Diversity in sound pressure levels and estimated active space of resident killer whale vocalizations

Signal source intensity and detection range, which integrates source intensity with propagation loss, background noise and receiver hearing abilities, are important characteristics of communication signals. Apparent source levels were calculated for 819 pulsed calls and 24 whistles produced by free-ranging resident killer whales by triangulating the angles-of-arrival of sounds on two beamforming arrays towed in series. Levels in the 1–20 kHz band ranged from 131 to 168 dB re 1 μPa at 1 m, with differences in the means of different sound classes (whistles: 140.2±4.1 dB; variable calls: 146.6±6.6 dB; stereotyped calls: 152.6±5.9 dB), and among stereotyped call types. Repertoire diversity carried through to estimates of active space, with “long-range” stereotyped calls all containing overlapping, independently-modulated high-frequency components (mean estimated active space of 10–16 km in sea state zero) and “short-range” sounds (5–9 km) included all stereotyped calls without a high-frequency component, whistles, and variable calls. Short-range sounds are reported to be more common during social and resting behaviors, while long-range stereotyped calls predominate in dispersed travel and foraging behaviors. These results suggest that variability in sound pressure levels may reflect diverse social and ecological functions of the acoustic repertoire of killer whales.

[1]  Donna Z. Mirkes,et al.  Advanced concepts in ocean measurements for marine biology , 1980 .

[2]  Christopher W. Clark,et al.  ESTIMATED SOURCE LEVELS OF FIN WHALE (BALAENOPTERA PHYSALUS) VOCALIZATIONS: ADJUSTMENTS FOR SURFACE INTERFERENCE , 2002 .

[3]  Vincent M. Janik,et al.  Animal Communication Networks: Underwater acoustic communication networks in marine mammals , 2005 .

[4]  J. Ford,et al.  Acoustic behaviour of resident killer whales (Orcinus orca) off Vancouver Island, British Columbia , 1989 .

[5]  D. Cato Simple methods of estimating source levels and locations of marine animal sounds. , 1998, The Journal of the Acoustical Society of America.

[6]  W. Watkins,et al.  Intense low-frequency sounds from an antarctic minke whalec Balaenoptera acutorostrata , 1972 .

[7]  Peter L. Tyack,et al.  A small towed beamforming array to identify vocalizing resident killer whales (Orcinus orca) concurrent with focal behavioral observations , 1998 .

[8]  J. Cynx,et al.  Amplitude regulation of vocalizations in noise by a songbird,Taeniopygia guttata , 1998, Animal Behaviour.

[9]  Jack W. Bradbury,et al.  Principles of Animal Communication , 1998 .

[10]  W. Dittus An Analysis of Toque Macaque Cohesion Calls from an Ecological Perspective , 1988 .

[11]  M. Ryan,et al.  Bat Predation and Sexual Advertisement in a Neotropical Anuran , 1982, The American Naturalist.

[12]  William A. Watkins,et al.  Listening to Hawaiian Spinner Porpoises, Stenella cf. Longirostris, with a Three-Dimensional Hydrophone Array , 1974 .

[13]  H. Brumm The impact of environmental noise on song amplitude in a territorial bird , 2004 .

[14]  W A Kuperman,et al.  Matched-field processing, geoacoustic inversion, and source signature recovery of blue whale vocalizations. , 2000, The Journal of the Acoustical Society of America.

[15]  P. O. Thompson,et al.  Underwater sounds of migrating gray whales, Eschrichtius glaucus (Cope). , 1968, The Journal of the Acoustical Society of America.

[16]  D. Todt,et al.  Male–male vocal interactions and the adjustment of song amplitude in a territorial bird , 2004, Animal Behaviour.

[17]  Torben Dabelsteen,et al.  Song and information about aggressive responses of blackbirds, Turdus merula: evidence from interactive playback experiments with territory owners , 1990, Animal Behaviour.

[18]  H. Carl Gerhardt,et al.  Sound pressure levels and radiation patterns of the vocalizations of some North American frogs and toads , 1975, Journal of comparative physiology.

[19]  Whitlow W. L. Au,et al.  The Sonar of Dolphins , 1993, Springer New York.

[20]  William A. Watkins,et al.  Observations of Peponocephala electra, the Melon-headed Whale, in the Southeastern Caribbean , 1997 .

[21]  J. Ford,et al.  Vocal traditions among resident killer whales (Orcinus orca) in coastal waters of British Columbia , 1991 .

[22]  A. Thode,et al.  Frequency beamforming of dolphin whistles using a sparse three-element towed array. , 2000, The Journal of the Acoustical Society of America.

[23]  P. Miller Mixed-directionality of killer whale stereotyped calls: a direction of movement cue? , 2002, Behavioral Ecology and Sociobiology.

[24]  P. Marler,et al.  Food-associated calls in rhesus macaques (Macaca mulatta): I. Socioecological factors , 1993 .

[25]  P. Miller,et al.  Within-pod variation in the sound production of a pod of killer whales, Orcinus orca , 2000, Animal Behaviour.

[26]  L. Miller,et al.  Sperm whale clicks: directionality and source level revisited. , 2000, The Journal of the Acoustical Society of America.

[27]  Alberto Leon-Garcia,et al.  Communication Networks , 2000 .

[28]  C. Mason,et al.  Release from masking due to spatial separation of sources in the identification of nonspeech auditory patterns. , 1998, The Journal of the Acoustical Society of America.

[29]  S. S. Stevens,et al.  The Masking of Pure Tones and of Speech by White Noise , 1950 .

[30]  R. Haven Wiley,et al.  5 – Adaptations for Acoustic Communication in Birds: Sound Transmission and Signal Detection , 1982 .

[31]  Frank Thomsen,et al.  On the communicative significance of whistles in wild killer whales (Orcinus orca) , 2002, Naturwissenschaften.

[32]  P. O. Thompson,et al.  Underwater Sounds from the Blue Whale, Balaenoptera musculus , 1971 .

[33]  Dietmar Todt,et al.  Noise-dependent song amplitude regulation in a territorial songbird , 2002, Animal Behaviour.

[34]  Douglas H. Cato,et al.  Sounds of a pygmy right whale (Caperea marginata) , 1992 .

[35]  V. Janik Source levels and the estimated active space of bottlenose dolphin (Tursiops truncatus) whistles in the Moray Firth, Scotland , 2000, Journal of Comparative Physiology A.

[36]  D B Moody,et al.  Regulation of voice amplitude by the monkey. , 1975, The Journal of the Acoustical Society of America.

[37]  L. H.,et al.  Communication Networks , 1936, Nature.

[38]  Behavioral biology of killer whales , 1986 .

[39]  W C Cummings,et al.  Passive acoustic location of bowhead whales in a population census off Point Barrow, Alaska. , 1985, The Journal of the Acoustical Society of America.

[40]  A. Foote,et al.  Environment: Whale-call response to masking boat noise , 2004, Nature.

[41]  Julia A. Mossbridge,et al.  AN “ACOUSTIC NICHE” FOR ANTARCTIC KILLER WHALE AND LEOPARD SEAL SOUNDS1 , 1999 .

[42]  A. A. Myrberg,et al.  OCEAN NOISE AND THE BEHAVIOR OF MARINE ANIMALS: RELATIONSHIPS AND IMPLICATIONS , 1978 .

[43]  Rex K. Andrew,et al.  Underwater Ambient Noise , 2007 .

[44]  Joseph W. Hall,et al.  Detection in noise by spectro-temporal pattern analysis. , 1984, The Journal of the Acoustical Society of America.

[45]  Jeanette A. Thomas,et al.  An ‘‘acoustic niche’’ for Antarctic killer whale and leopard seal sounds , 2001 .

[46]  P. Marler,et al.  Sound transmission and its significance for animal vocalization , 1977, Behavioral Ecology and Sociobiology.

[47]  M. Littlejohn,et al.  Mating‐Call Sound Intensities of Anuran Amphibians , 1971 .

[48]  C. W. Turl,et al.  Acoustic Source Levels of Four Species of Small Whales , 1976 .

[49]  R. Francois,et al.  Sound absorption based on ocean measurements. Part II: Boric acid contribution and equation for total absorption , 1982 .

[50]  W. Au,et al.  DIRECTIONALITY IN THE WHISTLES OF HAWAIIAN SPINNER DOLPHINS (STENELLA LONGIROSTRIS): A SIGNAL FEATURE TO CUE DIRECTION OF MOVEMENT? , 2003 .

[51]  V B Deecke,et al.  Quantifying complex patterns of bioacoustic variation: use of a neural network to compare killer whale (Orcinus orca) dialects. , 1999, The Journal of the Acoustical Society of America.

[52]  Philipp Goedeking,et al.  Primate Vocal Communication , 1988, Springer Berlin Heidelberg.

[53]  G. Klump,et al.  Comodulation masking release in a songbird , 1995, Hearing Research.

[54]  C. S. Johnson,et al.  Auditory Thresholds of a Killer Whale Orcinus orca Linnaeus , 1972 .

[55]  Robert J. Urick,et al.  Principles of underwater sound , 1975 .

[56]  David E. Bain,et al.  Effects of Masking Noise on Detection Thresholds of Killer Whales , 1994 .

[57]  C. Pytte,et al.  Regulation of vocal amplitude by the blue-throated hummingbird, Lampornis clemenciae , 2003, Animal Behaviour.

[58]  Eliot A. Brenowitz The active space of red-winged blackbird song , 1982, Journal of comparative physiology.

[59]  E Mercado,et al.  Environmental constraints on sound transmission by humpback whales. , 1998, The Journal of the Acoustical Society of America.

[60]  J. Ford,et al.  Characteristics of whistles from the acoustic repertoire of resident killer whales (Orcinus orca) off Vancouver Island, British Columbia. , 2001, The Journal of the Acoustical Society of America.

[61]  D. Todt,et al.  Acoustic communication in noise: regulation of call characteristics in a New World monkey , 2004, Journal of Experimental Biology.

[62]  K. Prestwich,et al.  Energy and Communication in Three Species of Hylid Frogs: Power Input, Power Output and Efficiency , 1989 .

[63]  William A. Watkins,et al.  Sound structure and directionality in Orcinus (killer whale) , 1966, Zoologica : scientific contributions of the New York Zoological Society..

[64]  R. Payne,et al.  Echolocation clicks of two free-ranging, oceanic delphinids with different food preferences: false killer whales Pseudorca crassidens and Risso's dolphins Grampus griseus , 2004, Journal of Experimental Biology.

[65]  D. Kroodsma,et al.  Ecology and evolution of acoustic communication in birds , 1997 .

[66]  J. Ford,et al.  Cultural transmission within maternal lineages: vocal clans in resident killer whales in southern Alaska , 2002, Animal Behaviour.

[67]  H. W. Marsh,et al.  Shallow‐Water Transmission , 1962 .

[68]  William A. Watkins,et al.  The harmonic interval : fact or artifact in spectral analysis of pulse trains , 1968 .

[69]  T. Loughlin Marine mammals and the Exxon Valdez , 1994 .

[70]  P. Marler,et al.  A comparison of vocalizations of red-tailed monkeys and blue monkeys, Ceropithecus ascanius and C. mitis, in Uganda. , 2010, Zeitschrift fur Tierpsychologie.

[71]  C. Erbe UNDERWATER NOISE OF WHALE‐WATCHING BOATS AND POTENTIAL EFFECTS ON KILLER WHALES (ORCINUS ORCA), BASED ON AN ACOUSTIC IMPACT MODEL , 2002 .

[72]  A. Solow,et al.  Call-type matching in vocal exchanges of free-ranging resident killer whales, Orcinus orca , 2004, Animal Behaviour.

[73]  K R Henry,et al.  Killer whale (Orcinus orca) hearing: auditory brainstem response and behavioral audiograms. , 1999, The Journal of the Acoustical Society of America.