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      When the noise goes on: received sound energy predicts sperm whale responses to both intermittent and continuous navy sonar

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          ABSTRACT

          Anthropogenic noise sources range from intermittent to continuous, with seismic and navy sonar technology moving towards near-continuous transmissions. Continuous active sonar (CAS) may be used at a lower amplitude than traditional pulsed active sonar (PAS), but potentially with greater cumulative sound energy. We conducted at-sea experiments to contrast the effects of navy PAS versus CAS on sperm whale behaviour using animal-attached sound- and movement-recording tags ( n=16 individuals) in Norway. Changes in foraging effort and proxies for foraging success and cost during sonar and control exposures were assessed while accounting for baseline variation [individual effects, time of day, bathymetry and blackfish (pilot/killer whale) presence] in generalized additive mixed models (GAMMs). We found no reduction in time spent foraging during exposures to medium-level PAS (MPAS) transmitted at the same peak amplitude as CAS. In contrast, we found similar reductions in foraging during CAS (d.f.=1, F=8.0, P=0.005) and higher amplitude PAS (d.f.=1, F=20.8, P<0.001) when received at similar energy levels integrated over signal duration. These results provide clear support for sound energy over amplitude as the response driver. We discuss the importance of exposure context and the need to measure cumulative sound energy to account for intermittent versus more continuous sources in noise impact assessments.

          Abstract

          Summary: Experimental exposure of sperm whales to continuous and pulsed sonar reveals that total pulse energy predicts cessation of foraging better than pulse amplitude or duty cycle.

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          Most cited references30

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          A digital acoustic recording tag for measuring the response of wild marine mammals to sound

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            A synthesis of two decades of research documenting the effects of noise on wildlife

            Global increases in environmental noise levels - arising from expansion of human populations, transportation networks, and resource extraction - have catalysed a recent surge of research into the effects of noise on wildlife. Synthesising a coherent understanding of the biological consequences of noise from this literature is challenging. Taxonomic groups vary in auditory capabilities. A wide range of noise sources and exposure levels occur, and many kinds of biological responses have been observed, ranging from individual behaviours to changes in ecological communities. Also, noise is one of several environmental effects generated by human activities, so researchers must contend with potentially confounding explanations for biological responses. Nonetheless, it is clear that noise presents diverse threats to species and ecosystems and salient patterns are emerging to help inform future natural resource-management decisions. We conducted a systematic and standardised review of the scientific literature published from 1990 to 2013 on the effects of anthropogenic noise on wildlife, including both terrestrial and aquatic studies. Research to date has concentrated predominantly on European and North American species that rely on vocal communication, with approximately two-thirds of the data set focussing on songbirds and marine mammals. The majority of studies documented effects from noise, including altered vocal behaviour to mitigate masking, reduced abundance in noisy habitats, changes in vigilance and foraging behaviour, and impacts on individual fitness and the structure of ecological communities. This literature survey shows that terrestrial wildlife responses begin at noise levels of approximately 40 dBA, and 20% of papers documented impacts below 50 dBA. Our analysis highlights the utility of existing scientific information concerning the effects of anthropogenic noise on wildlife for predicting potential outcomes of noise exposure and implementing meaningful mitigation measures. Future research directions that would support more comprehensive predictions regarding the magnitude and severity of noise impacts include: broadening taxonomic and geographical scope, exploring interacting stressors, conducting larger-scale studies, testing mitigation approaches, standardising reporting of acoustic metrics, and assessing the biological response to noise-source removal or mitigation. The broad volume of existing information concerning the effects of anthropogenic noise on wildlife offers a valuable resource to assist scientists, industry, and natural-resource managers in predicting potential outcomes of noise exposure.
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              Extreme diving of beaked whales.

              Sound-and-orientation recording tags (DTAGs) were used to study 10 beaked whales of two poorly known species, Ziphius cavirostris (Zc) and Mesoplodon densirostris (Md). Acoustic behaviour in the deep foraging dives performed by both species (Zc: 28 dives by seven individuals; Md: 16 dives by three individuals) shows that they hunt by echolocation in deep water between 222 and 1885 m, attempting to capture about 30 prey/dive. This food source is so deep that the average foraging dives were deeper (Zc: 1070 m; Md: 835 m) and longer (Zc: 58 min; Md: 47 min) than reported for any other air-breathing species. A series of shallower dives, containing no indications of foraging, followed most deep foraging dives. The average interval between deep foraging dives was 63 min for Zc and 92 min for Md. This long an interval may be required for beaked whales to recover from an oxygen debt accrued in the deep foraging dives, which last about twice the estimated aerobic dive limit. Recent reports of gas emboli in beaked whales stranded during naval sonar exercises have led to the hypothesis that their deep-diving may make them especially vulnerable to decompression. Using current models of breath-hold diving, we infer that their natural diving behaviour is inconsistent with known problems of acute nitrogen supersaturation and embolism. If the assumptions of these models are correct for beaked whales, then possible decompression problems are more likely to result from an abnormal behavioural response to sonar.
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                Author and article information

                Journal
                J Exp Biol
                J. Exp. Biol
                JEB
                jexbio
                The Journal of Experimental Biology
                The Company of Biologists Ltd
                0022-0949
                1477-9145
                1 April 2020
                6 April 2020
                6 April 2020
                : 223
                : 7
                : jeb219741
                Affiliations
                [1 ]Sea Mammal Research Unit, Scottish Oceans Institute, University of St Andrews , St Andrews, Fife KY16 8LB, UK
                [2 ]Faculty of Life and Environmental Sciences, University of Iceland , Askja, Sturlugata 7, 102 Reykjavik, Iceland
                [3 ]Acoustics and Sonar, Netherlands Organization for Applied Scientific Research (TNO) , PO Box 96864 The Hague, 2509 JG, The Netherlands
                [4 ]Defence Systems Division, Norwegian Defense Research Establishment (FFI) , NO-3191 Horten, Norway
                [5 ]LKARTS-Norway , NO-8290, Skutvik, Norway
                Author notes
                [* ]Author for correspondence ( si66@ 123456st-andrews.ac.uk )
                Author information
                http://orcid.org/0000-0002-2212-2135
                http://orcid.org/0000-0002-9894-2543
                http://orcid.org/0000-0001-9570-7567
                http://orcid.org/0000-0002-4210-8058
                http://orcid.org/0000-0003-2984-8606
                http://orcid.org/0000-0002-2550-1405
                http://orcid.org/0000-0002-7833-302X
                http://orcid.org/0000-0001-8028-9673
                Article
                JEB219741
                10.1242/jeb.219741
                7157582
                32107307
                edbd10fc-5631-4b38-98aa-f00b59155f0c
                © 2020. Published by The Company of Biologists Ltd

                This is an Open Access article distributed under the terms of the Creative Commons Attribution License ( https://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution and reproduction in any medium provided that the original work is properly attributed.

                History
                : 4 December 2019
                : 20 February 2020
                Funding
                Funded by: Defence Science and Technology Laboratory, http://dx.doi.org/10.13039/501100008812;
                Funded by: Netherlands Ministry of Defence;
                Funded by: U.S. Navy/Living Marine Resources Program;
                Funded by: Ministère de la Défense, http://dx.doi.org/10.13039/501100005900;
                Funded by: University of St Andrews, http://dx.doi.org/10.13039/501100000740;
                Categories
                Research Article

                Molecular biology
                anthropogenic noise,continuous active sonar,dtag,intermittent sound,time budget,time-series model

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