Experimental field studies to measure behavioral responses of cetaceans to sonar

Funding was provided by a variety of military and governmental funding sources from several nations acknowledged within referenced publications, notably the US Office of Naval Research, US Navy Living Marine Resources Program, and the navies of the USA, Norway, and the Netherlands. P.L.T. acknowledges the support of the MASTS pooling initiative (The Marine Alliance for Science and Technology for Scotland) in the completion of this study. MASTS is funded by the Scottish Funding Council (grant reference HR09011) and contributing institutions.Substantial recent progress has been made in directly measuring behavioral responses of free-ranging marine mammals to sound using controlled exposure experiments. Many studies were motivated by concerns about observed and potential negative effects of military sonar, including stranding events. Well-established experimental methods and increasingly sophisticated technologies have enabled fine-resolution measurement of many aspects of baseline behavior and responses to sonar. Studies have considered increasingly diverse taxa, but primarily odontocete and mysticete cetaceans that are endangered, particularly sensitive, or frequently exposed to sonar. This review focuses on recent field experiments studying cetacean responses to simulated or actual active military sonars in the 1 to 8 kHz band. Overall results demonstrate that some individuals of different species display clear yet varied responses, some of which have negative implications, while others appear to tolerate relatively high levels, although such exposures may have other consequences not measured. Responses were highly variable and may not be fully predictable with simple acoustic exposure metrics (e.g. received sound level). Rather, differences among species and individuals along with contextual aspects of exposure (e.g. behavioral state) appear to affect response probability. These controlled experiments provide critically needed documentation of identified behavioral responses occurring upon known sonar exposures, and they directly inform regulatory assessments of potential effects. They also inform more targeted opportunistic monitoring of potential responses of animals during sonar operations and have stimulated adaptations of field methods to consider increasingly realistic exposure scenarios and how contextual factors such as behavioral state and source proximity influence response type and probability.Publisher PDFPeer reviewe

Managing human activity and marine mammals: A biologically based, relativistic risk assessment framework

Presented here is a broadly applicable, transparent, repeatable analytical framework for assessing relative risk of anthropogenic disturbances on marine vertebrates, with the emphasis on the sound generating aspects of the activity. The objectives are to provide managers and action-proponents tools with which to objectively evaluate drivers of potential biological risk, to identify data gaps that limit assessment, and to identify actionable measures to reduce risk. Current regulatory assessments of how human activities (particularly those that produce sound) influence the likelihood of marine mammal behavioral responses and potential injury, rely principally on generalized characterizations of exposure and effect using simple, threshold-based criteria. While this is relatively straightforward in regulatory applications, this approach fails to adequately address realistic site and seasonal scenarios, other potential stressors, and scalable outcome probabilities. The risk assessment presented here is primarily based on a common and broad understanding of the spatial-temporal-spectral intersections of animals and anthropogenic activities, and specific examples of its application to hypothetical offshore wind farms are given. The resulting species- and activity-specific framework parses risk into two discrete factors: a population’s innate ‘vulnerability’ (potential degree of susceptibility to disturbance) and an ‘exposure index’ (magnitude-duration severity resulting from exposure to an activity). The classic intersection of these factors and their multi-dimensional components provides a relativistic risk assessment process for realistic evaluation of specified activity contexts, sites, and schedules, convolved with species-specific seasonal presence, behavioral-ecological context, and natural history. This process is inherently scalable, allowing a relativistic means of assessing potential disturbance scenarios, tunable to animal distribution, region, context, and degrees of spatial-temporal-spectral resolution

Species information in whistle frequency modulation patterns of common dolphins

Funding for this project was generously provided by the Office of Naval Research Marine Mammals and Biology program.The most flexible communication systems are those of open-ended vocal learners that can acquire new signals throughout their lifetimes. While acoustic signals carry information in general voice features that affect all of an individual's vocalizations, vocal learners can also introduce novel call types to their repertoires. Delphinids are known for using such learned call types in individual recognition, but their role in other contexts is less clear. We investigated the whistles of two closely related, sympatric common dolphin species, Delphinus delphis and Delphinus bairdii, to evaluate species differences in whistle contours. Acoustic recordings of single-species groups were obtained from the Southern California Bight. We used an unsupervised neural network to categorize whistles and compared the resulting whistle types between species. Of the whistle types recorded in more than one encounter, 169 were shared between species and 60 were species-specific (32 D. delphis types, 28 D. bairdii types). Delphinus delphis used 15 whistle types with an oscillatory frequency contour while only one such type was found in D. bairdii. Given the role of vocal learning in delphinid vocalizations, we argue that these differences in whistle production are probably culturally driven and could help facilitate species recognition between Delphinus species.Publisher PDFPeer reviewe

Mitigation of harm during a novel behavioural response study involving active sonar and wild cetaceans

Some studies of how human activities can affect wild free-ranging animals may be considered to have potential negative outcomes too severe to beethically studied. This creates a societal dilemma involving choices between continuing risky activities with high uncertainty about their potentialeffects on wildlife, often with considerable associated precaution or undertaking focused research to reduce uncertainty, but with some risk of harmfrom either strong response leading to potential stranding or direct physical injury from sound exposure. Recent and ongoing field experimentshave measured the conditions in which wild cetaceans respond to military sonar, and provided insight into the nature of responses. Here mitigationmeasures are reported for one of the first such experiments designed to measure fine-scale behavioural responses to controlled exposures of midfrequency(3–4 kHz) active sonar. The objective was to do so without causing the kinds of physical harm that have been previously observed (e.g.stranding events) and that motivated the study. A critical goal of this experimental study was to identify a response that was safe but that could beused as an indicator of the probability of risk from more extreme or sustained exposure from real military operations. A monitoring and mitigationprotocol was developed using a feedback control procedure for real-time mitigation of potential harm. Experimental protocols were modulatedrelative to indicators of potential risk with the explicit objective of detecting potentially harmful consequences of sound exposure and takingappropriate corrective action. Three categories of mitigation methods were developed and integrated within the experimental protocol incorporatingdesigned, engineered, and operational mitigation measures. Controlled exposure experiments involving free-ranging animals were conducted withoutany evident harm to the experimental subjects, while successfully eliciting behavioural responses that provided meaningful results to informmanagement decisions. This approach demonstrates the importance of careful design of protocols in exposure-response experiments, particularlyin pioneering studies assessing response where both the potential for harm and level of uncertainty may be high.Publisher PDFPeer reviewe

Context-dependent lateralized feeding strategies in blue whales.

Lateralized behaviors benefit individuals by increasing task efficiency in foraging and anti-predator behaviors [1-4]. The conventional lateralization paradigm suggests individuals are left or right lateralized, although the direction of this laterality can vary for different tasks (e.g. foraging or predator inspection/avoidance). By fitting tri-axial movement sensors to blue whales (Balaenoptera musculus), and by recording the direction and size of their rolls during lunge feeding events, we show how these animals differ from such a paradigm. The strength and direction of individuals' lateralization were related to where and how the whales were feeding in the water column. Smaller rolls (≤180°) predominantly occurred at depth (>70 m), with whales being more likely to rotate clockwise around their longest axis (right lateralized). Larger rolls (>180°), conversely, occurred more often at shallower depths (<70 m) and were more likely to be performed anti-clockwise (left lateralized). More acrobatic rolls are typically used to target small, less dense krill patches near the water's surface [5,6], and we posit that the specialization of lateralized feeding strategies may enhance foraging efficiency in environments with heterogeneous prey distributions

Marine seismic surveys and ocean noise : time for coordinated and prudent planning

Marine seismic surveys use intense (eg >= 230 decibel [dB] root mean square [RMS]) sound impulses to explore the ocean bottom for hydrocarbon deposits, conduct geophysical research, and establish resource claims under the United Nations Convention on the Law of the Sea. The expansion of seismic surveys necessitates greater regional and international dialogue, partnerships, and planning to manage potential environmental risks. Data indicate several reasons for concern about the negative impacts of anthropogenic noise on numerous marine species, including habitat displacement, disruption of biologically important behaviors, masking of communication signals, chronic stress, and potential auditory damage. The sound impulses from seismic surveys - spanning temporal and spatial scales broader than those typically considered in environmental assessments - may have acute, cumulative, and chronic effects on marine organisms. Given the international and transboundary nature of noise from marine seismic surveys, we suggest the creation of an international regulatory instrument, potentially an annex to the existing International Convention on the Prevention of Pollution from Ships, to address the issue.Publisher PDFPeer reviewe

Feeding performance by sympatric blue and fin whales exploiting a common prey resource

A major focus in macroecology is to understand the mechanisms that drive differences in habitat, morphology, and behavior in closely related species (Brown 1995). Within this paradigm, energy use and prey acquisition play a fundamental role in affecting behavior, especially with respect to foraging patterns and predator‐prey interactions. Because resources may occur in discrete regions and are often limited, sympatric organisms that feed in similar ways partition resources, when they are limited, both to avoid competition and maximize energy gain (Pianka 1974, Schoener 1983). Baleen whales (Mysticeti) comprise more than a dozen species that have evolved a bulk‐filter feeding strategy that enables the efficient exploitation of low trophic level oceanic resources. Although mysticetes are opportunistic predators that feed on diverse prey types, from schooling fish to small copepods, many sympatric baleen whale species, including blue (Balaenoptera musculus) and fin whales (Balaenoptera physalus), target the similar resources. What remains unknown, however, is whether resources are partitioned when baleen whales and prey overlap in both time and space.This is the publisher’s final pdf. The article is copyrighted by the Society for Marine Mammalogy and published by John Wiley & Sons Ltd. It can be found at: http://onlinelibrary.wiley.com/journal/10.1111/%28ISSN%291748-769

From individual responses to population effects : integrating a decade of multidisciplinary research on blue whales and sonar

Funding: Office of Naval Research (GrantNumber(s): N00014-19-1-2464).As ecosystems transform under climate change and expanding human activities, multidisciplinary integration of empirical research, conceptual frameworks and modelling methods is required to predict, monitor and manage the cascading effects on wildlife populations. For example, exposure to anthropogenic noise can lead to changes in the behaviour and physiology of individual marine mammals, but management is complicated by uncertainties on the long-term effects at a population level. We build on a decade of diverse efforts to demonstrate the strengths of integrating research on multiple stressors for assessing population-level effects. Using the case study of blue whales exposed to military sonar in the eastern north Pacific, we model how behavioural responses and environmental effects induced by climate change affect female survival and reproductive success. Environmental changes were predicted to severely affect vital rates, while the current regime of sonar activities was not. Simulated disturbance had a stronger effect on reproductive success than adult survival, as predicted by life-history theory. We show that information on prey resources is critical for robust predictions, as are data on baseline behavioural patterns, energy budgets, body condition and contextual responses to noise. These results will support effective management of the interactions between sonar operations and blue whales in the study area, while providing pragmatic guidance for future data collection to reduce key uncertainties. Our study provides important lessons for the successful integration of multidisciplinary research to inform the assessment of the effects of noise and other anthropogenic stressors on marine predator populations in the context of a changing environment.Publisher PDFPeer reviewe

Behavioral responses of fin whales to military mid-frequency active sonar

Funding. Primary funding for the SOCAL-BRS project was initially provided by the US Navy’s Chief of Naval Operations Environmental Readiness Division and subsequently by the US Navy’s Living Marine Resources (LMR) Program. Additional support for environmental sampling and logistics was also provided by the Office of Naval Research, Marine Mammal Program.The effect of active sonars on marine mammal behavior is a topic of considerable interest and scientific investigation. Some whales, including the largest species (blue whales, Balaenoptera musculus), can be impacted by mid-frequency (1-10 kHz) military sonars. Here we apply complementary experimental methods to provide the first experimentally controlled measurements of behavioral responses to military sonar and similar stimuli for a related endangered species, fin whales (Balaenoptera physalus). Analytical methods include: (1) Principal Component Analysis paired with Generalized Additive Mixed Models; (2) Hidden Markov Models; and (3) structured expert elicitation using response severity metrics. These approaches provide complementary perspectives on the nature of potential changes within and across individuals. Behavioral changes were detected in five of 15 whales during controlled exposure experiments (CEEs) using mid-frequency active sonar (MFAS) or pseudorandom noise (PRN) of similar frequency, duration, and source and received level. No changes were detected during six control (no noise) sequences. Overall responses were more limited in occurrence, severity, and duration than in blue whales and were less dependent upon contextual aspects of exposure and more contingent upon exposure received level. Quantifying the factors influencing marine mammal responses to sonar is critical in assessing and mitigating future impacts.Publisher PDFPeer reviewe