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The Ocean Cacophony

Detailed Analysis of Vessel Underwater Radiated Noise (URN), Its Bioacoustic Effects on Cetaceans, and Global Mitigation Strategies

Contents

Executive Summary

Underwater Radiated Noise (URN) from global vessel traffic constitutes a pervasive and chronic threat to Cetaceans, impacting critical life functions including foraging, migration, and reproduction. The noise is highly intense; source levels from large ships, when acoustically translated to human-equivalent airborne sound intensity, often correspond to levels exceeding 130 dB, a threshold associated with human pain and damage. The primary biological impact is chronic physiological stress, alongside acute behavioral changes (avoidance, increased energy expenditure) and severe acoustic masking, particularly for low-frequency vocalizing Mysticetes (baleen whales). Mitigation solutions are available across technical and operational domains. Technical innovations focus on propeller cavitation reduction and machinery isolation, which show high synergy with fuel efficiency efforts. Operationally, Vessel Speed Reduction (VSR) is highly effective, but regulatory frameworks, largely driven by non-mandatory IMO guidelines, struggle with compliance, necessitating a shift toward robust, mandatory implementation guided by advanced monitoring technologies like Passive Acoustic Monitoring (PAM).

Chapter 1: Foundations of Underwater Radiated Noise (URN)

1.1. The Anthropogenic Soundscape: Historical Context and Source Contribution

Cetaceans, including whales, dolphins, and porpoises, are fundamentally reliant on sound as their primary sense for interacting with the marine environment.1 Sound transmission in water is highly efficient, allowing cetaceans to communicate, navigate, find food, and locate mates over distances that can span hundreds of kilometers.2 This reliance on acoustic communication makes marine mammals acutely sensitive to alterations in their acoustic environment. Over the past six decades, the ocean soundscape has undergone a profound transformation due to human activity, with anthropogenic noise increasing tenfold in volume since the 1960s.2

This escalating acoustic contamination is largely driven by commercial shipping, which is the primary source of continuous, low-frequency anthropogenic noise in the ocean.5 This pervasive background noise fundamentally alters the acoustic range available to large whales. Compared to pre-industrial conditions, ambient noise levels in deep-water shipping lanes in the Northern Hemisphere have increased by approximately 20 dB in the frequencies below 100 Hz.6 This dramatic and chronic increase in background noise, which is directly linked to the tripling in the number and size of vessels in the world's merchant fleet7, forces whales into energetically costly compensatory behaviors, such as increasing the amplitude or frequency of their vocalizations, simply to maintain essential communication across shorter distances.9

1.2. Acoustic Signatures of the Global Fleet: Differentiation by Vessel Class

Vessel noise is generated by both large commercial ships and smaller recreational boats, but their acoustic characteristics and resulting ecological impacts differ significantly. Large commercial vessels, such as cargo carriers and tankers, generate powerful, low-frequency sound because they are large and typically move at faster speeds.10 The acoustic energy from these ships is primarily concentrated below 500 Hz, affecting indicator frequency bands like 63 Hz and 125 Hz.10 The broadband source level for a large ship measured at 1 m can reach levels up to 200 dB re 1 µPa.11 The principal mechanism for this intense noise is propeller cavitation, which occurs when low-pressure areas around the rotating propeller blades cause vapor bubbles to form and subsequently collapse violently, generating high-intensity noise across a broad frequency spectrum.12

In contrast, smaller recreational boats generate acoustic signatures characterized by clear tones derived from the rotational frequencies of their engines and propellers.14 While data on recreational boat noise is less standardized than that for commercial shipping14, studies of small vessels (e.g., 6 m length with 30 to 180 hp engines) show high localized radiated noise levels (RNLs) ranging between 153.4 and 166.1 dB re 1 µPa m.15 Importantly, these smaller vessels emit substantial acoustic energy at higher frequencies, sometimes reaching tens of kilohertz, which is highly relevant to toothed whales (Odontocetes).16 The distinction between these two primary sources is critical for policy development. Commercial shipping represents a low-frequency, global environmental contaminant requiring coordinated international regulation (e.g., through the IMO). Conversely, recreational boat noise acts as a high-frequency, localized contaminant, necessitating regional or national management, such as the implementation of noise emission standards within whale-watching regulations.17

Chapter 2: Quantifying the Acoustic Threat: Bridging the Air-Water Decibel Divide

Accurately describing the intensity of underwater sound in terms that relate to human experience requires addressing the fundamental physical and measurement differences between acoustic energy in water and air.

2.1. Inherent Discrepancies in Acoustic Reference Standards

The decibel unit (dB) is a logarithmic ratio measure requiring a standard reference pressure. In underwater acoustics, the universally accepted reference pressure is 1 microPascal (1 µPa), resulting in measurements reported as dB re 1 µPa.18 However, the human auditory system's threshold in air is near 20 µPa, which is the conventional reference standard for airborne sound.18 This discrepancy means that comparing the two scales directly is misleading; the 20 µPa reference used in air is already 20 log(20/1) ≈ 26 dB higher than the 1 µPa reference used underwater.20 Therefore, an initial step in pressure normalization requires subtracting 26 dB from the underwater measurement.

2.2. Physics of Sound Transmission: Impedance and Energy Flow Conversion

Normalization solely for reference pressure is insufficient, as it fails to account for the stark difference in the media through which the sound travels. Water is about 800 times denser than air and significantly less compressible. Consequently, a sound wave carrying the same pressure in air and water will transfer drastically different amounts of energy (intensity or power flow).19 This physical difference is quantified by the acoustic characteristic impedance of water, which is approximately 3600 times that of air.22

To determine a sound intensity level (SIL) equivalence that provides a relatable estimate of "loudness" to the human auditory system, the 26 dB pressure reference adjustment must be combined with the impedance correction, which is approximately 10 log(3600) ≈ 36 dB.22 The resultant generalized conversion factor used to translate underwater radiated noise (re 1 µPa) to an equivalent airborne sound pressure level (SPL) (re 20 µPa) requires a total subtraction of approximately 62 dB.22 This adjustment is vital because, without it, the raw underwater decibel value significantly overestimates the effective acoustic energy level that would be perceived by a human ear in air. This comprehensive acoustic translation ensures that the discussion of acute acoustic threats remains grounded in accurate physical comparison, allowing for non-sensational yet precise framing of the risk of near-source exposure.

2.3. Translation of Vessel Source Levels to Human-Equivalent Airborne Decibels

Applying this 62 dB conversion factor demonstrates the extreme intensity of URN. A large commercial ship generating 200 dB re 1 µPa at 1 m11 is roughly equivalent to a 138 dB sound in air (200 dB - 62 dB).22 This level is comparable to the noise produced by a jet engine at close range and significantly exceeds 130 dB, which is cited as the human pain threshold.22 For example, a supertanker radiating 190 dB underwater is equivalent to approximately 128 dB in air.22 While cetaceans possess distinct audiograms and hearing sensitivities, exposure to such intense sound levels poses serious risks. If a human were submerged and exposed to sound pressure levels of 200 dB re 1 µPa, physical hearing damage would be a certainty.21

Table 1: Underwater Sound Pressure Levels (dB re 1 µPa) and Approximate Air Equivalents

Noise Source (Underwater) Source Level (dB re 1 µPa @ 1m) Approximate Air Equivalent (dB re 20 µPa) Human Reference Equivalent (Air)
Seismic Air Gun (1m) 240 ~178 Severe hearing damage risk; Close-range explosion
Typical Active Sonar 220 ~158 Military aircraft takeoff
Large Ship, Broadband (1m) 200 ~138 Jet engine at 30m; Immediate human pain threshold
Supertanker 190 ~128 Loud rock concert or chainsaw
Ambient Noise, Sea State 4 100 ~38 Quiet residential area

Chapter 3: Critical Biological and Behavioral Impacts on Cetacean Populations

The consequences of elevated URN extend far beyond instantaneous physical damage, encompassing chronic stress, severe communicative interference, and detrimental behavioral changes that collectively diminish the fitness of marine mammal populations.

3.1. Mechanisms of Acoustic Masking: Critical Ratios and Communication Space

Acoustic masking is the degradation of an animal's ability to detect or recognize a sound of interest due to the presence of another sound source, the masker.23 Shipping noise, particularly its dominant low-frequency components, directly overlaps the vocalization frequencies used by baleen whales for long-distance communication.2 This constant interference functionally raises the auditory detection threshold for the whale's own biologically relevant sounds, effectively shrinking its communication space—the area over which signals can be detected.3

The effectiveness of masking varies significantly across frequencies and taxa, governed by the critical bandwidth (CB) of the listener.25 Odontocetes have narrow CBs at high frequencies (above 1 kHz), meaning only noise energy within a narrow spectral band will effectively mask their calls. However, for low-frequency signals, such as those used by Mysticetes, sound energy across a much wider band can cause substantial masking.25 While some research suggests that cetaceans may possess spectral or temporal processing capabilities enabling them to filter out non-threatening boat noise when the signals are biologically relevant (such as killer whale vocalizations)25, the overall degradation of the acoustic habitat necessitates compensatory behaviors. These behaviors, such as increasing call amplitude or frequency9, consume additional energy without fully restoring the historic communication range, highlighting a chronic ecological inefficiency.

3.2. Physiological Harm: Evidence of Noise-Induced Acute and Chronic Stress

The most compelling evidence of physical harm from URN relates to chronic physiological stress. A unique study tracking North Atlantic right whales (Eubalaena glacialis) found that a spontaneous 6 dB decrease in background noise following the maritime traffic halt after 9/11 correlated directly with decreased baseline levels of stress-related faecal hormone metabolites (glucocorticoids).2 This groundbreaking finding established that exposure to persistent low-frequency ship noise induces a chronic stress response in whales, indicating that noise is not merely an annoyance but has a definite physical effect.2

This discovery mandates a significant shift in management paradigms. Traditional regulatory frameworks often focus on avoiding high-intensity acoustic events that cause Temporary or Permanent Threshold Shifts (TTS/PTS), or instantaneous hearing damage. However, the evidence for chronic stress reveals that conservation efforts must prioritize the cumulative, long-term impact of persistent low-level noise.26 Managing acoustic habitat quality to alleviate widespread systemic stress becomes critical, particularly for endangered populations experiencing the accumulated impact of multiple stressors in heavy ship traffic areas.9

3.3. Behavioral Effects: Avoidance, Energetic Expenditure, and Displacement

Noise eliciting behavioral responses in cetaceans often represents avoidance strategies that come at the cost of increased energy expenditure.17 Studies on humpback whale mother-calf pairs subjected to high noise playbacks showed immediate and significant negative responses: the mother's resting time dropped by 30%, respiration rate doubled, and swim speed increased by 37%.17

Cetaceans engage in various forms of avoidance, moving away from sound sources27 or exhibiting strong startle responses, particularly to noise that has a sudden onset.25 This is especially true for small coastal dolphins, which show strong aversive behaviors toward high-speed recreational traffic like speedboats and jet skis.27 The combined effect of increased energy expenditure due to avoidance behavior and reduced foraging efficiency stemming from communication masking creates an "energetic deficit." For vulnerable species, this constant tax on metabolic reserves hinders recovery and population viability.

3.4. Long-Term Ecological Consequences: Migration and Habitat Abandonment

The disruption caused by URN can have significant consequences at the population level. Simulation studies suggest that the current acoustic environment, compared to pre-industrial conditions, may cause delays in whale migration arrivals of 3 to 4 days, translating to up to 20% added travel time.3

If noise contamination continues to rise, the consequences could be more severe. Noise generates an avoidance response and also lowers the detection of crucial environmental cues used for navigation.3 The model suggests that some whales may fail to reach their destinations, either by drifting off course or by being blocked by what effectively becomes an impenetrable "wall of noise" along their routes.3 This displacement from critical feeding, mating, or calving grounds poses a direct threat to population stability.

3.5. Indirect Impacts via Prey Species: The Trophic Cascade

The ecological damage from URN is not limited to direct effects on cetacean hearing and behavior; it extends systemically to prey species, including fish and invertebrates.1 Noise exposure can cause permanent and temporary hearing loss, stress, and behavioral reactions in fish, leading to reduced catch rates.28 For instance, hearing-specialist fishes, such as goldfish, exhibited Temporary Threshold Shifts (TTS) of up to 28 dB after just 24 hours of noise exposure.29 The time required for full recovery extended up to 14 days following a 21-day exposure period.29 The reduction in prey availability and the impairment of prey capture efficiency places an additional burden on cetacean populations, particularly those already facing resource limitations. This negative feedback loop—where noise both increases the predator's energy demand and reduces the available food supply—exacerbates the conservation challenge for apex marine species.

Chapter 4: Taxa-Specific Vulnerability: Mysticetes vs. Odontocetes

Vulnerability to URN is highly dependent on the characteristic hearing and vocalization frequencies of different cetacean groups.

4.1. Vulnerability of Baleen Whales (Mysticetes) to Low-Frequency URN

Mysticetes, which include blue, fin, and humpback whales, primarily use low-frequency sounds (pulses, moans, and whoops) ranging from tens to hundreds of hertz.4 These frequencies are highly susceptible to acoustic masking because they directly overlap with the low-frequency noise dominated by large commercial shipping (typically below 300 Hz).6 Baleen whales are considered more sensitive at these low frequencies than other marine mammals.16

The physiological adaptations that enable these whales to produce powerful, long-range sound underwater also constrain them. Recent research indicates that the evolved structure of baleen whale throats restricts their vocal range, preventing most species from shifting their vocalizations outside the persistent noise range generated by ships.30 Given that evolutionary adaptation requires thousands to millions of years, and anthropogenic noise pollution has dramatically increased over mere decades, baleen whales are unlikely to adapt quickly enough to the rapid acoustic shift.30 While some species, like humpback and right whales, possess specialized cartilage allowing them to produce higher frequency calls (up to 6,000 Hz), which can temporarily pierce the low-frequency ship noise, these calls travel shorter distances and may not replace the function of long-range acoustic signaling.30

4.2. Impacts on Toothed Whales (Odontocetes) and Coastal Species

Odontocetes (toothed whales, dolphins, and porpoises) utilize mid- to high-frequency whistles and clicks for both communication and echolocation.4 Their primary vulnerability stems from high-frequency URN, often generated by the higher-frequency energy radiated by both large ships and, critically, smaller recreational and coastal vessels.16

Coastal populations, such as the Southern Resident Killer Whales (SRKWs), are intensely exposed to localized small vessel traffic noise.16 Vessel noise is recognized as one of the three primary threats to SRKW recovery.31 The acoustic interference masks the whales' echolocation signals, which are essential for coordinated hunting behavior.25 By making it more difficult for these whales to find and catch salmon, the vessel noise effectively reduces the availability of prey.31 Consequently, regulatory bodies have begun implementing measures specific to these coastal populations, such as the new 1,000 yard mandatory buffer zone in Washington state waters, forcing boaters to minimize acoustic disturbance near the endangered SRKWs.31

Table 2: Comparative URN Vulnerability Profiles of Cetacean Taxa

Cetacean Group Key Communication Frequencies Dominant Vessel Noise Overlap Primary Impact Mechanism
Mysticetes (Baleen Whales) Low-frequency pulses/moans (tens to hundreds of Hz) Commercial Shipping (50-300 Hz) Chronic Stress; Long-range Communication Masking
Odontocetes (Toothed Whales) Mid- to High-frequency clicks/whistles (kHz) Propeller Cavitation, Small Vessels (tens of kHz) Echolocation Masking (Foraging); Acute Behavioral Avoidance

Chapter 5: Mitigation and Engineering Solutions for URN Reduction

Solutions for reducing URN are available across both technological design improvements and operational management practices.

5.1. Source Control: Propeller Hydrodynamics and Cavitation Minimization

Propeller cavitation remains the single largest contributor to noise pollution from large merchant ships.6 Consequently, technical mitigation is heavily focused on propeller design intended to minimize the formation and collapse of cavitation bubbles.13

Modern engineering efforts involve optimizing propeller blade shape and operating conditions to prevent large pressure drops.13 Crucially, efforts by the shipping industry to improve fuel efficiency often enhance hydrodynamic performance and reduce cavitation, suggesting a strong synergy between decarbonization goals and noise reduction.32 This was demonstrated during the "Radical Retrofit" of Maersk G-class container vessels, which focused on energy efficiency (increasing vessel draft and installing new propellers with boss cap fins). Post-retrofit data showed a significant reduction in source levels, with a median decrease of 6 dB in the low-frequency band and 8 dB in the high-frequency band.12 Such retrofits prove that large-scale source reduction is both technically viable and results in measureable acoustic benefits, especially at lower operating speeds.8 Furthermore, innovative designs, such as the trochoidal propeller that mimics a whale's tail motion33, and custom hybrid propellers capable of 8 to 13.8 dB noise reductions34, offer further specialized pathways for quieting the global fleet.

5.2. Machinery Noise Isolation and Hull Treatments

Internal machinery—including the engine, generator, and pumps—generates structure-borne vibration and airborne sound that propagates through the hull into the water.35 Effective source control must first focus on insulating and damping these internal components.35

Technical measures involve robust isolation systems. Resilient marine engine mounts utilize rubber in shear and compression to decouple the engine from the hull, reducing structure-borne vibration.36 For engine room treatments, a combination of materials is essential: sound-absorbing foams target mid- to high-frequency energy, while dense barriers like Mass Loaded Vinyl (MLV) block sound transmission.35 Beyond internal dampening, specialized hull treatments can also reduce URN. Air lubrication systems inject air bubbles beneath the hull to reduce frictional resistance, which cuts fuel consumption and incidentally reduces URN.33 A more aggressive system, historically used by navies, is the Masker system, which injects air bubbles near machinery spaces to actively muffle internal noise propagation.33

5.3. Operational Mitigation Programs: Vessel Speed Reduction (VSR)

The most immediate and scalable solution is the implementation of Vessel Speed Reduction (VSR) policies, often referred to as "slow steaming." Reducing speed dramatically reduces propeller cavitation and, consequently, URN.6 Quantified evidence supports VSR effectiveness: a 10% reduction in speed can decrease noise pollution by 40%, and a 20% speed reduction can cut noise pollution by 67%.39 This operational approach offers rapid relief to marine mammals in critical habitats.

Furthermore, integrating VSR with optimized routing—such as the voluntary recommended routes in place for North Atlantic right whale conservation—reduces the risk of mortality and collision.40 The implementation of specialized local operational measures, such as turning off high-frequency ultrasonic anti-fouling systems in critical areas, further optimizes the acoustic habitat quality for species like killer whales.41

Chapter 6: Policy, Governance, and Monitoring Frameworks

6.1. International Regulatory Landscape and the Mandate Gap

International governance of URN primarily rests with the International Maritime Organization (IMO), which began discussions on the issue in 2004.5 The IMO acknowledged the adverse effects of URN on marine life and approved guidelines for commercial ships on reducing noise in 2014, with revisions approved in 2023.5 These Guidelines for the Reduction of Underwater Noise are intended to provide advice to designers, shipbuilders, and operators on managing URN.32

However, the major impediment to widespread implementation is the non-mandatory status of these international URN guidelines.5 Unlike the mandatory SOLAS and MLC 2006 regulations governing noise protection for human personnel aboard ships42, the guidelines for marine life conservation rely purely on voluntary adoption and awareness.32 This regulatory gap means that despite the feasibility of technical solutions and the clear effectiveness of operational measures, adoption rates remain marginal across the global fleet. The lack of mandatory global standards hinders the potential to quiet the noisiest 10% of ships, which are believed to generate the majority of the total noise impact.6

6.2. Compliance and Enforcement of Operational Measures

The effectiveness of operational mandates varies dramatically depending on the regional context and enforcement mechanisms. In regions with dedicated management and strong collaboration, such as the right whale protection zones, combining reduced speeds with recommended routes saw compliance rates steadily increase to as high as 96% during successful study periods.40

In contrast, reports analyzing broader mandatory speed zones established by the National Oceanic and Atmospheric Administration (NOAA) indicated alarmingly low compliance, with only about 10% of vessels adhering to mandatory restrictions and 15% cooperating in voluntary areas between 2017 and 2020.44 This severe variance in compliance underscores that policy integration must prioritize robust real-time monitoring, enforcement, and the development of incentive programs to encourage fleet-wide adherence to URN reduction goals.45 The implementation of local, mandatory regulations, such as the 1,000 yard buffer zone for SRKWs, demonstrates that regional bodies are stepping in to enforce the noise reduction necessary for species recovery when global cooperation falters.31

6.3. Monitoring URN: The Role of Passive Acoustic Monitoring (PAM) Networks

The transition from non-binding advice to verifiable conservation outcomes depends on comprehensive monitoring frameworks. Passive Acoustic Monitoring (PAM) networks utilize hydrophones and hydrophone arrays (both fixed systems like SOSUS and mobile systems) to track vessels, locate cetaceans, and characterize the acoustic environment over vast scales.46

PAM provides the scientific basis required for regulatory action. For example, analysis of PAM data demonstrated that the disruption of commercial shipping during the COVID-19 pandemic led to a measurable reduction in ocean noise levels by as much as 2 dB (approximately 30%) across the North Pacific and Arctic Oceans.7 This empirical evidence directly supports the hypothesis that quietening measures result in ecological benefit.7 Furthermore, advanced PAM systems enable real-time decision support tools, allowing managers to dynamically track vessel compliance and relay real-time whale presence to mariners, facilitating immediate, targeted mitigation efforts.41 These monitoring networks are essential for transforming theoretical noise reduction objectives into measurable, verifiable ecological outcomes, thereby closing the loop between scientific understanding and regulatory action.

Conclusions and Recommendations

The acoustic integrity of the marine environment is fundamental to cetacean survival. Underwater Radiated Noise from shipping causes chronic stress, massive communication masking, and increased energy expenditure across whale populations, equivalent to continuous human exposure to sounds nearing or exceeding the threshold of physical pain.

Mandatory Standards Development

The non-mandatory nature of current international guidelines represents the largest barrier to effective URN mitigation. International bodies must move towards mandatory noise emission standards, perhaps initially targeting the noisiest 10% of the global fleet, to ensure systemic acoustic habitat improvement.

Incentivize Dual-Benefit Technologies

Given the proven synergy between propeller design optimizing fuel efficiency and reducing cavitation noise (6 to 8 dB reductions demonstrated), regulatory bodies should implement strong incentive programs that reward vessel owners for adopting technologies and retrofits that achieve both decarbonization and URN reduction goals.

Enhance Monitoring and Enforcement

The efficacy of operational measures like Vessel Speed Reduction is highly dependent on enforcement. Increased investment in Passive Acoustic Monitoring (PAM) networks is required to provide real-time data for compliance tracking, dynamic route management, and the validation of noise reduction measures, turning theoretical goals into verifiable ecological outcomes.

Prioritize Chronic Stress Mitigation

Policy must expand beyond managing acute hearing damage (TTS/PTS) to managing acoustic habitat quality. The established link between low-frequency URN and chronic physiological stress necessitates conservation measures aimed at reducing sustained background noise levels, particularly in critical habitats used for foraging, mating, and nursing.

Works Cited

  1. Weilgart, L. "The impacts of anthropogenic ocean noise on cetaceans and implications for management." Canadian Journal of Zoology, 2007. http://whitelab.biology.dal.ca/lw/publications/Weilgart%202007%20CJZ%20noise%20review.pdf
  2. "Shipping causes 'chronic stress' to whales." The Guardian, Feb 8, 2012. https://www.theguardian.com/environment/2012/feb/08/shipping-noises-chronic-stress-whales
  3. "Avoidance, confusion, solitude: whales react to rising noise pollution." University of Melbourne, Faculty of Science. https://science.unimelb.edu.au/about/news/-avoidance,-confusion,-solitude-whales-react-to-rising-noise-pollution
  4. "Marine Mammal Acoustics." NOAA Fisheries. https://www.fisheries.noaa.gov/new-england-mid-atlantic/science-data/marine-mammal-acoustics
  5. "IMO and anthropogenic underwater noise." United Nations. https://www.un.org/depts/los/consultative_process/contributions_19cp/IMO.pdf
  6. "A review of practical methods for reducing underwater noise pollution from large commercial vessels." Convention on Biological Diversity. https://www.cbd.int/doc/meetings/mar/mcbem-2014-01/other/mcbem-2014-01-submission-reducing-noise-en.pdf
  7. "A rare glimpse of a quieter ocean." NOAA Research. https://research.noaa.gov/a-rare-glimpse-of-a-quieter-ocean/
  8. "Retrofit-induced changes in the radiated noise and monopole source levels of container ships." NIH PMC. https://pmc.ncbi.nlm.nih.gov/articles/PMC10019734/
  9. "Evidence that ship noise increases stress in right whales." NIH PMC. https://pmc.ncbi.nlm.nih.gov/articles/PMC3350670/
  10. "Large Vessel Activity and Low-Frequency Underwater Sound Benchmarks in United States Waters." Frontiers in Marine Science. https://www.frontiersin.org/journals/marine-science/articles/10.3389/fmars.2021.669528/full

Note: Full citation list includes 48 sources. Complete references available upon request.