Vibrational and acoustic communication in fishes: The overlooked overlap between the underwater vibroscape and soundscape.

Substrate-borne communication via mechanical waves is widespread throughout the animal kingdom but has not been intensively studied in fishes. Families such as the salmonids and sculpins have been documented to produce vibratory signals. However, it is likely that fish taxa on or close to the substrate that produce acoustic signals will also have a vibratory component to their signal due to their proximity to substrates and energy transfer between media. Fishes present an intriguing opportunity to study vibrational communication, particularly in the context of signal production and detection, detection range, and how vibratory signals may complement or replace acoustic signals. It is highly likely that the vibrational landscape, the vibroscape, is an important component of their sensory world, which certainly includes and overlaps with the soundscape. With the wide range of anthropogenic activities modifying underwater substrates, vibrational noise presents similar risks as acoustic noise pollution for fishes that depend on vibrational communication. However, in order to understand vibrational noise, more empirical studies are required to investigate the role of vibrations in the fish environment.

Sound production biomechanics in three-spined toadfish and potential functional consequences of swim bladder morphology in the Batrachoididaea).

The relationship between sound complexity and the underlying morphology and physiology of the vocal organ anatomy is a fundamental component in the evolution of acoustic communication, particularly for fishes. Among vertebrates, the mammalian larynx and avian syrinx are the best-studied vocal organs, and their ability to produce complex vocalizations has been modeled. The range and complexity of the sounds in mammalian lineages have been attributed, in part, to the bilateral nature of the vocal anatomy. Similarly, we hypothesize that the bipartite swim bladder of some species of toadfish (family Batrachoididae) is responsible for complex nonlinear characters of the multiple call types that they can produce, supported by nerve transection experiments. Here, we develop a low-dimensional coupled-oscillator model of the mechanics underlying sound production by the two halves of the swim bladder of the three-spined toadfish, Batrachomoeus trispinosus. Our model was able to replicate the nonlinear structure of both courtship and agonistic sounds. The results provide essential support for the hypothesis that fishes and tetrapods have converged in an evolutionary innovation for complex acoustic signaling, namely, a relatively simple bipartite mechanism dependent on sonic muscles contracting around a gas filled structure.

Evidence of Atlantic midshipman (Porichthys plectrodon) vocalizations from an unmanned surface vehicle in the U.S. South Atlantica).

An unmanned surface vehicle (USV; Wave Glider) was deployed to study the coastal soundscape in shallow (less than 30 m) coastal waters off the coast of Cape Canaveral, FL, in July 2020 and January 2022. These surveys documented temporal and seasonal trends in biological sounds across a variety of habitats within an 812-km2 survey area, including sand shoals, sand-mud plains, and natural hardbottom. Among a broader diversity of identifiable and unidentifiable fish choruses recorded during the survey, a distinct and previously unidentified fish chorus was recorded; corroborating evidence suggests it and other sounds with similar spectral properties may be produced by Atlantic midshipman. Putative Atlantic midshipman sounds included an agnostic grunt and a seasonal chorus of persistent hums that peaked 3 h after sunset in the summer survey. While Atlantic midshipman have been demonstrated to have well-developed sonic muscles on their swim bladder, their acoustic behavior has not been previously described. Our use of a mobile passive acoustic platform combined with bottom sampling of fish communities highlights an important opportunity to identify previously undocumented biological sound sources in coastal habitats.

Offshore wind energy development: Research priorities for sound and vibration effects on fishes and aquatic invertebrates.

There are substantial knowledge gaps regarding both the bioacoustics and the responses of animals to sounds associated with pre-construction, construction, and operations of offshore wind (OSW) energy development. A workgroup of the 2020 State of the Science Workshop on Wildlife and Offshore Wind Energy identified studies for the next five years to help stakeholders better understand potential cumulative biological impacts of sound and vibration to fishes and aquatic invertebrates as the OSW industry develops. The workgroup identified seven short-term priorities that include a mix of primary research and coordination efforts. Key research needs include the examination of animal displacement and other behavioral responses to sound, as well as hearing sensitivity studies related to particle motion, substrate vibration, and sound pressure. Other needs include: identification of priority taxa on which to focus research; standardization of methods; development of a long-term highly instrumented field site; and examination of sound mitigation options for fishes and aquatic invertebrates. Effective assessment of potential cumulative impacts of sound and vibration on fishes and aquatic invertebrates is currently precluded by these and other knowledge gaps. However, filling critical gaps in knowledge will improve our understanding of possible sound-related impacts of OSW energy development to populations and ecosystems.

Limited vocal compensation for elevated ambient noise in bearded seals: implications for an industrializing Arctic Ocean.

Vocalizing animals have several strategies to compensate for elevated ambient noise. These behaviours evolved under historical conditions, but compensation limits are quickly being reached in the Anthropocene. Acoustic communication is essential to male bearded seals that vocalize for courtship and defending territories. As Arctic sea ice declines, industrial activities and associated anthropogenic noise are likely to increase. Documenting how seals respond to noise and identifying naturally occurring behavioural thresholds would indicate either their resilience or vulnerability to changing soundscapes. We investigated whether male bearded seals modified call amplitudes in response to changing ambient noise levels. Vocalizing seals increased their call amplitudes until ambient noise levels reached an observable threshold, above which call source levels stopped increasing. The presence of a threshold indicates limited noise compensation for seals, which still renders them vulnerable to acoustic masking of vocal signals. This behavioural threshold and response to noise is critical for developing management plans for an industrializing Arctic.

Phenological changes in North Atlantic right whale habitat use in Massachusetts Bay.

The North Atlantic right whale (Eubalaena glacialis) is one of the world's most highly endangered baleen whales, with approximately 400-450 individuals remaining. Massachusetts Bay (MB) and Cape Cod Bay (CCB) together comprise one of seven areas in the Gulf of Maine where right whales seasonally congregate. Here, we report on acoustically detected presence of right whales in MB over a nearly 6 year period, July 2007-April 2013, a time of both rapid ocean warming throughout the Gulf of Maine and apparent changes in right whale migratory dynamics. We applied an automated detection algorithm to assess hourly presence of right whale "up-calls" in recordings from a 19-channel acoustic array covering approximately 4,000 km2 in MB. Over the survey, up-calls were detected in 95% of 8 day periods. In each year, as expected, we observed a "peak season" of elevated up-call detections in late winter and early spring corresponding to the season when right whales congregate to feed in CCB. However, we also saw an increase in right whale occurrence during time periods thought to be part of the "off-season." With the exception of 2009-2010, when acoustic presence was unusually low, the mean percent of hours in which up-calls were detected increased every year, both during the peak season (from 38% in 2008 to 70% in 2012), and during the summer-fall season (from 2% in 2007 to 13% in 2012). Over the entire study, the peak season start date varied between 17 January and 26 February. Changes in right whale phenology in MB likely reflect broadscale changes in habitat use in other areas within the species range. This study demonstrates the value of continuous long-term survey datasets to detect and quantify shifts in cetacean habitat use as environmental conditions change and the long-term continued survival of right whales remains uncertain.

Exploring movement patterns and changing distributions of baleen whales in the western North Atlantic using a decade of passive acoustic data.

Six baleen whale species are found in the temperate western North Atlantic Ocean, with limited information existing on the distribution and movement patterns for most. There is mounting evidence of distributional shifts in many species, including marine mammals, likely because of climate-driven changes in ocean temperature and circulation. Previous acoustic studies examined the occurrence of minke (Balaenoptera acutorostrata) and North Atlantic right whales (NARW; Eubalaena glacialis). This study assesses the acoustic presence of humpback (Megaptera novaeangliae), sei (B. borealis), fin (B. physalus), and blue whales (B. musculus) over a decade, based on daily detections of their vocalizations. Data collected from 2004 to 2014 on 281 bottom-mounted recorders, totaling 35,033 days, were processed using automated detection software and screened for each species' presence. A published study on NARW acoustics revealed significant changes in occurrence patterns between the periods of 2004-2010 and 2011-2014; therefore, these same time periods were examined here. All four species were present from the Southeast United States to Greenland; humpback whales were also present in the Caribbean. All species occurred throughout all regions in the winter, suggesting that baleen whales are widely distributed during these months. Each of the species showed significant changes in acoustic occurrence after 2010. Similar to NARWs, sei whales had higher acoustic occurrence in mid-Atlantic regions after 2010. Fin, blue, and sei whales were more frequently detected in the northern latitudes of the study area after 2010. Despite this general northward shift, all four species were detected less on the Scotian Shelf area after 2010, matching documented shifts in prey availability in this region. A decade of acoustic observations have shown important distributional changes over the range of baleen whales, mirroring known climatic shifts and identifying new habitats that will require further protection from anthropogenic threats like fixed fishing gear, shipping, and noise pollution.

Dolphins simplify their vocal calls in response to increased ambient noise.

Ocean noise varies spatially and temporally and is driven by natural and anthropogenic processes. Increased ambient noise levels can cause signal masking and communication impairment, affecting fitness and recruitment success. However, the effects of increasing ambient noise levels on marine species, such as marine mammals that primarily rely on sound for communication, are not well understood. We investigated the effects of concurrent ambient noise levels on social whistle calls produced by bottlenose dolphins (Tursiops truncatus) in the western North Atlantic. Elevated ambient noise levels were mainly caused by ship noise. Increases in ship noise, both within and below the dolphins' call bandwidth, resulted in higher dolphin whistle frequencies and a reduction in whistle contour complexity, an acoustic feature associated with individual identification. Consequently, the noise-induced simplification of dolphin whistles may reduce the information content in these acoustic signals and decrease effective communication, parent-offspring proximity or group cohesion.

Validating automated click detector dolphin detection rates and investigating factors affecting performance.

Passive acoustic monitoring (PAM) is a widely used technique for studying the distribution and habitat use of cetaceans. The C-POD, an acoustic sensor with an onboard automated click detector, has been deployed in diverse acoustic environments, but studies verifying its offshore detection rates and factors affecting detection probability are scarce. To empirically evaluate the performance of C-PODs in detecting bottlenose dolphins (Tursiops truncatus), C-PODs were deployed alongside archival acoustic recorders 12-30 km offshore in the Northwest Atlantic Ocean. The C-POD and acoustic recordings, post-processed using PAMGUARD software, were compared for a period of 6852 h. C-POD false positive rates were very low (mean 0.003%), and positive hourly detection accuracy was very high (mean 99.6%). Analysis of the acoustic environment and dolphin click characteristics revealed that true positive detections by C-PODs were significantly more likely to occur when PAMGUARD detected more clicks and there was increased high frequency noise (>20 kHz), likely from distant or unclassified clicks. C-PODs were found to be reliable indicators of dolphin presence at hourly or greater time scales. These results support the application of C-PODs in PAM studies that aim to investigate patterns of dolphin occurrence, such as those related to offshore windfarms.

The evolution of jaw protrusion mechanics is tightly coupled to bentho-pelagic divergence in damselfishes (Pomacentridae).

Most species-rich lineages of aquatic organisms have undergone divergence between forms that feed from the substrate (benthic feeding) and forms that feed from the water column (pelagic feeding). Changes in trophic niche are frequently accompanied by changes in skull mechanics, and multiple fish lineages have evolved highly specialized biomechanical configurations that allow them to protrude their upper jaws toward the prey during feeding. Damselfishes (family Pomacentridae) are an example of a species-rich lineage with multiple trophic morphologies and feeding ecologies. We sought to determine whether bentho-pelagic divergence in the damselfishes is tightly coupled to changes in jaw protrusion ability. Using high-speed video recordings and kinematic analysis, we examined feeding performance in 10 species that include three examples of convergence on herbivory, three examples of convergence on omnivory and two examples of convergence on planktivory. We also utilized morphometrics to characterize the feeding morphology of an additional 40 species that represent all 29 damselfish genera. Comparative phylogenetic analyses were then used to examine the evolution of trophic morphology and biomechanical performance. We find that pelagic-feeding damselfishes (planktivores) are strongly differentiated from extensively benthic-feeding species (omnivores and herbivores) by their jaw protrusion ability, upper jaw morphology and the functional integration of upper jaw protrusion with lower jaw abduction. Most aspects of cranial form and function that separate these two ecological groups have evolved in correlation with each other and the evolution of the functional morphology of feeding in damselfishes has involved repeated convergence in form, function and ecology.

High-Resolution Analysis of Seismic Air Gun Impulses and Their Reverberant Field as Contributors to an Acoustic Environment.

In September and October 2011, a seismic survey took place in Baffin Bay, Western Greenland, in close proximity to a marine protected area (MPA). As part of the mitigation effort, five bottom-mounted marine acoustic recording units (MARUs) collected data that were used for the purpose of measuring temporal and spectral features from each impulsive event, providing a high-resolution record of seismic reverberation persistent after the direct impulse. Results were compared with ambient-noise levels as computed after the seismic survey to evidence that as a consequence of a series of repeating seismic impulses, sustained elevated levels create the potential for masking.

Novel underwater soundscape: acoustic repertoire of plainfin midshipman fish.

Toadfishes are among the best-known groups of sound-producing (vocal) fishes and include species commonly known as toadfish and midshipman. Although midshipman have been the subject of extensive investigation of the neural mechanisms of vocalization, this is the first comprehensive, quantitative analysis of the spectro-temporal characters of their acoustic signals and one of the few for fishes in general. Field recordings of territorial, nest-guarding male midshipman during the breeding season identified a diverse vocal repertoire composed of three basic sound types that varied widely in duration, harmonic structure and degree of amplitude modulation (AM): 'hum', 'grunt' and 'growl'. Hum duration varied nearly 1000-fold, lasting for minutes at a time, with stable harmonic stacks and little envelope modulation throughout the sound. By contrast, grunts were brief, ~30-140 ms, broadband signals produced both in isolation and repetitively as a train of up to 200 at intervals of ~0.5-1.0 s. Growls were also produced alone or repetitively, but at variable intervals of the order of seconds with durations between those of grunts and hums, ranging 60-fold from ~200 ms to 12 s. Growls exhibited prominent harmonics with sudden shifts in pulse repetition rate and highly variable AM patterns, unlike the nearly constant AM of grunt trains and flat envelope of hums. Behavioral and neurophysiological studies support the hypothesis that each sound type's unique acoustic signature contributes to signal recognition mechanisms. Nocturnal production of these sounds against a background chorus dominated constantly for hours by a single sound type, the multi-harmonic hum, reveals a novel underwater soundscape for fish.

Potential Bryde's whale (Balaenoptera edeni) calls recorded in the northern Gulf of Mexico.

Several marine autonomous recording units (MARUs) were deployed in northeastern Gulf of Mexico from 2010­2012 to study the acoustic ecology of Bryde's whales (Balaenoptera edeni) following the Deepwater Horizon oil spill. However, the acoustic repertoire of this sub-population is poorly documented, presently limiting the efficacy of acoustic monitoring applications. Numerous stereotyped, low-frequency signals from a putative biological sound source were found throughout the recordings. Sounds fell into three categories distinguished by spectral and temporal properties. Multiple calls overlapped temporally on individual MARUs, suggesting that multiple sources produced these sounds. The basic features are similar to those from other mysticetes, but they differ from any previously published sounds. Since Bryde's whales are the most common mysticete in the Gulf and have previously been observed within the recording area on multiple occasions, it is likely that Bryde's whales are the most probable source of these sounds. These results potentially identify a suite of previously undocumented calls from Bryde's whales, which could facilitate future passive acoustic monitoring efforts to better understand the population dynamics and status of this sub-population.

Hearing the seasSing Like Fish: How Sound Rules Life Under Water Amorina Kingdon Crown, 2024. 336 pp.

Rich underwater soundscapes await those who listen closely.

Global inventory of species categorized by known underwater sonifery.

A working group from the Global Library of Underwater Biological Sounds effort collaborated with the World Register of Marine Species (WoRMS) to create an inventory of species confirmed or expected to produce sound underwater. We used several existing inventories and additional literature searches to compile a dataset categorizing scientific knowledge of sonifery for 33,462 species and subspecies across marine mammals, other tetrapods, fishes, and invertebrates. We found 729 species documented as producing active and/or passive sounds under natural conditions, with another 21,911 species deemed likely to produce sounds based on evaluated taxonomic relationships. The dataset is available on both figshare and WoRMS where it can be regularly updated as new information becomes available. The data can also be integrated with other databases (e.g., SeaLifeBase, Global Biodiversity Information Facility) to advance future research on the distribution, evolution, ecology, management, and conservation of underwater soniferous species worldwide.

Acoustically detected year-round presence of right whales in an urbanized migration corridor.

Species' conservation relies on understanding their seasonal habitats and migration routes. North Atlantic right whales (Eubalaena glacialis), listed as endangered under the U.S. Endangered Species Act, migrate from the southeastern U.S. coast to Cape Cod Bay, Massachusetts, a federally designated critical habitat, from February through May to feed. The whales then continue north across the Gulf of Maine to northern waters (e.g., Bay of Fundy). To enter Cape Cod Bay, right whales must traverse an area of dense shipping and fishing activity in Massachusetts Bay, where there are no mandatory regulations for the protection of right whales or management of their habitat. We used passive acoustic recordings of right whales collected in Massachusetts Bay from May 2007 through October 2010 to determine the annual spatial and temporal distribution of the whales and their calling activity. We detected right whales in the bay throughout the year, in contrast to results from visual surveys. Right whales were detected on at least 24% of days in each month, with the exception of June 2007, in which there were no detections. Averaged over all years, right whale calls were most abundant from February through May. During this period, calls were most frequent between 17:00 and 20:00 local time; no diel pattern was apparent in other months. The spatial distribution of the approximate locations of calling whales suggests they may use Massachusetts Bay as a conduit to Cape Cod Bay in the spring and as they move between the Gulf of Maine and waters to the south in September through December. Although it is unclear how dependent right whales are on the bay, the discovery of their widespread presence in Massachusetts Bay throughout the year suggests this region may need to be managed to reduce the probability of collisions with ships and entanglement in fishing gear.

Seasonal and geographical patterns of fin whale song in the western North Atlantic Ocean.

Male fin whales, Balaenoptera physalus, produce a song consisting of 20 Hz notes at regularly spaced time intervals. Previous studies identified regional differences in fin whale internote intervals (INI), but seasonal changes within populations have not been closely examined. To understand the patterns of fin whale song in the western North Atlantic, the seasonal abundance and acoustic features of fin whale song are measured from two years of archival passive acoustic recordings at two representative locations: Massachusetts Bay and New York Bight. Fin whale 20 Hz notes are detected on 99% of recorded days. In both regions, INI varies significantly throughout the year as two distinct periods: a "short-INI" season in September-January (9.6 s) and a "long-INI" season in March-May (15.1 s). February and June-August are transitional-INI months, with higher variability. Note abundance decreases with increasing INI, where note abundance is significantly lower in April-August than in September-January. Short-INI and high note abundance correspond to the fin whale reproductive season. The temporal variability of INI may be a mechanism by which fin whale individuals encode and communicate a variety of behaviorally relevant information.

Nonlinear acoustic complexity in a fish 'two-voice' system.

Acoustic signals play essential roles in social communication and show a strong selection for novel morphologies leading to increased call complexity in many taxa. Among vertebrates, repeated innovations in the larynges of frogs and mammals and the syrinx of songbirds have enhanced the spectro-temporal content, and hence the diversity of vocalizations. This acoustic diversification includes nonlinear characteristics that expand frequency profiles beyond the traditional categorization of harmonic and broadband calls. Fishes have remained a notable exception to evidence for such acoustic innovations among vertebrates, despite their being the largest group of living vertebrates that also exhibit widespread evolution of sound production. Here, we combine rigorous acoustic and mathematical analyses with experimental silencing of the vocal motor system to show how a novel swim bladder mechanism in a toadfish enables it to generate calls exhibiting nonlinearities like those found among frogs, birds and mammals, including primates. By showing that fishes have evolved nonlinear acoustic signalling like all other major lineages of vocal vertebrates, these results suggest strong selection pressure favouring this mechanism to enrich the spectro-temporal content and complexity of vocal signals.

Differential geographic patterns in song components of male Albert's lyrebirds.

Geographic variation in bird song has received much attention in evolutionary studies, yet few consider components within songs that may be subject to different constraints and follow different evolutionary trajectories. Here, we quantify patterns of geographic variation in the socially transmitted "whistle" song of Albert's lyrebirds (Menura alberti), an oscine passerine renowned for its remarkable vocal abilities. Albert's lyrebirds are confined to narrow stretches of suitable habitat in Australia, allowing us to map likely paths of cultural transmission using a species distribution model and least cost paths. We use quantitative methods to divide the songs into three components present in all study populations: the introductory elements, the song body, and the final element. We compare geographic separation between populations with variation in these components as well as the full song. All populations were distinguishable by song, and songs varied according to the geographic distance between populations. However, within songs, only the introductory elements and song body could be used to distinguish among populations. The song body and final element changed with distance, but the introductory elements varied independently of geographic separation. These differing geographic patterns of within-song variation are unexpected, given that the whistle song components are always produced in the same sequence and may be perceived as a temporally discrete unit. Knowledge of such spatial patterns of within-song variation enables further work to determine possible selective pressures and constraints acting on each song component and provides spatially explicit targets for preserving cultural diversity. As such, our study highlights the importance for science and conservation of investigating spatial patterns within seemingly discrete behavioral traits at multiple levels of organization.

Comparative bioacoustics: a roadmap for quantifying and comparing animal sounds across diverse taxa.

Animals produce a wide array of sounds with highly variable acoustic structures. It is possible to understand the causes and consequences of this variation across taxa with phylogenetic comparative analyses. Acoustic and evolutionary analyses are rapidly increasing in sophistication such that choosing appropriate acoustic and evolutionary approaches is increasingly difficult. However, the correct choice of analysis can have profound effects on output and evolutionary inferences. Here, we identify and address some of the challenges for this growing field by providing a roadmap for quantifying and comparing sound in a phylogenetic context for researchers with a broad range of scientific backgrounds. Sound, as a continuous, multidimensional trait can be particularly challenging to measure because it can be hard to identify variables that can be compared across taxa and it is also no small feat to process and analyse the resulting high-dimensional acoustic data using approaches that are appropriate for subsequent evolutionary analysis. Additionally, terminological inconsistencies and the role of learning in the development of acoustic traits need to be considered. Phylogenetic comparative analyses also have their own sets of caveats to consider. We provide a set of recommendations for delimiting acoustic signals into discrete, comparable acoustic units. We also present a three-stage workflow for extracting relevant acoustic data, including options for multivariate analyses and dimensionality reduction that is compatible with phylogenetic comparative analysis. We then summarize available phylogenetic comparative approaches and how they have been used in comparative bioacoustics, and address the limitations of comparative analyses with behavioural data. Lastly, we recommend how to apply these methods to acoustic data across a range of study systems. In this way, we provide an integrated framework to aid in quantitative analysis of cross-taxa variation in animal sounds for comparative phylogenetic analysis. In addition, we advocate the standardization of acoustic terminology across disciplines and taxa, adoption of automated methods for acoustic feature extraction, and establishment of strong data archival practices for acoustic recordings and data analyses. Combining such practices with our proposed workflow will greatly advance the reproducibility, biological interpretation, and longevity of comparative bioacoustic studies.