB re 1 μPa rms) was evaluated to determine the modulation rate transfer function of a harbour porpoise. The ABR showed distinct envelope following responses up to a modulation rate of 1,900 Hz. The corresponding calculated equivalent rectangular duration of 263 μs indicates a good temporal resolution in the harbour porpoise auditory system similar to the one for the bottlenose dolphin. The results explain how the harbour porpoise can follow clicks and echoes during echolocation with very short inter click intervals.     

ECHOLOCATION IN DOLPHINS WITH A DOLPHIN-BAT COMPARISON

ABSTRACT

Dolphins possess a highly sophisticated auditory system and a keen capability for echolocation. Signals are emitted in the form of high intensity, short duration, broadband exponentially decaying pulses. The frequency spectra of echolocation signals used by many dolphins are dependent on the output intensity of the signals and not on any fine tuning by the animals. When the output intensity is low, the center frequency of the click tends to be low. As the output intensity increases, the center frequency also tends to increase. The pulses propagate from the dolphin's melon in a relatively narrow beam, and echoes are received via the lower jaw, with a slightly wider beam. Echo- locating dolphins can detect targets at ranges of approximately 100 plus meters, depending on the size of the targets. Target discrimination experiments have shown that dolphins can discriminate the shape, size, material composition and internal structure of targets from the echoes. The broadband short duration properties of the signal allow the echoes to have high temporal resolution, so that within the structure of the echoes a considerable amount of information on the properties of the target can be conveyed. A brief comparison between the bat and dolphin sonar system will also be made. Bats typically emit much longer signals and a wider variety of different types of signals than dolphins. Signals used by some bats are suited to detecting Doppler shift, whereas the dolphin signal is designed to be tolerant of Doppler effects.     

REACTIONS OF CAPTIVE HARBOR PORPOISES (PHOCOENA PHOCOENA) TO PINGER-LIKE SOUNDS

Pingers on gill nets can reduce bycatch of harbor porpoises. If harbor porpoises habituate to pingers, the effect may be reduced or lost. Two captive harbor porpoises were exposed to three sound types. All sounds were in the frequency band from 100 kHz to 140 kHz, 200 ms long, and presented once per 4 s. The source level was 153 dB re 1 μPa RMS at 1 m. Each session consisted of a 10‐min presound, a 5‐min sound, and a 10‐min postsound period. Behavior was recorded on video and on dataloggers placed on the dorsal fin of one animal. The loggers recorded heart rate, swimming speed, dive duration, and depth. The animals responded most strongly to the initial presentations of a sound. Surface time decreased, the heart rate dropped below the normal bradycardia, and echolocation activity decreased. The reactions of both animals diminished rapidly in the following sessions. Should the waning of responsiveness apply to wild animals, porpoises may adapt to the sounds but still avoid nets, or the bycatch may increase after some time. The success of long‐term use of pingers may then depend on the variety of sounds and rates of exposure.     

Echolocation of the big red bat Lasiurus egregius (Chiroptera: Vespertilionidae) and first record from the Central Brazilian Amazon

Lasiurus egregius (Peters, 1870) is a rare Neotropical vespertilionid bat and virtually no data on its ecology and echolocation calls are currently available. We report the capture of four individuals in the Central Amazon, representing the first record for the region and a significant (> 800 km) expansion of the species’ known range. Echolocation calls, recorded for the first time under natural conditions, were 1.5–8 ms in duration, and characterized by high mean bandwidth (18 kHz) and a mean frequency of maximum energy of 30 kHz.     

GEOGRAPHIC VARIATION IN RATES OF VOCAL PRODUCTION OF FREE-RANGING BOTTLENOSE DOLPHINS

Echolocation and whistle production, group sizes, and activities of free-ranging bottlenose dolphins were compared across four regions (Wilmington, NC Intracoastal Waterway [ICW]; Wilmington coastline; Southport, NC coastline; and Sarasota, FL inshore waters). Number of whistles and echolo-cation bouts differed significantly across sites. Dolphins whistled significantly more in Southport than in the other sites, independent of group size. Unlike at the other sites, dolphin vocalizations in Southport did not vary significantly across activities; this difference may be due to the fact that Southport animals were often found behind shrimp-trawling vessels, which may affect their behavior. Resident Sarasota dolphins vocalized significantly less than dolphins at the NC sites. At most sites, echolocation production per dolphin decreased as group size increased, supporting the idea that echolocation information is shared. In the ICW and Sarasota, echolocation production per dolphin was highest while feeding, indicating that echolocation is used in foraging. At all sites but Southport, whistle production per dolphin was highest while socializing, indicating that whistles are used in communication. Overall, these data show that dolphins have different vocal and activity patterns at different sites; thus, caution should be used when extrapolating results from one study site to another.     

Changes in the acoustic behavior of resident bottlenose dolphins near operating vessels

Maritime traffic is an issue of major ecological concern, and vessel noise may be an important source of disturbance for coastal cetaceans. In the Sado estuary, Portugal, core habitat areas of a small resident population of bottlenose dolphins (Tursiops truncatus) overlap with routes of intense maritime traffic, which presents an opportunity to assess vocal responses of these dolphins to specific vessel noise sources. Field recordings of dolphin vocalizations were made from April to November 2011, using a calibrated system. Dolphin behavior and group size were recorded, as well as the operating boat condition (no boats or specific boat type) in a 1,000 m radius. Spectral analyses of vocalizations allowed the categorization and quantitative analysis of echolocation click trains and social calls, including whistles. Mean overall call rates decreased significantly in the presence of operating vessels. Creaks (fast click trains) were significantly reduced in the presence of ferry boats. Significant differences were also observed in the whistles' minimum, maximum, and start frequencies. These changes in call emission rates and temporary shifts in whistles characteristics may be a vocal response to the proximity of operating vessels, facilitating communication in this busy, noisy estuary.