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Dolphins use their biosonar to discriminate objects with different features through the returning echoes. Cross-modal matching experiments were conducted with a resident bottlenose dolphin (Tursiops aduncus). Four types of objects composed of different materials (water-filled PVC pipes, air-filled PVC pipes, foam ball arrays, and PVC pipes wrapped in closed-cell foam) were used in the experiments, respectively. The size and position of the objects remained the same in each case. The data collected in the experiment showed that the dolphin’s matching accuracy was significantly different across the cases. To gain insight into the underlying mechanism in the experiments, we used finite element methods to construct two-dimensional target detection models of an echolocating dolphin in the vertical plane, based on computed tomography scan data. The acoustic processes of the click’s interaction with the objects and the surrounding media in the four cases were simulated and compared. The simulation results provide some possible explanations for why the dolphin performed differently when discriminating the objects that only differed in material composition in the previous matching experiments.

Echolocation signals emitted by odontocetes can be roughly classified into three broad categories: broadband echolocation signals, narrowband high-frequency echolocation signals, and frequency modulated clicks. Previous measurements of broadband echolocation signal propagation in the bottlenose dolphin (Tursiops truncatus) did not find any evidence of focusing as the signals travel from the near-field to far-field. Finite element analysis (FEA) of high-resolution computed tomography scan data was used to examine signal propagation of broadband echolocation signals of dolphins and narrowband echolocation signals of porpoises. The FEA results were used to simulate the propagation of clicks from phonic lips, traveling through the forehead, and finally transmission into the water. Biosonar beam formation in the near-field and far-field, including the amplitude contours for the two species, was determined. The finite element model result for the simulated amplitude contour in the horizontal plane was consistent with prior direct measurement results for Tursiops, validating the model. Furthermore, the simulated far-field transmission beam patterns in both the vertical and horizontal planes were also qualitatively consistent with results measured from live animals. This study indicates that there is no evidence of convergence for either Tursiops or Phocoena as the sound propagates from the near-field to the far-field.

Bio-logging devices are advancing the understanding of marine animal behavior, but linking sound production and behavior of individual baleen whales is still unreliable. Tag placement potentially within the near field of the sound source creates uncertainty about how tagged animal sounds will register on recorders. This study used data from a tagged singing humpback whale to evaluate this question of how sound levels present on a tag when calls are produced by a tagged animal. Root-mean-square (rms) received levels (RLs) of song units ranged from 112 to 164 dB re 1 μPa rms, with some, but not all, of the lower frequency units registering on the tag's 800 Hz accelerometer sensor. Fifty-nine percent of recorded units measured lower acoustic RLs than previously reported source levels for humpback song, but signal-to-noise ratios (SNRs) were 30–45 dB during periods of the dive with low noise. This research highlights that tag RL does not alone predict caller identity, argues for higher SNR thresholds if using SNR to inform decisions about the source of a call, and provides a baseline for future research identifying diagnostic properties of tagged animal calls in cetacean bioacoustic tag datasets.

Pelagic animals that form deep sea scattering layers (DSLs) represent an important link in the food web between zooplankton and top predators. While estimating the composition, density and location of the DSL is important to understand mesopelagic ecosystem dynamics and to predict top predators’ distribution, DSL composition and density are often estimated from trawls which may be biased in terms of extrusion, avoidance, and gear-associated biases. Instead, location and biomass of DSLs can be estimated from active acoustic techniques, though estimates are often in aggregate without regard to size or taxon specific information. For the first time in the open ocean, we used a DIDSON sonar to characterize the fauna in DSLs. Estimates of the numerical density and length of animals at different depths and locations along the Kona coast of the Island of Hawaii were determined. Data were collected below and inside the DSLs with the sonar mounted on a profiler. A total of 7068 animals were counted and sized. We estimated numerical densities ranging from 1 to 7 animals/m3 and individuals as long as 3 m were detected. These numerical densities were orders of magnitude higher than those estimated from trawls and average sizes of animals were much larger as well. A mixed model was used to characterize numerical density and length of animals as a function of deep sea layer sampled, location, time of day, and day of the year. Numerical density and length of animals varied by month, with numerical density also a function of depth. The DIDSON proved to be a good tool for open-ocean/deep-sea estimation of the numerical density and size of marine animals, especially larger ones. Further work is needed to understand how this methodology relates to estimates of volume backscatters obtained with standard echosounding techniques, density measures obtained with other sampling methodologies, and to precisely evaluate sampling biases.

Harbor porpoises (Phocoena phocoena) use narrow band echolocation signals for detecting and locating prey and for spatial orientation. In this study, acoustic impedance values of tissues in the porpoise's head were calculated from computer tomography (CT) scan and the corresponding Hounsfield Units. A two-dimensional finite element model of the acoustic impedance was constructed based on CT scan data to simulate the acoustic propagation through the animal's head. The far field transmission beam pattern in the vertical plane and the waveforms of the receiving points around the forehead were compared with prior measurement results, the simulation results were qualitatively consistent with the measurement results. The role of the main structures in the head such as the air sacs, melon and skull in the acoustic propagation was investigated. The results showed that air sacs and skull are the major components to form the vertical beam. Additionally, both beam patterns and sound pressure of the sound waves through four positions deep inside the melon were demonstrated to show the role of the melon in the biosonar sound propagation processes in the vertical plane.

A dual frequency identification sonar was used to estimate density and size of potential deep diving odontocetes' prey in deep sea scattering layers in Hawaii. Size distributions data showed a bimodal pattern, with a population of data above the 50 cm of length, possibly big squids, and a population of data below 50 cm, mainly micronekton. The number of animals bigger than 50 cm was correlated with the length of the peak of the frequency distribution. Data suggest that big predators, like squids that constitute prey for beaked, pilot, and sperm whales, are more likely to be found where they have better chance to find micronectonic prey.

Acoustic backscatter measurements were conducted on a stationary harbor porpoise (Phocoena phocoena) under controlled conditions. The measurements were made with the porpoise in the broadside aspect using three different types of signals: (1) a 475 μs linear frequency-modulated (FM) pulse with a frequency range from 23 to 160 kHz; (2) a simulated bottlenose dolphin (Tursiops “truncates”) click with a peak frequency of 120 kHz; and (3) a simulated killer whale (Orcinus orca) click with a peak frequency of 60 kHz. The measurement with the FM pulse indicated that the mean target strength at the broadside aspect decreased from −26 to −50 dB as the frequency increased from 23 to 120 kHz in a nearly linear fashion (on a logarithm plot). Target strength variation with frequency was similar to a previous backscatter measurement on a bottlenose dolphin over a comparable frequency range (23–80 kHz). The porpoise seems to be a stealth body with low backscatter properties. The target strength of the porpoise was also about 15−16 dB lower than that of the bottlenose dolphin. The difference in lung volume of the two species when expressed in dB was also approximately 15 dB.Acoustic backscatter measurements were conducted on a stationary harbor porpoise (Phocoena phocoena) under controlled conditions. The measurements were made with the porpoise in the broadside aspect using three different types of signals: (1) a 475 μs linear frequency-modulated (FM) pulse with a frequency range from 23 to 160 kHz; (2) a simulated bottlenose dolphin (Tursiops “truncates”) click with a peak frequency of 120 kHz; and (3) a simulated killer whale (Orcinus orca) click with a peak frequency of 60 kHz. The measurement with the FM pulse indicated that the mean target strength at the broadside aspect decreased from −26 to −50 dB as the frequency increased from 23 to 120 kHz in a nearly linear fashion (on a logarithm plot). Target strength variation with frequency was similar to a previous backscatter measurement on a bottlenose dolphin over a comparable frequency range (23–80 kHz). The porpoise seems to be a stealth body with low backscatter properties. The target strength of the porpoise was also...

The International Seabed Authority (ISA) is developing regulations to control the future exploitation of deep-sea mineral resources including sulphide deposits near hydrothermal vents, polymetallic nodules on the abyssal seafloor, and cobalt crusts on seamounts. Under the UN Convention on the Law of the Sea the ISA is required to adopt are taking measures to ensure the effective protection of the marine environment from harmful effects arising from mining-related activities. Contractors are required to generate environmental baselines and assess the potential environmental consequences of deep seafloor mining. Understandably, nearly all environmental research has focused on the seafloor where the most direct mining effects will occur. However, sediment plumes and other impacts (e.g., noise) from seafloor mining are likely to be extensive in the water column. Sediment plumes created on the seafloor will affect the benthic boundary layer which extends 10s to 100s of meters above the seafloor. Separation or dewatering of ore from sediment and seawater aboard ships will require discharge of a dewatering plume at some depth in the water column. It is important to consider the potential impacts of mining on the ocean’s midwaters (depths from ~200 m to the seafloor) because they provide vital ecosystem services and harbor substantial biodiversity. The better known epipelagic or sunlit surface ocean provisions the rest of the water column through primary production and export flux (This was not the focus at this workshop as the subject was considered too large and surface discharges are unlikely). It is also home to a diverse community of organisms including commercially important fishes such as tunas, billfish, and cephalopods that contribute to the economies of many countries. The mesopelagic or twilight zone (200-1000 m) is dimly lit and home to very diverse and abundant communities of organisms. Mesopelagic plankton and small nekton form the forage base for many deep-diving marine mammals and commercially harvested epipelagic species. Furthermore, detritus from the epipelagic zone falls through the mesopelagic where it is either recycled, providing the vital process of nutrient regeneration, or sinks to greater depths sequestering carbon from short-term atmospheric cycles. The waters below the mesopelagic down to the seafloor (both the bathypelagic and abyssopelagic) are very poorly characterized but are likely large reservoirs of novel biodiversity and link the surface and benthic ecosystems. Great strides have been made in understanding the biodiversity and ecosystem function of the ocean’s midwaters, but large regions, including those containing many exploration license areas and the greater depths where mining plumes will occur, remain very poorly studied. It is clear that pelagic communities are distinct from those on the seafloor and in the benthic boundary layer. They are often sampled with different instrumentation. The fauna have relatively large biogeographic ranges and they are more apt to mix freely across stakeholder boundaries, reference areas and other spatial management zones. Pelagic organisms live in a three-dimensional habitat and their food webs and populations are vertically connected by daily or lifetime migrations and the sinking flux of detritus from the epipelagic. The fauna do not normally encounter hard surfaces, making them fragile, and difficult to capture and maintain for sensitivity or toxicity studies. Despite some existing general knowledge, ecological baselines for midwater communities and ecosystems that likely will be impacted by mining have not been documented. There is an urgent need to conduct more research and evaluate the midwater biota (microbes to fishes) in regions where mining is likely to occur. Deep-sea mining activities may affect midwater organisms in a number of ways, but it is still unclear at what scale perturbations may occur. The sediment plumes both from collectors on the seafloor and from midwater discharge will have a host of negative consequences. They may cause respiratory distress from clogged gills or respiratory surfaces. Suspension feeders, such as copepods, polychaetes, salps, and appendicularians, that filter small particles from the water and form an important basal group of the food web, may suffer from dilution of their food by inorganic sediments and/or clogging of their fragile mucous filter nets. Small particles may settle on gelatinous plankton causing buoyancy issues. Metals, including toxic elements that will enter the food web, will be released from pore waters and crushed ore materials. Sediment plumes will also absorb light and change backscatter properties, reducing visual communication and bioluminescent signaling that are very important for prey capture and reproduction in midwater animals. Noise from mining activities may alter the behaviors of marine mammals and other animals. Small particles have high surface area to volume ratios, high pelagic persistence and dispersal and as a result greater potential to result in pelagic impacts. All of these potential effects will result in mortality, migration (both horizontal and vertical), decreased fitness, and shifts in community composition. Depending on the scale and duration of these effects, there could be reduction in provisioning to commercial fish species, delivery of toxic metals to pelagic food webs and hence human seafood supply, and alterations to carbon transport and nutrient regeneration services. After four days of presentations and discussions, the workshop participants came to several conclusions and synthesized recommendations. 1. Assuming no discharge in the epipelagic zone, it is essential to minimize mining effects in the mesopelagic zone because of links to our human seafood supply as well as other ecosystem services provided by the mesopelagic fauna. This minimization could be accomplished by delivering dewatering discharge well below the mesopelagic/bathypelagic transition (below ~1000 m depth). 2. Research should be promoted by the ISA and other bodies to study the bathypelagic and abyssopelagic zones (from ~1000 m depths to just above the seafloor). It is likely that both collector plumes and dewatering plumes will be created in the bathypelagic, yet this zone is extremely understudied and contains major unknowns for evaluating mining impacts. 3. Management objectives, regulations and management actions need to prevent the creation of a persistent regional scale “haze” (enhanced suspended particle concentrations) in pelagic midwaters. Such a haze would very likely cause chronic harm to deep midwater ecosystem biodiversity, structure and function. 4. Effort is needed to craft suitable standards, thresholds, and indicators of harmful environmental effects that are appropriate to pelagic ecosystems. In particular, suspension feeders are very important ecologically and are likely to be very sensitive to sediment plumes. They are a high priority for study. 5. Particularly noisy mining activities such as ore grinding at seamounts and hydrothermal vents is of concern to deep diving marine mammals and other species. One way to minimize sound impacts would be to minimize activities in the sound-fixing-and-ranging (SOFAR) channel (typically at depths of ~1000 m) which transmits sounds over very long distances. 6. A Lagrangian (drifting) perspective is needed in monitoring and management because the pelagic ecosystem is not a fixed habitat and mining effects are likely to cross spatial management boundaries. For example, potential broad-scale impacts to pelagic ecosystems should be considered in the deliberations over preservation reference zones, the choice of stations for environmental baseline and monitoring studies and other area-based management and conservation measures. 7. Much more modeling and empirical study of realistic mining sediment plumes is needed. Plume models will help evaluate the spatial and temporal extent of pelagic (as well as benthic) ecosystem effects and help to assess risks from different technologies and mining scenarios. Plume modeling should include realistic mining scenarios (including duration) and assess the spatial-temporal scales over which particle concentrations exceed baseline levels and interfere with light transmission to elucidate potential stresses on communities and ecosystem services. Models should include both near and far field-phases, incorporating realistic near field parameters of plume generation, flocculation, particle sinking, and other processes. It is important to note that some inputs to these models such as physical oceanographic parameters are lacking and should be acquired in the near-term. Plume models need to be complemented by studies to understand effects on biological components by certain particle sizes and concentrations.

The echolocation signals emitted by odontocetes can be roughly classified into three broad categories: broadband high-frequency echolocation signals, narrowband high-frequency echolocation signals, and frequency modulated clicks. Previous measurements of broadband echolocation signals propagation in the bottlenose dolphin (Tursiops truncatus) did not find any evidence of focusing as the signals travel from the near to far-field. Finite element analysis (FEA) of high-resolution CT scan data was used to examine signal propagation of broadband echolocation signals of dolphins and narrowband echolocation signals of porpoises. The FEA results were used to simulate the propagation of clicks from phonic lips, traveling through the forehead, and finally transmission into the seawater. Biosonar beam formation in the near-field and far-field including the amplitude contours for the two species was determined. The finite element model was validated by finding the simulated amplitude contour in the horizontal plane was consistent with prior direct measurement results for Tursiops. Furthermore, the simulated far-field transmission beam patterns in both vertical and horizontal planes were also qualitatively consistent with results measured from live animals. This study shows that there is no evidence of convergence for either Tursiops or Phocoena as the sound propagates from near-field to far-field.

Beaked and sperm whales are top predators living in the waters off the Kona coast of Hawai'i. Temporal and spatial analyses of the foraging activity of these two species were studied with passive acoustics techniques. Three passive acoustics recorders moored to the ocean floor were used to monitor the foraging activity of these whales in three locations along the Kona coast of the island of Hawaii. Data were analyzed using automatic detector/classification systems: M3R (Marine Mammal Monitoring on Navy Ranges), and custom-designed Matlab programs. The temporal variation in foraging activity was species-specific: beaked whales foraged more at night in the north, and more during the day-time off Kailua-Kona. No day-time/night-time preference was found in the southern end of the sampling range. Sperm whales foraged mainly at night in the north, but no day-time/night-time preference was observed off Kailua-Kona and in the south. A Generalized Linear Model was then applied to assess whether location and chlorophyll concentration affected the foraging activity of each species. Chlorophyll concentration and location influenced the foraging activity of both these species of deep-diving odontocetes.

Understanding the distribution of animals is of paramount importance for management and conservation, especially for species that are impacted by anthropogenic threats. In the case of marine mammals there has been a growing concern about the impact of human-made noise, in particular for beaked whales and other deep diving odontocetes. Foraging (measured via echolocation clicks at depth) was studied for Cuvier’s beaked whale ( Ziphius cavirostris ), sperm whale ( Physeter macrocephalus ), long-finned pilot whales ( Globicephala melas ) and Risso’s dolphin ( Grampus griseus ) using three passive acoustics recorders moored to the bottom of the ocean in a canyon area in the Ligurian Sea between July and December 2011. A Generalized Linear Model was used to test whether foraging was influenced by location and day of the year, including the possibility of interactions between predictors. Contrary to previous studies conducted by visual surveys in this area, all species were detected at all locations, suggesting habitat overlapping. However, significant differences were found in the occurrence of each species at different locations. Beaked and sperm whales foraged significantly more in the northern and western locations, while long-finned pilot whales and Risso’s dolphins hunted more in the northern and eastern location.

The biosonar signals of two free-swimming Atlantic bottlenose dolphins performing a complex sonar search for a bottom target in San Diego Bay were compared with the biosonar signals of a dolphin performing a target discrimination task in a net pen in the same bay. A bite-plate device carried by the free-swimming dolphins supported a hydrophone that extended directly in front of the dolphin. A biosonar measuring tool attached to the bite plate measured the outgoing biosonar signals while the dolphins conducted sonar searches. Each of the free-swimming dolphins used different biosonar search strategy in solving the problem and the dolphins' biosonar signals reflect the difference in strategy. The dolphin in the pen stationed in a hoop while echolocating on a target 6 m away and reported if the indentation on a spherical target was directed toward it. The signals were parameterized by determining the peak-to-peak source levels, source energy flux density, peak frequency, center frequency, root-mean-square (rms) bandwidth, rms duration, and the Q of the signals. Some parameters were similar for the free-swimming and stationary dolphins while some were significantly different, suggesting biosonar signals used by free-swimming animals may be different than signals used by dolphins in a pen.

The relative role of the various structures in the head of the baiji (Lipotes vexillifer) is examined. A finite element approach was applied to numerically simulate the acoustic propagation through a dolphin's head to examine the relative role of the skull, air sacs, and melon in the formation of the biosonar beam in the vertical plane. The beam pattern obtained with the whole head in place is compared with the beam pattern when the air sac is removed and the other structures (skull and melon) are in place, with only the skull removed, and finally with only the melon removed. The beam pattern with the air sacs and skull intact and the melon removed closely resembled the beam pattern for the complete head, suggesting that the melon has a minor role in the formation of the beam. The beam pattern for the other two cases had very little resemblance to the beam pattern for the whole head. The air sacs seem to have a role of directing propagation of the signal toward the front and the skull prevents the sound propagating below the rostrum. The beam patterns along with a correlation analysis showed that the melon had only a slight influence on the shape and direction of the beam. The resultant beam exiting the head of the dolphin is the result of complex reflection processes within the head of the animal.

Characterizing the trophic roles of deep-diving odontocete species and how they vary in space and time is challenged by our ability to observe foraging behavior. Though sampling methods are limited, foraging activity of deep-diving odontocetes can be monitored by recording their biosonar emissions. Daily occurrence of echolocation clicks was monitored acoustically for five months (July–December 2011) in the Ligurian Sea (Mediterranean Sea) using five passive acoustic recorders. Detected odontocetes included Cuvier's beaked whales ( Zipuhius cavirostris ), sperm whales ( Physeter macrocephalus ), Risso's dolphins ( Grampus griseus ), and long-finned pilot whales ( Globicephala melas ). The results indicated that the foraging strategies varied significantly over time, with sperm whales switching to nocturnal foraging in late September whereas Risso's dolphins and pilot whales foraged mainly at night throughout the sampling period. In the study area, winter nights are about five hours longer than summer nights and an analysis showed that pilot whales and Risso's dolphins adjusted their foraging activity with the length of the night, foraging longer during the longer winter nights. This is the first study to show that marine mammals exhibit diurnal foraging patterns closely correlated to sunrise and sunset.

Bottlenose dolphins project broadband echolocation signals for detecting and locating prey and predators, and for spatial orientation. There are many unknowns concerning the specifics of biosonar signal production and propagation in the head of dolphins and this manuscript represents an effort to address this topic. A two-dimensional finite element model was constructed using high resolution CT scan data. The model simulated the acoustic processes in the vertical plane of the biosonar signal emitted from the phonic lips and propagated into the water through the animal's head. The acoustic field on the animal's forehead and the farfield transmission beam pattern of the echolocating dolphin were determined. The simulation results and prior acoustic measurements were qualitatively extremely consistent. The role of the main structures on the sound propagation pathway such as the air sacs, melon, and connective tissue was investigated. Furthermore, an investigation of the driving force at the phonic lips for dolphins that emit broadband echolocation signals and porpoises that emit narrowband echolocation signals suggested that the driving force is different for the two types of biosonar. Finally, the results provide a visual understanding of the sound transmission in dolphin's biosonar.

Radiated noise from a high-speed jet-propelled watercraft (the M/V Alakai, 1,646 tons, length 117 m) was measured at hydrophone depths of 3, 6, and 10 m while the ship passed by at speeds of 6.17333 m/sec (12 knots), 12.3467 m/sec (24 knots), and 19.0344 m/sec (37 knots). Noise spectra were similar for all speeds and hydrophone depths. Spectra peaked below 100 Hz and dropped off continuously at higher frequencies. Calculated source level noise was 10 to 20 dB lower than noise from propeller-driven ships and much lower than for ships of similar speed. Although exposure to noise radiating from the M/V Alakai over short time periods is unlikely to cause hearing damage to whales, the combination of low radiated noise levels and high transit speeds leads to a shorter closing time (defined as time between when source level of the ship at a stationary receiver is greater than ambient noise and time that a ship traveling directly toward the receiver arrives at its location) between ship and whale. Compared with other types of ships traveling at similar speeds, closing time for the Alakai ranges from 20 sec shorter (at 6.17333 m/sec [12 knots]) to 22 min shorter (at 19.0344 m/sec [37 knots]). Shortest closing time for the Alakai is 89.1 sec at a speed of 6.17333 m/sec (12 knots). Shortened closing time might reduce successful detection and avoidance of high-speed jet-propelled ships by whales, and increased speed shortens the time during which whales have the opportunity to respond to this detection.

Sperm whales forage in the deep ocean, hunting for squid. An innovative approach for the study of sperm whale foraging behavior and habitat selection is reported in this letter. A DIDSON imaging sonar mounted on a profiler with a conductivity, temperature, and depth sensor was used to count and measure potential prey in the deep ocean during sperm whales' acoustical foraging encounters in Hawaii. Preliminary results show how this technique can be applied to the study of deep diving whale foraging and habitat selection. Sperm whales foraged where the density of prey decreased with depth and where the size of prey increased with depth.

The target detection capability of bottlenose dolphins in the presence of artificial masking noise was first studied by Au and Penner [J. Acoust. Soc. Am. 70, 687-693 (1981)] in which the dolphins' target detection threshold was determined as a function of the ratio of the echo energy flux density and the estimated received noise spectral density. Such a metric was commonly used in human psychoacoustics despite the fact that the echo energy flux density is not compatible with noise spectral density which is averaged intensity per Hz. Since the earlier detection in noise studies, two important parameters, the dolphin integration time applicable to broadband clicks and the dolphin's auditory filter shape, were determined. The inclusion of these two parameters allows for the estimation of the received energy flux density of the masking noise so that the dolphin target detection can now be determined as a function of the ratio of the received energy of the echo over the received noise energy. Using an integration time of 264 μs and an auditory bandwidth of 16.7 kHz, the ratio of the echo energy to noise energy at the target detection threshold is approximately 1 dB.

The baiji (Lipotes vexillifer) of the Yangtze River possesses a sophisticated biosonar system. In this study, a finite element approach was used to numerically investigate the propagation of acoustic waves through the head of the Yangtze River dolphin, which possesses an inhomogeneous and complex structure. The acoustic intensity distribution predicted from models with and without the melon and/or skull showed that the emitted sound beam was narrow and formed a highly directed acoustic beam, and the skull and melon significantly enhanced the directional characteristics of the emitted sound. Finally, for a short duration impulsive source, the emitted sound pressure distributions were also simulated at different propagation times. The results provide useful information for better understanding the operation of the biosonar system in this rare and perhaps extinct animal.

Ecological acoustic recorders (EARs) were moored off the bottom in relatively deep depths (609-710 m) at five locations around the island of Kauai. Initially, the EARs had an analog-to-digital sample rate of 64 kHz with 30-s recordings every 5 min. After the second deployment the sampling rate was increased to 80 kHz in order to better record beaked whale biosonar signals. The results of the 80 kHz recording are discussed in this manuscript and are the results of three deployments over a year's period (January 2010 to January 2011). Five categories of the biosonar signal detection of deep diving odontocetes were created, short-finned pilot whales, sperm whales, beaked whales, Risso's dolphins, and unknown dolphins. During any given day, at least one species of these deep diving odontocetes were detected. On many days, several species were detected. The biosonar signals of short-finned pilot whales were detected the most often with approximately 30% of all the signals, followed by beaked and sperm whales approximately 22% and 21% of all clicks, respectively. The seasonal patterns were not very strong except in the SW location with distinct peak in detection during the months of April-June 2010 period.

In 1974, Norris and Harvey measured the sound velocity profile of the melon of a just deceased bottlenose dolphin and found a low-velocity core in the melon with the velocity increasing towards the surface. This was the genesis of the “melon focusing” hypothesis in the formation of the biosonar beam. Aroyan solved the wave equation for a signal produced at the nasal plug using a finite difference approach and found that the skull was responsible for the biosonar beam. Aroyan’s results were never taken seriously and the notion that the melon was mainly responsible for the biosonar beam continues to this day. We used a finite element approach to solve the wave equation for a broadband signal generated at the phonic lips. CT scan data were used to provide a map of the acoustic impedance of the dolphin’s head and our results compared well with different measurements made around the dolphin’s head and in the far field. Our findings also showed that the skull and air sacs are the most influential structures in the formation of the biosonar beam and the melon was responsible for directing the axis of the beam. Our results should put to rest the “melon-focusing” hypothesis.In 1974, Norris and Harvey measured the sound velocity profile of the melon of a just deceased bottlenose dolphin and found a low-velocity core in the melon with the velocity increasing towards the surface. This was the genesis of the “melon focusing” hypothesis in the formation of the biosonar beam. Aroyan solved the wave equation for a signal produced at the nasal plug using a finite difference approach and found that the skull was responsible for the biosonar beam. Aroyan’s results were never taken seriously and the notion that the melon was mainly responsible for the biosonar beam continues to this day. We used a finite element approach to solve the wave equation for a broadband signal generated at the phonic lips. CT scan data were used to provide a map of the acoustic impedance of the dolphin’s head and our results compared well with different measurements made around the dolphin’s head and in the far field. Our findings also showed that the skull and air sacs are the most influential structures in ...

Sensitivity of echolocating dolphins to phase changes within echoes may be a vital piece of information when constructing echolocation models. Previous experiments have yielded ambiguous results leaving it unclear what cues might have been used by passively listening dolphins to discriminate between different phase altered signals. This study used a phantom echo generator to produce computer controlled echoes. The dolphin interacted with the system in a real echolocation task to discriminate between simulated targets that were unaltered and those that had a 180° phase shift. The frequency amplitude spectral content between the two targets was the same. There were no temporal differences between the two targets. The only cue that the dolphin could use to discriminate between them was the 180° phase shift. The dolphin preformed at a success level of 40% in discriminating the two echoes. This indicates that the 180° phase shift was not perceived.

Male humpback whales produce loud “songs” on the wintering grounds and some sing while escorting mother–calf pairs, exposing them to near-continuous sounds at close proximity. An Acousonde acoustic and movement recording tag deployed on a calf off Maui, Hawaii captured sounds produced by a singing male escort. Root-mean-square received levels ranged from 126 to 158 dB re 1 μPa. These levels represent rare direct measurements of sound to which a newly born humpback calf may be naturally exposed by a conspecific, and may provide a basis for informed decisions regarding anthropogenic sound levels projected near calves.

Grants for this study was provided by the National Natural Science Foundation (NNSF) of China (Grant No.31070347), the Ministry of Science and Technology of China (Grant No. 2011BAG07B05-3), the Knowledge Innovation Program of the Chinese Academy of Sciences (Grant No. KSCX2-EW-Z-4) and the Special Fund for Agro-scientific Research in the Public Interest of the Ministry of Agriculture of China (Grant No. 201203086) to DW, the State Oceanic Administration of China (Grant No. 201105011-3) and NNSF of China (Grant No. 31170501) to KXW and the China Scholarship Council (Grant No. (2014)3026) to ZTW. Background . Knowledge of species-specific vocalization characteristics and their associated active communication space, the effective range over which a communication signal can be detected by a conspecific, is critical for understanding the impacts of underwater acoustic pollution, as well as other threats. Methods. We used a two-dimensional cross-shaped hydrophone array system to record the whistles of free-ranging Indo-Pacific humpback dolphins (Sousa chinensis) in shallow-water environments of the Pearl River Estuary (PRE) and Beibu Gulf (BG), China. Using hyperbolic position fixing, which exploits time differences of arrival of a signal between pairs of hydrophone receivers, we obtained source location estimates for whistles with good signal-to-noise ratio (SNR ≥ 10 dB) and not polluted by other sounds and back-calculated their apparent source levels (ASL). Combining with the masking levels (including simultaneous noise levels, masking tonal threshold, and the Sousa auditory threshold) and the custom made site-specific sound propagation models, we further estimated their active communication space (ACS). Results. Humpback dolphins produced whistles with average root-mean-square ASL of 138.5 ± 6.8 (mean ± standard deviation) and 137.2 ± 7.0 dB re 1 µPa in PRE (N = 33) and BG (N = 209), respectively. We found statistically significant differences in ASLs among different whistle contour types. The mean and maximum ACS of whistles were estimated to be 14.7 ± 2.6 (median ± quartile deviation) and 17.1 ± 3.5 m in PRE, and 34.2 ± 9.5 and 43.5 ±12.2 m in BG. Using just the auditory threshold as the masking level produced the mean and maximum ACSat of 24.3 ± 4.8 and 35.7 ± 4.6 m for PRE, and 60.7 ± 18.1 and 74.3 ± 25.3 m for BG. The small ACSs were due to the high ambient noise level. Significant differences in ACSs were also observed among different whistle contour types. Discussion. Besides shedding some light for evaluating appropriate noise exposure levels and information for the regulation of underwater acoustic pollution, these baseline data can also be used for aiding the passive acoustic monitoring of dolphin populations, defining the boundaries of separate groups in a more biologically meaningful way during field surveys, and guiding the appropriate approach distance for local dolphin-watching boats and research boat during focal group following. Publisher PDF

The Station ALOHA Cabled Observatory (ACO) is an ocean-bottom observatory that allows continuous real-time monitoring of ocean processes including sounds produced by baleen whales. Baleen whales can be challenging to study using traditional visual methods due to their cryptic behavior and offshore ranges. Many baleen whales produce distinctive sounds that propagate well under water and so ocean-bottom hydrophones like the one at the ACO can be used to investigate the occurrence and acoustic behavior of these animals in locations that are difficult to access and study long-term using other methods. We examined 12 months of recordings from the ACO (February 2007–February 2008) and found that sounds produced by blue, sei, and minke whales all occurred seasonally between October and April. Low-frequency pulses produced by fin whales were detected year-round, although much less frequently during the summer months than during the winter months. These seasonal patterns matched those of humpback whales, who migrate to Hawai’ian waters to breed and give birth. Blue, minke, fin, and sei whales are probably using Hawai’ian waters for breeding, but further research is necessary to confirm this. The ACO has provided, and continues to provide, a long-term dataset for investigating seasonal and diurnal trends in the occurrence of baleen whales and other cetaceans at a location that would be difficult to study any other way.

The radiated noise from a high-speed water-jet-propelled catamaran was measured for catamaran speeds of 12, 24, and 37 kn. The radiated noise increased with catamaran speed, although the shape of the noise spectrum was similar for all speeds and measuring hydrophone depth. The spectra peaked at ~200 Hz and dropped off continuously at higher frequencies. The radiated noise was 10-20 dB lower than noise from propeller-driven ships at comparable speeds. The combination of low radiated noise and high speed could be a factor in the detection and avoidance of water-jet-propelled ships by baleen whales.

The use of passive acoustic monitoring (PAM) to detect aquatic life continues to increase because PAM devices can be deployed in remote areas and can operate for months or years at a time in a programmed manner to control the recording time, the duration of each recording, and the time to “sleep” to preserve battery power. This introduction will discuss the early history of these tools, their architecture, their uses, and the organization of this book. The architecture of almost all PAM devices is similar in that a microcontroller is used to manage the analog to digital conversion process, the flow of data from either a buffer or directly into storage, and the mode in which the PAM will be used. There are basically two main modes, a continuous mode in which data are collected continuously and a programmed or duty-cycled mode. Some acoustic tags are designed just for short time applications (hours or several days) and are attached by suction cups on swimming animals. This book contains chapters from different researchers discussing some of the interesting and exciting findings they have made by listening in the ocean.

Despite the importance of young animals to the proliferation of a species, logistic hur- dles often prevent the study of individuals' behavior and habitat requirements. This is particularly an issue in the case of cetaceans, which spend a large proportion of their time at depth. We con- ducted a study to describe the dive behavior of young humpback whale Megaptera novaeangliae calves on their breeding grounds in Hawaii, USA. We first implemented and evaluated strategies for approaching whale groups and deploying suction-cup tags (DTAGs), resulting in 3 successful attachments of DTAGs in the winter of 2011. The approach technique that was most successful while minimizing reactions from the whale groups was a passive drift approach. Tagged calves exhibited consistent dives to shallow depths when their groups were stationary, and some deeper dives that approached the ocean bottom, up to 78 m in one case. Mean dive durations ranged from 2.2 to 3.5 min, with calves spending 40% of their time within 3 m of the surface. This is the first study to collect tag data from baleen whale calves less than 6 mo in age and provides habitat use data important for management of this endangered species.

Biosonar signals radiated along the beam axis of an Atlantic bottlenose dolphin resemble short transient oscillations. As the azimuth of the measuring hydrophones in the horizontal plane progressively increases with respect to the beam axis the signals become progressively distorted. At approximately ±45°, the signals begin to divide into two components with the time difference between the components increasing with increasing angles. At ±90° or normal to the longitudinal axis of the animal, the time difference between the two pulses measured by the hydrophone on the right side of the dolphin's head is, on average, ∼11.9 μs larger than the time differences observed by the hydrophone on the left side of the dolphin's head. The center frequency of the first pulse is generally lower, by 33-47 kHz, than the center frequency of the second pulse. When considering the relative locations of the two phonic lips, the data suggest that the signals are being produced by one of the phonic lips and the second pulse resulting from a reflection within the head of the animal. The generation of biosonar signals is a complex process and the propagation pathways through the dolphin's head are not well understood.

The auditory filter shape of delphinid odontocetes was previously considered to be typically mammalian constant-quality in which filter bandwidths increase proportionally with frequency. Recent studies with porpoises demonstrate constant-bandwidth portions of the auditory filter. The critical ratios for a bottlenose dolphin were measured between 40 and 120 kHz by behaviorally determining the subject's ability to detect pure tones in the presence of white noise. Critical ratios as a function of frequency were constant, indicating the auditory filter acts as a constant-bandwidth system in this frequency range. Re-analysis of past studies supports these findings, and suggests the delphinid auditory system is best characterized as a constant-Q system below 40 kHz and a constant-bandwidth-like system between 40 kHz and 120 kHz before returning to a constant Q pattern at the highest frequencies.

The main Hawaiian Islands (MHI) are the principal breeding ground for humpback whales in the North Pacific. Over the past 3 decades, population recovery from whaling-era losses has resulted in a steady increase in the number of whales wintering in Hawaiian waters and a geo- graphic expansion of their distribution in the MHI. Until recently, no existing evidence showed that this expansion included the islands, atolls, and banks of the Northwestern Hawaiian Islands (NWHI). To better understand the occurrence of humpback whales in the NWHI, 9 ecological acoustic recorders (EARs) were deployed at sites throughout the archipelago to record the occurrence of humpback whale song, an indicator of winter breeding activity. Song was found to be prevalent at Maro Reef, Lisianski Island, and French Frigate Shoals but was also recorded at Kure Atoll, Midway Atoll, and Pearl and Hermes Atoll. Both the timing and abundance of song at several locations closely followed trends observed on Oahu, which is one of the MHI, strongly suggesting that humpback whales use the NWHI as a wintering area. This finding is of particular relevance in light of recent suggestions that, based on photo-identification data, a yet undocumented wintering area exists somewhere in the central North Pacific. We propose that the NWHI could be that area.

The "resident" ecotype of killer whales (Orcinus orca) in the waters of British Columbia and Washington State have a strong preference for Chinook salmon even in months when Chinook comprise less than about 10% of the salmon population. The foraging behavior of killer whales suggests that they depend on echolocation to detect and recognize their prey. In order to determine possible cues in echoes from salmon species, a series of backscatter measurements were made at the Applied Physics Laboratory (Univ. of Wash.) Facility on Lake Union, on three different salmon species using simulated killer whale echolocation signals. The fish were attached to a monofilament net panel and rotated while echoes were collected, digitized and stored on a laptop computer. Three transducer depths were used; same depth, 22° and 45° above the horizontal plane of the fish. Echoes were collected from five Chinook, three coho and one sockeye salmon. Radiograph images of all specimens were obtained to examine the swimbladder shape and orientation. The results show that echo structure from similar length but different species of salmon were different and probably recognizable by foraging killer whales.

Yangtze finless porpoises (Neophocaena asiaeorientalis) are known to use the narrowband signals for echolocation. In this study, a finite-element model was configured based on computed tomography imaging technique and tissue physical properties measurement to simulate biosonar signal emission and transmission processes through animal’s head. The roles of the main structures in the head such as the air sacs, melon, bony structures, connective tissue, blubber, mandibular fat on the biosonar beam formation were investigated, and the relative importance of these structures was compared. The biosonar beam properties of this neonate porpoise were compared with those of adult ones. The method in this paper suggested an effective way for investigating the acoustic processes in the heads of the neonate odontocetes.

Modern bioacoustic research with aquatic animals began in Hawaii in the 1968–1969 period when three independent programs started almost simultaneously. Two faculty members began studying various facets of hearing in fish at the University of Hawaii. Another faculty member from a different department established a laboratory in which dolphin hearing and echolocation were among some of the topics studied. The Navy started a dolphin facility to train dolphins to perform Navy related tasks. The Navy program also included research on dolphin echolocation including a study comparing the performance of two dolphins and a Straza-500 CTFM sonar in target detection. The Navy facility closed in 1993 but some of the bioacoustics research was transferred to the University’s Hawaii Institute of Marine Biology on Coconut Island about a mile from the former Navy facility in the same bay. The HIMB program expanded into field research with spinner dolphins and humpback whales, and passive acoustic monitoring of marine organisms. The local National Marine Fisheries Service of NOAA became a close partner in the PAM development and studies. This presentation will focus of the bioacoustic research performed with marine mammals highlighting some of the more important findings and contributions to the field.

Spinner dolphins (Stenella longirostris) exhibit different social behaviours at two regions in the Hawaiian Archipelago: off the high volcanic islands in the SE archipelago they form dynamic groups with ever-changing membership, but in the low carbonate atolls in the NW archipelago they form long-term stable groups. To determine whether these environmental and social differences influence population genetic structure, we surveyed spinner dolphins throughout the Hawaiian Archipelago with mtDNA control region sequences and 10 microsatellite loci (n = 505). F-statistics, Bayesian cluster analyses, and assignment tests revealed population genetic separations between most islands, with less genetic structuring among the NW atolls than among the SE high islands. The populations with the most stable social structure (Midway and Kure Atolls) have the highest gene flow between populations (mtDNA Phi(ST) < 0.001, P = 0.357; microsatellite F(ST) = -0.001; P = 0.597), and a population with dynamic groups and fluid social structure (the Kona Coast of the island of Hawai'i) has the lowest gene flow (mtDNA 0.042 < Phi(ST) < 0.236, P < 0.05; microsatellite 0.016 < F(ST) < 0.040, P < 0.001). We suggest that gene flow, dispersal, and social structure are influenced by the availability of habitat and resources at each island. Genetic comparisons to a South Pacific location (n = 16) indicate that Hawaiian populations are genetically depauperate and isolated from other Pacific locations (mtDNA 0.216 < F(ST) < 0.643, P < 0.001; microsatellite 0.058 < F(ST) < 0.090, P < 0.001); this isolation may also influence social and genetic structure within Hawai'i. Our results illustrate that genetic and social structure are flexible traits that can vary between even closely-related populations.

A psychoacoustic behavioral technique was used to determine the critical ratios (CRs) of two harbor porpoises for tonal signals with frequencies between 0.315 and 150 kHz, in random Gaussian white noise. The masked 50% detection hearing thresholds were measured using a "go/no-go" response paradigm and an up-down staircase psychometric method. CRs were determined at one masking noise level for each test frequency and were similar in both animals. For signals between 0.315 and 4 kHz, the CRs were relatively constant at around 18 dB. Between 4 and 150 kHz the CR increased gradually from 18 to 39 dB (∼3.3 dB/octave). Generally harbor porpoises can detect tonal signals in Gaussian white noise slightly better than most odontocetes tested so far. By combining the mean CRs found in the present study with the spectrum level of the background noise levels at sea, the basic audiogram, and the directivity index, the detection threshold levels of harbor porpoises for tonal signals in various sea states can be calculated. © 2009 Acoustical Society of America.

The roles of nocturnal light and lunar phase in the diel migration of micronekton from a nearshore scattering layer were examined. Migration patterns were measured over six complete lunar cycles using moored upwardlooking echosounders while nocturnal surface irradiance was recorded. We hypothesized that animals would remain at a constant isolume at night despite changes in nocturnal illumination between nights. The scattering layer migrated closer to the surface during dark nights than during well-lit ones. However, this movement was not enough to compensate for observed changes in light, and at night animals often remained at light levels higher than they experience at depth during the day. Light and lunar cycle were not completely coupled, allowing separation of the light and lunar phases. Contrary to the initial hypothesis, lunar phase accounted for substantially more of the variability in layer migration than surface irradiance, showing strong effects on the scattering layer’s depth and animal density within the layer. Changes in layer depth and animal density were amplified a small amount by variations in light level but were minimized by the seafloor in shallow areas. The horizontal component of the scattering layer’s migration was also affected by lunar phase, with animals remaining further offshore in deeper waters during nights near and during the full moon, even when these were not the nights with the highest light levels. These results suggest that moonlight may be a cue for an endogenous lunar rhythm in the process of diel migration rather than a direct cause. Diel vertical migration of zooplankton and nekton, active movement from deep, dark waters where animals reside during the day to near-surface waters at night (Longhurst 1976), is thought to be ultimately driven by maximization of survival controlled by food, predators, and physiological costs (Zaret and Suffern 1976; Enright 1977). However, individual animals have limited means to follow predator and prey distributions in the water column. Therefore, environmental cues must be used as proxies. Light is one of the strongest environmental cues available to migrating animals and is hypothesized to serve as a proxy for the risk of predation by visual predators (see a review in Forward 1988). Light is generally accepted as playing an important role in controlling the timing of diel vertical migration since most migrations occur at sunrise and sunset (Esterley 1911; see a review in Ringelberg 1995). Two mechanisms to explain how light may trigger vertical migration have been examined: absolute light intensity and rate of light change. One hypothesis about absolute light levels suggests that animals remain at an optimum light level, following this constant level, or isolume, up and down in the water column as the surface irradiance changes (Michael 1911; Longhurst 1976). Alternatively, animals may use an absolute threshold value of light to trigger upward movement when light becomes lower than the threshold (Esterley 1911; Pearre 1973). The absolute level of light may be difficult for animals to detect, however. Instead, animals may use the rate of the change in light intensity to cue migration (reviewed in Ringelberg 1995).

Noise levels in the ocean are increasing and are expected to affect marine mammals. To examine the auditory effects of noise on odontocetes, a bottlenose dolphin (Tursiops truncatus) was exposed to octave-band noise (4-8 kHz) of varying durations (2-30 min) and sound pressures (130-178 dB re 1 microPa). Temporary threshold shift (TTS) occurrence was quantified in an effort to (i) determine the sound exposure levels (SELs) (dB re 1 microPa(2) s) that induce TTS and (ii) develop a model to predict TTS onset. Hearing thresholds were measured using auditory evoked potentials. If SEL was kept constant, significant shifts were induced by longer duration exposures but not for shorter exposures. Higher SELs were required to induce shifts in shorter duration exposures. The results did not support an equal-energy model to predict TTS onset. Rather, a logarithmic algorithm, which increased in sound energy as exposure duration decreased, was a better predictor of TTS. Recovery to baseline hearing thresholds was also logarithmic (approximately -1.8 dB/doubling of time) but indicated variability including faster recovery rates after greater shifts and longer recoveries necessary after longer duration exposures. The data reflected the complexity of TTS in mammals that should be taken into account when predicting odontocete TTS.

The frequency band that an Atlantic bottlenose dolphin (Tursiops truncatus) used to perform an echolocation target discrimination task was determined using computer simulated phantom targets. The dolphin was trained to discriminate frequency filtered phantom targets from unfiltered ones in a go/no-go paradigm. The dolphin's performance indicated perception of echo alteration only when applied filters interfered with the frequency band between 29 and 42 kHz. The dolphin did not behaviorally convey perception of applied filters that affected frequencies outside this functional bandwidth, such as a low pass 43 kHz or a high pass 28 kHz filter. The upper limit of the functional bandwidth at 42 kHz corresponded with the dolphin's upper hearing limit of 45 kHz, as determined through auditory evoked potential measurements. The lower limit of the functional bandwidth corresponded to a drop in intensity below 30 kHz within the dolphin's echolocation clicks. The randomized presentation of different filters showed that the dolphin paid attention to the entire 29-42 kHz band for each trial, not just subsets. The absence of temporal cues between some of the targets the dolphin could discriminate indicated that in these cases the target discrimination cues were based solely on the frequency content.

Sonar techniques were used to quantitatively observe foraging predators and their prey simultaneously in three dimensions. Spinner dolphins foraged at night in highly coordinated groups of 16–28 individuals using strict four-dimensional patterns to increase prey density by up to 200 times. Herding exploited the prey’s own avoidance behavior to achieve food densities not observed otherwise. Pairs of dolphins then took turns feeding within the aggregation that was created. Using a proxy estimate of feeding success, it is estimated that each dolphin working in concert has more access to prey than it would if feeding individually, despite the costs of participating in the group maneuvers, supporting the cooperation hypothesis. Evidence of a prey density threshold for feeding suggests that feedback from the environment may be enough to favor the evolution of cooperation. The remarkable degree of coordination shown by foraging spinner dolphins, the very strict geometry, tight timing, and orderly turn taking, indicates the advantage conferred by this strategy and the constraints placed upon it. The consistent appearance of this behavior suggests that it may be a critical strategy for energy acquisition by spinner dolphins in energy poor featureless environments in the tropical Pacific Ocean. © 2009 Acoustical Society of America. DOI: 10.1121/1.2967480

A set of dolphin echolocation clicks collected from an Atlantic bottlenose dolphin in Kaneohe Bay, Hawai'i from a previous experiment is examined in terms of their time and frequency characteristics. The center frequency and rms bandwidth are calculated for the clicks and these are clustered into four classes by using a model based on the Bayesian information criterion. The echo signatures are attained from a solid, elastic homogeneous sphere for each class of clicks from an acoustic scattering model. The results from the scattering model are compared to experimental values. The joint time-frequency content of the resulting echo signals is obtained by the reduced interference distribution (RID). The RIDs are plotted and examined for each signal class for four spherical targets of different material compositions. RID correlation values are obtained for a standard target versus comparison targets by using a time-frequency correlator. The results suggest that dolphins may discriminate by auditory inspection of the time-frequency information returned by the targets. The modification of the outgoing clicks and examination of time-frequency target information may be fundamental to a dolphin's ability to identify and discriminate targets.