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Biodiversity Data Journal :
Forum Paper
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Corresponding author: Serge Gofas (sgofas@uma.es)
Academic editor: Christos Arvanitidis
Received: 29 Apr 2025 | Accepted: 11 Jul 2025 | Published: 31 Jul 2025
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Bailly N, Gofas S, Lonneville B (2025) A worldwide geographical scheme for recording the distribution of marine biota: proposal and call for feedback. Biodiversity Data Journal 13: e157371. https://doi.org/10.3897/BDJ.13.e157371
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This paper describes a project aimed at creating a worldwide set of polygons for recording marine distribution data, parallel to the current World Geographic Scheme for Recording Plant Distribution used on land. The countries’ Exclusive Economic Zones were either taken as recording units or subdivided according to Marine Ecosystems of the World or the IHO Limits of Oceans and Seas when appropriate; existing local schemes were adopted for Europe and Australia. A hierarchical set of five Level-1 units, 26 Level-2 units, 232 Level-3 units and 536 Level-4 units is presented for feedback and intended to be submitted as a standard to the Biodiversity Information Standards (TDWG). This project is expected to provide a means to instantly retrieve national checklists for any taxonomic group and also a valuable tool to handle imprecise country-level records from the old literature.
taxonomic databases, checklists, marine distributions, marine biota, biodiversity data, marine bioregions
More than thirty years ago, in the early stage of the Internet, the International Working Group on Taxonomic Databases (TDWG) recognised the need for an agreed system of geographical units at approximately “country” level and upwards for use in recording plant distributions. The deliberations over a period of three years of a committee involving taxonomists as well as applied botanists culminated in producing a standard World Geographic Scheme for Recording Plant Distributions (hereafter WGSRPD;
With an increasing wealth of data available in the marine realm, a similar standard is desirable for the sea. Geo-referenced data are accurately represented in such databases as the
With respect to land, two peculiarities of the world ocean make major differences. Firstly, it is a three-dimensional realm where the water column can span thousands of metres and harbours a vertical succession of thoroughly different biotic communities. The vertical distribution of these communities is driven by factors such as light penetration and the characteristics of water masses. In addition, the communities settled on the sea bottom (benthos) are radically different from those of the water column (plankton and nekton). On a map, those cannot be discriminated unless depth is added to geography. The other major difference with the terrestrial realm is that only ca. 42% of the world ocean surface is under the jurisdiction of coastal states, whereas the remaining 58% are so far “high seas” where there are no political boundaries. Therefore, the rationale of using political boundaries must be complemented by decisions regarding the treatment of “high seas”.
The present proposal is intended to parallel as much as possible the well-established terrestrial scheme, about which
Our endeavour is not starting from scratch. A wealth of marine (and also land) geographical units is stored, with associated descriptions and shapefiles, on the Marine Regions website (
In this paper, we review existing sets of geounits defined in the marine realm and discuss their suitability for the stated purpose, explain the rationale we intend to apply to the marine scheme and put forward a proposal for a comprehensive set of marine geounits, with also four levels of hierarchy. Following TDWG procedures, this proposal is posted on a GitHub repository where it is publicly accessible and open to comments. The main reason for presenting this communication is that, before implementing a marine equivalent of WGSRPD, the proposal would have better chances to be widely accepted if it is first exposed to the scrutiny and feedback from a scientific community interested in biogeography and conservation policy.
In this section, we discuss some of the existing schemes for subdivision of the marine realm and their suitability as potential recording units. Most of these are also listed by
The International Hydrographic Organization has published a detailed geographical scheme for subdivision of the world ocean (
The EEZs are jurisdictional waters declared by coastal states under the provisions of the United Nations Convention for the Law of the Sea (UNCLOS). For those states which have not declared so far an EEZ, the UNCLOS considers the concept of a “continental shelf” (not constrained to the physiographic continental shelf) which is granted with no need for a declaration and comprises the sea-bottom to the same extent of 200 nautical miles. In the Marine Regions website, “EEZ” is taken in the broad sense of a potential EEZ from shore to a distance of 200 NM and a complete set of shapefiles is publicly available (latest version 12 October 2023).
For many small or medium-sized coastal countries and territories, the corresponding EEZ provides a suitable polygon for a basic recording unit at Level 4. The EEZs of larger countries (e.g. the United States, South Africa, Indonesia, Japan) and/or countries with a complex maritime facade (e.g. Spain, Egypt, Costa Rica) must be split.
The Marine Regions website at VLIZ has developed a set of geounits with placetype “Marine Regions” (not to be confused with the name of the website as a whole) which is the intersection of EEZs with the IHO sea areas. These are straightforward for splitting some countries with two maritime facades (e.g. Costa Rica), but there are issues which preclude universally using this set of geounits when we need to subdivide EEZs:
(1) as a consequence of IHO severing adjacent sea areas from larger oceans or seas, some EEZ/IHO intersections result in awkward representations (e.g. the “Egyptian part of the Red Sea” in Marine Regions misses Gulf of Suez and Gulf of Aqaba, deemed to be different bodies of equal rank to “Red Sea” in the IHO Scheme).
(2) in other cases, the intersection of EEZ with major Oceans in the IHO scheme are too large (e.g. United States part of the Atlantic Ocean, spanning from Maine to Florida), therefore inadequate as basic recording units.
Under certain conditions, countries apply for an “Extended Continental Shelf” beyond the 200 miles, but no further from the baseline than either 350 miles or 100 miles beyond the 2500 m isobath (whichever is more favourable; see
Supported by the publication of
Nevertheless, their main drawback is that they ignore country boundaries and must therefore be adapted (intersecting a MEOW with an EEZ) if a country-based system is wanted. In this way, MEOW ecoregions are instrumental as a consistent criterion for subdividing large EEZs (e.g. Brazil, United States, Japan). At a higher hierarchical level, the MEOW Realms are also a consistent source for the definition of Level 2 units (Level 1 being the Oceans).
Coastal countries belonging to the European Union have commitments towards achieving a “Good Environmental Status” of their jurisdictional waters, as detailed in the Marine Strategies Framework Directive. This generates a need for environmental monitoring and elaboration of a suite of 11 “descriptors” (
Another well-established regional scheme is Australia’s “Integrated Marine and Coastal Regionalisation of Australia” (IMCRA 4.0). There are only seven Level 4 units (the states) in the terrestrial WGSRPD scheme, but the marine realm is complex. No less than 18 MEOW are defined in Australian waters (
IMCRA regions are used for display in the Australian Faunal Directory (
A further reason for adopting the IMCRA regions as Level 4 units is that, despite their number, their extension (average surface ca. 244,000 km2) is similar to that of Level 4 units in many other parts of the World.
A two-level scheme was established by
The development of remote sensing technology makes it possible to evaluate the concentration of chlorophyll, based on the solar radiation reflected by the oceans, this being “Ocean-Colour Radiometry” (OCR), for example, the Sea-viewing Wide Field-of-view sensor (SeaWiFS) project (
The Large Marine Ecosystem (LME) approach is aimed at improving the management of ecosystem goods and services. Large Marine Ecosystems (
These are arbitrary areas (
These areas span both EEZs and high seas and their limits are not systematically coincident with political borders so that, both because of this and because of their coarse resolution, we do not find them appropriate for a more fine-grained recording of distributions.
Global regionalisations of coastal waters are based on the analysis of actual distributions of marine species, usually those belonging to well-known groups such as fishes, which reliably cover the entire marine realm.
The marine bioregions designed by
With the increasing amount of Geo-referenced data contained in the Ocean Biodiversity Information System (OBIS) and the Global Biodiversity Information Facility (GBIF), attempts are being made for the categorisation of biogeographic realms, based on the statistical analysis of the distribution of occurrence data worldwide, without any a-priori grouping (e.g.
As mentioned earlier, 58% of the marine realm does not belong to “jurisdictional waters” and cannot be treated with the same rationale as the coastal areas. Under auspices of the Intergovernmental Oceanographic Commission (IOC) of UNESCO, the GOODS initiative presented (
From a pragmatic point of view, if major oceans are taken as Level 1 in the proposed scheme, “High Seas” of each ocean can be Level 2 geounits and the Watling et al. provinces (constrained to the high seas part) incorporated in Level 3 and Level 4.
A sensible way to subdivide the large abyssal provinces of
A geomorphic features map (GSFM) of the global ocean was elaborated by
For the pelagic realm, the difference between bathyal, abyssal and hadal is irrelevant to near-surface plankton. The mesopelagic biome was tackled by
TDWG (renamed Biodiversity Information Standards in 2006, but conserving the acronym which stands for “Taxonomic Databases Working Group”) is a distributed non-profit organisation, founded in 1985, with the mission to develop, adopt and promote standards and guidelines for the recording and exchange of data about organisms. Governance is assumed by an Executive Committee consisting of ten officers elected from and by the membership of the group. One of the means to achieve its mission is to promote the use of standards (give a definition) in several categories amongst which “data standards” for content of controlled vocabularies. A well-known standard from TDWG is DarwinCore for the interoperability of species occurrence data (
The adoption of TDWG standards goes through an elaborate process, which begins by the creation of an “Interest Group”, later consolidated as a “Task Group”, documented by a “charter” that defines the convenor (first contact), core members, motivation, guidance about how to become involved and a summary of what standard or guideline the group wants to achieve. The Geoschemes Interest group was approved in February 2021 by the TDWG Executive Committee and the respective Task Groups Geoschemes: Marine domain and Terrestrial domain (the update of WGSRPD) in November 2023. The corresponding charters are posted on the GitHub repository (Marine charter and Terrestrial charter).
Following section 3.3.3.1. of TDWG Standards Documentation Specification (
The terrestrial WGSRPD (second edition,
The development of Level 2 units is not straightforward. Global biogeographic schemes (e.g.
Taking inspiration from the terrestrial system, Level 3 may accommodate groups of Level 4 units which share political, biogeographical and/or geographical features (e.g. the Lesser Antilles in the Caribbean Sea, each island being a Level 4 unit; the Red Sea, Black Sea, Baltic Sea, Persian Gulf), but may be coincident with Level 4 units when no obvious grouping is suggested. Reasons for grouping at Level 3 include:
(1) Level 4 units belong to a single country and are situated within the same Level 2 unit;
(2) Level 4 units which are comprised in a clearcut geographical context.
For the marine coastal Level 4 units, “coastal” is understood as the 200 NM EEZ or potential EEZ, regardless of the extension of the physical continental shelf, regardless of non-claimed or partly claimed EEZs (e.g. Greece). Claims to an “extended continental shelf” in the sense of UNCLOS, which are currently in a state of flux, are not considered at this stage. Like in the WGSRPD, no attempt is made to represent separately any particular habitats (e.g. the rocky shore, coastal marshes, the bathyal level, the water column). Admittedly, it can be found shocking to represent the distribution of a limpet which is strictly coastal as a 200 miles fringe; however, we concluded that the advantages of using distinct sets of recording units for organisms living in different compartments (e.g. intertidal vs. subtidal, pelagic vs. benthic) are by far outweighed by the advantage of having a universal scheme. Our final consideration is that, if a species is reported in the “Basic Recording Unit”, it is understood to occur “somewhere” therein, not “everywhere” therein.
Level 4 units never encompass the jurisdictional waters of more than one country or territory. For those countries with large and/or complex jurisdictional waters, these are systematically split into as many units as there are Level 1 or Level 2 units involved (e.g. Costa Rica, Guatemala, albeit small, has one Pacific and one Caribbean unit; South Africa has one Atlantic and two Indian Ocean units). In addition to this, some large EEZs (e.g. Brazil, Indonesia) are subdivided even if they are entirely contained in the same Level 2 unit. A threshold for making subdivisions is met when such EEZs involve more than one of the ecoregions in the MEOW system of
Exceptions are made in places where there is a well-established local system of geounits with associated shapefiles: for Europe, the subregions defined for the Marine Strategy Framework Directive (MSFD;
We strived to design Level 4 units to match pre-existing schemes as far as possible (EEZ, IHO/EEZ intersection, MEOW). The rationale can be summarised as follows:
(1) EEZ is used wherever appropriate: not too large and contained within a single sea area [e.g. Belgium, Guinea, Uruguay];
(2) EEZ intersection with IHO is used as next choice if it has no issues [e.g. Costa Rica Caribbean and Pacific parts], using “Marine Regions” placetype (i.e. EEZ/IHO intersection) of VLIZ;
(3) if both EEZ and EEZ/IHO intersection has issues (units unwantedly split down by IHO [e.g. Egypt where Gulf of Suez and Gulf of Aqaba are not included in the IHO definition of Red Sea] or the resulting units still too large [e.g. United States part of the North Atlantic, spanning from Maine to Florida]), the EEZ is subdivided as follows:
- If an existing scheme is in usage and its units meet our requirements (no more than one country, no more than one sea area, not too large), it is adopted to subdivide the EEZ (this applies to the EU with MSFD and to Australia with IMCRA);
- Otherwise, by default, the MEOW boundaries are used to subdivide the EEZ into Level 4 units.
High seas need a specific rationale for the definition of basic recording units. Under auspices of the Intergovernmental Oceanographic Commission (IOC) of UNESCO, the GOODS initiative presented (
With the exceptions of Europe (MSFD subregions) and Australia (IMCRA regions), the default basis for coastal recording units will be version 12 of Maritime Boundaries (released October 2023) from Marine Regions (
When the limits of MEOW are used for the subdivision of large EEZs, the original shapefiles of the MEOWs (hosted on The Nature Conservancy website <https://geospatial.tnc.org/maps/marine-ecoregions-of-the-world-meow> and also at <https://www.arcgis.com/sharing/rest/content/items/903c3ae05b264c00a3b5e58a4561b7e6/data>) are not used. There are several issues: (1) the inner limit is placed way inland and must be trimmed against a shapefile of the baseline; (2) the outer limit is not 200 NM, but an approximate contour and (3) terms of use include the commitment that “... you are using the data for non-commercial purposes, and you don't alter them in any way” (our italics), whereas we want to adjust our recording units to EEZs. Therefore, we will subdivide EEZs (from Marine Boundaries v. 12 at VLIZ) along the MEOW limits when appropriate, but will not use the MEOW shapefiles as such.
For the high seas, we aim to use as Level 4 units the deep-sea basins (below 3500 m depth) and the seafloor elevations (above 3500 m). Source shapefiles are those of the abyssal provinces of
Following section 3.3.3.1. of TDWG Standards Documentation Specification, the term name is a controlled value that represents the class, property or concept [here, concept] described by the term definition. In the case of WGSRPD and the Geoschemes proposed standards, those names correspond to the codes designating the units (one digit or letter for Level 1, two for Level 2, three for Level 3 and Level 4 names derived from Level 3 names with two more letters appended).
Taking into account that the marine proposal follows in the footsteps of WGSRPD, care must be taken to assign them a code with the same format and without duplicates between the land and sea proposals. In the WGSRPD, Level 1 units are identified by a one-digit number (1 to 9), Level 2 regions by a two-digit number (10 to 91), Level 3 units by a three-letter code (not related to the ISO-3166 code of the country) and Level 4 units by a five-letter code in the format XXX-YY where XXX is the Level 3 code and YY an identifier of the level 4 unit. Where Level 3 and Level 4 are identical (the case of 294 out of 614 Level 4 units in WGSRPD i.e. nearly half), this is denoted by a Level 4 code in the format XXX-OO (the double letter ‘O’). In WGSRPD, Level 4 parts of large countries (e.g. Russia) or not so large, but situated at crossroads (e.g. Egypt) may belong to different Level 3, Level 2 and Level 1 units and this also happens in the marine system (e.g. Egypt, Costa Rica).
Developing a set of similar codes for the marine part poses some challenges in order to avoid homonymy with terrestrial codes. Taking into account that the system is alphanumerical and that numbers in the Level 1 and Level 2 are already used up by the terrestrial units, we are compelled to use letters for marine Level 1 units (we propose R for Arctic, A for Atlantic, I for Indian, P for Pacific and S for Southern Ocean), then one digit appended to those letters (A1, A2 etc.) for the marine Level 2 units.
At Level 3, codes must all be distinct from the WGSRPD codes because the Level 2 and Level 1 units above in the hierarchy are different. We propose that the Level 3 code be composed of two letters (by default, the ISO 3166-2 code of the country if appropriate) and one digit, the presence of that digit unambiguously denoting a marine code (there are three letters in all WGSRPD codes). The digit is ‘1’ by default, but must differ if more than one unit is defined for the country or if the letters are already used in another Level 3 unit. At Level 4, the convention of using the suffix ‘-OO’ is conserved, otherwise diagnostic letters are used.
For high seas, it would have been tempting to use the
The local names as discussed above are the same as the controlled value strings (required for controlled vocabularies according to the TDWG Standards Documentation Specification). Admittedly, those codes, be they WGSRPD or marine, are not user-friendly. However, they are machine-friendly provided that they are unique and, therefore, they are needed as part of the standard. With respect to the human user, we have focused on the full names, which are treated as labels, with the intention to make them as easy as possible to identify a geographic area. Following section 3.3.3.1. of TDWG Standards Documentation Specification, the label is a word or short phrase that serves as a human-readable name for the term. These labels of coastal recording units will always start with the adjacent country or dependency name, for example, “France - Bay of Biscay”, not “French part of the Bay of Biscay”. We gathered that many users dislike adjectives (e.g. “Panamanian part of the Pacific Ocean”), which are sometimes awkward to pick in a pull-down menu.
Remote parts and/or dependent states (e.g. Martinique, Cocos Island) are named on their own in the case where they are identified by a separate ISO 3166 code, for example, “Portugal – Azores EEZ” (ISO code PT), not “Azores EEZ”, “Spain – Demarcación marina Canaria” (ISO 3166-2 code SP), not “Canaries EEZ”, but “Martinique EEZ” (ISO code MQ), not “France – Martinique EEZ”. The mention of “EEZ” in the proposed name of geounits is needed because, otherwise, there would be a confusion with the names of terrestrial Level 4 units of the same country; it is omitted if the part is explicitly named a sea area (e.g. “France – Bay of Biscay”, “Saudi Arabia – Red Sea”).
Therefore, the labels read as following:
Country – EEZ (for the complete EEZ of the country). Like in the Marine Regions website, “EEZ” is used instead of “Jurisdictional waters” even where no EEZ is formally declared (Mediterranean), for the sake of brevity.
Country – IHOarea EEZ or Country – Ecoregion EEZ when a large or complex EEZ has been subdivided according to IHO, MEOW or other scheme, for example, “Norway – Barentz Sea EEZ”; “Brazil – Rio Grande EEZ”.
Country – MSFDsubzone where applicable in Europe (we omit “EEZ” in this case).
Australia – IMCRAprovince where applicable around Australia (we also omit “EEZ”).
The naming of high seas features involves no country name but High Seas as such have the ISO 3166 alpha2 code QP (albeit not officially designated). Taking into account that the regionalisation of
In the system here proposed, hierarchy involves the same four strictly hierarchical levels as in WGSRPD. However, the implementation of this set of recording units in the Marine Regions website offers unlimited access to alternative hierarchies, because geounits in Marine Regions can be set with multiple parents. For Level 4 units, possible parents are: (1) the Level 3 unit of this proposal; (2) the nation to which it belongs and (3) the relevant general sea area. Level 4 units in the Marine Regions website can also be set as “parent” for smaller features (e.g. a Marine Protected Area), making that the data associated with such features will be retrieved in a query targeting that Level 4 unit or a more comprehensive one.
Higher level units are here shown on Fig.
The proposal for a marine scheme features five Level 1 units, 26 Level 2 units, 232 Level 3 and 536 Level 4 units (436 coastal and 100 in high seas), comparable to 9 Level 1, 52 Level 2, 372 Level 3 and 609 Level 4 in the current terrestrial scheme. This draft proposal is posted (
Of the Level 4 basic recording units, 175 are coincident with the (potential) EEZ of a country or a dependent territory, 132 correspond to subdivisions of an EEZ following MEOW boundaries, 51 to subdivisions of an EEZ following IHO sea areas (i.e. the "Marine Region" placetype at VLIZ, not to be confused with the name of the website as a whole), 39 to Australian IMCRA areas, 18 to subdivisions according to European MSFD subareas and the 100 subdivisions of high seas are made according to the GOODS scheme.
In line with WGSRPD where Level 1 units are the continents, Level 1 units are the five oceans. Marginal seas (which are treated in the IHO scheme with the same rank as the major oceans) are appended to the major oceans following the boundaries set by NOAA (
Level 2 units in the coastal area correspond broadly to the realms of
Level 3 units are identical to Level 4 in 125 cases and the other 107 Level 3 units account for the remaining 411 Level 4 units. The Baltic Sea, Black Sea, Lesser Antilles and the tropical part of Brazil are examples of Level 3 groupings.
Level 4 units never encompass the jurisdictional waters of more than one country or territory. For those countries with large and/or complex jurisdictional water, these are systematically split into as many units as there are Level 1 or Level 2 units involved (e.g. Costa Rica, Guatemala, albeit small, has one Pacific and one Caribbean unit; South Africa has one Atlantic and two Indian Ocean units).
In addition to this, some large EEZs (e.g. Brazil, Indonesia) are subdivided even if they are entirely contained in the same Level 2 unit by default. By default, partitions were made along the limits of ecoregions in the MEOW system of Spalding et al. (2007). Exceptions are places where there is a well-established local system of geounits with associated shapefiles: for Europe, the subregions defined for the Marine Strategy Framework Directive (MSFD;
It must be emphasised here that the current project aims at establishing a geographic scheme to record distribution data, including those expressed as a text and is not primarily designed to provide a framework for biogeographic analyses. An immediate use for this set of basic recording units is its implementation in the “Distributions” factsheets of the World Register of Marine Species <www.marinespecies.org> (WoRMS). Actually, the present proposal was moved forward by WoRMS editors in a “Distributions Working Group” who recognised the lack of appropriate polygons to display distributions in the database and called for the selection of a coherent set of preferred geounits. WoRMS has stayed away from entering occurrence data based on unpublished datasets (leaving this endeavour to OBIS and GBIF) and, instead, relied on published sources and polygons within which at least one occurrence has been ascertained. The proposed set of geounits, compliant with political boundaries, as well as with current knowledge regarding biogeography, would enable the production of national or regional checklists of species for any country and any taxonomic group, once distribution factsheets are reasonably complete. In addition, the proposed recording units allow us to handle imprecise country-level records from the old literature. The completeness of mapping has so far been hampered by the lack of a consistent set of recording units, leading editors to use inappropriate one (too large, for example, “North West Atlantic” or countries displayed as land on the maps).
The proposal, as it stands, still needs polishing and this is why we strongly urge potential users to provide feedback and suggestions and to criticise or correct decisions we made in some parts of the World for which we do not have a first-hand knowledge. A few alterations were already made in this way, for example, the MEOW boundaries around Madagascar were disregarded because they do not reflect the uniqueness of the Madagascan deep South. The proposal, as it stands, is straightforward for many basic recording units, but we are not assertive for every detail and some points are still open for discussion. Following TDWG procedures, those points should be raised as “Issues” <https://github.com/tdwg/geoschemes/issues> on the proposal’s forum, which can be accessed by anyone using a GitHub account. We conclude this communication with warmly calling every interested scientist to give us feedback.
The authors are grateful to TDWG executives who supported the project in its early stage, particularly to Visotheary Ung who introduced SG to the TDWG formal procedures, to Stan Blum who helped setting up the GitHub repository at the time of our first step and to Steve Baskauf for useful advice on the structure of the Excel spreadsheets of the proposal. We thank Philippe Bouchet (MNHN, Paris) and Nelson Rios (Yale Peabody Museum, New Haven) for useful advice on some specific areas. We also acknowledge the support of the WoRMS community of editors, particularly those of us who endorsed the “Distributions Working Group”. To Francisco Pando, convenor of the Geoschemes terrestrial Task Group, for an always friendly endeavour to coordinate our proposals.
Not applicable
Background document (an illustrated and expanded version of section 2. “State of the art: Existing sets of geounitsin the marine realm” of this article), also posted on the project’s GitHub repository: https://github.com/tdwg/geoschemes/blob/main/marine/background_v3_2025-07.md.
Proposal document (an illustrated and expanded version of section 4. “Results: the proposal” of this article), also posted on the project’s GitHub repository: https://github.com/tdwg/geoschemes/blob/main/marine/proposal_v2_2024-12.md.
An Excel file describing all the proposed recording units in the TDWG Geoschemes marine proposal.