John A. Hildebrand’s research while affiliated with University of California System and other places

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Publications (405)


Evaluating Seasonal vs. Spatial Extrapolation for Cetacean Distribution Models in the California Current
  • Article

June 2025

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5 Reads

Marine Mammal Science

Elizabeth A. Becker

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Karin A. Forney

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Bruce J. Thayre

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[...]

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Mary Simeon

Species distribution models (SDMs) have been developed for multiple cetacean species within the California Current Ecosystem (CCE) from shipboard survey data collected by the Southwest Fisheries Science Center (SWFSC) in summer and fall, thus limiting the ability to inform management decisions in cool seasons when abundance and distribution patterns are substantially different. Winter and spring SDMs have been developed for a few species using California Cooperative Oceanic Fisheries Investigations (CalCOFI) shipboard survey data, but model predictions are limited to the waters off southern and central California. In this study, winter and spring density estimates for the entire CCE study area were made from SWFSC summer and fall model predictions (temporal extrapolation) and CalCOFI winter and spring model predictions (spatial extrapolation) for short-beaked common dolphin (Delphinus delphis delphis), Pacific white-sided dolphin (Lagenorhynchus obliquidens), Dall's porpoise (Phocoenoides dalli), and fin whale (Balaenoptera physalus). The performance of the models was compared based on available abundance estimates and documented distribution patterns in the cool seasons. Results reveal species-specific ecological factors to consider when extrapolating model predictions temporally or spatially, including whether a given study area includes a species “core habitat”, and whether static variables should be included when a species exhibits temporal distribution shifts.


Climatic and economic fluctuations revealed by decadal ocean soundscapes

June 2025

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9 Reads

The Journal of the Acoustical Society of America

Decadal variations of ocean soundscapes are intricately linked to large-scale climatic and economic fluctuations. This study draws on over 15 years of acoustic recordings at six sites within the Southern California Bight, investigating interannual, seasonal, and diel variations. By examining acoustic energy from fin and blue whales along with sounds from ships and wind, we identified changes in soundscape over time and space. This study reveals that sound levels associated with both biological and non-biological sound sources varied seasonally and correlated with large-scale climatic patterns and long-term oceanographic fluctuations. Baleen whale sound levels before, during, and after a marine heatwave were assessed; sound levels decreased in southern sites and increased in northern sites adjacent to the California Current, underscoring the potential for range shifts and habitat compression during warm years for these species. Ship-generated sound levels at high-traffic sites reflected economic events such as recessions, labor shortages and negotiations, and changes to port activities. Marine soundscapes offer an approach to assess the ocean's condition amid ongoing climatic and economic fluctuations.


Components of the gray whale tympanoperiotic complex (TPC) highlighted on a CT reconstruction of the right TPC (Specimen: SDNHM 25307). Orientation directions are shown in the lower left corner (D = dorsal, A = anterior, V = ventral, P = posterior). Gray: periotic bone - embedded in the skull. Teal: tympanic bulla - suspended from the periotic by two pedicles (Purple). Yellow dashes: outline of the ossicular chain. Solid Yellow: part of the ossicular chain not obscured by the tympanic bulla or periotic. Black star: fused junction between the malleus (first ossicular element) and the tympanic bulla. White star: location of the oval window, the interface that allows the stapes footplate (last ossicle) to push on the fluid channels in the cochlea of the inner ear.
Schematic of underwater test arrangement at TRANSDEC. The skull and sound source were each lowered halfway down the water-column (5.5–6 m depth). The projector was mounted on a pole in a fixed position, while the skull hung in a net suspended from a pole that was able to rotate to a specified angle relative to the transducer. The source to skull centroid separation was 1 m, putting the tip of the rostrum within 1 m of the sound source. An H-52 reference hydrophone was suspended from a bridge 12.5 m from the sound source and at 6 m water depth.
Accelerometer placement on skulls. (a) Plastic juvenile skull with aluminum plates attached to the plastic with J-B Weld MarineWeld epoxy. During experiments, accelerometers were superglued to the aluminum plates. Arrows show direction of measurement, and annotations describe the measurement that takes place at each location. (b) Sensors placed on natural juvenile skull. Accelerometers were superglued to the aluminum plates. In this case, the skull sensors are placed further forward on the skull to avoid remaining soft tissue. (c) Sensors on denuded juvenile skull. (d) Sensors on adult skull. In cases (c) and (d), sensors were glued directly to the bone (without aluminum plates).
Frequency response functions obtained with a low-frequency underwater source ensonifying the juvenile skull in each of its 3 forms at 0∘\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$0^\circ$$\end{document} (head-on) incidence. Smoothed curves are shown as thick lines atop the unsmoothed data. (a) Mediolateral velocity divided by incident pressure on the plastic skull. (b) Anteroposterior velocity divided by incident pressure on the plastic skull. (c) Mediolateral velocity divided by incident pressure on the natural skull. (d) Anteroposterior velocity divided by incident pressure on the natural skull. (e) Mediolateral velocity divided by incident pressure on the denuded skull. (f) Anteroposterior velocity divided by incident pressure on the denuded skull.
Frequency response ratios obtained with a low-frequency source ensonifying the plastic skull underwater between 170–1000 Hz at varying angles of incident ensonification (0∘\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$0^\circ$$\end{document} is head-on ensonification). (a) Ratio of left bulla to skull mediolateral velocity. (b) Ratio of right bulla to skull mediolateral velocity. (c) Ratio of left bulla to skull anteroposterior velocity. (d) Ratio of right bulla to skull anteroposterior velocity.

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Experimental observation of gray whale skull vibrations amplified in the bony hearing complex
  • Article
  • Full-text available

April 2025

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39 Reads

Mysticete whales have bilateral bony ear complexes (tympanoperiotic complexes) that amplify low frequency vibrations in proximity to their vocalization ranges. Understanding the functional mechanics would enable precise predictions of mysticete hearing sensitivity, which is currently unknown. We conducted experiments on a juvenile and an adult gray whale skull from deceased animals to measure the vibrational dynamics between the tympanic bullae and the skull. Relative motions between bullae and skull indicate sound transfer to the inner ear. For the juvenile, assessments were performed on (1) a 3D-printed plastic skull-replica, (2) the original skull after much of the soft tissue had been removed by dissection, and (3) the denuded skull after hydrogen peroxide was used to erode the remaining soft tissues. We excited vibrations in the juvenile skull underwater by projecting sound in a test pool, ranging from 170–1000 Hz. Additionally, we measured in-air vibrations of the plastic, denuded, and adult skulls using a mechanical shaker to drive vibrations anteroposteriorly (rostrum-to-tail) from 150–1000 Hz. The results consistently showed amplification of vibrations at the tympanic bullae compared to the base of the skull, demonstrating a mechanism by which low-frequency sound is transferred from the environment, through the skull, to the inner ear.

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Figure 1. a) Map of study area. Red dots represent HARP deployment sites. b) Instrument deployment configuration. One deployment of three passive acoustic recorders from site N shown as an example. Square markers represent 4-hydrophone small-aperture arrays, and circle marker represents the single hydrophone mooring. c) Diagram of instrumentation. Renderings of a 4-hydrophone array and a single hydrophone recorder.
Figure 4. a) Tracked whales by site. Tracks are colored by normalized track time, with the start of the track in purple and the end of the track in light blue. Square markers represent 4-hydrophone arrays, and circle markers represent single hydrophones. Black markers are used at sites with one deployment period and yellow markers are used at sites with multiple deployments, delineating the position of the first deployment. Bathymetric contour lines are every 100 m. b) Densities of tracked whales per site shown using 10 m by 10 m grid squares colored by the number of tracked whales within the squares. c) Probabilities of tracking an echolocating whale due to hydrophone array geometry based on Monte-Carlo simulation, normalized between 0 and 1.
Figure 6. a) Encounter with two whales at depth (solid lines) colored by time, illustrating distances with respect to time (pairs distance, purple dashed lines) and at closest points of approach without considering time (lane distance, blue dashed lines). For clarity, only a portion of calculated lane distances are plotted. b) Distance between pairs of whales in an encounter with respect to time (pairs distance, purple) and without respect to time (lane distance, blue) at all sites. Lines represent modal bins for each. c) Histogram of time lags between the closest points of approach for whale pairs per encounter at all sites.
Figure 7. a) Encounter with three whales forming two subgroups indicated by color (red and blue). Shading represents time, from light (start of track) to dark (end of track). Gaps in detections for an individual are represented as a dashed line. Black triangles denote positions of 4-hydrophone receivers. b) Number of subgroups per encounter for all deployments by total number of whales present, color represents counts. c) Distribution of mean number of whales across subgroups per encounter.
Long-term monitoring of Ziphius cavirostris behavior using 3D tracking from fixed hydrophone arrays off Southern California

March 2025

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39 Reads

Goose-beaked whales (Ziphius cavirostris) are a deep-diving toothed whale species and top predators in deep sea ecosystems. Much is yet to be learned about their social and foraging strategies due to their elusive behavior, but this information isincreasingly relevant given their demonstrated behavioral changes in association with anthropogenic sound. This study useddirection-of-arrival (DOA) localization to track the position of goose-beaked whales from echolocation clicks recorded onseafloor-mounted hydrophone arrays offshore Southern California. Overall, 2,738 tracks of diving goose-beaked whales wereprocessed from acoustic recordings collected at four long-term monitoring sites between 2018 and 2023. Results highlightdistinct spatial use patterns driven by bathymetric features at each site, with whales foraging closer to the seafloor at siteswith complex bathymetry and showing a preference for canyon slopes. Group sizes at depth ranged from 1 to 9 individualswith a mean of 2.34 and exhibited site-specific seasonal variability as well as a strong diel trend at one site. During manyof these encounters, individuals exhibited highly coordinated behaviors. This study demonstrates the value of long-termpassive acoustic tracking for studying elusive, deep-diving species and provides significant advancements in understandinggoose-beaked whale behavior at depth over long time scales.



Site location map
Orange circles indicate five sites monitored acoustically for toothed whale occurrence from 2010–2020. Site MC occupied two slightly different locations (1) and (2). The black triangle indicates the location of the Deepwater Horizon wellhead, with cumulative surface oil footprint⁷³ in red. Bathymetric contour depths are given in meters. Bathymetry obtained from GEBCO⁷⁴.
Density trends of deep diving odontocetes at deep monitoring locations from 2010 to 2020
Black dots represent acoustically-derived weekly density estimates. Error bars represent standard deviation. Median annual density change estimates (Δ) and associated interquartile ranges estimated using a bootstrap are noted in text, with seasonally-detrended Thiel-Sen fits of the time series denoted as red dotted lines. Pink bars indicate data gaps.
Density trends of delphinids at five monitoring locations from 2010 to 2020
Black dots represent acoustically-derived weekly density estimates. Error bars represent standard deviation. Median annual density change estimates (Δ) and associated interquartile ranges are noted in text, with seasonally-detrended Thiel-Sen fits of the time series denoted as red lines. Pink bars indicate data gaps.
Site and species-specific trends in marine mammal group densities over the 2010 to 2020 period
A decade of declines in toothed whale densities following the Deepwater Horizon oil spill

December 2024

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126 Reads

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3 Citations

Shortly after the Deepwater Horizon oil spill began in April 2010, a widely spaced passive acoustic monitoring array was deployed in the northeastern Gulf of Mexico to document the impacts of this unprecedentedly large and deep offshore oil spill on oceanic marine mammals. The array was subsequently maintained for over a decade. Here we document decadal density declines for seven of eight monitored species groups, including sperm whales (up to 31%), beaked whales (up to 83%), and small delphinids (up to 43%). Declines were observed both within and outside of the surface oil footprint. Though not conclusively linked to the oil spill, the broad spatial and temporal scale of these declines observed for disparate marine mammal species is consistent with Deepwater Horizon impacts. These declines have exceeded and outlasted post-spill damage assessment predictions, suggesting that the offshore ecosystem impacts of Deepwater Horizon may have been larger than previously thought.


FIG. 1. (Color online) Vessel traffic density [Automatic identification system (AIS) pings / month] for August 2017. Critical habitats are delineated by white lines including the Channel Islands NMS (dashed), the Blue Whale BIA (solid), and the Humpback Whale BIA (dotted).
FIG. 2. (Color online) Routes and percentage of vessel transits on northern and southern lanes modeled for space-centric noise reduction simulations. Multiroute includes both the SBC TSS (80% of vessels) and the Pt. Mugu Fairway (20% of vessels), as well as two altered Pt. Mugu Fairway options. The singleroutes have the proposed and altered Pt. Mugu Fairways supporting 100% of vessels.
FIG. 3. (Color online) The difference in monthly 50 Hz SPL between modern sound levels from August 2017 AIS data and source-centric simulations. Blue shading indicates areas with greater reduction and white shading indicates areas with no reduction. Speed reduction of all ships is predicted to achieve the greatest reduction in SPL.
FIG. 4. (Color online) Distributions of modeled 50 Hz SPLs within each critical habitat for modern (August 2017), pre-industrial, and simulated sound levels. The speed reduction (all) simulation allowed for the greatest reduction in SPL compared to modern ocean noise levels.
FIG. 6. (Color online) Distribution of SPLs within each critical habitat for seven space-centric simulations, in addition to pre-industrial ocean noise. The single-route techniques allowed for the greatest reduction in SPL compared to the multi-route techniques.
Assessing approaches for ship noise reduction within critical whale habitat

November 2024

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38 Reads

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1 Citation

The Journal of the Acoustical Society of America

Ship noise pollution significantly overlaps with critical habitats of endangered whales in the Santa Barbara Channel, prompting the need for effective noise reduction strategies. Various ship noise reduction approaches were assessed by simulating both source-centric (e.g., speed reduction or retrofit) and space-centric (e.g., routing changes) strategies to determine which would most effectively minimize noise within important marine habitats. Reducing the speeds of all ships achieved the highest noise reduction of the source-centric methods, although solely slowing cargo ships led to similar reductions. Implementing a single-route approach on the southern side of the Channel Islands achieved the greatest reduction of the space-centric strategies. For the multi-route approaches, some noise reduction was achieved by creating a buffer zone between the proposed shipping lanes and the critical habitat boundary. This simulation framework provides a mechanism for efficient exploration and assessment of noise reduction strategies across time and space. The framework can be updated to consider new approaches to changing ocean conditions.


Fig. 1. Acoustic monitoring sites in the northern Gulf of Mexico from 2010 to 2017, named for nearby oceanographic features: Green Canyon (GC), Mississippi Canyon (MC), and Dry Tortugas (DT). Deepwater Horizon wellhead is shown by a star, with cumulative surface oil footprint (Kobara 2019) in brown. Bathymetric contours at 1000 m depth increments are illustrated
Fig. 4. Comparison between received sound pressure levels (RLs) of measured (line) and predicted (bars) sperm whale echolocation clicks, based on cue (solid line) and group (dashed line) counting methods.. The number of clicks is displayed on a log scale. The model predicted RLs assuming animals have a uniform random position and orientation in the horizontal plane around the hydrophone. Refer to Fig. 1 for site abbreviations
Accounting for sperm whale population demographics in density estimation using passive acoustic monitoring

October 2024

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50 Reads

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2 Citations

Marine Ecology Progress Series

Sperm whales Physeter macrocephalus are highly sexually dimorphic, with adult males having larger bodies, more powerful echolocation clicks, and slower echolocation clicking rates compared to females. This study introduces methods for estimating sperm whale population densities in the Gulf of Mexico (GoMex) by accounting for the population demographics using passive acoustic monitoring and reveals that ignoring the differences between demographic segments can introduce bias in density estimates. Weekly densities were estimated per 3 demographic segments: social groups consisting of adult females and their offspring, mid-size animals, and adult males. Analysis revealed that the GoMex sperm whale population is primarily composed of social groups, which account for 92 to 98% of the overall population. Mid-size animals and adult males made up a small proportion of the population and were only intermittently present. Our 7 yr GoMex density estimates, including the 2010 Deepwater Horizon (DWH) oil spill period and subsequent years, revealed demographic-specific trends. Declines found at 2 north-central GoMex sites, coupled with increases at a southeastern site, may indicate population movements and potential impacts from the 2010 DWH oil spill and elevated noise levels from anthropogenic activities in the north-central GoMex.


Ocean soundscapes reveal climatic and economic oscillations

October 2024

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10 Reads

The Journal of the Acoustical Society of America

An accumulation of 26 859 days of acoustic recordings (spanning over 74 years at six sites) was processed and analyzed to investigate the interannual, seasonal, and diel patterns within six ocean soundscapes. Biological sound levels (18-25 Hz and 40–45 Hz) quantified acoustic energy from fin and blue whales, ambient sound levels (63-Hz and 800-Hz one-third-octave) quantified energy from ships and wind, and broadband spectra (15 Hz to 1 kHz) identified changes across time and space. Biological sound levels varied seasonally and correlated with large-scale climatic patterns and long-term ocean fluctuations. During marine heatwaves, baleen whale associated sound levels decreased in southern sites and increased in sites adjacent to the California Current. Ship sound levels at high-traffic sites reflected economic events such as the financial crisis, labor shortages and negotiations, and changes to port flows. Wind speeds and associated sound levels reflected on-shore/off-shore relationships and decreased during the morning hours. Understanding marine soundscapes aids in understanding the ocean’s ecological health amidst the ever-changing impact of climate change.


Sperm whale demographics in the Gulf of Alaska and Bering Sea/Aleutian Islands: An overlooked female habitat

July 2024

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152 Reads

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5 Citations

Sperm whales exhibit sexual dimorphism and sex-specific latitudinal segregation. Females and their young form social groups and are usually found in temperate and tropical latitudes, while males forage at higher latitudes. Historical whaling data and rare sightings of social groups in high latitude regions of the North Pacific, such as the Gulf of Alaska (GOA) and Bering Sea/Aleutian Islands (BSAI), suggest a more complex distribution than previously understood. Sperm whales are the most sighted and recorded cetacean in marine mammal surveys in these regions but capturing their demographic composition and habitat use has proven challenging. This study detects sperm whale presence using passive acoustic data from seven sites in the GOA and BSAI from 2010 to 2019. Differences in click characteristics between males and females (i.e., inter-click and inter-pulse interval) was used as a proxy for animal size/sex to derive time series of animal detections. Generalized additive models with generalized estimation equations demonstrate how spatiotemporal patterns differ between the sexes. Social groups were present at all recording sites with the largest relative proportion at two seamount sites in the GOA and an island site in the BSAI. We found that the seasonal patterns of presence varied for the sexes and between the sites. Male presence was highest in the summer and lowest in the winter, conversely, social group peak presence was in the winter for the BSAI and in the spring for the GOA region, with the lowest presence in the summer months. This study demonstrates that social groups are not restricted to lower latitudes and capture their present-day habitat use in the North Pacific. It highlights that sperm whale distribution is more complex than accounted for in management protocol and underscores the need for improved understanding of sperm whale demographic composition to better understand the impacts of increasing anthropogenic threats, particularly climate change.


Citations (58)


... The oil spill not only resulted in the immediate mortality of sperm whales (Williams et al. 2011) but also long-term health impacts from the 2.9 million l of chemical dispersants released in abundance after the spill (Kujawinski et al. 2011, J. P. Wise Sr. et al. 2011, C. F. Wise et al. 2014, J. P. Wise Jr. et al. 2014. In areas closest to the DWH well site, sperm whale densities, as estimated through passive acoustic monitoring, declined 24-31 % in the decade following the spill (Frasier et al. 2024). Models suggest that the overall GMx sperm whale population faced a decline of more than 10 % and will take up to 2 decades to recover (Marques et al. 2023). ...

Reference:

Change in matrilineal structure over time in an isolated population of sperm whales
A decade of declines in toothed whale densities following the Deepwater Horizon oil spill

... This included the southern DT site, where sperm whale densities increased, although densities there at the end of the monitoring period were still considerably lower than those observed at the northern sites, and lower than the more common species. A sperm whale demographic study over the 2010-2017 period suggests a greater increase in the occurrence of large, presumed transient males southern DT site than of matriarchal groups 34 . Density increases in the LF delphinid category require additional investigation, but explanations may include increased occupancy due to reduced competition or changes in prey composition 35 . ...

Accounting for sperm whale population demographics in density estimation using passive acoustic monitoring

Marine Ecology Progress Series

... Marine organisms are exposed to multiple noise sources simultaneously, making biologically relevant analyses important to understand the full spatiotemporal exposure of animals to noise in a region (ZoBell, 2023). With the understanding that noise reduction is achievable, it is important to investigate which approaches are most effective in reducing noise pollution in time and space. ...

Moving cargo, keeping whales: Investigating solutions for ocean noise pollution
  • Citing Article
  • March 2024

The Journal of the Acoustical Society of America

... months to years) regardless of oceanographic conditions. Analyses of such recordings can elucidate the seasonal distribution of sperm whales in a region [131], group sizes [132], demographic structure [133] and clan identity [32]. Recently developed sperm whale echolocation and coda detection software will dramatically reduce the time required to analyse large acoustic datasets [134], making the data analyses more feasible. ...

Sperm whale demographics in the Gulf of Alaska and Bering Sea/Aleutian Islands: An overlooked female habitat

... The study area covered a 220 Â 250 km grid encompassing latitude 33.26 N to 35.24 N and longitude 121.55 W to 118.83 W. This region includes three critical habitats: Channel Islands National Marine Sanctuary, Blue Whale BIA, and Humpback Whale BIA (Fig. 1). Sound pressure levels (SPLs) across this region were modeled using methods described in ZoBell et al. (2024), which are briefly outlined in the following. SPLs were modeled for pre-industrial and modern ocean noise as well as sourcecentric and space-centric simulations (Secs. ...

Comparing pre-industrial and modern ocean noise levels in the Santa Barbara Channel

Marine Pollution Bulletin

... southwestern parts of the Gulf of Mexico (Soldevilla, Debich, et al. 2022;Soldevilla et al. 2024). The rarity of the species and its open ocean environment make it difficult to robustly characterize its distribution and to obtain accurate abundance estimates, information that is critical for effective management and conservation strategies. ...

Rice's whale occurrence in the western Gulf of Mexico from passive acoustic recordings

Marine Mammal Science

... This redistribution of sampling effort may have influenced population trend assessments. Furthermore, the estimated population decline may also have been affected by the onset of La Niña oceanographic conditions in 2020, which continued through 2022, marking the longest consecutive stretch of negative Oceanic Niño Index since -2001(Thompson et al. 2024) and may have affected S. gigas behavior in nearshore waters. There was a La Niña event from 2016-2017 but it was considered weak (Wells et al. 2017). ...

State of the California Current Ecosystem report in 2022: a tale of two La Niñas

... In passive acoustic monitoring studies at fixed locations in Hawai'i, Ziegenhorn et al. (2023a,b) found that detections of Blainville's beaked whales Mesoplodon densirostris were significantly higher during La Niña periods at both sites studied (Manawai, Hawai'i Island), while goose-beaked whales Ziphius cavirostris had a similar relationship off Hawai'i Island and a weaker one at Manawai. An increase in detections for both species off of Hawai'i Island was documented during the 2010-2011 La Niña event, which corresponded with negative PDO and positive NPGO values (Ziegenhorn et al. 2023b). Therefore, the study hypothesized that a combination of the 3 events could lead to the most productive combination of these cycles for odontocetes in the main Hawaiian Islands. ...

Odontocete detections are linked to oceanographic conditions in the Hawaiian Archipelago

... Similarly, marine soundscapes study the effects of noise generated from shipping traffic, with noise pollution posing a significant threat to marine resources (Jones et al., 2023). In this case, underwater noise affects various coral reef organisms (Ferrier-Pagè s et al., 2021). ...

Impact of ship noise on the underwater soundscape of Eclipse Sound in the northeastern Canadian Arctic

Marine Pollution Bulletin

... Despite the paucity of winter density estimates, recordings collected year-round from static acoustic recorders suggest sperm whale are present off the eastern seaboard of the U.S. in all calendar months (Cohen et al., 2022;Kowarski et al., 2022). There is even evidence of a winter peak in occurrence off North Carolina (35 • N), with a subsequent peak in detections further north (37-40 • N) in spring (Stanistreet et al., 2018;Cohen et al., 2023). Winter research effort may therefore be viewed as a priority for refining sperm whale density estimates for the eastern seaboard of the U.S. ...

Spatial and temporal separation of toothed whales in the western North Atlantic

Marine Ecology Progress Series