Hearing Abilities of Baleen Whales

Defence R&D Canada

Hearing Abilities of Baleen Whales

Christine Erbe

128 Central Avenue, Unit 2Indooroopilly, Queensland 4068Australia

Contract Number: W7707-01-0828

Contract Scientific Authority: J. A. Theriault, Group Leader/TIAPS Data SystemsPhone (902) 426-3100 extension 376

Contractor Report

DRDC Atlantic CR 2002-065

October 2002

Copy No.________

Defence Research andDevelopment Canada

Recherche et développementpour la défense Canada

Copy No:__________

Hearing Abilities of Baleen Whales

Christine Erbe128 Central Avenue, Unit 2Indooroopilly, Queensland 4068Australia

Contract Number: W7707-01-0828

Contract Scientific Authority: J. A. Theriault, Group Leader/TIAPS Data SystemsPhone (902) 426-3100 extension 376

The scientific or technical validity of this Contract Report is entirely the responsibility ofthe contractor and the contents do not necessarily have the approval or endorsement of

Defence R&D Canada.

Defence R&D Canada – Atlantic

Contractor Report

DRDC Atlantic CR 2002-065

October 2002

DRDC Atlantic CR 2002-065 i

Abstract

Defence R&D Canada – Atlantic has an ongoing research program that requires the transmissionof acoustic energy in an undersea environment. Though the transmissions are generally at arelatively low level, every effort must be made to mitigate the potential for impact on marine life.

The hearing ability of toothed whales has been well studied and has been well publishedcompared to the ability of baleen whales. A literature review related to the hearing ability of,and potential for adverse impact on baleen whales was the focus of a contracted researchactivity. This report summarizes the results of that review.

Résumé

R et D pour la défense du Canada – Atlantique mène un programme permanent qui nécessitel'émission d'énergie acoustique dans un milieu sous-marin. Bien que les émissions soientgénéralement de niveau relativement faible, tous les efforts doivent être faits pour en atténuer leseffets possibles sur la vie marine.

La capacité auditive des baleines à dents a été bien étudiée et documentée par rapport à celle desbaleines à fanons. Un examen de la documentation relative à la capacité auditive des baleines àfanons, et des risques d'effets négatifs sur celles-ci, a fait l'objet d'une activité de recherche àcontrat. Le rapport résume les résultats de cet examen.

ii DRDC Atlantic CR 2002-065

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DRDC Atlantic CR 2002-065 iii

Executive summary

The Canadian Navy has a continuing requirement to acoustically detect both surface vessels andsubmarines. The Defence Research and Development Canada (DRDC) through the DRDCAtlantic has embarked on the first phase of development of the next generation of operationalsonars for the Canadian Navy: the Towed Integrated Active Passive Sonar (TIAPS). The TIAPSTechnology Demonstrator (TD) will demonstrate basic sonar capability and will serve as atestbed for concept development.

The TIAPS TD project will integrate both active and passive sonars in a single system. Bylowering the frequency from traditional hull-mounted sonar frequencies and by giving the sonara variable-depth capability, longer-range active detections can be achieved. Passive sonars havethe advantage of being covert and allowing for the possibility of positively identifying a class oftarget by its acoustic signature. Not only will both active and passive capabilities be included inTIAPS, but a high priority is placed on combining their capabilities.

Like other research areas at DRDC Atlantic, a number of sea trials are required in developingsuch a system. In order to minimize potential impact on marine mammals, impact mitigationprocedures are exercised during each trial. Mitigation efforts tend to evolve with time, asknowledge of potential impact is better understood.

The hearing ability of toothed whales has been well studied and has been well publishedcompared to the ability of baleen whales. A literature review related to the hearing ability of,and potential for adverse impact on baleen whales was the focus of a contracted researchactivity. This report summarizes the results of that review and as such adds to the understandingof potential impact.

Erbe, C. 2002 Hearing Abilities of Baleen Whales, DRDC Atlantic CR 2002-065

iv DRDC Atlantic CR 2002-065

Sommaire

La Marine canadienne a des besoins permanents en matière de détection acoustique de bâtimentsde surface et de sous-marins. R et D pour la défense du Canada (RDDC), par l'intermédiaire deRDDC Atlantique, a entrepris la première phase du développement de la prochaine génération desonars destinés aux opérations de la Marine canadienne : le sonar actif-passif intégré remorqué(TIAPS). Le démonstrateur technologique (TD) TIAPS fera la démonstration de la capacité debase du sonar et servira de banc d'essai pour le développement du concept.

Le projet TD TIAPS intégrera le sonar actif et le sonar passif en un système unique. L'utilisationde fréquences inférieures à celles des sonars de coque traditionnels et une capacité d'immersionvariable permettront d'accroître la portée de la détection active. Les sonars passifs ont l'avantaged'être discrets et de permettre la reconnaissance positive d'une classe de cibles à sa signatureacoustique. Non seulement le TIAPS exploitera-t-il les capacités actives et passives, maisl'intégration de ces capacités aura une priorité élevée.

Comme dans le cas d'autres recherches de RDDC Atlantique, un certain nombre d'essais en mersont nécessaires pour élaborer ce système. Afin de réduire le plus possible les potentiel effets surles mammifères marins, des procédures d'atténuation de ces effets sont mises en œuvre pourchaque essai. Les efforts d'atténuation tendent à évoluer avec le temps, à mesure que les effetséventuels sont mieux connus.

La capacité auditive des baleines à dents a été bien étudiée et documentée par rapport à celle desbaleines à fanons. Un examen de la documentation relative à la capacité auditive des baleines àfanons, et des risques d'effets négatifs sur celles-ci, a fait l'objet d'une activité de recherche àcontrat. Le rapport résume les résultats de cet examen et contribue ainsi à la compréhension desrépercussions possibles.

Erbe, C. 2002 Capacités auditives des baleines à fanons, RDDC AtlantiqueCR 2002-065

DRDC Atlantic CR 2002-065 v

Table of Contents

Abstract............................................................................................................................iResumé ............................................................................................................................iExecutive Summary .......................................................................................................iiiSommaire....................................................................................................................... ivTable of Contents ............................................................................................................vObjective.........................................................................................................................11. Introduction.................................................................................................................12. Baleen Whale Species .................................................................................................2

2.1 Species ..................................................................................................................2Right Whales...........................................................................................................2Family Balaenidae...................................................................................................2Pygmy Right Whale ................................................................................................2Family Neobalaenidae .............................................................................................2Rorquals ..................................................................................................................2Family Balaenopteridae ...........................................................................................2Grey Whale .............................................................................................................2Family Eschrichtiidae ..............................................................................................2

2.2 Distribution ...........................................................................................................23. Vocalizations...............................................................................................................2

Right Whales, Family Balaenidae................................................................................3Bowhead Whale, Balaena mysticetus.......................................................................3Northern Right Whale, Eubalaena glacialis..............................................................3Southern Right Whale, Eubalaena australis..............................................................3

Pygmy Right Whale, Family Neobalaenidae................................................................4Pygmy Right Whale, Caperea marginata..................................................................4

Rorquals, Family Balaenopteridae ...............................................................................4Blue Whale, Balaenoptera musculus........................................................................4Fin Whale, Balaenoptera physalus ...........................................................................5Sei Whale, Balaenoptera borealis.............................................................................5Bryde's Whale, Balaenoptera edeni..........................................................................6Minke Whale, Balaenoptera acutorostrata................................................................6Humpback Whale, Megaptera novaeangliae ............................................................6

Grey Whale, Family Eschrichtiidae .............................................................................7Grey Whale, Eschrichtius robustus ..........................................................................7

3.1 Echolocation..........................................................................................................74. Inner Ear Anatomy ......................................................................................................85. Behavioral Reaction to Underwater Sound ..................................................................8

Right Whales, Family Balaenidae................................................................................9Bowhead Whale, Balaena mysticetus.......................................................................9Northern Right Whale, Eubalaena glacialis..............................................................9Southern Right Whale, Eubalaena australis............................................................ 10

Pygmy Right Whale, Family Neobalaenidae.............................................................. 10

vi DRDC Atlantic CR 2002-065

Pygmy Right Whale, Caperea marginata................................................................ 10Rorquals, Family Balaenopteridae ............................................................................. 10

Blue Whale, Balaenoptera musculus...................................................................... 10Fin Whale, Balaenoptera physalus ......................................................................... 10Sei Whale, Balaenoptera borealis........................................................................... 10Bryde's Whale, Balaenoptera edeni........................................................................ 10Minke Whale, Balaenoptera acutorostrata.............................................................. 10Humpback Whale, Megaptera novaeangliae .......................................................... 11

Grey Whale, Family Eschrichtiidae ........................................................................... 12Grey Whale, Eschrichtius robustus ........................................................................ 12

6. Physiological Damage ............................................................................................... 127. Constructing an Audiogram....................................................................................... 13

Right Whales, Family Balaenidae.............................................................................. 14Bowhead Whale, Balaena mysticetus..................................................................... 14Northern Right Whale, Eubalaena glacialis............................................................ 14Southern Right Whale, Eubalaena australis............................................................ 14

Pygmy Right Whale, Family Neobalaenidae.............................................................. 15Pygmy Right Whale, Caperea marginata................................................................ 15

Rorquals, Family Balaenopteridae ............................................................................. 15Blue Whale, Balaenoptera musculus...................................................................... 15Fin Whale, Balaenoptera physalus ......................................................................... 15Sei Whale, Balaenoptera borealis........................................................................... 15Bryde's Whale, Balaenoptera edeni........................................................................ 15Minke Whale, Balaenoptera acutorostrata.............................................................. 15Humpback Whale, Megaptera novaeangliae .......................................................... 15

Grey Whale, Family Eschrichtiidae ........................................................................... 16Grey Whale, Eschrichtius robustus ........................................................................ 16

7.1 Estimating Acoustic Impact ................................................................................. 178. Conclusion ................................................................................................................ 18References .................................................................................................................... 19

DRDC Atlantic CR 2002-065 1

Objective

The objective of the current study is to derive an audiogram (a hearing curve) for large(baleen) whales, which can be used to determine potential bio-acoustic effects of DRDCAtlantic's marine acoustics program. The study is mainly based on a literature survey of baleenwhale audiology with subsequent derivations and conclusions.

1. Introduction

An audiogram is a function of auditory detection threshold versus frequency. Knowledgeof baleen whale audiograms is important to DRDC Atlantic, as DRDC Atlantic's marineactivities involve acoustic transmissions in the ocean, which are likely audible to baleen whalesand thus might affect them. With a baleen whale audiogram, ranges over which underwatersound is detectable by baleen whales and over which it interferes with their acousticcommunication and navigation can be estimated.

Baleen whales do not exist in captivity and have therefore been inaccessible forbehavioural audiogram measurements (involving training) or electrophysiological audiograms(measuring auditory evoked potentials AEP). A US military group is developing a portableelectrophysiological system that should ultimately be able to measure AEP audiograms on live,stranded or entangled animals within a few minutes. However, technical problems have causedthe idea to fail, and no audiogram has been recorded successfully. A gray whale calf, called JJ,spent a couple of months at SeaWorld San Diego. Audiogram measurements were attempted butwere unsuccessful due to logistic and technical difficulties (Ridgway and Carder (2001)).

As there is no direct data on baleen whale audiograms, hearing sensitivity has to beinferred indirectly. There are three ways to do this.

1) All animals can hear their own vocalizations and often the frequency bandwidth ofvocalizations overlaps with the frequency range of best hearing sensitivity. Therefore, a study ofbaleen whale vocalizations can yield an indication of the frequency range of best hearing.

2) A few dissections and subsequent anatomical studies of baleen ears taken from dead,stranded animals have been done. Such studies yield a relative audiogram, indicating thefrequency range of best hearing and relatively poorer hearing. Anatomical studies don't giveabsolute hearing thresholds.

3) The literature on observed behavioural reactions of baleen whales in the wild tobiological and industrial sounds is vast. Obviously, animals hear the sounds they react to. Thesestudies give absolute suprathresholds of hearing. Animals might not react to a sound that is justaudible, but only react to a sound that is a certain level louder. Reaction thresholds will dependon the current behavioural state of the animal; its previous experience with the particular (similaror other) sound (habituation versus sensitization); age, gender and health of the animal; groupcomposition (groups with calves appear more responsive); habitat and geographic location (e.g.close to shore vs offshore); season and time of day.

2 DRDC Atlantic 2002-065

2. Baleen Whale Species

Baleen whales belong to the Order Cetacea, Suborder Mysticeti. The following tablesummarizes the documented species to-date.

2.1 Species

Right Whales Family Balaenidae

Bowhead Whale Balaena mysticetus

Northern Right Whale Eubalaena glacialis

Southern Right Whale Eubalaena australis

Pygmy Right Whale Family Neobalaenidae

Pygmy Right Whale Caperea marginata

Rorquals Family Balaenopteridae

Blue Whale Balaenoptera musculus

Fin Whale Balaenoptera physalus

Sei Whale Balaenoptera borealis

Bryde's Whale Balaenoptera edeni

Minke Whale Balaenoptera acutorostrata

Humpback Whale Megaptera novaeangliae

Grey Whale Family Eschrichtiidae

Grey Whale Eschrichtius robustus

2.2 DistributionA good reference for a general distribution of the above species in the world's oceans is

Cawardine (1995).

3. Vocalizations

The following tables summarize the information available in the literature on the variousbaleen whale vocalizations.


Right Whales, Family Balaenidae

Bowhead Whale, Balaena mysticetus

Type Bandwidth∆∆∆∆f [Hz]

Dominantf [Hz]

Duration[s]

SL [dB re1µµµµPa @1m]

References

tonal FMmodulated

25-1200 100-400 0.4-3.8 129-178(C&H);156dB@100-150m ≈ 190dB@1m (C&J)

Clark and Johnson (1984);Wursig and Clark (1993);Ljungblad et al. (1982);Wursig et al. (1985); Clark etal. (1986); Cummings andHolliday (1987)

song 20-5000 ≈500 1min -hours

158-189(C&H)

Ljungblad et al. (1982);Cummings and Holliday(1987); Wursig and Clark(1993)

pulsive 25-3500 0.3-7.2(C&J)

152-185 Clark and Johnson (1984);Wursig et al. (1985);Cummings and Holliday(1987)

Northern Right Whale, Eubalaena glacialis

Type ∆∆∆∆f [Hz] Dom. f[Hz]

Dur. [s] SL [dBre 1µµµµPa@1m]

References

moans <400 100 0.5-3 Watkins and Schevill (1972),(1976); Thompson et al. (1979);Spero (1981)

Southern Right Whale, Eubalaena australis



References

tonal, FM,AM

30-1250 50-200 0.2-4.1 Payne and Payne (1971);Cummings et al. (1972) ;Clark (1982, 1983)

pulsive 30-2200 50-500 0.9-2.2 172-187 Cummings et al. (1972); Clark(1982, 1983); Wursig et al. (1982)


Pygmy Right Whale, Family Neobalaenidae

Pygmy Right Whale, Caperea marginata



References

thumps 60-135 0.1-0.3 159-173 Dawbin and Cato (1992)

Rorquals, Family Balaenopteridae

Blue Whale, Balaenoptera musculus


Dur.[s]

SL [dBre 1µµµµPa@1m]

References

110-25 0.5-3.1;1

- Thompson et al. (1996); Ljungbladet al. (1997); Norris et al. (2001)

tonal, FM

20-18; 56-32

17-22;8

Stafford et al. (2001)

doublets"A" and"B"

A: 16-24

B: 22-17

13-19each

185-190(T et al);195 @17Hz (Aet al)

Cummings and Thompson (1994);Rivers (1997); McDonald et al.(1995); Thompson et al. (1996);Thompson and Friedl (1982);Stafford et al. (1998, 1999, 2001);Aburto et al. (1997)

11-500 200 0.4-3 Ljungblad et al. (1997)moans(triplets,C&T;single, E)

12-400;20-60(C&T)

25-16;20-17

16-36(C&T);15-18;

188@14-222Hz(C&T)

Cummings and Thompson (1971);Edds (1982); Stafford et al. (1994);Hildebrand et al. (2001)

clicks 4000-10000;21000-31000

6000-8000;25000

127-132;159

Beamish and Mitchell (1971);Beamish (1979)


Fin Whale, Balaenoptera physalus

Type ∆∆∆∆f [Hz] Dom.f [Hz]

Dur.[s]

SL[dB re1µµµµPa@1m]

References

42-18 20 0.7-1 159-186;140;200

Watkins (1981); Watkins et al. (1987); Payneand Webb (1971); Thompson et al. (1992);Patterson and Hamilton (1964); Northrop et al.(1968); Cummings and Thompson (1994);Edds (1988); McDonald et al. (1995); Charif etal. (2002)

tonal,FM

150-10;310-20

34-75;70-80;129-150

0.5-1(E);0.2-4.7(T etal.)

155-165

Watkins (1981); Cummings et al. (1986); Edds(1988); Thompson et al. (1992)

doublets"A" and"B"

A: 26-16

B: 35-18

1 each 183-186

Cummings and Thompson (1994); Watkins etal. (1987)

(likely same signal as 20Hz tonal above)

clicks(?)

16000-28000

3mseach,9strain

Thompson et al. (1979)

Sei Whale, Balaenoptera borealis


Dur.[s]


References

tonal, FM,pulses

1500-3500

3000 4mseach, 0.5-0.8 train

Thompson et al. (1979); Knowltonet al. (1991)


Bryde's Whale, Balaenoptera edeni


Dur.[s]


References

pulsive 100-900 0.5-51;0.7-1.4

Edds et al. (1993)

moans 70-245 124 0.2-1.5 152-174 Cummings et al. (1986)

Minke Whale, Balaenoptera acutorostrata


Dur.[s]


References

tonal, FM 200-60 0.2-0.4 151-175 Schevill and Watkins (1972);Winn and Perkins (1976); Edds-Walton (2000)

song ? - - 2.4s, 0.3-15x persec

156-161 Gedamke et al. (2001)

pulsive 100-2000;200-400

100-200 0.04-0.07each,1mintrain

Winn and Perkins (1976);Mellinger et al. (2000)

clicks 3300-20000

4000-7500 0.5-1mseach

151 Beamish and Mitchell (1973);Winn and Perkins (1976)

Humpback Whale, Megaptera novaeangliae

Type ∆∆∆∆f [Hz] Dom.f [Hz]

Dur.[s]


References

moans 20-2000;360-1000

300;553

0.2-0.8;0.4-8.2

175-192 Thompson et al. (1986); Cerchioand Dahlheim (2001)

song 20-8000;>15000

120-4000

hours 144-174;171-189

Thompson et al. (1979); Payneand Payne (1985); Au et al.(2001a, 2001b); Cerchio et al.(2001)

pulsive 25-1800 179-185 Thompson et al. (1986)

clicks 2000-8200

Winn et al. (1970); Beamish(1979)


The study by Au et al. (2001b) indicates a common problem with "old" baleenrecordings. In earlier times, the recording bandwidth was often limited to below 8-10kHz andhigh-frequency components of vocalizations were not detected. As the main energy of baleenvocalizations lies at low frequencies, even nowadays researchers record baleen sounds only atlow frequencies. Au et al. (2001b) measured humpback song with 44kHz sampling frequencyand showed much higher song components than were previously reported. Energy at highfrequencies was about 18-24dB less than at low frequencies.

Grey Whale, Family Eschrichtiidae

Grey Whale, Eschrichtius robustus

Type ∆∆∆∆f[Hz]

Dom. f[Hz]

Dur.[s]


References

moans 20-1500

20-200;700-1200

0.1-2 <159;185

Cummings et al. (1968); Fish etal. (1974); Swartz and Cummings(1978); Crane and Lashkari(1996)

pulsive 45-4500

300-800 0.01-0.2(each);1-4(series)

>142 Dahlheim et al. (1984); Dahlheim(1987); Cummings et al. (1968);Moore and Ljungblad (1984);Crane and Lashkari (1996)

tonal, FM 100-350

300 Dahlheim et al. (1984); Mooreand Ljungblad (1984)

clicks 100-10000

1400;3400-4000

1-2 ms Fish et al. (1974); Norris et al.(1977)

3.1 EcholocationSome high-frequency clicks have been recorded on rare occasions in the presence of

baleen whales. It is controversial whether the clicks were actually emitted by the observed baleenwhales, or whether they were recording artifacts, or whether they were emitted by distantodontocetes or other sources (Watkins and Wartzok (1985)). In general, it is believed that a high-frequency echolocation system as used by odontocetes (toothed whales) is lacking in baleenwhales. However, some low-frequency vocalizations might be used for echolocation purposes.Although, echolocation as such has not yet been proven in baleen whales.

Beamish (1978) temporarily blindfolded a captive humpback whale that was thenallowed to swim through a maze of aluminum poles. No evidence of echolocation was obtained.

Ellison et al. (1987) speculated that reverberation of bowhead calls might providebowheads with information about ice conditions and aid in navigation. The possibility thatbaleen whales use their low-frequency sound for long-range echolocation has been discussed byothers, e.g. Clark (1993) and McDonald et al. (1995).


Frazer and Mercado (2000) surmise that male humpback whales might use their song asactive sonar to find females and they support their theoretical ocean acoustic calculations withbehavioral arguments. Au et al. (2001a) counter this publication stressing problems with ambientnoise interference and challenging some of the parameters and values Frazer and Mercado used.Frazer has since revisited his model and adheres to the humpback sonar hypothesis [pers. comm.2002].

4. Inner Ear Anatomy

Anatomical and paleontological evidence suggests that baleen whales are adapted to hearlow frequencies (Fleischer (1976, 1978, 1980); Norris and Leatherwood (1981)). Baleen whaleinner ear anatomy has been studied with dissected ears of dead, stranded animals. Ketten (1991,1992, 1994, 1997) concluded that baleen whales are most sensitive at low sonic to infrasonic(<20Hz) frequencies. The basilar membrane of the cochlea (inner ear) is much broader, thinnerand less rigidly supported than in odontocetes, who are high-frequency hearing specialists.

Houser et al. (2001) used such data from anatomical studies of humpback whale basilarmembranes in combination with psychoacoustic data and anatomical hearing indices of well-studied land-mammals (the cat and the human) to predict a humpback whale audiogram ofrelative hearing sensitivity. Since there is no data on absolute hearing thresholds in humpbackwhales, only relative frequency-dependent sensitivities could be predicted. The resulting U-shaped audiogram showed maximum sensitivity between 2-6 kHz, and a region of bestsensitivity (defined as relative sensitivity < 0.2 dB) between 700Hz and 10kHz. Reduction insensitivity was about 16dB/octave above 10kHz and 6dB/octave below 700Hz. (Though usingthe data and processing methods discussed in their paper, I computed about 20dB/octave above10kHz.) The range of best sensitivity corresponded to the range of humpback vocalizations aslisted above. However, maximum sensitivity occurred at slightly higher frequencies than whatone would have expected from humpback calls. Houser et al. suggested that this could be aninherent contribution of the cat and human audiograms used to predict the humpback audiogram.Houser et al. (2001) further applied 'evolutionary programming' to develop a bandpass filtermodel of the humpback ear, yielding a similar audiogram. The modelled bandpass filters shouldnot be used directly as indicators of critical bandwidths [pers. comm. 2002].

5. Behavioral Reaction to Underwater Sound

The literature on observed behavioral reactions of baleen whales to industrial noise isvast. However, many studies did not estimate received sound pressure levels. Reported reactionswere mostly avoidance behavior, but also include approach; a cessation of feeding, resting, socialinteraction; changes (sometimes cessation) of vocal calling; changes in travel paths and speed;longer dives; and shorter surfacings. The individual responses are not listed here. Differentreactions likely occur at different thresholds, not necessarily in any consistent order. In order toderive an audiogram, any reaction indicates that the sound was heard. If more than one reactionwas reported, I only listed the reactions observed at the lowest noise levels.




Sound f [Hz] received SPL[dB re 1µµµµPa]

Reaction References

13m diesel boat 3rd octave 84 (6dB aboveambient)

avoidance Koski and Johnson(1987)

icebreaker 3rd oct @≈80Hz

80-100 (0-10dB aboveamb)

avoidance LGL and Greeneridge(1995)

small ship broadband 110-115 avoidance Wartzok et al. (1989)approachingboat

20-1000 90 avoidance Richardson and Greene(1993)

Drillship 20-1000 >94-118 (0-20dB aboveambient,>87dB 3rd octband level @275Hz)

avoidance; shortersurfacing, fewerblows

Greene (1987);Richardson et al. (1990);Richardson and Greene(1993)

dredging 20-1000 113-131 (11-30dB aboveambient, >105dB 3rd oct bandlevel @400Hz)

avoidance; shortersurfacing, fewerblows

Richardson et al. (1990);Richardson and Greene(1993)

airgun 20-1000 124-131 shorter surfacing,fewer blows,longer intervalbetw. blows, orientaway

Richardson et al. (1986)

LGL and Greeneridge (1995) found that bowhead responses correlated best with 3rd

octave received levels and signal-to-ambient-noise ratios as opposed to broadband receivedlevels and signal-to-ambient-noise ratios. This seems plausible, because in broadbandmeasurements, part of the acoustic energy might not be audible. Furthermore, mammalian earsintegrate acoustic signals into a bank of auditory filters, called critical bandwidths. These arelikely a fraction of an octave wide in baleen whales, judged from critical band measurements intoothed whales (e.g. Erbe et al. (1999); Richardson et al. (1995)). It is the energy levels in thesecritical bands that determine audibility and response.


Male northern right whales approach mating calls of females; no received sound levelswere quoted (Parks et al. (2001)).



Northern and southern right whales respond to boats and ships, but appear less responsivethan e.g. bowhead whales (Richardson et al. (1994), Ch.9.3.3). No received sound pressurelevels and spectrum levels were found in the literature.



No acoustic response thresholds were found in the literature.



Aburto et al. (1997) reported that blue whales did not alter their vocalization patterns northeir movement patterns at ranges of 5nm from SURTASS LFA trials at full transmit power.Received levels were estimated to be 70-85dB below full transmit power (not quoted!). Croll etal. (2001) found no consistent and statistically significant response of actively foraging bluewhales to SURTASS LFA signals at received levels of 140dB re 1µPa, though vocal behaviorchanged occasionally.


Fin whales have been observed to swim towards distant calling conspecifics (Watkins1981). Croll et al. (2001) found no consistent and statistically significant response of activelyforaging fin whales to SURTASS LFA signals at received levels of 140dB re 1µPa.


No reports on sound levels producing a behavioral response were found in the literature.







Humpback whales react to boats and ships of all sizes (Richardson et al. (1994),Ch.9.3.3), but no received sound pressure levels and spectrum levels were found in the literature.

Sound f [Hz] ReceivedSPL [dBre 1µµµµPa]@ f

Reaction References

pingers 3500 - reducedentanglement

Lien et al. (1992)

pingers broadband,centered @4000

80-90 reducedentanglement

Maybaum (1993); Todd et al.(1992)

sonar 3300;3100-3600

- avoidance,faster swim

Maybaum (1990), (1993)

SURTASS LFAsonar

100-500 120-150 cessation ofsong (T; B etal); increasedsong duration(M et al)

Tyack (1998) Miller et al.(2000); Biassoni et al. (2000)

humpback calls 400-550 102 (16dBS/N); 100-115

approach Frankel and Herman (1993);Frankel et al. (1995); Tyackand Whitehead (1983)

synthetic FMsweep

10-1400 106 approach Frankel et al. (1995)

ATOC 75Hz 60-90 98-109;120-130

longer dives;change of swimdirection

Frankel and Clark (2000),(1998)

airgun broadband 150-169(Ma); 140,112 (Mc)

startle response(Ma);avoidance,startle (Mc)

Malme et al. (1985);McCauley et al. (2000)




Grey whales react to boats and ships of all sizes (Richardson et al. (1994), Ch.9.3.3), butno received sound pressure levels and spectrum levels were found in the literature.

Sound f [Hz] received SPL [dB re1µµµµPa]

Reaction References

drillship;drilling platform

broadband,dominant50-315Hz

108-120; 0dB aboveambient in loudest 3rd octband @ 250Hz

avoidance Malme et al. (1983),(1984)

underwaterplayback ofhelicopter sound

broadband,dominant50-250Hz

111-127; 0dB S/N inloudest 3rd oct band @100Hz

avoidance Malme et al. (1983),(1984)

underwaterplayback ofkiller whale call

loudest 3rd

oct @1kHz0dB signal-to-ambient-noise

avoidance Malme et al. (1983)

100cui singleairgun; 4000cui20-gun array

broadband 140-163 avoidance Malme et al. (1983,1984, 1988)

SURTASS LFAsonar

100-500 110-150 changes inmovement

Clark (1999)

pure tones 800, 900,1000,1500, 1800

95,100,108,135,142respectively; [-13dB]

changes inmovement,startleresponse

Dahlheim andLjungblad (1990)

Dahlheim and Ljungblad (1990) exposed wild grey whales to pure tones to estimatehearing, i.e. reaction, thresholds. Reported levels were source levels. Observed whales werewithin 20 m of the sound projector, and a maximum transmission loss of 13 dB was estimated.

Grey whales appear less responsive to impulsive noise than to continuous noise at thesame level. This is also the case in humans (Fidell et al. (1970)). Reasons might be eitherpsychological (Something that only happens once and is over fast is not worth reacting to.) orphysiological (Baleen whales might have long acoustic integration times, which would reducethe perceived energy of a brief, transient sound, and thus increase the reaction threshold overcontinuous sound.).

6. Physiological Damage

It is generally agreed that any sound at some level can cause physiological damage to theear and other organs and tissues. Data on physiological damage to marine mammals is scarce andis of no direct use to estimating a baleen whale audiogram, the topic of the current study.However, such data falls into the area of noise effects on marine mammals, and the few reportson physiological damage in baleen whales are therefore briefly mentioned here.


Two dead humpback whales with severe mechanical damage to the ears were found inNewfoundland in 1993 in the vicinity of subbottom blasting with 1700-5000kg charges (Kettenet al. (1993); Ketten (1995). The auditory damage was similar to that in humans exposed tosevere blast injury. It is unknown how close to the explosions the whales were at the time ofimpact.

On March 15-16, 2000, 17 cetaceans (Cuvier's beaked whales, Blainville's beakedwhales, minke whales and a spotted dolphin) stranded in the Bahamas. Ten animals werereturned to the water alive; seven are known to have died. Five of the dead beaked whales wereexamined by necropsy showing physiological damage to the auditory system, most likelyacoustically induced. The extent of damage was not considered immediately fatal, butdebilitating enough to compromise hearing and navigational abilities. The animals died of theside-effects of stranding rather than direct acoustic trauma. In the absence of other acousticsources in the area at that time, the mass stranding event was linked to the operation of US Navytactical mid-frequency sonar. Two types of sonar were used; one operating at 2.6 and 3.3kHz at235dB re 1µPa, the other operating at 6.8-8.2kHz at 223dB re 1µPa source level. (These sonarsare different from the SURTASS LFA sonar mentioned in the behavioral response section of thisreport.) It is unknown what received levels caused the physiological damage in the strandedanimals.

A very recent study explores the possibility of creating and measuring blast injury infresh post-mortem ears (Ketten et al. (2001)) using calibrated sources.

7. Constructing an Audiogram

All mammalian (terrestrial and marine) audiograms measured so far are U-shaped,identifying a frequency band of best hearing sensitivity, with decreasing sensitivity at lower andhigher frequencies. In many mammals, the low-frequency portion of the audiogram slopesupward gradually as frequency decreases, typically by 6-12dB per octave (Fay (1988)).

Thresholds of marine mammals, i.e. odontocetes (toothed whales) and pinnipeds (seals,sea lions, walrus), range between 40-70 dB re 1µPa at the frequencies of best sensitivity. It issometimes assumed that baleen whales have similar absolute thresholds at their frequencies ofbest hearing. It is interesting to note that ambient noise in the ocean at these frequencies is quitehigh. Even in quiet conditions (sea state 1) without any industrial activities nearby, averageambient noise levels below 1kHz are above 75dB re 1µPa (3rd octave band levels). Therefore,acoustic detection thresholds of biological and industrial sounds would be limited by ambientnoise rather than the baleen audiogram at those frequencies.

Behavioural response thresholds measured in the wild are usually fairly high (>100dB re1µPa received level). They were often just above ambient noise levels. In other words, fieldobservations of acoustic response thresholds have been limited by ambient noise rather thanbeing indicative of absolute hearing thresholds. Despite the fact that there is likely an offsetbetween the audibility threshold and the behavioural response threshold, this raises the questionof whether baleen hearing is noise-limited rather than threshold-limited at the frequencies of bestsensitivity.


Constructing a baleen audiogram based on recorded baleen vocalizations might be biasedtowards the frequencies of highest energy. The study by Au et al. (2001b) showed thathumpback whales emit much higher frequencies than had previously been recorded. This mightbe the case for other baleen species, too. Hearing bandwidth might thus extend to higherfrequencies than is concluded from low-frequency recordings of baleen sounds.

In my opinion, the most promising methods to obtain a true baleen audiogram in the nearfuture are the following:

1) A portable system to measure audiograms electrophysiologically on living entangled orstranded animals, within a few minutes. A major problem with baleen whales is the large amountof blubber surrounding the brain and attenuating brain waves. I do believe that technicalproblems can be solved in the near future. AEP audiograms don't yield minimum absolutethresholds, but rather give suprathresholds. With large enough signal amplification, these can bevery close to the 'true' absolute thresholds—which are defined and measured via behavioral testwith trained animals—(Szymanski et al. (1999)).2) Anatomical studies with subsequent software modeling of ear responses, such as the study byHouser et al. (2001) on humpback whales. This will only yield a relative audiogram. Absolutethresholds could be obtained with behavioral studies in the field using very band-limited signals(e.g. pure tones) as done by Dahlheim and Ljungblad (1990). This should be done in very quietambient noise conditions and perhaps at frequencies, where there is minimum ambient noise. Afew data points would be enough to 'fix' the relative audiogram in absolute pressure units. Thisapproach would still have the problem of the reaction threshold being potentially higher than theaudibility threshold.

The following is a best estimate of baleen hearing sensitivity based on the currentliterature survey, sorted by species.



Inferring from their vocalizations, bowheads should be most sensitive to frequenciesbetween 20 Hz-5 kHz, with maximum sensitivity between 100-500 Hz. The lowest reported 3rd

octave band level causing a behavioral response was 84dB, followed by 87, 90 and 94 dB.Assuming that the reaction threshold is somewhat higher than the audibility threshold, one mayconclude that the absolute hearing threshold falls indeed into the 40-70 dB range of other marinemammals.


The northern right whale is likely most sensitive between 100-400 Hz according to itsvocalizations. The southern right whale has been studied in more detail and given theirevolutionary relatedness, likely has very similar auditory capabilities.


From its own vocalizations, the southern right whale should be most sensitive between30 Hz - 2.2 kHz, with maximum sensitivity around 50-500 Hz.




Only one study on pygmy right whale calls was found, indicating best hearing sensitivityaround 60-135 Hz.



Blue whale calls have been studied extensively in recent years indicating best hearingsensitivity around 20 Hz, with good sensitivity up to 200 Hz, and perhaps some sensitivity up to8-25 kHz if the clicks reported by Beamish and Mitchell were original blue whale calls.


Similar to the blue whale, fin whale calls yield a region of best sensitivity around 20 Hz,up to 150 Hz. High-frequency clicks between 16-28 kHz were reported in Thompson et al.(1979) though are controversial.


Documentation of sei calls is rare, but indicates best sensitivity between 1.5-3.5 kHz.


Few recordings have been published; best sensitivity is likely around 70-900 Hz.


Minkes appear most sensitive between 100-200 Hz, with good sensitivity extending from60 Hz-2 kHz. High-frequency clicks were published in two studies, indicating some sensitivitybetween 4-7.5 kHz, up to 20 kHz.


Humpback calls have been studied extensively, indicating maximum sensitivity around120 Hz-4 kHz, with good sensitivity from 20 Hz-8 kHz and higher. Anatomical measurementsof humpback ears and subsequent software modeling predicted an audiogram with maximumsensitivity between 2-6 kHz, and good sensitivity between 700 Hz-10 kHz (Houser et al.(2001)). These ranges overlap well with the ranges of vocalizations, however the model mighthave yielded reduced sensitivity at low frequencies due to used psychoacoustic data of morehigh-frequency species (the cat and the human).

Fig. 1 shows the underwater audiogram for the humpback whale as predicted by Houseret al. (2001). Houser only predicted relative sensitivities. I plotted the audiogram twice andpositioned it on the y-axis such that the minimum threshold at the frequency of best hearing was40 dB re 1µPa for the first curve and 70 dB re 1µPa for the second curve. Assuming that the


humpback has absolute thresholds in the range of other non-baleen marine mammal species, thetrue humpback audiogram should lie somewhere between the two curves. For comparison, I alsoplotted the audiograms of two odontocetes, the beluga whale (average of 7 publishedaudiograms, see Erbe and Farmer (1998)) and the killer whale (Hall and Johnson (1972), andSzymanski et al. (1999) via Erbe (2002)). Finally, the in-air audiogram of man in units of dB re1µPa is shown as well (average of 9 data sets, see Fay (1988)).

The lowest reported behavioral thresholds for humpbacks were 80-90 dB received levelfrom pingers at 4 kHz. Assuming that the response threshold likely lies somewhat above theaudibility threshold, absolute sensitivity at 4kHz could well be near the upper range of bestsensitivity of other marine mammals: 70 dB.

102

103

104

105

20

40

60

80

100

120

140

frequency [Hz]

thre

shol

d [d

B r

e 1µ

Pa]

beluga killer human humpback

Figure 1: Audiograms of 2 odontocetes and the human. The 'true' humpback audiogram likelylies somewhere between the 2 humpback audiograms drawn.



Inferring from its own vocalizations, the grey whale should be sensitive to frequenciesbetween 20Hz-4.5kHz, with best sensitivity around 20Hz-1.2kHz. Two studies reported clicksup to 10kHz, with main energy between 1.4-4kHz. The lowest response threshold reported in theliterature was 82-95dB at 800Hz. This again supports the hypothesis that absolute hearingthresholds of baleen whales at the frequencies of best sensitivity overlap with minimumthresholds of other marine mammals.


7.1 Estimating Acoustic ImpactTo determine whether a sound is audible to a whale, one needs 1) the animal's audiogram

and 2) the width of its critical bands. Mammalian ears integrate acoustic energy into a bank ofadjacent and overlapping filters. The width of these filters is called the critical bandwidth.Critical bandwidths have not been measured in baleen whales, but in various odontocetes andpinnipeds. In general, the width increases with increasing frequency. A sound is audible if thereceived energy in any critical band surpasses the audiogram at the (center) frequency of thatcritical band. In the absence of critical band data in baleen whales, a 3rd octave band approach isoften used.

Sound audibility further depends on the level of ambient noise. Ambient noise will maskother sounds. Fletcher (1940) postulated that a sound is detectable if its energy is equal to orgreater than the noise energy in the same critical band. This equal-power-assumption is oftenused and has been corroborated with some (non-baleen) marine mammal experiments, e.g. Erbe(2000).

Mammalian ears not only integrate acoustic signals into frequency-filters, but alsointegrate brief signals over time. Audiograms are generally measured with tones long enough toyield a stable threshold. In terrestrial mammals, as well as odontocetes (toothed whales) andpinnipeds (seals and sea lions), the audibility threshold increases as signals become shorter than0.05-0.1s (see Richardson et al. (1995), Chapter 8.2.4, for review). Nothing is known aboutintegration times in baleen whales.

There is usually an offset between the sound level causing a reaction and the sound levelthat can be detected. Sound will have to be some dB above audibility (i.e. above the audiogramand above ambient noise) before an animal reacts to it. An example where animals react to asound nearly as soon as it becomes detectable is the case of beluga whales avoiding icebreakersin the Canadian Arctic as reported by Cosens and Dueck (1993) and Erbe and Farmer (2000).Temporary behavioral responses are usually not 'biologically significant' (affecting survival).Animals would have to be repeatedly disturbed during important behavior (e.g. nursing, mating,foraging) or be permanently scared away from critical habitat for the effects to be biologicallysignificant.

Industrial noise potentially interferes with communication between whales. It can maskenvironmental sounds that animals might use for navigation, as well as vocalizations used fornavigation and foraging. It can also mask the sounds of prey and the sounds of predators. Inorder to assess the likelihood of masking, one needs the animal audiogram and criticalbandwidth, as well as information on directional hearing and temporal resolution.

Any type of noise at some level has the ability to induce physiological damage to tissuesand organs, e.g. the ear. Hearing impairment can be either temporary (i.e. fully recoverable overtime) or permanent depending on the animal audiogram, the spectral characteristics of the noise(frequency and amplitude), the amount of energy per time for impulsive noise, the duration ofnoise exposure and the duty cycle or recovery time in between exposures. There are only threestudies in the literature on temporary hearing loss in marine mammals: Au et al. (1999) forbottlenose dolphins, Schlundt et al. (2000) for bottlenose dolphins and beluga whales, Kastak etal. (1999) for pinnipeds. This is an active field of research at the moment. Kastak et al (1999)showed that fairly low noise levels of 60-75dB octave band above the normal center-frequency


threshold can already cause a temporary threshold shift of 5dB. There is no data on thresholdsfor permanent hearing loss or other physiological damage in marine mammals.

Whales are indirectly affected by industrial noise if the noise reduces the amount ofavailable prey. A discussion of the effects of underwater noise on fish, invertebrates and marinebirds falls outside the framework of the current project, but could form a topic for a separateresearch project or literature review.

8. Conclusion

The lack of specific data on hearing abilities in baleen whales is a major limitation inevaluating the effects of man-made noise. Baleen whales appear more sensitive to and dependenton low-frequency sounds than odontocetes. Baleen whales are therefore more affected thanother marine mammals. Knowledge of absolute thresholds (i.e. absolute audiograms) is crucialfor estimating acoustic impact. The current study (Section 7) provides and summarizes the bestestimates for baleen hearing to-date, sorted by species.


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Hearing Abilities of Baleen Whales

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