Madsen Peter T.

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Madsen

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Peter T.

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0000-0002-5208-5259

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Estimated communication range and energetic cost of bottlenose dolphin whistles in a tropical habitat

(Acoustical Society of America, 2012-01) Jensen, Frants H. ; Beedholm, Kristian ; Wahlberg, Magnus ; Bejder, Lars ; Madsen, Peter T.

Bottlenose dolphins (Tursiops sp.) depend on frequency-modulated whistles for many aspects of their social behavior, including group cohesion and recognition of familiar individuals. Vocalization amplitude and frequency influences communication range and may be shaped by many ecological and physiological factors including energetic costs. Here, a calibrated GPS-synchronized hydrophone array was used to record the whistles of bottlenose dolphins in a tropical shallow-water environment with high ambient noise levels. Acoustic localization techniques were used to estimate the source levels and energy content of individual whistles. Bottlenose dolphins produced whistles with mean source levels of 146.7±6.2 dB re. 1 μPa(RMS). These were lower than source levels estimated for a population inhabiting the quieter Moray Firth, indicating that dolphins do not necessarily compensate for the high noise levels found in noisy tropical habitats by increasing their source level. Combined with measured transmission loss and noise levels, these source levels provided estimated median communication ranges of 750 m and maximum communication ranges up to 5740 m. Whistles contained less than 17 mJ of acoustic energy, showing that the energetic cost of whistling is small compared to the high metabolic rate of these aquatic mammals, and unlikely to limit the vocal activity of toothed whales.
Response to: the metabolic cost of whistling is low but measurable in dolphins

(Company of Biologists, 2020-06-08) Pedersen, Michael B. ; Fahlman, Andreas ; Borque-Espinosa, Alicia ; Madsen, Peter T. ; Jensen, Frants H.

Costs of sound production have been investigated only sparsely incetaceans, despite recent efforts to understand how increasinganthropogenic noise affects these animals that rely extensively onsound for communication and foraging. Theoretical estimates suggestthat metabolic costs of whistling for bottlenose dolphins should be<0.54% of resting metabolic rate (RMR) (Jensen et al., 2012),whereas empirical studies of a single whistling dolphin surprisinglyclaimed that sound production costs were around 20% of RMR (Holtet al., 2015; Noren et al., 2013). Addressing this discrepancy, wefound that costs of whistling were significantly less than 20% RMRand not statistically different from theoretical estimates (Pedersenet al., 2020). In their correspondence, Noren et al., 2020 argue thatthey did not claim whistling was‘costly’and questioned aspects ofour methods, and we address these points here.
Echolocation click source parameters of Australian snubfin dolphins (Orcaella heinsohni)

(Acoustical Society of America, 2018-04-30) de Freitas, Mafalda ; Smith, Joshua N. ; Jensen, Frants H. ; Beedholm, Kristian ; Madsen, Peter T.

The Australian snubfin dolphin (Orcaella heinsohni) is endemic to Australian waters, yet little is known about its abundance and habitat use. To investigate the feasibility of Passive Acoustic Monitoring for snubfin dolphins, biosonar clicks were recorded in Cygnet Bay, Australia, using a four-element hydrophone array. Clicks had a mean source level of 200 ± 5 dB re 1 μPa pp, transmission directivity index of 24 dB, mean centroid frequency of 98 ± 9 kHz, and a root-mean-square bandwidth of 31 ± 3 kHz. Such properties lend themselves to passive acoustic monitoring, but are comparable to similarly-sized delphinids, thus requiring additional cues to discriminate between snubfins and sympatric species.
Clicking for calamari : toothed whales can echolocate squid Loligo pealeii

(Inter-Research, 2007-11-27) Madsen, Peter T. ; Wilson, M. ; Johnson, Mark P. ; Hanlon, Roger T. ; Bocconcelli, Alessandro ; Aguilar De Soto, Natacha ; Tyack, Peter L.

Squid play an important role in biomass turnover in marine ecosystems and constitute a food source for ~90% of all echolocating toothed whale species. Nonetheless, it has been hypothesized that the soft bodies of squid provide echoes too weak to be detected by toothed whale biosonars, and that only the few hard parts of the squid body may generate significant backscatter. We measured the acoustic backscatter from the common squid Loligo pealeii for signals similar to toothed whale echolocation clicks using an energy detector to mimic the mammalian auditory system. We show that the dorsal target strengths of L. pealeii with mantle lengths between 23 and 26 cm fall in the range from –38 to –44 dB, and that the pen, beak and lenses do not contribute significantly to the backscatter. Thus, the muscular mantle and fins of L. pealeii constitute a sufficient sonar target for individual biosonar detection by toothed whales at ranges between 25 and 325 m, depending on squid size, noise levels, click source levels, and orientation of the ensonified squid. While epipelagic squid must be fast and muscular to catch prey and avoid visual predators, it is hypothesized that some deep-water squid may have adopted passive acoustic crypsis, with a body of low muscle mass and low metabolism that will render them less conspicuous to echolocating predators.
Sperm whale codas may encode individuality as well as clan identity

(Acoustical Society of America, 2016-05-19) Oliveira, Claudia ; Wahlberg, Magnus ; Silva, Monica A. ; Johnson, Mark P. ; Antunes, Ricardo ; Wisniewska, Danuta M. ; Fais, Andrea ; Goncalves, Joao M. A. ; Madsen, Peter T.

Sperm whales produce codas for communication that can be grouped into different types according to their temporal patterns. Codas have led researchers to propose that sperm whales belong to distinct cultural clans, but it is presently unclear if they also convey individual information. Coda clicks comprise a series of pulses and the delay between pulses is a function of organ size, and therefore body size, and so is one potential source of individual information. Another potential individual-specific parameter could be the inter-click intervals within codas. To test whether these parameters provide reliable individual cues, stereo-hydrophone acoustic tags (Dtags) were attached to five sperm whales of the Azores, recording a total of 802 codas. A discriminant function analysis was used to distinguish 288 5 Regular codas from four of the sperm whales and 183 3 Regular codas from two sperm whales. The results suggest that codas have consistent individual features in their inter-click intervals and inter-pulse intervals which may contribute to individual identification. Additionally, two whales produced different coda types in distinct foraging dive phases. Codas may therefore be used by sperm whales to convey information of identity as well as activity within a social group to a larger extent than previously assumed.
Following a foraging fish-finder : diel habitat use of Blainville's beaked whales revealed by echolocation

(Public Library of Science, 2011-12-07) Arranz, Patricia ; Aguilar De Soto, Natacha ; Madsen, Peter T. ; Brito, Alberto ; Bordes, Fernando ; Johnson, Mark P.

Simultaneous high resolution sampling of predator behavior and habitat characteristics is often difficult to achieve despite its importance in understanding the foraging decisions and habitat use of predators. Here we tap into the biosonar system of Blainville's beaked whales, Mesoplodon densirostris, using sound and orientation recording tags to uncover prey-finding cues available to echolocating predators in the deep-sea. Echolocation sounds indicate where whales search and encounter prey, as well as the altitude of whales above the sea-floor and the density of organisms around them, providing a link between foraging activity and the bio-physical environment. Tagged whales (n = 9) hunted exclusively at depth, investing most of their search time either in the lower part of the deep scattering layer (DSL) or near the sea-floor with little diel change. At least 43% (420/974) of recorded prey-capture attempts were performed within the benthic boundary layer despite a wide range of dive depths, and many dives included both meso- and bentho-pelagic foraging. Blainville's beaked whales only initiate searching when already deep in the descent and encounter prey suitable for capture within 2 min of the start of echolocation, suggesting that these whales are accessing prey in reliable vertical strata. Moreover, these prey resources are sufficiently dense to feed the animals in what is effectively four hours of hunting per day enabling a strategy in which long dives to exploit numerous deep-prey with low nutritional value require protracted recovery periods (average 1.5 h) between dives. This apparent searching efficiency maybe aided by inhabiting steep undersea slopes with access to both the DSL and the sea-floor over small spatial scales. Aggregations of prey in these biotopes are located using biosonar-derived landmarks and represent stable and abundant resources for Blainville's beaked whales in the otherwise food-limited deep-ocean.
Vessel noise effects on delphinid communication

(Inter-Research, 2009-12-03) Jensen, Frants H. ; Bejder, Lars ; Wahlberg, Magnus ; Aguilar De Soto, Natacha ; Johnson, Mark P. ; Madsen, Peter T.

Increasing numbers and speeds of vessels in areas with populations of cetaceans may have the cumulative effect of reducing habitat quality by increasing the underwater noise level. Here, we first use digital acoustic tags to demonstrate that free-ranging delphinids in a coastal deep-water habitat are subjected to varying and occasionally intense levels of vessel noise. Vessel noise and sound propagation measurements from a shallow-water habitat are then used to model the potential impact of high sound levels from small vessels on delphinid communication in both shallow and deep habitats, with bottlenose dolphins Tursiops sp. and short-finned pilot whales Globicephala macrorhynchus as model organisms. We find that small vessels travelling at 5 knots in shallow water can reduce the communication range of bottlenose dolphins within 50 m by 26%. Pilot whales in a quieter deep-water habitat could suffer a reduction in their communication range of 58% caused by a vessel at similar range and speed. Increased cavitation noise at higher speeds drastically increases the impact on the communication range. Gear shifts generate high-level transient sounds (peak– peak source levels of up to 200 dB re 1 µPa) that may be audible over many kilometres and may disturb close-range animals. We conclude that noise from small vessels can significantly mask acoustically mediated communication in delphinids and contribute to the documented negative impacts on animal fitness.
Potential for sound sensitivity in cephalopods

( 2010-07) Mooney, T. Aran ; Hanlon, Roger T. ; Madsen, Peter T. ; Christensen-Dalsgaard, Jakob ; Ketten, Darlene R. ; Nachtigall, Paul E.

Hearing is a primary sense in many marine animals and we now have a reasonable understanding of what stimuli generate clear responses, the frequency range of sensitivity, expected threshold values and mecha-nisms of sound detection for several species of marine mammals and fishes (Fay 1988; Au et al. 2000). For marine invertebrates, our knowledge of hearing capabilities is relatively poor and a definition or even certainty of sound detection is not agreed upon (Webster et al. 1992) despite their magnitude of biomass and often central role in ocean ecosystems. Cephalopods (squid, cuttlefish, octopods and nautilus) are particularly interesting subjects for inver-tebrate sound detection investigations for several reasons. Ecologically, they occupy many of the same niches as sound-sensitive fish (Budelmann 1994) and may benefit from sound perception and use for the same reasons, such as to detect predators, navigate, or locate conspecifics. Squid, for example, are often the prey of loud, echolocating marine mammals (Clarke 1996), and may therefore be expected to have evolved hearing to avoid predators. Anatomically, squid have complex statocysts that are considered to serve primarily as vestibular and acceleration detectors (Nixon and Young 2003). However, statocysts may also be analogs for fish otolithic organs, detecting acoustic stimuli (Budelmann 1992). Previous studies have debated the subject of squid hearing and recently there has been a revival of research on the subject. Here, we briefly review what is known about squid sound detection, revisit hearing definitions, discuss potential squid susceptibility to anthropogenic noise and suggest potential future research direc-tions to examine squid acoustic sensitivity.
Passive acoustic detection of deep-diving beaked whales

(Acoustical Society of America, 2008-11) Zimmer, Walter M. X. ; Harwood, John ; Tyack, Peter L. ; Johnson, Mark P. ; Madsen, Peter T.

Beaked whales can remain submerged for an hour or more and are difficult to sight when they come to the surface to breathe. Passive acoustic detection (PAD) not only complements traditional visual-based methods for detecting these species but also can be more effective because beaked whales produce clicks regularly to echolocate on prey during deep foraging dives. The effectiveness of PAD for beaked whales depends not only on the acoustic behavior and output of the animals but also on environmental conditions and the quality of the passive sonar implemented. A primary constraint on the range at which beaked whale clicks can be detected involves their high frequencies, which attenuate rapidly, resulting in limited ranges of detection, especially in adverse environmental conditions. Given current knowledge of source parameters and in good conditions, for example, with a wind speed of 2 m/s, a receiver close to the surface should be able to detect acoustically Cuvier's beaked whales with a high probability at distances up to 0.7 km, provided the listening duration exceeds the deep dive interval, about 2.5 h on average. Detection ranges beyond 4 km are unlikely and would require low ambient noise or special sound propagation conditions.
Beaked whales echolocate on prey

(Royal Society, 2004-12-07) Johnson, Mark P. ; Madsen, Peter T. ; Zimmer, Walter M. X. ; Aguilar De Soto, Natacha ; Tyack, Peter L.

Beaked whales (Cetacea: Ziphiidea) of the genera Ziphius and Mesoplodon are so difficult to study that they are mostly known from strandings. How these elusive toothed whales use and react to sound is of concern because they mass strand during naval sonar exercises. A new non-invasive acoustic recording tag was attached to four beaked whales (two Mesoplodon densirostris and two Ziphius cavirostris) and recorded high-frequency clicks during deep dives. The tagged whales only clicked at depths below 200 m, down to a maximum depth of 1267 m. Both species produced a large number of short, directional, ultrasonic clicks with no significant energy below 20 kHz. The tags recorded echoes from prey items; to our knowledge, a first for any animal echolocating in the wild. As far as we are aware, these echoes provide the first direct evidence on how free-ranging toothed whales use echolocation in foraging. The strength of these echoes suggests that the source level of Mesoplodon clicks is in the range of 200-220 dB re 1 μPa at 1 m. This paper presents conclusive data on the normal vocalizations of these beaked whale species, which may enable acoustic monitoring to mitigate exposure to sounds intense enough to harm them.
Classification of broadband echoes from prey of a foraging Blainville's beaked whale

(Acoustical Society of America, 2008-03) Jones, Benjamin A. ; Stanton, Timothy K. ; Lavery, Andone C. ; Johnson, Mark P. ; Madsen, Peter T. ; Tyack, Peter L.

Blainville's beaked whales (Mesoplodon densirostris) use broadband, ultrasonic echolocation signals with a −10 dB bandwidth from 26 to 51 kHz to search for, localize, and approach prey that generally consist of mid-water and deep-water fishes and squid. Although it is well known that the spectral characteristics of broadband echoes from marine organisms vary as a function of size, shape, orientation, and anatomical group, there is little evidence as to whether or not free-ranging toothed whales use spectral cues in discriminating between prey and nonprey. In order to study the prey-classification process, a stereo acoustic tag was deployed on a Blainville's beaked whale so that emitted clicks and the corresponding echoes from targets in the water could be recorded. A comparison of echoes from targets apparently selected by the whale and those from a sample of scatterers that were not selected suggests that spectral features of the echoes, target strengths, or both may have been used by the whale to discriminate between echoes. Specifically, the whale appears to favor targets with one or more nulls in the echo spectra and to seek prey with higher target strengths at deeper depths.
Three-dimensional beam pattern of regular sperm whale clicks confirms bent-horn hypothesis

(Acoustical Society of America, 2005-03) Zimmer, Walter M. X. ; Tyack, Peter L. ; Johnson, Mark P. ; Madsen, Peter T.

The three-dimensional beam pattern of a sperm whale (Physeter macrocephalus) tagged in the Ligurian Sea was derived using data on regular clicks from the tag and from hydrophones towed behind a ship circling the tagged whale. The tag defined the orientation of the whale, while sightings and beamformer data were used to locate the whale with respect to the ship. The existence of a narrow, forward-directed P1 beam with source levels exceeding 210 dBpeak re: 1 µPa at 1 m is confirmed. A modeled forward-beam pattern, that matches clicks >20° off-axis, predicts a directivity index of 26.7 dB and source levels of up to 229 dBpeak re: 1 µPa at 1 m. A broader backward-directed beam is produced by the P0 pulse with source levels near 200 dBpeak re: 1 µPa at 1 m and a directivity index of 7.4 dB. A low-frequency component with source levels near 190 dBpeak re: 1 µPa at 1 m is generated at the onset of the P0 pulse by air resonance. The results support the bent-horn model of sound production in sperm whales. While the sperm whale nose appears primarily adapted to produce an intense forward-directed sonar signal, less-directional click components convey information to conspecifics, and give rise to echoes from the seafloor and the surface, which may be useful for orientation during dives.
Whistling is metabolically cheap for communicating bottlenose dolphins (Tursiops truncatus)

(Company of Biologists, 2019-12-03) Pedersen, Michael B. ; Fahlman, Andreas ; Borque-Espinosa, Alicia ; Madsen, Peter T. ; Jensen, Frants H.

Toothed whales depend on sound for communication and foraging, making them potentially vulnerable to acoustic masking from increasing anthropogenic noise. Masking effects may be ameliorated by higher amplitudes or rates of calling, but such acoustic compensation mechanisms may incur energetic costs if sound production is expensive. The costs of whistling in bottlenose dolphins (Tursiops truncatus) have been reported to be much higher (20% of resting metabolic rate, RMR) than theoretical predictions (0.5–1% of RMR). Here, we address this dichotomy by measuring the change in the resting O2 consumption rate (V̇O2), a proxy for RMR, in three post-absorptive bottlenose dolphins during whistling and silent trials, concurrent with simultaneous measurement of acoustic output using a calibrated hydrophone array. The experimental protocol consisted of a 2-min baseline period to establish RMR, followed by a 2-min voluntary resting surface apnea, with or without whistling as cued by the trainers, and then a 5-min resting period to measure recovery costs. Daily fluctuations in V̇O2 were accounted for by subtracting the baseline RMR from the recovery costs to estimate the cost of apnea with and without whistles relative to RMR. Analysis of 52 sessions containing 1162 whistles showed that whistling did not increase metabolic cost (P>0.1, +4.2±6.9%) as compared with control trials (−0.5±5.9%; means±s.e.m.). Thus, we reject the hypothesis that whistling is costly for bottlenose dolphins, and conclude that vocal adjustments such as the Lombard response to noise do not represent large direct energetic costs for communicating toothed whales.
Single-click beam patterns suggest dynamic changes to the field of view of echolocating Atlantic spotted dolphins (Stenella frontalis) in the wild

( 2015-02) Jensen, Frants H. ; Wahlberg, Magnus ; Beedholm, Kristian ; Johnson, Mark P. ; Aguilar De Soto, Natacha ; Madsen, Peter T.

Echolocating animals exercise an extensive control over the spectral and temporal properties of their biosonar signals to facilitate perception of their actively generated auditory scene when homing in on prey. The intensity and directionality of the biosonar beam defines the field of view of echolocating animals by affecting the acoustic detection range and angular coverage. However, the spatial relationship between an echolocating predator and its prey changes rapidly, resulting in different biosonar requirements throughout prey pursuit and capture. Here we measured single click beam patterns using a parametric fit procedure to test whether free-ranging Atlantic spotted dolphins (Stenella frontalis) modify their biosonar beamwidth. We recorded echolocation clicks using a linear array of receivers and estimated the beamwidth of individual clicks using a parametric spectral fit, cross-validated with well-established composite beam pattern estimates. The dolphins apparently increased the biosonar beamwidth, to a large degree without changing the signal frequency, when they approached the recording array. This is comparable to bats that also expand their field of view during prey capture, but achieve this by decreasing biosonar frequency. This behaviour may serve to decrease the risk that rapid escape movements of prey take them outside the biosonar beam of the predator. It is likely that shared sensory requirements have resulted in bats and toothed whales expanding their acoustic field of view at close range to increase the likelihood of successfully acquiring prey using echolocation, representing a case of convergent evolution of echolocation behaviour between these two taxa.
Off-axis effects on the multipulse structure of sperm whale usual clicks with implications for sound production

(Acoustical Society of America, 2005-11) Zimmer, Walter M. X. ; Madsen, Peter T. ; Teloni, Valeria ; Johnson, Mark P. ; Tyack, Peter L.

Sperm whales (Physeter macrocephalus) produce multipulsed clicks with their hypertrophied nasal complex. The currently accepted view of the sound generation process is based on the click structure measured directly in front of, or behind, the whale where regular interpulse intervals (IPIs) are found between successive pulses in the click. Most sperm whales, however, are recorded with the whale in an unknown orientation with respect to the hydrophone where the multipulse structure and the IPI do not conform to a regular pulse pattern. By combining far-field recordings of usual clicks with acoustic and orientation information measured by a tag on the clicking whale, we analyzed clicks from known aspects to the whale. We show that a geometric model based on the bent horn theory for sound production can explain the varying off-axis multipulse structure. Some of the sound energy that is reflected off the frontal sac radiates directly into the water creating an intermediate pulse p1/2 seen in off-axis recordings. The powerful p1 sonar pulse exits the front of the junk as predicted by the bent-horn model, showing that the junk of the sperm whale nasal complex is both anatomically and functionally homologous to the melon of smaller toothed whales.
Narrow acoustic field of view drives frequency scaling in toothed whale biosonar

( 2018-10) Jensen, Frants H. ; Johnson, Mark P. ; Ladegaard, Michael ; Wisniewska, Danuta M. ; Madsen, Peter T.

Toothed whales are apex predators varying in size from 40-kg porpoises to 50-ton sperm whales that all forage by emitting high-amplitude ultrasonic clicks and listening for weak returning echoes [1, 2]. The sensory field of view of these echolocating animals depends on the characteristics of the biosonar signals and the morphology of the sound generator, yet it is poorly understood how these biophysical relationships have shaped evolution of biosonar parameters as toothed whales adapted to different foraging niches. Here we test how biosonar output, frequency, and directivity vary with body size to understand the co-evolution of biosonar signals and sound-generating structures. We show that the radiated power increases twice as steeply with body mass (P ∝ M1.47±0.25) than expected from typical scaling laws of call intensity [3], indicating hyperallometric investment into sound production structures. This is likely driven by a strong selective pressure for long-range biosonar in larger oceanic or deep-diving species to search efficiently for patchy prey. We find that biosonar frequency scales inversely with body size (F∝ M-0.19±0.03), resulting in remarkably stable biosonar beamwidth that is independent of body size. We discuss how frequency scaling in toothed whales cannot be explained by the three main hypotheses for inverse scaling of frequency in animal communication [3-5]. We propose that a narrow acoustic field of view, analogous to the fovea of many visual predators, is the primary evolutionary driver of biosonar frequency in toothed whales, serving as a spatial filter to reduce clutter levels and facilitate long-range prey detection.
Echolocation clicks of free-ranging Cuvier's beaked whales (Ziphius cavirostris)

(Acoustical Society of America, 2005-06) Zimmer, Walter M. X. ; Johnson, Mark P. ; Madsen, Peter T. ; Tyack, Peter L.

Strandings of beaked whales of the genera Ziphius and Mesoplodon have been reported to occur in conjunction with naval sonar use. Detection of the sounds from these elusive whales could reduce the risk of exposure, but descriptions of their vocalizations are at best incomplete. This paper reports quantitative characteristics of clicks from deep-diving Cuvier's beaked whales (Ziphius cavirostris) using a unique data set. Two whales in the Ligurian Sea were simultaneously tagged with sound and orientation recording tags, and the dive tracks were reconstructed allowing for derivation of the range and relative aspect between the clicking whales. At depth, the whales produced trains of regular echolocation clicks with mean interclick intervals of 0.43 s (±0.09) and 0.40 s (±0.07). The clicks are frequency modulated pulses with durations of ~200 µs and center frequencies around 42 kHz, –10 dB bandwidths of 22 kHz, and Q3 dB of 4. The sound beam is narrow with an estimated directionality index of more than 25 dB, source levels up to 214 dBpp re: 1 µPa at 1 m, and energy flux density of 164 dB re: 1 µPa2 s. As the spectral and temporal properties are different from those of nonziphiid odontocetes the potential for passive detection is enhanced.
Wind turbine underwater noise and marine mammals : implications of current knowledge and data needs

(Inter-Research, 2006-03-15) Madsen, Peter T. ; Wahlberg, Magnus ; Tougaard, Jakob ; Lucke, Klaus ; Tyack, Peter L.

The demand for renewable energy has led to construction of offshore wind farms with high-power turbines, and many more wind farms are being planned for the shallow waters of the world’s marine habitats. The growth of offshore wind farms has raised concerns about their impact on the marine environment. Marine mammals use sound for foraging, orientation and communication and are therefore possibly susceptible to negative effects of man-made noise generated from constructing and operating large offshore wind turbines. This paper reviews the existing literature and assesses zones of impact from different noise-generating activities in conjunction with wind farms on 4 representative shallow-water species of marine mammals. Construction involves many types of activities that can generate high sound pressure levels, and pile-driving seems to be the noisiest of all. Both the literature and modeling show that pile-driving and other activities that generate intense impulses during construction are likely to disrupt the behavior of marine mammals at ranges of many kilometers, and that these activities have the potential to induce hearing impairment at close range. The reported noise levels from operating wind turbines are low, and are unlikely to impair hearing in marine mammals. The impact zones for marine mammals from operating wind turbines depend on the low-frequency hearing-abilities of the species in question, on sound-propagation conditions, and on the presence of other noise sources such as shipping. The noise impact on marine mammals is more severe during the construction of wind farms than during their operation.
Recording and quantification of ultrasonic echolocation clicks from free-ranging toothed whales

( 2006-10-08) Madsen, Peter T. ; Wahlberg, Magnus

Toothed whales produce short, ultrasonic clicks of high directionality and source level to probe their environment acoustically. This process, termed echolocation, is to a large part governed by the properties of the emitted clicks. Therefore derivation of click source parameters from free-ranging animals is of increasing importance to understand both how toothed whales use echolocation in the wild and how they may be monitored acoustically. This paper addresses how source parameters can be derived from free-ranging toothed whales in the wild using calibrated multi-hydrophone arrays and digital recorders. We outline the properties required of hydrophones, amplifiers and analog to digital converters, and discuss the problems of recording echolocation clicks on the axis of a directional sound beam. For accurate localization the hydrophone array apertures must be adapted and scaled to the behavior of, and the range to, the clicking animal, and precise information on hydrophone locations is critical. We provide examples of localization routines and outline sources of error that lead to uncertainties in localizing clicking animals in time and space. Furthermore we explore approaches to time series analysis of discrete versions of toothed whale clicks that are meaningful in a biosonar context.
Studying the behaviour and sensory ecology of marine mammals using acoustic recording tags : a review

(Inter-Research, 2009-12-03) Johnson, Mark P. ; Aguilar De Soto, Natacha ; Madsen, Peter T.

Many marine animals use sound passively or actively for communication, foraging, predator avoidance, navigation, and to sense their environment. The advent of acoustic recording tags has allowed biologists to get the on-animal perspective of the sonic environment and, in combination with movement sensors, to relate sounds to the activities of the tagged animal. These powerful tools have led to a wide range of insights into the behaviour of marine animals and have opened new opportunities for studying the ways they interact with their environment. Acoustic tags demand new analysis methods and careful experimental design to optimize the consistency between research objectives and the realistic performance of the tags. Technical details to consider are the suitability of the tag attachment to a given species, the accuracy of the tag sensors, and the recording and attachment duration of the tag. Here we consider the achievements, potential, and limitations of acoustic recording tags in studying the behaviour, habitat use and sensory ecology of marine mammals, the taxon to which this technology has been most often applied. We examine the application of acoustic tags to studies of vocal behaviour, foraging ecology, acoustic tracking, and the effects of noise to assess both the breadth of applications and the specific issues that arise in each.

Madsen Peter T.

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