Sound use, sequential behavior and ecology of foraging bottlenose dolphins, Tursiops truncatus

Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy at the Massachusetts Institute of Technology and the Woods Hole Oceanographic Institution September 1999Odontocetes are assumed to use echolocation for navigation and foraging, but neither of these uses of biosonar has been conclusively demonstrated in free-ranging animals. Many bats are known to use echolocation throughout foraging sequences, changing the structure and timing of clicks as they progress towards prey capture. For odontocetes, however, we do not know enough about their foraging behavior to describe such sequences. To conduct detailed behavioral observations of any subject animal, the observer must be able to maintain continuous visual contact with the subject for a period commensurate with the duration of the behavior(s) of interest. Behavioral studies of cetaceans, which spend approximately 95% of their time below the water's surface, have been limited to sampling surface behavior except in special circumstances, e.g. clear-water environments, or with the use of technological tools. I addressed this limitation through development of an observation platform consisting of a remote controlled video camera suspended from a tethered airship with boat-based monitoring, adjustment, and recording of video. The system was used successfully to conduct continuous behavioral observations of bottlenose dolphins in the Sarasota Bay, FL area. This system allowed me to describe previously unreported foraging behaviors and elucidate functions for behaviors already defined but poorly understood. Dolphin foraging was modeled as a stage-structured sequence of behaviors, with the goal-directed feeding event occurring at the end of a series of search, encounter, and pursuit behaviors. The behaviors preceding a feeding event do not occur in a deterministic sequence, but are adaptive and plastic. A single-step transition analysis beginning with prey capture and receding in time has identified significant links between observed behaviors and demonstrated the stage-structured nature of dolphin foraging. Factors affecting the occurrence of specific behaviors and behavioral transitions include mesoscale habitat variation and individual preferences. The role of sound in foraging, especially echolocation, is less well understood than the behavioral component. Recent studies have explored the use of echolocation in captive odontocete foraging and presumed feeding in wild animals, but simultaneous, detailed behavioral and acoustic observations have eluded researchers. The current study used two methods to obtain acoustic data. The overhead video system includes two towed hydrophones used to record 'ambient' sounds of dolphin foraging. The recordings are of the 'ambient' sounds because the source of the sounds, i.e. animal, could not be localized. Many focal follows, however, were conducted with single animals, and from these records the timing of echolocation and other sounds relative to the foraging sequence could be examined. The 'ambient' recordings revealed that single animals are much more vocal than animals in groups, both overall and during foraging. When not foraging, single animals vocalized at a rate similar to the per animal rate in groups of ≥2 animals. For single foraging animals, the use of different sound types varies significantly by the habitat in which the animal is foraging. These patterns of use coupled with the characteristics of the different sound types suggest specific functions for each. The presence of multiple animals in a foraging group apparently reduces the need to vocalize, and potential reasons for this pattern are discussed. In addition, the increased vocal activity of single foraging animals lends support to specific hypotheses of sound use in bottlenose dolphins and odontocetes in general. The second acoustic data collection method records sounds known to be from a specific animal. An acoustic recording tag was developed that records all sounds produced by an animal including every echolocation click. The tag also includes an acoustic sampling interval controller and a sensor suite that measures pitch, roll, heading, and surfacing events. While no foraging events occurred while an animal was wearing an acoustic data logger, the rates of echolocation and whistling during different activities, e.g. traveling, were measured.This work was supported by the Education Office of the Woods Hole Oceanographic Institution, two grants from the Rinehart Coastal Research Center, the Ocean Ventures Fund; WHOI Sea Grant, ONR Grant #N00014-94-1-0692 to P. Tyack, and a Graduate Fellowship from the Office of Naval Research

Novel Bio-Logging Tool for Studying Fine-Scale Behaviors of Marine Turtles in Response to Sound

Increases in the spatial scale and intensity of activities that produce marine anthropogenic sound highlight the importance of understanding the impacts and effects of sound on threatened species such as marine turtles. Marine turtles detect and behaviorally respond to low-frequency sounds, however few studies have directly examined their behavioral responses to specific types or intensities of anthropogenic or natural sounds. Recent advances in the development of bio-logging tools, which combine acoustic and fine-scale movement measurements, have allowed for evaluations of animal responses to sound. Here, we describe these tools and present a case study demonstrating the potential application of a newly developed technology (ROTAG, Loggerhead Instruments, Inc.) to examine behavioral responses of freely swimming marine turtles to sound. The ROTAG incorporates a three-axis accelerometer, gyroscope, and magnetometer to record the turtle\u27s pitch, roll, and heading; a pressure sensor to record turtle depth; a hydrophone to record the turtle\u27s received underwater acoustic sound field; a temperature gauge; and two VHF radio telemetry transmitters and antennas for tag and turtle tracking. Tags can be programmed to automatically release via a timed corrodible link several hours or days after deployment. We describe an example of the data collected with these tags and present a case study of a successful ROTAG deployment on a juvenile green turtle (Chelonia mydas) in the Paranaguá Estuary Complex, Brazil. The tag was deployed for 221 min, during which several vessels passed closely (\u3c2 km) by the turtle. The concurrent movement and acoustic data collected by the ROTAG were examined during these times to determine if the turtle responded to these anthropogenic sound sources. While fine-scale behavioral responses were not apparent (second-by-second), the turtle did appear to perform dives during which it remained still on or near the sea floor during several of the vessel passes. This case study provides proof of concept that ROTAGs can successfully be applied to free-ranging marine turtles to examine their behavioral response to sound. Finally, we discuss the broad applications that these tools have to study the fine-scale behaviors of marine turtles and highlight their use to aid in marine turtle conservation and management

Humpback Whale Song and Foraging Behavior on an Antarctic Feeding Ground

The article of record as published may be located at http://dx.doi.org/10.1371/journal.pone.0051214Reports of humpback whale (Megaptera novaeangliae) song chorusing occuring outside the breeding grounds are becoming more common, but song structure and underwater behavior of individual singers on feeding grounds and migration routes remain unknown. Here, ten humpback whales in the Western Antarctic Peninsula were tagged in May 2010 with non-invasive, suction-cup attached tags to study foraging ecology and acoustic behavior. Background song was identified on all ten records, but additionally, acoustic records of two whales showed intense and continuous singing, with a level of organization and structure approaching that of typical breeding ground song. The songs, produced either by the tagged animalsor close associates, shared phrase types and theme structure with one another, and some song bouts lasted close to an hour. Dive behavior of tagged animals during the time of sound production showed song occurring during periods of active diving, sometimes to depths greater than 100 m. One tag record also contained song in the presence of feeding lunges identified from the behavioral sensors, indicating that mating displays occur in areas worthy of foraging. These data show behavioral flexibility as the humpbacks manage competing needs to continue to feed and prepare for the breeding season during late fall. This may also signify an ability to engage in breeding activities outside of the traditional, warm water breeding ground locations.This material is based upon work supported by the National Science Foundation under Grant No. ANT-07-39483. The authors also greatefully acknowledge funding support from the F.V. Hunt Fellowship of the Acoustical Society of America

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

Swimming kinematics and efficiency of entangled North Atlantic right whales

© The Author(s), 2017. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Endangered Species Research 32 (2017): 1-17, doi:10.3354/esr00781.Marine mammals are streamlined for efficient movement in their relatively viscous fluid environment and are able to alter their kinematics (i.e. fluke stroke frequency, amplitude, or both) in response to changes in force balance. Entanglement in fishing gear adds significant drag and buoyant forces that can impact swimming behaviors across a range of timescales. We deployed biologging tags during the disentanglement of 2 North Atlantic right whales Eubalaena glacialis to (1) examine how their kinematics changed in response to drag and buoyancy from entanglement in fishing gear, and (2) calculate resultant changes in swimming efficiency for one individual. We observed variable responses in dive behavior, but neither whale appeared to exploit added buoyancy to reduce energy expenditure. While some of the observed changes in behavior were individually specific, some swimming kinematics were consistently modulated in response to high drag and buoyancy associated with entangling gear, affecting thrust production. In high drag and buoyancy conditions, fluke strokes were significantly shorter and more variable in shape, and gliding was less frequent. Thrust and efficiency significantly differed among dive phases. Disentanglement reduced thrust coefficients ~4-fold, leading to 1.2 to 1.8-fold lower power (W). Ideal propulsive efficiency was significantly lower when entangled, though we detected no difference in observed propulsive efficiency between the conditions. Similar to carrying heavy objects or changing shoes, we present another condition where animals perceive unique movement constraints over seconds to minutes and develop compensatory strategies, altering their movement accordingly.J.M.v.d.H was supported by a postgraduate scholarship from the Natural Sciences and Engineering Research Council of Canada, the MIT Martin Family for Sustainability Fellowship, the Herrington Fitch Family Foundation, a NOAA Award #NA14OAR4320158 to The Cooperative Institute for the North Atlantic Region, and a WHOI-Duke Fellowship through the WHOI Marine Mammal Center

Hidden Markov models reveal complexity in the diving behaviour of short-finned pilot whales

This work was supported by award RC-2154 from the Strategic Environmental Research and Development Program and funding from the Naval Facilities Engineering Command Atlantic and NOAA Fisheries, Southeast Region. DS was supported by the United States Office of Naval Research grant N00014-12-1-0204, under the project entitled Multi-study Ocean acoustics Human effects Analysis (MOCHA).Diving behaviour of short-finned pilot whales is often described by two states; deep foraging and shallow, non-foraging dives. However, this simple classification system ignores much of the variation that occurs during subsurface periods. We used multi-state hidden Markov models (HMM) to characterize states of diving behaviour and the transitions between states in short-finned pilot whales. We used three parameters (number of buzzes, maximum dive depth and duration) measured in 259 dives by digital acoustic recording tags (DTAGs) deployed on 20 individual whales off Cape Hatteras, North Carolina, USA. The HMM identified a four-state model as the best descriptor of diving behaviour. The state-dependent distributions for the diving parameters showed variation between states, indicative of different diving behaviours. Transition probabilities were considerably higher for state persistence than state switching, indicating that dive types occurred in bouts. Our results indicate that subsurface behaviour in short-finned pilot whales is more complex than a simple dichotomy of deep and shallow diving states, and labelling all subsurface behaviour as deep dives or shallow dives discounts a significant amount of important variation. We discuss potential drivers of these patterns, including variation in foraging success, prey availability and selection, bathymetry, physiological constraints and socially mediated behaviour.Publisher PDFPeer reviewe

Dissipation processes in the Tongue of the Ocean

Author Posting. © American Geophysical Union, 2016. This article is posted here by permission of American Geophysical Union for personal use, not for redistribution. The definitive version was published in Journal of Geophysical Research: Oceans 121 (2016): 3159–3170, doi:10.1002/2015JC011165.The Tongue of the Ocean (TOTO) region located within the Bahamas archipelago is a relatively understudied region in terms of both its biological and physical oceanographic characteristics. A prey-field mapping cruise took place in the fall between 15 September 2008 and 1 October 2008, consisting of a series of transects and “clovers” to study the spatial and temporal variability. The region is characterized by a deep scattering layer (DSL), which is preyed on by nekton that serves as the food for beaked whale and other whale species. This study marks the first of its kind where concurrent measurements of acoustic backscatter and turbulence have been conducted for a nekton scattering layer well below the euphotic zone. Turbulence data collected from a Deep Microstructure Profiler are compared to biological and shear data collected by a 38 kHz Simrad EK 60 echo sounder and a hydrographic Doppler sonar system, respectively. From these measurements, the primary processes responsible for the turbulent production in the TOTO region are assessed. The DSL around 500 m and a surface scattering layer (SSL) are investigated for raised ε values. Strong correlation between turbulence levels and scattering intensity of prey is generally found in the SSL with dissipation levels as large as ∼10−7 W kg−1, 3 orders of magnitude above background levels. In the DSL and during the diel vertical migration, dissipation levels ∼10−8 W kg−1 were observed.U.S. Office of Naval Research Grant Number: N00014-08-1-1162-0

Does the marine biosphere mix the ocean?

Author Posting. © Sears Foundation for Marine Research, 2006. This article is posted here by permission of Sears Foundation for Marine Research for personal use, not for redistribution. The definitive version was published in Journal of Marine Research 64 (2006): 541-561, doi:10.1357/002224006778715720.Ocean mixing is thought to control the climatically important oceanic overturning circulation. Here we argue the marine biosphere, by a mechanism like the bioturbation occurring in marine sediments, mixes the oceans as effectively as the winds and tides. This statement is derived ultimately from an estimated 62.7 TeraWatts of chemical power provided to the marine environment in net primary production. Various approaches argue something like 1% (.63 TeraWatts) of this power is invested in aphotic ocean mechanical energy, a rate comparable to wind and tidal inputs

North Atlantic right whales (Eubalaena glacialis) ignore ships but respond to alerting stimuli

Author Posting. © Royal Society, 2004. This article is posted here by permission of Royal Society for personal use, not for redistribution. The definitive version was published in Proceedings of the Royal Society of London B 271 (2004): 227-231, doi:10.1098/rspb.2003.2570.North Atlantic right whales were extensively hunted during the whaling era and have not recovered. One of the primary factors inhibiting their recovery is anthropogenic mortality caused by ship strikes. To assess risk factors involved in ship strikes, we used a multi-sensor acoustic recording tag to measure the responses of whales to passing ships and experimentally tested their responses to controlled sound exposures, which included recordings of ship noise, the social sounds of conspecifics and a signal designed to alert the whales. The whales reacted strongly to the alert signal, they reacted mildly to the social sounds of conspecifics, but they showed no such responses to the sounds of approaching vessels as well as actual vessels. Whales responded to the alert by swimming strongly to the surface, a response likely to increase rather than decrease the risk of collision.Funding for this work was provided by the Fisheries Service of the US National Oceanic and Atmospheric Administration (contract no. NA87RJ0445), and was conducted under NOAA Fisheries permit to conduct scientific research no. 1014 issued to Dr Scott Kraus and Canadian Department of Fisheries and Oceans permits 2001-559 and 2002-568

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