Mooney T. Aran

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T. Aran

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  • Preprint
    Ambient noise and temporal patterns of boat activity in the US Virgin Islands National Park
    ( 2015-06) Kaplan, Maxwell B. ; Mooney, T. Aran
    Human activity is contributing increasing noise to marine ecosystems. Recent studies have examined the effects of boat noise on marine fishes, but there is limited understanding of the prevalence of this sound source. This investigation tracks vessel noise on three reefs in the US Virgin Islands National Park over four months in 2013. Ambient noise levels ranged from 106-129 dBrms re 1 μPa (100 Hz – 20 kHz). Boat noise occurred in 6-12% of samples. In the presence of boat noise, ambient noise in a low-frequency band (100-1000 Hz) increased by >7 dB above baseline levels and sound levels were significantly higher. The frequency with the most acoustic energy shifted to a significantly lower frequency when boat noise was present during the day. These results indicate the prevalence of boat noise and its overlap with reef organism sound production, raising concern for the communication abilities of these animals.
  • Article
    Auditory sensitivity in aquatic animals
    (Acoustical Society of America, 2016-06-08) Lucke, Klaus ; Popper, Arthur N. ; Hawkins, Anthony D. ; Akamatsu, Tomonari ; Andre, Michel ; Branstetter, Brian K. ; Lammers, Marc O. ; Radford, Craig A. ; Stansbury, Amanda L. ; Mooney, T. Aran
    A critical concern with respect to marine animal acoustics is the issue of hearing “sensitivity,” as it is widely used as a criterion for the onset of noise-induced effects. Important aspects of research on sensitivity to sound by marine animals include: uncertainties regarding how well these species detect and respond to different sounds; the masking effects of man-made sounds on the detection of biologically important sounds; the question how internal state, motivation, context, and previous experience affect their behavioral responses; and the long-term and cumulative effects of sound exposure. If we are to better understand the sensitivity of marine animals to sound we must concentrate research on these questions. In order to assess population level and ecological community impacts new approaches can possibly be adopted from other disciplines and applied to marine fauna.
  • Preprint
    Sonar-induced temporary hearing loss in dolphins
    ( 2009-03-10) Mooney, T. Aran ; Nachtigall, Paul E. ; Vlachos, Stephanie
    There is increasing concern that human-produced ocean noise is adversely affecting marine mammals, as several recent cetacean mass strandings may have been caused by animals’ interactions with naval “mid-frequency” sonar. However, it has yet to be empirically demonstrated how sonar could induce these strandings or cause physiological effects. In controlled experimental studies, we show that mid-frequency sonar can induce temporary hearing loss in a bottlenose dolphin (Tursiops truncatus). Mild behavioural alterations were also associated with the exposures. The auditory effects were only induced by repeated exposures to intense sonar pings with total sound exposure levels of 214 dB re: 1 μPa2∙s. Data support an increasing energy model to predict temporary noise-induced hearing loss and indicate that odontocete noise exposure effects bear trends similar to terrestrial mammals. Thus, sonar can induce physiological and behavioural effects in at least one species of odontocete; however, exposures must be of prolonged, high sound exposures levels to generate these effects.
  • Article
    Soundscapes as heard by invertebrates and fishes: particle motion measurements on coral reefs
    (Acoustical Society of America, 2022-07-14) Jones, Ian T. ; Gray, Michael D. ; Mooney, T. Aran
    Coral reef soundscapes are increasingly studied for their ecological uses by invertebrates and fishes, for monitoring habitat quality, and to investigate effects of anthropogenic noise pollution. Few examinations of aquatic soundscapes have reported particle motion levels and variability, despite their relevance to invertebrates and fishes. In this study, ambient particle acceleration was quantified from orthogonal hydrophone arrays over several months at four coral reef sites, which varied in benthic habitat and fish communities. Time-averaged particle acceleration magnitudes were similar across axes, within 3 dB. Temporal trends of particle acceleration corresponded with those of sound pressure, and the strength of diel trends in both metrics significantly correlated with percent coral cover. Higher magnitude particle accelerations diverged further from pressure values, potentially representing sounds recorded in the near field. Particle acceleration levels were also reported for boat and example fish sounds. Comparisons with particle acceleration derived audiograms suggest the greatest capacity of invertebrates and fishes to detect soundscape components below 100 Hz, and poorer detectability of soundscapes by invertebrates compared to fishes. Based on these results, research foci are discussed for which reporting of particle motion is essential, versus those for which sound pressure may suffice.
  • Book chapter
    Hearing in cetaceans : from natural history to experimental biology
    ( 2012-02) Mooney, T. Aran ; Yamato, Maya ; Branstetter, Brian K.
    Sound is the primary sensory cue for most marine mammals, and this is especially true for cetaceans. To passively and actively acquire information about their environment, cetaceans have perhaps the most derived ears of all mammals, capable of sophisticated, sensitive hearing and auditory processing. These capabilities have developed for survival in an underwater world where sound travels five times faster than in air, and where light is quickly attenuated and often limited at depth, at night, and in murky waters. Cetacean auditory evolution has capitalized on the ubiquity of sound cues and the efficiency of underwater acoustic communication. The sense of hearing is central to cetacean sensory ecology, enabling vital behaviors such as locating prey, detecting predators, identifying conspecifics, and navigating. Increasing levels of anthropogenic ocean noise appears to influence many of these activities. Here we describe the historical progress of investigations on cetacean hearing, with a particular focus on odontocetes and recent advancements. While this broad topic has been studied for several centuries, new technologies in the last two decades have been leveraged to improve our understanding of a wide range of taxa, including some of the most elusive species. This paper addresses topics including how sounds are received, what sounds are detected, hearing mechanisms for complex acoustic scenes, recent anatomy and physiology studies, the potential impacts of noise, and mysticete hearing. We conclude by identifying emerging research topics and areas which require greater focus.
  • Article
    Ontogenetic variation in the hearing sensitivity of black sea bass (Centropristis striata) and the implications of anthropogenic sound on behavior and communication
    (Company of Biologists, 2020-05-27) Stanley, Jenni A. ; Caiger, Paul E. ; Phelan, Beth ; Shelledy, Katharine ; Mooney, T. Aran ; Van Parijs, Sofie M.
    Black sea bass (Centropristis striata) is an important fish species in both commercial and recreational fisheries of southern New England and the mid-Atlantic Bight. Due to the intense urbanization of these waters, this species is subject to a wide range of anthropogenic noise pollution. Concerns that C. striata are negatively affected by pile driving and construction noise predominate in areas earmarked for energy development. However, as yet, the hearing range of C. striata is unknown, making it hard to evaluate potential risks. This study is a first step in understanding the effects of anthropogenic noise on C. striata by determining the auditory bandwidth and thresholds of this species using auditory evoked potentials (AEPs), creating pressure and acceleration audiograms. These physiological tests were conducted on wild-caught C. striata in three size/age categories. Results showed that juvenile C. striata significantly had the lowest thresholds, with hearing sensitivity decreasing in the larger size classes. Furthermore, Centropristis striata has fairly sensitive hearing relative to other related species. Preliminary investigations into the mechanisms of their hearing ability were undertaken with gross dissections and an opportunistic micro computed tomography image to address the auditory structures including otoliths and swimbladder morphology. Crucially, the hearing range of C. striata, and their most sensitive frequencies, directly overlap with high-amplitude anthropogenic noise pollution such as shipping and underwater construction.
  • Article
    Effects of frequency-dependent spatial variation in soundscape settlement cues for reef fish larvae
    (Inter Research, 2022-04-07) Salas, Andria K. ; Ballard, Megan S. ; Mooney, T. Aran ; Wilson, Preston S.
    The mechanisms that link reef soundscapes to larval fish settlement behaviors are poorly understood, yet the management of threatened reef communities requires we maintain the recruitment processes that recover and sustain populations. Using a field-calibrated sound propagation model, we predicted the transmission loss in the relevant frequency band as a function of range, depth, and azimuth to estimate the spatial heterogeneity in the acoustic cuescape. The model highlighted the frequency- and depth-dependence of the sound fields fishes may encounter, and we predict these complex spatial patterns influence how sounds function as settlement cues. Both modeling and field measurements supported a non-monotonic decline in amplitude with distance from the reef. We modeled acoustic fields created by sounds at frequencies from 2 common soniferous reef-based animals (snapping shrimps and toadfish) and estimated detection spaces of these sounds for larvae of 2 reef fish species. Results demonstrated that larval depth will influence cue availability and amplitude, and these spatial patterns of detection depend on cue frequency and the larval receiver’s auditory sensitivity. Estimated spatial scales of detection coupled with field measurements suggest cue amplitudes might allow some larvae to detect reef-based sounds at a range exposing them to the predicted spatial variation in the acoustic cuescape. In an individual-based model, cues available to even the shortest modeled distances improved settlement success. Our results emphasize the need to consider the frequency- and depth-dependence of the acoustic cues larval fishes encounter to increase understanding of the role of soundscapes in larval settlement.
  • Preprint
    Detection of low-frequency tones and whale predator sounds by the American sand lance Ammodytes americanus
    ( 2012-06) Strobel, S. M. ; Mooney, T. Aran
    Auditory evoked potentials (AEPs) were used to measure the hearing range and auditory sensitivity of the American sand lance Ammodytes americanus. Responses to amplitude modulated tone pips indicated that the hearing range extended from 50 to 400 Hz. Sound pressure thresholds were lowest between 200 and 400 Hz. Particle acceleration thresholds showed an improved sensitivity notch at 200 Hz but not substantial differences between frequencies and only a slight improvement in hearing abilities at lower frequencies. The hearing range was similar to Pacific sand lance A. personatus and variations between species may be due to differences in threshold evaluation methods. AEPs were also recorded in response to pulsed sounds simulating humpback whale Megaptera novaeangliae foraging vocalizations termed ‘megapclicks’. Responses were generated with pulses containing significant energy below 400 Hz. No responses were recorded using pulses with peak energy above 400 Hz. These results show that A. americanus can detect the particle motion component of low frequency tones and pulse sounds, including those similar to the low frequency components of megapclicks. Ammodytes americanus hearing may be used to detect environmental cues and the pulsed signals of mysticete predators.
  • Article
    Adverse effects of ocean acidification on early development of squid (Doryteuthis pealeii)
    (Public Library of Science, 2013-05-31) Kaplan, Maxwell B. ; Mooney, T. Aran ; McCorkle, Daniel C. ; Cohen, Anne L.
    Anthropogenic carbon dioxide (CO2) is being absorbed into the ocean, altering seawater chemistry, with potentially negative impacts on a wide range of marine organisms. The early life stages of invertebrates with internal and external aragonite structures may be particularly vulnerable to this ocean acidification. Impacts to cephalopods, which form aragonite cuttlebones and statoliths, are of concern because of the central role they play in many ocean ecosystems and because of their importance to global fisheries. Atlantic longfin squid (Doryteuthis pealeii), an ecologically and economically valuable taxon, were reared from eggs to hatchlings (paralarvae) under ambient and elevated CO2 concentrations in replicated experimental trials. Animals raised under elevated pCO2 demonstrated significant developmental changes including increased time to hatching and shorter mantle lengths, although differences were small. Aragonite statoliths, critical for balance and detecting movement, had significantly reduced surface area and were abnormally shaped with increased porosity and altered crystal structure in elevated pCO2-reared paralarvae. These developmental and physiological effects could alter squid paralarvae behavior and survival in the wild, directly and indirectly impacting marine food webs and commercial fisheries.
  • Article
    Field-based hearing measurements of two seabird species
    (Company of Biologists, 2019-02-18) Mooney, T. Aran ; Smith, Adam B. ; Larsen, Ole Naesbye ; Hansen, Kirstin Anderson ; Wahlberg, Magnus ; Rasmussen, Marianne H.
    Hearing is a primary sensory modality for birds. For seabirds, auditory data is challenging to obtain and hearing data are limited. Here, we present methods to measure seabird hearing in the field, using two Alcid species: the common murre Uria aalge and the Atlantic puffin Fratercula arctica. Tests were conducted in a portable semi-anechoic crate using physiological auditory evoked potential (AEP) methods. The crate and AEP system were easily transportable to northern Iceland field sites, where wild birds were caught, sedated, studied and released. The resulting data demonstrate the feasibility of a field-based application of an established neurophysiology method, acquiring high quality avian hearing data in a relatively quiet setting. Similar field methods could be applied to other seabirds, and other bird species, resulting in reliable hearing data from a large number of individuals with a modest field effort. The results will provide insights into the sound sensitivity of species facing acoustic habitat degradation.
  • Preprint
    Ocean acidification responses in paralarval squid swimming behavior using a novel 3D tracking system
    (Springer Nature, 2017-08-22) Zakroff, Casey ; Mooney, T. Aran ; Wirth, Colin
    Chronic embryonic exposure to ocean acidification (OA) has been shown to degrade the aragonitic statolith of paralarval squid, Doryteuthis pealeii, a key structure for their swimming behavior. This study examined if day-of-hatching paralarval D. pealeii from eggs reared under chronic OA demonstrated measurable impairments to swimming activity and control. This required the development of a novel, cost-effective, and robust method for 3D motion tracking and analysis. Squid eggs were reared in pCO2 levels in a dose-dependent manner ranging from 400 - 2200 ppm. Initial 2D experiments showed paralarvae in higher acidification environments spent more time at depth. In 3D experiments, velocity, particularly positive and negative vertical velocities, significantly decreased from 400 to 1000 ppm pCO2, but showed non-significant decreases at higher concentrations. Activity and horizontal velocity decreased linearly with increasing pCO2, indicating a subtle impact to paralarval energetics. Patterns may have been obscured by notable individual variability in the paralarvae. Responses were also seen to vary between trials on cohort or potentially annual scales. Overall, paralarval swimming appeared resilient to OA, with effects being slight. The newly developed 3D tracking system provides a powerful and accessible method for future studies to explore similar questions in the larvae of aquatic taxa.
  • Article
    Accelerating global ocean observing: monitoring the coastal ocean through broadly accessible, low-cost sensor networks
    (Marine Technology Society, 2021-05-01) Wang, Zhaohui Aleck ; Michel, Anna P. M. ; Mooney, T. Aran
    The global coastal ocean provides food and other critical resources to human societies. Yet this habitat, for which many depend, has experienced severe degradation from human activities. The rates of human-induced changes along the coast demand significantly improved coverage of ocean observations in order to support science-based decision making and policy formation tailored to specific regions. Our proposal envisions developing a global network of low-cost, easily produced and readily deployed oceanographic sensors for use on a wide variety of platforms in the coastal ocean. A substantially large number of these sensors can thus be installed on existing infrastructure, ships of opportunity, and fishing fleets, or even individually along the coast, particularly in vulnerable and disadvantaged regions. This would vastly increase the spatiotemporal resolution of the current data coverage along the coast, allowing greater equitable access. It would also offer significant opportunities for partnership with communities, NGOs, governments, and other stakeholders, as well as a wide range of commercial and industrial sectors to develop and deploy sensors in scalable networks transmitting data in near-real time. Finally, it presents a vastly lowered bar for participation by citizen scientists and other engaged members of the public to address location-specific coastal problems anywhere in the world.
  • Preprint
    Baseline hearing abilities and variability in wild beluga whales (Delphinapterus leucas)
    ( 2014-01) Castellote, Manuel ; Mooney, T. Aran ; Quakenbush, Lori T. ; Hobbs, Roderick ; Goertz, Caroline ; Gaglione, Eric
    While hearing is the primary sensory modality for odontocetes, there are few data addressing variation within a natural population. This work describes the hearing ranges (4-150 kHz) and sensitivities of seven apparently healthy, wild beluga whales (Delphinapterus leucas) during a population health assessment project that captured and released belugas in Bristol Bay, Alaska. The baseline hearing abilities and subsequent variations are addressed. Hearing was measured using auditory evoked potentials (AEPs). All audiograms showed a typical cetacean U-shape; substantial variation (>30 dB) was found between most and least sensitive thresholds. All animals heard well, up to at least 128 kHz. Two heard up to 150 kHz. Lowest auditory thresholds, 35-45 dB, were identified in the range 45-80 kHz. Greatest differences in hearing abilities occurred at both the high end of the auditory range and at frequencies of maximum sensitivity. In general, wild beluga hearing was quite sensitive. Hearing abilities were similar to belugas measured in zoological settings, reinforcing the comparative importance of both settings. The relative degree of variability across the wild belugas suggests that audiograms from multiple individuals are needed to properly describe the maximum sensitivity and population variance for odontocetes. Hearing measures were easily incorporated into field-based settings. This detailed examination of hearing abilities in wild Bristol Bay belugas provides a basis for a better understanding of the potential impact of anthropogenic noise on a noise-sensitive species. Such information may help design noise limiting mitigation measures that could be applied to areas heavily influenced and inhabited by endangered belugas.
  • Article
    Repeated call types in Hawaiian melon-headed whales (Peponocephala electra)
    (Acoustical Society of America, 2014-09) Kaplan, Maxwell B. ; Mooney, T. Aran ; Sayigh, Laela S. ; Baird, Robin W.
    Melon-headed whales are pantropical odontocetes that are often found near oceanic islands. While considered sound-sensitive, their bioacoustic characteristics are relatively poorly studied. The goal of this study was to characterize the vocal repertoire of melon-headed whales to determine whether they produce repeated calls that could assist in recognition of conspecifics. The first tag-based acoustic recordings of three melon-headed whales were analyzed. Tag records were visually and aurally inspected and all calls were individually extracted. Non-overlapping calls with sufficient signal-to-noise were then parameterized and visually grouped into categories of repeated call types. Thirty-six call categories emerged. Categories differed significantly in duration, peak and centroid frequency, and −3 dB bandwidth. Calls of a given type were more likely to follow each other than expected. These data suggest that repeated calls may function in individual, subgroup, or group recognition. Repeated call production could also serve to enhance signal detection in large groups with many individuals producing simultaneous calls. Results suggest that caution should be used in developing automatic classification algorithms for this species based on small sample sizes, as they may be dominated by repeated calls from a few individuals, and thus not representative of species- or population-specific acoustic parameters.
  • Article
    Ground-truthing daily and lunar patterns of coral reef fish call rates on a US Virgin Island reef
    (Inter Research, 2022-07-28) Ferguson, Sophie R. ; Jensen, Frants H. ; Hyer, Matthew D. ; Noble, Allison ; Apprill, Amy ; Mooney, T. Aran
    Coral reefs comprise some of the most biodiverse habitats on the planet. These ecosystems face a range of stressors, making quantifying community assemblages and potential changes vital to effective management. To understand short- and long-term changes in biodiversity and detect early warning signals of decline, new methods for quantifying biodiversity at scale are necessary. Acoustic monitoring techniques have proven useful in observing species activities and biodiversity on coral reefs through aggregate approaches (i.e. energy as a proxy). However, few studies have ground-truthed these acoustic analyses with human-based observations. In this study, we sought to expand these passive acoustic methods by investigating biological sounds and fish call rates on a healthy reef, providing a unique set of human-confirmed, labeled acoustic observations. We analyzed acoustic data from Tektite Reef, St. John, US Virgin Islands, over a 2 mo period. A subset of acoustic files was manually inspected to identify recurring biotic sounds and quantify reef activity throughout the day. We found a high variety of acoustic signals in this soundscape. General patterns of call rates across time conformed to expectations, with dusk and dawn showing important and significantly elevated peaks in soniferous fish activity. The data reflected high variability in call rates across days and lunar phases. Call rates did not correspond to sound pressure levels, suggesting that certain call types may drive crepuscular trends in sound levels while lower-level critical calls, likely key for estimating biodiversity and behavior, may be missed by gross sound level analyses.
  • Preprint
    Potential for sound sensitivity in cephalopods
    ( 2010-07) Mooney, T. Aran ; Hanlon, Roger T. ; Madsen, Peter T. ; Christensen-Dalsgaard, Jakob ; Ketten, Darlene R. ; Nachtigall, Paul E.
    Hearing is a primary sense in many marine animals and we now have a reasonable understanding of what stimuli generate clear responses, the frequency range of sensitivity, expected threshold values and mecha-nisms of sound detection for several species of marine mammals and fishes (Fay 1988; Au et al. 2000). For marine invertebrates, our knowledge of hearing capabilities is relatively poor and a definition or even certainty of sound detection is not agreed upon (Webster et al. 1992) despite their magnitude of biomass and often central role in ocean ecosystems. Cephalopods (squid, cuttlefish, octopods and nautilus) are particularly interesting subjects for inver-tebrate sound detection investigations for several reasons. Ecologically, they occupy many of the same niches as sound-sensitive fish (Budelmann 1994) and may benefit from sound perception and use for the same reasons, such as to detect predators, navigate, or locate conspecifics. Squid, for example, are often the prey of loud, echolocating marine mammals (Clarke 1996), and may therefore be expected to have evolved hearing to avoid predators. Anatomically, squid have complex statocysts that are considered to serve primarily as vestibular and acceleration detectors (Nixon and Young 2003). However, statocysts may also be analogs for fish otolithic organs, detecting acoustic stimuli (Budelmann 1992). Previous studies have debated the subject of squid hearing and recently there has been a revival of research on the subject. Here, we briefly review what is known about squid sound detection, revisit hearing definitions, discuss potential squid susceptibility to anthropogenic noise and suggest potential future research direc-tions to examine squid acoustic sensitivity.
  • Dataset
    Soundscape monitoring acoustic data collected in July of 2017 during an in situ larval coral settlement experiment in St. John, US Virgin Islands
    (Biological and Chemical Oceanography Data Management Office (BCO-DMO). Contact:, 2020-01-21) Apprill, Amy ; Mooney, T. Aran ; Lillis, Ashlee
    Matlab R2016 was used to process acoustic data from raw wave audio files. Mean power spectral densities were estimated (Hamming window, non-overlapping 0.5-sec windows, frequency resolution: 1.47 Hz) within 1-minute samples across the total experiment length (62 hours). For a complete list of measurements, refer to the full dataset description in the supplemental file 'Dataset_description.pdf'. The most current version of this dataset is available at:
  • Dataset
    Dates and locations of hydrophone deployments at coral reefs in St. John, U.S. Virgin Islands in 2016 and 2017
    (Biological and Chemical Oceanography Data Management Office (BCO-DMO). Contact:, 2020-12-15) Mooney, T. Aran ; Apprill, Amy ; Dinh, Jason
    A passive acoustic recorder was deployed at various coral reefs in St. John, U.S. Virgin Islands between 2016-03-28 and 2017-07-11. This dataset contains deployment dates and locations. For a complete list of measurements, refer to the full dataset description in the supplemental file 'Dataset_description.pdf'. The most current version of this dataset is available at:
  • Dataset
    Porites astreoides coral settlement counts collected in July of 2017 from an in situ larval coral settlement experiment in St. John, US Virgin Islands
    (Biological and Chemical Oceanography Data Management Office (BCO-DMO). Contact:, 2020-01-03) Apprill, Amy ; Mooney, T. Aran ; Lillis, Ashlee
    Brooding coral Porites astreoides colonies were collected on St. John, U.S. Virgin Islands on June 22nd, 2017 and used in an in-situ larval coral settlement experiment. Settlement counts were taken on June 28th, 2017. For a complete list of measurements, refer to the full dataset description in the supplemental file 'Dataset_description.pdf'. The most current version of this dataset is available at:
  • Dataset
    Acoustic summary data from hydrophone deployments at coral reefs in St. John, U.S. Virgin Islands in 2016 and 2017
    (Biological and Chemical Oceanography Data Management Office (BCO-DMO). Contact:, 2020-12-15) Mooney, T. Aran ; Apprill, Amy ; Dinh, Jason
    A passive acoustic recorder was deployed at various coral reefs in St. John, U.S. Virgin Islands between 2016-03-28 and 2017-07-11. This dataset contains average sound pressure level, low frequency sound pressure level (50 - 1500 Hz), high frequency sound pressure level (2 kHz - 20 kHz) and peak frequency of files at each site, sorted by files with and without boat noise. For a complete list of measurements, refer to the full dataset description in the supplemental file 'Dataset_description.pdf'. The most current version of this dataset is available at: