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Global inventory of species
Data Descriptor categorized by known underwatersonifery
OPEN

Audrey Looby 1,2 ✉, Christine Erbe 3, Santiago Bravo 4, Kieran Cox5, Hailey L. Davies
Lucia Di Iorio 7, Youenn Jézéquel 8, Francis Juanes 6, Charles W. Martin 2,9,T. Aran Mooney8, Craig Radford 10, Laura K. Reynolds 11, Aaron N. Rice 12,
Amalis Riera 6, Rodney Rountree 6,13, Brittnie Spriel 6, Jenni Stanley 14,
Sarah Vela 15,16 & Miles J. G. Parsons 17

6,

A working group from the Global Library of Underwater Biological Sounds effort collaborated with theWorld 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 tocompile a dataset categorizing scientific knowledge of sonifery for 33,462 species and subspecies acrossmarine 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 ofunderwater soniferous species worldwide.

Background & Summary

Numerous aquatic and semi-aquatic species, including amphibians, annelids, cephalopods, cetaceans, crustaceans, fishes, pinnipeds, reptiles, and sirenians, use sound underwater for information acquisition and dissemination1–11. Sounds produced intentionally for the purposes of communication or navigation (i.e., active
sounds), and incidentally through other activities (i.e., passive sounds) may both serve numerous ecological functions5,12,13, such as for environmental sensing14, predator or prey detection15–17, and competition18,19.
Additionally, the remote sensing of these sounds through passive acoustic monitoring (i.e., an observational
method to detect and characterize sounds) has applications for invasive species detection20,21, habitat and species
management22,23, ecological monitoring24, and optimization of aquaculture operations25, among other possible
uses. Underwater sounds and their uses nonetheless face increasing threats from human activities and resulting
environmental degradation26–30.
1Fisheries and Aquatic Sciences, Institute of Food and Agricultural Sciences, University of Florida, Gainesville, FL,
USA. 2Nature Coast Biological Station, Institute of Food and Agricultural Sciences, University of Florida, Cedar
Key, FL, USA. 3Centre for Marine Science and Technology, Curtin University, Perth, WA, Australia. 4Instituto
Oceanográfico, Universidade de São Paulo, São Paulo, SP, Brasil. 5Department of Biological Sciences, Simon Fraser
University, Burnaby, BC, Canada. 6Department of Biology, University of Victoria, Victoria, BC, Canada. 7Centre deFormation et de Recherche sur les Environnements Méditerranéens, CNRS, Université de Perpignan Via Domitia,
Perpignan, France. 8Biology Department, Woods Hole Oceanographic Institution, Woods Hole, MA, USA. 9Stokes
School of Marine and Environmental Sciences, University of South Alabama and Dauphin Island Sea Lab, Dauphin
Island, AL, USA. 10Institute of Marine Science, Leigh Marine Laboratory, University of Auckland, Warkworth, New
Zealand. 11Soil, Water, and Ecosystem Sciences Department, Institute of Food and Agricultural Sciences, University
of Florida, Gainesville, FL, USA. 12K. Lisa Yang Center for Conservation Bioacoustics, Cornell Lab of Ornithology,
Cornell University, Ithaca, NY, USA. 13The Fish Listener, Waquoit, MA, USA. 14Coastal Marine Field Station, School of
Science, University of Waikato, Tauranga, New Zealand. 15MERIDIAN, Halifax, NS, Canada. 16Dalhousie University,
Halifax, NS, Canada. 17Australian Institute of Marine Science, Perth, WA, Australia. ✉e-mail: alooby101@gmail.com
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Despite decades of research, documentations of underwater sonifery across different taxa remain highly variable. For example, the majority of whales, dolphins, and porpoises have been confirmed to produce sound31. In
contrast, fishes and underwater invertebrates have been relatively less studied when considering the thousandsof extant species5,32,33. There have been several recent efforts to review sound production information for larger
taxonomic groups globally (e.g., marine mammals31, marine invertebrates33, fishes34), but these have varied intheir approaches and definitions of sonifery. As such, a comprehensive inventory of all known underwater soniferous species remains lacking, limiting the ability to document, summarize, or quantify trends in global underwater sound production35.
To address previous data limitations, the effort to create a Global Library of Underwater Biological Sounds(GLUBS)35 in collaboration with the World Register of Marine Species (WoRMS) created a dataset of information on likely or documented underwater sonifery in aquatic species. The World Register of Marine Species(available at MarineSpecies.org) provides a comprehensive, searchable database of known marine organisms
and other taxa, featuring taxonomic information—including numerous accepted, synonymized, misspelled, andunaccepted classifications—along with a growing list of ecological attributes and traits36–38. Through its own
datasets and connections or associations with other global datasets (e.g., the Ocean Biogeographic Information
System, OBIS), the data presented on WoRMS can be used to answer a variety of questions related to distribution, evolution, ecology, management, conservation, and other fields worldwide36. The WoRMS database is
widely used among researchers, with consistent growth in visitors since its inception in 2007, and at least 7,000
publications that have cited or mentioned WoRMS and its related registers37,39. As part of ongoing initiatives
related to the United Nations Decade of Ocean Science for Sustainable Development (2021–2030) and Aboveand Beyond - Completing the World Register of Marine Species (ABC WoRMS), WoRMS is encouraging the
documentation of species traits to better aid ecological research and provide more comprehensive information
to its diverse user base40. Trait information on underwater sonifery by aquatic and semi-aquatic species is inservice of these goals.
Here, we present a global inventory of 33,462 aquatic and semi-aquatic species and subspecies categorizedbased on known underwater sound production. To create this dataset, we integrated existing sources for fishes,
mysticetes, and odontocetes, and we conducted additional literature surveys for other semi-aquatic and aquatictaxa (Fig. 1). This trait has six categories according to whether a species has been confirmed or is considered
likely to produce sound based on the available scientific literature (Table 1) following established definitionsof active and passive sound production in natural and unnatural conditions, among other terms (Table 2).
We found 729 species documented as producing active and/or passive sounds under natural conditions, withanother 21,911 species deemed likely to produce sounds based on evaluated taxonomic relationships (Fig. 2).
This dataset is available in the figshare data repository and WoRMS.

Methods

Trait and category definitions. The ecological trait, ‘species exhibits underwater soniferous behavior,’ andits respective categories were defined by a working group of seven scientists, all experts in bioacoustics and theresearch literature of a variety of aquatic taxa including mammals, fishes, and invertebrates. The various definitions were largely adapted from other recent works (e.g., ancestral state reconstruction41; active sounds5). We also
considered whether sounds were produced naturally, based on long-standing concerns of artificial instigation
creating unnatural or involuntary sound production behaviors42–44. We defined sounds produced under natural
conditions as those made in the absence of artificial manipulation, such as an electrical current or direct handling, but may have included sounds made in captive environments, such as tanks. The resulting table of the trait,categories, and definitions were then reviewed and approved by the WoRMS Data Management Team. The trait
and category definitions created are listed in Table 1 and additional definitions of terms are provided in Table 2.
Data collection.

Our data collection efforts centered on extant species and subspecies as listed on WoRMS.
We separated taxa into four groupings for the purposes of our data collection. These groupings allowed us tobetter respond to the wide disparities in species richness, numbers of publications on sound production, and theexistence of soniferous species reviews and inventories for the different taxa (Fig. 1).

Marine mammals. In this context, ‘marine mammals’ refers to fully aquatic marine mammal species and subspecies within the infraorder Cetacea or order Sirenia. All other mammal species for this dataset were examinedas ‘other tetrapods.’ We used WoRMS38 as our taxonomic authority and queried the WoRMS database for a listof all accepted, extant marine mammal species and subspecies to base our data collection on, encompassing 148
species and subspecies.
Marine mammal ecological trait categorizations were largely based on work conducted for an upcoming
Springer book on Marine Mammal Bioacoustics by Erbe and colleagues, which will provide an overview ofsound production in marine mammals. There are many hundreds of publications on the active sounds producedby marine mammals, and their sounds have been summarized in several reviews on specific species groups orregions (e.g., Erbe et al.45). Therefore, it was not necessary to undertake a comprehensive literature search forevery species. Rather, for each species and subspecies, we aimed to cite a recent review or summary of its sounds;a publication that documented a great variety of sounds; or one of the original or earliest publications of its
sounds—in this order. As WoRMS somewhat differed from other marine mammal taxonomic authorities, suchas the Society of Marine Mammalogy46, and the taxonomy associated with marine mammals is in ongoing flux,
we also provided geographic areas associated with each species or subspecies based on the publications cited. We
only focused on marine mammal active sound production for this review.
There were 19 species and subspecies—mostly offshore, cryptic species, such as some beaked whales (e.g.,
Berardius minimus)—whose sounds were not reported in available reviews or summaries. For these species, we
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Fig. 1 Conceptual diagram of the data collection methods used to create a dataset of species categorized by
known sonifery. For the marine mammals, other tetrapods, and fishes, species and subspecies in each taxa
were established based on the World Register of Marine Species (WoRMS) database, which were subsequently
categorized based on existing sources and additional online literature searching. For invertebrates, online
literature searching identified species that had been studied for their sound production, which were thenmatched to their appropriate listing on the WoRMS database. The resulting dataset was then published on
WoRMS and figshare.

searched Web of Science47 and Google Scholar48 for reports on their sound production in the peer-reviewed andgrey literature. No records of their sounds were found. Based on the documented sounds for species within thesame genera or families of these species, all these species with undocumented sound production were deemed‘likely to produce sound under natural conditions, but unconfirmed’ (i.e., Category 3).
Other tetrapods. For the purposes of this dataset, ‘other tetrapods’ can be summarized as aquatic andsemi-aquatic species and subspecies that occur in the following taxa groups: amphibians, hippopotamids,
murids, mustelids, penguins, pinnipeds, ursids, and reptiles including crocodilians, lacertids, serpents, and testudines. For each of these groups of taxa, we queried the WoRMS38 database for a list of all accepted, extant
marine species and subspecies. Though marine was specified, WoRMS defines ‘marine’ generously, so our taxa
also included species that may predominantly live in and around brackish or fresh water or that may only besemi-aquatic (e.g., Alligator mississippiensis, Hippopotamus amphibius, Rana aurora).
Using Google Scholar48 and Scopus49, for each species and subspecies, we searched for papers that included“[scientific name]” OR “[common name]” AND “sound” OR “call” OR “acoustic” OR “vocal” OR “noise” in thetitle or keywords. We confirmed that papers within the search list referred to underwater sound production of
the species in question and evaluated the content using the categories of sound production detailed in Table 1.
For each species and subspecies, when applicable, the oldest reference that identified sound production undernatural conditions was provided as the source of information for the WoRMS database. On occasion, a latersource was retained, instead, if it had more information on the repertoire or behavior of the species than theoldest reference. Papers that provided information on an alternative or additional species outside of the list produced from WoRMS were retained for our dataset and added to the WoRMS database.
There has been no formal assessment (e.g., evolutionary analysis) of the likelihood of underwater soundproduction for these taxa. Additionally, for these taxa, few studies have been conducted where researchers haveattempted to elicit underwater sound production. Therefore, for species that did not have a report of underwatersound production testing, the following actions were taken. If multiple species within the same family were
found to produce sound, the species in question was deemed likely to produce sound under natural conditions.

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Identifier

Presented information/field

Definition

Trait

Species exhibits underwater soniferous behavior

Does the species actively or passively produce sound under natural
conditions while submerged in water?

Category 1

Unknown or undetermined

There is no known acoustic, morphological, or physiological studyof sound production for this species and its likelihood of soniferous
behavior based on an ancestral state reconstruction analysis, lineage, or
evolutionary records has not been assessed, or this species has yet to be
categorized by an expert source.

Does not or is unlikely to produce sound undernatural conditions

Species has been the subject of one or more studies to assess its acoustic
behavior, morphology, and/or physiology, none of which have provided
evidence of sound production either actively or passively, and/or species
does not have a likelihood of soniferous behavior based on an ancestral
state reconstruction analysis that results in a probability below 0.5,
lineage, or evolutionary records.

Category 3

Likely to produce sound under natural conditions
but unconfirmed

There is no known acoustic documentation to confidently validate
sound production under natural conditions by this species, but it islikely to exhibit natural soniferous behavior based on an ancestral state
reconstruction analysis that results in a probability of 0.5 or higher,
lineage, evolutionary records, morphological characteristics, physiologicalcharacteristics, documented sound production behavior under artificial
conditions, and/or documented sound production behavior with someuncertainty.

Category 4

Produces passive sound under natural conditions

There is validated acoustic documentation that this species producespassive sound under natural conditions, but no validated documentation
to confirm that it actively produces sound.

Category 5

Produces active sound under natural conditions

There is validated acoustic documentation that this species produces
active sound under natural conditions, but no validated documentation toconfirm that it passively produces sound.

Category 6

Produces active and passive sound under natural
conditions

There is validated acoustic documentation that this species produces
active and passive sound under natural conditions.

Category 2

Table 1. Trait categories and their associated definitions used to describe soniferous behavior on the World
Register of Marine Species.

For example, most pinnipeds have reports of underwater sound production, but not all, and those that do nothave associated reports have not been fully investigated for this behavior. As such, it was deemed likely that all
pinniped species produce sound underwater, which can be confirmed when further investigations take place.
Similarly, while few penguins reportedly exhibit this trait, the ones that have been studied have produced active
sound underwater. Thus, all penguin species without an associated report have currently been assigned toCategory 3, as well. For taxa where studies and reports of sound production are few (e.g., crocodilians, mustelids, rodents), species that did not have an associated report of sound production were assigned to Category 1(i.e., unknown or unconfirmed).
Fishes. For the purposes of this dataset, fishes were defined as any extant, accepted species (not only marine)in the Subphylum Vertebrata, except for any species in the Megaclass Tetrapoda, as listed on WoRMS38. This
includes the taxa Agnatha, Chondrichthyes, Sarcopterygii, and Actinopterygii. A complete list of fish species
for use in the data collection was downloaded from WoRMS using either the WoRMS website or the R packages
taxize50 and worms51.
The categorizations for fishes were adapted from two existing data sources: FishSounds5,34,52 and Rice et al.53.
Both data sources used different criteria from the sonifery categories defined for WoRMS to determine the likelihood of sonifery in fishes. Nonetheless, they represent comprehensive resources of scientific knowledge about
soniferous fish diversity and therefore allowed us to circumvent the need to re-review the full scope of literature
on sound production in fishes, which comprise roughly 35,000 extant species54 and over 1,000 species that havebeen studied for sound production across more than 800 publications55.
The FishSounds website (available at FishSounds.net) presents a global inventory of fish species examined
for sound production in the scientific literature. For each species studied in each reference, FishSounds reports
whether the species has been found to produce sound, the sound production type as active and/or passive (asdefined in Table 2), the varying methodologies used to examine the species for sound production, and thepossibility for uncertainty (see Looby et al.5 for a complete description of the methods). The FishSounds dataset is therefore different in its sonifery determinations than the WoRMS categorizations described herein, but
it provided an effective starting point for our data collection. The FishSounds dataset was retrieved from the
FishSounds permanent data repository52. As the FishSounds dataset uses FishBase as its taxonomic source54,56,
the species in the FishSounds dataset were matched to their respective accepted species entries on WoRMS using
its Match Taxa tool38.
Using the FishSounds dataset, species were re-assigned to different WoRMS categories either automatedly
or following a re-review process. If a species had been examined morphophysiologically or auditorily for soundproduction as defined by Looby et al.5 and found to be soniferous but with some amount of doubt in the authors’conclusions or without visual confirmation of an auditory examination, then the species was placed in Category
3 and the FishSounds dataset was cited. To place fishes into Categories 4, 5, and 6 (Table 1), the FishSounds
dataset was filtered to produce a list of species and their associated references that had examined the fish species
for sound production auditorily with visual confirmation and had found the species to produce active and/or

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Fig. 2 Bar graphs showing the number of aquatic and semi-aquatic species and subspecies placed in eachsoniferous trait category. The panels separate each taxa grouping, with images included to represent them.

passive sounds, without doubt or uncertainty associated with their species identification, whether sounds were
produced, nor the type of sound produced—all as determined following Looby et al.5 The references in the
resulting list were then reviewed to determine whether the fish species produced the reported sounds undernatural conditions. A total of 819 species studied across 613 references were assessed this way.
For the species that needed to be re-reviewed, if sound production was reported as naturally occurring in atleast one reference in the FishSounds dataset, those species were placed in the appropriate Category 4, 5, or 6,
depending on the types of sounds produced as reported in the references. If not, they were placed in Category 3.
Because of the widespread use of direct manipulation to elicit fish sound production in the scientific literatureacross time and taxa42–44,57–62, authors within each reference had to explicitly specify that sound production
occurred without direct human manipulation to be considered natural. One or two references that documented
sound production were provided for each of the species that were reviewed with the methods described in thisand the previous paragraph. Not all references that studied a particular fish species for sound production werereviewed in this data collection effort, as in some cases a single reference was sufficient to determine a species
belonged in Category 4, 5, or 6. Additionally, the references cited for each category are not necessarily the oldest
or most comprehensive descriptors of sound production behaviors—they may have just been the first reviewedthat reported natural sound production. We recommend viewing the original FishSounds dataset for a moredetailed overview of the sound production studies conducted for each species52.
Rice et al.53 conducted an ancestral state reconstruction analysis of sound production in Actinopterygii basedon family-level sonifery data taken from the literature. Determinations made by Rice et al.53 of ancestral-state
sonifery at the family level were extracted from their Supplemental Figure S1. The families were then used todetermine whether species were likely to be soniferous (Category 3) or not likely to be soniferous (Category 2)
based on these taxonomic relationships, for any species not covered by the FishSounds dataset. Such categorizations cited the Rice et al.53 publication as their determining source.
Species that were not included in the Rice et al.53 dataset and were examined but not found to produce soundin the FishSounds dataset were placed in Category 2 and the FishSounds dataset was cited. While it is generally
assumed that all fish species are capable of making passive sound under certain conditions13, we did not include
such an assumption in our categorizations as it remains unclear whether all fishes are likely to produce passive
sounds naturally. Species that were not included in either dataset were placed in Category 1 and the FishSounds
dataset was cited. Any fish species not listed in the provided dataset should be considered Category 1, as they
were either absent from WoRMS, not listed as accepted species on WoRMS at the time of the review, or not covered under the definition of a fish species listed above.
Invertebrates. Since invertebrate species are so numerous with few that have been studied for their sound production, we only included species in our dataset for which we could find relevant publications. Any other species
not listed in the dataset should be considered in Category 1 (unknown or undetermined) until further data canbe collected or additional research is done.
We conducted both targeted and haphazard searches to compile data on invertebrate underwater soundproduction. We focused on several taxonomic groups that are more well-studied for their sound production. For
crustaceans, we used the Web of Science Core Collection database47 and searched using the string: “TS = (crustacean*) AND ((TS = ((sound* OR noise* OR *acoustic* OR vocal*) NEAR/2 (mak* OR made OR produc*
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Term

Definition

Active soundProduced deliberately in association with a particular behavior or situation, frequently withspecialized sonic organs or structures, and generally used for communication; also referred toas intentional, deliberate, or specialized sounds.

Passive soundMay not be associated with specialized sound-producing structures nor with specific behaviors
or situations, though may still serve some signal function; also referred to as incidental,
unspecialized, or mechanical sounds.

Natural conditions

In the absence of artificial manipulation, such as an electrical current or direct handling.

Acoustic study or documentation

Documentation in the peer-reviewed or grey scientific literature of the presence or absence ofaudible sounds through listening live or on a recording in captivity or the field.

Morphological or physiological study or documentation

Documentation in the peer-reviewed or grey scientific literature of the presence or absence ofmorphological and/or physiological structures with sound production functions.

Ancestral state reconstruction analysis

The extrapolation back in time from measured characteristics of individuals to their common
ancestors using statistical techniques to, in this case, recover information on the likelihood ofsoniferous behavior.

Artificial conditions

Involving human intervention or manipulation, such as an electrical current or direct handling.

Uncertainty

Doubt or uncertainty expressed by researchers conducting examinations for sound production
about their conclusions (e.g., that the species they examined was correctly identified, that theycorrectly identified active versus passive sounds).

Validated

The researchers’ conclusions have no associated uncertainty in the documentation (see‘Uncertainty’).

Table 2. Terms and their associated definitions used to describe soniferous behavior on the World Register ofMarine Species. See Looby et al.5, Joy et al.41 and Rice et al.53 for additional information.

OR emit* OR communicat* OR call*))) OR (TS = (soniferous)))” following Looby et al.5 For mollusks, wesearched by each class and group within the phylum (e.g., cephalopods) in Google Scholar using “[class name]”
OR “[group name]” AND “sound” OR “call” OR “acoustic” OR “vocal” OR “soniferous”. We also searched haphazardly in Web of Science47, Google Scholar48, and personal reference libraries for any other reports of underwater sound production in these or other invertebrate taxa. Each publication found was read to determine its
applicability to the data collection. The species studied were matched to their associated listing on WoRMS38 andplaced in their appropriate categories accordingly.

Data Records

The known sonifery data are available in the figshare data repository in a single Excel spreadsheet file63. The
spreadsheet contains three tabs: a data dictionary, the reference information, and the sonifery information. The
data dictionary provides information defining each column variable present in the other two spreadsheet tabs,
including the variable name, allowed values, variable definition, and any needed additional information. The
reference information provides the citation information (e.g., authors, publication name) for all the referencesused to categorize species by known underwater sonifery. The DOIs of the associated references were also provided whenever available to aid in locating the references. The sonifery information contains the taxon names,
the soniferous categories assigned to them, and any notes that were included about the categorizations. To relate
which reference was cited for each soniferous trait record, the reference information can be linked to the sonifery
information through the provided Reference Aphia IDs. The reference IDs were kept consistent with the Aphia
ID system utilized by WoRMS but can also be considered independently from WoRMS if using this dataset as astand-alone source. As scientific names may change with future taxonomic reclassifications, to improve dataset
sustainability, for each taxon, we also provided the Taxon Aphia ID as listed in WoRMS, though this may beancillary or unnecessary to the dataset’s use. The dataset as provided on figshare has also been integrated into
WoRMS, where it can be more readily updated with new soniferous information or taxonomic reclassifications.
Any R code and associated data files used to generate or summarize the dataset are also provided in figshare63.

Technical Validation

Data were collected by experts in their respective soniferous taxa using either existing data sources or throughnovel searches of the existing literature using systematized review methodology when possible64. The data collected were verified against other similar reviews65 or estimations of known soniferous species numbers66–68
when available to ensure the results provided herein were reasonable. Most of the data collected for the different animal groups identified in the methods were categorized by a single reviewer each, however, with limited
external validation at the time of publishing. This may have introduced omissions or mistakes in the dataset
based on human error or biases associated with the review methods. The WoRMS database allows for continual
editing and so the authors, as WoRMS thematic experts, will continue to update and validate the data as new
information becomes available or any corrections are needed. Any users of the compiled dataset should be awareof the possibility of errors and ongoing changes. Users are encouraged to verify any specific reports of soniferyusing the references provided and to contact the authors of this data publication if any discrepancies are found.
As both active and passive sounds have potential ecological signaling and monitoring applications5,12–25, thedataset attempted to capture both types of sonifery in its categories. The definitions provided for these soundtypes in Table 2 were adapted from previously published text and commonly used delineations in the scientific literature that the dataset was meant to encompass5,12,13,31,44,. The distinction between passive and active
sounds, however, may exist more along a spectrum as opposed to in stark contrasts5. For example, active soundproduction may be the result of exaptation in taxa where it is not an ancestral trait69, and passive sounds may
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be associated with particular behaviors or situations5,70. Determining the use of sounds for intentional communication can often be uncertain and may require further behavioral testing to designate conclusively5,71–73.
The dataset’s categorizations also relied on reviewers’ interpretations of sound production descriptions in thesurveyed references. This process may have simplified more nuanced sonifery descriptions in the source materials. Users are therefore highly encouraged to refer to the provided data sources for additional information on
the context surrounding the documented sound production. Because likely many species are capable of passive
sound production in certain contexts (e.g., through swimming), species were only categorized as passively soniferous in the dataset if their sounds were detectable in a natural context within a scientific study. We also did notinclude reports of passive sounds for the tetrapods in our data collection, though there are likely many species
that produce them31,71.
Users should be aware of potential limitations or qualifications associated with our categorizations. While
our trait and category definitions relied on widely used and established differentiations in the sound production
literature, they were relatively conservative in their inclusion of species in Categories 4, 5, and 6 and may therefore have led to the exclusion of species that are indeed naturally soniferous. For example, for the purposes ofour data collection, we emphasized species that were shown to produce sounds underwater naturally without
contact, electricity, tethering, or other forms of direct manipulation by humans, while still including sounds
spontaneously produced in artificial environments (e.g., tanks). This may have led to the exclusion of species
that do indeed produce sounds naturally, such as species who may make distress sounds in response to beingcaught by predators or those that readily produce sounds when handled but may have simply not yet been shownto do so in other agonistic situations7,43,59,74–76. Due to frequent uncertainty in studies of underwater sound production5, species may have been similarly excluded from Categories 4, 5, or 6 if their potential sonifery could
not be confirmed in the surveyed studies. Additionally, for some taxa, we relied on ancestral state reconstruction
analyses or assumptions of sonifery based on taxonomic relationships to determine species likely to be soniferous. There is, nonetheless, the possibility for sound production behaviors to be secondarily lost, thus the species
deemed likely soniferous should be treated with caution53,77–80. Users should be aware of the assumptions andsimplifications made by our categorizations, the potential for our understanding of sound production and likely
sound production to change, and that the data may later be updated to reflect new research even if the categories
themselves may not change.

Usage Notes

We envision numerous possible applications for the dataset described herein to study sound production andother topics related to aquatic and semi-aquatic species worldwide. By listing a soniferous behavior trait on
WoRMS, researchers will be able to easily search for and access known sonifery information for any species
encompassed by our review. This will aid in identifying both known soniferous species and species lacking
documented sound production. In addition to being listed on WoRMS, the soniferous trait data associated withthis review is available in a spreadsheet file in figshare that may be easier to use in analyses. The ecological trait
data can be easily integrated with other information available on WoRMS itself (e.g., taxonomy, environment)or its associated databases (e.g., conservation status, distribution, invasive status) to study global patterns ofsound production behavior and the use of passive acoustics for ecological monitoring. Sonifery has been studied in these contexts before, but only with more limited datasets5,53,65. Researchers interested in conducting
such analyses should be aware of the availability of software packages that could be used to easily access related
datasets (e.g., the rfishbase56, taxize50, worms51, and rinat81 packages in R). We hope our data will facilitate future
research on the distribution, evolution, ecology, management, and conservation of underwater soniferous species worldwide.

Code availability

Code and associated data files are available in the figshare data repository63.
Received: 3 July 2023; Accepted: 13 November 2023;
Published: xx xx xxxx

References

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https://doi.org/10.1098/rstb.2000.0685 (2000).
2. Ladich, F. & Winkler, H. Acoustic communication in terrestrial and aquatic vertebrates. J. Exp. Biol. 220, 2306–2317, https://doi.
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Acknowledgements

The conception of this project originated within the working group for a Global Library of Underwater BiologicalSounds (GLUBS), which includes a number of the co-authors of this dataset. The GLUBS working group is
supported by the Scientific Committee on Oceanic Research, Monmouth University Urban Coast Institute,and Rockefeller Program for the Human Environment. We thank them for their initiative and support. The
Richard Lounsbery Foundation has provided financial support for the continued development of this working
group. Thank you to the WoRMS Editorial Board and WoRMS Data Management Team, especially Leen
Vandepitte and Stefanie Dekeyzer, for allowing the inclusion of sonifery as a WoRMS trait and for their assistancein data definition and integration with WoRMS. Thank you also to Edward Urban for his assistance with the
WoRMS trait and category definitions. This work makes use of data and infrastructure provided by VLIZ andfunded by Research Foundation - Flanders (FWO) as part of the Belgian contribution to LifeWatch. The work of
the WoRMS Data Management Team is funded by FWO as part of the Belgian contribution to LifeWatch.

Author contributions

M.J.G.P. first conceived of the project. M.J.G.P. and A.L. coordinated with WoRMS. A.L., L.D.I., C.E., T.A.M.,
M.J.G.P., C.R., A.N.R. and J.S. drafted the data categorizations, with additional input from the other authors.
C.E. led the categorization of marine mammals. M.J.G.P. led the categorization of the other tetrapods. A.L. and
A.N.R. led the categorization of fishes with input from S.B., K.C., H.L.D., F.J., C.W.M., L.K.R., A.R., R.R., B.S.
and S.V. L.D.I., T.A.M., Y.J., C.R. and J.S. led the categorization of invertebrates. A.L. wrote the first outline of
the manuscript. All authors contributed to subsequent manuscript drafts and provided editorial feedback on the
manuscript.
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Competing interests

The authors declare no competing interests.

Additional information

Correspondence and requests for materials should be addressed to A.L.
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