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Aquatic Mammals, Volume 33, Number 4, 2007 ISSN 0167-5427
Marine Mammal Noise Exposure Criteria: Initial Scientific Recommendations
Brandon L. Southall, Ann E. Bowles, William T. Ellison, James J. Finneran, Roger L. Gentry,
Charles R. Greene Jr., David Kastak, Darlene R. Ketten, James H. Miller, Paul E. Nachtigall,
W. John Richardson, Jeanette A. Thomas, & Peter L. Tyack
Contents
Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 411
Chapter 1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 415
Objectives. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 415
Historical Perspective . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 416
Acoustic Measures and Terminology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 417
Sound Production and Use in Marine Mammals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 419
Responses to Sound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 420
Chapter 2. Structure of the Noise Exposure Criteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 427
Sound Types. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 427
Marine Mammal Functional Hearing Groups . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 430
Exposure Criteria Metrics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 434
Levels of Noise Effect: Injury and Behavioral Disturbance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 436
Chapter 3. Criteria for Injury: TTS and PTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 437
Effects of Noise on Hearing in Marine Mammals: TTS Data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 437
Injury from Noise Exposure: PTS-Onset Calculation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 441
Criteria for Injury from a Single Pulse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 442
Criteria for Injury from Multiple Pulses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 444
Criteria for Injury from Nonpulses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 444
Chapter 4. Criteria for Behavioral Disturbance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 446
Behavioral Response Data Analysis Procedures: Disturbance Criteria and Severity Scaling. . . . . . . . . . . . . 448
Criteria for Behavioral Disturbance: Single Pulse. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 451
Behavioral Response Severity Scaling: Multiple Pulses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 452
Behavioral Response Severity Scaling: Nonpulses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 456
Chapter 5. Research Recommendations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474
Measurements of Anthropogenic Sound Sources and Ambient Noise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474
Marine Mammal Auditory Processes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474
Behavioral Responses of Marine Mammals to Sound. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 477
Effects of Noise Exposure on Marine Mammal Hearing and Other Systems . . . . . . . . . . . . . . . . . . . . . . . . . 478
Particularly Sensitive Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 480
Necessary Progressions of Marine Mammal Noise Exposure Criteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 481
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 482
Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 482
Appendix A. Acoustic Measures and Terminology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 498
Appendix B. Studies Involving Marine Mammal Behavioral Responses to Multiple Pulses . . . . . . . . . . . . . . . . . 502
Appendix C. Studies Involving Marine Mammal Behavioral Responses to Nonpulses . . . . . . . . . . . . . . . . . . . . . 509
Aquatic Mammals
Papers dealing with all aspects of the care,
conservation, medicine, and science of aquatic mammals
Editor and Copyright: Jeanette A. Thomas
Department of Biological Sciences, Western Illinois UniversityQuad Cities, Moline, IL 61265, USA
Co-Editor: Kathleen Dudzinski
Dolphin Communication Project, Stonington, Connecticut 06378, USA
Prepared and published by Document and Publication Services, Western Illinois University, Macomb, IL 61455, USA
Marine Mammal Noise Exposure Criteria:
Initial Scientifc Recommendations
Supported through Joint Sponsorship by the European Association for
Aquatic Mammals, the Alliance of Marine Mammal Parks and Aquariums,
and the International Marine Animal Trainers Association
Founded by EAAM in 1974
Established by the EAAM in 1974
AQUATIC MAMMALS
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AQUATIC MAMMALS
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President Elect: Birgitta Mercera, Parc Asterix, BP 8, 60128 Plailly, France;
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Secretary/Treasurer: Sabrina Brando, Animal Concepts, Sea Mammal Research Unit, Gatty Marine Laboratory,
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MarineMammalNoiseExposureCriteria
Marine Mammal Noise Exposure Criteria:
Initial Scientific Recommendations
Brandon L. Southall,
1, 2
Ann E. Bowles,
3
Wllam T. Ellson,
4
James J. Fnneran,
5

Roger L. Gentry,
6
Charles R. Greene Jr.,
7
Davd Kastak,
2
Darlene R. Ketten,
8, 9

James H. Mller,
10
Paul E. Nachtgall,
11
W. John Rchardson,
12

Jeanette A. Thomas,
13
and Peter L. Tyack
8
1
NationalOceanicandAtmosphericAdministration(NOAA),NationalMarineFisheriesService,OfficeofScience
andTechnology,MarineEcosystemsDivision,NOAAsOceanAcousticsProgram,1315East-WestHighway#12539,
SilverSpring,MD20910-6233,USA;E-mail:Brandon.Southall@noaa.gov
2
LongMarineLaboratory,UniversityofCaliforniaatSantaCruz,100ShafferRoad,SantaCruz,CA95060,USA
3
Hubbs-SeaWorldResearchInstitute,2595IngrahamStreet,SanDiego,CA92109,USA
4
MarineAcoustics,Inc.,809AquidneckAvenue,Middletown,RI02842,USA
5
U.S.NavyMarineMammalProgram,SpaceandNavalWarfareSystemsCenterSanDiego,
53560HullStreet,SanDiego,CA92152-5000,USA
6
ProScienceConsulting,LLC,P.O.Box177,Dickerson,MD20842-0177,USA
7
GreeneridgeSciences,Inc.,4512ViaHuerto,SantaBarbara,CA93110,USA
8
WoodsHoleOceanographicInstitution,WoodsHole,MA02543,USA
9
HarvardMedicalSchool,DepartmentofOtologyandLaryngology,Boston,MA02114,USA
10
UniversityofRhodeIsland,DepartmentofOceanEngineering,SouthFerryRoad,Narragansett,RI02882,USA
11
HawaiiInstituteofMarineBiology,P.O.Box1346,Kaneohe,HI96744,USA
12
LGLLtd.,environmentalresearchassociates,P.O.Box280,22FisherStreet,KingCity,ONL7B1A6,Canada
13
WesternIllinoisUniversityQuadCities,DepartmentofBiologicalSciences,356160thStreet,Moline,IL61265,USA
Contents
Overvew . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 411
Chapter 1. Introducton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 415
Objectves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 415
Hstorcal Perspectve . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 416
Acoustc Measures and Termnology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 417
Sound Producton and Use n Marne Mammals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 419
Responses to Sound. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 420
Chapter 2. Structure of the Nose Exposure Crtera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 427
Sound Types. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 427
Marne Mammal Functonal Hearng Groups . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 430
Exposure Crtera Metrcs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 434
Levels of Nose Effect: Injury and Behavoral Dsturbance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 436
Chapter 3. Crtera for Injury: TTS and PTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 437
Effects of Nose on Hearng n Marne Mammals: TTS Data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 437
Injury from Nose Exposure: PTS-Onset Calculaton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 441
Crtera for Injury from a Sngle Pulse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 442
Crtera for Injury from Multple Pulses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 444
Crtera for Injury from Nonpulses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 444
Chapter 4. Crtera for Behavoral Dsturbance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 446
Behavoral Response Data Analyss Procedures: Dsturbance Crtera and Severty Scalng. . . . . . 448
Crtera for Behavoral Dsturbance: Sngle Pulse. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 451
Behavoral Response Severty Scalng: Multple Pulses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 452
Behavoral Response Severty Scalng: Nonpulses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 456
Southalletal.
Chapter 5. Research Recommendatons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474
Measurements of Anthropogenc Sound Sources and Ambent Nose . . . . . . . . . . . . . . . . . . . . . . . 474
Marne Mammal Audtory Processes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474
Behavoral Responses of Marne Mammals to Sound. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 477
Effects of Nose Exposure on Marne Mammal Hearng and Other Systems . . . . . . . . . . . . . . . . . . 478
Partcularly Senstve Speces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 480
Necessary Progressons of Marne Mammal Nose Exposure Crtera . . . . . . . . . . . . . . . . . . . . . . . 481
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 482
Lterature Cted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 482
Appendx A. Acoustc Measures and Termnology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 498
Appendx B. Studes Involvng Marne Mammal Behavoral Responses to Multple Pulses . . . . . . . . 502
Appendx C. Studes Involvng Marne Mammal Behavoral Responses to Nonpulses . . . . . . . . . . . . 509
MarineMammalNoiseExposureCriteria
Acronyms
Acronym Defnton
A-weghtng Frequency-selectve weghtng for aeral hearng n humans derved from the nverse of the dealzed
40-phon equal loudness hearng functon across frequences
ABR Audtory branstem response
ADD Acoustc deterrent devce
AEP Audtory evoked potentals
AHD Acoustc harassment devce
ANSI Amercan Natonal Standards Insttute
ASSR Audtory steady-state response
ATOC Acoustc Thermometry of Ocean Clmate program
CF Center frequency
C-weghtng Frequency-selectve weghtng for aeral hearng n humans derved from the nverse of the dealzed
100-phon equal loudness hearng functon across frequences
EFR Envelope followng response
EPA U.S. Envronmental Protecton Agency
ES Exploson smulator
fhgh Estmated upper functonal hearng lmt
flow Estmated lower functonal hearng lmt
HESS Hgh Energy Sesmc Survey
HPA Hypothalamc-ptutary-adrenal axs
IMAPS Integrated Marne Mammal Montorng and Protecton System
ISO Internatonal Standards Organzaton
JNCC U.K. Jont Nature Conservaton Commttee
LeqT Equvalent-contnuous sound level over perod T
LIeqT Impulse equvalent-contnuous sound level over perod T
LFA Low Frequency Actve (sonar)
M-weghtng Generalzed frequency weghtngs for varous groups of marne mammals, allowng for ther
functonal bandwdths and approprate n characterzng audtory effects of strong sounds
Mlf Frequency weghtng for low-frequency cetaceans (mystcetes)
Mmf Frequency weghtng for md-frequency cetaceans (most odontocetes)
Mhf Frequency weghtng for hgh-frequency cetaceans (odontocetes specalzed for use of very hgh
frequences)
Mpw Frequency weghtng for pnnpeds, lstenng n water
Mpa Frequency weghtng for pnnpeds, lstenng n ar
MMPA U.S. Marne Mammal Protecton Act
NIHL Nose-nduced hearng loss
NIPTS Nose-nduced permanent threshold shft
NIOSH U.S. Natonal Insttute for Occupatonal Safety and Health
NMFS U.S. Natonal Marne Fsheres Servce
v Southalletal.
Acronym Defnton
NOAA U.S. Natonal Oceanc and Atmospherc Admnstraton
NRC U.S. Natonal Research Councl
NRL U.S. Naval Research Laboratory
Pmax Maxmum sound pressure
OBN Octave-band nose
PCAD Natonal Research Councls Populaton Consequences of Acoustc Dsturbance Model
PICE Porpose ncdental catch elmnaton
PTS Permanent threshold shft
REFMS A computer program for predctng shock-wave propagaton from underwater explosons
RL Receved level
RMS Root-mean-square
SEL Sound exposure level
SL Source level (receved level measured or estmated 1 m from the source)
SLM Sound level meter
SPL Sound pressure level
TS Threshold shft
TTS Temporary threshold shft
USC Unted States Code
VAFB Vandenberg Ar Force Base
Overview
A group of experts n acoustc research from
behavoral, physologcal, and physcal dscplnes
was convened over a several year perod. The pur-
pose of ths panel was to revew the expandng lt-
erature on marne mammal hearng and on phys-
ologcal and behavoral responses to anthropogenc
sound, and to propose exposure crtera for certan
effects. The group employed all avalable relevant
data to predct nose exposure levels above whch
adverse effects on varous groups of marne mam-
mals are expected. Recent advances n these felds
and the pressng need for a scence-based para-
dgm to assess the effects of sound exposure were
the prmary motvatons for ths effort. Two cat-
egores of effects were consdered: (1) njury and
(2) behavoral dsturbance. The proposed crtera
for the onset of these effects were further segre-
gated accordng to the functonal hearng capa-
bltes of dfferent marne mammal groups, and
accordng to the dfferent categores and metrcs
of typcal anthropogenc sounds n the ocean. The
group acheved many of ts objectves but acknowl-
edges certan lmtatons n the proposed crtera
because of scarcty or complete absence of nfor-
maton about some key topcs. A major component
of these recommendatons s a call for specfc
research on crtcal topcs to reduce uncertanty
and mprove future exposure crtera for marne
mammals. Ths publcaton marks the culmnaton
of a long and challengng ntal effort, but t also
ntates a necessary, teratve process to apply and
refne nose exposure crtera for dfferent speces
of marne mammals.
The process of establshng polcy gudelnes
or regulatons for anthropogenc sound exposure
(.e., the applcaton of these exposure crtera) wll
vary among natons, jursdctons, and legal/polcy
settngs. Such processes should carefully consder
the lmtatons and caveats gven wth these pro-
posed crtera n decdng whether suffcent data
currently exst to establsh smplstc, broad crte-
ra based solely on exposure levels. In many cases,
especally for behavoral dsturbance, context-
specfc analyses consderng prevous studes on
speces and condtons smlar to those n queston
mght, at least for the foreseeable future, be more
approprate than general gudelnes.
StateofCurrentKnowledge
The avalable data on the effects of nose on
marne mammals are qute varable n quantty
and qualty. In many respects, data gaps severely
restrct the dervaton of scentfcally-based nose
exposure crtera and, n some cases, explct
threshold crtera for certan effects are not appro-
prate gven the amount and type of data avalable.
Scentfc nqury nto acoustc communcaton
among marne mammals extends back more than
half a century, but most of the specfc data rel-
evant to the proposed crtera have been publshed
wthn the last two decades. Owng to the mount-
ng publc, scentfc, and regulatory nterest n
conservaton ssues related to acoustcs, the aval-
able scence s progressng rapdly (e.g., see NRC,
2003, 2005).
Ths paper proposes, for varous marne mammal
groups and sound types, levels above whch there
s a scentfc bass for expectng that exposure
would cause audtory njury to occur. Controlled
measurements of hearng and of the effects of
underwater and aeral sound n laboratory settngs
have greatly expanded the ablty to assess aud-
tory effects. Whle understandng of the hearng
capactes among all marne mammals remans
admttedly rudmentary, there s a farly detaled
understandng of some key aspects of underwater
and aeral hearng n a few representatve speces
of odontocetes, pnnpeds, and srenans, although
hearng n mystcetes remans untested. Avalable
data, along wth the compellng evdence of smlar
audtory processes among all mammals, enables
some reasonable extrapolatons across speces for
estmatng audtory effects, ncludng the exposure
levels of probable onset of njury. Recent evdence
suggests that exposure of beaked whales to under-
water nose may, under certan (generally unknown)
condtons, result n non-audtory njury as well
(e.g., Fernndez et al., 2005). At present, however,
there are nsuffcent data to allow formulaton of
quanttatve crtera for non-audtory njures.
There are many more publshed accounts of
behavoral responses to nose by marne mammals
than of drect audtory or physologcal effects.
Nevertheless, the avalable data on behavoral
responses do not converge on specfc exposure
condtons resultng n partcular reactons, nor do
they pont to a common behavoral mechansm.
Even data obtaned wth substantal controls,
precson, and standardzed metrcs ndcate hgh
varance both n behavoral responses and n expo-
sure condtons requred to elct a gven response.
It s clear that behavoral responses are strongly
AquaticMammals 2007, 33(4), 411-414, DOI 10.1578/AM.33.4.2007.411
412 Southalletal.
affected by the context of exposure and by the an-
mals experence, motvaton, and condtonng.
Ths realty, whch s generally consstent wth
patterns of behavor n other mammals (nclud-
ng humans), hampered our efforts to formulate
broadly applcable behavoral response crtera for
marne mammals based on exposure level alone.
Frequency-WeightingFunctions
In humans, hearng processes n a large number
of male and female subjects of dfferent ages
have been tested to determne a basc audomet-
rc curve, equal-loudness curve, and the levels and
exposure duratons needed to nduce ether recov-
erable hearng loss (called temporary threshold
shft or TTS) or permanent threshold shft (PTS).
In addton, the manner n whch successve expo-
sures to nose contrbute to TTS growth has been
well-documented n humans (e.g., Kryter, 1994;
Ward, 1997). In assessng the effects of nose
on humans, ether an A- or C-weghted curve
s appled to correct the sound-level measure-
ment for the frequency-dependent hearng func-
ton of humans. Early on, the panel recognzed
that smlar, frequency-weghted hearng curves
were needed for marne mammals; otherwse,
extremely low- and hgh-frequency sound sources
that are detected poorly, f at all, mght be subject
to unrealstc crtera.
One of the major accomplshments n ths
effort was the dervaton of recommended fre-
quency-weghtng functons for use n assessng
the effects of relatvely ntense sounds on hearng
n some marne mammal groups. It s abundantly
clear from measurements of marne mammal hear-
ng n the laboratory, call characterstcs, and aud-
tory morphology that there are major dfferences
n audtory capabltes across marne mammal
speces (e.g., Wartzok & Ketten, 1999). Most pre-
vous assessments of acoustc effects ether faled
to account for dfferences n functonal hearng
bandwdth among marne mammal groups or dd
not recognze that the nomnal audogram mght
be a relatvely poor predctor of how the audtory
system responds to relatvely strong exposures.
The authors delneated fve groups of marne
mammals based on smlartes n ther hearng, and
they developed a generalzed frequency-weght-
ng (called M-weghtng) functon for each.
The fve groups and the assocated desgnators are
(1) mystcetes (baleen whales), desgnated
as low-frequency cetaceans (Mlf); (2) some
odontocetes (toothed whales), desgnated as
md-frequency cetaceans (Mmf); (3) odontocetes
specalzed for usng hgh frequences (.e., por-
poses, rver dolphns, and the genera Kogia and
Cephalorhynchus) (Mhf); (4) pnnpeds (.e., seals,
sea lons, and walruses) lstenng n water (Mpw);
and (5) pnnpeds lstenng n ar (Mpa). These cr-
tera do not specfcally address srenans, the sea
otter, or the polar bear, n part because of the lack
of key data n these speces.
The M-weghtng functons were defned based
on known or estmated audtory senstvty at df-
ferent frequences rather than vocal characterstcs
per se. Owng to the paucty of relevant data, these
audtory functons are ntentonally precauton-
ary (wde) and lkely overestmate the functonal
bandwdth for most or all speces. Ther prmary
applcaton s n predctng audtory damage rather
than levels of detecton or behavoral response.
Consequently, t s more approprate to use flat-
ter functons than would be obtaned by employ-
ng a smple nverse-audogram functon.
ExposureCriteriaMetrics
To further complcate the dervaton of nose expo-
sure crtera, sounds can be descrbed wth varous
acoustc metrcs, ncludng sound pressure levels
and sound exposure levels. The latter s a measure
of receved sound energy. Avalable lterature pro-
vdes a mxture of both measures, but many sound
sources have prmarly been descrbed n pressure
level unts. To accommodate these two measures,
and to account for all relevant acoustc features
that may affect marne mammals, we developed
dual crtera for nose exposures n each of the fve
functonal hearng groups, usng both sound pres-
sure and sound exposure levels.
ExposureCriteriaforInjury
Another area n whch we provde substantve
conclusons s n the determnaton of sound
exposures beleved to cause drect audtory njury
to marne mammals. By all accounts, the nner
ear s the organ system most drectly senstve to
sound exposure and, thus, the most susceptble to
sound-derved damage. We defne the mnmum
exposure crteron for njury as the level at whch
a sngle exposure s estmated to cause onset of
permanent hearng loss (PTS). Data on TTS n
marne mammals, and on patterns of TTS growth
and ts relaton to PTS n other mammals, were
used to estmate thresholds for njury. Owng to
the lmted avalablty of relevant data on TTS
and PTS, the extrapolaton procedures underlyng
these estmatons are necessarly precautonary.
To account for all of the potentally njurous
aspects of exposure, dual crtera for njury were
establshed for each functonal marne mammal
hearng group based on nstantaneous peak pres-
sure (unweghted) and total energy (M-weghted).
Exposure crtera for njury are gven for two types
of sounds, pulse and nonpulse, and for sngle and
multple exposures. The term pulse s used here to
descrbe bref, broadband, atonal, transents (ANSI,
MarineMammalNoiseExposureCriteria 413
1986; Harrs, 1998, Chapter 12), whch are charac-
terzed by a relatvely rapd rse-tme to maxmum
pressure followed by a decay that may nclude a
perod of dmnshng and oscllatng maxmal and
mnmal pressures. Examples of pulses are sounds
from explosons, gunshots, sonc booms, sesmc
argun pulses, and ple drvng strkes. Nonpulse
(ntermttent or contnuous) sounds can be tonal,
broadband, or both. They may be of short dura-
ton but wthout the essental propertes of pulses
(e.g., rapd rse-tme). Examples of anthropogenc,
oceanc sources producng such sounds nclude
vessels, arcraft, machnery operatons such as
drllng or wnd turbnes, and many actve sonar
systems. As a result of propagaton, sounds wth
the characterstcs of a pulse at the source may lose
pulsatle characterstcs at some (varable) dstance
and can be characterzed as a nonpulse by certan
recevers.
Regardless of the anthropogenc sound, f a
marne mammals receved exposures exceed the
relevant (pulse or nonpulse) crteron, audtory
njury (PTS) s assumed to be lkely. Chapter 3,
Crtera for Injury, provdes detals regardng
the exposure levels requred to cause TTS-onset
and the extrapolaton of those results to estmate
levels above whch PTS-onset may occur. For all
fve functonal hearng groups, we propose dual
exposure crtera above whch audtory njury s
lkely.
ExposureCriteriaforBehavior
One challenge n developng behavoral crtera
s to dstngush a sgnfcant behavoral response
from an nsgnfcant, momentary alteraton n
behavor. For example, the startle response to a
bref, transent event s unlkely to persst long
enough to consttute sgnfcant dsturbance. Even
strong behavoral responses to sngle pulses, other
than those that may secondarly result n njury
or death (e.g., stampedng), are expected to ds-
spate rapdly enough as to have lmted long-term
consequence. Consequently, upon exposure to a
sngle pulse, the onset of sgnfcant behavoral
dsturbance s proposed to occur at the lowest
level of nose exposure that has a measurable
transent effect on hearng (.e., TTS-onset). We
recognze that ths s not a behavoral effect per
se, but we use ths audtory effect as a de facto
behavoral threshold untl better measures are
dentfed. Lesser exposures to a sngle pulse are
not expected to cause sgnfcant dsturbance,
whereas any compromse, even temporarly, to
hearng functons has the potental to affect vtal
rates through altered behavor.
For other anthropogenc sound types (multple
pulses, nonpulses), we conducted an extensve
revew of the avalable lterature but were unable
to derve explct and broadly applcable numer-
cal threshold values for delneatng behavoral
dsturbance. We dd develop a quanttatve scor-
ng paradgm that numercally ranks, as a severty
scalng, behavoral responses observed n ether
feld or laboratory condtons. We appled ths
approach to the approprate behavoral data for
multple pulses and nonpulses. Some of these data
suffer from poor statstcal power, lmted nfor-
maton on receved sound levels and background
nose, nsuffcent measurements of all potentally
mportant contextual varables, and/or nsuffcent
controls. Some such data are analyzed here solely
for llustratve purposes. Most behavoral studes
suffered from at least some of these problems.
Therefore, we do not ntend to gve unform sc-
entfc credence to all of the cted data, and we
expect future studes to gve greater attenton and
rgor to these crtcal requrements.
Ths revew and scorng process, whle not a
formal meta-analyss for normalzng and pool-
ng dsparate observatons, corroborated certan
nterestng aspects of marne mammal behavoral
responses to sound exposure. Foremost was that
a behavoral response s determned not only by
smple acoustc metrcs, such as receved level
(RL), but also by contextual varables (e.g., labo-
ratory vs feld condtons, anmal actvty at the
tme of exposure, habtuaton/senstzaton to
the sound, etc.). Also mportant s the presence
or absence of acoustc smlartes between the
anthropogenc sound and bologcally relevant
natural sgnals n the anmals envronment (e.g.,
calls of conspecfcs, predators, prey). Wthn
certan smlar condtons, there appears to be
some relatonshp between the exposure RL and
the magntude of behavoral response. However,
n many cases, such relatonshps clearly do not
exst, at least when response data are pooled
across multple speces and contexts. Ths argues
for a context-based approach to dervng nose
exposure crtera for behavoral responses. That
concept, along wth our revew and scalng of the
avalable observatonal data, provdes a founda-
ton for establshng dose-response relatonshps
for some specfc crcumstances and a startng
pont for future analyses when addtonal data are
avalable.
ConclusionsandResearchRecommendations
Ths process has resulted n several sgnfcant
advances. These nclude a revew and nterpre-
taton of the avalable lterature on njury and
behavoral data usng precautonary extrapola-
ton procedures, dervaton of marne mammal
frequency-weghtng functons, specfcaton of
quanttatve crtera for audtory njury, and derva-
ton of a severty scale for behavoral responses.
414 Southalletal.
The nablty to dentfy broadly applcable,
quanttatve crtera for behavoral dsturbance n
response to multple-pulse and nonpulse sounds s
an acknowledged lmtaton.
Our efforts to derve marne mammal nose
exposure crtera clearly llustrate the fact that,
at present, research n ths feld remans lmted
n many areas. The need for extrapolaton pro-
cedures and precautonary assumptons ponts
drectly to research needs n a varety of areas on a
varety of speces. In certan condtons, proposed
crtera for an entre marne mammal group are
based on the most precautonary measurement
or observaton for a speces wthn that group,
despte the fact that, for other speces wthn that
group, there are emprcal data ndcatng that
hgher exposures are requred to nduce the same
effect. We beleve t s approprate to use the most
precautonary data n proposng group-wde crte-
ra applcable for speces where there are no drect
measurements. We also feel t s approprate on
a case-by-case bass to apply the most relevant
emprcal data (.e., from the speces or genus of
concern) n settng the exposure thresholds spec-
fed n polcy gudelnes.
Fnally, we emphasze that exposure crtera for
sngle ndvduals and relatvely short-term (not
chronc) exposure events, as dscussed here, are
nsuffcent to descrbe the cumulatve and eco-
system-level effects lkely to result from repeated
and/or sustaned human nput of sound nto the
marne envronment and from potental nterac-
tons wth other stressors. Also, the njury crtera
proposed here do not appear to predct what may
have been ndrect njury from acoustc exposure
n several cases where cetaceans of several spe-
ces mass-stranded followng exposure to mltary
sonar.
The extensve research recommendatons gven
here (see Chapter 5) represent our collectve
vew of the concerted effort that wll be requred
over the comng decades. Hgh prorty catego-
res of research nclude (1) contnued expanson
of knowledge on basc marne mammal hearng
capabltes, ncludng sound localzaton, the
detecton of realstc sound sgnals, commun-
caton maskng, and audtory scene analyss;
(2) contnued expanson of knowledge on baselne
marne mammal behavoral patterns; (3) well-
controlled, drect measurements (usng appropr-
ate, standardzed acoustc metrcs) of the effects
of sound exposure on marne mammal hearng,
behavor, and physology; and (4) rsk-assessment
studes of the cumulatve and synergstc effects
of nose and other exposure(s) on ndvduals and
populatons.
Understandng and managng the effects
of nose on marne lfe wthout unjustfably
constranng mportant human actvtes n the
oceans wll contnue to be challengng for the
foreseeable future. Wth sustaned and focused
research n key areas, future scentsts wll be
equpped to make nformed mprovements to the
ntal scentfc recommendatons presented here.
These mprovements should deally be ntegrated
nto scence-based rsk assessment models that
consder all aspects of sound exposure and other
potental stressors on ndvdual marne mammals,
populatons, and marne ecosystems.
1. Introduction
Objectives
Recent nterest and concern about the effects
of anthropogenc nose on marne mammals
has trggered consderable new research (e.g.,
Costa et al., 2003; Frstrup et al., 2003; Fnneran
et al., 2005a), summares of avalable nformaton
(Rchardson et al., 1995; Wartzok & Ketten, 1999),
and recommendatons for specfc acton (NRC,
1994, 2000, 2003, 2005). Systematc, objectve,
scence-based nterpretaton of the avalable data s
crtcally needed to nform management agences
charged wth mtgatng adverse effects of anthro-
pogenc nose on protected speces. In response to
ths need, we use here the full body of scentfc
data on marne mammal hearng and the effects of
nose on hearng and behavor, augmented where
approprate by nterpretatons of terrestral mammal
(ncludng human) data, to develop proposed expo-
sure crtera that are as comprehensve, defensble,
and precse as s currently possble. The scope of
these crtera ncludes njurous and behavoral
effects of a sngle nose exposure event on an nd-
vdual cetacean (whales, dolphns, and porposes)
or pnnped (seals, sea lons, and walruses).
The recommended nose exposure crtera are
scence-based, developed wthout addressng the
commercal, socetal, or practcal ramfcatons
of mplementng the conclusons reached here.
We ntend to mrror the process used n the devel-
opment of damage rsk crtera for humans (see
Crocker, 1997). Polcy gudelnes developed for
regulatory and socetal purposes are based both on
scentfc evdence (as summarzed n ths paper
for marne mammals) and on other consderatons
(e.g., economc, practcal, socal, and ethcal) not
dealt wth here. Thus, on certan ponts, polcy
gudelnes that are developed separately for the
purposes of varous jursdctons, natons, or users
of these crtera may dffer from the scence-based
crtera recommended here.
All forms of anthropogenc nose receved by
marne mammals were consdered, whether pro-
duced under water or n ar, and we adopted a
comparatve approach, whch we regard as essen-
tal to any crtera-settng process for nonhuman
anmals. For most of the ~128 marne mammal
speces and subspeces (Rce, 1998) consdered
here, no emprcal data were avalable on nomnal
hearng characterstcs or on the effects of nose
on hearng or behavor. Practcal, ethcal, and
legal consderatons lmt the level of scentfc
nformaton that s avalable for dervng crtera
applcable to ether humans or marne mammals.
Consequently, certan assumptons and crtera
proposed here were based on nformaton from
other mammalan groups, where justfed. Where
such data present a varety of optons, we made
ntentonally precautonary decsons (.e., lower
proposed exposure levels) to reduce the rsk of
assumng no effect when one was actually present.
The term precautonary s used here wthout ref-
erence to any regulatory or polcy mplcaton of
ths word. Scentsts would more conventonally
use the term conservatve n ths regard rather
than the more bureaucratc precautonary, but n
certan complex nstances here, the term conser-
vatve would be potentally ambguous, depend-
ng on the perspectve of the reader. When nfor-
maton was lmted, extrapolatons were made
cautously to mnmze the rsk of falng to recog-
nze an effect when one actually occurs (Type-II
statstcal error) as can occur wth small sample
szes or mprecse measurements.
Each generalzaton/extrapolaton was dent-
fed, all precautonary decsons were noted, and
the logc leadng to each proposed crteron was
specfed. Thus, when new data become avalable,
approprate modfcatons can be made readly.
Studes that are needed to resolve the uncertan-
tes encountered n developng the current crtera
are dscussed n detal (see Chapter 5, Research
Recommendatons). Realstcally, however, the
generalzaton of nformaton between related
speces wll reman essental n many cases for the
foreseeable future.
Our ntent was to derve recommended nose
exposure crtera usng the best nformaton cur-
rently avalable, dentfy weaknesses n the present
approach, call for relevant research, and structure
the crtera such that future mprovements can be
ncorporated easly. Lack of data lmted the pro-
posed nose exposure crtera to ndvdual marne
mammals exposed to acute exposure events (such
as the passage of one vessel or a seres of actve
sonar transmssons). Also, the proposed crtera
are lmted to cetaceans and pnnpeds. We expect
that nose exposure crtera for other marne
mammals (manatees, dugongs, polar bears, and
sea otters), as well as other marne taxa, wll be
developed as addtonal data become avalable and
are evaluated. In fact, a separate expert panel (S3/
AquaticMammals 2007, 33(4), 415-426, DOI 10.1578/AM.33.4.2007.415
416 Southalletal.
WG92: Effects of Sound on Fsh and Turtles) has
been establshed under the Standards Commttee
(S3) of the Acoustcal Socety of Amerca to con-
sder nose exposure crtera for fsh and turtles.
Addtonally, crtera are clearly needed for cumu-
latve effects and for effects at speces or even eco-
system levels, but data to support those types of
crtera do not currently exst.
The present recommended crtera represent a
major step n ntatng a lengthy, systematc pro-
cess to predct and dentfy acoustc exposure con-
dtons (natural or anthropogenc) assocated wth
varous effects on marne mammals. Ths paper s
delberately structured n a somewhat formulac
and report-lke manner so that the logc underly-
ng certan assumptons and extrapolatons (as
well as the data needed to test and/or strengthen
them) s self-evdent. We expect there wll be an
teratve process of mprovng and expandng the
complexty of the exposure crtera, smlar to the
decades-long development of human nose expo-
sure crtera (see Crocker, 1997). Because of the
matrx structure of the proposed crtera, thresh-
olds n specfc cells can be updated ndependently
as new nformaton becomes avalable.
There s an extensve hstory and dversty of
exposure crtera for humans wth varous knds of
acoustc exposure. A full dscusson of these crte-
ra s beyond the scope of ths paper, but examples
nclude workplace nose standards (e.g., NIOSH,
1998), standards for the protecton of mltary
personnel (U.S. DoD, 1997), and natonal polcy
gudelnes (e.g., EPA, 1974; BG PPG, 1994).
Several addtonal examples were also consdered,
whether receved under water or n ar, n varous
decsons underlyng the marne mammal crtera
proposed here. The process of establshng human
nose exposure crtera has been dffcult and con-
tentous, but establshng nose exposure crtera
for marne mammals s consderably more daunt-
ng gven the dversty of marne mammal speces
across three orders, the complexty of aeral and
underwater acoustc exposures, and profound data
lmtatons.
Historical Perspective
Concerns about potental adverse effects of anthro-
pogenc nose on marne lfe began n the 1970s
(e.g., Payne & Webb, 1971) and expanded n the
1980s. Experments durng the 1980s wth sesmc
arguns ndcated that bowhead whales (Balaena
mysticetus) and gray whales (Eschrichtiusrobus-
tus) exhbted clear, sustaned avodance of opera-
tonal areas at dstances where pulse root-mean-
square (RMS) sound pressure levels (SPLs) were
160 to 170 dB re: 1 Pa (Malme et al., 1983, 1984,
1986, 1988; Rchardson et al., 1986; Ljungblad
et al., 1988). In contrast, early observatons of
bowhead and gray whales exposed to contnu-
ous ndustral sounds, such as those assocated
wth drllng operatons, suggested 120 dB re:
1 Pa as the approxmate threshold for behavoral
dsturbance of these baleen whales (Malme et al.,
1984; Rchardson et al., 1990a, 1995 [pp. 286-
287]). Sgnfcant ndvdual varablty was noted
n typcal behavoral responses, however, wth
some ndvdual whales respondng only when
very close to sound sources and others reactng
at much longer dstances (and to lower receved
sound levels). Ths varablty rases questons as
to whether behavoral responses are most appro-
prately descrbed by the exposure receved level
(RL) of the stmulus at the anmal, the sgnal-to-
ambent nose dfferental, the rate of change of
the sgnal, or smply to the presence of the human
actvty as ndcated by acoustc cues and/or vsual
stmul.
Concern about the effects of acoustc pulses
from sesmc exploraton and contnuous sound
from other ndustral actvtes resulted n the
mposton of mtgaton requrements on some
ndustral actvtes n certan jursdctons by
the early- to md-1980s. Subsequent events,
such as the Heard Island Feasblty Test n
1991 (Baggeroer & Munk, 1992), the Acoustc
Thermometry of Ocean Clmate (ATOC) pro-
gram n the late-1990s (see NRC, 1994, 2000; Au
et al., 1997; Costa et al., 2003), and the U.S.
Navys low-frequency actve sonar program (e.g.,
Croll et al., 2001) resulted n popular and govern-
mental nterest n settng crtera for safe levels of
sound for marne mammal exposure (NRC, 1994,
2000, 2003; Rchardson et al., 1995). Ths nterest
has expanded wth the fndng that tactcal, md-
frequency, mltary sonar transmssons are some-
tmes correlated, n specfc condtons, wth mass
strandng events of (predomnantly) several beaked
whale speces, ncludng Cuvers (Ziphiuscaviro-
stris), Blanvlles (Mesoplodondensirostris), and
Gervas (Mesoplodoneuropaeus) beaked whales
(see Evans & England, 2001; Fernndez et al.,
2005; Cox et al., 2006).
In 1995, the U.S. Natonal Marne Fsheres
Servce (NMFS) set underwater do not exceed
crtera for exposure of marne mammals to
underwater pulses from sesmc arguns. These
crtera were 190 dB re: 1 Pa for pnnpeds and
most odontocete cetaceans and 180 dB re: 1 Pa
for mystcetes and sperm whales (Physeter mac-
rocephalus) (and, by nference, for pygmy and
dwarf sperm whales [Kogia spp.]). These exposure
lmts were ntended as precautonary estmates of
exposures below whch physcal njury would not
occur n these taxa. There was no emprcal ev-
dence as to whether exposure to hgher levels of
MarineMammalNoiseExposureCriteria 417
pulsed sounds would or would not cause audtory
or other njures. Gven the lmted data then aval-
able, however, t could not be guaranteed that
marne mammals exposed to hgher levels would
not be njured. Further, t was recognzed that
behavoral dsturbance could, and n some cases
lkely would, occur at lower RLs.
In June 1997, the Hgh Energy Sesmc Survey
(HESS) team (1999, Appendx 5) convened a
panel of experts to assess nose exposure crtera
for marne mammals exposed to sesmc pulses.
The consensus was that, gven the best avalable
data at that tme, exposure to argun pulses wth
RLs above 180 dB re: 1 Pa (averaged over the
pulse duraton) was lkely to have the potental to
cause serous behavoral, physologcal, and hear-
ng effects. The panel noted the potental for 10
dB varablty around the 180 dB re: 1 Pa level,
dependng on speces, and that more nformaton
was needed.
The NMFS has contnued to use a do not
exceed exposure crteron of 180 dB re: 1 Pa for
mystcetes and (recently) all odontocetes exposed
to sequences of pulsed sounds, and a 190 dB re:
1 Pa crteron for pnnpeds exposed to such
sounds. Hgher thresholds have been used n the
U.S. for sngle pulses such as explosons used n
naval vessel-shock trals. Behavoral dsturbance
crtera for pulsed sounds have typcally been set
at an SPL value of 160 dB re: 1 Pa, based manly
on the earler observatons of mystcetes reactng
to argun pulses (e.g., Malme et al., 1983, 1984;
Rchardson et al., 1986). The relevance of the 160
dB re: 1 Pa dsturbance crteron for odontocetes
and pnnpeds exposed to pulsed sounds s not
at all well-establshed, however. Although these
crtera have been appled n varous regulatory
actons (prncpally n the U.S.) for more than a
decade, they reman controversal, have not been
appled consstently n the U.S., and have not been
wdely accepted elsewhere.
More recently, a consderable body of data has
accumulated on the levels at whch transent and
more prolonged sounds cause the onset of tempo-
rary threshold shft (TTS) and varous behavoral
reactons. Some of these data are not consstent
wth the aforementoned defacto crtera used n
recent years n the Unted States.
One man purpose of ths paper s to synthe-
sze and apply all avalable nformaton to derve
proposed objectve nose exposure crtera for a
large subset of marne mammals. The effect levels
consdered (njury and sgnfcant behavoral
dsturbance) were generally consstent wth the
defntons of levels A and B harassment, respec-
tvely, of the U.S. Marne Mammal Protecton Act
(MMPA) of 1972 (16 USC, 1361); however,
many of the behavors consdered at the lower end
of our severty scalng paradgm would almost
certanly not consttute bologcally sgnfcant
dsturbance (or consequently level B harassment
under the MMPA). However, our exposure crtera
were derved wthout regard for polcy decsons
of the U.S. or any naton and should therefore not
be assumed to correspond wth regulatory catego-
res or defntons of effects. Snce harassment
defntons under the MMPA are not unform for
all human actvtes and are subject to change,
addtonal nterpretaton of the nformaton pre-
sented would be requred to evaluate effects wth
regard to ths (or any other) statute.
Acoustic Measures and Terminology
Ths secton brefly consders those acoustc mea-
sures and termnology that are drectly relevant
to these marne mammal exposure crtera. More
detaled descrptons of some of the terms gven
n ths and other sectons, ncludng equatons
relevant to many of the defntons, are gven n
Appendx A. Basc acoustc termnology s pre-
sented n numerous other sources (e.g., Knsler
et al., 1982; ANSI, 1986, 1994; Rchardson et al.,
1995; Harrs, 1998; NRC, 2003).
Sound s approprately descrbed as havng two
components: (1) a pressure component and (2) a
partcle moton component. Partcle motonthe
oscllatory dsplacement, velocty, or acceleraton
of the actual partcles of the medum at a par-
tcular locatons drectonal and best descrbed
by a 3-dmensonal vector. Marne mammal sen-
stvty to partcle moton s poorly understood,
but t appears to be functonally lmted (Fnneran
et al., 2002a) n contrast to the sensory capabl-
tes of most or all fsh (see Popper et al., 2003).
Conversely, as compared to fsh, marne mammals
generally have greater senstvty to sound pres-
sure (lower detecton thresholds) and much wder
functonal hearng bandwdths (see Fay, 1988;
Rchardson et al., 1995; Popper et al., 2003).
Consequently, n consderng the potental effects
of sound on marne mammals, partcle moton s
rarely dscussed. Except for specal crcumstances
(e.g., plane and sphercal waves), there s no
smple relatonshp between pressure and partcle
velocty. The vast majorty of studes of hearng
n captve marne mammals have been conducted
n relatvely small enclosed volumes of water,
makng the plane wave assumpton (and a priori
knowledge of the relatonshp between pressure
and velocty) nvald.
It s mportant to dstngush between the source
level (SL), or level measured 1 m from the source,
vs the receivedlevel (RL), whch s the level mea-
sured at the recever (usually a marne mammal
heren).
418 Southalletal.
The term ntensty s often used generally
wth respect to subjectve acoustc parameters
(.e., loudness), but t s used here n a strct sense.
Sound ntensty s normally defned as the tme-
averaged actve ntensty (Knsler et al., 1982;
Fahy, 1995); ths quantty corresponds to local
net transport of sound energy and s related to
the product of the sound pressure and the partcle
velocty component n-phase wth the sound pres-
sure. In the majorty of laboratory studes, complex
sound felds typcally create complex, spatally
varyng relatonshps between pressure and veloc-
ty. In these crcumstances, sound ntensty cannot
be estmated from pressure measurements alone
(whch assume that pressure and partcle velocty
are n-phase), and specfc measurements of the
sound partcle velocty (or pressure gradent) are
requred n order to characterze ntensty.
We dstngushed two basc sound types:
(1) pulse and (2) nonpulse. Our operatonal def-
ntons of sound types are gven n Chapter 2,
Structure of the Nose Exposure Crtera, and
are dscussed at greater length n Appendx A. The
pulse/nonpulse dstncton s mportant because
pulses generally have a dfferent potental to cause
physcal effects, partcularly on hearng (e.g.,
Ward, 1997).
Peak sound pressure (Pmax) s the maxmum
absolute value of the nstantaneous sound pressure
durng a specfed tme nterval and s denoted n
unts of Pascals (Pa). It s n no sense an averaged
pressure. Peak pressure s a useful metrc for ether
pulse or nonpulse sounds, but t s partcularly
mportant for characterzng pulses (ANSI, 1986;
Harrs, 1998, Chapter 12). Peak-to-peak sound
pressure s the algebrac dfference between the
maxmum postve and maxmum negatve nstan-
taneous peak pressure. The mean-squared pres-
sure s the average of the squared pressure over
some duraton. Sound pressure levels are gven as
the decbel (dB) measures of the pressure metrcs
defned above. The RMS SPL s gven as dB re:
1 Pa for underwater sound and dB re: 20 Pa
for aeral sound. Peak sound pressure levels are
denoted hereafter as dB re: 1 Pa (peak) n water
and dB re: 20 Pa (peak) n ar. Peak-to-peak
sound pressure levels are dB re: 1 Pa (peak-to-
peak) n water and dB re: 20 Pa (peak-to-peak)
n ar.
Duration s the length of a sound n seconds.
Duraton s mportant because t affects other
sound measures, specfcally mean-square and/or
RMS sound pressure (Madsen, 2005). Because of
background nose and reverberaton, duraton can
be dffcult to specfy precsely, but a functonal
defnton (see Appendx A) s used here.
Sound exposure level (SEL) s a measure of
energy. Specfcally, t s the dB level of the
tme ntegral of the squared-nstantaneous sound
pressure normalzed to a 1-s perod. It can be an
extremely useful metrc for assessng cumulatve
exposure because t enables sounds of dfferng
duraton, sometmes nvolvng multple expo-
sures, to be compared n terms of total energy.
Several methods exst for summng energy over
multple exposures to generate a sngle exposure
equvalent value. The relatvely straghtforward
approach used here s descrbed n Appendx A (eq.
5). Ths summaton procedure essentally generates
a sngle exposure equvalent value that assumes
no recovery of hearng between repeated expo-
sures. As dscussed below, recovery functons for
marne mammal TTS durng and followng mult-
ple exposures are stll unknown; however, consd-
erng nomnal TTS recovery functons n terrestral
mammals when exposures occur mnutes to hours
apart (see Kryter, 1994; Ward, 1997), the above
summaton procedure would lkely overestmate
the effect of multple exposures n many cond-
tons. Ths summaton procedure was ntentonally
selected as a precautonary measure n the absence
of emprcal nformaton, although note the tem-
poral condtons gven n the Sound Types sec-
ton of Chapter 2. The approprate unts are dB re:
1 Pa
2
-s for underwater SEL and dB re: (20 Pa)
2
-s
for aeral SEL.
Frequency-selectiveweighting s often employed
to measure (as a sngle number) sound pressure or
energy n a specfc frequency band of sound, wth
emphass or de-emphass on partcular frequences
as a functon of the relatve senstvty of a recever.
For aeral hearng n humans, A-weghtng s derved
from the nverse of the dealzed 40-phon equal
loudness hearng functon across frequences, stan-
dardzed to 0 dB at 1 kHz (Harrs, 1998). Ths pro-
vdes level measures denoted as dB(A). C-weght-
ng s determned from the nverse of the dealzed
100-phon equal loudness hearng functon (whch
dffers n several regards from the 40-phon func-
ton), standardzed to 0 dB at 1 kHz (Harrs, 1998).
Ths provdes level measures denoted as dB(C). In
the absence of equal-loudness contours for marne
mammals, specal frequency-weghtng functons
based loosely on human C-weghtng and general
knowledge of functonal hearng bandwdth were
developed here for functonal marne mammal hear-
ng groups (see the Marne Mammal Functonal
Hearng Groups secton of Chapter 2).
Other measures of nose nterference wth
crtcal functons n humans, ncludng the
Artculaton Index (French & Stenberg, 1947)
and the more recent Speech Interference Level
(see Beranek & Ver, 1992), focused on the percep-
ton of speech and effects of nose. Consequently,
exposure crtera geared toward speech percep-
ton (e.g., Beranek, 1989) focus on a frequency
MarineMammalNoiseExposureCriteria 419
bandwdth narrower than the audble bandwdth.
For a detaled dscusson of speech ntellgblty
and nose mpacts, see Chapter 6 n Kryter (1994).
It s clear that the percepton of conspecfc vocal
sgnals n marne mammals s crtcally mportant
n varous lfe hstory functons (dscussed below;
see Wartzok & Ketten, 1999) and that nterference
wth these functons may have partcularly nega-
tve consequences.
The hypothess that vocalzatons concde
wth the range of hearng s based on an adaptve
argument that vocal energy should be selected to
le wthn the range of hearng for maxmum eff-
cency of communcaton. However, several lnes
of evdence suggest that other adaptve pressures
may shape the vocal range. Frst, vocal anatomy
may produce energy at other frequences as a
byproduct of producng sound wthn the hearng
range. If there s no pressure to elmnate these
frequences, they can be expected to persst. An
example s the ultrasonc components of hummng-
brd song, whch le well outsde the range of brd
hearng (Pytte et al., 2004). Second, to promote
long-range transmsson, the vocal range may be
adapted to produce greater energy at the low end
of the range than would be expected based on the
audtory threshold functon (Larom et al., 1997).
Greater relatve energy at low frequences s also
seen n a number of prmate speces as a byprod-
uct of producng the formant structure of ther calls
(Ftch & Hauser, 1995). Fnally, anmals may pro-
duce sounds wth dsproportonate low-frequency
nformaton to sgnal greater sze, potentally tar-
getng predators rather than conspecfcs (Ftch,
1999; Matrosova et al., 2007). Thus, a number of
selectve forces can drve the development of an
emphass on low-frequency energy n vocalzatons
not matched by the shape of the audtory threshold
functon. Whle vocal range can be expected to cor-
relate wth hearng range to some degree, gvng a
rough ndcaton of the frequency range of hearng,
t cannot be used to estmate ether the shape of the
audtory threshold functon or to assgn upper and
lower frequency lmts.
We lack suffcent emprcal data on whether
vocal frequency range suffcently predcts all
frequences that are bologcally sgnfcant,
however.
Certan marne mammal responses to anthropo-
genc sounds, such as the sometmes strong reac-
tons by beaked whales to md-frequency sonar,
would not be expected f only sounds wthn the
bandwdth of vocal output were mportant n pre-
dctng a behavoral response. Hence, our precau-
tonary frequency-weghtng approach assumes
that the full audble band s relevant. As addtonal
data become avalable on both hearng capa-
bltes (specfcally, equal-loudness contours)
and behavoral responses to natural (ncludng
conspecfc) and anthropogenc sounds, a more
refned means of frequency-weghtng than the
ntentonally precautonary (broad) M-weghtng
functons may be recommended.
Kurtosis s a statstcal measure of a probablty
dstrbuton often appled to descrbe the shape of
the ampltude dstrbuton (Hamernk & Hsueh,
1991; Le et al., 1994; Hamernk et al., 2003). In
some regards, t appears to be a hghly relevant
metrc n that mpulsve sound wth hgh nega-
tve kurtoss, rapd onset, and hgh nstantaneous
peak-pressure may be partcularly njurous to
some mammals (Hamernk et al., 2003).
Sound Production and Use in Marine Mammals
As a general statement, all studed marne mam-
mals can produce sounds n varous mportant con-
texts. They use sound n socal nteractons as well
as to forage, to orent, and to respond to predators.
Interference wth these functons, through the var-
ous effects of nose on hearng and/or behavor
dentfed below, thus has the potental to nterfere
wth vtal rates dentfed by the NRC (2005) as
partcularly sgnfcant effects of exposure.
The nose exposure crtera gven here are
focused on current knowledge of hearng and
the effects of nose on hearng and/or behavor n
marne mammals. Thus, a detaled dscusson and
revew of the expansve lterature on the produc-
ton and the uses of sound s beyond the scope of
ths paper; nterested readers are referred to the
many revews of marne mammal acoustc sgnals
(e.g., Schusterman, 1981; Watkns & Wartzok,
1985; Au, 1993; Rchardson et al., 1995; Wartzok
& Ketten, 1999; Clark & Ellson, 2004). Because
of the extreme mportance of detectng conspecfc
socal sgnals n marne mammal lfe hstory func-
tons, however, a bref and very general dscus-
son of sound output characterstcs n the major
marne mammal groups s gven here.
The large whales (mystcete cetaceans, as
descrbed below) generally produce low-fre-
quency sounds n the tens of Hz to the several kHz
band, wth a few sgnals extendng above 10 kHz.
These sounds appear to serve predomnantly socal
functons, ncludng reproducton and mantanng
contact, but they may also play some role n spa-
tal orentaton.
The dolphns and porposes (odontocete ceta-
ceans, also descrbed below) produce sounds
across some of the wdest frequency bands that
have been observed n anmals. Ther socal sounds
are generally n the range audble to humans, from
a few hundreds of Hz to several tens of kHz, but
specalzed clcks used n bosonar (echolocaton)
420 Southalletal.
systems for prey detecton and navgaton extend
well above 100 kHz.
Pnnpeds (seals, sea lons, and walruses) also
produce a dversty of sounds, though generally
over a lower and more restrcted bandwdth (gen-
erally from 100 Hz to several tens of kHz). Ther
sounds are used prmarly n crtcal socal and
reproductve nteractons. Pnnpeds spend tme
both at sea and on land, however, and thus pro-
duce sounds n both water and ar.
Because sound producton n marne mam-
mals s ntegral to so many mportant behavors,
nterference wth these communcatve functons
s consdered to be partcularly adverse (see sever-
ty scalng descrbed n Chapter 4, Crtera for
Behavoral Dsturbance). As dscussed n Chapter
5, consderable addtonal research s needed to
dentfy condtons n whch anthropogenc nose
exposure nterferes wth acoustc communcaton
as well as ways n whch marne mammals cope
wth maskng nose to overcome nterference n
detectng real-world sgnals n complex, 3-dmen-
sonal marne envronments.
Responses to Sound
Anmals exposed to ether natural or anthropo-
genc sound may experence physcal and psycho-
logcal effects, rangng n magntude from none to
severe. Ths bref dscusson consders the range
of potental mpacts, whch depend on spatal rela-
tonshps between a sound source and the anmal
recever; senstvty of the recever; receved expo-
sure level, duraton, and duty cycle; and many
other factors (see also Rchardson et al., 1995).
The same acoustc source may have radcally
dfferent effects dependng on operatonal and
envronmental varables, and on the physologcal,
sensory, and psychologcal characterstcs of
exposed anmals. It s mportant to note that these
anmal varables may dffer (greatly n some cases)
among ndvduals of a speces and even wthn
ndvduals dependng on varous factors (e.g.,
sex, age, prevous hstory of exposure, season, and
anmal actvty). Responses elcted can depend
both on the context (feedng, matng, mgratng,
etc.) n whch an ndvdual s ensonfed and
on a host of experental varables (see Wartzok
et al., 2004). Consequently, certan effects may
be poorly descrbed wth smple measures such
as SPL alone, and may only be predctable when
addtonal varables are consdered. We consd-
ered all known factors n developng the nose
exposure crtera proposed here, but data lmta-
tons precluded the dervaton of explct exposure
crtera for all of the effects dscussed below.
Audibility
When a sound can be perceved amdst background
nose, t s consdered to be audble. Audblty can
dffer from detectablty n that a recevng system
may detect a sgnal at some level even when t s
ncapable of meanngful percepton. Audblty
s determned by the characterstcs of receved
sound, characterstcs of the recevng system, and
background nose condtons (ether external or
nternal). Audton (hearng) s a well-developed
and prmary sensory modalty for most, f not all,
marne vertebrates (Schusterman, 1981; Tyack,
1998; Fay & Popper, 2000). It nvolves codng,
processng, ntegratng, and respondng to sound
n a varety of ways, some not outwardly evdent
(Yost, 2000). Lke other anmals, marne mam-
mals have multple sound-recepton pathways and
rely on sgnal processng at multple levels nte-
grated wthn the cochlea and nervous system to
optmze percepton.
Marne mammal hearng capabltes are
quantfed n lve subjects usng behavoral aud-
ometry and/or electrophysologcal technques
(e.g., Schusterman, 1981; Au, 1993; Kastak &
Schusterman, 1998; Wartzok & Ketten, 1999;
Nachtgall et al., 2000, 2007; Fnneran & Houser,
2006; Andr & Nachtgall, 2007; Supn & Popov,
2007). For speces not studed wth in vivo aud-
ometry, some audtory characterstcs can be est-
mated based on sound producton frequences; on
observatons of sound characterstcs that ether do
or do not elct behavoral responses n untraned
anmals (e.g., Rchardson et al., 1995; Erbe, 2002);
or on audtory morphology, ncludng bomechan-
cal propertes of the baslar membrane and other
characterstcs (Wartzok & Ketten, 1999).
Behavioralaudiograms are obtaned from cap-
tve, traned anmals usng standard psychometrc
testng procedures. Wth approprate controls and
suffcent tranng, behavoral data are presently
consdered to most accurately represent hearng
capabltes of a test subject. Behavoral audo-
metrc studes are tme-consumng, however, and
the results depend on the tranng and attenton of
subjects as well as the background nose cond-
tons n captve settngs. Because marne mam-
mals are large and dffcult to mantan, behav-
oral audograms representng an entre speces
are typcally based on a few ndvduals (often
one anmal). Addtonally, subjects are generally
obtaned opportunstcally (e.g., ndvduals reha-
bltated after strandng) rather than by random
samplng of ndvduals from wld populatons.
Ths may provde a somewhat based representa-
ton of normal hearng for the speces f reha-
bltated anmals have compromsed hearng
capabltes (see Andr et al., 2007). Indvdual
dfferences n hearng senstvty among subjects,
MarineMammalNoiseExposureCriteria 421
and methodologcal dfferences among nvestga-
tors, can lead to mproper conclusons when nom-
nal speces audograms are based on data from
a sngle anmal (e.g., compare Hall & Johnson,
1972, wth Szymansk et al., 1999). Hearng sen-
stvty has been measured usng behavoral meth-
ods n fewer than 20 of the ~128 cetacean and
pnnped speces (based on the taxonomy of Rce,
1998).
Electrophysiologicalaudiometry nvolves mea-
surng small electrcal voltages (audtory evoked
potentals [AEPs]) produced by neural actvty
when the audtory system s stmulated by sound.
Wth ths technque, neural responses are typ-
cally averaged whle many relatvely short dura-
ton sgnals are presented. Ths technque s com-
paratvely fast and less senstve to factors such as
subject experence and reproductve, behavoral,
or motvatonal states that affect behavoral aud-
ometry. Whereas behavoral audograms can only
be made wth traned, captve anmals, AEP mea-
sures of sound detecton can also be made wth
untraned ndvduals that are stranded, tempo-
rarly restraned, or n rehabltaton (see Cook
et al., 2006; Andr et al., 2007; Delory et al., 2007;
Taylor et al., 2007).
AEP and behavoral technques measure dffer-
ent features of the audtory system and may gener-
ate somewhat dfferent measured results. Relevant
comparsons of AEP and behavoral audograms
are lmted and are the subject of ongong scen-
tfc nvestgaton. Besdes the need to obtan both
types of data on the same ndvduals, there are
complcatons due to dfferences n the types of
test stmul used by dfferent researchers, prob-
lems n estmatng the true RL at the relevant
sensory organ(s), and the dffculty of determn-
ng absolute sgnal ampltudes that barely elct
neural responses. Even so, Yuen et al. (2005),
Fnneran et al. (2007b), and Schlundt et al. (2007)
demonstrated that, wth carefully calbrated and
repeated measurements, the two procedures can
produce comparable detecton thresholds n at
least a few cetacean speces.
An audtory threshold, estmated by ether
behavoral or electrophysologcal responses, s
the level of the quetest sound audble n a spec-
fed percent of trals. An audtory threshold s not
an nvarant crtcal value above whch a sound s
always heard and below whch t s never heard.
Instead, t s a sound level at whch there s an
explct sgnal detecton probablty (often 50%;
determned a priori). Ths probablty depends
on a number of ntrnsc factors (Green & Swets,
1974; Egan, 1975; McMllan & Creelman, 1991).
In all speces tested thus far, the hearng response
n relaton to frequency s a generally U-shaped
curve wth a frequency range of best senstvty
(lowest hearng thresholds) and frequences both
below and above ths range where senstvty s
relatvely poor (hgher threshold values). Speces
dffer n absolute senstvty and functonal fre-
quency bandwdth (see Fay, 1988; Rchardson
et al., 1995), such that dentcal sounds may be
perceved radcally dfferently by ndvduals of
dfferent speces. Indvdual dfferences wthn
speces have also been demonstrated n some ter-
restral speces (see Fay, 1988) and, to a lesser
extent, n marne mammals as well (see Houser &
Fnneran, 2006b, for the most defntve example
of ths). Sounds whose levels barely exceed back-
ground nose levels may be detectable but may
or may not elct changes n ndvdual behavor.
Ideally, absolute or unmasked hearng thresh-
olds should be measured n low background nose
condtons such as anechoc testng enclosures.
Whle ths s standard practce n human aud-
ometry, very few of the marne mammal hearng
data obtaned to date have been measured n such
condtons. Lmted recent data obtaned wth pn-
npeds tested n a hem-anechoc testng chamber
n ar (descrbed n Kastak et al., 2005) suggest
that maskng from envronmental nose n testng
enclosures may have sgnfcantly affected mea-
surements of absolute hearng; thresholds n a
harbor seal (Phocavitulina) were n fact 30 dB
lower n very low background nose condtons
(Holt et al., 2001).
Whle the above concepts and studes are essen-
tal n understandng general hearng capabltes
(e.g., functonal bandwdth, range of best hearng
senstvty) of marne mammals, anmals n the
real world rarely lsten for smple acoustc sg-
nals from pont sources and do not lve n a nose-
controlled envronment. Rather, they are presented
wth spatally complex and tme-varyng streams
of acoustc nformaton n often nosy envron-
ments. Measurements usng smple sound stmul
have ndcated that marne mammals are generally
qute adept at localzng acoustc sources n labo-
ratory condtons (Mhl, 1964; Gentry, 1967;
Terhune, 1974; Moore & Au, 1975; Renaud &
Popper, 1975; Holt et al., 2004, 2005). Many of
the behavoral observatons dscussed n Chapter
4 (and n Appendces B & C) ndcated relatvely
precse orentaton behavors to sound sources (or
sound localzaton) n the feld as well. Lmted
laboratory data are also avalable regardng how
marne mammals detect relatvely smple stm-
ul over background maskng nose (dscussed
below). A more complex perceptual matter related
to localzaton and detecton over maskng nose
s the manner n whch vertebrates process com-
plex nformaton to perceve the acoustc (or aud-
tory) scenethat s, gan useful nformaton from
422 Southalletal.
the sute of sounds around them n the real world
(e.g., Fay & Popper, 2000).
Bregman (1990) consdered how the human
audtory system constructs a perceptual acoustc
mage of the surroundng envronment and events
occurrng n that envronment. He posts that, as n
vsual percepton, hearng systems are organzed
n such a manner that related acoustc events (such
as the frequency structure of a harmonc sgnal or
a repeated sgnal from the same source n a 3-
dmensonal space) are grouped perceptually n
a meanngful way. Accordng to the process of
auditorysceneanalysis, the audtory system sorts-
out related elements of a complex natural acous-
tc envronment nto those arsng from dfferent
sound sources. Furthermore, prevous experence
can have powerful effects on the processng and
nterpretaton of sounds. Ths too s smlar to psy-
chologcal processes underlyng vsual percepton
n whch the range to an object may be nferred
from knowledge of an objects general sze and
physcal appearance.
Presumng such capabltes occur n marne
vertebrates, whch s logcal gven the mportance
of sound to marne mammals, t seems lkely that
they could perceve range and the general nature
(e.g., movement) of sound sources. Acoustic
streamsegregation, the dentfcaton of relatvely
smple stmul from dfferent, overlappng patterns,
has been demonstrated n several brd and bat spe-
ces (MacDougall-Shackleton et al., 1998; Moss
& Surlykke, 2001). Nether acoustc stream seg-
regaton nor audtory scene analyss has yet been
nvestgated n marne mammals (but see Madsen
et al., 2005a). Each of these processes, along wth
more data on sound localzaton, may be relevant n
the contnued development of approprate marne
mammal nose exposure crtera (see the Marne
Mammal Functonal Hearng Groups secton of
Chapter 5, for research recommendatons).
AuditoryMasking
Nose may partally or entrely reduce the aud-
blty of sgnals, a process known as auditory
masking. The extent of nterference depends on
the spectral, temporal, and spatal relatonshps
between sgnals and maskng nose, n addton
to other factors. Human audtory systems per-
form frequency-based assessment (smlar to
Fourer analyss) on ncomng sgnals such that,
for most exposure levels, sgnfcant maskng of
tonal sgnals s almost exclusvely by nose n a
narrow band (called the crtcal band) of smlar
frequences (Wegel & Lane, 1924; Fletcher, 1940;
Greenwood, 1961). Wth ncreasng masker level,
however, there s an asymmetrcal spread n the
maskng effect such that detecton of frequences
above those of the maskng stmulus s more sg-
nfcantly mpeded (see Buus, 1997; Yost, 2000).
Because of common bomechancal cochlear
propertes across taxa (Echteler et al., 1994),
maskng s expected to follow smlar prncples n
other mammals (ncludng marne mammals). The
structure and functon of the outer and mddle ear
dffer profoundly between terrestral and marne
mammals (Wartzok & Ketten, 1999); however,
the characterstcs of audtory maskng are strk-
ngly smlar among nonspecalzed mammals n
general (Fay, 1988; Echteler et al., 1994), nclud-
ng marne mammals tested n ar and n water
(Turnbull & Terhune, 1990; Southall et al., 2000,
2003). Smlartes n morphology and mamma-
lan cochlear functonal dynamcs (as revealed by
maskng studes) suggest that audtory data from
terrestral mammals may be relably used n some
stuatons where marne mammal data are lackng.
Data on audtory maskng n marne mammals are
not presented n detal here because they are not
drectly used n formulatng the recommended
nose exposure crtera (but see Southall et al.,
2000, 2003, for revews).
AuditoryThresholdShift
Anmals exposed to suffcently ntense sound
exhbt an ncreased hearng threshold (.e., poorer
senstvty) for some perod of tme followng
exposure; ths s called a noise-induced thresh-
old shift (TS). Factors that nfluence the amount
of TS nclude the ampltude, duraton, frequency
content, temporal pattern, and energy dstrbuton
of nose exposure. The magntude of TS normally
decreases over tme followng cessaton of the
nose exposure. The amount of TS just after expo-
sure s called the ntal TS.
If TS eventually returns to zero (.e., the thresh-
old returns to the pre-exposure value), t s called
TTS. The followng physologcal mechansms
are thought to play some role n nducng TTS,
also referred to as audtory fatgue: effects on sen-
sory har cells n the nner ear that reduce ther
senstvty, modfcaton of the chemcal envron-
ment wthn sensory cells, resdual mddle-ear
muscular actvty, dsplacement of certan nner
ear membranes, ncreased blood flow, and post-
stmulatory reducton n both efferent and sensory
neural output (Kryter, 1994; Ward, 1997). Where
these effects result n TTS rather than a permanent
change n hearng senstvty, they are wthn the
nomnal bounds of physologcal varablty and
tolerance and do not represent physcal njury
(Ward, 1997). Recovery of nomnal hearng func-
ton may occur quckly, and the amount of TTS
measured depends on the tme elapsed snce the
cessaton of nose exposure; subscrpts are used
to ndcate the tme n mnutes after exposure. For
MarineMammalNoiseExposureCriteria 423
example, TTS2 means TTS measured 2 mn after
exposure cessaton.
If TS does not return to zero after a relatvely
long nterval (on the order of weeks), the resdual
TS s called a nose-nduced permanent threshold
shft (PTS). The dstncton between PTS and TTS
depends on whether there s a complete recovery
of TS followng nose exposure. PTS s consdered
to be audtory njury. Some of the apparent causes
of PTS n mammals are severe extensons of
effects underlyng TTS (e.g., rreparable damage
to the sensory har cells). Others nvolve dfferent
mechansms, such as exceedng the elastc lmts
of certan tssues and membranes n the mddle
and nner ears and resultant changes n the chem-
cal composton of nner ear fluds (Ward, 1997;
Yost, 2000). The relatonshp between TTS and
PTS depends on a hghly complex sute of var-
ables concernng the study subject and the expo-
sure. Ths relatonshp remans poorly understood,
even for humans and small terrestral mammals n
whch ths topc has been nvestgated ntensvely
(see Kryter, 1994; Yost, 2000).
In addton to the potental for dscrete, ntense
sounds to result n TTS or PTS, chronc sound
exposure, common n ndustralzed socetes, can
result n nose-nduced PTS n humans as they age
(see Kryter, 1994). Reduced hearng senstvty
as a smple functon of development and agng
(presbycusis) has been demonstrated n both chl-
dren (Roche et al., 1978) and adults (e.g., Brant
& Fozard, 1990). In the long-term, nose-nduced
hearng loss and presbycuss appear to result n
a progressve PTS that s a complex, nonlnear
process and partcularly affects hgh-frequency
hearng. Lmted research n cetaceans and pn-
npeds has revealed patterns of presbycuss that
are smlar to those observed n humans (Rdgway
& Carder, 1997; Brll et al., 2001; Schusterman
et al., 2002; Houser & Fnneran, 2006b; Rechmuth
et al., 2007), further underscorng certan gen-
eral smlartes n audtory processes across
mammals.
PTS and TTS data from humans and non-
human terrestral mammals were used to develop
safe exposure gudelnes for human work envron-
ments (e.g., NIOSH, 1998). For marne mammals,
recent data are avalable regardng sounds that
cause modest TTS (generally < 20 dB decrease
n senstvty) n a few speces of odontocetes and
pnnpeds. No data exst on exposures that would
cause PTS n these taxa, however (see Chapter 2
for detaled dscussons). Consequently, the only
current opton for estmatng exposure condtons
that would cause PTS-onset n marne mammals
s to use the avalable marne mammal TTS data
combned wth data from terrestral mammals
on TTS growth rates wth ncreasng acoustc
exposure (see the Crtera for Injury: TTS and
PTS secton of Chapter 3).
BehavioralReactionstoSound
Behavoral responses to sound are hghly varable
and context-specfc (see Wartzok et al., 2004, for
a dscusson). Some sounds that are audble to an-
mals may elct no overt behavoral response. Ths
s most common when the sound does not greatly
exceed the mnmum detectable level and s not
ncreasng or fluctuatng (Rchardson et al., 1995).
Inablty to detect an overt response does not nec-
essarly mean that there s no subtle behavoral (or
other) effect, however.
When observable reactons do occur, they may
nclude orentaton or attracton to a sound source;
ncreased alertness; modfcaton of characterstcs
of ther own sounds; cessaton of feedng or socal
nteracton; alteraton of movement/dvng behav-
or; temporary or permanent habtat abandonment;
and, n severe cases, panc, flght, stampede, or
strandng, sometmes resultng n njury or death
(e.g., Rchardson et al., 1995; Evans & England,
2001; Gordon et al., 2004; Schefele et al., 2005;
Cox et al., 2006; Nowacek et al., 2007). Mnor
or temporary behavoral effects are often smply
evdence that an anmal has heard a sound and
may not ndcate lastng consequence for exposed
ndvduals. For the purposes of settng crte-
ra, the effects of greatest concern are those that
may negatvely mpact reproducton or survval.
Ultmately, t s the bologcal relevance of the
reacton n terms of vtal parameters that must be
determned. In proposng nose exposure crtera,
one must clearly and explctly dfferentate trv-
al effects from those wth the potental to affect
vtal rates. However, t has proven to be exceed-
ngly challengng to dstngush among and rank
the varous effects and to establsh a generally
accepted defnton of bologcally meanngful
behavoral dsturbance (see NRC, 2005).
Except for nave ndvduals, behavoral
responses depend crtcally on the prncples
of habituation and sensitization. An anmals
exposure hstory wth a partcular sound affects
whether t s subsequently less lkely (habtua-
ton) or more lkely (senstzaton) to respond to
a stmulus such as sound exposure. The processes
of habtuaton and senstzaton do not necessar-
ly requre an assocaton wth a partcular adverse
or bengn outcome. Rather, ndvduals may be
nnately predsposed to respond to certan stmul
n certan ways. These responses may nteract
wth the processes of habtuaton and senstza-
ton for subsequent exposure. Where assocatve
learnng occurs, ndvduals lnk a partcular expo-
sure wth a known outcome (postve, negatve,
or neutral) and use that nformaton n gudng
424 Southalletal.
future decsons on whether and how to respond
to smlar stmul. The relatonshp between these
two categores of learnng (non-assocatve and
assocatve) can be hghly complex, partcularly
for experenced ndvduals (see Deecke et al.,
2002).
Many contextual varables may be power-
ful contrbutors to an anmals percepton of and
reacton to the acoustc scene. These nclude the
percepton of source proxmty (nearness), relatve
movement (encroachment or retreat), and general
novelty or famlarty, all of whch may affect the
type and magntude of the resultng behavoral
response(s). In terms of proxmty, the presence
of hgh-frequency components n a sound and the
lack of reverberaton, both of whch are ndca-
tve of proxmty, may be more relevant acoustc
cues of spatal relatonshp than smply exposure
level alone (see P. Mller, 2002). If a source s per-
ceved to be approachng, the response s often
stronger. In addton, the actvty of the ndvdual
and ts fdelty to a current locaton often affect
the response.
Thus, n addton to source characterstcs,
other factors that may be crtcal n determnng
behavoral effects nclude past experence, stu-
atonal varables, recever audtory systems, and
the extent to whch the sound resembles famlar
bengn or noxous stmul (e.g., Irvne et al., 1981;
NRC, 2005). Anmals that fal to exhbt general
avodance when exposed to a certan sound source
may stll detect the sound but are ether habtuated
to exposure or may dsplay less dramatc behav-
oral responses (e.g., alterng vocal behavor,
modfyng orentaton/movement patterns).
The magntude of a gven behavoral response
may not be a drect functon of exposure levels
or even of the anmals experental hstory. If
the sound trggers an ant-predator response
n the subject (e.g., Irvne et al., 1981; Fnley
et al., 1990), the response magntude may reflect
the ndvduals underlyng physologcal con-
dton, the relatve costs n ftness of falng to
respond, the avalablty of alternatve refuges,
and other factors specfc to predator defense (Gll
& Sutherland, 2000; Frd & Dll, 2002; Beale &
Monaghan, 2004).
For all these reasons, behavoral responses
to anthropogenc sounds are hghly varable.
Meanngful nterpretaton of behavoral response
data (and bologcally relevant conservaton dec-
sons) must consder not only the relatve mag-
ntude and apparent severty of behavoral reac-
tons to human dsturbance but also the relevant
acoustc, contextual, and ecologcal varables.
In many cases, specfc acoustc features of the
sound and contextual varables (e.g., proxmty,
subject experence and motvaton, duraton, or
recurrence of exposure) may be of consderably
greater relevance to the behavoral response than
smple acoustc varables such as exposure RL.
For example, f an anthropogenc sound s per-
ceved as ndcatng the presence of a predator,
t s lkely to trgger a strong defensve reacton
at relatvely low RLs. On the other hand, sounds
that resemble conspecfc sgnals may be gnored
or nduce approach or avodance, dependng upon
the context. Further, typcally neutral sounds may
cause ncreasng annoyance reactons (such as
avodance) as a functon of exposure level. Ths
makes t dffcult or mpossble to justfy basng
broad, objectve determnatons of mpact thresh-
olds on RL alone. Ths s the prmary reason why
ths paper does not propose explct behavoral
dsturbance crtera levels for certan sound types.
Rather, we collated avalable data relatng acous-
tc exposure to the severty of observed behav-
oral response n a form that allows a varety of
relatonshps to be estmated (Chapter 4). When
research allows the separaton of annoyance from
cases where an anmal nterprets sounds as sg-
nals from predators, prey, or conspecfcs, t may
become possble to classfy sgnals and predct
responses more precsely.
Non-AuditoryEffects
The audtory system appears to nclude the organs
most susceptble to nose exposure, at least n
humans (e.g., Ward, 1997). The lmted data on
captve marne mammals exposed to varous
knds of nose support a smlar concluson, sug-
gestng that TTS-onset occurs at levels whch may
be below those requred for drect non-audtory
physologcal trauma (but see dscusson of deep-
dvng speces below). Nose exposure does have
the potental to nduce a range of drect or nd-
rect physologcal effects on non-audtory struc-
tures. These may nteract wth or cause certan
behavoral or audtory effects, or they may occur
entrely n the absence of those effects.
Nose exposure may affect the vestbular and
neurosensory systems. For nstance, n humans,
dzzness and vertgo can result from exposure
to hgh levels of nose, a condton known as nys-
tagmus (see Oosterveld et al., 1982; Ward, 1997;
Halmagy et al., 2005). Lttle s known about ves-
tbular functons n marne mammals. There are
sgnfcant dfferences n vestbular structures n
some marne mammal speces compared to most
land mammals (Wartzok & Ketten, 1998; Ketten,
2000). In cetaceans n partcular, the vestbular
components are suffcently reduced and have
such low neural representaton that the prnc-
pal functon may be essentally to provde lm-
ted gravtatonal and lnear acceleraton cues.
Pnnpeds by contrast have a well-developed,
MarineMammalNoiseExposureCriteria 425
more conventonal vestbular apparatus that
lkely provdes multple sensory cues smlar to
those of most land mammals. Both pnnpeds and
cetaceans retan the drect couplng through the
vestbule of the vestbular and audtory systems;
therefore, t s possble, albet not known, that
marne mammals may be subject to nose-nduced
effects on vestbular functon as has been shown
n land mammals and humans. Responses to
underwater sound exposures n human dvers and
other mmersed land mammals suggest that ves-
tbular effects are produced from ntense under-
water sound at some lower frequences (Steevens
et al., 1997). Theoretcal effects on the human ves-
tbular system as well as other organs (e.g., lungs)
from underwater sound exposures also have been
explored through models (Cudahy & Ellson,
2002); however, there are no comparable mea-
surements or models for marne mammals at ths
pont from whch to estmate such effects. Data
are clearly needed for all major marne mammal
taxa to more fully assess potental mpacts on non-
audtory systems.
Relatvely low-level physologcal responses
nclude changes n cardac rate (bradycardia or
tachycardia) and respratory patterns, whch may
lead to changes n metabolsm. Stress reactons
n humans and other vertebrates nclude varous
physologcal changes to pulmonary, cardac,
metabolc, neuro-endocrne, mmune, and repro-
ductve functons (e.g., Hales, 1973; Lee, 1992;
Vrjkotte et al., 2000). Studes of nose-nduced
stress n marne mammals are very lmted, but
endocrne secretons of glucocortcods and altered
cardovascular functon have been documented n
odontocetes exposed to hgh-level sound (Romano
et al., 2004; cf. Thomas et al., 1990c). Nose expo-
sure also often leads to changes n surfacng-res-
praton-dve cycles of cetaceans (e.g., Rchardson
& Malme, 1993), whch may have varous phys-
ologcal effects. Assumng that effects n marne
and terrestral mammals are smlar, ntermedate
physologcal responses to stressors (ncludng
nose) may accompany avodance or aggres-
sve behavors and nclude sngle audtory star-
tle responses, the ntaton and sustenance of
the catecholamne response, and physologcal
preparaton for fght or flght. The most severe
physologcal responses would nclude multple
or repeated audtory startle responses, trgger-
ng of the hypothalamc-ptutary-adrenal (HPA)
axs and assocated elevated blood glucocortcod
level, substantally altered metabolsm or energy
reserves, lowered mmune response, dmnshed
reproductve effort, and potental tssue trauma
(e.g., Sapolsky et al., 2000). [The ssue of stress
responses to nose exposure has been dscussed
recently by Wrght et al. (n press).]
Sound at certan frequences can cause an ar-
flled space to vbrate at ts resonant frequency
(acoustc resonance), whch may ncrease the lke-
lhood of mechancal trauma n the adjacent or sur-
roundng tssue. The resonant frequences of most
marne mammal lungs are below the operatng
frequences of many anthropogenc sound sources
(Fnneran, 2003). Further, bologcal tssues are
heavly damped, estmated tssue dsplacement
at resonant frequences s predcted to be exceed-
ngly small, and lung tssue damage s generally
uncommon n acoustc-related marne mammal
strandng events. For these reasons, specalsts do
not regard lung resonance as a lkely sgnfcant
non-audtory effect for marne mammals exposed
to anthropogenc nose sources that operate above
100 Hz (U.S. Department of Commerce, 2002).
Ths concluson mght not apply to lower-fre-
quency sources that operate at a partcular fre-
quency for a sgnfcant duraton.
The non-audtory effect now beng most
actvely dscussed n marne mammalogy s ntro-
gen gas bubble growth, resultng n effects smlar
to decompresson sckness n humans. Jepson et al.
(2003) and Fernndez et al. (2004, 2005) hypoth-
eszed that lesons (gas and fat embol) observed
n ndvdual beaked whales found stranded after
mltary sonar exercses were somehow caused by
invivo ntrogen bubble formaton. Osteonecross
n sperm whales has further been suggested as
a chronc result of ntrogen bubble formaton
(Moore & Early, 2004).
To date, the gas bubble hypothess remans
untested, and the acoustc causatve mechansm
for formaton of embol, f any, s unknown.
Theoretcally, bubble precursors n supersaturated,
homogenzed tssue may ncrementally enlarge
durng the successve passage of compresson and
rarefacton portons of acoustc waves that exceed
statc pressure (rectfed dffuson; Crum & Mao,
1996). Alternatvely, a sngle acoustc exposure
could actvate bubble precursors, allowng them
to grow by gradual expanson nto bubbles n
ntrogen-supersaturated tssue (statc dffuson;
see Potter, 2004). The dvng patterns of some
marne mammals ncrease gas-tssue saturaton
and potentally could ncrease the susceptblty of
nose-exposed anmals to bubble growth va ether
mechansm (Rdgway & Howard, 1979; Houser
et al., 2001b). Ntrogen supersaturaton levels for
deep-dvng speces of nterest, ncludng beaked
whales, are based on theoretcal models, however
(Houser et al., 2001b). No unequvocal support for
ether pathway presently exsts.
The evdence for bubble formaton as a causal
mechansm between certan types of acoustc
exposure and strandng events remans equvo-
cal. At a mnmum, scentfc dsagreement and/or
426 Southalletal.
complete lack of nformaton exsts regardng the
followng mportant ponts: (1) receved acous-
tc exposure condtons for anmals nvolved n
strandng events; (2) pathologcal nterpretaton
of observed lesons n stranded marne mammals
(Fernndez et al., 2004; Pantados & Thalmann,
2004); (3) acoustc exposure condtons requred
to nduce such physologcal trauma drectly;
(4) whether nose exposure may cause behav-
oral reactons (e.g., atypcal dvng behavor) that
secondarly nduce bubble formaton and tssue
damage (Jepson et al., 2003; Fernndez et al.,
2005; Zmmer & Tyack, 2007); and (5) the extent
that postmortem artfacts ntroduced by decompo-
ston before samplng, handlng, freezng, or nec-
ropsy procedures affect nterpretaton of observed
lesons. Tests of the gas bubble hypothess may
yeld data pertnent to future marne mammal
nose exposure crtera, but too lttle s currently
known to establsh explct exposure crtera for
ths proposed mechansm.
Courtesy: A. Fredlander
2. Structure of the Noise Exposure Criteria
When de facto nose exposure gudelnes are
used by management agences, they generally are
based on a small number of categores of marne
mammals and sound types. Though t would be
convenent to have a sngle exposure crteron
for all speces and sound sources, such a smpl-
fed approach s not supported by avalable sc-
ence. However, some categorzaton of speces
and sources s warranted based on current nfor-
maton. The many anthropogenc sound sources
used n marne envronments can be categorzed
based on certan acoustc and operatonal features.
Smlarly, there s great dversty n hearng and
n the bologcal effects of nose among marne
mammals, but current knowledge supports some
functonal and/or phylogenetc groupngs.
It s also nether possble nor desrable to derve
dstnct exposure crtera for every speces and
sound source. Important generalzatons across
taxa would be mssed even f resources and tme
were adequate to study each speces and expo-
sure condton. Further, t s mpractcal to apply
numerous, speces-specfc crtera when predct-
ng and/or attemptng to mtgate effects.
A standard scentfc approach n such stua-
tons s to categorze anmals based on functonal
characterstcs and sound sources based on phys-
cal smlartes, and to summarze the nformaton
n a matrx format. We subdvde cetaceans and pn-
npeds nto fve functonal hearng categores based
on the frequences they hear. Other methods of cat-
egorzaton are, of course, possble. For nstance,
Verboom (2002) reled heavly on drect measure-
ments of nose mpacts on hearng to quantfy the
effects of nose exposure on marne mammals. Some
of hs proposed crtera are comparable wth those
presented here. The present effort makes broader
use of laboratory and feld behavoral and audomet-
rc data, addtonal recent data, and extrapolatons
from terrestral mammals not used by Verboom. We
dvde sound sources nto three types accordng to
acoustc characterstcs defned at the source. Note
that at a dstance, a sound may have sgnfcantly
dfferent features; categorzng sounds based on
source characterstcs s a precautonary and prag-
matc approach (as s descrbed n the next secton).
The justfcatons for and assumptons underlyng
our categorzaton of functonal hearng groups and
sound types are descrbed here. The number of sub-
dvsons n future nose exposure crtera wll lkely
ncrease as more supportng data are acqured.
The format of the recommended marne
mammal nose exposure crtera s thus a matrx of
15 cells that systematcally consders three sound
types (see next secton) and fve functonal marne
mammal hearng groups (see the Marne Mammal
Functonal Hearng Groups secton of ths chap-
ter). Wthn each of those 15 cells, we consder
two general acoustc metrcs (see the Exposure
Crtera Metrcs secton) and two levels of expo-
sure effect (Levels of Nose Effect: Injury and
Behavoral Dsturbance secton of ths chapter).
Sxty possble crtera result (.e., 3 sound types
5 marne mammal groups 2 metrcs 2 mpact
levels), although fewer than 60 are reported due to
data lmtatons. Whereas sound types are defned
by source features, crtera values represent levels
receved by ndvdual marne mammals.
Sound Types
Three sound types are used: (1) a sngle pulse,
(2) multple pulses, and (3) nonpulses. The separa-
ton between pulses and nonpulses s supported by
data on audtory fatgue and acoustc trauma n ter-
restral mammals (e.g., Dunn et al., 1991; Hamernk
et al., 1993) and s generally consstent wth the
sound types dstngushed for damage rsk crtera
n humans (e.g., U.S. DoD, 1997; NIOSH, 1998).
Pulses and nonpulses are dstngushed by
numerous defntons and mathematcal dstnc-
tons (e.g., Burdc, 1984). The emprcal dstnc-
ton used here s based on a measurement proce-
dure usng several temporal weghtngs. Varous
exponental tme-weghtng functons appled n
measurng pulse and nonpulse sounds may yeld
dfferent measured receved levels (RLs) (see
Harrs, 1998). Most sound level meters (SLM) pro-
vde optons for applyng ether a slow or fast
tme constant (1,000 or 125 ms, respectvely) for
measurng nonpulses or an mpulse tme con-
stant (35 ms) approprate for measurng pulses.
For a sound pulse, the slow or fast SLM settngs
result n lower sound pressure level (SPL) mea-
surements than those obtaned usng the mpulse
settng. Each of these tme constants s selected
based on propertes of the human audtory system.
These may be at least generally relevant for other
mammalan audtory systems, although further
emprcal data on temporal resoluton n marne
mammals are needed (see Chapter 5, Research
Recommendatons).
AquaticMammals 2007, 33(4), 427-436, DOI 10.1578/AM.33.4.2007.427
428 Southalletal.
Harrs (1998) proposed a measurement-based
dstncton of pulses and nonpulses that s adopted
here n defnng sound types. Specfcally, a 3-dB
dfference n measurements between contnuous
and mpulse SLM settngs ndcates that a sound s
a pulse; a < 3-dB dfference ndcates that a sound
s a nonpulse. We note the nterm nature of ths
dstncton for underwater sgnals and the need for
an explct dstncton and measurement standard
such as exsts for aeral sgnals (ANSI, 1986).
Harrss (1998) defntons assumed use of A-
weghtng as do most human-orented defntons
of acoustcal measurements; however, dfferent
frequency-weghtng functons should be used for
varous anmal taxa (as dscussed below). Leavng
that queston asde temporarly, t s nstructve to
compare the mpulse equvalent-contnuous sound
level (LIeqT) for a sound that ncreases n level wth
the correspondng equvalent-contnuous level
(LeqT). Here, LIeqT has an mpulse ntegraton tme of
35 ms and LeqT, defned as sound exposure dvded
by T, s expressed as a level. As an example, sup-
pose that a source s examned over a 2-s perod
(T = 2 s). The hghest LAIeq2s (A here denotes A-
weghtng) durng ths perod s 75.2 dB, and the
hghest LALeq2s s 65.1 dB. The dfference of 10.1
dB s greater than the 3-dB crteron gven by
Harrs (1998); therefore, the sound s consdered
to be a pulse.
The dstncton between pulses and nonpulses s
not always clear n practce. For nstance, certan
sgnals (e.g., acoustc deterrent and harassment
devces) have characterstcs of both pulses and
nonpulses. Also, certan sound sources (e.g., ses-
mc arguns and ple drvng) may produce pulses
at the source but, through varous propagaton
effects, may meet the nonpulse defnton at greater
dstances (e.g., Greene & Rchardson, 1988). Ths
means that a gven sound source mght be subject
to dfferent exposure crtera, dependng on the
dstance to the recever and ntervenng propaga-
ton varables. Whle ths s certanly realstc for
many real-world exposures, measurements at the
anmal are often not practcal. Changes n sound
characterstcs wth dstance generally result n
exposures becomng less physologcally damag-
ng wth ncreasng dstance because sharp tran-
sent peaks become less promnent. Therefore,
these crtera use a precautonary approach and
classfy sound types based on acoustc character-
stcs at the source. Addtonal emprcal measure-
ments are needed to advance our understandng of
sound type classfcaton as a functon of source,
range, and envronmental varables. We empha-
sze that the use of source parameters to classfy
sound types does not negate our decson to rec-
ommend exposure crteron levels relatve to RLs
at the anmal.
Treatng pulses and nonpulses as dscrete sound
types s justfed by data on mammals n general and
several cetacean speces n partcular (Dunn et al.,
1991; Hamernk et al., 1993; also see the Effects
of Nose on Hearng n Marne Mammals TTS
Data secton n Chapter 3). Mammalan hearng
s most readly damaged by transent sounds wth
rapd rse-tme, hgh peak pressures, and sustaned
duraton relatve to rse-tme (for humans: Thery
& Meyer-Bsch, 1988; for chnchllas [Chinchilla
lanigera], Dunn et al., 1991). Consstent wth these
results, those odontocetes tested thus far have been
shown to experence TTS-onset at lower respectve
exposure levels f the sound s a pulse rather than a
nonpulse (Fnneran et al., 2002b, 2005a).
Mammals are also apparently at greater rsk
from rapdly repeated transents and those wth
hgh mpulse ampltude kurtosis (Erdrech, 1986).
Hamernk et al. (1993, 2003) argued that the ds-
tncton between exposures wth relatvely hgh
and low peakedness s to some extent an over-
smplfcaton. Hghly varable threshold shfts
can result from exposures of varable peaked-
ness but comparable overall levels, dependng on
a host of factors. Hamernk et al. (1993, 2003)
also noted that peak pressure levels suffcent to
exceed mechancal lmts of the cochlea, and thus
more lkely to nduce acoustc trauma, tend to be
more typcal of pulses than nonpulses.
The present crtera also categorze sound types
based on repetton. For mammals, sngle and mul-
tple nose exposures at varous levels and dura-
tons generally dffer n ther potental to nduce
audtory fatgue or trauma. Ths results prncpally
from the temporal nteracton between exposure
and recovery perods (e.g., Kryter, 1994) and df-
ferences n receved total acoustc energy. Further,
multple exposures may ncrease the lkelhood of
behavoral responses because of ncreased prob-
ablty of detecton and the (generally) greater
bologcal sgnfcance of contnued exposure as
opposed to a sngle, transent event (although see
dscusson of habtuaton n the Responses to
Sound secton of Chapter 1).
Sngle exposures are consdered here as ds-
crete acoustc events n whch receved sound
levels exceed ambent nose n at least some por-
ton of the frequency band of functonal marne
mammal hearng once n a 24-h perod; mult-path
receptons of a sngle exposure are not consd-
ered multple exposures. Multple exposures are
consdered to be acoustc events causng RLs to
exceed ambent nose wthn the functonal band-
wdth more than once, wth an ntervenng quet
perod not exceedng 24 h. If the exposure event
s nterrupted, even brefly (other than as a result
of the anmals own actone.g., breachng), t s
consdered a multple exposure.
MarineMammalNoiseExposureCriteria 429
Exposures should be categorzed as ether pulsed
or nonpulsed sounds as descrbed above. Sngle and
multple exposures to ether pulse or nonpulse sounds
(or both) are possble. Examples of sngle pulses and
sngle nonpulses are sounds from a sngle frng of
an argun or a sngle vessel passage, respectvely.
Multple pulse or multple nonpulse sounds are
more dffcult to delneate, gven the dversty and
complexty of sound sources. A seres exclusvely
consstng of two or more nonpulses would clearly be
a multple nonpulse exposure (e.g., multple vessel
passages). A multple pulse exposure would sm-
larly be descrbed as a seres exclusvely contanng
pulses (e.g., repeated ple strkes) or a combnaton
of pulses and nonpulses (e.g., the combned vessel
nose and argun transmssons of a sesmc vessel).
One justfcaton for treatng combned pulses and
nonpulses as pulses s that the proposed exposure
crtera for njury are more precautonary (lower)
n the case of pulses than for nonpulses. Specfc
consderaton should be gven, on a case-by-case
bass, as to whether such a dstncton would neces-
sarly be the more precautonary. For nstance, f a
compound exposure ncluded relatvely hgh-level
nonpulses as well as relatvely low-level pulses, the
more approprate and protectve dstncton mght
be to classfy t as a nonpulse exposure.
The proposed exposure crtera for njury from
sngle and multple exposures to both sound types
are numercally dentcal (Chapter 3). Ths s another
precautonary decson, arsng from the fact that no
marne mammal data were avalable regardng the
effects of nter-exposure nterval on recovery from
audtory effects (e.g., TTS). A summaton procedure
s appled to quantfy the fatgung effects of mult-
ple exposures wth an equvalent SEL value (Chapter
1; also Appendx A, eq. 5). The SEL metrc takes
account of the pressure waveform and duraton of
ether sngle or multple sound events; t represents
cumulatve receved energy. Ths approach effectvely
negates the need for numercally dfferent njury cr-
tera for sngle and multple exposures at the expense
of neglectng assumed, but as-yet poorly understood
recovery phenomena durng ntervals between expo-
sures. Ths s a precautonary approach, pendng
avalablty of data on acoustc recovery by marne
mammals durng ntervals between exposures.
When consderng behavoral responses, sngle
and multple nonpulse exposures are consdered as
a sngle category. Insuffcent nformaton exsts to
assess the use of SEL as a relevant metrc n the con-
text of marne mammal behavoral dsturbance for
anythng other than a sngle pulse exposure. Future
nose exposure crtera for behavoral dsturbance
may dstngush SPL and SEL exposure crtera for
addtonal condtons, but for most sound types (the
excepton beng sngle pulses), the avalable data
are best assessed n relaton to SPL (dscussed n
detal n Chapter 4). Consequently, the structure of
the exposure crtera matrx ncludes a categorcal
dstncton between sngle and multple pulses gven
that numercal SEL thresholds are recommended for
a sngle pulse, but not for multple pulses. No such
dstncton s made for nonpulses where the avalable
data do not (at least currently) support dfferental
behavoral crtera for sngle vs multple exposures.
Thus, the current state of scentfc knowledge
regardng mammalan hearng and varous nose
mpacts supports three dstnct sound types as
relevant for marne mammal nose exposure cr-
tera: (1) sngle pulse, (2) multple pulses, and
(3) nonpulses. Examples of sound sources belong-
ng n each of these categores (based on character-
stcs of the sound emtted at the source) are gven
n Table 1. A smplstc measurement procedure
usng source characterstcs (the 3-dB dstncton
based on Harrs, 1998, descrbed above) s used
here to dstngush a pulse from a nonpulse, whle
the smple defntons above dstngush sngle
and multple exposures.
Table 1. Sound types, acoustc characterstcs, and selected examples of anthropogenc sound sources; note sound types are
based on characterstcs measured at the source. In certan condtons, sounds classfed as pulses at the source may lack these
characterstcs for dstant recevers.
Sound type Acoustc characterstcs (at source) Examples
Sngle pulse Sngle acoustc event; > 3-dB dfference between
receved level usng mpulse vs equvalent
contnuous tme constant
Sngle exploson; sonc boom; sngle argun,
watergun, ple strke, or sparker pulse; sngle png
of certan sonars, depth sounders, and pngers
Multple pulses Multple dscrete acoustc events wthn 24 h;
> 3-dB dfference between receved level usng
mpulse vs equvalent contnuous tme constant
Seral explosons; sequental argun, watergun,
ple strkes, or sparker pulses; certan actve sonar
(IMAPS); some depth sounder sgnals
Nonpulses Sngle or multple dscrete acoustc events wthn
24 h; < 3-dB dfference between receved level
usng mpulse vs equvalent contnuous tme
constant
Vessel/arcraft passes; drllng; many construc-
ton or other ndustral operatons; certan sonar
systems (LFA, tactcal md-frequency); acoustc
harassment/deterrent devces; acoustc tomogra-
phy sources (ATOC); some depth sounder sgnals
430 Southalletal.
Marine Mammal Functional Hearing Groups
Speces of cetaceans and pnnpeds were assgned
to one of fve functonal hearng groups based on
behavoral psychophyscs, evoked potental aud-
ometry, audtory morphology, and (for pnnpeds)
the medum n whch they lsten. Cetaceans and
pnnpeds are broadly separable based on phylo-
genetc and functonal dfferences (Reynolds &
Rommel, 1999). Cetaceans were further subd-
vded accordng to dfferences n ther measured
or estmated hearng characterstcs and not neces-
sarly accordng to ther phylogeny (as n Wartzok
& Ketten, 1999). Pnnpeds are consdered a sngle
group, but as amphbous mammals, ther hearng
dffers n ar and n water (Kastak & Schusterman,
1998); separate crtera were requred for each
medum. The taxa n each functonal hearng
group (based on Rce, 1998) are gven n Table 2.
MarineMammalHearing
All marne mammals evolved from terrestral, ar-
adapted ancestors (Domnng et al., 1982; Barnes
et al., 1985) and, at least n part, retan the nomnal
mammalan trpartte perpheral audtory system
(.e., external audtory meatus, ar-flled mddle
ear, and spral-shaped cochlea). Most of the mech-
ansms of mammalan hearng are also conserved
such as the basc lever structure of the osscles and
the tonotopc organzaton of the har cells along
the nner ears baslar membrane.
However, marne mammal audtory systems dffer
n havng some adaptatons that seem to be related
to pressure, hydrodynamcs, and sound recepton n
water (see Wartzok & Ketten, 1999). For nstance,
the pnna has been reduced or elmnated n most
speces, owng to hydrodynamc adaptatons.
Tssue modfcatons may enable the reducton or
elmnaton of gas spaces n the mddle ear of some
marne mammals. Consequently, bone conducton,
rather than the conventonal osscular chan, may be
an addtonal (or prmary) sound transmsson path
to the cochlea (e.g., Repennng, 1972; Au, 1993).
There are mportant dfferences n these adaptatons
wthn and between marne mammal taxa.
Knowledge of marne mammal hearng vares
wdely among groups, but for most speces t s
qute lmted compared to knowledge of terrestral
mammal hearng. Because of the sheer sze, lm-
ted and dsproportonate avalablty n captve
Table 2. Functonal marne mammal hearng groups, audtory bandwdth (estmated lower to upper frequency hearng
cut-off), genera represented n each group, and group-specfc (M) frequency-weghtngs
Functonal hearng
group
Estmated audtory
bandwdth
Genera represented
(Number speces/subspeces)
Frequency-weghtng
network
Low-frequency
cetaceans
7 Hz to 22 kHz Balaena,Caperea,Eschrichtius,
Megaptera,Balaenoptera
(13 speces/subspeces)
Mlf
(lf: low-frequency cetacean)
Md-frequency
cetaceans
150 Hz to 160 kHz Steno,Sousa,Sotalia,Tursiops,Stenella,
Delphinus,Lagenodelphis,Lagenorhynchus,
Lissodelphis,Grampus,Peponocephala,
Feresa,Pseudorca,Orcinus,Globicephala,
Orcaella,Physeter,Delphinapterus,
Monodon,Ziphius,Berardius,
Tasmacetus,Hyperoodon,Mesoplodon
(57 speces/subspeces)
Mmf
(mf: md-frequency
cetaceans)
Hgh-frequency
cetaceans
200 Hz to 180 kHz Phocoena,Neophocaena,
Phocoenoides,Platanista,Inia,Kogia,
Lipotes,Pontoporia,Cephalorhynchus
(20 speces/subspeces)
Mhf
(hf: hgh-frequency
cetaceans)
Pnnpeds n water 75 Hz to 75 kHz Arctocephalus,Callorhinus,
Zalophus,Eumetopias,Neophoca,
Phocarctos,Otaria,Erignathus,Phoca,
Pusa,Halichoerus,Histriophoca,
Pagophilus,Cystophora,Monachus,
Mirounga,Leptonychotes,Ommatophoca,
Lobodon,Hydrurga,and Odobenus
(41 speces/subspeces)
Mpw
(pw: pnnpeds n water)
Pnnpeds n ar 75 Hz to 30 kHz Same speces as pnnpeds n water
(41 speces/subspeces)
Mpa
(pa: pnnpeds n ar)
MarineMammalNoiseExposureCriteria 431
settngs, and, for many speces and jursdctons,
the protected status of marne mammals, there
are lmtatons n obtanng hearng data for many
speces. Behavoral or electrophysologcal audo-
grams exst for fewer than 20 marne mammal
speces (of ~128 speces and subspeces; Rce,
1998). By combnng these data wth comparatve
anatomy, modelng, and response measured n
ear tssues from speces that are dffcult to study,
however, t s possble to descrbe the frequency
senstvty and crtcal adaptatons for underwa-
ter hearng n each of the fve functonal hearng
groups of marne mammals consdered here.
Low-frequency cetaceans consst of 13 speces
and subspeces of mystcete (baleen) whales n
fve genera (based on Rce, 1998; see Table 2). No
drect measurements of hearng exst for these an-
mals, and theores regardng ther sensory capabl-
tes are consequently speculatve (for a detaled
assessment by speces usng the lmted avalable
nformaton, see Erbe, 2002). They are too large to
mantan n the laboratory for psychophyscal test-
ng. The lmted evoked potental measurements
on anmals of ths sze have not yet yelded hearng
thresholds (Rdgway & Carder, 2001), but techno-
logcal advances may soon enable evoked poten-
tal audometry on relatvely small and/or young
mystcetes. In these speces, hearng senstvty
has been estmated from behavoral responses
(or lack thereof) to sounds at varous frequences,
vocalzaton frequences they use most, body sze,
ambent nose levels at the frequences they use
most, and cochlear morphometry (Rchardson
et al., 1995; Wartzok & Ketten, 1999; Houser
et al., 2001a; Erbe, 2002; Clark & Ellson, 2004).
Untl better nformaton s avalable regard-
ng the relatonshp between audtory senstvty
and marne envronmental nose, the senstvty
of mystcetes cannot be easly nferred from the
acoustc envronment.
The combned nformaton strongly suggests
that mystcetes are lkely most senstve to sound
from perhaps tens of Hz to ~10 kHz. However,
recent data ndcated that humpback whales
(Megaptera novaeangliae) produce some sgnals
wth harmoncs extendng above 24 kHz (Au
et al., 2006). These harmoncs have consderably
lower levels than occur at lower frequences, and
ther presence does not necessarly ndcate they
are audble to the whales. Nonetheless, some
hgh-frequency energy s present. [Addtonally,
some recent anatomcal modelng work by
Ketten et al. (2007) suggested that some myst-
cetes may have functonal hearng capabltes at
frequences as hgh as 30 kHz.] Whle we do not
nclude these recent results at ths tme, we note
ther presence and the possblty that the upper
frequency lmt of the M-weghtng functon
for mystcetes may need to be revsted based
on emergng knowledge. At present, we est-
mate the lower and upper frequences for func-
tonal hearng n mystcetes, collectvely, to be
7 Hz and 22 kHz (Ketten et al., 2007).
Md- and hgh-frequency cetaceans are all
odontocetes (toothed whales). Unlke the mystce-
tes, all odontocete cetaceans appear to have hghly
advanced echolocaton (bosonar) systems that
use ntermedate to very hgh frequences (tens of
kHz to 100+ kHz: see Au, 1993; Rchardson et al.,
1995; Wartzok & Ketten, 1999). They also produce
socal sounds n a lower-frequency band, ncludng
generally low to ntermedate frequences (1 kHz to
tens of kHz). Consequently, ther functonal hear-
ng would be expected to cover a wder absolute
frequency range than s assumed for mystcetes or
has been demonstrated for pnnpeds (dscussed
below). Ths has been expermentally confrmed
n the odontocete speces whose hearng has been
measured (dscussed below); however, ther best
hearng senstvty typcally occurs at or near the
frequency where echolocaton sgnals are stron-
gest. Based on the dfferental characterstcs of
echolocaton sgnals n two groups of odontocetes
(see Au, 1993) and on the hearng data descrbed
below, odontocetes were dvded nto md- and
hgh-frequency functonal groups (as seen gener-
ally n Wartzok & Ketten, 1999).
Md-frequency cetaceans nclude 32 speces
and subspeces of dolphns, sx speces of larger
toothed whales, and 19 speces of beaked and bot-
tlenose whales (see Table 2). Functonal hear-
ng n ths group was estmated to occur over a
wde range of low to very hgh frequences. Based
on the combned avalable data, md-frequency
speces are estmated to have lower and upper
frequency lmts of nomnal hearng at approx-
mately 150 Hz and 160 kHz, respectvely. As for
the other hearng groups, there s varablty wthn
and among speces, ntense sgnals below and
above the stated bounds may be weakly detect-
able, and there s a progressve rather than nstan-
taneous reducton n hearng senstvty near these
lmts. Md-frequency cetaceans generally do not
appear well-adapted to detect or to dscrmnate
sgnals outsde ths frequency band, however. The
scarcty (and varablty) of emprcal data pre-
cludes a fner subdvson of ths relatvely dverse
and large group of marne mammals, though t s
acknowledged that some md-frequency speces
lkely have a narrower functonal hearng band
than the range gven above.
Behavoral hearng data are avalable for the
followng md-frequency cetacean speces: bot-
tlenose dolphn (Tursiops truncatus: Johnson,
1967; Ljungblad et al., 1982; Fnneran et al.,
2005a), beluga (Delphinapterus leucas: Whte
432 Southalletal.
et al., 1978; Awbrey et al., 1988; Johnson, 1992;
Rdgway et al., 2001; Fnneran et al., 2005b),
kller whale (Orcinus orca: Hall & Johnson,
1972; Szymansk et al., 1999), false kller whale
(Pseudorca crassidens: Thomas et al., 1988,
1990a; Au et al., 1997), Rssos dolphn (Grampus
griseus: Nachtgall et al., 1995; Au et al., 1997);
and Pacfc whte-sded dolphn (Lagenorhynchus
obliquidens: Tremel et al., 1998).
Audograms derved usng audtory evoked
potental (AEP) methodology (Supn et al., 2001)
have been obtaned for a number of cetacean spe-
ces. Specfc AEP technques, whch nvolve
measurng electrophysologcal responses to
sound, nclude those measurng transent evoked
responses, such as the audtory branstem response
(ABR) or md-latency response, and those mea-
surng steady-state evoked responses such as the
envelope followng response (EFR) or audtory
steady-state response (ASSR). Md-frequency
cetacean speces tested nclude the bottlenose
dolphn (Bullock et al., 1968; Seeley et al., 1976;
Popov & Supn, 1990; Houser & Fnneran, 2006b;
Fnneran et al., 2007a; Hernandez et al., 2007;
Popov et al., 2007), kller whale (Szymansk
et al., 1999), beluga (Popov & Supn, 1990;
Klshn et al., 2000), common dolphn (Delphinus
delphis: Popov & Klshn, 1998), Rssos dolphn
(Dolphn, 2000; Nachtgall et al., 2005, 2007),
tucux dolphn (Sotaliafluviatilis: Popov & Supn,
1990), strped dolphn (Stenella coeruleoalba:
Kastelen et al., 2003), Pacfc whte-sded dol-
phn (Au et al., 2007), false kller whale (Supn
et al., 2003), and Gervas beaked whale (Cook
et al., 2006). Addtonally, Yuen et al. (2005) con-
ducted a comparatve study of behavoral and AEP
thresholds for the false kller whale, and Fnneran
& Houser (2006), Houser & Fnneran (2006a),
and Fnneran et al. (2007b) have compared behav-
oral and AEP thresholds n multple bottlenose
dolphns.
The hgh-frequency cetaceans nclude eght
speces and subspeces of true porposes, sx spe-
ces and subspeces of rver dolphns plus the fran-
cscana, Kogia, and four speces of cephalorhyn-
chds (see Table 2). Functonal hearng n ths
group was estmated to occur between 200 Hz and
180 kHz. Behavoral audograms are avalable for
the followng hgh-frequency cetacean speces:
harbor porpose (Phocoena phocoena: Andersen,
1970; Kastelen et al., 2002a), Chnese rver dol-
phn (Lipotes vexillifer: Wang et al., 1992), and
Amazon rver dolphn (Iniageoffrensis: Jacobs &
Hall, 1972). Audograms usng AEP methodol-
ogy have been obtaned for three speces: harbor
porpose (Popov et al., 1986, 2006; Beedholm
& Mller, 2007; Lucke et al., 2007b); fnless
porpose (Neophocaena phocaenoides: Popov
et al., 2006); and Amazon rver dolphn (Popov &
Supn, 1990).
The pnnpeds nclude 16 speces and subspeces
of sea lons and fur seals (otards), 23 speces and
subspeces of true seals (phocds), and two sub-
speces of walrus (odobends). Pnnpeds produce
a wde range of socal sgnals, most occurrng at
relatvely low frequences. They lack the hghly-
specalzed actve bosonar systems of odontocete
cetaceans, possbly as a result of ther amphbous
lfestyle (see Schusterman et al., 2000). Because
of ths aspect of ther lfe hstory, pnnpeds com-
muncate acoustcally n ar and water, have sg-
nfcantly dfferent hearng capabltes n the
two meda, and may be subject to both aeral and
underwater nose exposure (Schusterman, 1981;
Kastak & Schusterman, 1998, 1999). These df-
ferences necesstate separate nose exposure crte-
ra for pnnpeds n each medum.
For pnnpeds n water, behavoral measures
of hearng are avalable for the northern fur seal
(Callorhinus ursinus: Moore & Schusterman,
1987; Babushna et al., 1991), Calforna sea
lon (Zalophuscalifornianus: Schusterman et al.,
1972; Moore & Schusterman, 1987; Kastak &
Schusterman, 1998, 2002; Southall et al., 2004),
northern elephant seal (Mirounga angustirostris:
Kastak & Schusterman, 1998, 1999; Southall
et al., 2004), Hawaan monk seal (Monachus
schauinslandi: Thomas et al., 1990b), harp seal
(Pagophilus groenlandicus: Terhune & Ronald,
1972), rnged seal (Phoca hispida: Terhune
& Ronald, 1975), harbor seal (Mhl, 1967,
1968; Terhune & Turnbull, 1995; Kastak &
Schusterman, 1995, 1998; Southall et al., 2004),
and walrus (Odobenus rosmarus: Kastelen
et al., 2002b). Rdgway & Joyce (1975) measured
the gray seals (Halichoerus grypus) underwater
hearng usng evoked potental audometry.
For pnnpeds n ar, behavoral measures of hear-
ng are avalable for the northern fur seal (Moore
& Schusterman, 1987; Babushna et al., 1991),
Calforna sea lon (Schusterman, 1974; Kastak &
Schusterman, 1998; Kastak et al., 2004b), north-
ern elephant seal (Kastak & Schusterman, 1998,
1999; Kastak et al., 2004b), harp seal (Terhune
& Ronald, 1971), and harbor seal (Mhl, 1968;
Kastak & Schusterman, 1998; Kastak et al.,
2004b). Aeral hearng n pnnpeds has also been
measured usng evoked potental audometry n
the gray seal (Rdgway & Joyce, 1975), Calforna
sea lon (Bullock et al., 1971; Rdgway & Joyce,
1975; Mulsow & Rechmuth, 2007; Rechmuth
et al., 2007), harbor seal (Thorson et al., 1998;
Wolsk et al., 2003; Mulsow & Rechmuth, 2007;
Rechmuth et al., 2007), and northern elephant
seal (Houser et al., 2007; Mulsow & Rechmuth,
2007; Rechmuth et al., 2007).
MarineMammalNoiseExposureCriteria 433
The combned results of these studes nd-
cate that pnnpeds are senstve to a broader
range of sound frequences n water than n ar.
The data further suggest dfferences n the func-
tonal hearng range among otards, phocds,
and odobends, especally under water (Kastak &
Schusterman, 1998; Kastelen et al., 2002b). For
these proposed nose exposure crtera, however,
pnnpeds are consdered a sngle functonal hear-
ng group because the data are too lmted, both n
terms of absolute hearng data and TTS measure-
ments (see The Effects of Nose on Hearng n
Marne Mammals: TTS Data secton n Chapter
3), to support fner subdvsons. We estmate that
pnnpeds have functonal underwater hearng
between 75 Hz and 75 kHz and functonal aeral
hearng between 75 Hz and 30 kHz. These ranges
are essentally based on data for phocd seals,
whch have the broadest audtory bandwdths of
the pnnpeds. Ths approach results n a precau-
tonary functonal bandwdth for estmatng fre-
quency-weghtng functons (below) and nose
mpacts on pnnpeds.
In summary, based on current knowledge
of functonal hearng n marne mammals, fve
dstnct, functonal hearng categores were
defned: (1) low-frequency cetaceans (.e., mys-
tcetes), (2) md-frequency cetaceans (.e., most
odontocetes), (3) hgh-frequency cetaceans (.e.,
porposes, rver dolphns, pygmy sperm whale,
and Cephalorhynchus), (4) pnnpeds n water,
and (5) pnnpeds n ar. The genera n each group,
and the estmated lower and upper frequency
hearng lmts, are shown n Table 2. Because
the fve functonal hearng groups of marne mam-
mals dffer n hearng bandwdth, each may be
affected dfferently by dentcal nose exposures.
Therefore, frequency-weghtng functons are
requred to develop marne mammal nose expo-
sure crtera.
Frequency-WeightingFunctions
As a general statement, anmals do not hear
equally well at all frequences wthn ther func-
tonal hearng range. Frequency weghtng s a
method of quanttatvely compensatng for the
dfferental frequency response of sensory sys-
tems. Generalzed frequency-weghtng functons
were derved for each functonal hearng group of
marne mammals usng prncples from human
frequency-weghtng paradgms, wth adjustments
for the dfferent hearng bandwdths of the varous
marne mammal groups.
For humans, substantal mprovement n dose-
response models s obtaned by flterng nose
through equal-loudness functons, partcularly
the 40-phon, equal-loudness functon (A-weght-
ng) and the 100-phon functon (C-weghtng).
These frequency-weghtng functons take nto
account both the frequency bandwdth of human
hearng and loudness percepton. For use as fre-
quency flters, the functons are nverted; normal-
zed to 0 dB n the frequency range of best hearng
(specfcally at 1,000 Hz for humans); and deal-
zed for mplementaton n hearng ads, sound
level meters, and other measurement devces.
At mnmum, metrcs used for anmals should
elmnate naudble frequences both below and
above the range of functonal hearng. The abso-
lute audtory threshold functon (audogram) has
been suggested as a frequency-weghtng func-
ton for marne speces exposed to underwater
sound (e.g., Malme et al., 1989; Thorson et al.,
1998; Heathershaw et al. 2001; Nedwell et al.,
2007) as well as for terrestral anmals (Delaney
et al., 1999; Bjork et al., 2000). However, the
audtory threshold functon does not characterze
the flattenng of equal-loudness percepton wth
the ncreasng stmulus level that has been dem-
onstrated n humans (Fletcher & Munson, 1933).
Acoustc njury would only be expected to occur
at levels far above the detecton thresholdthat
s, levels for whch the flattenng effect would be
expected. Consequently, t s unclear how useful
or approprate the audtory threshold functon s
n dervng frequency-weghtng flters n marne
mammals for whch psychophyscal equal-loud-
ness measurements are generally unavalable
(although see prelmnary measurements by
Rdgway & Carder, 2000). Further, the lmted
TTS data for cetaceans exposed to tones at dffer-
ent frequences (dscussed below) suggest that an
audogram-based frequency-weghtng functon
would produce too much flterng at lower fre-
quences (.e., the weghtng functon for hearng
effects should be flatter than the nverted audo-
gram procedure would ndcate).
Therefore, a precautonary procedure was used
to derve frequency-specfc, marne mammal
weghtng functons. Each was based on an algo-
rthm that requres only the estmated (as ~80 dB
above best hearng senstvty) lower and upper
frequences of functonal hearng as gven n the
above descrpton of each marne mammal group
and n Table 2. The resultng functons were
desgned to reasonably represent the bandwdth
where acoustc exposures can have audtory effects
and were desgned to be most accurate for descrb-
ng the adverse effects of hgh-ampltude nose
where loudness functons are expected to flatten
sgnfcantly. The weghtng functons (desgnated
M for marne mammal) are analogous to the
C-weghtng functon for humans, whch s com-
monly used n measurng hgh-ampltude sounds.
In the general absence of emprcal data, however,
the upper and lower frequency roll-offs of the
434 Southalletal.
M-weghtng functons are symmetrcal, whereas
C-weghtng admts more energy at the lower than
at the upper frequency lmts (ANSI, 2001).
The M-weghtng functons assume a logarth-
mc reducton n audtory senstvty outsde of the
range of best hearng senstvty, wth the functon
beng 6 dB down from peak senstvty at the lower
and upper frequency lmts. Audtory detecton
thresholds at these lmts (see above dscusson
of lower and upper frequency cut-offs) can be
80 dB hgher (less senstve) than those at the fre-
quences of best hearng senstvty. Consequently,
these frequency flters are much flatter than
audograms and probably qute precautonary
even consderng the expected flattenng of equal-
loudness contours at hgh exposure levels. The
M-weghtng functons are also precautonary n
that regons of best hearng senstvty for most
speces are lkely consderably narrower than the
M-weghtng functons (desgned for the overall
marne mammal group) would suggest. The gen-
eral expresson for M-weghtng (M[f]), usng
the estmated lower and upper functonal hear-
ng lmts (flow and fhgh) for each of the fve func-
tonal marne mammal hearng groups, s gven n
Appendx A (eq. 7 & 8). These frequency-weght-
ng functons are dentfed n Table 2, and each s
depcted graphcally n Fgure 1.
The M-weghtng functons de-emphasze fre-
quences that are near the lower and upper fre-
quency ends of the estmated hearng range as
ndcated by negatve relatve values (Fgure 1).
Ths de-emphass s approprate because, to have
a gven audtory effect, sound at these frequences
must have hgher absolute ampltude than sound n
the regon of best hearng senstvty. As a corol-
lary, sound at a gven level wll have less effect f
t s near or (especally) beyond the lower or upper
bounds of the functonal hearng range than f t s
well wthn that frequency range. It s mportant
to note the ncremental nature of the frequency-
weghtng functons, whch approxmate the
gradual reducton n audtory effect at frequences
outsde the range of greatest senstvty.
Use of such M-frequency-weghtng functons
s superor to flat weghtng across all frequences
because t accounts for known or estmated dffer-
ences n the frequency response characterstcs for
each functonal hearng group. At least n the context
of njury crtera, t s superor to frequency-weght-
ng va the nverse-audogram method as t takes
nto account the expected flattenng of equal-
loudness curves at the hgh exposure levels where
TTS and PTS are expected. It s also superor to a
boxcar-type step functon because t more closely
approxmates the gradual roll-off of senstvty
below and above the range of optmum senstvty.
Furthermore, each of the recommended shallow
frequency-weghtng functons ncludes, wthn ts
relatvely flat porton, the full audble range for
each speces for whch audtory data are avalable.
In other words, none of the speces ncluded wthn
each functonal hearng group has been shown or
s expected to have any porton of ts best hearng
senstvty outsde the flat porton of the relevant
frequency-weghtng functon. Thus, the functons
are qute precautonary, whch s approprate gven
that data are lmted or lackng for most speces.
Exposure Criteria Metrics
Many acoustc metrcs (e.g., RMS or peak SPL,
SEL, kurtoss) could be consdered n relaton
to nose mpacts on anmals. It s mpossble to
predct unequvocally whch one s best assoc-
ated wth the lkelhood of njury or sgnfcant
behavoral dsturbance across all taxa because of
speces dfferences and the fact that real-world
sound exposures contan many wdely dfferng
temporal patterns and pressure sgnatures. To
account for such dfferences and to allow for cur-
rent scentfc understandng of tssue njury from
nose exposure, the proposed njury crtera ncor-
porate a dual-crtera approach based on both peak
pressure and energy. For an exposed ndvdual,
whchever crteron s exceeded frst (.e., the more
precautonary of the two measures) s used as the
operatve njury crteron. Smlarly, a dual-crte-
ron approach (peak sound pressure and energy)
s also proposed for behavoral dsturbance from
a sngle pulse.
The pressure crtera for njury are defned as
those peak SPLs above whch tssue njury s pre-
dcted to occur, rrespectve of exposure duraton.
Any sngle exposure at or above ths peak pressure
s consdered to cause tssue njury, regardless of
the SPL or SEL of the entre exposure. For each
marne mammal group, the recommended pres-
sure-based njury crtera are the same for all sound
types and are based on the crteron for a sngle
pulse. Ths s a precautonary procedure; pressure
crtera based on TTS data for nonpulses would
yeld much hgher estmates of the exposure nec-
essary for PTS-onset. By proposng, for all cases,
pressure crtera approprate to a sngle pulse, we
protect aganst the possblty that, for some sound
sources, one or more ntense pulses may occason-
ally be embedded n nonpulse sounds.
For exposures lackng ntense peak pressure
components, avalable data ndcate that measure-
ments ntegratng nstantaneous pressure squared
over the duraton of sound exposure are well corre-
lated wth the probablty of TTS-onset and tssue
njury. Consequently, for exposures other than
those contanng ntense peak pressure transents,
SEL s the (or at least one of the) approprate
MarineMammalNoiseExposureCriteria 435
(A)
(A)
(B )
W
e
i
g
h
t
i
n
g

(
d
B
)
W
e
i
g
h
t
i
n
g

(
d
B
)
Frequency (kHz)
Frequency (kHz)
M-weighting
M-weighting
pinnipeds in water, M
pw
pinnipeds in air, M
pa
low-frequency cetaceans, M
lf
mid-frequency cetaceans, M
mf
high-frequency cetaceans, M
hf
(B)
(A)
(B )
W
e
i
g
h
t
i
n
g

(
d
B
)
W
e
i
g
h
t
i
n
g

(
d
B
)
Frequency (kHz)
Frequency (kHz)
M-weighting
M-weighting
pinnipeds in water, M
pw
pinnipeds in air, M
pa
low-frequency cetaceans, M
lf
mid-frequency cetaceans, M
mf
high-frequency cetaceans, M
hf
Figure 1. The M-weighting functions for (A) low-, mid-, and high-frequency cetaceans, as well as for (B) pinnipeds in water
and air.
436 Southalletal.
metrc(s) for estmatng TTS-onset and predctng
PTS-onset n humans (ISO, 1990).
Ths use of SEL s based on the assumpton that
sounds of equvalent energy wll have generally
smlar effects on the audtory systems of exposed
human subjects, even f they dffer n SPL, dura-
ton, and/or temporal exposure pattern (Kryter,
1970; Nelsen et al., 1986; Yost, 1994; NIOSH,
1998). Under the equal-energy assumpton, at
exposure levels above TTS-onset, each doublng
of sound duraton s assocated wth a 3 dB reduc-
ton n the SPL theoretcally requred to cause the
same amount of TTS. Ths relatonshp has been
used n the dervaton of exposure gudelnes for
humans (e.g., NIOSH, 1998). Numerous authors
have questoned the predctve power of usng
a smplstc total energy approach n all cond-
tons. It fals to account for varyng levels and
temporal patterns of exposure/recovery, among
other factors, and wll thus lkely overestmate
the TTS resultng from a complex nose exposure
(Hamernk & Hsueh, 1991; Hamernk et al., 1993,
2002; Ahroon et al., 1993; Ward, 1997; Strasser
et al., 2003). A comparatve assessment of TTS as
a functon of exposure level n mammals, fsh, and
brds suggests that there are drect relatonshps
but that the slopes vary among taxa (Smth et al.,
2004). The debate over the valdty of the equal
energy rule of nose exposure remans unre-
solved, even for humans.
Some lmted evdence favorng an SEL
approach exsts for marne mammals, how-
ever. Specfcally, an equal-energy relatonshp
for TTS-onset appears to hold reasonably well
for certan nose exposure types wthn sev-
eral md-frequency cetacean speces (Fnneran
et al., 2002b, 2005a; see Effects of Nose
on Hearng n Marne Anmals: TTS Data
secton n Chapter 3). A recent study of n-
ar TTS n a Calforna sea lon (Kastak
et al., 2007) llustrates some condtons n whch
exposures wth dentcal SEL result n consder-
ably dfferent levels of TTS. Nevertheless, because
the very lmted marne mammal data agree rea-
sonably well (at least as a frst-order approxma-
ton) wth equal-energy predctons, and predc-
tons based on SEL wll be precautonary for
ntermttent exposures, we regard t as approprate
to apply the SEL metrc for certan nose exposure
crtera untl future research ndcates an alter-
nate and more specfc course. In certan applca-
tons, there s much more scentfc justfcaton
for use of SEL-based crtera than for prevous
ad hoc SPL crtera (dscussed n the Hstorcal
Perspectve secton n Chapter 1). In applcatons
nvolvng audtory effects, SEL-based crtera
wll lkely more relably dstngush cases where
phenomena of concern (TTS, PTS, etc.) wll and
wll not lkely occur.
Levels of Noise Effect:
Injury and Behavioral Disturbance
Drect audtory tssue effects (njury) and behav-
oral dsrupton are the two categores of nose
effect that are consdered n these marne mammal
exposure crtera. Chapter 3 summarzes all
avalable data on the effects of nose on marne
mammal hearng. It also descrbes how these data
are appled and extrapolated usng precautonary
measures to predct audtory njury and to derve
thresholds and proposed crtera for njury.
In Chapter 4 and Appendces B & C, we summa-
rze the current understandng and avalable data
regardng marne mammal behavoral responses
to nose. Chapter 4 ncludes a quanttatve sever-
ty scale based generally on the NRCs (2005)
Populaton Consequences of Acoustc Dsturbance
(PCAD) Model. Chapter 4 also ncludes a lmted
and cautous entry of behavoral-response data
nto a matrx of severty scalng as a functon of
RL. Currently avalable data, pooled by functonal
hearng group, do not support specfc numercal
crtera for the onset of dsturbance. Rather, they
ndcate the context-specfcty of behavoral reac-
tons to nose exposure and pont to some general
conclusons about response severty n certan,
specfc condtons.
3. Criteria for Injury: TTS and PTS
The crtera for njury for all marne mammal
groups and sound types are receved levels (fre-
quency-weghted where approprate) that meet the
defnton of PTS-onset used here (40 dB-TTS,
descrbed below). Crtera were derved from mea-
sured or assumed TTS-onset thresholds for each
marne mammal group plus TTS growth rate est-
mates (gven below). Avalable TTS data for two
md-frequency cetacean speces and three speces
of pnnpeds are used as the bass for estmatng
PTS-onset thresholds n all cetaceans (cetacean
procedure descrbed below; see PTS-Onset for
Pulses) and n all pnnpeds (see PTS-Onset
for Nonpulse Sounds), respectvely. The pro-
posed njury crtera are presented by sound type
because, for a gven sound type, many of the same
extrapolaton and summaton procedures apply
across marne mammal hearng groups.
A dual-crteron approach was used for the rec-
ommended njury crtera. That s, any receved
nose exposure that exceeds ether a peak pressure
or a SEL crteron for njury s assumed to cause
tssue njury n an exposed marne mammal. Of
the two measures of sound exposure, peak pres-
sures are to be unweghted (.e., flat-weghted),
whereas SEL metrcs are to be M-weghted for the
relevant marne mammal group (Fgure 1). In prac-
tce, the receved nose condtons should be com-
pared to the two exposure crtera for that sound
type and functonal hearng group, and the more
precautonary of the two outcomes accepted.
Effects of Noise on Hearing in
Marine Mammals: TTS Data
Nose exposure crtera for audtory njury deally
should be based on exposures emprcally shown to
nduce PTS-onset; however, no such data presently
exst for marne mammals. Instead, PTS-onset must
be estmated from TTS-onset measurements and
from the rate of TTS growth wth ncreasng expo-
sure levels above the level elctng TTS-onset. PTS
s presumed to be lkely f the threshold s reduced
by 40 dB (.e., 40 dB of TTS). We used avalable
marne mammal TTS data and precautonary extrap-
olaton procedures based on terrestral mammal
data (see Level of Nose Effect n Chapter 2)
to estmate exposures assocated wth PTS-onset.
Exstng TTS measurements for marne mammals
are revewed n detal here snce they serve as the
quanttatve foundaton for the njury crtera.
To date, TTSs measured n marne mammals
have generally been of small magntude (mostly
< 10 dB). The onset of TTS has been defned as
beng a temporary elevaton of a hearng thresh-
old by 6 dB (e.g., Schlundt et al., 2000), although
smaller threshold shfts have been demonstrated to
be statstcally sgnfcant wth a suffcent number
of samples (e.g., Kastak et al., 1999; Fnneran
et al., 2005a). Normal threshold varablty wthn
and between both expermental and control ses-
sons (no nose) does warrant a TTS-onset crte-
ron at a level that s always clearly dstngush-
able from that of no effect. We consdered a 6 dB
TTS suffcent to be recognzed as an unequvo-
cal devaton and thus a suffcent defnton of
TTS-onset.
Most of the frequences used n TTS exper-
ments to date are wthn the flat portons of the
M-weghtng functons gven here, but not nec-
essarly wthn the regons of greatest hearng
senstvty. Wthn the range of best hearng sen-
stvty for a gven ndvdual, detecton thresholds
are generally smlar. Wthn ths band, exposures
wth the same absolute level but dfferent fre-
quency are thus smlar n terms of ther effectve
sensaton level. Sensationlevel s the amount (n
dB) by whch an RL exceeds the threshold RL
for that sgnal type wthn a prescrbed frequency
band (Yost, 2000). If two exposures wth dentcal
absolute level are both audble, but one s outsde
the frequency range of best hearng senstvty,
sensaton level wll be less for the latter exposure,
and ts potental effects wll be dmnshed. By
creatng frequency-weghted functons that are
flat across vrtually the entre functonal hearng
band, rather than just the regon of best senstvty,
we have made another precautonary decson n
the absence of underlyng data on equal-loudness
functons.
Audtory fatgue (.e., TTS) n md-frequency
cetaceans has been measured after exposure to
tones, mpulsve sounds, and octave-band nose
(OBN). In pnnpeds, t has been measured upon
exposure to constructon nose and OBN n both
ar and water.
CetaceanTTS
The sound exposures that elct TTS n cetaceans
have been measured n two md-frequency spe-
cesbottlenose dolphn and beluga (specfc ref-
erences gven below)wth at least lmted data
AquaticMammals 2007, 33(4), 437-445, DOI 10.1578/AM.33.4.2007.437
438 Southalletal.
beng avalable for exposures to a sngle pulse and
to nonpulsed sounds rangng from 1-s to ~50-mn
duraton. There are no publshed TTS data for any
other odontocete cetaceans (ether md- or hgh-
frequency) or for any mystcete cetaceans (low-
frequency). Ths revew s organzed accordng to
the duraton of the fatgung stmulus, wth short-
est exposures dscussed frst.
Fnneran et al. (2000) exposed two bottlenose
dolphns and one beluga to sngle pulses from an
exploson smulator (ES). The ES conssted of
an array of pezoelectrc sound projectors that
generated a pressure waveform resemblng that
from a dstant underwater exploson. The pressure
waveform was generally smlar to waveforms
predcted by the Navy REFMS model (Brtt et al.,
1991). The ES faled to produce realstc energy
at frequences below 1 kHz, however. No substan-
tal (.e., 6 dB) threshold shfts were observed
n any of the subjects exposed to a sngle pulse at
the hghest receved exposure levels (peak: 70 kPa
[10 ps]; peak-to-peak: 221 dB re: 1 Pa (peak-to-
peak); SEL: 179 dB re: 1 Pa
2
-s)].
Fnneran et al. (2002b) repeated ths experment
usng a sesmc watergun that produced a sngle
acoustc pulse. Expermental subjects conssted of
one beluga and one bottlenose dolphn. Measured
TTS2 was 7 and 6 dB n the beluga at 0.4 and 30
kHz, respectvely, after exposure to ntense sngle
pulses (peak: 160 kPa [23 ps]; peak-to-peak: 226
dB re: 1 Pa (peak-to-peak); SEL: 186 dB re: 1
Pa
2
-s). Thresholds returned to wthn 2 dB of
the pre-exposure value wthn 4 mn of exposure.
No TTS was observed n the bottlenose dolphn
at the hghest exposure condton (peak: 207 kPa
[30 ps]; peak-to-peak: 228 dB re: 1 Pa (peak-to-
peak); SEL: 188 dB re: 1 Pa
2
-s). These studes
demonstrated that, for very bref pulses, hgher
sound pressures were requred to nduce TTS
than had been found for longer tones (dscussed
below).
Schlundt et al. (2000) reported TTS n fve bot-
tlenose dolphns and two belugas exposed to 1-s
pure tones (nonpulses). Ths paper also ncluded
a re-analyss of TTS data from a techncal report
by Rdgway et al. (1997). At frequences of 3 kHz,
10 kHz, and 20 kHz, SPLs necessary to nduce
TTS-onset were 192 to 201 dB re: 1 Pa (SEL:
192 to 201 dB re: 1 Pa
2
-s). The mean exposure
SPL for TTS-onset was 195 dB re: 1 Pa (195
dB re: 1 Pa
2
-s). Note the approprately dffer-
ent metrcs for the nonpulse sources used n ths
study and those nvolvng pulses. Also note that
the SPL and SEL values are dentcal n ths spe-
cal case because of the 1-s duraton fatgung
stmul. At 0.4 kHz, no subjects exhbted shfts
after exposures up to SPL exposures of 193 dB
re: 1 Pa (193 dB re: 1 Pa
2
-s). Data at 75 kHz
were nconclusve: one dolphn exhbted a TTS
after exposure at 182 dB SPL re: 1 Pa (182 dB
re: 1 Pa
2
-s) but not at hgher exposure levels. The
other dolphn experenced no threshold shft after
exposure to maxmum SPL levels of 193 dB re:
1 Pa (193 dB re: 1 Pa
2
-s). The shfts occurred
most often at frequences above the fatgung
stmulus.
Fnneran et al. (2005a) measured TTS n bot-
tlenose dolphns exposed to 3 kHz tones wth
duratons of 1, 2, 4, and 8 s and at varous SPL
values. Tests were conducted n a quet pool n
contrast to prevous studes n San Dego Bay,
where thresholds were masked by broadband
nose. Small amounts of TTS (3 to 6 dB) occurred
n one dolphn followng exposures wth SELs of
190 to 204 dB re: 1 Pa
2
-s. These results are con-
sstent wth those of Schlundt et al. (2000), nd-
catng that ther results had not been sgnfcantly
affected by the use of masked hearng thresholds
n quantfyng TTS. In general, the SEL necessary
for TTS-onset was relatvely consstent across the
range of exposure duratons, whereas exposure
SPL values causng TTS-onset tended to decrease
wth ncreasng exposure duraton. These results
confrmed that, for these testng condtons (bot-
tlenose dolphns exposed to 8-s tones of varable
SPL), TTS magntude was best correlated wth
exposure SEL rather than SPL.
Schlundt et al. (2006) reported on the growth
and recovery of TTS n a bottlenose dolphn
exposed to 3 kHz tones wth SPLs up to 200 dB
re 1 Pa and duratons up to 128 s. The maxmum
exposure SEL was 217 dB re 1 Pa
2
-s, whch pro-
duced a TTS4 of ~23 dB. All thresholds recovered
to baselne values wthn 24 h, most wthn 30
mn. The growth of TTS4 wth ncreasng expo-
sure SEL was ~1 dB TTS per dB SEL for TTS4 of
~15 to 18 dB.
Fnneran et al. (2007b) measured TTS n a
bottlenose dolphn after sngle and multple expo-
sures to 20 kHz tones. Hearng thresholds were
estmated at multple frequences (10 to 70 kHz)
both behavorally and electrophysologcally (by
measurement of multple audtory steady-state
responses). Three experments were performed.
The frst two featured sngle exposures (20 kHz,
64-s tones at 185 and 186 dB re 1 Pa). The thrd
featured three 20 kHz, 16-s exposures separated
by 11 and 12 mn, wth a mean SPL of 193 dB re
1 Pa (SD = 0.8 dB). Hearng loss was frequency-
dependent, wth the largest TTS occurrng at 30
kHz, less at 40, and then 20 kHz, and lttle or no
TTS at other measured frequences. AEP thresh-
old shfts reached 40 to 45 dB and were always
larger than behavoral shfts, whch were 19 to 33
dB. Complete recovery requred up to 5 d, wth
the recovery rate at 20 kHz beng ~2 dB/doublng
MarineMammalNoiseExposureCriteria 439
of tme and the rate at 30 and 40 kHz ~5 to 6 dB/
doublng of tme.
Nachtgall et al. (2003) measured TTS (ca. 20
mn after nose cessaton) n a bottlenose dolphn
and found an average 11 dB shft followng a 30-
mn net exposure to OBN wth a 7.5 kHz center
frequency (CF) (max SPL: 179 dB re: 1 Pa; SEL:
~212 to 214 dB re: 1 Pa
2
-s). The net exposure
tme was calculated as the total expermental tme
mnus the tme requred for the subject to surface
to breathe. Exposure durng breathng perods was
measured and factored nto the SEL measurement.
No TTS was observed after exposure to the same
OBN at maxmum SPL values of 165 and 171 dB
re: 1 Pa (SEL: ~198 to 200 dB re: 1 Pa
2
-s and
204 to 206 dB re: 1 Pa
2
-s, respectvely).
Usng AEP methods, Nachtgall et al. (2004)
found TTS5 of ca. 4 to 8 dB followng nearly 50-
mn exposures to OBN wth a CF of 7.5 kHz (max
SPL: 160 dB re: 1 Pa; SEL: ~193 to 195 dB re: 1
Pa
2
-s). The dfference n results between the two
Nachtgall et al. studes (slghtly lower TTS after
exposure to much lower exposure energy) was
attrbuted to measurng TTS at a shorter nterval
after the exposure ended (5 vs ~20 mn), and thus
allowng less opportunty for hearng recovery.
Further, Nachtgall et al. (2004) repeatedly mea-
sured hearng untl recovery had occurred. TTS
recovery was shown to occur wthn mnutes or
tens of mnutes, dependng on the amount of the
threshold shft. Generally, the recovery rate was
1.5 dB of recovery per doublng of tme and was
consstent n both studes (Nachtgall et al., 2003,
2004).
The Natonal Research Councl (NRC) (1994)
dentfed the need to know whether marne mam-
mals experence greatest TTS at a frequency
1
2-
octave above the frequency of exposure when
exposed to loud tones as has been shown n terres-
tral mammals. Nachtgall et al. (2004) observed
an average threshold shft of 4 dB at 8 kHz but 8
dB shft at 16 kHz followng the exposure to OBN
centered at 7.5 kHz as descrbed above. A smlar
upward frequency shft also has been observed by
Schlundt et al. (2000) and Fnneran et al. (2007b)
for md-frequency cetaceans. These fndngs pro-
vde strong evdence for fundamental smlartes
n cochlear mcromechancs n marne and land
mammals (NRC, 1994, p. 51) and further justfy
the judcous extrapolaton of TTS data wthn
marne mammal functonal hearng groups and
from terrestral to marne mammals.
The above results provde emprcal measures of
exposure condtons assocated wth TTS-onset n
md-frequency cetaceans exposed to sngle pulses
and nonpulses. Combned, these data demonstrate
that, as compared wth the exposure levels neces-
sary to elct TTS when exposure duraton s short,
lower SPLs (but smlar SEL values) are requred
to nduce TTS when exposure duraton s longer.
These fndngs are generally consstent wth mea-
surements n humans and terrestral mammals
(Kryter, 1970; Harrs, 1998; NIOSH, 1998) and
support the use of SEL to approxmate the aud-
tory effects of varable exposure level/duraton
condtons. Although there are certan (possbly
many) condtons under whch an explct equal-
energy rule may fal to adequately descrbe the
audtory effects of varable and/or ntermttent
nose exposure, the combned cetacean TTS data
presented above generally support the use of SEL
as a frst-order approxmaton, at least untl add-
tonal data are avalable.
For cetaceans, publshed TTS data are lmted
to the bottlenose dolphn and beluga (Fnneran
et al., 2000, 2002b, 2005a; Schlundt et al., 2000;
Nachtgall et al., 2003, 2004). Where data exst for
both speces, we use the more precautonary result
(usually for beluga) to represent TTS-onset for all
md-frequency cetaceans. No publshed data exst
on audtory effects of nose n ether low- or hgh-
frequency cetaceans (an area of needed research
as dscussed n Chapter 5); therefore, data from
md-frequency cetaceans are used as surrogates
for these two other groups (cetacean proce-
dure). [We are aware of some very recent TTS
measurements for an ndvdual harbor porpose
exposed to sngle pulses (Lucke et al., 2007a)
but lack suffcent detals regardng methodology
and data analyss to drectly consder those data
quanttatvely.]
Low-frequency cetaceans (mystcetes), based
on ther audtory anatomy (Wartzok & Ketten,
1999) and ambent nose levels n the frequency
ranges they use (Clark & Ellson, 2004), almost
certanly have poorer absolute senstvty (.e.,
hgher thresholds) across much of ther hearng
range than do the md-frequency speces (but
see earler dscusson). Md-frequency cetaceans
experence TTS-onset at relatvely hgh levels
compared wth ther absolute hearng senstvty
at smlar frequences (.e., hgh sensaton levels),
although t s not known that ths s smlarly char-
acterstc of low-frequency cetaceans. Our use of
TTS data from md-frequency cetaceans as a sur-
rogate for low-frequency cetaceans presumes that
the two groups have smlar audtory mechansms
and are not radcally dfferent n relatve senstv-
ty to fatgung nose, and that relatve dfferences
n absolute senstvty between the two groups are
generally as expected.
For hgh-frequency speces, data from md-
frequency cetaceans are currently used as a sur-
rogate n the absence of avalable group-specfc
data. Asde from ther extended upper-frequency
hearng, hgh-frequency cetaceans appear to be
440 Southalletal.
generally smlar n audtory anatomy and hear-
ng capabltes to md-frequency speces, though
there are some general dfferences between the
groups n sound producton. Based on avalable
nformaton and our extrapolaton procedures,
slghtly lower estmates of TTS-onset may be war-
ranted for hgh-frequency cetaceans exposed to
very hgh-frequency sounds ( 100 kHz). [Also,
prelmnary measurements of TTS n a harbor
porpose exposed to a sngle argun pulse (Lucke
et al., 2007a) suggest that ths speces may exper-
ence TTS-onset at levels lower than would be sug-
gested by extrapolatng from md-frequency ceta-
ceans. Those results, f confrmed, may provde a
more emprcal bass for estmatng TTS-onset n
hgh-frequency cetaceans and dervng group-spe-
cfc njury crtera.]
PinnipedTTS(UnderWater)
Sound exposures that elct TTS n pnnpeds under
water have been measured n ndvdual subjects of
three pnnped speces (harbor seal, Calforna sea
lon, and northern elephant seal). Avalable data
nvolved exposures to ether broadband or octave-
band nonpulse nose over duratons rangng from
~12 mn to several hours, plus lmted data on
exposure to underwater pulses. Interestngly,
there were consstent among-speces dfferences
n the exposure condtons that elcted TTS under
water. For the condtons tested, the harbor seal
experenced TTS at lower exposure levels than dd
the Calforna sea lon or northern elephant seal.
There are no underwater TTS data for any other
pnnped speces.
The followng revew frst consders expo-
sure to nonpulses, organzed chronologcally,
followed by a bref dscusson of the lone study
on exposure to pulses. All but one of the studes
(Fnneran et al., 2003) came from one laboratory
and from the same ndvdual test subjects. Kastak
& Schusterman (1996) reported a TTS of ~8 dB
(measured under water at 100 Hz) n a harbor seal
followng exposure to broadband arborne, non-
pulse nose from nearby constructon. Under con-
trolled condtons, Kastak et al. (1999) measured
TTS of ca. 4 to 5 dB n a harbor seal, Calforna
sea lon, and northern elephant seal followng 20-
to 22-mn exposure to underwater OBN centered
at frequences from 100 Hz to 2 kHz. Exposures
were normalzed to octave-band levels 60 to 75
dB above each subjects hearng threshold (.e., 60
to 75 dB sensaton level) to present smlar effec-
tve exposure condtons to each of the three sub-
jects. Because of ths approach, absolute exposure
values (n terms of both SPL and SEL) were qute
varable dependng on subject and test frequency.
Subsequently, Kastak et al. (2005) made TTS
measurements on the same subjects usng 2.5
kHz OBN, hgher sensaton levels (up to 95 dB),
and longer exposure duratons (up to 50-mn net
exposure). These data largely corroborate prev-
ous fndngs concernng TTS-onset n these pn-
npeds. They also support sensaton level as a rele-
vant metrc for normalzng exposures wth smlar
duratons across speces havng dfferent absolute
hearng capabltes. Comparatve analyses of
the combned underwater pnnped data (Kastak
et al., 2005) ndcated that, n the harbor seal, a
TTS of ca. 6 dB occurred wth 25-mn exposure to
2.5 kHz OBN wth SPL of 152 dB re: 1 Pa (SEL:
183 dB re: 1 Pa
2
-s). Under the same test cond-
tons, a Calforna sea lon showed TTS-onset at
174 dB re: 1 Pa (SEL: 206 dB re: 1 Pa
2
-s), and
a northern elephant seal experenced TTS-onset at
172 dB re: 1 Pa (SEL: 204 dB re: 1 Pa
2
-s).
Data on underwater TTS-onset n pnnpeds
exposed to pulses are lmted to a sngle study.
Fnneran et al. (2003) exposed two Calforna
sea lons to sngle underwater pulses from an
arc-gap transducer. They found no measurable
TTS followng exposures up to 183 dB re: 1 Pa
(peak-to-peak) (SEL: 163 dB re: 1 Pa
2
-s). Based
on the Kastak et al. (2005) measurements usng
nonpulse sounds, the absence of TTS for the sea
lons followng such exposures s generally not
surprsng.
PinnipedTTS(InAir)
Audtory fatgue has been measured followng
exposure of pnnpeds to sngle pulses of n-ar
sound and to nonpulse nose.
Bowles et al. (unpub. data) measured TTS-
onset for harbor seals exposed to smulated sonc
booms at peak SPLs of 143 dB re: 20 Pa (peak)
(SEL: 129 dB re: [20 Pa]
2
-s). Hgher exposure
levels were requred to nduce TTS-onset n both
Calforna sea lons and northern elephant seals n
the same test settng, consstent wth the results
for nonpulse sound both under water and n ar.
Audtory fatgue to arborne sound has also
been measured n the same three speces of pnn-
peds after exposure to nonpulse nose, specfcally
2.5 kHz CF OBN for 25 mn (Kastak et al., 2004a).
The harbor seal experenced ca. 6 dB of TTS at
99 dB re: 20 Pa (SEL: 131 dB re: [20 Pa]
2
-s).
Onset of TTS was dentfed n the Calforna
sea lon at 122 dB re: 20 Pa (SEL: 154 dB re:
[20 Pa]
2
-s). The northern elephant seal exper-
enced TTS-onset at 121 dB re: 20 Pa (SEL: 163
dB re: [20 Pa]
2
-s). The subjects n these tests
were the same ndvduals tested n water (Southall
et al., 2001; Kastak et al., 2005).
Kastak et al. (2007) measured TTS-onset and
growth functons for the same Calforna sea lon
exposed to a wder range of nose condtons. A
total of 192 exposure sequences were conducted
MarineMammalNoiseExposureCriteria 441
wth OBN (centered at 2.5 kHz) at levels 94 to
133 dB re: 20 Pa and duratons 1.5 to 50 mn
net exposure duraton. In these more ntense nose
exposures, TTS magntudes up to 30 dB were
measured at the 2.5 KHz test frequency. Full
recovery was observed followng all exposures;
ths occurred rapdly (lkely wthn tens of mn-
utes) for small shfts but took as long as 3 d n the
case of the largest TTS. The estmated SEL value
concdng wth TTS-onset across these vared
exposure condtons was 159 dB re: (20 Pa)
2
-s
wth a TTS growth functon of ~2.5 dB TTS/dB
nose. For TTS exceedng 20 dB, a recovery rate
of ~2.6 dB/doublng of tme was calculated. These
results generally agree wth those of Kastak et al.
(2004a) but provde a larger data set, across a
wder range of exposure condtons wth whch
to derve an emprcal TTS-growth functon. They
also support the concluson that patterns of TTS
growth and recovery are generally smlar to those
of terrestral mammals and that sensaton level for
the partcular speces and medum (water or ar) s
the approprate metrc for comparng the effects of
underwater and aeral nose exposure.
Injury from Noise Exposure:
PTS-Onset Calculation
As dscussed n Chapter 1, PTS s an rreversble
elevaton of the hearng threshold (.e., a reducton
n senstvty) at a specfc frequency (Yost, 2000).
Ths permanent change followng ntense nose
exposure results from damage or death of nner
or outer cochlear har cells. It s often followed by
retrograde neuronal losses and persstent chemcal
and metabolc cochlear abnormaltes (Saunders
et al., 1991; Ward, 1997; Yost, 2000).
Nose-nduced PTS represents tssue njury, but
TTS does not. Although TTS nvolves reduced
hearng senstvty followng exposure, t results
prmarly from the fatgue (as opposed to loss)
of cochlear har cells and supportng structures
and s, by defnton, reversble (Nordmann et al.,
2000). Many mammals, ncludng some pnnpeds
(Kastak et al., 1999, 2005) and cetaceans (e.g.,
Schlundt et al., 2000; Nachtgall et al., 2004),
demonstrate full recovery even after repeated
TTS. Snce TTS represents a temporary change n
senstvty wthout permanent damage to sensory
cells or support structures, t s not consdered to
represent tssue njury (Ward, 1997). Instead, the
onset of tssue njury from nose exposure s con-
sdered here as PTS-onset.
PTS as a functon of age (presbycusis; dscussed
n Chapter 1) generally appears to be a normal pro-
cess of agng n mammals (ncludng humans and
marne mammals), but no specfc allowance for
ths s ncluded n our proposed exposure crtera.
Data that would be needed to support alternate
crtera allowng for presbycuss are lackng. Our
approach, whch uses TTS data from subjects pre-
sumed to have normal hearng as the startng
pont for estmatng PTS-onset, s precautonary.
It s expected to overestmate damagng effects for
those ndvduals wth dmnshed absolute hear-
ng senstvty and/or functonal bandwdth pror
to the exposure.
Data on the effects of nose on terrestral mam-
mals can be useful n consderng the effects on
marne mammals n certan condtons (as ds-
cussed n Chapter 1) because of smlartes n
morphology and functonal dynamcs among
mammalan cochleae. Under that premse, t s
assumed that a nose exposure capable of nduc-
ng 40 dB of TTS wll cause PTS-onset n marne
mammals. Based on avalable data for terrestral
mammals, ths assumpton s lkely somewhat
precautonary as there s often complete recov-
ery from TTS of ths magntude or greater. Such
precauton s approprate, however, because the
precse relatonshp between TTS and PTS s not
fully understood, even for humans and small ter-
restral mammals despte hundreds of studes (see
Kryter, 1994; Ward, 1997). For marne mammals,
ths presumably complex relatonshp s unknown,
and lkely wll reman so. The avalable marne
mammal TTS data provde a bass for establsh-
ng a maxmum allowable amount of TTS up to
whch PTS s unlkely, however, and for conclud-
ng that PTS s ncreasngly lkely to occur above
ths pont. In usng TTS data to estmate the expo-
sure that wll cause PTS-onset, our approach s to
acknowledge scentfc uncertanty and to err on
the sde of overestmatng the possblty of PTS
(.e., on the sde of underestmatng the exposure
requred to cause PTS-onset).
In humans, when TTS2 magntude for a sngle
exposure exceeds ca. 40 dB, the lkelhood of
PTS begns to ncrease substantally (Kryter
et al., 1966; Kryter, 1994). Threshold shfts
greater than 40 dB have been demonstrated to
be fully recoverable after some perod of tme n
some terrestral mammal speces (human: Ward,
1959; Ahroon et al., 1996; chnchlla: Mller
et al., 1971; Mongolan gerbl [Merionesunguicu-
latus]: Boettcher, 1993). Generally, however, TTS
exceedng 40 dB requres a longer recovery tme
than smaller shfts, suggestng a hgher probablty
of rreversble damage (Ward, 1970) and possbly
dfferent underlyng mechansms (Kryter, 1994;
Nordman et al., 2000).
Our dervaton of proposed njury crtera for
marne mammals begns wth measured or est-
mated nose exposure condtons assocated wth
TTS-onset n cetaceans and pnnpeds. Procedures
for estmatng PTS-onset, assumed to occur n
442 Southalletal.
condtons causng 40 dB of TTS, were derved by
combnng (1) measured or estmated TTS-onset
levels n marne mammals and (2) the estmated
growth of TTS n certan terrestral mammals
exposed to ncreasng nose levels. The general
PTS-onset procedures dffer accordng to sound
type (pulses and nonpulses), the extent of avalable
nformaton, and requred extrapolaton. To est-
mate exposure condtons that wll result n PTS-
onset, SEL and SPL were consdered separately.
PTS-OnsetforPulses
Henderson & Hamernk (1986) reported that n
chnchllas exposed to pulses up to a certan level,
for each dB of added exposure above that whch
caused TTS-onset, a further TTS of about 0.5 dB
resulted. For the hghest exposure levels, as much
as 3 dB of addtonal TTS was found per addtonal
dB of nose. Thus, n extrapolatng TTS growth
functons from terrestral to marne mammals, a
precautonary approach s justfed such as usng
a slope nearer the upper extreme of ths range to
estmate the growth of TTS wth exposure level.
When dealng wth pulsed sound, to estmate SEL
exposures concdent wth PTS-onset, we assume
a slope of 2.3 dB TTS/dB nose. Ths s relatvely
precautonary n relaton to the data by Henderson
& Hamernk (1986) on chnchllas. Ths slope trans-
lates to an njury crteron (for pulses) that s 15
dB above the SEL of exposures causng TTS-onset
(defned above as 6 dB TTS). That s, PTS-onset
(40 dB TTS) s expected to occur on exposure to an
M-weghted SEL 15 dB above that assocated wth
TTS-onset ([40 dB TTS 6 dB TTS] / [2.3 dB TTS/
dB nose exposure] 15 dB nose exposure above
TTS-onset).
In terms of sound pressure, TTS-onset thresh-
olds n marne mammals, partcularly cetaceans,
are qute hgh (see above). The predcted PTS-
onset values would be very hgh (perhaps unreal-
stcally so as they would approach the cavtaton
lmt of water) f the aforementoned 15 dB df-
ference between TTS-onset and PTS-onset were
assumed. Consequently, an addtonal precauton-
ary measure was appled by arbtrarly assumng
that the pressure dfference between TTS-onset
and PTS-onset for pulses mght be just 6 dB. Ths
results n a TTS growth relatonshp of 6 dB
TTS/dB nose (.e., [40 dB TTS 6 dB TTS] / [6
dB TTS/dB nose exposure] 6 dB nose expo-
sure above TTS-onset). That s an extremely con-
servatve slope functon gven that t s double the
hghest rate found n chnchllas by Henderson &
Hamernk (1986). Ths 6 dB of added exposure,
above the exposure elctng TTS-onset, essen-
tally establshes a proposed (unweghted) peak-
pressure celng value for all sound types.
PTS-OnsetforNonpulseSounds
The peak pressure values assumed to be assocated
wth onset of njury (PTS-onset) are numercally
equvalent for nonpulse and pulse sounds. Among
other consderatons, ths allows for the possblty
that solated pulses could be embedded wthn the
predomnantly nonpulse sound.
To estmate the SEL value that would cause
PTS-onset for nonpulse sounds, we used the fol-
lowng procedure. In humans, each added dB
of nonpulse nose exposure above TTS-onset
results n up to 1.6 dB of addtonal TTS (Ward
et al., 1958, 1959). Assumng ths relatonshp
apples to marne mammals, ~20 dB of addtonal
nose exposure above that causng TTS-onset s
requred to nduce PTS-onset (.e., [40 dB TTS
6 dB TTS] / [1.6 dB TTS/dB nose exposure] =
21.3 dB of addtonal nose exposure). We rounded
ths down to a slghtly more precautonary value
of 20 dB of addtonal nose exposure above TTS-
onset. Consequently, to estmate PTS-onset and
derve the SEL njury crtera for nonpulses, we
add 20 dB to the M-weghted SEL values est-
mated to cause TTS-onset. The lone excepton
to ths approach s for pnnpeds n ar (dscussed
below) where a more precautonary TTS growth
rate was used based on a relatvely large emprcal
data set (Kastak et al., 2007).
Criteria for Injury from a Single Pulse
As per the PTS-Onset Calculaton secton of ths
chapter, the recommended crtera for njury from
exposure to a sngle pulse, expressed n terms of
peak pressure, are TTS-onset levels plus 6 dB of
addtonal exposure. In terms of SEL, the recom-
mended crtera are TTS-onset levels plus 15 dB
of addtonal exposure.
For all cetaceans exposed to pulses, the data
of Fnneran et al. (2002b) were used as the bass
for estmatng exposures that would lead to TTS-
onset (and, consequently, PTS-onset). They est-
mated that, n a beluga exposed to a sngle pulse,
TTS-onset occurred wth unweghted peak levels
of 224 dB re: 1 Pa (peak) and 186 dB re: 1 Pa
2
-s.
The latter s equvalent to a weghted (Mmf) SEL
exposure of 183 dB re: 1 Pa
2
-s as some of the
energy n the pulse was at low frequences to
whch the beluga s less senstve. Addng 6 dB
to the former (224 dB) values, the pressure cr-
teron for njury for md-frequency cetaceans s
therefore 230 dB re: 1 Pa (peak) (Table 3, Cell
4). Addng 15 dB to the latter (183 dB) value,
the M-weghted SEL njury crteron s 198 dB
re: 1 Pa
2
-s (Table 3, Cell 4). These results are
assumed to apply (see cetacean procedure, p. 439)
to low- and perhaps hgh-frequency cetaceans
(Table 3, Cells 1 & 7, respectvely) as well as to
MarineMammalNoiseExposureCriteria 443
md-frequency cetaceans. These njury crtera,
expressed n SEL, are slghtly more precautonary
than, but generally consstent wth, Kettens 1998
predcton (pers. comm.) that 30% of ndvdual
cetaceans exposed to pulses wth an SEL of 205
dB re: 1 Pa
2
-s would experence PTS.
For pnnpeds n water, there are no emprcal
data concernng the levels of sngle pulses that
would lead to TTS-onset. At least for the Calforna
sea lon, the requred exposure s expected to be
greater than 183 dB re: 1 Pa (peak) and 163 dB
re: 1 Pa
2
-s) because Fnneran et al. (2003) found
no TTS n two Calforna sea lons followng such
exposures. In the absence of specfc data on the
level of a sound pulse that would cause TTS-onset
for pnnpeds n water, we used a three-step pro-
cess to estmate ths value:
(1) We began wth the Fnneran et al. (2002b)
data on TTS-onset from sngle pulse expo-
sures n a md-frequency cetacean. TTS-
onset occurred wth a peak pressure of 224
dB re: 1 Pa (peak) and Mmf-weghted SEL
of 183 dB re: 1 Pa
2
-s.
(2) We assumed that the known pnnped-to-
cetacean dfference n TTS-onset upon
exposure to nonpulse sounds would also
apply (n a relatve sense) to pulses.
Specfcally, wth nonpulse sounds, harbor
seals experence TTS-onset at ca. 12 dB
lower RLs than do belugas (.e., 183 vs
195 dB re: 1 Pa
2
-s; Kastak et al., 1999,
2005; Southall et al., 2001; Schusterman
et al., 2003 vs Fnneran et al., 2000, 2005a;
Schlundt et al., 2000; Nachtgall et al., 2003,
2004). Assumng that ths dfference for
nonpulse sounds exsts for pulses as well,
TTS-onset n pnnpeds exposed to sngle
underwater pulses s estmated to occur at
a peak pressure of 212 dB re: 1 Pa (peak)
and/or an SEL exposure of 171 dB re: 1
Pa
2
-s. Each of these metrcs s 12 dB less
than the comparable value for md-frequency
cetaceans (see Fnneran et al., 2002b, and
above).
(3) As per the PTS-onset Procedure (dscussed
earler), we added 6 dB to the former (212
dB) value to derve the recommended njury
pressure crteron of 218 dB re: 1 Pa (peak)
(unweghted) for pnnpeds n water exposed
to a sngle pulse. Smlarly, we added 15 dB
to the latter value (171 dB) to derve the rec-
ommended M-weghted SEL njury crteron
of 186 dB re: 1 Pa
2
-s (Table 3, Cell 10).
These proposed crtera are lkely precauton-
ary because the harbor seal s the most sen-
stve pnnped speces tested to date, based
on results from a sngle ndvdual (Kastak
et al., 1999, 2005).
For pnnpeds n ar exposed to a sngle sound
pulse, the proposed crtera for njury were
based on measurements by Bowles et al. (unpub.
data), whch ndcated that TTS-onset n harbor
Table 3. Proposed njury crtera for ndvdual marne mammals exposed to dscrete nose events (ether sngle or multple
exposures wthn a 24-h perod; see Chapter 2)
Sound type
Marne mammal group Sngle pulses Multple pulses Nonpulses
Low-frequency cetaceans Cell 1 Cell 2 Cell 3
Sound pressure level 230 dB re: 1 Pa (peak) (flat) 230 dB re: 1 Pa (peak) (flat) 230 dB re: 1 Pa (peak) (flat)
Sound exposure level 198 dB re: 1 Pa
2
-s (Mlf) 198 dB re: 1 Pa
2
-s (Mlf) 215 dB re: 1 Pa
2
-s (Mlf)
Md-frequency cetaceans Cell 4 Cell 5 Cell 6
Sound pressure level 230 dB re: 1 Pa (peak) (flat) 230 dB re: 1 Pa (peak) (flat) 230 dB re: 1 Pa (peak) (flat)
Sound exposure level 198 dB re: 1 Pa
2
-s (Mmf) 198 dB re: 1 Pa
2
-s (Mmf) 215 dB re: 1 Pa
2
-s (Mmf)
Hgh-frequency cetaceans Cell 7 Cell 8 Cell 9
Sound pressure level 230 dB re: 1 Pa (peak) (flat) 230 dB re: 1 Pa (peak) (flat) 230 dB re: 1 Pa (peak) (flat)
Sound exposure level 198 dB re: 1 Pa
2
-s (Mhf) 198 dB re: 1 Pa
2
-s (Mhf) 215 dB re: 1 Pa
2
-s (Mhf)
Pnnpeds (n water) Cell 10 Cell 11 Cell 12
Sound pressure level 218 dB re: 1 Pa (peak) (flat) 218 dB re: 1 Pa (peak) (flat) 218 dB re: 1 Pa (peak) (flat)
Sound exposure level 186 dB re: 1 Pa
2
-s (Mpw) 186 dB re: 1 Pa
2
-s (Mpw) 203 dB re: 1 Pa
2
-s (Mpw)
Pnnpeds (n ar) Cell 13 Cell 14 Cell 15
Sound pressure level 149 dB re: 20 Pa (peak) (flat) 149 dB re: 20 Pa (peak) (flat) 149 dB re: 20 Pa (peak) (flat)
Sound exposure level 144 dB re: (20 Pa)
2
-s (Mpa) 144 dB re: (20 Pa)
2
-s (Mpa) 144.5 dB re: (20 Pa)
2
-s (Mpa)
Note: All crtera n the Sound pressure level lnes are based on the peak pressure known or assumed to elct TTS-onset,
plus 6 dB. Crtera n the Sound exposure level lnes are based on the SEL elctng TTS-onset plus (1) 15 dB for any type
of marne mammal exposed to sngle or multple pulses, (2) 20 dB for cetaceans or pnnpeds n water exposed to nonpulses,
or (3) 13.5 dB for pnnpeds n ar exposed to nonpulses. See text for detals and dervaton.
444 Southalletal.
seals occurs followng exposure to 143 dB re:
20 Pa (peak) and 129 dB re: (20 Pa)
2
-s. As for
underwater exposures to nonpulse sounds (Kastak
et al., 1999, 2005), hgher exposure levels were
requred to nduce TTS n Calforna sea lons and
northern elephant seals. Consequently, usng harbor
seal TTS data to establsh njury crtera for expo-
sure to a sngle aeral pulse n pnnpeds s lkely a
precautonary approxmaton. Based on these est-
mates of peak pressure and SEL assocated wth
TTS-onset, plus 6 dB and 15 dB, respectvely, to
estmate PTS-onset, the njury crtera for pnn-
peds exposed to a sngle aeral pulse are 149 dB re:
20 Pa (peak) (unweghted) and 144 dB re: (20 Pa)
2
-s,
M-weghted (Table 3, Cell 13).
Criteria for Injury from Multiple Pulses
For all marne mammal groups, the recommended
crtera for exposure to multple pulses, expressed
n both SPL and SEL unts, were numercally
dentcal to the crtera for a sngle pulse. Any
exposure n a seres that exceeds the peak pressure
crteron would be consdered potentally njur-
ous. In addton, the cumulatve SEL for multple
exposures should be calculated usng the summa-
ton technque descrbed n Chapter 1 (Appendx
A, eq. 5). The resultng SEL value for multple
pulses s then compared to the SEL njury crte-
ron for a sngle pulse n the same functonal hear-
ng group. As for the sngle pulse crtera, peak
pressures are unweghted (.e., flat-weghted),
but SEL should be weghted by the approprate
M-weghtng functon (Fgure 1).
For cetaceans, the proposed crtera for njury
by multple pulses are therefore 230 dB re:
1 Pa (peak) and, followng summaton, 198
dB re: 1 Pa
2
-s n terms of SEL (Table 3, Cells
2, 5 & 8). As for sngle pulses, ths approach s
consdered precautonary for md- and low-fre-
quency speces, but some cauton s warranted n
applyng t to hgh-frequency speces (cf. Lucke
et al., 2007a).
Followng the same logc, the proposed njury
pressure crteron for pnnpeds n water exposed
to multple pulses s 218 dB re: 1 Pa (peak) and
the njury SEL crteron s 186 dB re: 1 Pa
2
-s
(Table 3, Cell 11). For pnnpeds n ar, the pro-
posed njury pressure crteron for multple pulses
s 149 dB re: 20 Pa (peak) and the njury SEL cr-
teron s 144 dB re: (20 Pa)
2
-s (Table 3, Cell 14).
Criteria for Injury from Nonpulses
SPL and SEL appear to be approprate metrcs
for quantfyng exposure to nonpulse sounds. But
because SPL measures nvolve averagng over
some duraton, they may not adequately quantfy
hgh peak pressure transents embedded wthn
exposures of longer duraton but lower-pressure
magntude. There are related lmtatons wth SEL
n that temporal ntegraton s nvolved.
To account for the potentally damagng aspects
of hgh-pressure transents embedded wthn
nonpulse exposures, a precautonary approach
was taken, and the same peak pressure crteron
for njury proposed for sngle pulses s also rec-
ommended as the crteron for multple pulses n
all functonal hearng groups. Thus, f any compo-
nent of a nonpulse exposure (unweghted) exceeds
the peak pressure crteron, njury s assumed to
occur. We expect that only rarely wll the njury
pressure crteron for nonpulse sound be exceeded
f the njury SEL crteron s not exceeded (.e.,
the SEL crteron wll be the effectve crteron n
most exposure condtons).
For nonpulsed sounds, the recommended SEL
crtera for njury (PTS-onset) are M-weghted
exposures 20 dB hgher than those requred
for TTS-onset (see PTS-Onset Calculaton:
Nonpulses). Injury SEL crtera for multple non-
pulses are numercally dentcal to those for sngle
nonpulses for all hearng groups. We make no
dstncton between sngle and multple nonpulses
except that the cumulatve SEL for multple expo-
sures s calculated as descrbed n Chapter 1 and
Appendx A, eq. 5.
For all cetaceans exposed to nonpulses, the rec-
ommended pressure crteron for njury s 230 dB
re: 1 Pa (peak) (Table 3, Cells 3, 6, & 9), the same
crteron as for sngle pulses n these functonal
hearng groups. Injury SEL crtera are based on
TTS data for md-frequency speces and extrapo-
lated to the other cetacean groups (see cetacean
procedure, p. 439). The SEL crteron for non-
pulse njury n cetaceans s calculated to be an M-
weghted exposure of 215 dB re: 1 Pa
2
-s (Table 3,
Cells 3, 6 & 9). Ths s based on 195 dB re: 1 Pa
2
-s
as an estmate of TTS-onset n md-frequency ceta-
ceans (Fnneran et al., 2002b, 2005a; Schlundt
et al., 2000; Nachtgall et al., 2003, 2004) plus 20
dB to estmate PTS-onset. Applyng ths approach
to low-frequency cetaceans s consdered pre-
cautonary, but some cauton may be warranted
n extrapolatng to hgh-frequency cetaceans (cf.
sngle-pulse data of Lucke et al., 2007a).
We note that specal njury crtera, dfferent
from those shown n Cell 6 of Table 3, are lkely
needed for exposure of beaked whale speces
to nonpulses. Under certan condtons, beaked
whales of several speces (prmarly Cuvers,
Blanvlles, and Gervas beaked whales) have
stranded n the presence of sound sgnals from
tactcal md-frequency mltary sonars (Frantzs,
1998; Evans & England, 2001; Fernndez et al.,
2005; Cox et al., 2006). There have been other
MarineMammalNoiseExposureCriteria 445
ncdents (e.g., NMFS, 2005; Hohn et al., 2006)
where marne mammal strandngs or other anom-
alous events nvolvng other marne mammal
speces have occurred n assocaton wth
md-frequency sonar operatons. They are, how-
ever, much more ambguous, dffcult to nterpret,
and appear fundamentally dfferent than the spe-
cfc beaked whale events. Lttle s known about
the exposure levels, or about the postons or reac-
tons of other marne mammals n the areas durng
md-frequency sonar tranng operatons. The
most extreme, ultmate response of some beaked
whales n specfc condtons (strandng and sub-
sequent death) does not appear to be typcal of
other marne mammals.
Sound felds resultng from sonar operatons
have been modeled n several of the above cases
(e.g., the 1996 event n Greece and the 2000
event n the Bahamas), and t s possble to at
least roughly bound the estmated exposures for
some of the ndvduals that stranded (DSpan
et al., 2006). Whle the specfc exposure levels
wll never be quanttatvely known, t does appear
lkely that the exposures for some of the beaked
whales that stranded were below the crtera for
tssue njury proposed above.
Consequently, the general njury crtera do not
seem suffcently precautonary for beaked whales
exposed to some nonpulse sounds under certan
condtons. Emprcal data to support dscrete,
scence-based njury crtera specfc to beaked
whales exposed to tactcal, md-frequency, ml-
tary sonar are lackng, however. Regulatory agen-
ces should consder adoptng provsonal njury
crtera for beaked whales exposed to actve, md-
frequency, mltary sonars that are lower (n terms
of RL) than the crtera used for md-frequency
cetaceans and nonpulse sources generally. Of
foremost mportance, specfc studes are needed
to better defne the mechansm of njury n these
apparently senstve speces (see Chapter 5).
For pnnpeds n water, the recommended pres-
sure crteron for njury from exposure to nonpulse
sounds s the same value as appled to pulses:
an unweghted value of 218 dB re: 1 Pa (peak)
(Table 3, Cell 12). To derve the assocated SEL
crteron, we began wth the measured nonpulse
exposure elctng TTS-onset n a harbor seal, 183
dB re: 1 Pa
2
-s (Kastak et al., 1999, 2005). Ths s
lkely a precautonary choce because SEL values
~10 to 20 dB hgher were requred to nduce TTS-
onset n a Calforna sea lon and a northern ele-
phant seal. We assume that 20 dB of addtonal
nose exposure wll elct PTS-onset (see Effects
of Nose on Hearng secton of ths chapter),
resultng n an Mpw-weghted SEL crteron of 203
dB re: 1 Pa
2
-s for pnnpeds exposed to nonpulse
sound n water (Table 3, Cell 12).
For pnnpeds n ar exposed to nonpulse sound,
the njury pressure crteron s a flat-weghted value
of 149 dB re: 20 Pa (peak) (Table 3, Cell 15), con-
sstent wth that for pulses. The SEL crteron s
based on occurrence of TTS-onset n a harbor seal
exposed n ar to 131 dB re: (20 Pa)
2
-s (Kastak
et al., 2004a). In estmatng the exposure that
would cause PTS-onset, we use emprcal mea-
surements of TTS growth as a functon of expo-
sure SEL n a Calforna sea lon. Kastak et al.
(2007) found a TTS growth rate of 2.5 dB TTS/dB
nose based on nearly 200 exposure sequences
nvolvng varable exposure level and duraton
condtons. Ths growth rate mples a 13.5 dB df-
ference between TTS- and PTS-onset as opposed
to the 20 dB value used for marne mammals n
water. When the 13.5 dB fgure s added to the
TTS-onset value for harbor seals (131 dB re: [20
Pa]
2
-s), we obtan a proposed Mpa-weghted SEL
crteron of 144.5 dB re: (20 Pa)
2
-s for pnnpeds
n ar (Table 3, Cell 15).
The use for all pnnpeds of harbor seal TTS
data combned wth the sea lon growth functon
would be an exceedngly precautonary procedure.
Ths PTS-onset estmate s consderably below
the TTS-onset estmates for both the northern ele-
phant seal (163 dB re: [20 Pa]
2
-s; Kastak et al.,
2004a) and the Calforna sea lon (159 dB re: [20
Pa]
2
-s; Kastak et al., 2007). Applyng the TTS
growth functon of 2.5 dB TTS/dB nose from
Kastak et al. (2007) to these TTS-onset estmates
would yeld PTS-onset values of 172.5 and 176.5
dB re: (20 Pa)
2
-s for the Calforna sea lon and
northern elephant seal, respectvely. As noted n
the Overvew, where specfc data are avalable
for the speces or genus of concern, t s appropr-
ate for crtera to be based on those data rather than
the generalzed crtera that are recommended for
the overall group of marne mammals.
4. Criteria for Behavioral Disturbance
Behavoral reactons to acoustc exposure are
generally more varable, context-dependent, and
less predctable than effects of nose exposure on
hearng or physology. Anmals detectng one knd
of sgnal may smply orent to hear t, whereas
they mght panc and flee for many hours upon
hearng a dfferent sound, potentally even one
that s queter, but wth some partcular sgnf-
cance to the anmal. The conservaton of cochlear
propertes across mammals justfes judcous
applcaton of audtory data from terrestral mam-
mals where data on marne mammals are mssng.
However, the context-specfcty of behavoral
responses n anmals generally makes extrapola-
ton of behavoral data napproprate. Assessng
the severty of behavoral dsturbance must conse-
quently rely more on emprcal studes wth care-
fully controlled acoustc, contextual, and response
varables than on extrapolatons based on shared
phylogeny or morphology.
Consderable research has been conducted
to descrbe the behavoral responses of marne
mammals to varous sound sources. Fortunately,
at least lmted data are avalable on behavoral
responses by each of the fve functonal marne
mammal groups to each sound type consdered
here. As evdent n the extensve lterature revew
summarzed below and descrbed n detal n
Appendces B & C, however, very few studes
nvolvng suffcent controls and measurements
exst. In addton, the nfluence of experence wth
the expermental stmulus or smlar sounds has
usually been unknown.
To assess and quantfy adverse behavoral
effects of nose exposure, a metrc for the mpact
such changes mght have on crtcal bologcal
parameters such as growth, survval, and reproduc-
ton s needed. Behavoral dsturbances that affect
these vtal rates have been dentfed as partcularly
mportant n assessng the sgnfcance of nose
exposure (NRC, 2005). Unfortunately, as Wartzok
et al. (2004) ponted out, no such metrc s cur-
rently avalable, and t s lkely to take decades of
research to provde the analytcal framework and
emprcal results needed to create such a metrc, f
one n fact s ultmately even vable.
In humans, a common and useful means of est-
matng behavoral dsturbance from nose expo-
sure s to ask ndvduals to rate or descrbe the
degree to whch varous sounds are bothersome.
Subjectve percepton of nose annoyance has
been quantfed (e.g., Schultz, 1978; Angerer
et al., 1991) and used to develop dose-response
relatonshps for nose exposure n human com-
munty nose applcatons (see Kryter, 1994,
Chapter 10). Practcal ssues (e.g., dffcultes n
tranng nonverbal speces to provde nterpretable
responses and questons about the applcabl-
ty of captve data to free-rangng anmals) have
prevented ths or smlar approaches from beng
appled to marne mammals. Instead, most efforts
have focused on analyses of observable reactons
to known nose exposure.
For most free-rangng marne mammals, behav-
oral responses are often dffcult to observe. Also,
precse measurements of receved nose exposure
and other relevant varables (e.g., movement of
source, presence of hgh-frequency harmoncs
ndcatng relatve proxmty, and pror experence
of exposed ndvduals) can be dffcult to obtan.
Only a subset of dsturbance studes have est-
mated receved sound levels, and only a very small
number have actually measured RLs at the subject.
Further, exposures are often complcated by mul-
tple contextual covarants such as the presence of
vessels and/or humans close to subjects ether for
observaton or to deploy playback sources (e.g.,
Frankel & Clark, 1998). Interpretaton of the
observed results s hghly lmted by uncertanty
as to what does and does not consttute a mean-
ngful response. Also, most behavoral-response
studes have concentrated on short-term and local-
zed behavoral changes whose relevance to nd-
vdual well-beng and ftness, let alone populaton
parameters, s lkely to be low.
A further complcaton s that observatons from
laboratory and feld settngs cannot be drectly
equated. Laboratory studes are usually precse n
quantfyng exposures and responses. The expo-
sure condtons very rarely approxmate those n
the feld, however, and measured behavor may
have lttle or no relevance to the ways n whch
unconstraned, untraned wld anmals respond.
Conversely, feld measurements may address
responses of free-rangng mammals to a specfc
sound source but often lack adequate controls and
precson n quantfyng acoustc exposures and
responses. Clearly, there s a need for a framework
to ntegrate laboratory and feld data, despte the
challenges n constructng that framework.
Another dffcult ssue concerns the appropr-
ate nose exposure metrc for assessng behavoral
AquaticMammals 2007, 33(4), 446-473, DOI 10.1578/AM.33.4.2007.446
MarineMammalNoiseExposureCriteria 447
reactons. Most boacoustcans recommend
reportng several dfferent measures of acoustc
exposure, such as SPL and SEL (as n Blackwell
et al., 2004a, 2004b). Of the many studes that
report source SPL, relatvely few specfy whether
RMS, peak, peak-to-peak, or other sound pressure
measurements were made. Addtonally, relatvely
few papers provde suffcent relevant nforma-
ton about sound transmsson loss n the study
area. A small number of papers report estmates or
drect measurements of receved SPL, but very few
report SEL. The approprate measure for predct-
ng probablty of a behavoral response s lkely
to vary dependng upon the behavoral context. For
example, f an anmal nterprets a sound as ndcat-
ng the presence of a predator, a short fant sgnal
may evoke as strong a response as a longer, strong
sound. But f an anmal s respondng to a context-
neutral stmulus that s merely annoyng, the prob-
ablty of response may well scale wth duraton and
level of exposure.
It s dffcult to defne the SEL for ndvdual
anmals n the wld exposed to a specfc sound
source. Ideally, receved SEL over the anmals
full duraton of exposure would be measured
(Madsen et al., 2005a). We expect that the prob-
ablty and severty of some knds of response wll
vary wth duraton as well as level of exposure;
for those stuatons, an SEL metrc may be most
approprate. However, the most practcal way to
look for consstent patterns of response as a func-
ton of RL and duraton, gven the current state
of scence, s to evaluate how dfferent anmals
respond to smlar sound sources used n smlar
contexts. For example, the relatonshp between
acoustc exposure and anmal responses s lkely
to be qute dfferent for mammals exposed to
sounds from a slow-movng sesmc survey vessel
operatng n a gven habtat for many weeks as
compared wth a torpedo transmttng drectonal
hgh-frequency sonar pngs as t transts an area
once at many tens of knots. Smlarly, an acous-
tc harassment devce placed n a habtat for years
s lkely to evoke a dfferent severty of response
than would several short pulses at a comparable
SPL. Untl more controlled studes become aval-
able wth calbrated measurements of RLs and
ambent nose measurements (ncludng sgnal-to-
nose rato), the best way to predct lkely effects
wll be a common-sense approach that assesses
avalable data from stuatons smlar to the stu-
aton of concern.
Consderng all of these lmtatons and the
nature of the avalable data, as a practcal matter,
we use SPL as the acoustc metrc for the behav-
oral analyses gven below. Where necessary and
approprate, smple assumptons regardng trans-
msson loss were appled to predct RLs. Ths
was done only for studes that provded suffcent
nformaton on source and envronmental charac-
terstcs. Our approach does not presume that SPL
s necessarly the acoustc metrc best correlated
wth behavoral changes (sgnfcant or otherwse).
In partcular, SPL fals to account for the dura-
ton of exposure whereas ths s captured usng
SEL. SPL s the metrc that has most often been
measured or estmated durng dsturbance studes,
however. Thus, t s currently the best metrc wth
whch to assess the avalable behavoral response
data. Future studes should report the full range of
standard acoustc measurements approprate to the
sound source n queston and should also nclude
measurements of background nose levels n order
to assess sgnal-to-nose ratos. These addtonal
data should eventually clarfy whch exposure
metrcs best predct dfferent knds of behavoral
responses and whch are most approprate for use
n polcy gudelnes applcable to dfferent types
of nose exposures.
Beyond the dscusson of whch metrc s most
approprate to quantfy the exposure level of a
sound, t s recognzed that many other varables
affect the nature and extent of responses to a par-
tcular stmulus. Wartzok et al. (2004) dscussed
n detal the hghly varable response of belugas
exposed to smlar sounds n dfferent locatons
for example, Frost et al. (1984) vs Fnley et al.
(1990). In those cases, t appears that the context
(recent experence of the belugas wth the sound
stmulus, ther current actvty, and ther motva-
ton to reman or leave) was much more sgnfcant
n governng ther behavoral responses. Smlarly,
reactons of bowhead whales to sesmc argun
sounds depend on whether the whales are feedng
(Rchardson et al., 1986; Mller et al., 2005) vs
mgratng (Rchardson et al., 1999). Reactons of
bowheads and other cetaceans to boats depend on
whether the boats are movng or statonary, and on
the relatve movement of the boat and the whale
(see Rchardson et al., 1995; Wartzok et al., 2004).
In these and some other cases, smple metrcs of
exposure (wthout consderng context) wll not
relably predct the type and severty of behav-
oral response(s). Our analyses here, whch use
exposure SPL alone, are admttedly rudmentary
and lmted by the fact thatfor most speces and
stuatonscurrent data do not support a more
sophstcated approach.
Another key consderaton nvolves dffer-
entatng bref, mnor, bologcally unmportant
reactons from profound, sustaned, and/or bo-
logcally meanngful responses related to growth,
survval, and reproducton. The bologcal rel-
evance of a behavoral response to nose expo-
sure may depend n part on how long t perssts.
Many mammals perform vtal functons (e.g.,
448 Southalletal.
feedng, restng, travelng, socalzng) on a del
cycle. Repeated or sustaned dsrupton of these
functons s more lkely to have a demonstrable
effect on vtal rates than a sngle, bref dsturbance
epsode. The NRC (2005) argued that, although
the duraton of behavors lkely to affect vtal rates
s beleved to be partcularly sgnfcant, current
scentfc knowledge s nsuffcent to support an
analytcal treatment of bologcal sgnfcance and
ad hoc crtera are needed n the nterm. Here,
substantve behavoral reactons to nose expo-
sure (such as dsrupton of crtcal lfe functons,
dsplacement, or avodance of mportant habtat)
are consdered more lkely to be sgnfcant f they
last more than one del perod, or recur on subse-
quent days. Consequently, a reacton lastng less
than 24 h and not recurrng on subsequent days s
not regarded as partcularly severe unless t could
drectly affect survval or reproducton.
In the absence of an overarchng means of quan-
tfyng the bologcal sgnfcance of an effect, we
had to adopt a more descrptve method of assess-
ng the range of possble responses and the sever-
ty of behavoral response. To do ths, we took
two dfferent approaches. For the unusual case of
exposure to a sngle pulse, where the exposure s
very bref and responses are usually bref as well,
a procedure for determnng recommended crtera
s dentfed and appled. For all other condtons,
an ordnal and subjectve response severty scal-
ng was developed and appled to those data on
marne mammal behavoral responses for whch
estmates of receved SPL were avalable. These
analyses were lmted to peer-revewed lterature
(publshed or n press) and peer-revewed techn-
cal reports, wth some exceptons on a case-by-
case bass.
The severty scale was desgned to provde
some analytcal bass for assessng bologcal
sgnfcance, but t had to be rooted n the knds
of descrptons provded n the avalable scen-
tfc lterature. Our current understandng of the
nfluences of contextual varables on behavoral
responses n free-rangng marne mammals s
very lmted. The analyses presented here should
be consdered wth these cautons and caveats n
mnd. Our goal was to revew the relevant scen-
tfc lterature, tally behavoral effects by the type
of acoustc exposure for each category of marne
mammal and sound type, and draw what conclu-
sons were approprate based on the nformaton
avalable.
The general procedures for determnng behav-
oral response exposure crtera for a sngle pulse,
and for conductng the severty analyses of nd-
vdual behavoral responses vs receved SPL, are
dscussed n the next secton. Subsequent sectons
dscuss the exposure crteron levels for sngle
pulses and summarze the lterature consdered n
the severty scalng analyses for multple pulses
and nonpulse sources. More detaled dscus-
sons of ths lterature are gven n Appendx B
for multple pulses and Appendx C for nonpulse
sources.
Behavioral Response Data Analysis Procedures:
Disturbance Criteria and Severity Scaling
SinglePulse
Due to the transent nature of a sngle pulse, the
most severe behavoral reactons wll usually be
temporary responses, such as startle, rather than
prolonged effects, such as modfed habtat utl-
zaton. A transent behavoral response to a sngle
pulse s unlkely to result n demonstrable effects
on ndvdual growth, survval, or reproducton.
Consequently, for the unque condton of a sngle
pulse, an audtory effect s used as a defacto ds-
turbance crteron. It s assumed that sgnfcant
behavoral dsturbance mght occur f nose expo-
sure s suffcent to have a measurable transent
effect on hearng (.e., TTS-onset). Although TTS
s not a behavoral effect per se, ths approach s
used because any compromse, even temporar-
ly, to hearng functons has the potental to affect
vtal rates by nterferng wth essental commun-
caton and/or detecton capabltes. Ths approach
s expected to be precautonary because TTS at
onset levels s unlkely to last a full del cycle or to
have serous bologcal consequences durng the
tme TTS perssts. Because ths approach s based
on an audtory phenomenon, the exposure crtera
can reasonably be developed for entre functonal
hearng groups (as n the njury crtera) rather
than on a speces-by-speces bass. The extrapo-
laton procedures used to estmate TTS-onset for
sngle pulse exposures for each hearng group are
descrbed n Chapter 3 (see the Injury from Nose
Exposure: PTS-Onset Calculaton secton).
A dual-crteron approach (usng both SPL
[peak] and SEL) was used to determne behavoral
crtera for a sngle pulse exposure. Consstent
wth the njury crtera, whch also were based on
audtory fatgue data, RLs that exceed the crteron
for ether metrc are consdered to have greater
potental to elct a bologcally sgnfcant behav-
oral response. Proposed crtera for exposure to
a sngle pulse for each functonal hearng group
are gven n the next secton. These crtera are the
TTS-onset thresholds dscussed n Chapter 3.
An excepton was made n any case where
behavoral data ndcate that a sngle pulse expo-
sure may elct a sustaned and potentally sgnf-
cant response when the RL s below that requred
for TTS-onset. Ths can apply to hauled-out pn-
npeds, whch sometmes stampede from a beach
MarineMammalNoiseExposureCriteria 449
upon exposure to a sonc boom and may not return
for many hours (e.g., Holst et al., 2005a, 2005b).
In cases where such behavoral responses may
result n the njury or death of pups or other nd-
vduals, exposure levels should be consdered n
the context of njury crtera. Conversely, f aval-
able behavoral data ndcate that the response
threshold for exposure to a sngle pulse s above
the level requred for TTS-onset, then the TTS-
onset level s retaned as the behavoral crteron
as a further precautonary procedure.
MultiplePulsesandNonpulses
For all other sound types than sngle pulses, we
expect that sgnfcant behavoral effects wll occur
more commonly at levels below those nvolved n
temporary or permanent losses of hearng sens-
tvty. Ths argues aganst basng threshold crtera
exclusvely on TTS and ndcates the need for a
paradgm to predct the probablty of sgnfcant
behavoral response as a functon of nose expo-
sure. However, because of the extreme degree
of group, speces, and ndvdual varablty n
behavoral responses n varous contexts and con-
dtons, t s less approprate to extrapolate behav-
oral effects as opposed to audtory responses.
The avalable data on marne mammal behavoral
responses to multple pulse and nonpulse sounds
are smply too varable and context-specfc to jus-
tfy proposng sngle dsturbance crtera for broad
categores of taxa and of sounds.
Ths should not, however, lead to the concluson
that there are nsuffcent data to conduct a system-
atc assessment of the lkelhood that certan sound
exposures wll nduce behavoral effects of varable
serousness n marne mammals. On the contrary,
ths feld has seen many and acceleratng strdes
n characterzng how certan knds of sounds can
affect marne mammal behavor. Quantfcaton
of the severty or sgnfcance of these effects wll
contnue to be challengng. However, based on
the NRC (2005) model descrbed above n whch
behavoral reactons wth a greater potental to
affect vtal rates are of partcular concern, a sm-
plstc scalng paradgm n whch to consder the
avalable data appears to provde the most justf-
able way forward at present.
Frst, we developed an ordnal rankng of
behavoral response severty (see Table 4). The
ntent of ths scalng was to delneate those behav-
ors that are relatvely mnor and/or bref (scores
0-3); those wth hgher potental to affect forag-
ng, reproducton, or survval (scores 4-6); and
those consdered lkely to affect these vtal rates
(scores 7-9). Ths s an admttedly smplstc
way of scalng the strkngly complex and poorly
understood behavoral patterns of marne mam-
mals n real-world condtons. It does provde a
rudmentary framework for assessng the relatve
bologcal mportance of behavoral responses and
s lkely a closer approxmaton of realty than pre-
vous step-functon thresholds (as dscussed n the
Hstorcal Perspectves secton of Chapter 1).
Ths approach emphaszes that dsturbance s a
graduated, rather than a yes-or-no, phenomenon
and that some nose-nduced changes n behavor
are more sgnfcant than others. We expect that
future studes nvolvng multvarate analyss of
multple behavoral response varables, multple
measures of acoustc exposure, and multple con-
textual varables wll provde a foundaton for
more sophstcated nterpretatons.
Second, we revewed avalable research and
observatons for each of the fve marne mammal
functonal hearng groups exposed to ether mul-
tple pulse or nonpulse sounds (.e., Cells 2, 3, 5,
6, 8, 9, 11, 12, 14 & 15 n our matrx of sound
type by anmal group). We consdered measure-
ments of behavoral response both n the feld
and n the laboratory accordng to the behavoral
severty scale. Studes wth nsuffcent nforma-
ton on exposures and/or responses were con-
sdered but not ncluded n the severty analyss.
Where ndvdual (and/or groups consdered as an
ndvdual; see below) behavoral responses and
assocated receved sound levels were reported,
the observatons were assgned the approprate
behavoral score from Table 4 and the case was
ncluded n a severty scorng table for the relevant
matrx cell. One dmenson n ths type of table
was the behavoral score (defned n Table 4);
the other dmenson was the receved SPL wthn
10-dB ranges. Where multple responses were
reported for the same ndvdual and/or group n a
study (or where t was possble that ths had been
donepseudoreplcaton), approprate fractons
of a sngle observaton were assgned to relevant
cells n the scorng table. As a result, there are frac-
tonal responses for some ndvdual and/or group
responses n the tabular severty-scalng forms.
For example, a sngle behavoral observaton for
one ndvdual was weghted as equvalent to ten
observatons for another ndvdual by assgnng
each observaton (some potentally n dfferent
RL/severty score bns) of the second ndvdual a
relatve weght of 0.1.
Many observatons of marne mammals nvolve
multple ndvduals because many speces occur
n large socal groups and are followed as a group.
In ths case, f one ndvdual responds to a sound,
the other group members may respond to the
response as opposed to the sound. In such obser-
vatons, the full group was consdered to repre-
sent an ndvdual (.e., the group became the
unt of analyss). As a precautonary approach, the
most severe response by any ndvdual observed
450 Southalletal.
Table 4. Severty scale for rankng observed behavoral responses of free-rangng marne mammals and laboratory subjects
to varous types of anthropogenc sound
Response
score
1
Correspondng behavors
(Free-rangng subjects)
2
Correspondng behavors
(Laboratory subjects)
2
0 - No observable response - No observable response
1 - Bref orentaton response (nvestgaton/vsual orentaton) - No observable response
2 - Moderate or multple orentaton behavors
- Bref or mnor cessaton/modfcaton of vocal behavor
- Bref or mnor change n respraton rates
- No observable negatve response; may
approach sounds as a novel object
3 - Prolonged orentaton behavor
- Indvdual alert behavor
- Mnor changes n locomoton speed, drecton, and/or dve
profle but no avodance of sound source
- Moderate change n respraton rate
- Mnor cessaton or modfcaton of vocal behavor (duraton
< duraton of source operaton), ncludng the Lombard Effect
- Mnor changes n response to traned
behavors (e.g., delay n statonng,
extended nter-tral ntervals)
4 - Moderate changes n locomoton speed, drecton, and/or dve
profle but no avodance of sound source
- Bref, mnor shft n group dstrbuton
- Moderate cessaton or modfcaton of vocal behavor (duraton
duraton of source operaton)
- Moderate changes n response to
traned behavors (e.g., reluctance to
return to staton, long nter-tral
ntervals)
5 - Extensve or prolonged changes n locomoton speed, drecton,
and/or dve profle but no avodance of sound source
- Moderate shft n group dstrbuton
- Change n nter-anmal dstance and/or group sze (aggregaton
or separaton)
- Prolonged cessaton or modfcaton of vocal behavor
(duraton > duraton of source operaton)
- Severe and sustaned changes n
traned behavors (e.g., breakng away
from staton durng expermental
sessons)
6 - Mnor or moderate ndvdual and/or group avodance of sound
source
- Bref or mnor separaton of females and dependent offsprng
- Aggressve behavor related to nose exposure (e.g., tal/flpper
slappng, fluke dsplay, jaw clappng/gnashng teeth, abrupt
drected movement, bubble clouds)
- Extended cessaton or modfcaton of vocal behavor
- Vsble startle response
- Bref cessaton of reproductve behavor
- Refusal to ntate traned tasks
7 - Extensve or prolonged aggressve behavor
- Moderate separaton of females and dependent offsprng
- Clear ant-predator response
- Severe and/or sustaned avodance of sound source
- Moderate cessaton of reproductve behavor
- Avodance of expermental stuaton
or retreat to refuge area ( duraton of
experment)
- Threatenng or attackng the sound
source
8 - Obvous averson and/or progressve senstzaton
- Prolonged or sgnfcant separaton of females and dependent
offsprng wth dsrupton of acoustc reunon mechansms
- Long-term avodance of area (> source operaton)
- Prolonged cessaton of reproductve behavor
- Avodance of or senstzaton to exper-
mental stuaton or retreat to refuge
area (> duraton of experment)
9 - Outrght panc, flght, stampede, attack of conspecfcs, or
strandng events
- Avodance behavor related to predator detecton
- Total avodance of sound exposure
area and refusal to perform traned
behavors for greater than a day
1
Ordnal scores of behavoral response severty are not necessarly equvalent for free-rangng vs laboratory condtons.
2
Any sngle response results n the correspondng score (.e., all group members and behavoral responses need not be
observed). If multple responses are observed, the one wth the hghest score s used for analyss.
MarineMammalNoiseExposureCriteria 451
wthn a group was used as the rankng for the
whole group.
A specfc category of behavoral studes was
one n whch marne mammal dstrbutons were
measured around a sound source durng quet and
actve perods. The avalable data typcally nvolve
comparsons of the dstrbuton of anmals before
exposure (control or reference) vs durng expo-
sure (expermental); the dfference n dstrbuton
of the group was the behavoral response. Usng
ths method, and gven equvalent range measure-
ments for control and expermental observatons,
phantom RLs for mammals detected durng
control perods (RLs that would have exsted f n
fact the source was actve) can be calculated and
compared to actual RLs durng expermental con-
dtons. In ths way, the percentage of avodance
responses by ndvduals durng the exposure was
then calculated.
For the studes used n ths analyss, nose
exposure (ncludng source and RL, frequency,
duraton, duty cycle, and other factors) was ether
drectly reported or was reasonably estmated
usng smple sound propagaton models deemed
approprate for the sources and operatonal env-
ronment. Because of the general lack of precson
n many studes and the dffcultes n poolng the
results from dsparate studes here, we pooled
ndvdual exposure SPL nto 10-dB bns.
Our analyss of the avalable behavoral
response studes presents raw, ndvdual obser-
vatons of reactons to multple pulses and non-
pulses as a functon of exposure RL. The basc
output of ths procedure s a seres of tables, one
for each combnaton of the fve marne mammal
functonal hearng groups and these two sound
types (multple pulses and nonpulses). The over-
all tally wthn each cell represents the number of
ndvduals and/or ndependent group behavoral
responses wth estmated and/or measured RL n
the specfed 10-dB category.
Ths analyss s ntended to provde some
foundaton for judgng the degree to whch aval-
able data suggest the exstence of dose-response
relatonshps between nose exposure and marne
mammal behavor. An example of such a dose-
response functon s the Schultz (1978) curve
used to predct growth of human annoyance wth
ncreasng nose level. The reader should note,
however, that the substantal, acknowledged cave-
ats and lmtatons of the current approach, partc-
ularly those related to contextual varables other
than smply exposure level. Any applcaton of
the severty analyses gven below should carefully
consder the nature of the avalable nformaton
regardng sound source, speces, sex/age class,
sound-propagaton envronment, and especally
the overall context of exposure relatve to that
shown n the studes revewed here. The results
from pror behavoral studes n whch these var-
ables are farly smlar to those n the antcpated
exposure stuaton wll very lkely be the most rel-
evant. Informaton from those studes should be
most strongly weghted n assessng the lkelhood
of sgnfcant behavoral dsturbance.
Criteria for Behavioral Disturbance: Single Pulse
For all cetaceans exposed to sngle pulses, the
crtera were based on the Fnneran et al. (2002b)
results for TTS-onset n a beluga exposed to a
sngle pulse. The unweghted peak sound pressure
values of 224 dB re: 1 Pa (peak) and weghted
(Mmf) SEL values of 183 dB re: 1 Pa
2
-s are rec-
ommended as behavoral dsturbance crtera
for md-frequency cetaceans (Table 5, Cell 4). By
extrapolaton (see cetacean procedure, Chapter
3, p. 439), the same values were also proposed
for low- and hgh-frequency cetaceans (Table 5,
Cells 1 & 7, respectvely). The only dfference n
the applcaton of these crtera to the three ceta-
cean groups s the nfluence of the respectve fre-
quency-weghtng functons for SEL crtera (Mlf
and Mhf vs Mmf).
For pnnpeds exposed to sngle pulses n water,
the proposed behavoral dsturbance crtera are
also the estmated TTS-onset values. For pnn-
peds as a whole, these are 212 dB re: 1 Pa (peak)
and weghted (Mpw) SEL of 171 dB re: 1 Pa
2
-s
(Table 5, Cell 10) as dscussed n Chapter 3.
For pnnpeds n ar, the proposed behavoral
crtera are based on the strong responses (stam-
pedng behavor that could njure some nd-
vduals or separate mothers from pups) of some
speces, especally harbor seals, to sonc booms
from arcraft and mssle launches n certan
condtons (Berg et al., 2001, 2002; Holst et al.,
2005a, 2005b). No responses resultng n njury
were observed n these specfc studes, but the
behavoral responses were, n some cases, among
those that would be consdered relatvely severe
n regards to vtal rates. It was therefore deter-
mned approprate to use results from these stud-
es rather than TTS-based thresholds for behav-
oral response crtera. The proposed crtera are
109 dB re: 20 Pa (peak) and frequency-weghted
(Mpa) SEL of 100 dB re: (20 Pa)
2
-s (Table 5,
Cell 13). These levels are substantally below
TTS-onset values. They are also probably qute
precautonary as behavoral response crtera for
the group as a whole, especally for speces other
than harbor seals where hgher exposures were not
observed to nduce strong (or n some cases any)
responses.
452 Southalletal.
Behavioral Response Severity Scaling:
Multiple Pulses
Low-FrequencyCetaceans/MultiplePulses(Cell2)
Numerous feld observatons have been made
of low-frequency cetaceans reactng to multple
pulses ether ncdentally durng ongong opera-
tons or ntentonally durng experments. A mod-
erate number of speces and expermental cond-
tons have been consdered, but the sources have
usually been sesmc argun arrays. Some of the
studes focused on mgratng whales seen from
fxed observaton platforms or n/near mgratory
corrdors. Ths approach mnmzes pseudorepl-
caton wthout the need for dentfyng ndvduals
because ndvduals are unlkely to pass observers
more than once.
Table 6 summarzes the methods used to obtan
acoustc measurements and observatons of behav-
oral or dstrbutonal responses (see Appendx B
for more detals). As n most cells, a number of
reported observatons were not scored or reported
here due to lack of some key nformaton and, n
some cases, dffcultes n accountng for varous
contextual varables. A few of these excluded
studes are lsted at the bottom of Table 6. Table
7 shows the results of the severty scalng analy-
ses of ndvdual and/or group responses, con-
sderng the studes deemed to contan suffcent
data on exposure condtons and behavoral
responses. For mgratng bowhead whales, the
onset of sgnfcant behavoral dsturbance from
multple pulses occurred at RLs (RMS over pulse
duraton) around 120 dB re: 1 Pa (Rchardson
et al., 1999). For all other low-frequency cetaceans
(ncludng bowhead whales not engaged n mgra-
ton), ths onset was at RLs around 140 to 160 dB
re: 1 Pa (Malme et al., 1983, 1984; Rchardson
et al., 1986; Ljungblad et al., 1988; Todd et al.,
1996; McCauley et al., 1998, 2000) or perhaps
hgher (Mller et al., 2005). There s essentally no
overlap n the RLs assocated wth onset of behav-
oral responses by members of these two groups
based on the nformaton currently avalable.
Mid-FrequencyCetaceans/MultiplePulses(Cell5)
A lmted number of behavoral observatons have
been made of md-frequency cetaceans exposed to
Table 5. Proposed behavoral response crtera for ndvdual marne mammals exposed to varous sound types; specfc
threshold levels are proposed for sngle pulses. See the referenced text sectons and tables for severty scale analyses of
behavoral responses to multple pulses and nonpulses.
Sound type
Marne mammal group Sngle pulses Multple pulses Nonpulses
Low-frequency cetaceans Cell 1 Cell 2
1
Cell 3
6
Sound pressure level 224 dB re: 1 Pa (peak) (flat) Tables 6 & 7 Tables 14 & 15
Sound exposure level 183 dB re: 1 Pa
2
-s (Mlf) Not applcable Not applcable
Md-frequency cetaceans Cell 4 Cell 5
2
Cell 6
7
Sound pressure level 224 dB re: 1 Pa (peak) (flat) Tables 8 & 9 Tables 16 & 17
Sound exposure level 183 dB re: 1 Pa
2
-s (Mmf) Not applcable Not applcable
Hgh-frequency cetaceans Cell 7 Cell 8
3
Cell 9
8
Sound pressure level 224 dB re: 1 Pa (peak) (flat) [Tables 18 & 19] Tables 18 & 19
Sound exposure level 183 dB re: 1 Pa
2
-s (Mhf) Not applcable Not applcable
Pnnpeds (n water) Cell 10 Cell 11
4
Cell 12
9
Sound pressure level 212 dB re: 1 Pa (peak) (flat) Tables 10 & 11 Tables 20 & 21
Sound exposure level 171 dB re: 1 Pa
2
-s (Mpw) Not applcable Not applcable
Pnnpeds (n ar) Cell 13 Cell 14
5
Cell 15
10
Sound pressure level 109 dB re: 20 Pa (peak) (flat) Tables 12 & 13 Tables 22 & 23
Sound exposure level 100 dB re: (20 Pa)
2
-s (Mpa) Not applcable Not applcable
1
Low-Frequency Cetaceans/Multple Pulses (Cell 2) secton
2
Md-Frequency Cetaceans/Multple Pulses (Cell 5) secton
3
Hgh-Frequency Cetaceans/Multple Pulses (Cell 8) secton
4
Pnnpeds n Water/Multple Pulses (Cell 11) secton
5
Pnnpeds n Ar/Multple Pulses (Cell 14) secton
6
Low-Frequency Cetaceans/Nonpulses (Cell 3) secton
7
Md-Frequency Cetaceans/Nonpulses (Cell 6) secton
8
Hgh-Frequency Cetaceans/Nonpulses (Cell 9) secton
9
Pnnpeds n Water/Nonpulses (Cell 12) secton
10
Pnnpeds n Ar/Nonpulses (Cell 15) secton
MarineMammalNoiseExposureCriteria 453
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454 Southalletal.
multple pulses. Feld observatons have nvolved
sperm whales and a few other odontocete spe-
ces exposed to sesmc arguns and explosves.
Laboratory nvestgatons have consdered behav-
oral responses to varous knds of multple pulse
sources. Agan, some observatons were excluded
due to lack of relevant nformaton. Four studes
of ndvdual md-frequency cetacean responses
to multple pulse exposures contaned suffcent
acoustc and behavoral nformaton for ncluson
n ths analyss. These nclude feld observatons of
free-rangng sperm whales and belugas studed by
Madsen & Mhl (2000), Madsen et al. (2002), and
Mller et al. (2005), as well as laboratory observa-
tons of captve false kller whales by Akamatsu
et al. (1993). The nformaton from these studes
s summarzed n Table 8 and dscussed n detal
n Appendx B; the companon severty scalng
analyss s shown n Table 9.
The combned data for md-frequency ceta-
ceans exposed to multple pulses do not ndcate
a clear tendency for ncreasng probablty and
severty of response wth ncreasng RL. In cer-
tan condtons, multple pulses at relatvely low
RLs (~80 to 90 dB re: 1 Pa) temporarly slence
ndvdual vocal behavor for one speces (sperm
whales). In other cases wth slghtly dfferent
stmul, RLs n the 120 to 180 dB re: 1 Pa range
faled to elct observable reacton from a sgnf-
cant percentage of ndvduals ether n the feld or
n the laboratory.
High-FrequencyCetaceans/MultiplePulses(Cell8)
Based on our source type dstncton (see Chapter
2), vrtually all sources of transent sound used n
quanttatve behavoral studes of hgh-frequency
cetaceansfor example, acoustc harassment
devces (AHDs) and acoustc deterrent devces
(ADDs)would be characterzed as nonpulse
sounds. Whle ndvdual elements produced by
some of these sources could be characterzed as
pulses, and sequences of them as multple pulses,
they are generally emtted n such rapd fashon
that some mammalan audtory systems lkely
perceve them as nonpulses. Further, some AHDs
and ADDs, and most other sources used n behav-
oral studes wth hgh-frequency cetaceans, lack
the characterstcs of pulses such as extremely fast
rse-tme, correspondngly broad frequency band-
wdth, and hgh kurtoss. Due to uncertanty over
the extent to whch some of these sgnals may be
perceved and the overarchng paucty of data, t
s not possble to present any data on behavoral
responses of hgh-frequency cetaceans as a func-
ton of receved levels of multple pulses. Avalable
data for nonpulse sounds are consdered below
(see the Hgh-Frequency Cetaceans/Nonpulses
[Cell 9] secton). We note the need for emprcal
behavoral research n these anmals usng sound
sources (such as argun or ple-drvng stmul)
unequvocally classfed as multple pulses (see
Chapter 5).
Table 7. Number (n bold) of low-frequency cetaceans (ndvduals and/or groups) reported as havng behavoral responses
to multple pulse nose; responses were categorzed nto 10-dB RL bns, ranked by severty of the behavoral response (see
Table 4 for severty scalng), and combned wth other observatons havng the same RL/severty score. A summary of the
ndvdual studes ncluded n ths table s gven n the Low-Frequency Cetaceans/Multple Pulses (Cell 2) secton of ths
chapter. Parenthetcal subscrpts ndcate the reference reportng the observatons as lsted n Table 6.
Receved RMS sound pressure level (dB re: 1 Pa)
Response
score
80 to
< 90
90 to
< 100
100 to
< 110
110 to
< 120
120 to
< 130
130 to
< 140
140 to
< 150
150 to
< 160
160 to
< 170
170 to
< 180
180 to
< 190
190 to
< 200
9
8
7 1.0
(6)
6 9.5
(7)
47.4
(7)
2.2
(7)
3.4
(4, 6, 8)
5.8
(1, 2, 3, 6)
4.5
(1, 2, 3, 4, 6)
8.3
(1, 2, 4, 8, 9)
5 1.0
(7)
1.0
(4)
1.0
(1, 2)
4
3 1.0
(1, 2)
1.0
(1, 2)
2
1 5.0
(7)
6.0
(7)
1.0
(7)
2.5
(1, 2, 3)
3.0
(5)
0 59.8
(7)
17.7
(7)
1.1
(7, 9)
0.1
(9)
0.6
(3, 9)
6.8
(1, 2, 3, 9)
6.3
(1, 2, 9)
MarineMammalNoiseExposureCriteria 455
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456 Southalletal.
PinnipedsinWater/MultiplePulses(Cell11)
Informaton on behavoral reactons of pnnpeds
n water to multple pulses nvolves exposures to
small explosves used n fsheres nteractons,
mpact ple drvng, and sesmc surveys. Several
studes lacked matched data on acoustc expo-
sures and behavoral responses by ndvduals. As
a result, the quanttatve nformaton on reactons
of pnnpeds n water to multple pulses s very
lmted (see Table 10). The severty scalng analy-
ss for ndvdual behavoral responses for Cell 11
s gven n Table 11.
Our general fndng s that, based on the lmted
data on pnnpeds n water exposed to multple
pulses, exposures n the ~150 to 180 dB re: 1 Pa
range (RMS values over the pulse duraton) gen-
erally have lmted potental to nduce avodance
behavor n pnnpeds. RLs exceedng 190 dB
re: 1 Pa are lkely to elct responses, at least n
some rnged seals (Harrs et al., 2001; Blackwell
et al., 2004b; Mller et al., 2005). Note that the
SEL assocated wth a sngle 190 dB re: 1 Pa
(RMS) pulse from an argun s typcally ca. 175
dB re: 1 Pa
2
-s. That exceeds the estmated TTS
threshold for the closely related harbor seal (171
dB re: 1 Pa
2
-s; see Chapter 3). Thus, n the case
of rnged seals exposed to sequences of argun
pulses from an approachng sesmc vessel, most
anmals may show lttle avodance unless the RL
s hgh enough for mld TTS to be lkely.
PinnipedsinAir/MultiplePulses(Cell14)
How multple pulses produced n ar affect pnn-
peds was among the least well-documented of the
condtons we consdered. Most of the avalable
data on responses to pulses were from sngle pulse
events (e.g., rocket launches) over populatons of
pnnpeds exposed to such sgnals repeatedly (e.g.,
Thorson et al., 1998, 1999, 2000a, 2000b; Berg
et al., 2001, 2002, 2004). These events do not occur
frequently enough for the exposures to be consd-
ered multple pulses, and many of them contaned
nonpulse as well as pulse exposures. They are
dscussed n some detal n Appendx B (as well
as n Appendx C when nonpulses are nvolved).
Appendx B also dscusses several other studes
potentally relevant to Cell 14 but ultmately not
used n ths analyss. Consequently, the quantta-
tve nformaton analyzed for reactons of pnn-
peds n ar exposed to multple pulses (see Tables
12 & 13) focused on the aeral data by Blackwell
et al. (2004b). These extremely lmted data sug-
gest very mnor, f any, observable behavoral
responses by pnnpeds exposed to arborne pulses
wth RLs 60 to 80 dB re: 20 Pa.
Behavioral Response Severity Scaling:
Nonpulses
Low-FrequencyCetaceans/Nonpulses(Cell3)
Whle there are clearly major areas of uncertanty
remanng, there has been relatvely extensve
behavoral observaton of low-frequency ceta-
ceans exposed to nonpulse sources. As summa-
rzed n Table 14 (and dscussed n greater detal
n Appendx C), these feld observatons nvolve
the majorty of low-frequency cetacean speces
exposed to a wde range of ndustral, actve sonar,
and tomographc research actve sources (Baker
et al., 1982; Malme et al., 1983, 1984, 1986;
Table 9. Number (n bold) of md-frequency cetaceans (ndvduals and/or groups) reported as havng behavoral responses
to multple pulse nose; responses were categorzed nto 10-dB RL bns, ranked by severty of the behavoral response (see
Table 4 for severty scalng), and combned wth other observatons havng the same RL/severty score. A summary of the
ndvdual studes ncluded n ths table s gven n the Md-Frequency Cetaceans/Multple Pulses (Cell 5) secton of ths
chapter. Parenthetcal subscrpts ndcate the reference reportng the observatons as lsted n Table 8.
Receved RMS sound pressure level (dB re: 1 Pa)
Response
score
80 to
< 90
90 to
< 100
100 to
< 110
110 to
< 120
120 to
< 130
130 to
< 140
140 to
< 150
150 to
< 160
160 to
< 170
170 to
< 180
180 to
< 190
190 to
< 200+
9
8
7
6 0.17
(3)
0.17
(3)
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(3)
1.3
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5
4
3
2
1
0 0.25
(3)
0.25
(3)
3.0
(2)
4.0
(2)
6.7
(1, 4)
MarineMammalNoiseExposureCriteria 457
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458 Southalletal.
Rchardson et al., 1990b; McCauley et al., 1996;
Basson et al., 2000; Croll et al., 2001; Palka &
Hammond, 2001; Nowacek et al., 2004).
The combned nformaton generally ndcates
no (or very lmted) responses at RLs 90 to 120
dB re: 1 Pa and an ncreasng probablty of
avodance and other behavoral effects n the 120
to 160 dB re: 1 Pa range (severty scalng: Table
15). However, these data also ndcated consd-
erable varablty n RLs assocated wth behav-
oral responses. Contextual varables (e.g., source
proxmty, novelty, operatonal features) appear to
have been at least as mportant as exposure level
n predctng response type and magntude.
Mid-FrequencyCetaceans/Nonpulses(Cell6)
A relatvely large number of md-frequency cetaceans
have been observed n the feld and n the laboratory
respondng to nonpulse sounds, ncludng vessels
and watercraft (LGL & Greenerdge, 1986; Gordon
et al., 1992; Palka & Hammond, 2001; Buckstaff,
2004; Morsaka et al., 2005), pulsed pngers and
AHDs/ADDs (Watkns & Schevll, 1975; Morton &
Symonds, 2002; Montero-Neto et al., 2004), ndus-
tral actvtes (Awbrey & Stewart, 1983; Rchardson
et al., 1990b), md-frequency actve sonar (NRL,
2004a, 2004b; NMFS, 2005), and tones or bands
of nose n laboratory condtons (Nachtgall et al.,
2003; Fnneran & Schlundt, 2004). Summary nfor-
maton on these studes s gven n Table 16; detaled
descrptons are gven n Appendx C. As n other
condtons, a number of potentally relevant feld
studes are not ncluded n the severty scalng anal-
yss due to lack of suffcently detaled nformaton.
An addtonal challenge n nterpretng many
of the feld data for ths condton s solatng
the effect of RL from the effects of mere source
presence (as possbly ndcated by vsual stmul
or other aspects of acoustc exposure such as the
presence of hgh-frequency components) and
other contextual varables. For ths reason, several
studes were consdered but not ntegrated nto the
analyss. The laboratory observatons are of cap-
tve cetaceans exposed to precsely controlled and
known nose exposures n the context of hearng
and TTS experments. However, the relevance of
behavoral reactons of traned, food-renforced
captve anmals exposed to nose to the reactons
of free-rangng marne mammals s debatable.
Ths s dscussed n greater detal n Appendx C.
The combned feld and laboratory data for md-
frequency cetaceans exposed to nonpulse sounds
do not lead to a clear concluson about RLs conc-
dent wth varous behavoral responses (see sever-
ty scalng, Table 17). In some settngs, ndvduals
n the feld showed behavoral responses wth hgh
severty scores to exposures from 90 to 120 dB re:
1 Pa, whle others faled to exhbt such responses
for exposure RLs from 120 to 150 dB re: 1 Pa.
Contextual varables other than exposure RL, and
probable speces dfferences, are the lkely rea-
sons for ths varablty n response. Context may
also explan why there s great dsparty n results
from feld and laboratory condtonsexposures
n captve settngs generally exceeded 170 dB re:
1 Pa before nducng behavoral responses.
Table 11. Number (n bold) of pnnpeds n water (ndvduals and/or groups) reported as havng behavoral responses to
multple pulse nose. Responses were categorzed nto 10-dB RL bns, ranked by severty of the behavoral response (see
Table 4 for severty scalng), and combned wth other observatons havng the same RL/severty score; a summary of the
ndvdual studes ncluded n ths table s gven n the Pnnpeds n Water/Multple Pulses (Cell 11) secton of ths chapter.
Parenthetcal subscrpts ndcate the reference reportng the observatons as lsted n Table 10.
Receved RMS sound pressure level (dB re: 1 Pa)
Response
score
80 to
< 90
90 to
< 100
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< 130
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9.9
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460 Southalletal.
High-FrequencyCetaceans/Nonpulses(Cell9)
Numerous controlled studes have been conducted
on the behavoral reactons of hgh-frequency
cetaceans to varous nonpulse sound sources
both n the feld (Culk et al., 2001; Olesuk
et al., 2002; Johnston, 2002) and n labora-
tory settngs (Kastelen et al., 1997, 2000, 2005,
2006a). However, only one hgh-frequency spe-
ces (harbor porpose) has been extensvely
studed and that speces provded all the aval-
able data on behavoral response magntude vs
receved exposure condtons. The orgnal stud-
es were attempts to reduce harbor porpose by-
catch by attachng warnng pngers to fshng gear.
More recent studes consder whether AHDs and
ADDs also exclude harbor porposes from crt-
cal habtat areas, and whether these devces affect
harbor porpose behavor n controlled laboratory
condtons.
The combned wld and captve anmal data
(summarzed n Table 18 and dscussed n detal n
Appendx C) clearly support the observaton that
harbor porposes are qute senstve to a wde range
of human sounds at very low exposure RLs (~90 to
120 dB re: 1 Pa), at least for ntal exposures. Ths
observaton s also evdent n the severty scalng
analyss for Cell 9 (Table 19). All recorded expo-
sures exceedng 140 dB re: 1 Pa nduced profound
and sustaned avodance behavor n wld harbor
porposes. Whether ths apparently hgh degree of
behavoral senstvty to anthropogenc acoustc
sources extends to other hgh-frequency cetacean
speces (or nonpulse sources other than AHDs and
ADDs) s unknown. Gven the lack of nforma-
ton to the contrary, however, such a relatonshp
should be assumed as a precautonary measure.
Habtuaton to sound exposure was noted n some
but not all studes. Strong ntal reactons of hgh-
frequency cetaceans at relatvely low levels may n
some condtons wane wth repeated exposure and
subject experence.
PinnipedsinWater/Nonpulses(Cell12)
The effects of nonpulse exposures on pnn-
peds n water are poorly understood. Studes
for whch enough nformaton was avalable for
analyss nclude feld exposures of harbor seals
to AHDs (Jacobs & Terhune, 2002) and exposure
of translocated freely dvng northern elephant
seals to a research tomography source (Costa
et al., 2003), as well as responses of captve harbor
seals to underwater data communcaton sources
(Kastelen et al., 2006b). These lmted avalable
data (see Table 20 & Appendx C) suggested that
exposures between ~90 and 140 dB re: 1 Pa
generally do not appear to nduce strong behav-
oral responses n pnnpeds exposed to nonpulse
sounds n water; no data exst regardng exposures
at hgher levels. The severty scale results for Cell
12 are gven n Table 21.
It s mportant to note that among these stud-
es of pnnpeds respondng to nonpulse exposures
n water, there are some apparent dfferences n
responses between feld and laboratory cond-
tons. Specfcally, n ths case, captve subjects
responded more strongly at lower levels than dd
anmals n the feld. Agan, contextual ssues are
the lkely cause of ths dfference. Captve sub-
jects n the Kastelen et al. (2006b) study were not
renforced wth food for remanng n nose felds,
n contrast to the laboratory studes for md-fre-
quency cetaceans descrbed above. Subjects n the
Table 13. Number (n bold) of pnnpeds n ar (ndvduals and/or groups) reported as havng behavoral responses to
multple pulse nose; responses were categorzed nto 10-dB RL bns, ranked by severty of the behavoral response (see
Table 4 for severty scalng), and combned wth other observatons havng the same RL/severty score. A summary of the
ndvdual studes ncluded n ths table s gven n the Pnnpeds n Ar/Multple Pulses (Cell 14) secton of ths chapter.
Parenthetcal subscrpts ndcate the reference reportng the observatons as lsted n Table 12.
Receved RMS sound pressure level (dB re: 20 Pa)
Response score 50 to < 60 60 to < 70 70 to < 80 80 to < 90 90 to < 100 100 to < 110 110 to < 120
9
8
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MarineMammalNoiseExposureCriteria 461
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MarineMammalNoiseExposureCriteria 463
feld may have been more tolerant of exposures
because of motvaton to return to a safe locaton
(Costa et al., 2003) or motvaton to approach
enclosures holdng prey tems (Jacobs & Terhune,
2002).
PinnipedsinAir/Nonpulses(Cell15)
There has been consderable effort to study the
effects of aeral nonpulse sounds on pnnped
behavor, prmarly nvolvng rocket launches,
arcraft overflghts, powerboat approaches, and
constructon nose. Unfortunately, as dscussed n
Appendx C, many of the studes are dffcult to
nterpret n terms of exposure RL and ndvdual
or group behavoral responses. In many cases,
t was dffcult or mpossble to dscern whether
the reported behavoral response was nduced by
the nose from a specfc operaton or some cor-
related varable such as ts vsual presence. For
these reasons, most of the observatonal studes
of behavoral dsturbance were not approprate for
scorng behavoral responses relatve to exposure
RL. However, a number of the techncal reports
and analyses of rocket launches are relevant for
ths cell and contan suffcently detaled nfor-
maton regardng estmated RLs. These observa-
tons are, however, complcated by the fact that
all studes were conducted n the same general
area wth subjects lkely habtuated to the pres-
ence of launch nose. Further, n many cases,
exposures contaned both a nonpulse component
and a pulse component (descrbed below). Only
those observatons (Thorson et al., 1999, 2000b;
Berg et al., 2002) for whch there was clearly just
nonpulse exposure were consdered n the severty
scalng analyses for ths condton.
The lmtatons of these and other potentally
applcable studes resulted n a very lmted data
set for use n ths analyss (see summary n Table
22 and severty scalng analyss n Table 23). As a
general statement from the avalable nformaton,
pnnpeds exposed to ntense (~110 to 120 dB re:
20 Pa) nonpulse sounds tended to leave haulout
areas and seek refuge temporarly (mnutes to a few
hours) n the water, whereas pnnpeds exposed to
dstant launches at RLs ~60 to 70 dB re: 20 Pa
tended to gnore the nose. It s dffcult to assess
the relevance of ether of these observatons to
nave ndvduals, however, gven the repeated
exposure of study colones to such nose events and
the potental that observed ndvduals were habtu-
ated. Due to the lmtatons of avalable data, t s
not currently possble to make any further general
characterzatons regardng ths condton.
Table 15. Number (n bold) of low-frequency cetaceans (ndvduals and/or groups) reported as havng behavoral responses
to nonpulses; responses were categorzed nto 10-dB RL bns, ranked by severty of the behavoral response (see Table 4
for severty scalng), and combned wth other observatons havng the same RL/severty score. A summary of the nd-
vdual studes ncluded n ths table s gven n the Low-Frequency Cetaceans/Nonpulses (Cell 3) secton of ths chapter.
Parenthetcal subscrpts ndcate the reference reportng the observatons as lsted n Table 14.
Receved RMS sound pressure level (dB re: 1 Pa)
Response
score
80 to
< 90
90 to
< 100
100 to
< 110
110 to
< 120
120 to
< 130
130 to
< 140
140 to
< 150
150 to
< 160
160 to
< 170
170 to
< 180
180 to
< 190
190 to
< 200
9
8
7 2.5
(10)
1.5
(10)
6 4.9
(2)
7.4
(1, 2, 4)
16.2
(1, 2, 3, 5)
13.6
(2, 5)
4.2
(1, 2)
0.8
(2)
5
4 3.0
(5, 7)
1.0
(7)
1.0
(7)
3 1,117
(9)
0.27
(6)
2 0.5
(7)
4.0

(7)
5.0
(7)
2.0
(7)
1.0
(7)
1
0 1.1
(2)
82.6
(2, 3, 4)
33.9
(1, 2, 3, 4)
7.08
(2, 4, 6, 10)
7.2
(4, 10)
1.45
(2, 8, 10)
464 Southalletal.
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d


d
o
l
p
h

n
s
V
e
s
s
e
l

n
o

s
e

a
n
d

p
r
e
s
e
n
c
e
R
L

e
s
t

m
a
t
e
s

b
a
s
e
d

o
n

s
o
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r
c
e

a
n
d

e
n
v

r
o
n
m
e
n
t
a
l

c
h
a
r
a
c
t
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s
t

c
s
V

s
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a
l

o
b
s
e
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v
a
t

o
n
s

o
f

n
d

d
u
a
l

a
n
d

g
r
o
u
p

m
o
v
e
m
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n
t
s

a
n
d

b
e
h
a
v

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r
a
l

p
a
t
t
e
r
n
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d
u
r

n
g

v
e
s
s
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l

a
p
p
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o
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c
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o
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R
L
s

1
1
0
-
1
2
0

d
B

S
P
L
;

s
e
v
e
r
-

t
y

s
c
o
r
e
:

3
M
o
r
t
o
n

&

S
y
m
o
n
d
s

(
2
0
0
2
)
7
K

l
l
e
r

w
h
a
l
e
s
V
a
r

o
u
s

A
H
D
s
R
L

e
s
t

m
a
t
e
s

b
a
s
e
d

o
n

s
o
u
r
c
e

a
n
d

e
n
v

r
o
n
m
e
n
t
a
l

c
h
a
r
a
c
t
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r

s
t

c
s
C
e
n
s
u
s

d
a
t
a

f
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r

n
d

d
u
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l

a
n
d

g
r
o
u
p

s

g
h
t

n
g
s

u
s
e
d

t
o

e
s
t

m
a
t
e

e
x
c
l
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s

o
n

z
o
n
e
s
Y
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E
x
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o
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R
L
s

1
4
0
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5
0

d
B

S
P
L
;

s
e
v
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r
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y

s
c
o
r
e
:

8
B
u
c
k
s
t
a
f
f

(
2
0
0
4
)
8
B
o
t
t
l
e
n
o
s
e

d
o
l
p
h

n
s
V
e
s
s
e
l

n
o

s
e

a
n
d

p
r
e
s
e
n
c
e

(
a
p
p
r
o
a
c
h
e
s
)
C
a
l

b
r
a
t
e
d

R
L

m
e
a
s
u
r
e
m
e
n
t
s

m
a
d
e

i
n

s
i
t
u

n
e
a
r

a
r
e
a
s

o
f


e
x
p
o
s
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a
s
s

v
e

a
c
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t

c

m
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t
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f

n
d

d
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l

v
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c
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d
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v
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s
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l

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p
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p
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1
1
0
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1
2
0

d
B

S
P
L
;

s
e
v
e
r
-

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y

s
c
o
r
e
:

2
N
R
L

(
2
0
0
4
a
,

2
0
0
4
b
)
;

N
M
F
S

(
2
0
0
5
)
9
K

l
l
e
r

w
h
a
l
e
s
M

d
-

f
r
e
q
u
e
n
c
y

a
c
t

v
e

m

-
t
a
r
y

s
o
n
a
r
S
o
m
e

c
a
l

b
r
a
t
e
d

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L

m
e
a
s
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r
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m
e
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t
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a
n
d

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L

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t

m
a
t
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s

f
r
o
m

m
o
d
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l

n
g

s
o
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e

a
n
d


e
n
v

r
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n
m
e
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t
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c
h
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s
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s
V

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b
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d
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l

a
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d

g
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m
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m
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a
n
d

b
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h
a
v

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l

p
a
t
t
e
r
n
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b
e
f
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r
e
,

d
u
r

n
g
,

a
n
d

a
f
t
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r

n
c

d
e
n
t
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e
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7
0

d
B

S
P
L
;

s
e
v
e
r
-

t
y

s
c
o
r
e
:

6
MarineMammalNoiseExposureCriteria 465
S
t
u
d
y
R
e
f
e
r
e
n
c
e

n
u
m
b
e
r

(
f
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r

T
a
b
l
e

1
7
)
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u
b
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t

s
p
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d

s
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f

a
c
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c


m
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a
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m
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t
s
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y
p
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f

n
d

d
u
a
l

a
n
d
/
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r


g
r
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c
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c
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a
n
a
l
y
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s

(
s
e
e

T
a
b
l
e

1
7
)
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o
n
t
e

r
o
-
N
e
t
o

e
t

a
l
.

(
2
0
0
4
)
1
0
T
u
c
u
x

(
r

v
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d
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l
p
h

n
s
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k
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n
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t
m
a
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D
D
s
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m
a
t
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b
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m
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c
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g
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m
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d
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x
p
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a
n
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c
o
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t
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l

t
r

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l
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e
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0

d
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s
e
v
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y

s
c
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e
:

6
M
o
r

s
a
k
a


e
t

a
l
.

(
2
0
0
5
)
1
1
I
n
d
o
-
P
a
c

c

d
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l
p
h

n
s
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e
s
s
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l

n
o

s
e

a
n
d

p
r
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s
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c
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C
a
l

b
r
a
t
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d

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L

m
e
a
s
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m
e
n
t
s

m
a
d
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n

s
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u

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r

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a
s

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f


e
x
p
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a
c
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c

m
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t
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g

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f

n
d

d
u
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l


v
o
c
a
l

o
u
t
p
u
t

d
u
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g

v
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s
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l

a
p
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s

1
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0
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3
0

d
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S
P
L
;

s
e
v
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r
-

t
y

s
c
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r
e
:

5
N
a
c
h
t

g
a
l
l


e
t

a
l
.

(
2
0
0
3
)
1
2
B
o
t
t
l
e
n
o
s
e

d
o
l
p
h

n
s


(
c
a
p
t

v
e
)
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o
n
p
u
l
s
e

n
o

s
e

(
b
a
n
d
s
)
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a
l

b
r
a
t
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d

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L

m
e
a
s
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m
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n
t
s

i
n

s
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t
u

w

t
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n

t
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t

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n
c
l
o
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s
u
a
l

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b
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t

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b
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h
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v

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p
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w

t
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p
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m
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l
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c
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t
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t

Y
e
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E
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p
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1
7
0
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1
8
0

d
B

S
P
L
;

s
e
v
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r
-

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y

s
c
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r
e
:

6
F

n
n
e
r
a
n

&

S
c
h
l
u
n
d
t

(
2
0
0
4
)
1
3
B
o
t
t
l
e
n
o
s
e

d
o
l
p
h

n
s


(
c
a
p
t

v
e
)
N
o
n
p
u
l
s
e

n
o

s
e

(
t
o
n
e
s
)
C
a
l

b
r
a
t
e
d

R
L

m
e
a
s
u
r
e
m
e
n
t
s

m
a
d
e

i
n

s
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t
u

w

t
h

n

t
e
s
t

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n
c
l
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s
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V

s
u
a
l

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b
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v
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t

o
n
s

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f

b
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h
a
v

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l

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s
p
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n
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w

t
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n

e
x
p
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m
e
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t
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l

c
o
n
t
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x
t

n

l
a
b
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t
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y

c
o
n
d

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n
s
Y
e
s
E
x
p
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s
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R
L
s

1
8
0
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2
0
0

d
B

S
P
L
;

s
e
v
e
r
-

t
y

s
c
o
r
e
s
:

0

&

8
R
e
n
d
e
l
l

&

G
o
r
d
o
n

(
1
9
9
9
)
N
o
t

n
c
l
u
d
e
d
L
o
n
g
-
f

n
n
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d

p

l
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t

w
h
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s
A
c
t

v
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m

-
t
a
r
y

s
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n
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r
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n
s
u
f
f

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t

d
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t
a

f
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t
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s

a
n
a
l
y
s

s
P
a
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s

v
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a
c
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c

m
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m
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f

w
h

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t
l
e

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a
t
e
s
N
o
N
/
A
C
h

l
v
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r
s

&

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o
r
k
e
r
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n

(
2
0
0
1
)
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o
t

n
c
l
u
d
e
d
B
o
t
t
l
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e

d
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h

n
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n
d

p
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e
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n
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f
f

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n
t

d
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a

f
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s

a
n
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l
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V

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l

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b
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t

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d

d
u
a
l

f
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g

n
g

b
e
h
a
v

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r
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o
N
/
A
B
o
r
d

n
o

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t

a
l
.

(
2
0
0
2
)
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o
t

n
c
l
u
d
e
d
F
r
a
n
c

s
c
a
n
a

d
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l
p
h

n
s
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D
D
s
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n
s
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f
f

e
n
t

d
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f
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s

a
n
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l
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n
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f

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n
t

d
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N
/
A
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l

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m
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t

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l
.

(
2
0
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2
)
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t

n
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l
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d
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d
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d

p
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c
e
A
c
o
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s
t

c

m
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s
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m
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f

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e

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s

b
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m
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m
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d

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g

b
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N
/
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t

a
l
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(
2
0
0
3
)
N
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t

n
c
l
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d
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d
B
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t
l
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D
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d
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m
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m
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t

a
n
d

d

n
g

b
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h
a
v

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r
N
o
N
/
A
H
a
s
t

e

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t

a
l
.

(
2
0
0
3
)
N
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t

n
c
l
u
d
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d
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t
t
l
e
n
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d
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h

n
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e
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l

n
o

s
e

a
n
d

p
r
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n
c
e
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n
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466 Southalletal.
Table 17. Number (n bold) of md-frequency cetaceans (ndvduals and/or groups) reported as havng behavoral responses
to nonpulses; responses were categorzed nto 10-dB RL bns, ranked by severty of the behavoral response (see Table 4
for severty scalng), and combned wth other observatons havng the same RL/severty score. A summary of the nd-
vdual studes ncluded n ths table s gven n the Md-Frequency Cetaceans/Nonpulses (Cell 6) secton of ths chapter.
Parenthetcal subscrpts ndcate the reference reportng the observatons as lsted n Table 16.
Receved RMS sound pressure level (dB re: 1 Pa)
Response
score
80 to
< 90
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< 100
100 to
< 110
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120 to
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130 to
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140 to
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< 160
160 to
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180 to
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190 to
< 200
9
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134
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Courtesy: A. Fredlander
MarineMammalNoiseExposureCriteria 467
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MarineMammalNoiseExposureCriteria 469
Table 19. Number (n bold) of hgh-frequency cetaceans (ndvduals and/or groups) reported as havng behavoral responses
to nonpulses; responses were categorzed nto 10-dB RL bns, ranked by severty of the behavoral response (see Table 4
for severty scalng), and combned wth other observatons havng the same RL/severty score. A summary of the nd-
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Parenthetcal subscrpts ndcate the reference reportng the observatons as lsted n Table 18.
Receved RMS sound pressure level (dB re: 1 Pa)
Response
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MarineMammalNoiseExposureCriteria 471
Table 21. Number (n bold) of pnnpeds n water (ndvduals and/or groups) reported as havng behavoral responses to
nonpulses; responses were categorzed nto 10-dB RL bns, ranked by severty of the behavoral response (see Table 4 for
severty scalng), and combned wth other observatons havng the same RL/severty score. A summary of the ndvdual
studes ncluded n ths table s gven n the Pnnpeds n Water/Nonpulses (Cell 12) secton of ths chapter. Parenthetcal
subscrpts ndcate the reference reportng the observatons as lsted n Table 20.
Receved RMS sound pressure level (dB re: 1 Pa)
Response
score
80 to
< 90
90 to
< 100
100 to
< 110
110 to
< 120
120 to
< 130
130 to
< 140
140 to
< 150
150 to
< 160
160 to
< 170
170 to
< 180
180 to
< 190
190 to
< 200
9
8
7
6 1.0
(3)
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(2)
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(2)
2.0
(2)
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1
0 1.0
(3)
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(1, 2)
Courtesy: A. Fredlander
472 Southalletal.
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MarineMammalNoiseExposureCriteria 473
Table 23. Number (n bold) of pnnpeds n ar (ndvduals and/or groups) reported as havng behavoral responses to non-
pulses; responses were categorzed nto 10-dB RL bns, ranked by severty of the behavoral response (see Table 4 for sever-
ty scalng), and combned wth other observatons havng the same RL/severty score. A summary of the ndvdual studes
ncluded n ths table s gven n the Pnnpeds n Ar/Nonpulses (Cell 15) secton n ths chapter. Parenthetcal subscrpts
ndcate the reference reportng the observatons as lsted n Table 22.
Receved RMS sound pressure level (dB re: 20 Pa)
Response
score
50 to
< 60
60 to
< 70
70 to
< 80
80 to
< 90
90 to
< 100
100 to
< 110
110 to
< 120
9
8
7
6 1.0
(1, 2, 3)
5
4
3
2
1
0 1.0
(2)
Courtesy: Peter M. Schefele
5. Research Recommendations
The marne mammal nose exposure crtera
proposed here represent a synthess and precau-
tonary applcaton of current scentfc nforma-
ton. Clearly, the relance on extrapolaton proce-
dures, extreme data gaps and lmtatons n many
areas, and precautonary assumptons throughout
pont to the need for targeted research to fll spe-
cfc gaps n support of subsequent crtera. We
consder the current nose exposure crtera to be
merely an ntal step n an teratve process to
understand and better predct the effects of nose
on marne mammal hearng and behavor.
Research recommendatons are gven below
n several broad categores relevant to mprovng
marne mammal nose exposure crtera. No pr-
ortzaton s mpled n the orderng of these areas
or research topcs wthn them, however, and ths
s by no means an exhaustve lst. We present, n
abbrevated form, what we regard as crtcal, tar-
geted research needs to mprove future teratons
of these exposure crtera. Some of the most mpor-
tant research recommendatons are summarzed n
Table 24; each s dscussed n more detal n the
relevant secton below. Many of these research
recommendatons are smlar to recommendatons
made prevously (NRC, 1994, 2000, 2003, 2005;
Rchardson et al., 1995). Although there has been
progress n the last decade, much mportant work
remans to be done.
Measurements of Anthropogenic
Sound Sources and Ambient Noise
Comprehensve and systematc measurements
are needed of all relevant anthropogenc sound
sources that have a reasonable lkelhood of
adversely affectng marne mammal hearng or
behavor. Emprcal measures of sound felds
enable more accurate estmaton of RLs usng
propagaton models and valdate the selecton of
dfferent propagaton models as approprate. Such
studes must report the full range of relevant stan-
dard acoustc measurements and should nclude
detaled nformaton about equpment calbraton
and/or propagaton modelng methods used (e.g.,
Goold & Fsh, 1998; Wales & Hetmeyer, 2002;
Blackwell et al., 2004a). Measurements are also
needed to descrbe condtons where sounds clas-
sfed as pulses at the source transton to non-
pulse exposures. To measure in situ exposures
from specfc sound sources, archval acoustc
tags should be deployed on free-rangng marne
mammals and/or platforms near the anmals n
controlled exposure condtons.
If future nose exposure crtera are to consder
the mportant matters of audtory maskng, cumu-
latve exposure effects on ndvduals, and ecosys-
tem effects (dscussed below), addtonal data are
needed concernng ambent ocean nose on var-
ous spatal and temporal scales. These data should
be used to determne how ambent nose budgets
vary as a functon of natural and human actv-
tes. These data wll need to be ntegrated wth
expanded nformaton on marne mammal abun-
dance and dstrbuton. The NRC (2003) recom-
mended that a systematc effort be made to obtan
passve acoustc data, ncludng average (steady-
state) ambent nose from 1 Hz to 200 kHz, and
ncludng transent human sources not dentfed n
classcal ambent nose measurements. We concur
and call for wde-rangng acoustc measurements
desgned to test explct hypotheses about spatal
and temporal varablty n marne ambent nose.
Marine Mammal Auditory Processes
AbsoluteHearingData
Future teratons of these crtera wll be sg-
nfcantly mproved by ncreased knowledge of
hearng senstvty derved from behavoral and
electrophysologcal measurements and anatom-
cal models. The most pressng needs are for data
on deep-dvng cetaceans such as beaked whales
and on low-frequency specalsts (mystcetes).
Better nformaton on nter-speces dfferences
s also needed to valdate the functonal hearng
groups used here or alternatvely to dentfy other
relevant subdvsons (e.g., phocd vs otard pn-
npeds or potental parttonng of md-frequency
cetaceans). The number of ndvduals tested
should be ncreased n all speces, wth the pos-
sble excepton of the bottlenose dolphn, n order
to better understand ndvdual dfferences wthn
speces. Hearng senstvty across the full func-
tonal hearng range should be measured, where
possble, rather than just those frequences con-
taned wthn the communcaton sgnals of spe-
ces beng nvestgated.
Improvements are needed n both electro-
physologcal and behavoral testng methods to
ncrease the number of ndvduals of each spe-
ces that can be tested, and to dstngush absolute
AquaticMammals 2007, 33(4), 474-497, DOI 10.1578/AM.33.4.2007.474
MarineMammalNoiseExposureCriteria 475
Table 24. Research recommendatons n varous subject areas needed to enhance future marne mammal nose exposure
crtera (as dscussed n Chapter 5)
Research topc General descrpton Crtcal nformaton needs
Acoustic
measurementsof
relevantsound
sources
Detaled measurements needed of
source levels, frequency content, and
radated sound felds around ntense
and/or chronc nose sources.
Comprehensve, calbrated measurements of the propertes
of human-generated sound sources, ncludng frequency-
dependent propagaton and receved characterstcs n
dfferent envronments.
Ambientnoise
measurements
Systematc measurements of underwa-
ter ambent nose are needed to quan-
tfy how human actvtes are affectng
the acoustc envronment.
Comprehensve, calbrated measurements of amb-
ent nose, ncludng spectral, temporal, and drectonal
aspects, n dfferent oceanc envronments; ambent nose
budgets ndcatng relatve contrbuton of natural and
anthropogenc sources and trends over tme.
Absolutehearing
measurements
Audometrc data are needed to deter-
mne functonal bandwdth, speces and
ndvdual dfferences, dynamc hearng
ranges, and detecton thresholds for
realstc bologcal stmul.
Carefully controlled behavoral and electrophysologcal
measurements of hearng senstvty vs frequency for more
ndvduals and speces, partcularly for hgh-prorty spe-
ces, such as beaked whales and mystcetes. Also, detec-
ton thresholds for complex bologcal sgnals.
Auditoryscene
analysis
Measurements to determne the sophs-
tcated perceptual and processng capa-
bltes of marne mammals that enable
them to detect and localze sources n
complex, 3-D envronments.
Measurements of stream segregaton, spatal percepton,
multdmensonal source localzaton, frequency dscrm-
naton, temporal resoluton, and feedback mechansms
between sound producton and hearng systems.
Marinemammal
behavioralresponses
tosoundexposure
Measurements of behavoral reactons
to varous sound types are needed,
ncludng all relevant acoustc, contex-
tual, and response varables.
Carefully constructed observatonal and exposure exper-
ments that consder not only RL but also source range,
moton, sgnal-to-nose rato, and detaled nformaton on
recevers, ncludng baselne behavor, pror experence
wth the sound, and responses durng exposure.
Effectsofsound
exposureonmarine
mammalhearing:
masking,TTS,and
PTS
Contnued effort s needed on the
smultaneous and resdual physolog-
cal effects of nose exposure on marne
mammal hearng.
Masked hearng thresholds for smple stmul n more spe-
ces and ndvduals, as well as complex bologcal sgnals
and realstc maskers; allowance for drectonal effects;
comparatve data on TTS-onset and growth n a greater
number of speces and ndvduals for nonpulse and pulsed
anthropogenc sources; recovery functons after exposures
and between repeated exposures.
Effectsofsound
exposureonmarine
mammal
non-auditory
systems
Physologcal measurements are needed
for both acute and chronc sound expo-
sure condtons to nvestgate effects on
non-audtory systems.
Varous baselne and exposure-condton measurements,
ncludng ntrogen saturaton levels; bubble nucle; the
formaton of hemorrhages, embol, and/or lesons; stress
hormones; and cardovascular responses to acute and
chronc nose exposure.
Particularly
sensitivespecies:
beakedwhales
Baselne and exposure data on these
poorly understood taxa to assess ther
apparent senstvty to certan anthropo-
genc sound sources.
Varous studes, ncludng measurements and modelng
related to (1) hearng senstvty, (2) dvng and vocalza-
ton parameters, (3) tssue propertes, (4) gas/fat embol
formaton and sgnfcance, (5) advanced detecton capa-
bltes for localzng and trackng them, and (6) behav-
oral reactons to varous anthropogenc and natural sound
sources.
476 Southalletal.
from masked thresholds. Audtory evoked poten-
tal (AEP) technques should contnue to be
mproved and standardzed for pnnpeds and
small cetaceans. Researchers should contnue to
develop procedures applcable to stranded nd-
vduals of speces generally not represented n
captve settngs, partcularly for speces that may
be especally senstve to certan types of acous-
tc exposure. The massve body sze of mystcetes
may requre that AEP studes begn usng smaller
speces (e.g., mnke whale) that may be stranded,
trapped n tdal fshng enclosures (wers), or tem-
porarly avalable n a holdng faclty. Behavoral
audometrc methods, whch nvestgate the effect
of the overall perceptual and cogntve system on
detecton, should also contnue to be employed
and mproved, partcularly those that ncrease the
speed wth whch results are obtaned wthout sac-
rfcng precson of measurements.
Addtonally, behavoral methods should be
developed to measure hearng characterstcs
that requre a subjectve judgment of percepton
such as evaluaton of equal loudness between two
acoustc stmul. Equal-loudness hearng contours
for marne mammals are needed to refne the broad
frequency-weghtng networks derved here.
A fnal consderaton s that behavoral audo-
metrc research should eventually move beyond
the use of relatvely smple artfcal stmul (e.g.,
pure tones, nose bands, broadband clcks, tone
pps). Such stmul can be precsely controlled
and can be used to clearly ndcate whch acoustc
feature trggers the response n the whole anmal
or ts audtory system. Anmals n nature, how-
ever, rarely encounter such sounds. Whle some
bologcal sgnals consst of combnatons of tonal
elements, most are exceedngly complex. Marne
mammal detecton thresholds for complex, bo-
logcally relevant stmul may be poorly predcted
by experments usng smple artfcal stmul.
Humans, for example, are partcularly adept at
dentfyng speech-lke sounds n nose (Yost,
2000). Anmals are expected to be smlarly sens-
tve to mportant natural sounds. To base future
nose crtera on more relevant audometrc data,
research s needed on detecton thresholds for bo-
logcally meanngful sounds, such as vocalzatons
of conspecfcs, prey, and predators, and sounds
needed for actve or passve acoustc navgaton.
Such measurements wll further be useful n nves-
tgatng the potental actve space (detecton range
n three dmensons) for acoustc communcaton
(e.g., Brenowtz, 1982; Jank, 2000; Au et al.,
2004) and the effects of anthropogenc sound on
the actve space. Feld studes usng bologcally
relevant sounds would be more relevant to real-
world communcaton and maskng than studes
nvolvng smple, artfcal test stmul.
AuditorySceneAnalysis
Whle baselne hearng nformaton s clearly
needed, urgently n some cases, more advanced,
comprehensve, and nnovatve measurements are
also needed that provde nsght nto the ways n
whch anmals use ther audtory sense to derve
detaled nformaton about ther surroundng env-
ronment. For future teratons of nose exposure
crtera to consder multple stmul and cumula-
tve effects, addtonal data wll be needed on
sound localzaton n three-dmensonal audtory
space, frequency dscrmnaton, temporal resolu-
ton, and, specfcally, detecton of bologcal sg-
nals n complex sound felds.
Several studes of terrestral anmals
(MacDougall-Shackleton et al., 1998; Moss &
Surlykke, 2001) have nvestgated how subjects
process multple acoustc stmul that are smul-
taneously present but dffer n acoustc sgnature
ether temporally or spatally. The acoustc scene
concept, owng largely to the work of Bregman
(1990), has the potental to play a major role n
the development and progresson of acoustc expo-
sure crtera. Bregman draws powerful analoges
between modaltes of percepton, ncludng the
fundamental ways n whch hgher processng sys-
tems assocate common elements of complex stm-
ul n hghly cluttered perceptual envronments.
One analogy that Bregman (1990) makes wth
regard to the nnate power of vsual scene analyss
s the ablty of the vsual processng porton of the
human bran to estmate object sze wthout regard
to dstance. The mplcaton s that the reverse s
true as wellf the sze of somethng s known,
ts dstance can be nferred from vsual appear-
ance. Extendng ths ablty to anmals that rely on
underwater hearng to determne dstance, smlar
perceptual processes may occur. If so, mammals
may determne range by usng varous effects of
the propagaton medum on sound transmsson
(e.g., presence of structured mult-path sgnal
spreadng, frequency dependent mult-path losses,
and absorpton effects n partcular; Ellson &
Wexel, 1994). Further, both loudness modulaton
and source movement relatve to the recever pro-
vde sgnfcant clues as to the dstance and general
nature of the sound source. If one consders sound
to play a role n the lfe of marne wldlfe smlar
to that of sght n terrestral anmals, then context
clues such as tempo, encroachment, and proxmty
must take on a powerful role n determnng an an-
mals response to any gven sound. These hypoth-
eses need to be studed n marne mammals.
MarineMammalNoiseExposureCriteria 477
Behavioral Responses of
Marine Mammals to Sound
There s an urgent need for better and more exten-
sve data on behavoral responses to sound, nclud-
ng measurement of the specfc acoustc features
of exposures and consderaton of prevous expe-
rence wth the sound and all relevant contextual
varables. The current behavoral exposure crtera
are qute lmted n several ways. Insuffcent data
exst to support crtera other than those based on
SPL alone, and ths metrc fals to account for the
duraton of exposure beyond the separaton of
pulse from nonpulse sounds. Also, there s much
varablty n responses among speces of the same
functonal hearng group and also wthn speces.
Because of the poorly understood modfy-
ng nfluences of numerous varables, behavoral
responses usually cannot be predcted a priori
wth much confdence gven present nformaton.
In addton, the bologcal sgnfcance of any
observed behavoral response s even more dff-
cult to assess (NRC, 2005).
Research s needed to quantfy behavoral reac-
tons of a greater number of free-rangng marne
mammal speces to specfcally controlled or well-
characterzed exposures from dfferent human
sound sources. The most drect way to obtan these
knds of extremely detaled data would be to attach
acoustc dosmeter tags to ndvduals and drectly
measure nose exposure, behavoral response,
and physologcal changes, f any. It s essental
that future research nvestgates responses n con-
texts as smlar as possble to those of nterest.
Responses of both nave and prevously exposed
ndvduals should be studed and dstngushed to
the greatest extent possble.
Further, such experments must ensure that all
relevant acoustc measurements of sound exposure
be reported more systematcally than n many pre-
vous studes. Specfcally, behavoral responses
need to be drectly correlated wth the physcal
parameters (e.g., SPL, SEL) of the stmul most
lkely to evoke the responses. Such research
clearly requres greater knowledge of exposure
parameters (ncludng SPL over some duraton)
than currently exsts for most studes.
The relatonshp between exposure SPL and/or
SEL and behavoral reacton should be determned
for representatve speces wthn each functonal
hearng group. Whether the relatonshp follows
a dose-response-lke functon for varous sound
types, and under what condtons, s a sgnfcant
and pressng open queston.
We need more data on the magntude and tme
course of behavoral responses to known nose
exposures to test the valdty of concepts outlned
here, and to make progress toward dentfyng
specfc behavoral crtera. Duty cycle (the pro-
porton of tme when the nose s present) s also
lkely to be mportant. Magntude and duraton of
response are the most readly quantfed param-
eters that may be useful n determnng whether
a behavoral response s lkely to have a bolog-
cally meanngful outcome. Nose exposure crtera
should attempt to dstngush between mnor, tem-
porary behavoral changes and those wth greater
sgnfcance. Ths s necessary n order to focus
on bologcally sgnfcant behavoral responses
(see NRC, 2005) and the exposure condtons that
elct them.
Consderng the many contextual cues that free-
rangng anmals use to perceve and characterze
sound sources and to determne a response, t s
not surprsng that our analyss revealed a hgh
degree of varablty n behavoral responses as a
functon of RL. Consequently, the logc of relyng
solely on exposure RL as the metrc for behav-
oral responses s substantally dmnshed. A host
of varables addtonal to RL may be mportant
to marne mammals n assessng a sound and
determnng how to react. Ths argues for care-
ful desgn and executon of controlled exposure
experments to replcate the sgnal of nterest n
as many dmensons as possble. Serous con-
sderaton should be gven to developng a broad
mult-varable approach to behavoral research
that takes nto account not only source type and
exposure level but also dstance, moton, and rela-
tve sgnal-to-nose rato. Some studes are already
developng data of the scale and qualty needed
for such an approach. Ths ncludes studes pro-
vdng broad, long-term measurements of amb-
ent sounds n areas cohabted by anthropogenc
sources and marne wldlfe. Where these studes
nclude remotely deployed passve acoustc sen-
sors and tagged anmals, they approach what may
become the new standard. As addtonal nfor-
maton becomes avalable, future nose exposure
crtera may assess behavoral reactons not only
accordng to RL measured wth multple acoustc
parameters, range (near and far), relatve moton
(towards, parallel, etc.), and rate of change, but
also n relaton to the anmals actvty or per-
ceptual stuaton (e.g., neutral; threatened, as by
a predator; or postve, related to food, matng,
etc.).
The role of habtuaton and senstzaton n
behavoral reactons to nose exposure s a crt-
cal subject for future research. These processes
can only be studed under controlled or well-
defned condtons (as n Deecke et al., 2002). A
key queston s how habtuaton and senstzaton
develop wth repeated exposure n specfc eco-
logcally relevant crcumstances. For example,
the pattern of habtuaton to a neutral stmulus
478 Southalletal.
s lkely to follow qute a dfferent pattern from
selectve habtuaton to a harmless stmulus that
s ntally perceved as a threat (Deecke et al.,
2002). Furthermore, t would be desrable to know
f there are common acoustc features n sounds
to whch marne mammals become senstzed. For
example, to whch acoustc features of a threat,
such as a vessel used to hunt anmals, does an
anmal become senstzed?
Analyses of the behavor of varous anmal spe-
ces n the presence of predators suggest that they
have evolved ant-predator responses that mrror
ther responses to human dsturbance. Accordng
to predaton rsk theory, varous ecologcal con-
sderatons beyond smply dsturbance magntude
are very lkely nvolved n determnng and pre-
dctng behavoral response (Frd & Dll, 2002).
The bologcal relevance of behavoral changes
can only be determned n natural populatons n
whch vtal lfe hstory parameters (e.g., reproduc-
ton, growth, and survval rates) can be measured
before and after nose exposure and n condtons
where other potental stressors have been controlled
(NRC, 2005). One mportant queston s whether
these lfe hstory parameters are the same n popu-
latons that have apparently habtuated to expo-
sure and reman n relatvely nosy envronments
as they are n populatons lvng n queter cond-
tons. Because of the apparently major nfluence
of experence and the strong context-specfcty of
behavoral responses to nose, feld measurements
must be made for long perods followng repeated
or contnual exposure. Longtudnal studes should
be conducted to assess the tme course of expo-
sure to varous exstng sound sources known or
suspected to cause relatvely long-term (seasonal)
habtat abandonment. Where possble, parallel
studes should be done n neghborng areas wth
dfferent levels of nose exposure. Such studes
should allow for other non-acoustc factors lkely
to affect dstrbuton such as predators, prey, and
other mportant envronmental covarates. These
studes wll often need to extend over long per-
ods (many years) n order to be effectve, and they
should be planned and funded recognzng that.
Ideally, such a study should start collectng data
well n advance of the ntroducton of anthropo-
genc nose, and contnue throughout the perod of
antcpated mpact and for long enough thereafter
to observe return to baselne.
Effects of Noise Exposure on Marine Mammal
Hearing and Other Systems
AuditoryMasking
Audtory maskng s lkely the most wdespread
effect of anthropogenc nose on populatons of
marne mammals. The prncples of maskng are
reasonably well-known from laboratory studes n
mammals, ncludng marne mammals. To enable
maskng to be ncluded n subsequent nose expo-
sure crtera, however, data are needed on mask-
ng and ts effects n real-world condtons for
all functonal hearng groups. Data are needed
on the maskng effects of natural and anthropo-
genc nose sources; on detecton of smple, artf-
cal stmul; and, ncreasngly, on more complex,
bologcally meanngful sgnals. Drectonalty n
the maskng sound and/or the sgnal of nterest s
very lkely to affect the severty of maskng and
needs to be consdered. Baselne measurements
are needed on functonal communcaton ranges
for dfferent acoustc sgnals and on the reducton
of those ranges caused by ether natural or anthro-
pogenc maskers. Also needed are addtonal feld
measurements of the behavoral adjustments that
marne mammals make to offset maskng effects
(e.g., Lesage et al., 1999; Serrano & Terhune,
2002; Foote et al., 2004; Schefele et al., 2005).
TemporaryThresholdShift(TTS)
TTS studes n marne mammals reman lmted
to a very few speces and ndvduals, lmtng the
certanty wth whch they may be extrapolated
wthn and among groups. A number of specfc
TTS studes are needed to mprove future crte-
ra. For nstance, t s crtcal to future teratons
of these nose exposure crtera that research on
TTS-onset, TTS growth wth nose exposure, and
recovery rates expands to larger numbers of nd-
vduals and speces, and to speces n the low- and
hgh-frequency cetacean groups. Presently, extrap-
olaton procedures must be used because TTS
data are unavalable for certan functonal hearng
groups. Addtonally, certan hghly precautonary
procedures are used here n the estmaton of PTS-
onset because the growth rate of TTS wth ncreas-
ng exposure level s generally poorly understood,
even for the few marne mammal speces n whch
TTS has been studed. The relatonshp between
audtory senstvty and susceptblty to TTS/PTS
should be determned by group.
To the extent possble, electrophysologcal
technques should be used to obtan these TTS
data to ncrease sample sze and knowledge of
recovery functons.
More data for pnnpeds also are needed, par-
tcularly for pulse exposures where extrapolatons
of cetacean data currently must be used. Partcular
emphass should be placed on determnng
whether harbor seals have ncreased senstvty to
nose exposure relatve to other pnnped speces,
as current nformaton suggests, and f so, whether
speces closely related to the harbor seal also are
more senstve than are other pnnpeds.
MarineMammalNoiseExposureCriteria 479
To mnmze the need for such extrapolaton
and reduce the assumptons requred to predct
PTS-onset, emprcal data are needed on TTS
growth rates up to greater shft magntudes (10 to
30 dB). These data are needed for both pulse and
nonpulse sound types, at a varety of exposure fre-
quences, n both sngle and multple exposures.
These results should further elucdate whether,
and n what condtons, the equal energy hypoth-
ess may be approprate for comparng the effects
of varable nose exposures n marne mammals.
For pulse exposures, partcular attenton should
be pad to whether TTS growth s drectly related
to overall nose energy, and whether the kurtoss
of exposure s also a factor (see Erdrech, 1986;
Thery & Meyer-Bsch, 1988; Dunn et al., 1991;
Hamernk et al., 1993, 2003).
A further topc for future research s deter-
mnng whether usng 40 dB of TTS as a proxy
for PTS-onset s a precautonary approach, and
whether TTSs on the order of 25 to 35 dB are fully
recoverable n marne mammals as expected from
terrestral mammal data. To avod any possblty
of njury, such studes should contnue to take a
precautonary approach, usng gradual ncreases
n exposure level and duraton.
A related queston s how TTS recovers fol-
lowng nose cessaton n varable condtons.
Data on recovery functons and TTS magntude
are needed for representatve speces from each
functonal hearng group. Electrophysologcal
technques may be partcularly useful n ths
regard. These data may be useful n comparng
basc audtory system responses to nose exposure
and determnng how summaton procedures for
multple exposures should be modfed to more
precsely consder exposure ntermttence. Levels
of relatvely long duraton nose exposure causng
asymptotc TTS, n whch TTS values do not con-
tnue to ncrease n magntude wth exposure but
may have longer-lastng effects, should be deter-
mned. Recovery functons from asymptotc TTS
of varous levels should be compared wth recov-
ery functons from non-asymptotc TTS.
Fnally, the exstence of a stapedal reflex n
marne mammals and ts possble role n mtgat-
ng TTS and other effects of ntense nose expo-
sure are areas of needed research. For certan
nose exposures, partcularly those wth relatvely
low frequences and long duraton, the mddle ear
muscles (tensor tympan and stapedal) of terres-
tral mammals may contract and reduce the ampl-
fyng functon of the osscular chan (Yost, 2000).
Ths muscular contracton reduces the amount of
acoustc energy transmtted nto the cochlea va
the stapes. Ths stapedal reflex has been demon-
strated n humans exposed to ntense sound (Davs
et al., 1955) as well as echolocatng bats exposed
to ther own ntense outgong clcks (Henson,
1965). The mddle ears of marne mammals have
some specalzed adaptatons relatve to terrestral
mammals (see Wartzok & Ketten, 1999). In water,
f bone conducton (rather than osscular conduc-
ton) s the predomnant transmsson path, t s
possble that a stapedal reflex, f present, may
have lmted or no protectve functon for ntense
acoustc exposures. Research s also needed on the
role of meatal closure n pnnpeds durng nose
exposure. Such closures could be an alternatve
or addtonal way of reducng audtory senstvty.
Ether mechansm could also affect the nterpreta-
ton of threshold f performed durng audometrc
measurements.
PermanentThresholdShift(PTS)
Sound exposures causng PTS-onset, used here
to defne njury from acoustc exposure, have
not been measured n marne mammals. Instead,
exposures that would cause PTS-onset are est-
mated from measured TTS-onset usng assump-
tons about the growth of TTS wth nose expo-
sure level. Drect measurements of PTS n marne
mammals are hghly desrable for establshng
future njury crtera, but they are unlkely to be
obtaned due to ethcal, legal, and/or practcal
consderatons. Data from modelng and exposure
of cadavers to very ntense acoustc stmul gve
some ndcaton of condtons causng PTS but do
not reveal the exposure condtons that produce
PTSinvivo, nor actve processes that affect bas-
lar membrane dsplacement. Consequently, our
research recommendatons for mprovng PTS-
onset predctons for marne mammals nvolve
more ndrect measures.
One recommended type of ndrect measure s
to compare age-related hearng changes n captve
ndvduals that have been nvolved n TTS exper-
ments wth those that have not. Ths comparson
may provde some nsght nto the complex rela-
tonshp between repeated TTS and PTS, whch
remans poorly understood for terrestral mam-
mals, ncludng humans. One man mpedment,
however, s that confoundng varables lkely exst
other than controlled nose exposure. For captve
ndvduals used n TTS studes, absolute hear-
ng should be tested both durng and followng
sequences of nose exposure experments. For
captve ndvduals not used n TTS experments,
absolute hearng should be measured at regular
ntervals over extended perods. The latter group
may more readly dsplay natural age-related hear-
ng loss (presbycusis) than the former, as well as
potental sex dfferences. For both groups, efforts
should be made to characterze long-term ambent
nose condtons experenced by test anmals.
480 Southalletal.
Non-AuditoryEffectsofNoiseExposure
Lack of specfc data on acoustc exposures caus-
ng non-audtory effects n marne mammals cur-
rently prevents dervng explct exposure crtera
for such effects. Research s underway, however,
that may make ths possble n future versons
of the crtera. Non-audtory effects of nose are
potentally sgnfcant but reman generally poorly
understood.
A current hypothess regardng non-aud-
tory effects s that acoustc exposure may pro-
duce ntrogen bubbles n blood or other tssues.
Hemorrhages, gas and fat embol, and other
lesons have been reported n some marne mam-
mals exposed to md-frequency mltary sonar
(Jepson et al., 2003; Fernndez et al., 2004, 2005).
Substantal emprcal questons reman, however.
Frst among these s whether ntrogen bubbles are
n fact responsble for the hemorrhages, embol,
and other lesons reported. Conversely, are enough
ntrogen bubbles produced to pose a rsk of related
tssue njures, under any set of crcumstances,
arsng from hgh ntrogen supersaturaton levels,
acoustc exposure, and/or drastc changes n behav-
or? Do hgh levels of ntrogen supersaturaton or
gas or fat embol occur n dvng mammals that
have not been exposed to ntense anthropogenc
sound? Do these or related phenomena occur n
speces other than beaked whales? If bubble for-
maton s acoustcally medated, does t occur
as a drect result of acoustc exposure of bubble
precursors (nucle) n tssue, or ndrectly through
changes n dvng behavor? If the pathway s
drect, how does bubble formaton and/or growth
occur? A more thorough understandng s needed
of lpd bochemstry n tssues that may be par-
tcularly senstve to acoustcally medated bubble
formaton (e.g., acoustc fats). Modelng studes
are needed on tssue propertes and ther relevance
to ntrogen bubble formaton at specfc frequen-
ces of nterest. These studes should consder the
growth of dscrete bubbles from precursors n var-
ous tssues, and the nteracton among coalesced
aggregatons of acoustcally actvated bubbles.
If the pathway s ndrect and medated by behav-
or, s rapd surfacng more rsky than remanng
submerged too long and exceedng physologcal
lmts? How does the dve profle affect the lmts of
ntrogen supersaturaton n normal dvng? Do hgh
levels of ntrogen supersaturaton and gas embol
occur n marne mammals that have voluntary
control over depth, dvng profle, and nter-dve
nterval? Resoluton of these questons s lkely to
requre nterplay between modelng and emprcal
measurements (Zmmer & Tyack, 2007).
In conjuncton wth the above physologcal
modelng and measurements, controlled expo-
sure experments should be conducted wth
deep-dvng marne mammals to determne behav-
oral responses to sound sources, ncludng sonar.
These experments should use realstc source and
receved levels. If responses are dentfed, ths
may dentfy stuatons where t would be useful
to conduct observatonal studes of responses
durng uncontrolled use of anthropogenc sound
sources. Research should characterze the changes
n dvng behavor and should determne what they
mean n terms of bubble formaton or growth wth
contnued exposure.
Other possble non-audtory effects of acoustc
exposure should be nvestgated as well. Stress
hormone levels assocated wth nose exposure
should be more fully nvestgated. As of now,
they have been nvestgated followng exposure of
captve odontocetes to hgh-level sound (Thomas
et al., 1990c; Romano et al., 2004). The ablty of
anmals to recrut effectve stress responses should
also be studed durng chronc exposuresfor
example, n captve anmals that lve permanently
n nosy vs quet envronments. Effects of nose
exposure on marne mammal vestbular and car-
dovascular systems should also be studed.
Particularly Sensitive Species
In rare crcumstances, marne mammals (prmar-
ly beaked whales) have been known to strand
and ultmately de followng exposure to tactcal,
md-frequency actve sonar (see Cox et al., 2006;
Nowacek et al., 2007). Our knowledge of these
knds of extreme reactons to acute exposures
remans poor. However, the avalable nforma-
ton suggests that at least some speces of beaked
whales are partcularly senstve to ths one spe-
cfc category of sound sources.
Gas bubble formaton s a hypotheszed path-
way of ths effect (e.g., Fernndez et al., 2005),
but t remans poorly understood and the precse
mechansm underlyng these strandngs remans
unknown (e.g., Cox et al., 2006). The controlled
exposure experments outlned above are essental
to revealng the condtons and responses underly-
ng ths effect. Untl such research s conducted,
dervng scence-based exposure crtera specf-
cally for beaked whales or other deep-dvng ceta-
ceans exposed to actve sonar wll prove dffcult
or mpossble.
One current hypothess s that behavoral reac-
tons nfluence beaked whale dvng patterns n a
way that nduces physcally debltatng or dsor-
entng njures (Cox et al., 2006). Both the specf-
cs of ths potental mechansm and whether t s
specfc to beaked whales remans unknown, how-
ever. Mammals, ncludng some marne mammals,
show strong avodance responses when evadng
predators. Sounds from tactcal md-frequency
MarineMammalNoiseExposureCriteria 481
sonars somewhat resemble, n frequency band
and modulaton, the socal sgnals of one of the
only predators of large marne mammals, the
kller whale. If beaked whales nhert a broad tem-
plate for acoustc detecton of these predators, as
waterfowl do for vsual detecton of aeral preda-
tors (Lorenz, 1939; Tnbergen, 1948), they mght
respond to sonar as f t were a predator. Learnng
s requred for selectve habtuaton to safe stm-
ul that resemble those from predators (Deecke
et al., 2002). Many of the strandngs that concde
wth sonar exercses have occurred n stes where
kller whales are rare. Possbly these stranded
anmals have not had enough experence wth
ether sonar or kller whales to learn the dffer-
ence. Propagaton of sound n the ocean may also
degrade acoustc features that help dfferentate
the two classes of stmul at a dstance. It s plau-
sble that ths type of reacton could occur at rela-
tvely long dstances from the source f the sound
s alarmng based on propertes other than hgh
RL.
Whether beaked whales n certan condtons
mstake tactcal md-frequency sonar sgnals
for kller whales and consequently change ther
behavor n a way that njures them s an empr-
cal queston. Ths should be carefully nvestgated
usng controlled experments that take nto account
the relevant contextual varables dscussed above.
Addtonal baselne data on beaked whale phys-
ology, lfe hstory, and behavor are also needed
to approprately address questons regardng the
apparent senstvty of these anmals to certan
knds of anthropogenc sound. Fnally, n some
specfc condtons, such as sonar tranng ranges,
where sophstcated lstenng arrays make t pos-
sble to detect marne mammals over large ranges
before and durng actve sonar operatons, actve
or passve detecton of marne mammal behav-
oral patterns may become ncreasngly possble.
Whle these observatons have lmtatons, gven
that they may be able to detect more ndvduals
wthout requrng taggng efforts, they may be an
mportant complement to drected experments.
Some other speces of marne mammals are
unusually responsve to certan anthropogenc
sounds, ether generally or under partcular cond-
tons, and ths can result n strong and sometmes
large-scale avodance. Examples nclude harbor
porposes and, n some but not all stuatons,
beluga and bowhead whales (Fnley et al., 1990;
Rchardson et al., 1999; Olesuk et al., 2002;
Mller et al., 2005). There s a need for addtonal
behavoral and acoustc nformaton to better char-
acterze these extreme responses, the stuatons n
whch they occur, and whether smlar responses
can occur n other related speces or n response to
other smlar stmul.
Necessary Progressions of Marine Mammal
Noise Exposure Criteria
The currently proposed nose exposure crtera
are for ndvdual sound exposures and ndvdual
marne mammals. The research recommended
above s needed to substantate and mprove future
teratons of these types of crtera. Future tera-
tons of behavoral dsturbance crtera may derve
dose-response functons based on an ordnal scor-
ng paradgm smlar to that provded. Ths may
occur for subcategores of sound sources wthn
the general categores here (e.g., sesmc sgnals
as a subset of multple pulses, vessel nose as a
subset of nonpulses). It may also occur for sub-
groups of speces wthn the broad categores rec-
ognzed here (e.g., phocd vs otard pnnpeds) and
for other types of marne mammals not addressed
here (e.g., srenans, sea otters, polar bears).
Future teratons of these nose exposure crtera
should also perhaps dstngush several dfferent
categores of response that are expected, for both
theoretcal and emprcal reasons, to vary wth
RL n dfferent ways. For example, f an anmal
responds to a sound as f t were from a predator
(Frd & Dll, 2002), one would expect the dose-
response functon to have a very dfferent shape
as compared to that f the anmal responds based
on nterference wth the anmals ablty to com-
muncate acoustcally or echolocate. Predctng
whether a sound mght trgger an ant-predator
response would requre more detaled analyses of
acoustc parameters of the anthropogenc sound
compared to sgnals of predators. Further, n some
non-marne taxa, dfferent ant-predator responses
may be trggered dependng on levels and other
characterstcs of acoustc stmul (Spangler, 1988;
Hoy, 1989) and may be modulated by the cost of
the response as well as the perceved rsk (Frd
& Dll, 2002). Behavoral ecologsts hypothesze
that ant-predator behavor should balance rsk of
predaton aganst cost of response, ncludng cost
of foregone benefts from alternatve actvtes
(Frd & Dll, 2002). These non-acoustc param-
eters must be taken nto account n order to under-
stand dsturbance responses. The acoustc param-
eters affectng ant-predator behavor may nvolve
detecton thresholds, ambent nose condtons,
source dstance and source movement, as well
as the more drect measures of receved sound.
In future studes, most or all of these parameters
should be measured.
Addtonally, further exposure crtera are
needed to fully consder the effects of anthropo-
genc nose on other types of marne lfe, nclud-
ng the effects of sngle and multple exposures
on ndvdual nvertebrates, fsh, and sea turtles as
well as srenans, sea otters, and polar bears. There
482 Southalletal.
are fewer data to support crtera for marne bota
other than cetaceans and pnnpeds, and crtera
are perhaps as urgently (or more urgently) needed
for some other groups. Some fsh and most sea
turtle speces are consdered threatened or endan-
gered. The effects of anthropogenc nose on
fsh are also of partcular mportance gven ther
central role as both predators and prey n many
marne ecosystems and because of human depen-
dence on fsheres.
Addtonal crtera are also needed for the cumu-
latve effects of repettve or long-term nose expo-
sure on marne mammals. Ideally, spatotemporal
data on marne ambent nose and long-term expo-
sure hstores of ndvduals should be ntegrated
wth vtal rate data for marne mammal popula-
tons to address ths queston. Consderably more
data are needed on how nose mpacts n sngle
anmals can be extended to the populaton level.
Such measurements wll lkely requre extensve
measurements on a few representatve speces and
conservatve extrapolatons wthn and between
functonal hearng groups.
Nose exposure crtera that consder ecosystem-
level effects are needed as well. It s possble that
the effects of nose exposure on some elements of
local food webs may have a cascade effect to other
elements wthn the web. No data are avalable on
the ecologcal effects of underwater nose, even
at a local scale. However, gven the upward trend
n human actvtes n many nearshore areas, such
ecologcal effects should be antcpated.
Progress n each of these research areas wll
nvolve teratve processes that depend on the
avalablty of relevant scentfc data. Lke the
process of mprovng and expandng future nose
exposure crtera, our ablty to understand and
predct the effects of anthropogenc nose expo-
sure on marne ecosystems wll contnue to evolve
over a perod of many decades.
Acknowledgments
The authors acknowledge three former mem-
bers of the marne mammal nose exposure cr-
tera group: Whtlow Au, Sam Rdgway, and
Ronald Schusterman. Partal fnancal support
for meetngs of the nose exposure crtera group
was provded by the U.S. Natonal Oceanc and
Atmospherc Admnstraton, Natonal Marne
Fsheres Servce, Ocean Acoustcs Program.
The authors also apprecate the expertse,
revew, and assstance of the followng people:
Laure Allen, Robyn Anglss, Dan Costa, Tara
Cox, Ned Cyr, Adam Frankel, Ron Kastelen,
Steve Leathery, Rob McCauley, Davd Mellnger,
Rchard Merrck, Douglas Nowacek, Deborah
Palka, Mchael Payne, Arthur Popper, Andrew
Solow, Sofe Van Parjs, Wllem Verboom, Donna
Wetng, and fve anonymous revewers.
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Appendix A. Acoustic Measures and Terminology
Ths appendx provdes a more detaled descrp-
ton of many key acoustc measurements and
terms used throughout the nose exposure crte-
ra. It s not ntended as an exhaustve or nstruc-
tve text on these exceedngly complex ssues
(for more detaled treatments, see Knsler et al.,
1982; ANSI, 1986, 1994; Rchardson et al., 1995;
Harrs, 1998; NRC, 2003). Rather, t s ntended
to provde farly straghtforward defntons and
equatons related to the marne mammal nose
exposure crtera.
Pulses and Nonpulse Sounds
The dstncton between these two general sound
types s mportant because they have dfferng
potental to cause physcal effects, partcularly
wth regard to hearng (e.g., Ward, 1997).
Pulses, as used n the context of ths paper,
are defned as bref, broadband, atonal, tran-
sents (ANSI, 1986; Harrs, 1998, Chapter 12).
Examples of pulses (at least at the source) are
explosons, gunshots, sonc booms, sesmc argun
pulses, and ple drvng strkes. These sounds are
all characterzed by a relatvely rapd rse from
ambent pressure to a maxmal pressure value fol-
lowed by a decay perod that may nclude a perod
of dmnshng, oscllatng maxmal and mnmal
pressures. The rapd rse-tme characterstc of
these sounds ensures that they are also broad-
band n nature, wth the hgher-frequency com-
ponents beng related to the rapdty of the rse-
tme. Pulses, ether as solated events or repeated
n some successon, generally have an ncreased
capacty to nduce physcal njury as compared
wth sounds that lack these features.
Nonpulse (intermittent or continuous) sounds
can be tonal, broadband, or both. Some of these
nonpulse sounds can be transent sgnals of short
duraton but wthout the essental propertes of
pulses (e.g., rapd rse-tme). Examples of sources
producng nonpulse sounds nclude vessels; ar-
craft; machnery operatons, such as drllng or
wnd turbnes; and many actve sonar systems. The
duraton of such sounds, as receved at a dstance,
can be greatly extended n hghly reverberant env-
ronments. It s crtcal to note that a sound that has
characterstcs of a pulse at the source may, as a
result of propagaton effects, lose those charac-
terstcs at some (varable) dstance and could be
characterzed as a nonpulse for certan recevers.
Pulses and nonpulses are dstngushed here by
an emprcal approach based on several temporal
weghtngs. Varous exponental tme-weghtng
functons appled n measurng pulse and nonpulse
sounds may yeld dfferent measured receved
levels (RLs) (see Harrs, 1998). By way of llus-
traton, most sound level meters (SLM) provde
optons for applyng ether a slow or fast tme
constant (1,000 or 125 ms, respectvely) for mea-
surng nonpulses, or an mpulse tme constant (35
ms) approprate for measurng pulses. If appled
to a sound pulse, the slow or fast SLM settngs
result n lower sound pressure level (SPL) mea-
surements than those obtaned usng the mpulse
settng. Each of these tme constants was selected
based on the physcal propertes of the human
audtory system. It s clear that further empr-
cal measures of temporal resoluton n marne
mammals are needed, partcularly for anmal taxa
whose hearng extends to sgnfcantly hgher or
lower frequences than n humans (see Chapter 5,
Research Recommendatons). Future nose cr-
tera are expected to nclude dstnctons between
pulse and nonpulse sounds that may be more spe-
cfcally approprate for marne mammals than
s ths current smple approach. We note also the
need for an explct dstncton and measurement
standard, such as exsts for aeral sgnals (ANSI,
1986).
Peak sound pressure s the maxmum absolute
value of the nstantaneous sound pressure durng
a specfed tme nterval and s denoted as Pmax n
unts of Pascals (Pa). It s not an averaged pressure.
Peak pressure s a useful metrc for ether pulse or
nonpulse sounds, but t s partcularly mportant
for characterzng pulses (ANSI, 1986; Harrs,
1998, Chapter 12). Because of the rapd rse-tme
of such sounds, t s mperatve to use an adequate
samplng rate, especally when measurng peak
pressure levels (Harrs, 1998, Chapter 18).
Peak-to-peak sound pressure s the algebrac
dfference between the maxmum postve and
maxmum negatve nstantaneous peak pressure.
The mean-squared pressure s the average
of the squared pressure over some duraton. For
nonpulse sounds, the averagng tme s any con-
venent perod suffcently long enough to permt
averagng the varablty nherent n the type of
sound. Note that some of the varablty of the
receved sound typcally arses smply from the
AquaticMammals 2007, 33(4), 498-501, DOI 10.1578/AM.33.4.2007.498
MarineMammalNoiseExposureCriteria 499
relatve movement of a free-rangng anmal and a
source, whether the latter s movng or statonary.
Sound pressure levels (SPLs) are gven as the
decbel (dB) measures of the pressure metrcs
defned above. The root-mean-square (RMS)
SPL s gven as dB re: 1 Pa for underwater sound
and dB re: 20 Pa for aeral sound. Peak SPLs are
gven as dB re: 1 Pa (peak) n water and dB re:
20 Pa (peak) n ar. Peak-to-peak SPLs are dB
re: 1 Pa (peak-to-peak) n water and dB re: 20
Pa (peak-to-peak) n ar. Source level (SL) s the
receved level measured or estmated 1 m from the
source.
Duration s the length of a sound, generally n
seconds. Duraton s mportant because t affects
varous acoustc metrcs, ncludng mean-square
and/or RMS sound pressure (Madsen, 2005).
Because of background nose and reverbera-
ton, duraton can be dffcult to defne precsely.
Varous defntons of duraton exst n the ltera-
ture such as the tme between the ponts on the
pressure-tme waveform P(t) determned to be
ether 10 dB (0.316 tmes) or 20 dB (0.1 tmes)
below the nstantaneous peak pressure (Hamernk
& Hsueh, 1991). Malme et al. (1983, 1984) used a
smlar approach. Harrs (1998, Chapter 11) sug-
gested alternatve constructs, ncludng exponen-
tal tme weghtng. Ths topc s dscussed below
wth regard to updatng measurement standards
for mpulse sounds. Greene (1997) descrbed a
practcal defnton of pulse duraton based on
the nterval over whch 90% of the sound energy
arrved at the recever. Ths nterval could also be
used as the averagng tme for mean-square pres-
sure (Madsen, 2005). Ths approach has been
wdely used n measurng exposure duraton and
SPL values for sesmc argun and ple drvng sg-
nals (e.g., McCauley et al., 1998; Blackwell et al.,
2004b). Defned as such, duraton s the nterval
between the 5% and 95% bounds of the tme-nte-
gral of the nstantaneous sound-pressure squared
(sound exposure [E(t)] as defned below) whle
accountng for background nose and low-level
reverberaton (assumed to be contnuous). That s,
the background nose s measured over a perod of
tme before the pulse occurs and then s subtracted
from the cumulatve sum-of-square pressures to
determne the sum-of-square pressures from the
mpulsve sound alone. Ths s done by manually
dentfyng a perod of tme (t1, t2) precedng the
event, deemed to be representatve of ambent
nose. The mean-square pressure (n Pa
2
) of the
ambent (Pamb)
2
s determned wth the followng
relatonshp:
}

=
2
1
) (
1
2
1 2
2
t
t
amb
dt t P
t t
P (1)
( )
}
=
t
t
amb temp
dt P t P t E
2
2 2
) ( ) ( (2)
( )
}
=
t
t
amb
a
dt P t P t E
2 2
) ( ) ( (3)
( )
}
=
t
t
a
dt t P t E ) ( ) (
2
(4)

}
=
2
1
0
2
10
) (
) (
log 10
ref
N
n
T
n
p
dt t p
SEL (5)
( ) | |
4
4
) (
o

=
X O
X kurt (6)
} ) ( max{
) (
log 20 ) (
10
f R
f R
f M = (7)
) )( (
) (
2 2 2 2
2 2
low high
high
f f f f
f f
f R
+ +
=
(8)
(1) eq.
The temporal (or event) sound exposure
Etemp (t) (n Pa
2
-s) s then calculated as
}

=
2
1
) (
1
2
1 2
2
t
t
amb
dt t P
t t
P (1)
( )
}
=
t
t
amb temp
dt P t P t E
2
2 2
) ( ) ( (2)
( )
}
=
t
t
amb
a
dt P t P t E
2 2
) ( ) ( (3)
( )
}
=
t
t
a
dt t P t E ) ( ) (
2
(4)

}
=
2
1
0
2
10
) (
) (
log 10
ref
N
n
T
n
p
dt t p
SEL (5)
( ) | |
4
4
) (
o

=
X O
X kurt (6)
} ) ( max{
) (
log 20 ) (
10
f R
f R
f M = (7)
) )( (
) (
2 2 2 2
2 2
low high
high
f f f f
f f
f R
+ +
=
(8)
(2) eq.
The 0% sound exposure pont (ta) sgnfes the
start of the acoustc event and the 100% sound
exposure pont (tb) sgnfes the end of the event.
These two ponts are where the E(t) curve begns
to rse and where t levels off, respectvely. Ther
selecton can be dffcult due to varaton n amb-
ent nose precedng (and overlappng) the acoustc
event, as well as reverberaton plus ambent nose
followng the event. Consequently, many nvest-
gators dentfy these ponts subjectvely.
The sound exposure E(t) (n Pa
2
-s), where t
tb, s then calculated as
}

=
2
1
) (
1
2
1 2
2
t
t
amb
dt t P
t t
P (1)
( )
}
=
t
t
amb temp
dt P t P t E
2
2 2
) ( ) ( (2)
( )
}
=
t
t
amb
a
dt P t P t E
2 2
) ( ) ( (3)
( )
}
=
t
t
a
dt t P t E ) ( ) (
2
(4)

}
=
2
1
0
2
10
) (
) (
log 10
ref
N
n
T
n
p
dt t p
SEL (5)
( ) | |
4
4
) (
o

=
X O
X kurt (6)
} ) ( max{
) (
log 20 ) (
10
f R
f R
f M = (7)
) )( (
) (
2 2 2 2
2 2
low high
high
f f f f
f f
f R
+ +
=
(8)
(3) eq.
where E100 = E(tb) s 100% of the sound expo-
sure. For the 5% pont, E(t) s determned as E5
= 0.05E100 = E(t5), whle E(t) for the 95% energy
pont s determned as E95 = 0.95E100 = E(t95).
Thus, E90 = E95 E5 and duraton (Td) = t95 t5 (s)
where the receved pressure level greatly exceeds
the ambent level, eq. 3 can be reduced to
}

=
2
1
) (
1
2
1 2
2
t
t
amb
dt t P
t t
P (1)
( )
}
=
t
t
amb temp
dt P t P t E
2
2 2
) ( ) ( (2)
( )
}
=
t
t
amb
a
dt P t P t E
2 2
) ( ) ( (3)
( )
}
=
t
t
a
dt t P t E ) ( ) (
2
(4)

}
=
2
1
0
2
10
) (
) (
log 10
ref
N
n
T
n
p
dt t p
SEL (5)
( ) | |
4
4
) (
o

=
X O
X kurt (6)
} ) ( max{
) (
log 20 ) (
10
f R
f R
f M = (7)
) )( (
) (
2 2 2 2
2 2
low high
high
f f f f
f f
f R
+ +
=
(8)
(4) eq.
Sound exposure level (SEL) s the decbel
level of the cumulatve sum-of-square pressures
over the duraton of a sound (e.g., dB re: 1 Pa
2
-s)
for sustaned nonpulse sounds where the exposure
s of a constant nature (.e., source and anmal
postons are held roughly constant). However,
ths measure s also extremely useful for pulses
and transent nonpulse sounds because t enables
sounds of dfferng duraton to be characterzed
n terms of total energy for purposes of assessng
exposure rsk.
The SEL metrc also enables ntegratng sound
energy across multple exposures from sources
such as sesmc arguns, ple drvng, and most
sonar sgnals. Several methods exst for summng
energy over multple exposures. We use a rela-
tvely straghtforward approach here, specfcally
500 Southalletal.
}

=
2
1
) (
1
2
1 2
2
t
t
amb
dt t P
t t
P (1)
( )
}
=
t
t
amb temp
dt P t P t E
2
2 2
) ( ) ( (2)
( )
}
=
t
t
amb
a
dt P t P t E
2 2
) ( ) ( (3)
( )
}
=
t
t
a
dt t P t E ) ( ) (
2
(4)

}
=
2
1
0
2
10
) (
) (
log 10
ref
N
n
T
n
p
dt t p
SEL (5)
( ) | |
4
4
) (
o

=
X O
X kurt (6)
} ) ( max{
) (
log 20 ) (
10
f R
f R
f M = (7)
) )( (
) (
2 2 2 2
2 2
low high
high
f f f f
f f
f R
+ +
=
(8)
(5) eq.
where nstantaneous sound-pressure (p) s mea-
sured n Pa for n exposures and the reference
pressure (pref) s 1 Pa under water and 20 Pa n ar.
Ths summaton procedure essentally generates a
sngle exposure equvalent value that assumes
no recovery of hearng between repeated expo-
sures. The approprate unts for underwater SEL
are dB re: 1 Pa-s, and the approprate unts for
aeral SEL are dB re: (20 Pa)
2
-s.
Kurtosis s a statstcal measure of the prob-
ablty dstrbuton of sound pressure ampltudes
(Hamernk & Hsueh, 1991; Le et al., 1994;
Hamernk et al., 2003) that descrbes the shape
of the ampltude dstrbuton. In some regards,
t appears to be a hghly relevant metrc n that
mpulsve sound wth hgh kurtoss and hgh
nstantaneous peak pressure may be partcularly
njurous to some mammals (Hamernk et al.,
2003). Kurtoss s related to the fourth central-
moment and s defned for random varable X as
}

=
2
1
) (
1
2
1 2
2
t
t
amb
dt t P
t t
P (1)
( )
}
=
t
t
amb temp
dt P t P t E
2
2 2
) ( ) ( (2)
( )
}
=
t
t
amb
a
dt P t P t E
2 2
) ( ) ( (3)
( )
}
=
t
t
a
dt t P t E ) ( ) (
2
(4)

}
=
2
1
0
2
10
) (
) (
log 10
ref
N
n
T
n
p
dt t p
SEL (5)
( ) | |
4
4
) (
o

=
X O
X kurt (6)
} ) ( max{
) (
log 20 ) (
10
f R
f R
f M = (7)
) )( (
) (
2 2 2 2
2 2
low high
high
f f f f
f f
f R
+ +
=
(8)
(6) eq.
where O s the expectaton operator, s the mean,
and S s the standard devaton. When kurtoss s
hgh, ampltude dstrbuton s generally more cen-
trally peaked and may have broader tals. Normal
dstrbutons have a kurtoss value of 3 ndepen-
dent of the mean or standard devaton.
Frequency-selective weighting s often employed
to measure (as a sngle number) sound pressure or
energy n a specfc frequency band, wth emphass
or de-emphass on partcular frequences as a func-
ton of the senstvty to those frequences. For aeral
hearng n humans, A-weghtng s derved from the
nverse of the dealzed 40-phon equal loudness hear-
ng functon across frequences standardzed to 0 dB
at 1 kHz (Harrs, 1998), provdng level measures
denoted as dB(A). C-weghtng s determned from
the nverse of the dealzed 100-phon equal loudness
hearng functon (whch dffers n several regards
from the 40-phon functon) standardzed to 0 dB at
1 kHz (Harrs, 1998); level measures are denoted as
dB(C).
Absent equal-loudness contours for marne
mammals, specal weghtng functons based
loosely on human weghtng functons and
general knowledge of functonal hearng band-
wdth, were developed here for the fve functonal
marne mammal hearng groups (see the Marne
Mammal Functonal Hearng Groups secton
n Chapter 2). M-weghtng has a mathematcal
structure smlar to the C-weghtng used n human
hearng, whch reflects the fact that sounds must
be more ntense at hgh and low frequences for
them to have equal audtory effect. C-weghtng
s most approprate for determnng the effects of
ntense soundsthat s, those wth loudness equal
to that of a tone 100 dB above threshold at 1,000
Hz. The M-weghtng was desgned to do much
the same for the dfferent marne mammal groups
wth the only dfference beng ther varyng low-
and hgh-frequency cutoffs. The M-weghtng for
marne mammals, lke the C-weghtng used n
humans, rolls off at a rate of 12 dB per octave.
The general expresson for M-weghtng (M[f]),
usng estmated frequency cutoffs for each func-
tonal marne mammal hearng group, s gven as
}

=
2
1
) (
1
2
1 2
2
t
t
amb
dt t P
t t
P (1)
( )
}
=
t
t
amb temp
dt P t P t E
2
2 2
) ( ) ( (2)
( )
}
=
t
t
amb
a
dt P t P t E
2 2
) ( ) ( (3)
( )
}
=
t
t
a
dt t P t E ) ( ) (
2
(4)

}
=
2
1
0
2
10
) (
) (
log 10
ref
N
n
T
n
p
dt t p
SEL (5)
( ) | |
4
4
) (
o

=
X O
X kurt (6)
} ) ( max{
) (
log 20 ) (
10
f R
f R
f M = (7)
) )( (
) (
2 2 2 2
2 2
low high
high
f f f f
f f
f R
+ +
=
(8)
(7) eq.
where
}

=
2
1
) (
1
2
1 2
2
t
t
amb
dt t P
t t
P (1)
( )
}
=
t
t
amb temp
dt P t P t E
2
2 2
) ( ) ( (2)
( )
}
=
t
t
amb
a
dt P t P t E
2 2
) ( ) ( (3)
( )
}
=
t
t
a
dt t P t E ) ( ) (
2
(4)

}
=
2
1
0
2
10
) (
) (
log 10
ref
N
n
T
n
p
dt t p
SEL (5)
( ) | |
4
4
) (
o

=
X O
X kurt (6)
} ) ( max{
) (
log 20 ) (
10
f R
f R
f M = (7)
) )( (
) (
2 2 2 2
2 2
low high
high
f f f f
f f
f R
+ +
=
(8)
(8) eq.
The estmated lower and upper functonal hear-
ng lmts (flow and fhgh) for each of the fve func-
tonal marne mammal hearng groups and the
names of the frequency-weghtng functons are
gven n Table 2. The weghtng functons de-
emphasze frequences that are near the lower and
upper frequency ends of the estmated hearng
range as ndcated by the negatve relatve values
n Fgure 1.
Audition (hearing) s a well-developed and pr-
mary sensory modalty for most, f not all, marne
vertebrates (Schusterman, 1981; Tyack, 1998; Fay
& Popper, 2000). The vertebrate hearng system
nvolves codng, processng, ntegratng, and
respondng to sound n a varety of ways, some
not outwardly evdent (Yost, 2000).
Hearing (auditory) threshold s most com-
monly measured by behavoral or electrophys-
ologcal responses and s defned as the SPL of
the quetest sound audble n some percentage
of expermental trals. In ar, measurements are
often conducted n specally constructed sound
chambers. When that s not possble, tests must be
conducted n low background nose condtons to
yeld meanngful threshold data.
Sensation level represents the dfference (n
dB) between the overall level of a sound and the
recevers audtory threshold at smlar sound fre-
quences. It s partcularly useful as a means of
comparng the relatve exposure level of a sound
MarineMammalNoiseExposureCriteria 501
for ndvduals that may have dfferent hearng
capabltes. Sensaton level s sometmes abbre-
vated SL, but ths s not done n ths document to
avod confuson wth the very dfferent concept of
source level.
Auditory masking s the partal or complete
reducton n the audblty of sgnals due to the
presence of nterferng nose (see Buus, 1997).
The degree of maskng depends on the spectral
and temporal relatonshps between sgnals and
maskng nose as well as ther respectve RLs
(e.g., Fletcher, 1940).
Sound localization s the determnaton of
source locaton based on features of receved
sounds. Ths crtcal, complex process of the
audtory system can nvolve the detecton of
sounds produced drectly by a source (passve ls-
tenng) or the detecton of echoes reflected off a
target (as n the case of bosonar).
Auditory scene analysis s the process by whch
the audtory system sorts out related elements of
a complex acoustc envronment nto those arsng
from dscrete sound sources. Ths process s sm-
lar to psychologcal processes underlyng vsual
percepton whereby many dfferent vsual mages
are perceved as dscrete elements wthn a vsual
scene.
Temporary Threshold Shift (TTS) s a revers-
ble elevaton n hearng threshold (.e., a non-
permanent reducton n hearng senstvty) most
commonly resultng from nose exposure.
Permanent Threshold Shift (PTS) s a perma-
nent elevaton n hearng threshold (.e., an unre-
coverable reducton n hearng senstvty). PTS
can occur from a varety of causes, but t s most
often the result of ntense and/or repeated nose
exposures. In that case t s also referred to as nose
nduced hearng loss (NIHL) or nose nduced per-
manent threshold shft (NIPTS).
Courtesy: A. Fredlander
Appendix B. Studies Involving Marine Mammal Behavioral
Responses to Multiple Pulses
Low-Frequency Cetaceans/Multiple Pulses
(Cell 2)
Numerous feld observatons have been made
of low-frequency cetaceans reactng to multple
pulses, ether opportunstcally exposed to ongo-
ng operatons or by ntentonal exposures. A mod-
erate number of speces and expermental cond-
tons have been consdered, but the source was
usually a sesmc argun or arrays of these ntense
sources. Some studes focused on mgratng an-
mals observed from fxed observaton platforms or
n/near mgraton corrdors.
The general results of the severty scalng
analyss for ths condton suggest the onset of
more sgnfcant behavoral dsturbances from
multple pulses for mgratng bowhead whales at
RLs (RMS over pulse duraton) around 120 dB
re: 1 Pa (Rchardson et al., 1999). For all other
low-frequency cetaceans (ncludng feedng bow-
head whales), ths onset was at RLs around 150
to 160 dB re: 1 Pa (Malme et al., 1983, 1984;
Rchardson et al., 1986; Ljungblad et al., 1988;
Todd et al., 1996; McCauley et al., 1998). There s
essentally no overlap n the RLs assocated wth
the onset of behavoral responses by members of
these two groups based on the nformaton cur-
rently avalable.
Sesmc arguns operated near bowhead whales
generally ntate avodance reactons as well as
changes n locomoton and respraton (Reeves
et al., 1984; Rchardson et al., 1985, 1986, 1999;
Ljungblad et al., 1988). Durng the autumn mgra-
ton, avodance behavor has been observed at rel-
atvely great (20+ km) ranges from source opera-
tons (Kosk & Johnson, 1987; Rchardson et al.,
1999). Ljungblad et al. (1988) dd not nvestgate
behavoral reactons over such ranges. Durng
the summer, feedng bowheads exhbted subtle
behavoral responses but not actve avodance
at dstances beyond 6 km from argun sources
(Rchardson et al., 1986; see also Mller et al.,
2005).
Rchardson et al. (1999) studed autumn-
mgratng bowhead whale and found avodance
by most ndvdual whales at dstances up to 20
km and some avodance at 20 to 30 km. Sesmc
surveys usng argun arrays wth 6 to 16 guns and
total volumes of 560 to 1,500 n
3
were conducted
n shallow (generally < 20 m) water of the Alaskan
Beaufort Sea. Whales n ther westward autumn
mgraton over three seasons (1996 to 1998) were
detected wth aeral surveys on days wth and
wthout sesmc survey actvty. Usng the obser-
vatons of dozens of mgratng whales durng per-
ods when arguns were not actve, we were able
to calculate the percentage of observed whales
durng sesmc surveys that demonstrated avod-
ance behavor at varous RLs (see Table 7). These
results ndcate that mgratng bowhead whales n
the Rchardson et al. (1999) study often avoded
areas where RLs exceeded 120 to 130 dB re: 1 Pa
(RMS over pulse duraton).
In contrast, Rchardson et al. (1986) observed
qute dfferent movement patterns of bowhead
whales exposed to sesmc argun sounds on
ther summer feedng grounds n the Canadan
Beaufort Sea. Receved levels from a sngle ses-
mc argun (0.66-L) were measured in situ near
ndvdual whales beng observed 3 to 5 km from
the sound source, and ranged from 118 to 133 dB
re: 1 Pa. Vsual orentaton by groups of whales
durng argun exposure was observed on two of
fve occasons; only mnor changes n swm-
mng and respraton patterns were observed.
Rchardson et al. (1986) also made opportunstc
observatons of groups of bowhead whales near
a sesmc vessel operatng an argun array. At the
hghest RLs, some measurements exceeded the
dynamc range of the recordng equpment and are
consdered exposure mnma, although ths was
not the case for most measurements relevant to
the behavoral observatons. From these observa-
tons and the controlled exposure to sounds from a
sngle argun, Rchardson et al. (1986) concluded
that some whales responded subtly by changng
dvng and breathng patterns at relatvely low
RLs (ca. 120 to 140 dB re: 1 Pa), but that avod-
ance and other more profound behavoral changes
were generally not observed unless the RL was
160 dB re: 1 Pa.
Ljungblad et al. (1988) conducted a seres of
acoustc experments on behavoral reactons of
bowhead whales exposed to sounds from shps
wth operatng argun(s). Experment 1 was con-
ducted on a group of eght whales. When a ses-
mc vessel approached to wthn 3.5 km (max.
RL near observed ndvduals was 142 dB re: 1
Pa), the bowhead whales coalesced and moved
n a tght group away from the approachng vessel.
AquaticMammals 2007, 33(4), 502-508, DOI 10.1578/AM.33.4.2007.502
MarineMammalNoiseExposureCriteria 503
Experment 2 nvolved a group of three bowhead
whales that demonstrated startle responses at the
onset of sounds from an argun 7 km away (max.
measured RL was 165 dB re: 1 Pa). Behavor
returned to pre-exposure values shortly after the
operaton was termnated. Experment 3 nvolved a
group of seven bowhead whales that demonstrated
avodance behavor at ranges of ~3.5 km (max.
measured RL of 178 dB re: 1 Pa). Experment
4 nvolved a group of 50 bowhead whales.
Behavoral reactons were frst observed at ranges
of about 8 km (max. measured RLs of 157 dB re:
1 Pa) and avodance behavor was noted at ~3
km (RLs ~165 dB re: 1 Pa). Avodance behavor
n ths nstance smlarly abated shortly followng
cessaton of exposure (and was thus assgned a
behavoral score of 6).
Recent work on summerng bowhead whales
by Mller et al. (2005) also found that avodance
responses were lmted to dstances of at most a
few klometers and RLs exceedng 160 dB re: 1
Pa. Mller et al. conducted a montorng program
over two summers for varous marne mammals
offshore of the Mackenze Delta n the Southeast
Beaufort Sea before and durng sesmc surveys.
They presented observatonal data from both
vessel-based and aeral observatons of bowhead
whales, belugas, and several pnnped speces.
The general methodology s brefly dscussed
here as well as data on behavoral responses by
low-frequency cetaceans (bowhead whales) and
the correspondng rank on the severty scale.
The argun operatons nvolved 3-D sesmc
proflng from a 67-m vessel usng two dent-
cal 2,250 n
3
sleevegun arrays, each wth 24 ar-
guns. Shots were at 8-s ntervals and at a depth of
5 m below the surface of the water. Surveys were
conducted n very shallow water (13 m average).
Acoustc montorng wth calbrated hydrophones
across the 10 Hz to 24 kHz bandwdth was con-
ducted whle sesmc operatons were underway.
Physcal propertes of the operatonal envron-
ment, and hence sound propagaton n the shal-
low water envronments, were hghly varable,
but RLs as a functon of range from actve argun
arrays were measured. Vessel-based observers
and aeral surveyors used lne-transect methods
to montor marne mammals n and adjacent to
sesmc operatonal areas, both before and durng
shootng. Bowhead whales observed durng the
perods concdent wth sesmc operatons were
presumed to be feedng (.e., not mgratng). Many
bowheads (355 ndvduals n 232 groups) were
seen by marne mammal observers aboard the ses-
mc vessel. Sghtng rates were lower and mean
sghtng dstances were somewhat larger durng
sesmc operatons than at tmes when the arguns
were not operatng, but the zone of avodance
around actve arguns was very lmted. The
approxmate dfference n mean sghtng dstance
was ~600 m. Smlarly, the aeral surveyors dd
not detect any large-scale avodance of the argun
operatons by bowheads. These observatons were
generally consstent for both years n whch mea-
surements were made and are generally consstent
wth the observatons of Rchardson et al. (1986)
n the same regon and season (summer). Anmals
not exhbtng observable behavoral reactons
(response score: 0) were consstently sghted n
areas where RLs very lkely ranged from 130 to
180 dB re: 1 Pa. The general lack of sghtngs
wthn a small area around the sesmc vessel sug-
gests behavoral avodance (response score: 6) at
RLs exceedng 180 dB re: 1 Pa. Exposures were
not estmated to exceed 190 dB re: 1 Pa. The
entre study was treated as a sngle observaton for
the purposes of the behavoral analyss. Half of
the observaton was scored as avodance behav-
or and half as no response, wth exposure RL bns
from 130 to 190 dB re: 1 Pa (Table 6).
The combned data for bowhead whale avod-
ance of argun sounds (Rchardson et al., 1986,
1999; Ljungblad et al., 1988; Mller et al., 2005)
ndcated that, when mgratng, these anmals can
be partcularly prone to behavoral dsturbance,
wth the onset of sgnfcant responses occur-
rng at approxmately 120 dB re: 1 Pa (RMS
over pulse duraton) (Table 6). In contrast, when
feedng, they may show subtle effects at low RLs
but only tend to dsplay actve avodance at RLs
exceedng 160 dB re: 1 Pa.
Low-frequency cetaceans, other than mgratng
bowhead whales, appear to be much more toler-
ant of exposure to multple pulses, although data
are lmted to a few speces and (prmarly) argun
sources. Avalable data for speces other than bow-
heads nclude reactons to opportunstc and nten-
tonal exposures of humpback whales (Malme
et al., 1985; Todd et al., 1996; McCauley et al.,
1998, 2000) and gray whales (Malme et al., 1983,
1984, 1986, 1988; also see revew by Moore &
Clarke, 2002). Todd et al. (1996), Malme et al.
(1983, 1984), and McCauley et al. (1998) are
ncluded n the behavoral scorng analyss here
because they contan suffcent nformaton on
exposures and ndvdual responses of low-fre-
quency cetaceans other than bowhead whales.
Todd et al. (1996) analyzed the mpact of con-
structon actvty (explosons and drllng) on the
entanglement of three foragng humpback whales
off Newfoundland. They conducted observatons
of whale behavor durng and followng explo-
sons and obtaned acoustc measurements of
underwater sound sgnatures. The data suggest
few short-term changes n movement and behav-
or patterns n response to dscrete exposures;
504 Southalletal.
however, repeated exposures to hgh levels may
have resulted n sensory mparment n whales and
perhaps greater susceptblty to entanglement n
fshng gear.
Malme et al. (1983, 1984) documented behav-
oral reactons of mgratng gray whales to sesmc
pulses from both sngle arguns and an array. Only
land-based observers were used, whch meant that
the observers could not have affected the whales
behavor. Both phases of the nvestgaton yelded
the general concluson that RLs exceedng 160 dB
re: 1 Pa (on an approxmate RMS bass) were
requred to cause mgratng gray whales to avod
argun sounds, although statstcally sgnfcant
reactons that were less profound occurred at
much larger ranges and lower levels. From ther
emprcal phase II results, Malme et al. (1984)
calculated 10, 50, and 90% probabltes of gray
whale avodance reactons n these condtons to
be 164, 170, and 180 dB re: 1 Pa, respectvely.
McCauley et al. (1998) made behavoral obser-
vatons of mgratng humpback whales off western
Australa durng sesmc operatons wth a sngle
argun and several argun array confguratons.
Sesmc track lnes were orented perpendcular to
the mgraton paths of humpback whales movng
through the area. Aeral surveys were conducted
to determne the presence of humpback whales
movng through the survey area. Detaled obser-
vatonal data were presented for ndvduals and
groups of whales; RLs were measured at var-
able ranges. The sesmc survey dd not appear to
grossly affect the mgraton of humpback whales
through the area; however, avodance behav-
or was observed to begn at ranges from 5 to 8
km and to be almost unversal at ranges of 1 to
4 km. Exposures to a sngle argun (20 n
3
) were
extrapolated to equvalent ranges for exposure to
full arrays based on emprcal measurements. The
data ndcated an onset of behavoral avodance at
~159 dB re: 1 Pa (peak-to-peak), roughly equva-
lent to the full array at 5 km. General behavoral
avodance (most ndvduals) occurred at a range
of about 1 km for the sngle gun (~168 dB re: 1
Pa [peak-to-peak]), equvalent to the full array at
about 3 km. Some ndvdual whales dd approach
closer than the typcal 3-km stand-off range; these
may have been males nvestgatng the presence of
the low-frequency source.
In addton to presentng agan the results gven
n the McCauley et al. (1998) paper, McCauley
et al. (2000) provde addtonal behavoral observa-
tons of 16 humpback whale pods that approached
as a sngle argun (Bolt PAR 600b 20-n
3
) was
operated. These whales were also observed after
termnaton of argun operatons. These trals were
conducted n a large embayment (Exmouth Gulf)
as the anmals were engaged n a varety of restng
and socal behavors. Fve trals were excluded
from consderaton n our analyss, but behavoral
observatons were reported for the remanng 11.
Of these, ten ncluded cow pods of varous szes,
and one was a lone male. Snce the cow pods were
not mgratng and were not ndvdually dent-
fed, a sngle behavoral observaton s ncluded
n Table 7 for the ten observatons. The results
for the cow pods were very consstent, ndcatng
clear avodance (severty score = 6) of the argun
at exposures n the 140 to 150 dB re: 1 Pa range
(RMS over pulse duraton). The lone male essen-
tally gnored the argun untl wthn ca. 100 m,
when the receved level approached 180 dB re:
1 Pa (RMS); ths response may have had as much
or more to do wth the presence of the vessel than
exposure to the argun sound. Notng ths con-
textual complexty here, a sngle observaton for
ths ndvdual s reported n the 170 to 180 dB re:
1 Pa exposure bn n Table 7 as general avod-
ance (severty score = 6).
Mid-Frequency Cetaceans/Multiple Pulses
(Cell 5)
A lmted number of behavoral observatons have
been made of md-frequency cetaceans exposed to
multple pulses. Feld observatons have nvolved
sperm whales and a few other odontocete speces
exposed to sesmc arguns and small explosves
(Madsen & Mhl, 2000; Madsen et al., 2002;
Mller et al., 2005). Laboratory nvestgatons have
consdered behavoral responses to varous knds
of multple pulse sources (Akamatsu et al., 1993).
As n most crtera cells, a number of reported
observatons were not scored and reported here
due to lack of relevant nformaton and dffcultes
n accountng for varous contextual varables. A
summary of those studes used and others consd-
ered s gven n Table 8; the severty scalng analy-
ss for Cell 5 s shown n Table 9.
The combned data for md-frequency ceta-
ceans exposed to multple pulses do not ndcate
a clear pattern of ncreasng probablty and sever-
ty of response wth ncreasng RLs. In certan
condtons, multple pulses at relatvely low RLs
(~80 to 90 dB re: 1 Pa) temporarly slence nd-
vdual acoustc behavor for one speces (sperm
whales). In other cases wth slghtly dfferent
stmul, RLs n the 120 to 180 dB re: 1 Pa range
faled to elct observable reactons from a sgnf-
cant percentage of ndvduals of the same speces,
both n the feld and n the laboratory.
FieldObservations(Cell5)
Madsen & Mhl (2000) nvestgated sperm whale
responses to small underwater detonators that
ncluded 1-g TNT charges, producng a 1-ms
MarineMammalNoiseExposureCriteria 505
broadband (300 Hz to 15 kHz) pulse; several
charges were trggered per day. Echolocaton clck
behavor was montored, and one whale was local-
zed acoustcally. Ths ndvdual demonstrated no
modulaton of vocal behavor when exposed to an
RMS-equvalent RL of ~173 dB re: 1 Pa. There
was also one observaton of a whale exposed to
179 dB re: 1 Pa; t contnued breathng normally
wth no vsble response.
Madsen et al. (2002) studed responses of sperm
whales n Norway to sounds assocated wth ds-
tant sesmc survey operatons. Calbrated RLs
for ndvduals and correlated acoustc behavor
are reported for three dscrete sghtngs over a 5-d
perod. The frst observaton nvolved three sperm
whales tracked by acoustc localzaton wthn a
dspersed array of calbrated hydrophones, whch
also recorded argun sounds from an array operat-
ng 40 km away. RL at the poston of the whale
was estmated to be 123 dB re: 1 Pa. The second
observaton (3 d later) nvolved a sngle sperm
whale recorded before, durng, and after argun
exposure at a range of 86 km; measured RL was
130 dB re: 1 Pa. The thrd observaton (2 d later)
nvolved three ndvduals; the survey vessel was
94 km away and measured RL was 130 dB re:
1 Pa. No change n sperm whale acoustc behav-
or was noted n any of these cases. The authors
also played artfcal codas and notced that two
whales drected ther sonar beams at the speaker,
but nsuffcent nformaton s gven to assocate
ths response wth a partcular RL.
Mller et al. (2005) documented behavoral reac-
tons of varous marne mammal speces, nclud-
ng belugas, to argun operatons. The general
methodology s detaled above (see the Cell 2
secton). Owng to ther normal seasonal patterns
n the Beaufort Sea, belugas were most abundant
n the Mller et al. (2005) study area pror to the
start of sesmc operatons. There were relatvely
few vessel-based sghtngs, most of whch were
made when arguns were not actve. Many belugas
were observed durng aeral surveys, however, and
these data were used to compare beluga sghtngs
wthn concentrc 10-km bands around the actve
sesmc source wth sghtng rates n non-argun
condtons. Durng argun operatons, Mller et al.
detected sgnfcantly fewer anmals 10 to 20 km
from sesmc operatons and an unexpectedly hgh
number of sghtngs n the 20- to 30-km zone. Ths
was suggestve of behavoral avodance of sesmc
operatons at dstances up to 20 km. These observa-
tons may n part explan why so few anmals were
observed by shpboard marne mammal observers.
Mller et al. noted that the apparent avodance of
sesmc operatons was much greater than expected
f the whales were respondng to non-argun
sounds assocated wth vessel operaton. For the
purposes of our behavoral analyses, the combned
beluga results were treated as a sngle observa-
ton that was subdvded equally nto ether avod-
ance behavor or no observable response. Belugas
exposed to RLs of 100 to 120 dB re: 1 Pa (RMS
over pulse duraton) were determned to have had
no observable reacton (response score: 0) to ses-
mc exposures. RLs between 120 and 150 dB re: 1
Pa were determned to have nduced temporary
avodance behavor (response score: 6) n belugas,
based on the vessel-based and aeral observatons.
Based on both the vessel-based and aeral surveys,
exposures apparently dd not exceed 150 dB re:
1 Pa. Weghted behavoral response scores for
each of these fve exposure RL bns are gven n
Table 7.
Several studes nvolved behavoral reactons
of free-rangng, md-frequency cetaceans but
lacked specfc measures to be ncluded drectly
n our analyses. Andr et al. (1997) exposed sperm
whales to varous stmul, ncludng two pulse
sounds (recorded coda playbacks and a 10-kHz
pulse). A sgnfcant number of exposed whales
exhbted vocal modulatons and modfed dvng
behavor, but nsuffcent nformaton s avalable
on receved exposures of ndvdual whales. Stone
(2003) compled a large database of sghtng
data of several md-frequency cetacean speces
observed from sesmc survey vessels. Sghtng
rates of small odontocetes were sgnfcantly lower
when arguns were frng, and they were sghted at
greater dstances from vessels, ndcatng avod-
ance behavor. The study sponsors (JNCC) kndly
provded raw data for use n our quanttatve
avodance analyses, but they are not ncluded due
to dffcultes n estmatng exposure RL for nd-
vdual sghtngs. (See also Stone & Tasker, 2006,
for a recently publshed account.)
LaboratoryObservations(Cell5)
Akamatsu et al. (1993) nvestgated avod-
ance behavor n two captve false kller whales
exposed to 15 dfferent knds of sounds, ncludng
pulse sequences (manual strkes on a metal ppe
once every 2 s) n the 24 to 115 kHz range. For
ths stmulus, no avodance was seen followng
the frst exposure (174 dB re: 1 Pa), but tempo-
rary avodance behavor (response score: 6) was
observed for successve exposures at 174 and 178
dB re: 1 Pa.
Fnneran et al. (2000) observed behavoral
responses of two captve bottlenose dolphns and
a beluga whale durng TTS experments nvolvng
a seres of mpulsve exposures desgned to rep-
lcate dstant explosons. Each anmal exhbted
alteratons of nomnal traned behavors (reluc-
tance to return to expermental statons) durng the
experment; the onset of behavoral dsturbance
506 Southalletal.
occurred n the beluga at 220 dB re: 1 Pa (peak-
to-peak) and n the two bottlenose dolphns at 196
and 209 dB re: 1 Pa (peak-to-peak), respectvely.
In a related study, Fnneran et al. (2002b) observed
behavoral responses of a bottlenose dolphn and a
beluga whale after exposure to mpulsve sounds
produced by a water gun. Both ndvduals showed
a smlar reluctance to return to expermental sta-
tons (beluga at 202 dB re: 1 Pa (peak-to-peak);
bottlenose dolphn at 229 dB re: 1 Pa [peak-to-
peak]). Romano et al. (2004) studed physolog-
cal responses to these exposures n these same
anmals. They observed clear neuro-mmune
responses n the beluga at exposures above 222
dB re: 1 Pa (peak-to-peak) and sgnfcant dffer-
ences n aldosterone and monocyte counts n the
dolphn for exposures exceedng 225 dB re: 1 Pa
(peak-to-peak).
High-Frequency Cetaceans/Multiple Pulses
(Cell 8)
Based on our source type dstncton (see Chapter
2), vrtually all sound sources used n behavoral
studes of hgh-frequency cetaceans (e.g., acoustc
harassment devces [AHDs] and acoustc deterrent
devces [ADDs]) would be characterzed as non-
pulses. Whle ndvdual elements produced by
some of these sources would be characterzed as
pulses, and sequences of them as multple pulses,
they are generally emtted n such rapd fashon
that mammalan audtory systems are lkely to
perceve them as nonpulses. Further, some AHDs,
ADDs, and all other sources used n behavoral
studes wth hgh-frequency cetaceans lack the
characterstcs of pulses. Due to the lack of data, t
s not possble to present any behavoral response
data on multple pulses for hgh-frequency ceta-
ceans; avalable data for nonpulse sounds are
consdered elsewhere (see the Hgh-Frequency
Cetaceans/Nonpulses [Cell 9] sectons of Chapter 4
and Appendx C). We note the need for behavoral
research on these anmals usng sound sources
unequvocally classfed as pulses.
Pinnipeds in Water/Multiple Pulses (Cell 11)
Informaton on behavoral reactons of pnnpeds
n water to multple pulses s derved from studes
usng small explosves smlar to those used n fsh-
eres nteractons, constructon actvty, and ses-
mc surveys. Several studes lacked matched data
on acoustc exposures and behavoral responses by
ndvduals. As a result, the quanttatve nforma-
ton on reactons of pnnpeds n water to multple
pulses s very lmted. Our general fndng s that
exposures n the ~150 to 180 dB re: 1 Pa range
(RMS over pulse duraton) generally have lmted
potental to nduce avodance behavor n pnn-
peds, whereas RLs exceedng 190 dB re: 1 Pa are
lkely to elct responses, at least n some rnged
seals (Harrs et al., 2001; Blackwell et al., 2004b;
Mller et al., 2005).
Harrs et al. (2001) documented responses
of pnnpeds (prmarly rnged seals, but a few
bearded and spotted seals) and obtaned calbrated
measures of RLs wthn defned spatal zones
durng operaton of a sngle argun, an 11-argun
array totalng 1,320 n
3
, and durng control per-
ods. Vsual observatons from the sesmc vessel
were lmted to the area wthn a few hundred
meters, and 79% of the seals observed were wthn
250 m of the vessel. Durng daylght, seals were
observed at nearly dentcal rates wth no arguns,
one argun, or when a full argun array was frng.
Seals were sgnfcantly further away durng full
array operatons compared to the other two con-
dtons. Also, there was some avodance wthn
150 m of the vessel n these condtons (0.37 seals
seen per hour n control perods compared to 0.21
seals/h durng full array operatons). Sesmc
operatons were not beleved to cause many, f
any, seals to desert the operatonal area.
Blackwell et al. (2004b) nvestgated behav-
oral reactons of rnged seals to mpact sounds
assocated wth the drvng of steel ppes n the
constructon of an ol producton faclty. Multple
strkes were recorded under water at dstances up
to 3 km from the source. Unweghted peak pres-
sure level, SPL, and SEL measurements were
made at varous dstances. At the closest pont (63
m), RLs were 151 dB re: 1 Pa (RMS), 157 dB re:
1 Pa (peak), and 145 dB re: 1 Pa
2
-s (SEL). Pulses
had measurable components extendng to over 10
kHz, although more than 95% of the energy n the
sgnals was below 225 Hz. A frequency-weght-
ng metrc somewhat smlar to that proposed here
was appled to the recorded sgnals n estmatng
audblty ranges. Indvduals demonstrated no
or low-level behavoral responses to ple-drvng
sounds, but were somewhat responsve to helcop-
ter overflghts. Blackwell et al. noted, however,
that ther data were collected after a prolonged
perod of ntensve constructon actvty and may
reflect the least responsve part of the orgnal
populaton of seals that may have already habtu-
ated to the nose source. Indvdual observatons
n whch helcopters were not present are consd-
ered n our behavoral analyss, weghted by the
total number of relevant observatons (Table 11).
Aeral measurements of multple pulse exposures
were also obtaned n ths study and are consd-
ered n the relevant condton below.
Mller et al. (2005) documented behavoral
reactons of varous marne mammal speces,
ncludng pnnpeds n water, to sesmc argun
MarineMammalNoiseExposureCriteria 507
operatons. The general methodology s detaled
above (see the Cell 2 secton). The vast major-
ty (> 90%) of the seals were rnged seals and
the remander were bearded seals. Vessel-based
observers saw seals around the vessel, and often
qute close to t, throughout the perod of ses-
mc operatons. Seals were observed sgnfcantly
further away durng argun operatons n the frst
summer, whereas the reverse pattern was actu-
ally the case n the second season. Combned, the
results suggest essentally no observable behav-
oral response n pnnpeds exposed to sesmc
sgnals n these specfc condtons. Based on the
acoustc measurements that were conducted and
the areas n whch these pnnpeds were observed,
RLs were lkely 170 to 200 dB re: 1 Pa (RMS
over pulse duraton). A sngle observaton of no
reacton (response score: 0) for pnnpeds n water
s reported for ths study and s weghted equally
across these exposure RL bns (Table 8).
Several other studes were deleted from our
analyss due to a lack of certan nformaton.
Two studes nvestgated small frecracker-lke
explosves (called seal bombs) and ther effect
on the underwater behavor of pnnpeds around
fshng gear (Shaughnessy et al., 1981; Mate &
Harvey, 1987). Intally, these explosves tend to
nduce the desred avodance behavor, but ths
response fades quckly due to habtuaton (see
Rchardson et al., 1995). Mate & Harvey (1987)
reported farly extensve descrptons of startle
and temporary avodance data as well as some
nformaton on exposure condtons. Besdes the
challengng matter of nterpretng the apparently
rapd habtuaton to ths sound source, however,
data are lackng that relate dscrete exposures wth
defned behavoral responses of specfc ndvdual
pnnpeds. For these reasons, we excluded data
on responses to seal bombs from our analyss.
Moulton et al. (2003, 2005) conducted surveys
of rnged seal dstrbuton before and durng the
constructon and operaton of the same ol produc-
ton faclty descrbed by Blackwell et al. (2004a,
2004b). Sound sources ncluded nonpulse as well
as multple pulse sources (ncludng mpact ple-
drvng). Ther observatons across multple sea-
sons ndcated lttle or no behavoral avodance of
the area n response to varous ndustral actv-
tes. Due to dffcultes wth control observatons
across seasons and the lack of nformaton about
dscrete exposures and ndvdual reactons, how-
ever, we excluded the Moulton et al. (2003, 2005)
data from our analyss. A fnal study for whch
avalable data were nsuffcent for ncluson here
s Thompson et al. (1998). That telemetry study
seemed to show much hgher responsveness of
gray and harbor seals to argun sounds than has
been demonstrated n other studes, whch reled
on vsual observatons. Thus, future studes may
show some seals to be more responsve to multple
pulses than Table 11 would suggest.
Pinnipeds in Air/Multiple Pulses (Cell 11)
The effects of multple aeral pulses on pnnpeds
are among the least well-documented of the cond-
tons we consdered. Most of the avalable data on
responses to pulses are from sngle-pulse events
(e.g., rocket launches) over populatons of pn-
npeds exposed to such sgnals repeatedly (e.g.,
Thorson et al., 1998, 1999, 2000a, 2000b; Berg
et al., 2001, 2002, 2004). These launches are not
repeated so frequently that the exposure can be
consdered as nvolvng multple pulses, and many
of the exposures nclude nonpulse components.
However, they are dscussed n some detal n ths
appendx (as well as n Appendx C for nonpulses
wthn these studes) along wth several other stud-
es potentally relevant to Cell 14 but ultmately
not used n the analyss here. Consequently, the
quanttatve nformaton analyzed for reactons of
pnnpeds n ar exposed to multple pulses (see
Table 12 for summary and Table 13 for severty
scalng analyss) focuses on the aeral data of
Blackwell et al. (2004b). These extremely lmted
data suggest very mnor, f any, observable behav-
oral responses for exposures rangng from 60 to
80 dB re: 20 Pa.
Blackwell et al. (2004b) reported behavoral
reactons of rnged seals to aeral mpact sounds
from ple-drvng (descrbed above). Multple
strkes were recorded n ar at dstances up to
500 m from the source. Unweghted SPL, peak
sound pressure levels, and SEL measurements
were made at varous dstances. At the closest pont
(63 m) average RLs were 93 dB re: 20 Pa (RMS),
111 dB re: 20 Pa (peak), and 87 dB re: (20 Pa)
2
-
s (SEL). Mean pulse duratons were between
0.17 and 0.63 s, wth measurable energy to over
10 kHz, but wth 95% of the energy occurrng
between 89 and 3,534 Hz. A frequency-weght-
ng metrc somewhat smlar to that proposed here
was appled to the recorded sgnals n estmat-
ng audblty ranges. Indvduals demonstrated
very lmted behavoral responses to ple-drvng
sounds n some condtons (most appeared ether
ndfferent or curous) but were more respon-
sve to helcopter overflghts. Data were collected
after prolonged constructon actvtes, and some
habtuaton probably had taken place already.
Indvdual observatons for whch helcopters
were not present are consdered n the behavoral
analyss here and weghted by the total number of
relevant observatons (Table 13) to equal a sngle
observaton for the study.
508 Southalletal.
Perry et al. (2002) measured the effects of
repeated (0 to 5/d) sonc booms from Concorde
arcraft on harbor and gray seals on Sable Island,
Nova Scota. They measured the number of an-
mals on shore before and after booms as well as
the frequency of varous behavors. Addtonally,
they compared heart rates n exposure and control
condtons usng recordng devces deployed on
the anmals. They reported receved sound over-
pressure of booms on the breedng beaches of
both pnnped speces. Observed effects on anmal
presence, behavor, and heart rate were generally
mnor and not statstcally sgnfcant; anmals
were largely tolerant of the sounds but became
somewhat more alert followng them. However,
Perry et al. (2002) note that there s a long hs-
tory of sonc booms from arcraft n the area and
the anmals are lkely habtuated to ther presence.
Due to ths complcaton and the lack of explct
receved SPL measures at exposed ndvduals,
we dd not score the results of Perry et al. (2002)
here.
Appendix C. Studies Involving Marine Mammal Behavioral
Responses to Nonpulses
Low-Frequency Cetaceans/Nonpulses
(Cell 3)
Whle there are clearly major areas of uncertanty
remanng, there has been relatvely extensve
behavoral observaton of low-frequency ceta-
ceans exposed to nonpulse sources. As sum-
marzed n Table 14, these feld observatons
nvolve the majorty of low-frequency cetacean
speces exposed to a wde range of ndustral,
actve sonar, and tomographc research actve
sources (Baker et al., 1982; Malme et al., 1983,
1984, 1986; Rchardson et al., 1990b; McCauley
et al., 1996; Frankel & Clark, 1998; Borggaard
et al., 1999; Basson et al., 2000; Croll et al.,
2001; Palka & Hammond, 2001; Nowacek et al.,
2004). Observatons from several related studes
(Dahlhem, 1987; Frankel & Clark, 2000, 2002;
Schck & Urban, 2000; Moore & Clarke, 2002;
Jahoda et al., 2003; Mobley, 2005) were revewed
brefly but not analyzed here because key nfor-
maton was lackng.
These papers generally ndcate no (or very
lmted) responses at RLs 90 to 120 dB re: 1 Pa
and an ncreasng probablty of avodance and
other behavoral effects n the 120 to 160 dB re:
1 Pa range (Table 14). However, the data also
ndcate consderable varablty n RLs assoc-
ated wth behavoral responses. Contextual var-
ables (e.g., source proxmty, novelty, operatonal
features) appear to have been at least as mportant
as exposure level n predctng response type and
magntude.
Baker et al. (1982) nvestgated behavoral
responses of ndvdual humpback whales to vessel
traffc n southeast Alaska. Indvdual RLs were
not reported, but suffcent nformaton regardng
ndvdual ranges was obtaned to approxmate
exposures gven that the acoustc characterstcs
of dentcal classes of vessel classes nvolved
were measured n smlar condtons by Mles &
Malme (1983). Results ndcate some behavoral
avodance when RL was n the 110 to 120 dB re:
1 Pa range and clear avodance at 120 to 140 dB
re: 1 Pa.
Malme et al. (1983, 1984) used playback meth-
ods to document behavoral reactons of mgrat-
ng gray whales to ntermttent sounds of hel-
copter overflghts and contnuous sounds from
drllng rgs and platforms. Both phases of the
nvestgaton yelded the general concluson that
RLs exceedng 120 dB re: 1 Pa nduced demon-
strable behavoral reactons (avodance). Malme
et al. (1984) calculated 10%, 50%, and 90%
probabltes of gray whale avodance reactons
n these condtons as 110, 120, and 130 dB re:
1 Pa. Malme et al. (1986) observed the behavor
of feedng gray whales durng four expermental
playbacks of drllng sounds (50 to 315 Hz; 21-
mn overall duraton and 10% duty cycle; source
levels 156 to 162 dB re: 1 Pa-m). In two cases
for RLs 100 to 110 dB re: 1 Pa, there was no
observed behavoral reacton. Avodance behavor
was observed n two cases where RLs were 110 to
120 dB re: 1 Pa.
Rchardson et al. (1990b) performed 12 play-
back experments n whch bowhead whales n the
Alaskan Arctc were exposed to drllng sounds.
Low-frequency source characterstcs and trans-
msson loss were well-characterzed, enablng
RL estmates to be made for ndvdual cases.
Whales generally dd not respond to exposures n
the 100 to 130 dB re: 1 Pa range, although there
was some ndcaton of mnor behavoral changes
n several nstances.
Usng dfferent detecton and samplng tech-
nques, McCauley et al. (1996) reported several
cases of humpback whales respondng to vessels
n Hervey Bay, Australa, along wth measure-
ments of nose RL. Not all cases reported provded
suffcent nformaton to assocate a response or
lack of response wth exposure, but n three cases,
ndvdual responses and nose RL were reported.
Results ndcated clear avodance at RLs between
118 to 124 dB re: 1 Pa.
Palka & Hammond (2001) analyzed lne transect
census data n whch the orentaton and dstance
off transect lne were reported for large numbers of
mnke whales. General addtve models were used
to estmate the range at whch cetaceans respond
to the nose of the research vessel by approach or
avodance. The typcal avodance dstance for 272
mnke whales n the Gulf of Mane was 717 m;
for 352 mnke whales n the North Sea, t was 563
m; and for 493 mnke whales n the Northeastern
Atlantc, t was 695 m. Receved levels were est-
mated based on a nomnal source level for that
class of research vessel (ca. 170 to 175 dB re:
1 Pa-m) and an assumpton of sphercal (20 log
R) spreadng loss (54 dB loss @ 500 m; lkely
AquaticMammals 2007, 33(4), 509-521, DOI 10.1578/AM.33.4.2007.509
510 Southalletal.
reasonable for these condtons). These data are
represented n Table 14 by the 110 to 120 dB re:
1 Pa exposures and a relatvely low (less severe)
behavoral response score of three (.e., mnor
changes n locomoton speed, drecton, and/or
dvng profle).
Several addtonal studes have used playback
experments wth actve sound sources to nvest-
gate the behavoral reactons of low-frequency ceta-
ceans to nonpulse sources. Basson et al. (2000) and
Mller et al. (2000) report behavoral observatons
for humpback whales exposed to a low-frequency
sonar stmulus (160- to 330-Hz frequency band;
42-s tonal sgnal repeated every 6 mn; source levels
170 to 200 dB re: 1 Pa-m). Measured RLs ranged
from 120 to 150 dB re: 1 Pa. In nne cases, nd-
vdual whales contnued sngng throughout expo-
sures, whle n four nstances, ndvduals ceased
callng when they joned another whale. The cessa-
ton of song and jonng another ndvdual s typ-
cal of normal mystcete socal nteractons (Tyack,
1981). Consequently, these events were not scored
as a vocal response to the playback but as a mod-
erate orentng behavor (severty score = 2). For
the remanng fve playbacks, ndvdual whales
stopped sngng durng exposure wthout jonng
other whales (severty scale = 4). Although sngers
also stop spontaneously under control condtons,
the latter fve expermental trals were consdered
vocal cessaton resultng from sound exposure
(Basson et al., 2000). However, there are nsuf-
fcent data to compare control and expermental
cases for spontaneous rates of cessaton. Analyss
of all sngers ndcated an ncrease n song dura-
ton durng exposure due to ncreased repetton
of elements of the song. Snce t was possble that
some ndvdual whales were represented multple
tmes wthn the playbacks, the Basson et al.
(2000) and Mller et al. (2000) data were scored as
a sngle behavoral observaton. The 18 ndvdual
observatons were weghed nversely by the total
number (1/18) n Table 15.
Croll et al. (2001) nvestgated responses of for-
agng fn and blue whales to the same LFA sonar
stmulus off southern Calforna. Unlke the pre-
vous two studes, where ndvdual expermental
subjects were tracked on a behavoral scale, ths
study used sghtng data on an ecologcal scale.
Playbacks and control ntervals wth no trans-
msson were used to nvestgate behavor and
dstrbuton on tme scales of several weeks and
spatal scales of tens of klometers. Sghtngs and
whale dvng behavor were not random but were
related to envronmental features such as the con-
tnental shelf break and ts effects on prey abun-
dance rather than operaton and locaton of the
nonpulse sonar source. The general concluson
was that whales remaned feedng wthn a regon
for whch 12 to 30% of exposures exceeded 140
dB re: 1 Pa. A sngle observaton was scored for
ths study because ndvdual responses were not
reported.
Frankel & Clark (1998) conducted playback
experments wth wnterng humpback whales
around the Bg Island of Hawa. The sound source
was a sngle speaker producng a low-frequency
M-sequence (sne wave wth multple-phase
reversals) sgnal n the 60 to 90 Hz band. Ths was
smlar n bandwdth to the ATOC source, but had
a much lower output level (172 dB re: 1 Pa @
1 m). A vertcal lne array of calbrated hydrophones
was deployed from a spar buoy to measure receved
sgnals in situ. Detaled observatons of many
behavoral patterns (ncludng respraton, dvng,
and general movements) were recorded before,
durng, and after playback (n = 50) and control
(n = 34) sequences. A sngle tral also nvolved
playback of humpback foragng sounds. Most of
the playback sequences nvolved very low-level
RLs, ca. 90 to 120 dB re: 1 Pa, though not spec-
fed n suffcent detal to nclude n the analyss
here. For 11 playbacks, exposures were between
120 and 130 dB re: 1 Pa and ncluded suff-
cent nformaton regardng ndvdual responses.
Durng eght of the trals, there were no measur-
able dfferences n tracks or bearngs relatve to
control condtons, whereas on three occasons,
whales ether moved slghtly away from (n = 1) or
towards (n = 2) the playback speaker durng expo-
sure. Because t was not possble to determne
whether the same ndvdual whales were repre-
sented more than once n the playback sequences,
a sngle observaton was recorded for Frankel &
Clark (1998), wth 0.73 of ths observaton (8/11)
scored as a 0 (no response) and 0.27 (3/11) scored
as a 3 (mnor changes n locomoton speed, drec-
ton, and/or dvng). A fnal mportant observaton
from the detaled statstcal analyss by Frankel &
Clark was that the presence of the source vessel
tself had a greater effect than dd the M-sequence
playback.
Fnally, Nowacek et al. (2004) used controlled
exposures to demonstrate behavoral reactons of
northern rght whales to varous nonpulse sounds.
Playback stmul ncluded shp nose; socal
sounds of conspecfcs; and a complex, 18-mn
alert sound consstng of repettons of three
dfferent artfcal sgnals (alternatng 1-s pure
tones [500 and 850 Hz]; a 2-s, tonal, frequency
downsweep [4,500 to 500 Hz]; and a par of 1-
s pure tones [1,500 Hz and 2,000 Hz] ampltude
modulated at 120 Hz). A total of ten whales were
tagged wth calbrated nstruments that measured
receved sound characterstcs and concurrent
anmal movements n three dmensons. Fve out
of sx exposed whales reacted strongly to alert
MarineMammalNoiseExposureCriteria 511
sgnals at measured RLs between 130 and 150 dB
re: 1 Pa (.e., ceased foragng and swam rapdly
to the surface; severty scale = 7). Two of these
ndvduals were not exposed to shp nose and
are gven as a dscrete observaton n Table 15,
whereas the other four were exposed to both stm-
ul and thus weghted as 0.5 (1/2) observatons for
the respectve RL and severty score. These whales
reacted mldly to conspecfc sgnals (not scored
here because of bologcal sgnals). Seven whales,
ncludng the four exposed to the alert stmulus,
had no measurable response to ether shp sounds
or actual vessel nose. Ths study by Nowacek
et al. ncluded the careful expermental desgn,
controls, and detaled nformaton on exposure and
ndvdual behavoral response that were requred
for behavoral analyss. More studes of ths type
and rgor are urgently needed (see Chapter 5).
We revewed addtonal studes concernng low-
frequency cetaceans and nonpulse sounds but dd
not nclude them n the analyss here, generally due
to the absence of key nformaton. Dahlhem (1987)
exposed gray whales to playbacks of outboard
nose, gray whale calls, and tonal sounds. Whales
sgnfcantly ncreased callng rate and modfed
call structure for sources other than the test tone
(the latter caused all vocalzaton to cease). Durng
and followng longer duraton playbacks of ol
drllng and kller whale sounds wth more precse
trackng of gray whale locatons, ndvduals spent
more tme mllng, and whales remaned farther off-
shore durng kller whale playbacks. Unfortunately,
nsuffcent nformaton s presented to assocate
changes wth specfc RLs. Borggaard et al. (1999)
measured the effects of ndustral actvty on several
mystcete speces n Newfoundland, but nsuffcent
nformaton s reported on ndvdually dscernble
responses. Schck & Urban (2000) appled statst-
cal methods to assess spatal avodance of actve
drllng rgs by bowhead whales, but no acoustc
data are reported. Moore & Clarke (2002) synthe-
szed prevously publshed data (all consdered
separately above) on numerous nonpulse sources,
n order to assess the avodance probablty of gray
whales for varous exposure RLs. Jahoda et al.
(2003) studed ndvdual responses of fn whales
(n = 25) to close rapd approaches of small vessels;
18 observatons ncluded control and expermental
data. Clear behavoral responses were observed,
but nether RL nor range from source to ndvdu-
als were gven. Results are further complcated by
whale taggng attempts from the vessel. Frankel
& Clark (2000) and Mobley (2005) nvestgated
the dstrbuton of humpback whales n Hawa n
relaton to the operaton of a low-frequency tomo-
graphc source (ca. 75 Hz; 37.5-Hz nomnal band-
wdth; 20-mn duraton every 2 h durng daylght
hours; source level: 195 dB re: 1 Pa-m). Frankel
& Clark (2000) observed whales from a land staton
and determned that the average dstance between
the sound source and the whale groups sghted was
sgnfcantly greater durng source operaton. These
and other data were also consdered n the context
of other factors affectng humpback whale dstrbu-
ton off the sland of Kaua. Mobley (2005) con-
ducted aeral surveys n each of three years (2001,
source off; 2002 & 2003, source on) durng the
peak season of humpback resdency. Abundance
and dstrbuton of whales were very smlar n the
area surroundng the source over all three years;
small dfferences n sghtng rates, sghtng loca-
ton depth, and dstances from the source and shore
were not statstcally sgnfcant. Frankel & Clark
(2002) and Mobley (2005) lack explct data on
RLs assocated wth ndvdual behavoral observa-
tons, whch precludes ther ncluson here.
Mid-FrequencyCetaceans/Nonpulses(Cell6)
A relatvely large number of md-frequency
cetaceans have been observed n the feld and n
the laboratory respondng to nonpulse sounds,
ncludng vessels and watercraft (LGL &
Greenerdge, 1986; Gordon et al., 1992; Palka
& Hammond, 2001; Buckstaff, 2004; Morsaka
et al., 2005), pulsed pngers and ADD/AHDs
(Watkns & Schevll, 1975; Morton & Symonds,
2002; Montero-Neto et al., 2004), ndustral
actvtes (Awbrey & Stewart, 1983; Rchardson
et al., 1990b), md-frequency actve mltary
sonar (NRL, 2004a, 2004b; NMFS, 2005), and
tones or bands of nose n laboratory condtons
(Nachtgall et al., 2003; Fnneran & Schlundt,
2004). Summary nformaton on these stud-
es s gven n Table 16. As n other condtons,
a number of potentally relevant feld studes are
not ncluded n the severty scalng analyss due to
lack of suffcently detaled nformaton.
An addtonal challenge n nterpretng many of
the feld data for ths condton s solatng the effect
of RL from the effects of mere source presence (as
possbly ndcated by vsual stmul or other aspects
of acoustc exposure such as the presence of hgh-
frequency components) and other contextual var-
ables. For ths reason, several studes were consd-
ered but not ntegrated nto the analyss.
The laboratory observatons are of captve ceta-
ceans exposed to precsely controlled and known
nose exposures n the context of hearng and TTS
experments. The relevance of behavoral reac-
tons of traned, food-renforced captve anmals
exposed to nose n assessng reactons of free-rang-
ng marne mammals s not well-known, however
(dscussed below).
The combned feld and laboratory data for
md-frequency cetaceans exposed to nonpulse
sounds do not lead us to a clear concluson about
512 Southalletal.
RLs concdent wth varous behavoral responses
(see severty scalng, Table 17). In some settngs,
ndvduals n the feld showed profound (and
what we regard here as sgnfcant) behavoral
responses to exposures from 90 to 120 dB re:
1 Pa, whle others faled to exhbt such responses
for exposure RLs from 120 to 150 dB re: 1 Pa.
Contextual varables other than exposure RL, and
probable speces dfferences, are the lkely reasons
for ths varablty. Context, ncludng the fact that
captve subjects were often drectly renforced
wth food for toleratng nose exposure, may also
explan why there s great dsparty n results
from feld and laboratory condtonsexposures
n captve settngs generally exceeded 170 dB re:
1 Pa before nducng behavoral responses.
FieldObservations(Cell6)
The most extensve seres of observatons regard-
ng vessels and watercraft s from LGL and
Greenerdge (1986) and Fnley et al. (1990), who
documented belugas and narwhals (Monodon
monoceros) congregated near ce edges reactng
to the approach and passage of ce-breakng shps.
Over a 3-y perod (1982 to 1984), they used both
ce-based local observatons of whales and aeral
surveys, and also made detaled acoustc measure-
ments. The survey method made t dffcult to
assess ndependent groups of anmals. Some large-
scale groupngs could be dentfed for several df-
ferent dsturbance perods, however. Pre-ds-
turbance group sze was ~3; we dvded reported
numbers of dsturbed herds by three to estmate
the number of ndependent groups. Aeral surveys
n 1984 lumped sghtngs by mnute, whch cor-
responded to about 3.4 km n dstance. We consd-
ered ths dstance suffcent to treat each mnute as
an ndependent unt for avodance analyss. The
responses of both speces over a 3-y perod were
generally smlar to responses they make to preda-
tors as descrbed by Inut hunters.
Beluga whales responded to oncomng ves-
sels by (1) fleeng at speeds of up 20 km/h from
dstances of 20 to 80 km, (2) abandonng normal
pod structure, and (3) modfyng vocal behavor
and/or emttng alarm calls. Narwhals, n contrast,
generally demonstrated a freeze response, lyng
motonless or swmmng slowly away (as far as
37 km down the ce edge), huddlng n groups,
and ceasng sound producton. There was some
evdence of habtuaton and reduced avodance 2
to 3 d after onset. Due to the detaled and exten-
sve nature of these observatons, data from each
season, and how they are nterpreted here, are
gven n detal.
The 1982 season observatons by LGL &
Greenerdge (1986) nvolved a sngle passage
of an cebreaker wth both ce-based and aeral
measurements on 28 June 1982. Four groups of
narwhals (n = 9 to 10, 7, 7, and 6) responded when
the shp was 6.4 km away (exposure RLs of ~100
dB re: 1 Pa n the 150- to 1,150-Hz band). At
a later pont, observers sghted belugas movng
away from the source at > 20 km (exposure RLs
of ~90 dB re: 1 Pa n the 150- to 1,150-Hz band).
The total number of anmals observed fleeng was
about 300, suggestng approxmately 100 nde-
pendent groups (of three ndvduals each), whch
s the sample sze used here. No whales were
sghted the followng day, but some were sghted
on 30 June, wth shp nose audble at spectrum
levels of approxmately 55 dB re 1 Pa/Hz (up to
4 kHz).
Observatons durng 1983 (LGL & Greenerdge,
1986) nvolved two ce-breakng shps wth aeral
survey and ce-based observatons durng seven
samplng perods. As the frst vessel approached
at a dstance of about 65 km, ce-based observ-
ers noted reactons from both narwhals (seven
groups) and belugas (eght groups) (exposure
RLs of ~101 to 105 dB re: 1 Pa n the 20- to
1,000-Hz band). After 22 h wthout operaton, the
vessel commenced ce-breakng, and a second
cebreaker approached (exposure RLs of ~120
dB re: 1 Pa n the 20- to 1,000-Hz band). Ths
resulted n the rapd movement of > 225 belugas
(estmated as a sample sze of 75 for ths analy-
ss); belugas were nether seen nor heard for
the remander of the second observaton perod.
Behavoral responses were also observed for
10 groups of narwhals. A total of 73 narwhals
were seen and/or heard, but ther reactons are
not clearly reported and are thus excluded from
analyss here. At the onset of the thrd samplng
perod, followng a 4.5-h slent nterval, four
narwhal groups were observed n nomnal socal
behavor (dvng and vocalzng). An ce-breakng
vessel operated ntermttently, but no change was
observed n narwhal behavor. Belugas n the area
dd modfy vocalzaton parameters durng opera-
tons (exposure RLs of ~116 dB re: 1 Pa n the
20- to 1,000-Hz band). A 6-h quet nterval was
followed by 10.5 h of ce-breakng operaton, but
bad weather precluded anmal observatons. After
an addtonal 9-h hatus, ce-breakng commenced
agan by both vessels (exposure RLs of ~121 dB
re: 1 Pa n the 20- to 1,000-Hz band). Ice-based
observers documented 14 narwhals and 11 belu-
gas leavng the area, and aeral surveys ndcated
80% of 673 belugas movng away from sound
sources (estmated number of groups calculated
as [.8]*[673/3] = 179.5). As nose levels from ce-
breakng operatons dmnshed, a total of 45 nar-
whals returned to the area and engaged n dvng
and foragng behavor. The sxth observaton
perod followed 6.5 h wthout a vessel n the area,
MarineMammalNoiseExposureCriteria 513
durng whch 30 belugas (estmated as 10 groups)
and 15 narwhals (estmated as fve groups) were
observed dvng n the area (exposure RLs of
~105 dB re: 1 Pa n the 20- to 1,000-Hz band).
A sngle beluga vocal response was noted at RL
= 116 dB re: 1 Pa n the 20- to 1,000-Hz band.
Aeral surveys ndcated dense concentratons of
narwhals (n = 50) and belugas (n = 400) appar-
ently foragng well away from the dsturbance ste.
Durng the fnal samplng perod, followng an
8-h quet nterval, no reactons were seen from 28
narwhals and 17 belugas (exposure RLs rangng
up to 115 dB re: 1 Pa).
The fnal season (1984) reported n LGL &
Greenerdge (1986) nvolved aeral surveys before,
durng, and after the passage of two ce-breakng
shps. The lack of ce camps precluded acoustc
measurements as well as behavoral observatons.
A prelmnary survey was conducted the day
before operatons, and an addtonal aeral survey
was conducted as both shps commenced operat-
ng. Durng operatons, no belugas and few nar-
whals were observed n an area approxmately 27
km ahead of the vessels, and all whales sghted
over 20 to 80 km from the shps were swmmng
strongly away. Addtonal observatons confrm
the remarkable spatal extent of avodance reac-
tons to ths sound source n ths context. In the
absence of acoustc measurements, however, t
was necessary to estmate RLs from the detaled
data from the same ce-breakng vessel durng the
prevous season.
Behavoral responses at farly low exposure
RLs are suggested by studes of some other md-
frequency cetaceans as well. Gordon et al. (1992)
conducted opportunstc vsual and acoustc mon-
torng of sperm whales n New Zealand exposed
to nearby whale-watchng boats (wthn 450 m).
Indvduals could not be used as the unts of
analyss because t was dffcult to re-sght spe-
cfc ndvduals durng both exposure and control
condtons. Sperm whales respred sgnfcantly
less frequently, had shorter surface ntervals, and
took longer to start clckng at the start of a dve
descent when boats were nearby than when they
were absent. Nose spectrum levels of whale-
watchng boats ranged from 109 to 129 dB re:
1 Pa/Hz. Over a bandwdth of 100 to 6,000 Hz,
equvalent broadband source levels are ~157 dB
re: 1 Pa-m; RLs at a range of 450 m are ~104 dB
re: 1 Pa.
Palka & Hammond (2001) appled a General
Addtve Model to lne transect data to estmate
the range at whch md-frequency cetaceans typ-
cally responded to the nose of research vessels.
The subjects were Atlantc whte-sded dolphns
n the Gulf of Mane and whte-beaked dolphns
(Lagenorhynchus albirostris) n the North Sea.
The whte-sded dolphns exhbted smple avod-
ance behavor (as ndcated by ther orentatons)
out to an estmated range of 592 m based on 85
group sghtngs (n > 1). Whte-beaked dolphns
actually approached vessels between 150 and 300
m away, but demonstrated avodance at dstances
of 300 to 700 m. Typcal avodance dstance was
estmated as 716 m based on 48 groups sghted.
Buckstaff (2004), usng repeated samples
of the behavor of 14 ndvdual bottlenose dol-
phns, observed 1,233 vessel approaches (wthn
400 m) near Sarasota, Florda. Dolphn whstle
rates became elevated before vessel nose was
detectable to the researcher lstenng va towed
hydrophones. Vessel RLs measured near dolphn
subjects ranged from 113 to 138 dB re: 1 Pa.
Dolphn vocal responses were observed before
vessel sounds were audble, and apparently
occurred wth RLs n the 110 to < 120 dB re: 1
Pa category.
Morsaka et al. (2005) compared whstles from
three populatons of Indo-Pacfc bottlenose dol-
phns (Tursiops aduncus). One populaton was
exposed to vessel nose wth spectrum levels of
~85 dB re: 1 Pa/Hz n the 1- to 22-kHz band
(broadband RL ~128 dB re: 1 Pa) as opposed to
~65 dB re: 1 Pa/Hz n the same band (broadband
RL ~108 dB re: 1 Pa) for the other two stes.
Dolphn whstles n the noser envronment had
lower fundamental frequences and less frequency
modulaton, suggestng a shft n sound param-
eters as a result of ncreased ambent nose.
Morton & Symonds (2002) used census data
on kller whales n Brtsh Columba to evaluate
avodance of nonpulse AHD sources. They con-
sdered unusually long tme scales, comparng
pre-exposure data from 1985 to 1992, exposure
from 1993 to 1998, and post-exposure from 1999
to 2000. The response data were smply pres-
ence or absence, makng t dffcult to assess RLs.
Usng some montorng and reasonable assump-
tons, however, they estmated audblty ranges
throughout the complex study area. Avodance
ranges were ca. 4 km. Also, there was a dramatc
reducton n the number of days resdent kller
whales were sghted durng AHD-actve perods
compared to pre- and post-exposure perods and
a nearby control ste. Morton & Symonds dd not
ndcate how many pods were nvolved n ther
analyss. Consequently, we assume a sngle nde-
pendent group n our analyss.
Montero-Neto et al. (2004) studed avodance
responses of tucux (Sotaliafluviatilis) to Dukane

Netmark ADDs. Source characterstcs are not
gven, but dentcal devces were used by Culk et
al. (2001), and acoustc parameters are reported n
detal there (and n the Cell 9 secton). In a total
of 30 exposure trals, ~5 groups each demonstrated
514 Southalletal.
sgnfcant avodance compared to 20 pnger off
and 55 no-pnger control trals over two quad-
rats of about 0.5 km
2
. Nether avodance range
nor RLs are gven, but based upon a central ds-
tance from the quadrat of 10 m, and assumng 15
log R transmsson loss n ths shallow envron-
ment (water depth 1 to 5 m), estmated exposure
RLs were ~115 dB re: 1 Pa.
The only specfc stuaton nvolvng exposure
of wld marne mammals to actve md-frequency
mltary sonar for whch exposure condtons are
known wth any degree of specfcty nvolved
ncdental exposure of kller whales to sounds
from the naval vessel USS Shoup (NRL, 2004a,
2004b; NMFS, 2005). A group (J-pod) of south-
ern resdent kller whales n the eastern Strat of
Juan de Fuca and Haro Strat, Washngton, was
observed by researchers before, durng, and after
the approachng USSShoup transmtted sonar sg-
nals from ts 53C sonar at a source level of ca. 235
dB re: 1 Pa-m once every ca. 28 s for several
hours. At ts pont of closest approach, the mean
drect-path RL wthn a specfed area around the
anmals was ca. 169 dB re: 1 Pa (NRL, 2004a,
2004b). As ndcated by NMFS (2005), there s
some dscrepancy n nterpretaton of the behav-
oral responses among researchers who were
ether on the water or who observed vdeo record-
ngs of behavoral responses. The lead researcher
followng and observng the anmals durng the
event ndcated that ndvduals n the group dem-
onstrated abnormal avodance behavor, most
dramatcally at the pont of closest approach.
However, the behavor of the whales apparently
returned to normal wthn a short perod followng
cessaton of sonar transmssons. A severty score
of 6 (mld/moderate avodance) s subsequently
reported n the 160 to 170 dB re: 1 Pa bn for ths
sngle observaton of the group.
Awbrey & Stewart (1983) played back sem-sub-
mersble drllshp sounds (source level: 163 dB re:
1 Pa-m) to belugas n Alaska. They reported avod-
ance reactons at 300 and 1,500 m and approach
by groups at a dstance of 3,500 m (RLs ~110 to
145 dB re: 1 Pa over these ranges assumng a
15 log R transmsson loss). Smlarly, Rchardson
et al. (1990b) played back drllng platform
sounds (source level: 163 dB re: 1 Pa-m) to belu-
gas n Alaska. They conducted aeral observatons
of eght ndvduals among ~100 spread over an
area several hundred meters to several klometers
from the sound source and found no obvous reac-
tons. Moderate changes n movement were noted
for three groups swmmng wthn 200 m of the
sound projector.
A number of addtonal studes (Rendell &
Gordon, 1999; Chlvers & Corkeron, 2001;
Bordno et al., 2002; Wllams et al., 2002; Cox et
al., 2003; Haste et al., 2003; Lusseau, 2003; Foote
et al., 2004; Schefele et al., 2005) were revewed
n detal. The results were excluded from Table 17
due to lmted or no nformaton on anmal num-
bers and/or locaton relatve to the source, acous-
tc propertes of sources, propagaton varables, or
receved exposure condtons. The general obser-
vatons of each study are gven here brefly. Haste
et al. (2003) documented ncreased swmmng
and dvng synchrony of bottlenose dolphns off
northern Scotland n the presence of vessel traf-
fc. Lusseau (2003) observed effects on behavor
of New Zealand bottlenose dolphns wthn 400 m
of boats. Chlvers & Corkeron (2001) consdered
dfferences n behavor of bottlenose dolphns that
do and do not forage around trawlers. Wllams
et al. (2002) observed that some kller whales adopt
erratc movement patterns, suggestve of avod-
ance, when whale-watchng vessels accelerate to
ntersect the whales course. RLs of vessel sound
ncreased approxmately 14 dB wth ncreased
speed assocated wth leapfroggng. Bordno et al.
(2002) determned that ADDs were ntally effec-
tve at reducng by-catch of Francscana dolphns
n Argentne subsstence gllnet fsheres. Cox
et al. (2003) nvestgated reactons of bottle-
nose dolphns to Dukane

NetMark 1000 ADDs

attached to commercal gllnets and found very
lmted to no behavoral avodance. A group of
long-fnned plot whales (Globicephala melas)
demonstrated sgnfcant elevatons of whstle
rates followng each exposure to md-frequency
mltary sonar reported to be at a hgh level
(Rendell & Gordon, 1999).
Fnally, two recent papers deal wth mportant
ssues relatng to changes n marne mammal vocal
behavor as a functon of varable background
nose levels. Foote et al. (2004) found ncreases n
the duraton of kller whale calls over the perod
1977 to 2003, durng whch tme vessel traffc n
Puget Sound, and partcularly whale-watchng
boats around the anmals, ncreased dramatcally.
Schefele et al. (2005) demonstrated that belugas
n the St. Lawrence Rver ncreased the levels of
ther vocalzatons as a functon of the background
nose level (the Lombard Effect). (See also
Parks et al., 2007, for a related new paper on mys-
tcetes.) These papers demonstrate some degree of
plastcty n the vocal sgnal parameters of marne
mammals n response to the ambent condton
(lkely affected by the presence of human sound
sources). These studes were not partcularly
amenable to the knd of analyss conducted n the
severty scalng. We note the partcular mpor-
tance of drect measurements of nose mpacts on
marne mammal vocalzaton and communcaton
systems.
MarineMammalNoiseExposureCriteria 515
LaboratoryObservations(Cell6)
Several researchers conductng laboratory exper-
ments on hearng and the effects of nonpulse
sounds on hearng n md-frequency cetaceans
have reported concurrent behavoral responses.
Nachtgall et al. (2003) reported that nose expo-
sures up to 179 dB re: 1 Pa and 55-mn duraton
affected the traned behavors of a bottlenose dol-
phn partcpatng n a TTS experment. Fnneran
& Schlundt (2004) provded a detaled, compre-
hensve analyss of the behavoral responses of
belugas and bottlenose dolphns to 1-s tones (RLs
160 to 202 dB re: 1 Pa) n the context of TTS
experments. Romano et al. (2004) nvestgated the
physologcal responses of a bottlenose dolphn
and a beluga exposed to these tonal exposures and
demonstrated a decrease n blood cortsol levels
durng a seres of exposures between 130 and 201
dB re: 1 Pa. Collectvely, the laboratory observa-
tons suggested the onset of behavoral response
at hgher RLs than dd feld studes (see Table 16).
The dfferences were lkely related to the very df-
ferent condtons and contextual varables between
untraned, free-rangng ndvduals vs laboratory
subjects that were rewarded wth food for tolerat-
ng nose exposure.
High-Frequency Cetaceans/Nonpulses (Cell 9)
Numerous controlled studes have been con-
ducted recently on the behavoral reac-
tons of hgh-frequency cetaceans to var-
ous nonpulse sound sources both n the feld
(Culk et al., 2001; Johnston, 2002; Olesuk
et al., 2002) and n laboratory settngs (Kastelen
et al., 1997, 2000, 2005, 2006a). However, only
one hgh-frequency speces (harbor porpose) has
been extensvely studed. For that speces, suf-
fcent data are avalable to estmate behavoral
response magntude vs receved exposure cond-
tons. The orgnal studes were attempts to reduce
harbor porpose by-catch by attachng warnng
pngers to fshng gear. More recent studes con-
sder whether ADDs and AHDs also exclude
harbor porposes from crtcal habtat areas and
whether these devces affect harbor porpose
behavor n controlled laboratory condtons.
The combned wld and captve anmal data
(summarzed n Table 18) clearly support the
observaton that harbor porposes are qute sens-
tve to a wde range of human sounds at very low
exposure RLs (~90 to 120 dB re: 1 Pa), at least for
ntal exposures. Ths observaton s also evdent
n the severty scalng analyss for Cell 9 (Table
19). All recorded exposures exceedng 140 dB
re: 1 Pa nduced profound and sustaned avod-
ance behavor n wld harbor porposes. Harbor
porposes also tend to avod boats, although
Dalls porposes do not (Rchardson et al., 1995).
Whether ths apparently hgh degree of behavoral
senstvty by harbor porposes to anthropogenc
sounds extends to other hgh-frequency cetacean
speces (or to nonpulse sources other than ADDs,
AHDs, and boats) s unknown. However, gven
the lack of nformaton to the contrary, such a
relatonshp should be assumed as a precautonary
measure.
Habtuaton to sound exposure was noted n
some but not all studes. In certan feld cond-
tons, strong ntal reactons of hgh-frequency
cetaceans at relatvely low RLs appeared to wane
rather rapdly wth repeated exposure (Cox et al.,
2001). In contrast, several laboratory observatons
showed lttle or no ndcaton of reduced behav-
oral senstvty as a functon of exposure exper-
ence (Kastelen et al., 1997, 2005).
FieldObservations(Cell9)
Kraus et al. (1997) found (and Barlow & Cameron,
2003, later confrmed) that ADDs can affect by-
catch rates of harbor porposes n commercal
fshng applcatons. Kraus et al. (1997) found that
nets wth Dukane

pngers (10-kHz fundamental

frequency, 300-ms duraton, 132 dB re: 1 Pa
source level) were suffcently avoded that sg-
nfcantly fewer porposes were entangled than n
nets lackng pngers. Ther observatons suggest
an ADD avodance range of at least 10 m (expo-
sure RL ~110 dB re: 1 Pa) but are not explct
enough n documentng exposure condtons or
ndvdual responses to nclude n the behavoral
scorng analyss here.
Culk et al. (2001) conducted behavoral
observatons of groups of harbor porposes near
Vancouver Island before, durng, and after the
removal of a PICE pnger (eght dfferent wde-
band swept frequency sgnals between 20 and
160 kHz; 300-ms duraton at random ntervals
[5 to 30 s]; max. broadband SL = 145 dB re:
1 Pa @ 1 m). Source characterstcs of the alarm
were known, but propagaton measurements were
not made insitu. Exposure RLs are estmated here
based on source characterstcs and assumptons
regardng propagaton, allowng for measures
of smlar sources n smlar condtons. A large
excluson zone of approxmately 530-m radus
surroundng actve acoustc alarms was observed
(correspondng to exposure RLs of ~90 to 100 dB
re: 1 Pa). Indvdual sghtng and avodance data
durng CEE actve and control condtons were
scored for ndvduals wthn and outsde ths
excluson zone (see Table 18).
Johnston & Woodley (1998) conducted an exten-
sve survey of AHDs used n the Bay of Fundy to
exclude pnnpeds from salmon aquaculture stes.
Based on the behavoral observatons of Olesuk
516 Southalletal.
et al. (1996), Johnston & Woodley (1998) deter-
mned that harbor porposes were lkely beng
excluded from extensve areas of mportant habtat
as a result of overlappng AHD deployments. Ths
study lacked the dscrete observatonal data neces-
sary for analyss here, but two subsequent studes
contaned such measurements for harbor porposes
exposed to AHDs.
Olesuk et al. (2002) conducted a controlled
exposure n whch they deactvated an array of
four Armar

AHDs for 3 wk and then actvated

the array for three 3-wk ntervals over an 18-wk
perod. Source characterstcs are known (10-kHz
fundamental frequency; 194 dB re: 1 Pa-m (peak-
to-peak) source level; seres of 1.8-ms pulses,
repeated at 40-ms ntervals grouped nto 2.3-s
trans separated by 2.1-s quet perods). However,
no emprcal acoustc measurements were obtaned.
Exposure RLs were estmated here based on source
characterstcs and smple assumptons about the
propagaton of hgh-frequency sounds n shallow-
water envronments. Actve AHDs resulted n clear
avodance behavor by ndvduals and groups of
harbor porposes. The sghtng rate durng actve
perods as a percent of that durng control perods
was only 1.4% at ranges from 400 to 599 m, 2.5
to 3.3% at 600 to 2,499 m, and 8.1% at 2.5 to 3.5
km. RLs at 3.5 km were estmated to be ~123 dB
re: 1 Pa. Avodance data durng actve and control
perods were scored here for ndvduals wthn the
varous exposure zones (Table 18).
Johnston (2002) observed smlar harbor por-
pose behavoral avodance of actve AHDs. They
used an Armar

dB II Plus AHD System (10-kHz

fundamental frequency; 180 dB re: 1 Pa-m source
level, producng a short tran of 2.5-ms sgnals
repeated every 17 s) deployed from a small boat.
They sghted fewer anmals when the AHD was
actve, and these anmals were sgnfcantly fur-
ther away than durng control phases. Approxmate
exposure RLs at the pont of closest approach were
estmated here as ~128 dB re: 1 Pa; mean clos-
est approach dstance was consstent wth exposure
RLs of ~125 dB re: 1 Pa.
Addtonal feld observatons of harbor por-
poses suggest that ther apparently hgh degree
of behavoral senstvty extends to sources other
than ADDs and AHDs. Koschnsk et al. (2003)
observed behavoral responses of harbor porposes
to smulated wnd turbne nose (max. energy
between 30 and 800 Hz; spectral densty source
levels of 128 dB re: 1 Pa/Hz at 80 and 160 Hz).
They sghted harbor porposes at greater ranges
durng playbacks of smulated wnd turbne nose
and observed that anmals more frequently used
echolocaton sgnals durng ndustral actvty.
These data are not scored here, however, due to
lmted avalable nformaton about nose exposure
condtons and ndvdual behavoral responses.
Fnally, whle ther study was not consdered n
the severty scalng here, we note the mportance
of the Cox et al. (2001) observatons regardng
harbor porpose habtuaton. They found that wld
porposes were ntally dsplaced by approx-
mately 208 m from actve ADDs, but ths dsplace-
ment decreased by 50% n 4 d, and reached control
levels n 10 to 11 d. Because of the potental for
habtuaton, t should be noted that many or most
of the feld observatons reported here, other than
those that nvolve long-duraton deployments, are
lkely most relevant for nave ndvduals.
LaboratoryObservations(Cell9)
Relatvely extensve laboratory data are avalable
on captve, ndvdual hgh-frequency cetaceans
exposed to some of the same acoustc alarms
(ADDs and AHDs) and scarng devces deployed
n feld applcatons. We appled our behavoral
scorng paradgm to data from each of the captve
studes conducted by Kastelen and colleagues,
whch ncluded relatvely detaled nformaton on
ndvdual responses and drectly measured expo-
sure RLs.
Kastelen et al. (1997) recorded behavoral
responses (locaton, swmmng speed, and
respraton patterns) of a nave, captve harbor
porpose exposed to a varety of clcks, tones, and
frequency sweeps. All of the relatvely low expo-
sure RLs (~90 to 115 dB re: 1 Pa) resulted n
strong behavoral avodance (subjects bascally
swam rapdly as far from the devces as pos-
sble wthn the enclosure) as well as changes n
swmmng speed and breathng patterns. Although
ths response quckly abated followng nose cessa-
ton, no habtuaton was observed across multple
exposure events. Data from ndvdual exposure
trals were presented by Kastelen et al. and are
analyzed here. To avod pseudoreplcaton, these
data are nversely weghted by the total number
of trals to approxmate a sngle exposure for the
ndvdual. Based on harbor porpose hearng mea-
surements (Andersen, 1970) and the Kastelen et
al. (1997) data on behavoral reactons, Taylor
et al. (1997) estmated zones of nose nfluence
(audblty, behavoral dsturbance, and hearng
damage) for free-rangng harbor porposes.
Subsequently, Kastelen et al. (2000) exposed
two nave subjects to three dfferent nonpulse
sources and observed generally smlar behavoral
avodance n all condtons. Pooled data for each
subject were scored and reported here; pooled
data for each alarm n the dose-response analyss
were weghted to equate wth a sngle exposure
event for each ndvdual. Kastelen et al. (2001)
later measured smlar behavoral responses of
MarineMammalNoiseExposureCriteria 517
the same two ndvdual harbor porposes to three
dfferent acoustc alarms, but these data were not
ncluded n ths analyss because subjects were no
longer nave to controlled nose exposures.
Kastelen et al. (2005) exposed two addtonal
nave harbor porposes to varous sounds assoc-
ated wth underwater data transmsson systems
(clcks, tones, sweeps, and mpulsve dstance
sensors wth a range of source characterstcs).
They drectly measured source levels of each
sound type and RLs at numerous postons wthn
the expermental pool. Observed behavoral
responses (avodance and changes n swmmng
and respraton patterns) were very smlar to
those durng the prevous Kastelen et al. (1997,
2000, 2001) studes. Pooled data for each nd-
vdual response and source type were analyzed
here n the same manner as we appled to the
Kastelen et al. (2000) measurements. Kastelen
et al. (2006a) exposed yet another nave ndvdual
harbor porpose and reported very smlar fnd-
ngs, whch we ncorporated as a sngle pooled
result, wth all exposures equally weghted.
Pinnipeds in Water/Nonpulses (Cell 12)
The effects of nonpulse exposures on pnn-
peds n water are poorly understood. Studes for
whch enough nformaton was avalable for our
analyss nclude feld exposures of harbor seals to
AHDs (Jacobs & Terhune, 2002) and of translo-
cated dvng northern elephant seals to a research
tomography source (Costa et al., 2003), as well
as responses of captve harbor seals to underwa-
ter data communcaton sources (Kastelen et al.,
2006b). These lmted data (see Table 20) suggest
that exposures between ~90 and 140 dB re: 1 Pa
generally do not appear to nduce strong behav-
oral responses n pnnpeds exposed to nonpulse
sounds n water; no data exst regardng exposures
at hgher levels. The severty scalng for Cell 12 s
gven n Table 21.
It s mportant to note that among these stud-
es of pnnpeds respondng to nonpulse exposures
n water, there are some apparent dfferences n
responses between feld and laboratory condtons.
In contrast to the md-frequency odontocetes, cap-
tve pnnpeds responded more strongly at lower
levels than dd anmals n the feld. Agan, contex-
tual ssues are the lkely cause of ths dfference.
Captve subjects n the Kastelen et al. (2006b)
study were not renforced wth food for remanng
n nose felds, whereas free-rangng subjects may
have been more tolerant of exposures because
of motvaton to return to a safe locaton (Costa
et al., 2003) or to approach enclosures holdng
prey tems (Jacobs & Terhune, 2002).
FieldObservations(Cell12)
Jacobs & Terhune (2002) observed harbor seal
reactons to Armar

dB plus II AHDs (general

source characterstcs gven n the Cell 9 secton
above; source level n ths study was 172 dB re:
1 Pa-m) deployed around aquaculture stes. From
1 to 10 AHDs were deployed around nne dffer-
ent stes. Jacobs & Terhune measured receved
SPLs around the AHDs and measured the behav-
or of seals n the surroundng area. Seals n ths
study were generally unresponsve to sounds from
the AHDs. Durng two specfc events, ndvdu-
als came wthn 43 and 44 m of actve AHDs and
faled to demonstrate any measurable behavoral
response; estmated exposure RLs based on the
measures gven were ~120 to 130 dB re: 1 Pa.
These ndvdual observatons are weghted to rep-
resent a sngle observaton for ths study, scored
(as 0), and reported n Table 21.
Costa et al. (2003) measured receved nose
levels from an ATOC sound source off north-
ern Calforna usng acoustc data loggers placed
on translocated elephant seals. Subjects were
captured on land, transported to sea, nstrumented
wth archval acoustc tags, and released such that
ther transt would lead them near an actve ATOC
source (at 939-m depth; 75-Hz sgnal wth 37.5-
Hz bandwdth; 195 dB re: 1 Pa-m max. source
level, ramped up from 165 dB re: 1 Pa-m over
20 mn) on ther return to a haulout ste. Costa et
al. provded a wde range of detaled quanttatve
measures of ndvdual dvng behavor, responses,
and exposure RLs n well-characterzed contexts;
ths knd of nformaton was deal for the present
purposes. Dve depth and duraton, descent/ascent
velocty, surface nterval, and exposure RL were
recorded from a total of 14 seals. An addtonal
three seals were exposed to the ATOC source
durng translocatons and behavoral observatons
were made, but exposure RLs were unavalable.
Seven control seals were nstrumented smlarly
and released when the ATOC source was not actve.
Receved exposure levels of the ATOC source for
expermental subjects averaged 128 dB re: 1 Pa
(range 118 to 137) n the 60- to 90-Hz band. None
of the nstrumented anmals termnated dves or
radcally altered behavor upon exposure, but
some statstcally sgnfcant changes n dvng
parameters were documented n nne ndvduals.
The behavoral scores assgned here for statst-
cally sgnfcant responses were ether three or four
dependng on the change n dvng behavor durng
exposure relatve to mean values for the same nd-
vduals before and after exposure (< 50% change
scored 3; > 50% change scored 4). Translocated
northern elephant seals exposed to ths partcular
nonpulse source (ATOC) began to demonstrate
518 Southalletal.
subtle behavoral changes at ~120 to 140 dB re: 1
Pa exposure RLs (Table 21).
Several other feld studes (dscussed brefly
below) were consdered but not ncluded n the
behavoral analyses due to lmted nformaton
about source and/or propagaton characterstcs,
ndvdual responses durng and/or n the absence
of exposure, or both. Whle studyng cetaceans,
Rchardson et al. (1990b, 1991) made some
observaton of rnged and bearded seal responses
to playbacks of underwater drllng sounds. Ther
fndngs generally suggested a farly hgh degree
of tolerance by exposed pnnpeds to these sounds.
Ths contrasts to some extent wth the results of
Frost & Lowry (1988) who found some reducton
n rnged seal denstes around slands on whch
drllng was occurrng. Norberg & Ban (1994)
made detaled acoustc measurements of several
arrays of Cascade Appled Scences

AHDs (11.9-
to 14.7-kHz frequency sweeps; 195 dB re: 1 Pa-
m source level; 1-ms pulse produced n 57 to 58
dscrete pulse chrps of 2.3-s total duraton). These
devces were placed on the Chttenden Locks n
Puget Sound, Washngton, n an effort to dssuade
predaton of wld steelhead trout by Calforna
sea lons. Behavoral responses of ndvdual an-
mals, however, were not reported. Norberg (2000)
evaluated the behavoral responses of Calforna
sea lons to Armar

AHDs (10-kHz fundamen-

tal frequency; 195 dB re: 1 Pa-m source level;
short tran of 2.5-ms sgnals repeated every 17
s) ntended to reduce predaton on salmonds n
aquaculture facltes. Behavoral observatons
suggested lmted behavoral deterrence by the
devces (predaton rates were smlar n exper-
mental and control condtons), but measures of
RLs and ndvdual response behavor are absent.
Yurk (2000) also observed pnnpeds exposed to
AHDs n the context of fsheres nteractons. He
determned that actve AHDs were more effec-
tve than a mechancal barrer or altered lghtng
condtons n dssuadng harbor seals from prey-
ng on fsh under brdges. Agan, however, nsuf-
fcent nformaton regardng receved sounds
and ndvdual responses s avalable to consder
these observatons explctly here. Koschnsk
et al. (2003) observed harbor seals durng under-
water playbacks of smulated wnd turbne nose
(maxmum energy between 30 and 800 Hz; spec-
tral densty source levels of 128 dB re: 1 Pa/Hz
at 80 and 160 Hz). Harbor seals were sghted at
greater dstances durng playbacks than durng
control condtons. However, lmted nformaton
on receved exposures and ndvdual behavoral
responses precluded ncluson n our analyss.
Moulton et al. (2003, 2005) studed rnged seals
before and durng the constructon and operaton
of an ol producton faclty. They found lttle or
no avodance of the area around the varous ndus-
tral sources, most of whch emtted nonpulses.
Because of the contnuous exposure to multple
sound sources at varyng dstances, ths study
dd not produce data on dscrete exposures and
responses.
LaboratoryObservations(Cell12)
Kastelen et al. (2006b) exposed nne captve
harbor seals n a ~25 30 m enclosure to non-
pulse sounds used n underwater data commun-
caton systems (smlar to acoustc modems). Test
sgnals were dentcal to those used by Kastelen
et al. (2005) n harbor porpose exposure studes
(frequency modulated tones, sweeps, and bands
of nose wth fundamental frequences between
8 and 16 kHz; 128 to 130 [ 3] dB re: 1 Pa-m
source levels; 1- to 2-s duraton [60-80% duty
cycle]; or 100% duty cycle). They recorded seal
postons and the mean number of ndvdual
surfacng behavors durng control perods (no
exposure), before exposure, and n 15-mn exper-
mental sessons (n = 7 exposures for each sound
type). Background nose and exposure RLs (n
terms of Leq; 32-s total tme) were measured at
numerous postons around the enclosure for
each acoustc source. Acoustc dscomfort was
recognzed based on movement out of areas that
anmals used durng control perods. An acoustc
dscomfort threshold was calculated for the group
of seals for each source type, and for each sound
source ths was ca. 107 dB re: 1 Pa. Seals gener-
ally swam away from each source, avodng t by
~5 m, although they dd not haul out of the water
or change surfacng behavor. Seal reactons dd
not appear to wane over repeated exposure (.e.,
there was no obvous habtuaton), and the colony
of seals generally returned to baselne condtons
followng exposure.
For the behavoral analyss conducted here, the
Kastelen et al. (2006b) results were nterpreted as
follows. Because the behavor of ndvduals wthn
the same pool at the same tme cannot be consd-
ered ndependent, the group of nne harbor seals
was consdered a sngle observaton. Because of
smlarty of sources and exposure condtons and
the close temporal tmng of exposures, we com-
bned observatons across the four sound types and
nclude a sngle observaton wthn each appropr-
ate 10-dB bn. Exposures between ~80 and 107 dB
re: 1 Pa seemed nsuffcent to nduce behavoral
avodance n the colony of seals, but hgher expo-
sures were consdered suffcent. Consequently,
sngle observatons ndcatng no response (0)
appear n the 80 to 90 and n the 90 to 100 dB re:
1 Pa exposure bns, and a sngle observaton
ndcatng avodance behavor (6) s shown n the
100 to 110 dB re: 1 Pa condton (Table 21).
MarineMammalNoiseExposureCriteria 519
Pinnipeds in Air/Nonpulses (Cell 15)
There has been consderable effort to study the
effects of aeral nonpulse sounds on pnnped
behavor, prmarly nvolvng rocket launches, ar-
craft overflghts, power-boat approaches, and con-
structon nose. Unfortunately, many of the studes
are dffcult to nterpret n terms of exposure RL
and ndvdual or group behavoral responses. In
many cases, t was dffcult or mpossble to ds-
cern whether the reported behavoral response was
nduced by the nose from a specfc operaton or
some correlated varable such as ts vsual presence.
For these reasons, most of the observatonal studes
of behavoral dsturbance are not approprate for
quanttatve analyses relatng exposure level and
scored behavoral response. However, a number
of the techncal reports and analyses of rocket
launches are relevant for ths cell and contan suf-
fcently detaled nformaton regardng estmated
RLs. These observatons are complcated, how-
ever, by the fact that all studes were conducted n
the same general area wth subjects lkely habtu-
ated to the presence of launch nose. Further, n
many cases, exposures contaned both a nonpulse
component and a pulse component (descrbed
below). Only those observatons for whch there
was clearly just nonpulse exposure were con-
sdered n the severty scalng analyss (Thorson
et al., 1999, 2000b; Berg et al., 2002).
The lmtatons of these and other potentally
applcable studes resulted n a very lmted data
set for use n ths analyss (see summary n Table
22 and severty scalng analyss n Table 23). As a
general statement from the avalable nformaton,
pnnpeds exposed to ntense (~110 to 120 dB re:
20 Pa) nonpulse sounds often leave haulout areas
and seek refuge temporarly (mnutes to a few
hours) n the water. In contrast, pnnpeds exposed
to dstant launches at RLs ~60 to 70 dB re: 20 Pa
tend to gnore the nose. It s dffcult to assess the
relevance of ether of these observatons to nave
ndvduals, however, gven the repeated exposure
of colones studed to such nose events. Also,
there are strong speces dfferences, wth harbor
seals beng much more responsve than northern
elephant seals (e.g., Holst et al., 2005a, 2005b).
Due to the lmtatons of the avalable data, t s
not currently possble to make any further general
characterzatons regardng ths condton.
A seres of hghly detaled, quanttatve analy-
ses on the behavor of pnnpeds exposed to the
sounds of varous large mssle launches were
revewed. These sources generally produce sus-
taned, generally low-frequency (lttle energy
above 1,000 Hz) rumblng sounds lastng tens
of seconds (nonpulse) assocated wth launch
boosters, as well as a sonc boom (pulse) n flght
as the rocket goes supersonc. Extensve research
has been conducted on the effects of both sound
types on pnnpeds. Nonpulse exposures are
consdered n ths secton, whereas behavoral
responses to the pulse component of some of the
same launches are consdered n Appendx B.
Because many measurements were made on the
same few colones of pnnpeds that were exposed
to multple launches, t s lkely that some of the
same ndvduals were resampled. Therefore, we
weghted the combned results across studes for
each speces and breedng locaton nto a sngle
observaton for the behavoral analyss here. That
s, we consdered each speces n an ndvdual
breedng colony a sngle unt of observaton
across studes. The results were pooled accord-
ngly n Table 22, but the studes are dscussed
longtudnally below. The studes dscussed below
reported exposure condtons on or near pnnped
breedng rookeres durng launches of dfferent
types of rockets usng a varety of metrcs, nclud-
ng A-weghted values and a frequency-weghtng
functon derved from the harbor seal audogram;
we used unweghted SPL values for the analyss
here.
Thorson et al. (1998) measured harbor seal
responses and conducted AEP measurements on
seals exposed to a Ttan IV A-18 launch from
Vandenberg Ar Force Base (VAFB), Calforna.
They studed colones both on the manland at VAFB
and on nearby Santa Cruz Island. Unfortunately,
the launch occurred at nght and durng a perod
of relatvely hgh tde, lmtng both the number of
seals present on the rookeres and the observaton
of ndvduals. However, behavoral montorng
over several days after the launch dd not ndcate
any abandonment of the breedng rookeres at
ether ste. Hearng measures (AEP) on ndvduals
tested before and several hours after the launch dd
not ndcate any loss of senstvty.
Thorson et al. (1999) conducted smlar observa-
tons of harbor seals at VAFB and also observed
northern elephant seals, Calforna sea lons, and
northern fur seals at nearby San Mguel Island.
Followng the launch (of an Athena 2 IKONOS-1
mssle), 33 harbor seals (ncludng sx pups) at the
VAFB rookery entered the water. They began to
return to the beach begnnng 16 mn after the launch,
and no pups were observed to have ded as a result of
the event. Ths behavor was consdered to represent
both mnor avodance and a bref/mnor potental
or actual separaton of females and dependent off-
sprng (scored 6 here). The maxmum unweghted
SPL value was 119 dB re: 20 Pa. Indvduals
of the three pnnped speces montored on
San Mguel Island reacted smlarly. However, ther
responses were to the sonc boom generated by the
rocket once arborne rather than to the nonpulse
520 Southalletal.
sounds assocated wth the launch per se, and thus
are not scored here.
Thorson et al. (2000a) conducted observatons
of harbor seal abundance, dstrbuton, and haulout
patterns at VAFB for several days before and after
the launch of a Ttan II G-13 mssle from VAFB.
Ths launch occurred durng the mddle of the nght,
precludng drect observaton of seal reactons (and
behavoral scorng here), although observatons
on subsequent days ndcated generally nomnal
harbor seal presence and dstrbuton n the area.
Thorson et al. (2000b) measured behavoral and
audtory responses of harbor seals at VAFB and
behavoral responses of northern elephant seals
and Calforna sea lons on San Mguel Island to
the launch of a Ttan IV B-28 mssle from VAFB.
They observed all 54 harbor seals at the VAFB ste
movng from the breedng rookery nto the water
wthn 2 mn of the onset of the launch (47 entered
the water mmedately). The maxmum unweghted
SPL value near the rookery was 116 dB re: 20 Pa;
ths exposure was consdered here to be consstent
wth a behavoral score of 6 for ths group of seals.
The sound perssted for several mnutes, and the
unweghted SEL value was 127 dB re: (20 Pa)
2
-
s. There was no dfference n the hearng capabl-
tes of three young seals tested usng AEP meth-
ods before and after the mssle launch. Nether
the Calforna sea lons nor elephant seals on San
Mguel Island were observed to respond at all to
the fant nose assocated wth the launch, cor-
respondng to a severty scalng score of 0 (Table
23). These sounds were from the launch boosters
(nonpulses) rather than sonc booms and were est-
mated here as ~60 to 70 dB re: 20 Pa based on the
measurements and descrptons gven.
Berg et al. (2001) obtaned smlar measure-
ments of behavoral responses of harbor seals at
VAFB and Calforna sea lons and northern ele-
phant seals at San Mguel Island to a Delta II EO-1
mssle launch from VAFB. Observatons were also
made of southern sea otter (Enhydralutrisnereis)
and Calforna brown pelcan (Pelecanusocciden-
taliscalifornicus) responses. No harbor seals were
hauled out on the VAFB rookery durng ths launch.
Berg et al. note that subsequent harbor seal abun-
dance and dstrbuton n the days after the launch
were wthn normal varablty, and there appeared
to be no lastng behavoral reactons. Elephant seals
and Calforna sea lons at San Mguel Island dd
not notceably respond to sounds assocated wth
the launch, whch n ths case were predomnantly
the sonc boom (pulse) component.
Berg et al. (2002) measured behavoral responses
of harbor seals on VAFB rookeres to the launch
of a Ttan IV B-34 mssle from a launch pad at
VAFB ~8.6 km away. At the tme of the launch, 38
seals were present at two haulout stes, all of whch
entered the water mmedately followng the onset
of launch nose. More seals (n = 56) were present
at the locatons 90 mn after the launch event, nd-
catng the temporary and mnor nature of the ds-
turbance, and no njured anmals were located. The
avodance behavor was concdent wth a max-
mum unweghted SPL value near the rookeres of
119 dB re: 20 Pa (unweghted aeral SEL value
was 130 dB re: [20 Pa]
2
-s).
Fnally, Berg et al. (2004) observed behavoral
responses of Calforna sea lons, northern elephant
seals, and northern fur seals on San Mguel Island
to the launch of an Atlas IIAS MLV-14 mssle
from VAFB. Receved sgnals were sonc booms
whch had lttle to no effect on the behavor of the
pnnpeds, other than mnor orentng behavors
and movements n some of the Calforna sea lons.
These results are not scored here, n part because
the sounds ncluded pulses.
Other researchers have nvestgated the effects
of other knds of human actvtes (e.g., arcraft,
motorboats, general human presence) as well as
rocket launches on the haulout behavor, ncludng
avodance, of pnnpeds (Allen et al., 1984; Suryan
& Harvey, 1998; Born et al., 1999; Moulton et al.,
2002). The combned results ndcated that hauled-
out pnnpeds n certan condtons can be dsturbed,
sgnfcantly n some cases, by the presence of var-
ous human actvtes. However, these studes lack
ether specfc estmates of receved nose expo-
sure condtons or ndvdual-specfc behavoral
responses or both. Addtonally, multple stmul
were generally smultaneously present, ncludng
the vsual presence of sources, whch preclude ther
ncluson here. Gentry et al. (1990) determned that
northern fur seals were generally tolerant of under-
ground explosons and other quarryng operatons
n relatvely close proxmty; only a few orentng
behavors were observed n response to the largest
blasts. Some acoustc measurements were made,
but ndvdual behavors or group responses and
receved exposure levels were not reported and were
thus not scored here.
Holst et al. (2005a, 2005b) observed behavoral
responses n three speces of pnnpedsharbor seal,
Calforna sea lon, and northern elephant sealon
San Ncolas Island to 47 small- and md-szed ms-
sle launches over a 4-y perod. They observed
anmal presence and dstrbuton before launches
and behavor durng and followng launches. Some
of the mssles generated sonc booms, but the
majorty of the exposures were relatvely low-fre-
quency, long-duraton rumblng sounds that would
be categorzed as nonpulses. Durng many launches,
acoustc measurements were made near the anmals
whose behavor was vdeotaped. Peak, SPL, and SEL
exposures were reported. Ths dataset has not been
ncorporated nto the present analyss. However,
MarineMammalNoiseExposureCriteria 521
results indicated that California sea lions had mixed
reactions to rocket launches, with some individuals
exhibiting startle responses and increased vigilance
and others showing virtually no reaction. Northern
elephant seal reactions were minimal, consisting
only of minor movements and orienting responses
that rapidly subsided. Conversely, harbor seals were
by far the most responsive of the pinnipeds observed,
with many individuals entering the water from
haulout sites following rocket launches and failing
to return for periods of hours. No cases of long-
term pup separation or of injury were documented.
If those phenomena had occurred, they would be
considered relatively severe in terms of the behav-
ioral scoring paradigm given here and should also be
considered as they relate to injury criteria. In
California sea lions and northern elephant seals,
there were significant correlations between behav-
ioral responses and both the missiles closest dis-
tance and the RL of the launch sound near the
pinnipeds (SEL). Corresponding relationships for
harbor seals were weaker. Holst et al. (2005b) con-
cluded that the temporary behavioral responses,
even the relatively severe ones observed in harbor
seals, do not appear to have substantial adverse
effects on pinniped populations. This conclusion
is based on the decades-long occurrence of missile
launches and the presence of increasing numbers
of pinnipeds of all three species in the area.
(LefttoRight): David Kastak, Roger Gentry, Peter Tyack, Brandon Southall, Darlene Ketten, W. John Richardson, Jeanette
Thomas, James Miller, Ann Bowles, James Finneran, Charles Greene, Jr., Paul Nachtigall, and William Ellison
Thismanuscriptisrespectfullydedicatedtoourco-author,DavidKastak.
Davewasabrilliantscientist,butevenmoreimportantly,hewasamanwith
greatsincerity,integrity,andasharpwit.Hewasaninspirationandmentortomany,
andhissignificant,incisiveresearchonmarinemammalcognitionandsensory
systemsoverthepasttwodecadesprovidedadvancesthatshapedandwill
continuetoguidethefutureofthesefields.Davewasavaluedcolleagueand
treasuredfriendtoallofus.Hewillbemissedbutneverforgotten.
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