Marine Biology (2024) 171:44

https://doi.org/10.1007/s00227-023-04300-6

REVIEW, CONCEPT, AND SYNTHESIS

Towards global traceability for sustainable cephalopod seafood

Ian G. Gleadall1 · Hassan Moustahfid2 · Warwick H. H. Sauer3 · Lahsen Ababouch4 ·
Alexander I. Arkhipkin5,21 · Jilali Bensbai6 · Isa Elegbede7 · Abdelmalek Faraj6 · Pedro Ferreiro‑Velasco8 ·
Roberto González‑Gómez9 · Carmen González‑Vallés8 · Unai Markaida10 · Piedad S. Morillo‑Velarde9 ·
Graham J. Pierce11 · Stacy Pirro12 · Cristina Pita13 · Katina Roumbedakis14 · Yasunori Sakurai15 ·
David Scheel16 · Paul W. Shaw17 · Pedro Veiga8,18 · Demian A. Willette19 · Andreas Winter5 ·
Tadanori Yamaguchi20

Received: 14 February 2023 / Accepted: 7 September 2023

© The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature 2023

Abstract
Cephalopods are harvested in increasingly large quantities but understanding how to control and manage their stocks, and
tracking the routes of the consumption that exploits them, lag behind what has been developed for exploiting finfish. This
review attempts to redress the imbalance by considering the status of the major cephalopod stock species and the traceability
of cephalopod seafood along the trade value chain. It begins with a general overview of the most important exploited cepha-
lopods, their stock status and their market. Four major cephalopod resources are identified: the three squid species Todarodes
pacificus, Dosidicus gigas and Illex argentinus; and one species of octopus, Octopus vulgaris. The techniques and problems of
stock assessment (to assess sustainability) are reviewed briefly and the problems and possible solutions for assessing benthic
stock such as those of octopuses are considered. An example of a stock well managed in the long term is presented to illustrate
the value of careful monitoring and management: the squid Doryteuthis gahi available in Falkland Islands waters. Issues sur-
rounding identification, mislabelling and illegal, unreported and unregulated (IUU) fishing are then reviewed, followed by a
discussion of approaches and techniques of traceability as applied to cephalopods. Finally, some of the mobile apps currently
available and in development for tracking seafood are compared. This review concludes with observations on the necessity
for the strengthening and international coordination of legislation, and more rigorous standards for seafood labelling and for
taxonomic curation of DNA sequences available in public databases for use in seafood identification.

Keywords Barcoding · eDNA · Fisheries management · Genomics · IUU fishing · Machine learning · Metabarcoding ·
Stock assessment · Traceability app · Octopus · Squid · Amphioctopus · Doryteuthis gahi · Dosidicus gigas · Illex
argentinus · Octopus vulgaris · Todarodes pacificus

This paper is dedicated to the memory of Dr Yuri M. Froerman,

who died on 9 March, 2023. Active particularly in the 1970s
and 1980s at AtlantNIRO, Kaliningrad, he was an outstanding
researcher, working his way up from technician to become Head
of the Laboratory of Commercial Invertebrates. In the mid-
1970s, he formulated a working hypothesis concerning the life
cycle of the squid Illex illecebrosus in the Northwest Atlantic,
overcoming many difficulties to organize expeditions to study the
ecology of the early stages of the development of this species,
brilliantly verifying his hypothesis. In the 1980s, he made great
contributions to studies on the population ecology of Illex
argentinus in the Southwest Atlantic, and the fishery ecology of
Dosidicus gigas in the Costa Rica Dome area. As Chief Scientist
of a research cruise in 1988, he identified dense concentrations
of D. gigas and organized the Soviet Jumbo Squid Trawl Fishery
in this area in 1989–1991. Unfortunately, he had to retire from
cephalopod research in 1992.

Responsible Editor: R. Rosa.

13
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 44 Page 2 of 46
Introduction: sustainability and traceability
Seafood is the world’s most highly traded food commodity
(Asche et al. 2015) and globalization of food supplies now
brings a much wider variety of seafood to the table than
in the past. However, it is also among the foodstuffs most
prone to illegal practices, since, after oil, it is the second
most common food product affected by fraud (Guardone
et al. 2017). The importance of the sustainable production
of seafood is therefore coming under scrutiny during the cur-
rent UN Decade of Ocean Science for Sustainable Develop-
ment, from 2021 to 2030, which is focussing attention onto
a number of key challenges facing humanity (UN General
Assembly 2015). While there have been some major changes
in learning how to ensure seafood sustainability, relatively
little attention has been focussed on cephalopods (i.e., squid,
octopus and cuttlefish), although they are the third largest
marine group consumed by the human population (the larg-
est two being finfish and crustaceans). A major requirement
to achieve sustainability is to ensure that seafood can be
tracked reliably through the sales chain from catch to con-
sumer, fisher to fork (e.g., Gleadall et al. 2022; GDST 2023;
SALT 2023; for these and other abbreviations, see Table 1).
This review was motivated by the recognition that (i)
there is currently insufficient information available on the
issues of traceability and sustainability as applied to cepha-
lopods, and (ii) one way to address and solve these problems
is through the collaborative efforts of experts from various
parts of the world, many of whom attended the most recent
CIAC meeting in Sesimbra, Portugal, in April 2022. That
meeting was therefore a timely opportunity to begin this
collaboration.
Information on many aspects of cephalopods is insuffi-
cient and there are signs of instability in annual catches of
some key species of interest (Arkhipkin et al. 2015a; Sauer
et al. 2021), suggesting that current fishing practices prob-
ably are not sustainable. With this realization, the aims of
this review are to draw attention to and address the root
issues as a step towards achieving reliable, effective and
meaningful traceability of cephalopods, a prerequisite for
reliable, effective and meaningful monitoring and regulation
of sustainable production (Nielsen 2016; Lewis and Boyle
2017; Boero 2021a, b). These root issues include an under-
standing of regulation and certification, (mis)identification,
stock assessment and fisheries management.
What is traceability and what are its functions?
Traceability is defined as the ability to follow the movement
of food through specified stages of production, processing
and distribution (Codex Alimentarius Commission: FAO
and WHO 2018; ISO 22005 2007). Each stage is confined
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Marine Biology (2024) 171:44
to one-step forward or one-step backward in the chain. ‘Trac-
ing’ involves searching the previous (upstream) history of
the product origin, and ‘tracking’ the subsequent (down-
stream) history after the product has been shipped onwards
(e.g., Food Standards Australia New Zealand 2017; Future
of Fish 2020). Traceability requires proper documentation
and record keeping, along with proper handling protocols
during receiving, processing and shipping, to ensure that
the product can be tracked accurately (FishWise 2018).
Ultimately, end-to-end traceability also implies that a con-
sumer unit of seafood at a restaurant or retailer can be traced
throughout the supply chain back to its point of harvest by a
vessel or on a fish farm.
The principles involved parallel those of food safety
embodied in the Hygiene Analysis and Critical Control
Points (HACCP) system, a concept whereby the origin of
food is tracked through each consecutive transaction, so that,
in the event of a food safety compromise, the source of the
problem can be determined quickly and efficiently (by trac-
ing back up the chain), allowing it to be remedied with a
minimum of delay (devised originally to provide safe food
for astronauts on the Apollo Space Program; e.g., Mortimore
and Wallace 2013). One aim of seafood traceability is to
prevent harm to the resource by ensuring that the ultimate
sources are legitimate and involve fishers and other con-
tributors who abide by internationally agreed regulations
put in place to ensure the sustainability of seafood resources
(Global Dialogue on Seafood Traceability (GDST) 2023;
Seafood Alliance for Legality and Traceability (SALT)
2023).
While traceability itself is a key action, it is important to
place it in context when applied to the seafood value chain.
The ultimate aim is to use traceability to monitor the extrac-
tion of seafood from each provenance to ensure that the
resource is not over-exploited and to inform resource stock
assessment to guide appropriate management strategies and
enable harvesting to be maintained indefinitely, which is the
meaning of ‘sustainable’. To verify the status of traceability,
then, it is necessary to understand the status of the resource
itself and the effectiveness of stock assessment and fisheries
management into which the traceability data are to be fed to
realize sustainability. For stock assessment, species identi-
fication (ID) is crucial: misidentification compromises stock
assessment, since the assessment may end up being of a spe-
cies different from the actual species of interest. Therefore,
addressing global traceability of cephalopods is intertwined
with issues of (mis)identification, stock assessment and fish-
eries management.
Traceability systems vary from simple paper traceabil-
ity records to increasingly common electronic traceability
systems that use electronic means such as computerized
or cloud-based databases, barcodes, software solutions,
or other tools to capture and record product traceability
Marine Biology (2024) 171:44 Page 3 of 46 44

Table 1  Key to abbreviations and acronyms

AI Artificial intelligence
ADFG Alaska Department of Fish and Game
ANN Artificial Neural Network
BLAST Basic Local Alignment Search Tool
BOLD Barcode of Life Database
CDS Catch documentation scheme
CephRes The Association for Cephalopod Research. ETS (a non-profit ‘third sector’ company, ‘Ente del Terzo
Settore’), Naples, Italy
CIAC Cephalopod International Advisory Council
CPUE Catch per unit effort
CV Computer vision
DL Deep learning
EEZ Exclusive Economic Zone (as prescribed by the 1982 UN Convention on the Law of the Sea)
EU European Union
FDA US Food and Drug Administration
FAO Food and Agricultural Organization of the United Nations
FIP Fishery Improvement Project
FISH-BOL Fish Barcode of Life (incorporated into BOLD)
FoS Friends of the Sea
GDST Global Dialogue on Seafood Traceability
GPO Giant Pacific octopus
GSSI Global Sustainable Seafood Initiative
HoReCa Hotel, restaurant and catering
ID Species taxonomic identification
IUCN International Union for Conservation of Nature
IUU fishing Illegal, unregulated and unreported fishing
MAPMDREF Le Ministère de l'Agriculture, de la Pêche Maritime, du Développement Rural et des Eaux et Forêts
(Moroccan Ministry of Agriculture, Maritime Fisheries, Rural Development, and Waters and Forests)
MCS Monitoring, control, and surveillance
MSC Marine Stewardship Council (international, non-profit; headquartered in London, England)
ML Machine learning
NCBI National Center for Biotechnology Information
NGS Next-generation sequencing
NOAA US National Oceanic and Atmospheric Administration
‘NOAA Fisheries’ (part of NOAA; also known as NMFS, US National Marine Fisheries Service)
NWA Northwest Africa
OLE Office of Law Enforcement (part of NOAA)
PSMA Port State Measures Agreement
RFMO Regional Fisheries Management Organization
SALT Seafood Alliance for Legality and Traceability
SDGs UN Sustainable Development Goals
SFP Sustainable Fisheries Partnership Foundation
SIMP US Seafood Import Monitoring Program
SNPs Single nucleotide polymorphisms
SPRFMO Commission of the South Pacific Regional Fisheries Management Organisation
SPS Sanitary and Phytosanitary Measures
STS Seafood Traceability System
TREMs Trade Restrictive Measures
UN United Nations
US or USA United States of America
WTO World Trade Organization

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 44 Page 4 of 46
information. To be fully interoperable, systems must be
able to utilize a common data format (syntactic interop-
erability) and they must interpret information based on
shared definitions (semantic interoperability) (Future of
Fish 2016).
Traceability is especially important in today’s context
of increasingly complex supply and distribution systems
as food products pass through multiple hands, in multiple
countries before reaching the final retail shelf. Robust trace-
ability and chain-of-custody mechanisms can also prevent
fraud, or non-certified products being passed off as certified
ones. There are multiple drivers for traceability in the food
sector generally including mandatory food safety require-
ments, private safety and quality certifications, sustainabil-
ity claims, and business-related drivers such as inventory
control, promoting efficiencies, and communication along
the supply chain.
In the USA, food safety regulation initially was a major
driver of the development and investment in traceability
practices and technologies across most food sectors during
the past few decades. More recently, various drivers within
the seafood sector have proved to be incentives for apply-
ing traceability to issues beyond food safety and inventory
management. These include increased media attention on the
legal and social risks within seafood supply chains, govern-
mental traceability requirements, and private-sector sustain-
ability commitments.
Increased media coverage of the environmental, social,
and legal issues associated with seafood has led to sig-
nificant shareholder concerns, potential impacts on brand
value, and challenges to the corporate social responsibility
initiatives of companies. This recent attention has also cre-
ated an opportunity for companies with full traceability to
actively promote the many benefits of their products such as
social and fair-trade compliance and engagement in fishery
improvements. Consequently, companies are increasingly
committing to sustainable seafood sourcing policies, and the
challenge is now for those companies to be able to trace the
origin of their products to ensure that species and attributes
of the products are meeting their policies and communicated
to the customer accurately. For companies that buy and sell
seafood, the lack of product origin information and supply-
chain transparency can pose significant risks. The first step
towards mitigating and eventually eliminating these risks is
to ensure that end to-end, electronic, interoperable traceabil-
ity systems are in place throughout the supply chain. This
work is already under way with some companies that are
instituting traceability policies and setting goals, often with
the assistance of NGOs, government bodies, and technology
companies. However, much more work is needed (Lewis and
Boyle 2017).
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Marine Biology (2024) 171:44
Regulation and certification: necessary
prerequisites for traceability?
Seafood traceability has been born out of the realization
that, for seafood to be sustainable, it is important to regu-
late its exploitation and demonstrate that it has come from
a properly managed resource. However, guaranteeing that a
resource is sustainable relies on a complex of certification
schemes and regulatory mechanisms. Many of these have
yet to be applied effectively to cephalopod seafood but are
briefly summarized here to understand what is required to
bring sustainable cephalopod exploitation up to speed. Much
of the impetus for seafood regulation has its origin in the
USA, in view of its reliance on large amounts of imported
seafood.
There are several important and influential factors in the
US seafood industry. Regulatory institutions (NOAA Fisher-
ies and the FDA) aim to ensure that the US consumer can
enjoy safe and sustainable seafood. Through NOAA Fisher-
ies, the consumer should be secure in the knowledge that,
first, it comes from a sustainable fishery (whether harvested
and processed locally or imported from another country,
either directly or via processing in a third country and re-
exported to the USA); and, second, it was fished legally in
full respect of US requirements concerning factors such as
gear, by-catch, discards and transparency. In addition, the
FDA determines whether or not seafood is safe to consume,
being free of infection or intoxication. US importers, proces-
sors and retailers are concerned to meet these NOAA Fisher-
ies and FDA requirements, because the burden of proof is
on them, so traceability is of paramount importance to their
business.
The Marine Stewardship Council (MSC) is a leader in
seafood eco-labelling and certification, although the organi-
zation Friends of the Sea (FoS) has established its promi-
nence in southern Europe. The limitations of MSC and FoS
labelling and certification schemes have become apparent,
because, despite 20 years of activity, less than 20% of the
seafood traded internationally is certified. As a result, many
major seafood firms have started recognizing fisheries that
are under a Fishery Improvement Project (FIP) as a move
towards sustainability for sourcing, even though the seafood
is not yet eco-labelled or certified. Currently, there are six
cephalopod fisheries MSC-certified or in assessment and
nine cephalopod fisheries included in FIPs (Table 2). Note,
however, that a major problem with fishery certifications is
that they may involve only a limited number of fishers and
producers in a given area and do not include most of the
fishery. An example is the FIP for Mexico Yucatan octopus
which includes only part of the total fishing fleet and fish-
ing plants. Sometimes, there may be more than one FIP for
the same fishery (note the two different US certified squid
fisheries at the top of Table 2; see footnote a).
Marine Biology (2024) 171:44 Page 5 of 46 44

Table 2  Cephalopod fisheries certified or under full assessment by the Marine Stewardship Council (MSC) and included in Fishery Improve-
ment Projects (FIPs) (Information updated from Roumbedakis et al. 2021 and MSC 2023)
Name Type & Status Species Country

Cephalopod Fisheries MSC-Certified or in Assessment

a
US Northeastern Coast Longfin Inshore Squid Certified Illex illecebrosus US
and Northern Shortfin Squid Bottom Trawl Doryteuthis (Amerigo) pealeii
Fishery
a
US Northeast Squid Bottom Trawl Fishery Certified Illex illecebrosus US
Doryteuthis (Amerigo) pealeii
Western Australia octopus Certified Octopus djinda Australia
Western Asturias Octopus Trap Fishery of Certified Octopus vulgaris Spain
Artisanal Cofradias
California Market Squid Producers Alliance In assessment Loligo opalescens US
purse seine fishery
California market squid purse seine fishery In assessment Loligo opalescens US
Cephalopod Fisheries included in FIPs
Chile Patagonian red octopus—hookah Basic, Active Enteroctopus megalocyathus Chile
Indonesia Nusa Tenggara Timur Day Basic, Active Octopus cyanea Indonesia
octopus—diver-caught, gleaning, hand
gathered, handline, jig & spear
Mexico Bahia de Los Angeles octopus—trap/ Basic, Active Octopus bimaculatus Mexico
diver-caught/hand gathered Octopus hubbsorum
Southwest Atlantic Argentine shortfin squid— Basic, Active Illex argentinus Falkland Islands
jig (TSSFA)
Mexico Yucatan octopus—drift rod and line Comprehensive, Active Octopus maya Mexico
Octopus americanus
Southwest Madagascar octopus—diving & Comprehensive, Active Octopus cyanea Madagascar
gleaning
Peru jumbo flying squid—jig Comprehensive, Active Dosidicus gigas Peru
Indonesia North Sumatra squid—handline Comprehensive, Active Uroteuthis chinensis Indonesia
India Kerala shrimp and cephalopods—trawl Comprehensive, Active Amphioctopus neglectus India
Uroteuthis duvauceli
Sepia pharaonis
Argentina shortfin squid—jig Prospective Illex argentinus Argentina
Shantou-Taiwan Chinese common squid and Basic, ­InactiveA Uroteuthis chinensis China
short arm octopus—jig/trawl Amphioctopus fangsiaob
Shantou-Taiwan shortarm octopus—jig Basic, ­InactiveB Amphioctopus fangsiaob Taiwan
Mexico Gulf of California jumbo squid—jig Comprehensive, ­InactiveC Dosidicus gigas Mexico
East China Sea and Yellow Sea Japanese flying Comprehensive, ­InactiveD Todarodes pacificus China
squid—trawl
Japan Tomamae giant Pacific octopus—barrel Comprehensive, ­InactiveE Enteroctopus dofleini Japan
flowing

See: https://​fishe​ries.​msc.​org/​en/​fishe​ries/@@​search?​q=​squid​&​search = currently, both certification units have merged to reduce costs
a
Note that there are two different certification units for the same squid fishery: two business groups applied to certify their respective fleets,
which is permitted by the MSC certification program. The first is shown as ‘certified’ on the MSC web page and the ‘US Northeast squid bottom
trawl fishery’ as ‘combined with other certificate’
b
Originally as the junior synonym Octopus ocellatus
Causes of inactivity of FIPs: Athree years without environmental progress; Bmerged with another FIP; Cpaused until this fishery resumes and
improvements can continue to be implemented; Dat MSC’s request (major changes in the nature of the fishery have led to the need for re-assess-
ing the scope of improvements and work plan implementation); Emissed deadline for 3-year audit report and failed to meet social responsibility
policy

A problem with certification is to understand which is available evidence, but it is important that these schemes
the most trustworthy and reliable scheme. All eco-label- are assessed and that benchmarks are provided, which is
ling certification schemes claim that they respect interna- the function of the Global Sustainable Seafood Initiative
tional requirements and are based on science and the best (GSSI), a public–private partnership supporting its partners

13
 44 Page 6 of 46
in accelerating the implementation of the United Nations
Sustainable Development Goals (UN SDGs; UN General
Assembly 2015).
NOAA Fisheries, the FDA, and US importers, proces-
sors and retailers are bound by the international rules of
trade in seafood, which have been driven hard politically and
economically by the US to promote free trade and a liberal
economy. This culminated with the creation of the World
Trade Organization (WTO) and its various binding rules
and agreements including the Sanitary and Phytosanitary
Measures (SPS) and Technical Barriers to Trade agreements
for market access and entry. As a result, NOAA Fisheries,
the FDA, and importers to the USA must implement market
access requirements (including those related to traceability)
that are in conformity with internationally agreed require-
ments. For seafood, this is facilitated by Globefish, which
is a multi-donor-funded project within the Food and Agri-
cultural Organization (FAO) Fisheries Division, established
in 1984. Globefish is responsible for providing up-to-date
trade and market information on fish and fishery products,
promoting and easing trade-related information exchange
among the seafood industry, governments, academia and
stakeholders worldwide, particularly for developing coun-
tries and countries in transition. Globefish gathers, assesses,
analyses, and disseminates up-to-date fish marketing and
trade information worldwide, through key publications and
its website (FAO 2021b).
Certification and regulation of cephalopods vary greatly
according to species, the locality and distribution of each
currently playing a large part in dictating the extent to which
they can be monitored and tracked as a seafood product. This
in turn has a large impact on the sustainability of the stock
of each particular species on the market.
Cephalopods and their market
Cephalopods have been harvested in large amounts in recent
decades, reaching a peak global catch of 4.9 Mt in 2014,
decreasing to 3.7 Mt in 2020 (FAO 2022a). They are mostly
wild caught, supplied mainly from China, India, Morocco
and Peru (although a substantial proportion of these catches,
especially of squid, is taken on the high seas, i.e., in inter-
national waters). Although their consumption is attaining
high popularity in many countries, they remain culturally
linked mostly to southern European countries (Portugal,
Spain, Italy and Greece), some Latin American countries
(Mexico, Argentina and Uruguay) and several Pacific and
Asian countries, particularly Japan (Markaida and Gilly
2016). They are widely traded internationally. Over 62% of
production is exported, generating a revenue of US$ 10.2
billion in 2020, representing about 7% of the global sea-
food trade value. The most important import markets for
cephalopods are China, the European Union (particularly
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Marine Biology (2024) 171:44
Italy and Spain), Japan and the Republic of Korea. In 2020,
global trade of octopus was estimated at 283,577 t, valued at
US$ 2.1 billion. Major producers are China, Japan, Mexico,
Morocco and Mauritania (FAO 2022a). Countries key to the
trade network for live, fresh or chilled octopus are France,
Germany, Italy and Spain (Ospina-Alvarez et al. 2022).
The USA is the largest seafood importing country with
imports accounting for 65% of domestic seafood consump-
tion and 15% of the global total seafood import value in
2020 (FAO 2022a). This includes 65,411 t of imported
cephalopods valued at US$ 383 million, of which 17,278
t was octopus valued at US$ 118 million. The same year,
the USA exported around 18,000 t of cephalopods valued
at US$ 110 million and negligible amounts of octopus.
Imports of octopus have increased significantly over the
years, with the volume doubling and the value increasing
several-fold between 2010 and 2021 (NOAA 2023).
To enter the lucrative US market, industrial and artisa-
nal fisheries that harvest octopus, along with the import-
ers and processors with whom they work, must ensure
that harvests comply with strict technical regulations
and standards of resource sustainability and food safety,
regardless of whether they are harvested in the US Exclu-
sive Economic Zone (EEZ) or imported. Compliance
requires reliable and robust traceability technologies but
enables smoother seafood trading and distribution. Equally
important, making reliable traceability technologies avail-
able and affordable for exporters can help small-scale har-
vesters in developing countries to conform to regulations,
which otherwise can become technical barriers to trade,
blocking access to this lucrative market. Ideally, reliable
traceability will help to promote fair-trade practices and
economic development. For example, the main export
from Mauritania is octopus, estimated at 68% of the value
of all its seafood exports, with almost 80% of the octopus
harvested by artisanal Mauritanian fishers. However, the
route of this harvest through to arrival in the US, and the
monetary value obtained by the local Mauritanian fishers,
are currently obscure.
Status of cephalopod resources
To comment on and point the way ahead for the traceability
of cephalopods, it is first necessary to understand the status
of the exploited stocks, the fishing industry and the value
chain including the major species involved, the condition of
the stocks, how (and to what extent) the stocks and fisher-
ies are monitored and managed, and to recognize the main
problems.
Marine Biology (2024) 171:44
Fig. 1  Fluctuations in the annual catch of Todarodes pacificus in
Japan and South Korea. (Data from Fisheries Agency of Japan and
Japan Fisheries Research and Education Agency (2021), a public-
domain resource which is updated periodically)
Fig. 2  Global capture for the major cephalopod straddling stock spe-
cies Dosidicus gigas, Illex argentinus, Todarodes pacificus from 1950
to 2020. (Source: FAO 2022b)
Major cephalopod stocks
More than half the world’s annual cephalopod catch consists
of just three species of squid in the family Ommastrephidae:
Todarodes pacificus from the Northwest Pacific (Figs. 1, 2),
Dosidicus gigas from the eastern Pacific, and Illex argenti-
nus from the Southwest Atlantic (Fig. 2; FAO 2021a, 2022a).
Unfortunately, these are all transboundary, straddling and/
or highly migratory stocks (Maguire et al. 2006; Howard
et al. 2013) which, in the absence of binding international
regulations or agreements, lack unified stock assessment and
management strategies (Arkhipkin et al. 2022).
The last 2 decades have seen a large expansion of distant-
water squid-fishing fleets, mainly from China, which have
unrestricted access to such stocks. This increased pressure
on these three squid species, especially on the high seas,
renders these resources highly vulnerable to over-exploita-
tion. The relevant coastal states implement their own fish-
eries management measures within their respective EEZs
Page 7 of 46 44
where these transboundary, straddling and highly migratory
resources of squid occur (Arkhipkin et al. 2015a). However,
data exchange between these states tends to be limited and
this hinders joint management of these resources. Further,
unregulated high-seas squid fisheries with little-to-no con-
trol on fishing activities, and poor or non-existent monitor-
ing (e.g., see Welch et al. 2022), pose a significant threat to
the sustainability of these squid resources. In addition, these
squid appear in markets around the world often without reli-
able information on where they were caught or by whom.
Todarodes pacificus, the Pacific flying squid
The Japanese flying squid, Todarodes pacificus (Steenstrup,
1880), has a 1-year life cycle and is distributed in the north-
western Pacific Ocean, the Sea of Japan, the Yellow Sea
and the East China Sea (Ishii 1925; Sasaki 1929; Soeda
1950; Hamabe and Shimizu 1966; Araya 1967; Kasahara
1978; Okutani 1983). The distribution range shifts season-
ally with changes in water temperature (Soeda 1950; Araya
1967; Okutani 1983; Sakurai et al. 2000): the northern
limit of its range is around 50°N in September and around
40°N in April. Three or four cohorts can be distinguished,
based on the spawning season, of which the autumn and
winter cohorts are the largest in terms of biomass (Araya
1967; Okutani 1983; Osako and Murata 1983; Kidokoro
2009; Kidokoro et al. 2010; Sakurai et al. 2013).
Catches of this species have long been known to fluctu-
ate: the subject of intensive research effort as long ago as the
early 1920s (Ishii 1925). Total annual landings of T. pacifi-
cus were less than 200,000 t before the 1930s and increased
in the 1940s with the development of commercial fisheries.
Annual landings were usually 400,000–500,000 t during the
1950s and 1960s, but decreased markedly during the 1970s,
dropping to 100,000 t in the mid-1980s. In the 1990s, land-
ings increased and rebounded to about 300,000–400,000 t,
but annual landings have decreased steadily since the year
2000 (Kidokoro et al. 2010) falling to a mere 13,416 t in
2019 (Fig. 1, Fisheries Agency of Japan and Japan Fisheries
Research and Education Agency 2021).
The decline in the stock size of T. pacificus during the
1970s and 1980s was considered to have been caused by
excessive fishing effort at that time (Murata 1989), but
changing environmental conditions may also strongly affect
the stock size (Sakurai et al. 2000). The drastic declines in
recent years are suspected to be a result of IUU fishing. The
catch-per-unit-effort (CPUE) of T. pacificus caught in the
northwestern Tatar Strait by vessels of the Russian Federa-
tion in the 2010s generally remained stable (mean CPUE
between 2003 and 2019 was 582 ± 45.8 kg.vessel−1 ­day−1),
but mean body weight increased from 205 ± 4.0 g in 2004
to 256 ± 3.5 g in 2012, and 297 ± 6.3 g in 2019, which
13
 44 Page 8 of 46
apparently is an effect of climate warming in the Sea of
Japan (Dulenina et al. 2020).
South Korean vessels targeting T. pacificus operate in the
Sea of Japan, Tsushima Strait and the Yellow Sea. Total
annual landings of this species were less than 100,000 t in
the 1980s and until 1992. Increasing suddenly, the landings
were almost the same as those of Japan for several years. In
the 2000s and 2010s, South Korean landings exceeded those
of Japan by about 50%. However, the landings decreased
from around 2000 in the same way as seen in Japan (Fig. 1,
Fisheries Agency of Japan and Japan Fisheries Research
and Education Agency 2021). The number of North Korean
fishing vessels operating in the Sea of Japan increased from
2017 to 2019 (Park et al. 2020), but their landings of T.
pacificus are unknown.
Vessels from China targeting T. pacificus operate in
the Yellow Sea and East China Sea, but since 2004 have
appeared in the exclusive economic zone (EEZ) of North
Korea in the Sea of Japan (Wang et al. 2018). These Chinese
vessels were reportedly operating in the Sea of Japan from
May to November with a peak between August and Octo-
ber (as observed through night-time satellite images of the
lights used to attract the squid, since this is an area of high
incidence of vessels switching off their automatic identifica-
tion system (AIS; Park et al. 2020; Welch et al. 2022). Catch
amounts for T. pacificus landings in China are unclear.
In view of the ongoing serious decline in catches, it is
important that ways are found to regulate fishing effort, for
which it is necessary first to find ways of improving methods
of stock assessment. This will not be possible until there is
a better understanding of total catch volumes aggregated
across all the countries fishing this species, and to achieve
this reliable traceability is key.
Dosidicus gigas, the Southeast Pacific Humboldt or jumbo
flying squid
Dosidicus gigas (d’Orbigny, 1835) is the most abundant
commercial squid species in the Southeast Pacific. This
nerito-oceanic squid inhabits open waters of the tropical
part of the Eastern Pacific from the shelf edge of South
America to approximately 125°W along the equator (Jereb
and Roper 2010). On the shelf and continental slope, D.
gigas occurs in the EEZs of practically all Pacific-coast
American States, from Chile in the south to Alaska (USA)
in the north (Arkhipkin et al. 2015a). Dosidicus gigas is
the largest representative of the family Ommastrephidae,
attaining 100–120 cm mantle length and 65 kg weight
(Nigmatullin et al. 2001). This species is fished in very
large amounts, even though its flesh contains ammonia as a
buoyancy aid (Denton 1974), giving it a noticeably acid taste
and a distinct, characteristic ammoniacal smell. These can
be disguised, however (see Sect. “Deliberate mislabelling”).
13
Marine Biology (2024) 171:44
Two metapopulations of this squid inhabiting the two
hemispheres are genetically different, with equatorial
currents acting as a natural barrier (Staaf et al. 2010).
Currently, two phenotypic groups or size forms are
distinguished within each ‘hemispheric’ metapopulation:
medium (< 49 cm ML) and large (> 49 m ML; Keyl et al.
2008; Tafúr et al. 2010). The existence of these two size
groups of squid is the result of water temperature impact on
D. gigas juveniles. Warmer waters induce early maturation
with animals having a 1-year (or even shorter) life span
attaining small or medium sizes. Colder waters delay
maturation until the second year, with squid attaining large
sizes and having a life span of 1.5–2 years (Arkhipkin et al.
2015b).
Detailed information about the life cycle of D. gigas is still
lacking. It is known that it undergoes pelagic spawning in
open-ocean tropical waters of temperatures ranging between
15 and 25 °C (Staaf et al. 2008). An unknown proportion
of recruitment remains within the spawning grounds and
feeds there. The rest of the recruitment migrates eastwards
from the open ocean to the tropical and subtropical regions
of the continental slope and shelf and feeds extensively
on myctophids and other pelagic organisms. An unknown
proportion of these migrating recruits performs extensive
latitudinal seasonal migrations from subtropical waters to
cold-temperate regions of both hemispheres. These recruits
use productive areas of temperate and subpolar shelves and
continental slopes for intensive feeding and growth, and
then return to their subtropical/tropical spawning grounds
to spawn (Gretchina and Zúñiga 2019). Squid that had stayed
in the shelf/slope tropical feeding areas upon maturation also
perform ‘reverse’ migrations westward back to the oceanic
spawning grounds, benefiting from lesser pressure by
nektonic predators in less-productive oceanic waters (Alegre
et al. 2014; Csirke et al. 2018).
Dosidicus gigas has been exploited since the 1980s. Until
1999, the majority of the total catch was taken from a rather
small area in the northern part of Baja California in Mexico,
with annual catches attaining over 120,000 t in 1997. After
the series of strong El Niño events (up to 1997) and two
sequential years of La Niña (in 1998–1999), the large size
group of D. gigas appeared in great numbers in the southern
hemisphere where annual catches of 20,000–30,000 t
increased sharply in 1996–1997, reaching ~ 300,000 t in
2005 (Arkhipkin et al. 2015a; Csirke et al. 2018).
Targeted fishing for D. gigas in the northern hemisphere
declined dramatically after 2013 and was almost non-
existent after 2015, as the large form disappeared from the
catches. In the last decade, the specialized fishery for D.
gigas has taken place only in the Southeast Pacific where
the fleets of three countries (Peru, Chile and China) take
more than 97% of the total annual catch (FAO 2021a, b). In
Chile, from 19 to 35 industrial pelagic trawl or purse seine
Marine Biology (2024) 171:44
vessels and about 2,000 small artisanal boats fish primarily
the large form of D. gigas, which migrates to Chilean waters
mainly from February to September (Gretchina and Zúñiga
2019). The total annual catch in Chile has not fluctuated
strongly over the last decade, varying between 106,000 and
181,000 t. However, recent catches have fallen significantly,
to 58,000 t in 2019 and 55,000 t in 2020, as a consequence
of smaller squid size and possibly because of substantial
changes in the fishing fleet: since 2020, only artisanal vessels
have been allowed to target D. gigas there (SPRFMO 2021).
Since 2011, the Peruvian fishing fleet consists of more than
4600 small artisanal vessels fishing mainly within 40 nm
off the coast. Artisanal vessels fish for squid using large
hand jigs year-round and target immature squid during their
feeding period, as well as maturing and mature squid just
before their migrations to the pelagic spawning grounds
(Yamashiro et al. 2018). Annual catches by the Peruvian
fleet are the largest among the three main fishing states,
ranging between 300,000 and 600,000 t (FAO 2021b).
The high-seas fishery for D. gigas in the Southeast
Pacific consists of Asian industrial jigging fleets fishing in
tropical waters from the equator to 20°S and from 120°W
to the boundary of the EEZs of Peru, Ecuador and north-
ern Chile. The great majority of the fleet is represented
by Chinese vessels that increased their numbers from 276
jiggers in 2016 to a maximum of 557 jiggers in 2020 (Li
et al 2021). A dozen Taiwanese and South Korean jiggers
also fish in this area. The fleet targets D. gigas in different
areas throughout the year. In summer, the Asian fleet oper-
ates in equatorial waters in the open ocean targeting small,
immature squid, belonging to medium and large groups.
In autumn, the Chinese fleet fishes closer to the EEZs of
Ecuador, Peru and northern Chile and targets a high pro-
portion of mature squids also belonging to both size groups
(Li et al 2021). The total annual catch of the Chinese fleet
has been relatively stable during the last five years, rang-
ing between 223,300 and 358,000 t (FAO 2021b). How-
ever, increased fishing effort caused a significant decline
in CPUE from 3.9 to 4.0 t ­vessel−1 ­day−1 in 2017–2018
to 2.8–3.0 t ­vessel−1 ­day−1 in 2019–2020 (Li et al 2021).
As a result, the Chinese Government issued a unilateral
3-month fishing ban from 1 September to 30 November
that coincided with the main spawning peak of D. gigas in
the Eastern Pacific (5° N–5° S, 110°–95° W) (Nigmatullin
et al. 2001). Further, the Chinese Government restricted the
number of jigging vessels fishing north of 8°S to 420 ves-
sels, and to 400 vessels fishing south of 8°S for 2022–2023
(Undercurrent News 2021).
The Dosidicus gigas fishery is relatively stable in the
Southeast Pacific with the total annual catch exceeding
750,000 t every year of the last decade (FAO 2022b).
However, it is not known what proportion of the D. gigas
metapopulation migrates between the EEZs and high seas,
Page 9 of 46 44
making it difficult to estimate an impact of fishing pressure
on specific phases of squid ontogeny by different fleets.
The high-seas fisheries in the Eastern Pacific are controlled
by the Commission of the South Pacific Regional Fisheries
Management Organisation (SPRFMO) with all fishing ves-
sels required to follow the conservation and management
measures adopted by the Commission. However, it seems
that SPRFMO members are reluctant to support and accept
management measures that may constrain their high-seas
fishing operations, in particular for the D. gigas fishery.
There are no accepted regulations on the amount of fish-
ing effort, uniformity of the reported data or independent
catch verifications, a situation which has prevented the
SPRFMO from undertaking any adequate assessment of
the D. gigas stocks in the Convention area or EEZs of
adjoining coastal states. It seems currently unlikely that
the SPRFMO will be able to impose more stringent regula-
tions or to achieve closer cooperation for stock assessment
on the high seas and within each EEZ, rendering stocks of
this squid vulnerable to over-exploitation (Arkhipkin et al
2022). Reliable traceability would clearly be of great help
in resolving many of these shortcomings.
Illex argentinus, the Southwest Atlantic shortfin squid
Illex argentinus (Castellanos, 1960), sometimes called the
Argentine shortfin squid, is the most abundant commercial
squid species in the Southwest Atlantic, occurring in the
EEZs of Brazil, Uruguay, Argentina and the Falkland
Islands, and also in the high-seas areas of the Patagonian
Shelf at 42° S and 45–47° S (Jereb and Roper 2010). The
population structure of I. argentinus is complex, but a
predominant 95% comprises the South Patagonian Stock
(Hatanaka 1988).
The total annual catch of I. argentinus exceeds 1 Mt in
some years (e.g., 1999 and 2015; FAO 2022b). Currently,
this squid is caught by three main fleets in different parts of
its range. The first fleet fishes exclusively within the Argen-
tinean EEZ and consists of ~ 80–90 Argentinean domes-
tic jiggers and trawlers. The second fleet consists of ~ 105
Taiwanese and South Korean jigging vessels and ~ 10–15
Spanish and Falkland-flagged trawlers, starting to fish on
the high seas in January–February and then moving into
the northern part of the Falkland Conservation Fishing
Zones (FICZ/FOCZ), where the licenced fishery takes place
between February and middle of June. The third and largest
fleet consists of 300–400 jigging vessels and several dozen
trawlers (predominantly Chinese) and fishes exclusively in
the unregulated areas of the high seas, mainly from Decem-
ber until July (Arkhipkin et al. 2022). The catches and sub-
sequent trading figures for this species should correspond to
the amounts reported by these countries in the international
production and trade databases (FAO 2022b; UN 2023).
13
 44 Page 10 of 46 Marine Biology (2024) 171:44

In the last 20 years, total annual catches of I. argentinus

have experienced ‘boom and bust’ variability with almost an
order of magnitude difference between the poorest (146,645
t in 2016) and richest (1,011,331 t in 2015) years (FAO
2022b). The most stable catches are observed for the first
fleet in Argentina, which has the advantage of exploiting the
large Patagonian and Argentinean Shelves within the Argen-
tinean EEZ. In contrast, catches of the second fleet have
experienced the greatest variability, up to 20-fold during the
last decade (FAO 2022b). Such variation in catches is prob-
ably caused by the variability in southward migrations of Fig. 3  Octopus vulgaris annual landings on the NW African coast
the South Patagonian Stock squid, the only stock exploited (FAO, CECAF 2020)
by this fleet (Arkhipkin et al. 2015a). One of the causes of
such variability could be oceanographic conditions in the
southern part of the Patagonian Shelf, with colder waters
preventing squid migrations further south. Another cause
could be the fishing activities by the numerous vessels of the
third fleet in the high seas taking up a significant proportion
of the South Patagonian Stock during their southward feed-
ing migrations (Waluda et al. 2001a, b). On the high seas,
the variability in catches taken by the third fleet has been
intermediate, as these vessels exploit not only migrating
South Patagonian Stock, but also squid stocks belonging to
other less numerous cohorts such as summer-spawning and Fig. 4  Octopus vulgaris abundance index (kg per 30 min.) from
Bonaerensis North Patagonian Stocks (Brunetti et al. 1998). Moroccan demersal surveys, Cap Boujdor-Lagouira (FAO, CECAF
The Southwest Atlantic is unique as it does not have a 2020). See text for explanation of the observed fluctuations
Regional Fisheries Management Organization (RFMO)
responsible for management and conservation of all trans-
boundary and straddling stocks including I. argentinus. importers from NWA. Octopus is fished by both small-scale
Without coordinated management and regulation of the fish- and industrial fleets, which use different gears that impact
eries, straddling stocks of I. argentinus are highly vulnerable the stock differently (Sauer et al. 2021).
to overfishing, especially in years of naturally occurring low Landings and abundance in NWA (Figs. 3, 4) show inter-
recruitment. Reliable traceability of the provenance of fished annual variability due to the biological characteristics of
I. argentinus is a prerequisite for resolving this vulnerability. being a short-lived species. The fluctuations in abundance
and catches are influenced by annual variations in recruit-
Octopus vulgaris, the common octopus of the Eastern ment and environmental condition changes. In Morocco, the
Central Atlantic landings between 1991 and 1997 showed a decline that could
be attributed to the increased effort resulting from the entry
Although squid account for the majority of cephalopod of artisanal boats and coastal trawlers to the fishery. Since
landings, the common octopus, O. vulgaris, stands out as 1997, an improvement in landings has been observed in
the octopus species of major fishing and trading interest. Morocco after the biological rest period was increased from
The Eastern Central Atlantic (FAO Major Fishing Area 34), one period to two, which reduced fishing effort by about
especially off Northwest Africa (NWA; i.e., Morocco, Mau- 30% compared with 1996 and the departure of the Euro-
ritania, Senegal and The Gambia), is considered the most pean trawling fleet targeting cephalopods in 1999. In 2003,
productive fishing ground of this species with 89,000 t in the catch declined dramatically, as a result of the uncon-
2019 (FAO, CECAF 2020), representing 32% of total demer- trolled increase of artisanal fishing that was structured sub-
sal catches in this region. Even though the catch is small sequently, followed by a long, gradual recovery and stability.
compared to other fish species, it has a high economic value. Octopus in Morocco are generally caught mixed with
In Mauritania, for example, it is the top fishery resource in other demersal species including other cephalopods (e.g.,
terms of value in a country where GDP is currently heav- squid and cuttlefish). These other species have shown a
ily reliant on fishing. In the last few decades, the number gradual increase recently due to the management measures
of countries importing octopus has increased (Sauer et al. being implemented, mainly indirect measures associated
2021), although Japan, Spain and Italy remain the major with octopus management plans (MAPMDREF/DPM 2004)

13
Marine Biology (2024) 171:44
and the establishment of protected areas that are presumably
contributing to protecting essential habitats and biodiversity.
Another management measure that is contributing to
sustainability of octopus is the quota control system for the
coastal and offshore fishing fleets, established since 2001,
which requires all catches to be landed at the local fishing
ports for control by the local commissions.
Stock assessment
To understand whether or not fishing on a given stock is
sustainable, some form of stock assessment is necessary.
Stock assessment is an aspect of the quality of manage-
ment of a stock and hence speaks to both sustainability and
traceability (although not assuring either). Where fisheries
are assessed and managed rigorously, compliance is also
excellent. Compliance may be achieved by a top–down pro-
cess, being required to obtain and maintain licences (e.g., as
required in the Falkland Islands Doryteuthis gahi fishery; see
Sect. “Doryteuthis gahi, a model squid stock”), or it may be
achieved through a more bottom–up, co-management-based
process, and in any case is necessary for certification (e.g., in
the Western Asturias Octopus Trap Fishery for O. vulgaris;
Table 2). Where there is little assessment, but there are many
rules and regulations, there is probably less compliance; for
example, with octopus being sold directly to restaurants and
thus bypassing controls (as in parts of southern Europe).
Globally, many cephalopod stocks, especially those
supporting important fisheries, are assessed using a vari-
ety of methods, ranging from depletion models (see the
Sect. “Assessment based on modelling: depletion models
and other approaches”), which are mainly used in squid fish-
eries, to production models, which have largely fallen out of
fashion for assessing finfish stocks but seem to produce use-
ful results even if the assumption of fixed carrying capacity
is unrealistic for short-lived species such as cephalopods.
Small-scale cephalopod fisheries such as European fisheries
for O. vulgaris, often lack formal assessment even if fish-
ing is strictly controlled. In small-scale fisheries, the role of
fishers in assessment and management may be especially
important (Sect. “Octopus stock assessment: the role of fish-
ers”) and citizen science can also play a role (Sect. “Octopus
stock assessment: the role of citizen science”). However,
many fished stocks (including most European fished cepha-
lopod stocks) are not routinely assessed and, indeed, often
the stocks are not formally defined.
Assessment based on modelling: depletion models
and other approaches
Stock assessments of cephalopods are constrained by their
life-cycle characteristics of weak spawner–recruit rela-
tionships, high sensitivity to environmental factors, and
Page 11 of 46 44
particularly their short, circa-annual, lifespan. An annual
stock is effectively a new population every year and its abun-
dance cannot be determined from standing biomass carried
over from prior years (Rosenberg et al. 1990; Pierce and
Guerra 1994). Many squid fisheries therefore carry out stock
assessments with depletion models, which can be calculated
using in-season time series of catch and effort to estimate
levels of abundance that would sustain the observed rates
of catch (Arkhipkin et al. 2021a). This modelling approach
is applied in Spain in western Asturias for the small-scale
octopus fishery there (Roa-Ureta et al. 2021), and also in
Morocco since 2011 for the assessment of the Total Allowa-
ble Catch Octopus and for fisheries indicators’ simulation on
a weekly basis (https://r​ drr.i​ o/g​ ithub/b​ en-m
​ hamed/D
​ EPLr/).
A depletion model in its most basic form assumes a
closed population in a fixed area with individuals removed
only through catch (DeLury 1947). Cephalopod stocks, how-
ever, do not always meet the assumption of being closed,
and further advances of the model have been implemented.
For assessing short-lived species over longer periods (e.g.,
the duration of a fishing season), natural mortality, while
difficult to measure (Caddy 1996), needs to be factored in,
in addition to catch, as a source of removal (Rosenberg et al.
1990; Agnew et al. 1998). In some stocks, immigration or
recruitment pulses recur during the depletion period and
need to be incorporated in the depletion model (Brodziak
and Rosenberg 1993; Payá 2007). Immigrations during the
depletion period are typically evident from sudden increases
in catch-per-unit-effort, but may not be distinguishable from
localized aggregations that were already there, undetected
by fishing vessels. A forthcoming challenge for depletion-
model stock assessment is then how to evaluate catch fluc-
tuations more objectively and, if possible, by automated
algorithms. Other challenges include the use of Bayesian
priors to stabilize the model computation, non-linearity of
the relationships between catch, abundance, and fishing
effort (an issue for all assessment models that use fishery
data), and varying natural mortality rates (Arkhipkin et al
2021a). With worldwide increasing trends in cephalopod
catches (Rodhouse et al. 2014; Doubleday et al. 2016),
continuing development of the depletion model is of high
importance. However, the need for intensive monitoring to
permit stock depletion (and within-season immigration) to
be followed in real-time results in this method being costly
and time-consuming to apply and it may not be feasible for
all fisheries.
Various other assessment approaches commonly applied
to finfish stocks have been used for assessing squid, octo-
pus and cuttlefish stocks (with the description of population
dynamics being simplified to reflect the lack of multiple
year classes) including cohort analysis, virtual population
analysis and yield-per-recruit models (e.g., Lange and Sis-
senwine 1983; Csirke 1987; Rosenberg et al. 1990; Roel and
13
 44 Page 12 of 46
Butterworth 2000; Royer et al. 2002, 2006; Hendrickson and
Hart 2006; Gras et al. 2014). In other cases, putative cohorts
have been identified using length–frequency data [which
carried the risk that within-year microcohorts (recruitment
pulses arising from asynchronous laying and hatching of
eggs) are interpreted as annual cohorts] prior to application
of such models (e.g., Agnesi et al. 1998; Orsi Relini et al.
2006; Giordano et al. 2010).
Survey catch rate is used as an abundance index in
several cephalopod fisheries (e.g., Kidokoro and Mori 2004;
Hendrickson and Showell 2016), while in the Saharan Bank
area, production models have been used routinely (Sato
and Hatanaka 1983). More recent variants of production
models (ASPIC, A Stock Production Model Incorporating
(environmental) Covariates; and SPiCT, Surplus Production
in Continuous Time) have also been applied to cephalopod
stocks (e.g., Abella et al. 2010; ICES 2020; Geraci et al.
2021) and simplified variants using only catch data have
been applied to multiple data-poor stocks in Europe
including various cephalopod stocks (Froese et al. 2018;
Tsikliras et al. 2021).
Finally, numerous forecasting models based on
environmental correlates of abundance and with varying
degrees of sophistication have been proposed (e.g., Osako
and Murata 1983; Coelho and Rosenberg 1984; Fogarty
1989; Otero et al. 2008) and this remains a promising
area for future development, even if success ultimately
depends on understanding the mechanisms underlying
empirical relationships between abundance and environment
(Moustahfid et al. 2021).
In assessing stock, a vital component is to quantify fish-
eries catches to calculate the biomass removed from the
stock by harvesting. Fishing vessel catch data are therefore
a fundamental component of stock assessment. While this
can be based on ship and observer logs for pelagic harvests
such as squid, most octopus catches rely on small-scale
artisanal fishers. Having a reliable system of traceability in
place could play a large part in helping to assess stocks more
objectively and accurately, whatever the physical character-
istics of the animals and methods of fishing.
Octopus stock assessment: the role of fishers
Stock assessment for squid is generally performed using
information from commercial fishing vessel logs and
manifests plus market data, and research fishing, in
combination with mathematical modelling. However,
octopuses are not usually captured on a large volumetric
scale by methods such as trawling (with some exceptions
such as Morocco) but by manually intensive methods using
the laying of inshore coastal traps, jigs, or pots by artisanal
fishers, or harvesting octopuses by hand in the intertidal
zone. Reliable information from artisanal fisheries has
13
Marine Biology (2024) 171:44
tended to be difficult to obtain in most areas: the fishers
are often secretive about the places they visit to ply their
skills, they may violate fishing regulations (i.e., use fishing
methods deemed illegal or ignoring limits on the number of
traps deployed), and in many cases, they are relatively poor
and therefore may be fearful of declaring their entire catch.
Also, they may be subsistence fishers who consume or barter
much of what they catch. However, in recent years, data on
standing stocks are being supplemented by observations in
the field from fishers in partnership with local authorities and
by harnessing the help of recreational divers. Participation of
fishers in European octopus artisanal fisheries management
ranges through co-management in Galicia (Otero et al.
2005) and active participation in Andalucia and Sardinia, to
top–down management systems in Algarve and the Thracian
Sea (Pita et al. 2021). However, sensibly, the involvement
of fishers has been increasing over the last few years, which
is desirable to successfully manage this species (Pita et al.
2021). A similar approach of management in collaboration
with fishers has been used in Morocco since the crisis of the
octopus fishery in 2003–2004 (MAPMDREF/DPM 2004).
Octopus taken by small-scale fishers is a component
of coastal subsistence culture in many parts of the world.
Examples include ‘Octopus’ tehuelchus in Argentina
(Storero and Narvarte 2024); and the giant Pacific octopus,
Enteroctopus dofleini (Wülker, 1910), the large size of which
allows this species a particular importance throughout its
range. Alaska Native and British Columbia First Nations
subsistence harvest of octopus traditionally occurs in the
intertidal during low tides, where octopuses are harvested
from their dens (Fall 1997). However, subsistence harvest
can also occur when octopuses are attracted to bait targeting
other species such as halibut fishing via hook and line, or
crab harvest via baited pots. As a typically nearshore fishery,
the octopus harvest may be sensitive to shore-based impacts
such as heavy use of an area by beachcombers, or oil spills
that directly coat the intertidal environment (Juday and Fos-
ter 1990; Benedetti-Cecchi et al. 2001).
On parts of the British Columbia coast, the cultural
importance of octopuses is revealed in the naw náaGalang
(or constructed octopus houses, maintained by the Haida
Gwaii), just one form of mariculture practised by coastal
indigenous people (Smith et al. 2019; Pacific Sea Garden
Collective 2022). Shelter and food can both be critical to
octopuses after they have settled from the planktonic stage,
both in terms of survival while growing and for brooding
eggs. Construction of octopus houses involves moving rocks
around on the beach to redirect or slow the outflow of water
with the tide, and form extensive tide pools. Rock piles
erected within the area are shaped to accommodate an octo-
pus, and a strategically placed removable stone on each pile
is used to make it easier to harvest an octopus from within.
Marine Biology (2024) 171:44
Once harvested, new octopuses move into the same suitable
habitat, often within a short period of time.
The impact of subsistence fishing for octopus is currently
of minor importance at the global scale, but most fishing for
Octopus vulgaris is by large numbers of artisanal fishers
who then sell their catch to middlemen. Off the coast of West
Africa, fishers currently are unlikely to cooperate in provid-
ing catch information at the point of capture, so traceability
must begin at the point of landing the catch and selling it on.
In parts of Spain and Portugal, however, local govern-
ments and scientists are beginning to work in partnership
with fishers, but currently this is at a pilot stage and is a
development for the future which, if successful, will be a
significant advance towards much more reliable traceability.
Such partnerships can be used to encourage consumer feed-
back and the establishment of niche seafood products, which
in turn may be a stimulus to provide more stock assessment
data to chase the increased value of popular niche harvesting.
Octopus stock assessment: the role of citizen science
In attempting to assess cephalopod stocks, typical fisheries
sampling methods may not provide an ideal approach: for
example, octopuses tend to be den-occupying ambush preda-
tors, spending much of their time inaccessible within rock
clefts, crevices and small caves, which cannot be sampled
easily. With practice, SCUBA divers can learn how to locate
octopus dens but scientific surveys based on such a strategy
are impractical for most scientists. Volunteer-based citizen
science, however, is an ideal approach to such surveys.
Citizen science involves members of the public volunteer-
ing their help to conduct scientific research: citizen scien-
tists may design experiments, collect data, analyze results,
and sometimes help to resolve particular research problems
(Bonney et al. 2009; Haklay et al. 2021; NOAA 2021).
Several attributes of scientific research projects are ideally
suited for enlisting the help of citizen science: scientific pro-
tocols are well designed and easy to learn and execute, and
in modern times, the Internet enables easy data submission
and acquisition of results. Some research is not practical
for scientists alone to execute. For example, large data sets
may be needed, but data collection can be labour intensive
and involve collection in the field of quantitative measure-
ments and observations over broad spatial and/or temporal
extents. Such projects can be pursued much more efficiently
by harnessing the help of volunteers in the broader human
population. An example of citizen science contributions to
research involving octopuses is the work by Scheel (2015)
and Scheel and Johnson (2021), who used data collected by
volunteer dive programmes (Seattle Aquarium and REEF,
respectively) to examine the impact of temperature and cli-
mate change on recruitment of E. dofleini in Washington
State waters.
Page 13 of 46 44
Traceability has already benefited from the application
of citizen science in a marine context (Giuffredi et al. 2021;
Agersnap et al. 2022), so adapting it to cephalopod stock
assessment is clearly feasible. Possible stock assessment
tasks for citizen scientists could include the collection of
images and their metadata (locality, time and ID) and quanti-
tative monitoring of sightings. Applications to traceability in
the future might involve eDNA sampling (e.g., to ascertain
distribution limits of certain species, particularly of octo-
pus), the testing and practical use of portable eDNA kits and
of traceability apps, which contribute to the enhancement of
traceability.
Fisheries management: achieving sustainability
In order for a species to be exploited sustainably, stock
assessment is fundamental (see the above parts of Sect.
“Status of cephalopod resources”), because this provides
a way of finding an appropriate estimate of a stock size
for which the fishery can be considered to be sustainable.
Most of the examples above have illustrated some of the
difficulties involved in stock assessment (e.g., a straddling,
migratory stock; or the problems of assessing numbers
of a cryptic benthic animal such as a species of octopus).
When reasonably accurate stock assessments are possible,
though, fisheries management becomes a more confident
process, presenting the means to safely exploit a seafood
resource which is sustainable in the long term, providing a
reliable source of nourishment, employment and profit for
the burgeoning human population. One of the best examples
of a well-managed cephalopod stock is the Falklands stock
of the squid Doryteuthis gahi.
Doryteuthis gahi, a model squid stock
The Falkland Islands economy is highly dependent on
two squid species: the ommastrephid Illex argentinus
(Sect. “Illex argentinus, the Southwest Atlantic shortfin
squid”) and the loliginid Doryteuthis gahi (d’Orbigny,
1835), accounting for 41% and 28%, respectively, of fish-
ery catches between 2012 and 2021. Together, they com-
prise 60–80% of annual commercial fishery catches from
Falkland Island waters (Agnew et al. 2005; Arkhipkin et al.
2013). Fisheries for both species are licenced in the Falkland
Islands Conservation Zones, and both have shown resilience
of their stock abundance and life-history parameters through
more than 3 decades of heavy exploitation (Arkhipkin et al.
2021b). However, management implementation differs sig-
nificantly between these two species, because I. argentinus
(captured mostly by vessels from South Korea, China and
Taiwan as part of international agreements; Falkland Islands
Government 2022) suffers from the problems of being a
straddling and transboundary stock (cf. also D. gigas and
13
 44 Page 14 of 46
Fig. 5  Exemplary label of a seafood package (from Vigo, Spain)
providing accurate information about the contents, the ‘calamar
patagónico’ (Patagonian squid) Doryteuthis gahi from the vicinity
of the Falkland Islands (which are situated on an extension of the
T. pacificus; see Sect. “Major cephalopod stocks”; Fig. 2).
Doryteuthis gahi has a more restricted coastal distribution
typical of squids in Family Loliginidae. This species is here
considered in a little more detail to demonstrate a species
that can be tracked through the system from being caught
in the Falklands through to various destinations around the
world; sales-chain information that, ideally, should be avail-
able for all species. Predictably, seafood products based on
Doryteuthis gahi are labelled clearly and correctly (Fig. 5),
in contrast to the sadly common occurrence of mislabelling
(typically fraudulent) which is characteristic of some other
squid species (cf. Sect. “Deliberate mislabelling”).
Doryteuthis gahi is a ‘domestic’ squid species completing
its entire life cycle in Falkland Islands waters (Arkhipkin et al.
2013), so the Doryteuthis gahi fishery is fully controlled by
the Falkland Islands Government. Major facets of Doryteuthis
gahi management include: spatial restriction of the target fish-
ery to one delineated area on the shelf break (as the only area
where small-mesh trawls, suitable for catching small squid, are
permitted); temporal restriction to two separate seasons per
year (corresponding to two annual cohorts) timed to end when
Doryteuthis gahi reaches spawning maturity; effort limitation
13
Marine Biology (2024) 171:44
Patagonian continental shelf). The ingredients label (top left) pro-
vides the species name as Loligo gahi (an older name for this species,
using the genus name Loligo instead of Doryteuthis) and the fishing
area, FAO 41, which is the SW Atlantic Ocean (Image: PFV)
to 16 vessels per season; and in-season monitoring to ensure
an escapement biomass of minimum 10,000 t at the end of
each season (Arkhipkin et al. 2008; Barton 2002). A survey
is conducted prior to the opening of each season to estimate
the Doryteuthis gahi recruitment stock and initiate in-season
assessment. Every survey includes fixed-station trawls cover-
ing the entire fishing area, and adaptive-station trawls selected
to increase the precision of biomass estimates in high-density
or school-edge locations. The dual approach ensures that the
scientific requirements of randomization and repeatability are
met, via fixed stations; and the spatio-temporal variability of
the Doryteuthis gahi population is captured via adaptive sta-
tions (Gawarkiewicz and Mercer 2019). As a single-jurisdic-
tion fishery, Doryteuthis gahi management can also respond
quickly to occurrences such as an exceptional ingression of
I. argentinus into the Doryteuthis gahi fishing zone (Winter
2015), or a sudden increase in the presence of pinnipeds (Iri-
arte et al. 2020).
The Falkland Islands Doryteuthis gahi stock has shown
increasing catch trends in recent years (Arkhipkin et al.
2021b), indicative of a successful management structure.
However, while accurate statistics about the stock itself are
Marine Biology (2024) 171:44
Fig. 6  Spider chart illustrating the approximate export footprint
tracking the trade in Doryteuthis gahi for 2018–2021. Mean annual
exports (tonnes) by the Falkland Islands (blue lines; line thickness
related to exported volume) and Spain (orange lines). Values are
‘squid exports’ data (UN COMTRADE database; UN 2023), com-
bining all commodity codes where squid can be reported (HS 6-digit:
useful (and exceptional compared to stocks of many other
species), it is important to keep track of the harvest throughout
the value chain and to understand more about the presence of
Doryteuthis gahi within the global market for cephalopods.
Falkland Islands flagged vessels reported that Doryteuthis
gahi represented 86% of their total production in 2020, so
squid exports accounted for most (c. 90%) of the seafood
exported from the Falkland Islands in the same year (FAO
2022b). Therefore, even though Falkland Islands trade sta-
tistics are not identified to species level, it can be inferred
that > 95% of ‘squid’ exports are Doryteuthis gahi. Destina-
tion market countries between 2018 and 2021 (Fig. 6) were
Spain (90%), Morocco (5%), South Africa (1%), and Ger-
many (1%). Namibia has also been a relevant importer (2%
of the Falkland squid exports in 2020), but did not import
from the Falklands in 2021 (S&P Global 2023).
In 2021, there was a considerable increase (64%) in Dory-
teuthis gahi production compared with the period 2012–20.
This surplus resulted in a corresponding increase in exports,
with trade flow affected at two main levels: (a) predictably,
there was an increase of imports by each of the main trade
partners: Spain (+ 20%), Morocco (+ 106%), South Africa
Page 15 of 46 44
030741, 030742, 030743, 030749, 160554) and may be a mixture
of data for several squid species (including other loliginids, ommas-
trephids and cuttlefish), particularly for Spanish exports. However,
Falkland Islands vessel reports of Doryteuthis gahi production repre-
sent > 95% of total Falklands squid production (FAO 2022b). Source:
UN (2023)
(+ 448%), and Germany (+ 11%); (b) there was a significant
increase in exports to some new markets or markets that
traditionally have not been key importers of this commodity
including 435 t to Senegal (a new importer in 2021) and 464
t to Russia, + 167% from 220 t in 2020 (UN 2023).
The data available in the international databases do not
fully depict the trade flows of this commodity, known for
being re-exported by Spanish processors both within and
outside the EU. For instance (and similar to what happens
with other related commodities such as I. argentinus; Harte
et al. 2018), it is likely that some of the Doryteuthis gahi
imported by Spain is processed and re-exported to other
Spanish key end markets within the EU (e.g., France, Italy,
Portugal, Croatia, Germany and Greece), or even to third
countries such as Morocco for processing and re-exporting.
According to US imports data, there was a considerable
increase in Spanish squid imports in 2021, but there are no
details of the specific amounts per squid species, because
squid imports are reported as either ‘Loligo’ or ‘squid’
(NOAA 2023).
The successful Doryteuthis gahi fishery stock manage-
ment model developed by the Falkland Islands Fisheries
13
 44 Page 16 of 46
Department has been accompanied by a successful mar-
keting strategy led by Spanish seafood companies feeding
southern European retail and food service supply chains.
Besides efforts to find new markets in Latin America and
Southeast Asia, the majority of the extra production of 64%
in 2021 and 78% in 2022 (compared with mean annual pro-
duction over the period 2012–20) has been absorbed by
southern EU food service and retail sectors due to several
characteristics confirmed by Spanish industry members of
the Global Squid Supply Chain Roundtable run by the Sus-
tainable Fisheries Partnership (SFP). These characteristics
are as follows.
1. Volume aggregation The entire production of the fish-
ery comes from 16 industrial vessels fishing for Doryteu-
this gahi in Falkland Islands waters with strong ties to, or
belonging to, just a few Spanish seafood companies, which
have been able to establish long-term business agreements
with major southern EU food services and retailers, thereby
optimizing marketing planning due to an annually assured
supply.
2. Increased processing Cleaning and gutting squid is
manual-labour intensive, which is a limiting factor for some
hotel, restaurant and catering (HoReCa) businesses that do
not have the necessary capacity or resources. This challenge
has been taken up through increased processing capacity for
cleaning and gutting at Spanish facilities, or in third-country
facilities owned by Spanish suppliers (e.g., in Morocco, Sen-
egal, Namibia and South Africa). This has broadened the
spectrum of HoReCa establishments willing to buy from
food service companies operating in the southern EU. At
the same time, this extra basic processing capacity has ena-
bled the development of new products that better meet the
needs of the retail sector, fostering the supply to southern
EU retailers as well as steadily contributing to open new
retail markets in the central and northern EU.
3. Seafood inflation During 2022, the EU experienced
high price inflation on all food products due to increased
production and distribution costs. Fortunately, in the Dory-
teuthis gahi fishery of the Falkland Islands, this has been
balanced out by increased efficiency of the fleet (probably
as a result of changes since 2016, when equal duration of
the fishing season was introduced for both seasons, exploit-
ing both cohorts: a strategy which almost doubled CPUE
which has not fallen back since, i.e., the ‘efficiency’ of every
vessel has almost doubled; Arkhipkin, pers obs). This has
helped to increase the competitiveness of Doryteuthis gahi
products in the EU, ultimately making them more attractive
to consumers and HoReCa businesses, and therefore a good
option both for household consumption and as a replacement
of other seafood items on restaurant menus.
4. Traceability The vertical integration of the fishing
fleet with Spanish processors, together with the EU import
requirements, enables full traceability of Doryteuthis gahi
13
Marine Biology (2024) 171:44
products from catch to consumer. Similar squid products
that can compete with Doryteuthis gahi in the EU and US
markets are known for having higher risks of IUU fishing
input and human rights violations due to the involvement of
unregulated distant-water fleets, bad practices, and opaque
processing, particularly in China (Asche et al. 2022; Selig
et al. 2022).
5. Sustainability performance Seafood sustainability is
now a major aspect of purchasing policy for any contempo-
rary retailer and food service company operating in EU or
North American markets. Evidence from independent third-
party sustainability assessments (such as those developed
by FishSource for this fishery; SFP 2022) that the fishery is
considered well managed and sustainably exploited is also
an important factor from a market perspective. This has
helped to build trust among producers, suppliers and major
EU and North American market operators.
These characteristics of the D. gahi industry have enabled
it to navigate the crisis brought about by the current COVID-
related effects on business much better than other seafood
industries, and also by the shortly preceding Brexit fiasco
(via the uncertainties which affected trading conditions
among the UK, Falkland Islands and the EU). Traceability
and sustainability included, these characteristics have also
aided this industry to expand its market penetration, reach-
ing new clients and creating new opportunities throughout
the EU and into North American countries (typically indi-
cated as USA, although Canada is involved via the free trade
agreement between them, with US importers of squid also
supplying Canadian food service and retail operators).
Where traceability is not as transparent and
­straightforward as in this Doryteuthis gahi industry, a
­fundamental requirement is to have clear criteria for ­species
or provenance identification.
Identification
Taxonomic identification of individual squid or octopus
species is important but tricky, and more so for those who
are not taxonomic experts such as behavioural ecologists,
fishers, or citizen scientists. Consider, for example, one of
the world’s most widely known and distinctive octopuses,
the giant Pacific octopus (GPO), Enteroctopus dofleini. The
genus name, Enteroctopus, has been used widely only since
the turn of the century (based on the revision by O’Shea
(1999), although it was first coined by Rochebrune and
Mabille 1889). Using museum specimens, Pickford (1964)
proposed that this species (as ‘Octopus dofleini’) is com-
posed of three subspecies, but by the 1990s that was not
supported (Gleadall 1993; Hochberg 1998). Now, however,
rather than several subspecies, it is becoming clear that the
GPO includes more than one species. Among the northern
Pacific Alaskan population, studies using DNA sequencing
Marine Biology (2024) 171:44
(Toussaint et al. 2012a, b; Hollenbeck and Scheel 2012,
2017) confirmed the presence of a second species, the
‘frilled giant Pacific octopus’ (scientific name pending).
This previously unnoticed species can be recognized visu-
ally by its body pattern characteristics and in some samples
it is more common than the well-known E. dofleini (Scheel
unpublished data from incidental catches in Prince William
Sound spot shrimp surveys by the Alaska Department of
Fish and Game; ADFG). However, the two species are often
confused by non-specialists and currently fisheries data
­consider only a single GPO, all under the name E. dofleini.
In an informal, unpublished trial, Scheel examined the
success rate at distinguishing between the two species of
GPO shown in pictures of the catch from the ADFG shrimp
survey (Scheel unpublished data from ADFG) depicting the
live octopuses in situ, out of water, on the deck of a ves-
sel or against some other neutral background. Two trained
observers agreed with each other and with genetic ID of
species for all but a single image, for which the opinion of
the two observers differed. People experienced in octopus
ID can be used to train artificial intelligence (AI; see Sect.
“Artificial intelligence, computer vision, machine learning
and deep learning”) to identify octopuses in images and,
where possible, IDs can be validated using DNA sampling
(but note the caveats in Sect. “DNA barcoding for species
identification”: it is important that DNA reference sam-
ples themselves have been identified correctly; Vecchione
et al. 2022). Such principles are involved in the construc-
tion of certain tracer apps built around the identification of
­cephalopod harvests (see Sect. “Traceability apps”).
Obstacles to successful identification have their history
in taxonomic practice to work from preserved specimens,
since preservation can destroy or obscure the differences
most obvious in life. Other reasons for (accidental) misi-
dentification include: misprints and incorrect cues in spe-
cies keys (e.g., Hochberg and Dorsey 1987, Fig. 14.5,
inadvertently illustrating E. dofleini as an ocellate species,
perhaps Octopus bimaculoides); the unique challenges of
naming and describing highly and continuously variable
octopus body patterns (Packard and Hochberg 1977; Roper
and Hochberg 1988; Hanlon and Messenger 2018); and our
incomplete understanding of cephalopod diversity. Many
species, including the frilled GPO, have yet to be identi-
fied unambiguously through publication of a scientifically
rigorous taxonomic description.
Even for the specialist, identifying organisms can be
subject to some ambiguity. Scientific ID traditionally relies
on morphological appearance but size and colour of ceph-
alopod individuals within a species may vary considerably,
and often it can be difficult to distinguish between varia-
tion among individuals within the same species and differ-
ences among species (Laptikhovsky et al. 2002; Anderson
et al. 2008; Gebhardt and Knebelsberger 2015; Alujević
Page 17 of 46 44
et al. 2022). Additionally, the use of common names is
a source of even more confusion (especially since they
may vary from place to place even within the same coun-
try), and is a major reason why the binomial (‘two-name’)
‘genus + species’ Latin-based system of Linnaeus is pre-
ferred for scientific purposes (Winston 1999).
In northern European cultures, cephalopods have entered
the diet relatively recently and in past centuries they were
used only as bait for catching finfish, so until the last few
decades there has been little interest in ID at the species
level: Simmonds (in Bartsch 1931), for example, recorded
long ago that squid species of genus Loligo are a favourite
food of the cod, Gadus morhua, and that half of the cod
caught by fishers off Newfoundland was taken using squid
as bait. A large amount of squid is still used today as finfish
bait by sport fishers in USA but typically the squid species
being used is unknown: as a bait item, fishers are not inter-
ested in which squid species they have (Moustahfid, pers
obs). Indeed, packages containing octopus arms labelled as
‘calamari’ (= ‘squid’) are not questioned by people unfamil-
iar with cephalopods (see Sect. “Misidentification”).
Taxonomic identification is gradually becoming easier
and more reliable with the advent of various databases freely
available on the Internet which provide some guidance for
both scientists and laypeople in how to recognize species.
However, it is difficult for one person to be expert at iden-
tifying any particular animal group because of the large
number of species, so it is easy for errors to creep in when
database compilers attempt to identify species with which
they are less familiar. Errors tend to be repeated and eventu-
ally find their way into all the available databases and there
is a time lag in updates and revisions to these databases.
There is also some inertia in accepting and incorporating
new advances in ID. In the field of octopus ID, for exam-
ple, recent improvements include more detailed descriptions
(e.g.Gleadall et al. 2010; Leite et al. 2008, 2021); clear rec-
ommendations to ensure the best conditions necessary for
preserving reference (‘type’) specimens (e.g., Gleadall et al.
2010); and emphasizing the importance of ensuring a direct
connection between DNA sequences and morphological
data using the same reference specimen (e.g., a holotype
specimen) for both data sources (Gleadall 2013). Improve-
ments are also underway (González-Gómez et al. 2023, this
special issue) to enhance the information available for the
Americas, which have long suffered from erroneous Euro-
centric IDs of the endemic American species of octopus
in view of similarities with species described in Europe,
such as Octopus vulgaris Cuvier, 1797, and Pinnoctopus
macropus (Risso, 1832). Some journal editors, reviewers
and researchers clearly are unaware of such improvements,
allowing the publication of inadequate descriptions of little
help to subsequent researchers, and which fail to present
essential basic morphometric data (even in supplementary
13
 44 Page 18 of 46
files), misinterpret previous studies, omit important and rel-
evant recent literature, and consequently cite outdated pub-
lications, and propose IDs which are fundamentally ques-
tionable because of the absence of any comparisons with
taxonomic ‘type’ specimens.
Over the last 30 years or so, there has been a trend to try
to circumvent the more difficult and ‘subjective’ IDs based
on morphology in favour of the use of ‘more objective’
gene sequence comparisons: the so-called ‘barcode’ tech-
niques such as species differences in the DNA sequence of
mitochondrial gene Cytochrome oxidase 1 (COI). However,
IDs based on sequence comparisons are only as good as the
morphological ID of the specimen from which the refer-
ence sequence was taken, and it has been recognized that
the number of errors of names associated with sequences
deposited in repositories such as GenBank is much larger
than one might expect (Groenenberg et al. 2009; Vecchione
et al. 2022). Although IDs are not guaranteed by the owners
of databases such as GenBank, many users seem to accept
unquestioningly the ID accompanying each registered
sequence. This lack of reliability in DNA sequence data-
base curation has consequences outside of pure science: it
has resulted in at least one recorded instance where a sus-
pected species substitution fraud was rendered inconclusive,
because DNA sequences from publicly available databases
indicated close matches to more than three different squid
species, preventing any criminal prosecution from taking
place (Moore 2020). (That is, samples identified by three
different species names in the ‘barcode’ database had clearly
been taken from individuals of the same species.)
For both octopus and squid, discrimination of species
can be challenging, even when appropriately paired mor-
phological and sequence data are available. The close simi-
larity among species that actually are different may well be a
property of the rapid speciation considered to have occurred
recently (and probably still in progress) in modern cepha-
lopods (Nesis 1999). Therefore, for example, the morpho-
logical differences between the squid species Alloteuthis
media and A. subulata (distinguished on the basis of their
morphology by Naef 1923) overlap to the extent that for
some authors they are regarded as variations among a single
species (Laptikhovsky et al. 2002), although DNA analysis
suggests that they are indeed different (Anderson et al. 2008;
Gebhardt and Knebelsberger 2015; Alujević et al. 2022).
Among octopuses, ‘common’ octopuses around the world
were all identified until recently as Octopus vulgaris but
are now recognized as a complex of different species with
their own discrete biogeographical limits, as confirmed by
analyzing differences in nuclear and mitochondrial COI
gene sequence, as well as subtle morphological differences
detected subsequently. Off the American Atlantic coasts,
for example, there are two species: Octopus insularis Leite
and Haimovici, 2008, in warmer shallower waters; and O.
13
Marine Biology (2024) 171:44
americanus Froriep, 1806, in cooler deeper waters. In East
Asia, the common octopus is now identified as O. sinen-
sis d’Orbigny, 1841 (Gleadall 2016), and a similar species
in the Kermadec Islands as O. jollyorum Reid and Wilson
2015. However, despite a gap between their populations of
nearly 9,000 km and a distinctive morphological difference
between males of O. sinensis and O. jollyorum, they are
recognized by some researchers as but one species, based
on the (apparently objective and infallible) DNA sequence
similarity of the COI gene (Amor et al. 2016; but see Amor
and Hart 2021). Possible cryptic diversity (that is, the pres-
ence of more than one species where only one species has
been detected morphologically) has been proposed also for
the squids Loliolus beka, Uroteuthis edulis; and octopuses
Octopus minor, Amphioctopus fangsiao, and Hapalochlaena
lunulata from Chinese waters (Xu et al. 2022).
Identification of cephalopods in commercial products
In the drive to improve species ID (especially to overcome
the widespread problem of species substitution in the mar-
ket; see Sect. “Misidentification”), seafood species have
been identified by a range of different techniques (Silva and
Hellberg 2021). However, techniques such as the analysis
of allozyme polymorphisms have serious limitations when
applied to food, because fresh tissue is necessary. Identifi-
cation techniques based on DNA analysis overcome such
limitations, because DNA molecules are more stable to
different treatments (heating, salting and smoking) than
proteins (Chapela et al. 2002). Table 3 is a list of those
techniques used for cephalopod species in commercial
products. More than half of these studies have relied on
genetic barcoding using the COI gene, amplifying, and
BLAST searching of new sequences against public DNA
databases using BOLD and the NCBI web pages (Table 3).
Detecting the provenance of samples is another approach
that is increasing in use, because it is a way to trace a sea-
food item without restriction to species that may be difficult
to identify taxonomically at the species level. Beginning in
2022, chemical signatures are used to trace the origin of
cephalopod products on both domestic and international
scales, based on analysis of fresh or frozen soft tissues and
particular octopus parts such as statoliths or ink. Stable
carbon and oxygen isotope analyses (isotope-ratio mass
spectrometry) of statoliths and elemental analyses (X-ray
fluorescence using Itrax) of soft tissue of octopus have
been combined successfully to deduce the provenance of
four octopus species at international and domestic scales
(Martino et al. 2022). Total X-ray fluorescence-based multi-
elemental profiling of O. vulgaris ink has successfully dis-
tinguished among four collection areas in Portugal (Duarte
et al. 2022). Spectral fingerprints obtained with near-infra-
red (NIR) spectroscopy have been patterned by different
Marine Biology (2024) 171:44 Page 19 of 46 44

Table 3  Methods used for cephalopod species identification and provenance identification in market products
Method Target gene Species number* N tested, mislabeling% References

Species identification
­AFLPa 2 S, 1 O, 1 C Maldini et al. (2006)
PCR-RFLPb 16SrRNA 5S Colombo et al. (2002)
­FINSc 16SrRNA 9 S, 1 O 8 Chapela et al. (2002)
cytb 13 O, 11 C, 4 S 20 (30%) Espiñeira et al. (2010)
PCR–RFLP + FINS cytb 8S 18 Chapela et al. (2003)
cytb 21 S, 2 O, 1 C 15 (33%) Santaclara et al. (2007)
cytb 4 S, 1 O 5 (60%) Pardo et al. (2018)
Real-time PCR COIII, 16SrRNA, ITS1 26 S, 16 C, 15 O 34–42 Espiñeira and Vieites 2012);
Herrero et al. (2012)
COI, cytb, ATPase 6, 12SrDNA 4S Ye et al. (2016)
COI, 16SrRNA, 12SrRNA 5 S, 5 O, 1 C Chung et al. (2018)
COI 2S 48 Shi et al. (2020)
COI 11 S, 5 O, 4 C 17 (25%) Velasco et al. (2020)
Multiplex PCR 16SrRNA 5 S, 2 O 8 Kim et al. (2015b)
COI 3O 30 Lee et al. (2022)
DNA barcoding COI, cob, 16SrRNA 1 S, 1 O Nicolè et al. (2012)
COI, cytb 5O 51 (55%) Debenedetti et al. (2014)
COI 6 (33%) Armani et al. (2015)
COI 1O 1 (0%) Khaksar et al. (2015)
COI, 16SrRNA 7O 59 Martin (2015)
COI, 16SrRNA 3 C, 2 S 19–18 (52–22%) Galal-Khallaf et al. (2016)
COI 2 O, 1 S Shen et al. (2016)
COI, 16SrRNA 6 S, 4 C, 2 O 66(48%) Guardone et al. (2017)
COI, 16SrRNA 5 S, 4 C, 1 O 95 (2%) Wen et al. (2017)
COI 2 S, 1 O 8 (0%) Marin et al. (2018)
COI 1 S, 1 C 7(57%) Sarmiento et al. (2018)
COI 2 S, 1 O 11 (27%) Tinacci et al. (2018a)
COI 4 S, 3 C, 2 O 16 (19%) Tinacci et al. (2018b)
COI 1O 17 (53%) EJFEFJ (2019)
COI 4 S, 3 C, 2 O 59 (31%) Fernández et al. (2020)
COI 3 O, 1 S 6 (0%) Wallstrom et al. (2020)
COI 1S 35 (23%) Sun et al. (2021)
“ + NanoTracer COI 5 O, 4 S, 3 C 150 (33%) Maggioni et al. (2020)
­LAMPd COI 1S Ye et al. (2017)
COI 3S Tatulli et al. (2020)
RPA-LFAe COI 8 S, 5 O, 1 C 32 (3%) Velasco et al. (2021)
Nanopore COI 2C Ho et al. (2020)
Sequencing COI, cytb 2S Toxqui-Rodríguez et al. (2023)
Provenance identification
­NIRSf 2 C, 5 areas Curró et al. (2021)
NIRS 2 O, 2 areas Varrá et al. (2022)
­XRFg, ­IRMSh 4 O, 5 areas Martino et al. (2022)
XRF 1 O, 4 areas Duarte et al. (2022)

Species numbers are subdivided as C, cuttlefish; O, octopus; and S, squid

*Included here are only those papers working with commercial samples. There are many other publications dealing with genetic studies of
cephalopod species taken from the wild using barcoding genes but these are not relevant to commercial harvest sampling
a
Amplified fragment length polymorphism; bPolymerase chain reaction—restriction fragment length polymorphism; cForensically informative
nucleotide sequencing; dLoop-mediated isothermal amplification; eRecombinase polymerase amplification and lateral flow assay; fNear-infrared
reflectance spectroscopy; gX-ray fluorescence spectrometry; hIsotope-ratio mass spectrometry

13
 44 Page 20 of 46
traditional and modern (e.g., ML) tools to identify the origin
of two species of octopus from two different areas (Varrá
et al. 2022). Indeed, an array of tools has now been used
to test provenance traceability in marketable cephalopods
(Table 3).
Legislation for identification and labelling
Current legislation on ID for seafood products worldwide
still has many gaps (see Sect. “IUU fishing and a summary
of measures in place to combat it”). Mandatory seafood
labelling provisions such as for scientific and common
names is still quite limited (D’Amico et al. 2016). The fol-
lowing brief review of cephalopod product legislation con-
siders only countries most relevant to cephalopod trade and
consumption including China, Korea, the EU and the US.
‘The Seafood List’ is the US FDA ‘Guide to Acceptable
Market Names for Seafood’ sold in interstate commerce.
It contains around 40 cephalopod common and scientific
names. However, the market names considered acceptable
are just ‘squid’ or ‘calamari’, ‘cuttlefish’ and ‘octopus’,
which are ambiguous generic names providing incomplete
information for consumers (Wallstrom et al. 2020).
A recent review on the labelling of seafood products
(D’Amico et al. 2016) includes the most recent EU regula-
tions. Mandatory information for any product containing
live, fresh, chilled, frozen, dried, salted or in-brine cepha-
lopods include scientific name and provenance. However,
for prepared or preserved (canned) cephalopods, no such
information is mandatory. Neither is it mandatory for res-
taurants or other mass caterers, which may indicate cepha-
lopods on their menu by generic or otherwise incomplete
names.
Each EU country has its own legislation regarding cepha-
lopod commercial names (Herrero et al. 2012; Xiong et al.
2016). The list for Spain established in 1986 had many short-
comings such as labelling canned cuttlefish (Sepia spp.) as
longfin squid ‘calamar’ (Herrero et al. 2012). Currently,
individual countries draw up and publish a list of the com-
mercial designations accepted in their territory including
accepted local or regional names (European Commission
2022). Mediterranean countries list more cephalopod species
than the rest of Europe but they still contain scientific names
that are now unaccepted, such as Octopus dollfusi (currently
identified as Amphioctopus aegina).
The Chinese fishery sector suffers from significant leg-
islative and managerial shortcomings and, unlike many
other countries, lacks specific provisions for the labelling
of fishery products and an official reference list of seafood
trade names. In fact, except for establishing some basic
compulsory labelling requirements, only a few standards
(some mandatory and some voluntary) have been issued.
13
Marine Biology (2024) 171:44
Indication of the scientific name or provenance of seafood
is optional (Xiong et al. 2016). The popularity in China of
dried cephalopod products, which are even more difficult to
identify, paves the way for species substitution (Wen et al.
2017; Shi et al. 2020).
South Korea has sought to introduce a Seafood Trace-
ability System (STS) providing a record of a product briefly
using barcode- and QR code-based information technology.
However, its application is voluntary and does not include
any of the imported species. Lack of strong implementation
measures from the government, resistance from the industry
and producers, low incentive and limited interest from con-
sumers have all contributed to the current failure of the STS.
Thus, the Korean market is still vulnerable to IUU products
that lack proper hygiene and regulatory standards, which
has serious implications considering the nation’s heavy
dependence on seafood (EJF 2019). A number of funda-
mental improvements in regulations and laws for labelling
are clearly needed (see Sect. “Misidentification”).
The fishery sector fishing octopus from particular areas
has sought to obtain an official certificate of designation
of origin. Unofficial designations such as ‘Polbo das Rias’
(estuary octopus) for Octopus vulgaris from Galicia might
legally protect the traditional way of preparing octopus in
typical local fairs and pilgrimages, thus adding value to the
product, and providing access to national and international
markets (Espiñeira and Vieites 2012). A similar effort (still
pending) has also been included in the management plan
for O. maya, a Caribbean endemic of the Campeche Bank,
Yucatán Peninsula, Mexico (DOF 2014).
Misidentification
Given the technical difficulties of ID to distinguish species
with the naked eye (Lee et al. 2022; Sect. “Identification”),
the occurrence of accidental misidentifications is not sur-
prising. Because of the difficulties (even for experts) of iden-
tifying species correctly, misidentifications and consequent
unintentional substitutions can and do occur. These may be
due to similar geographic distribution of species such as the
common octopus, Octopus vulgaris, and the lesser octopus,
Eledone cirrhosa, which have overlapping distributions and
similar morphological characteristics. Also, the juveniles of
different octopus species can be misidentified at any level in
the value chain, even at the point of capture.
The complexity of cephalopod trade flows and variations
in, or lack of, labelling systems and official lists of seafood
trade names in different countries can make it difficult to
accurately identify the origin of species used in cephalopod
products, especially in processed preparations where poten-
tially identifiable anatomical features have been removed.
These problems, along with often-inadequate traceability
measures, create opportunities for exploitation through
Marine Biology (2024) 171:44
product mislabelling or substitution with species of lower
commercial value, as well as abusive practices such as the
addition of water to artificially increase product weight and,
ultimately, the price (Ospina-Alvarez et al. 2022).
Misidentifications can contribute to miscalculation of
stock assessment for both the actual species (for which, con-
sequently, the stock will be under-assessed) and the species
whose name is used to identify the organism (for which the
stock will be over-assessed). For both organisms, this results
in inaccuracies in tracking the species from capture to con-
sumer, and ultimately errors in stock management (based on
errors of stock assessment) which compromise the sustain-
ability of both resources. For these reasons, misidentification
must be minimized but currently the problem is massive and
can account for up to half the estimated stock of particular
species (Naaum et al. 2016).
Academic reports on specimen substitution are some-
times susceptible to technical problems when attempting to
assess the extent of misidentification and mislabelling. For
example, there are two reports on frozen products imported
to Italy (Debenedetti et al. 2014; Guardone et al. 2017) that
include labels identifying some of the samples as ‘Octopus
membranaceus’. This is a poorly characterized species of
uncertain distribution, but which is the type species of the
small octopus genus Amphioctopus (i.e., the correct identi-
fication is Amphioctopus membranaceus). The species iden-
tification reference used in these reports was the first edition
of the FAO catalogue of world cephalopods (Roper et al.
1984), published during a period when genus Amphiocto-
pus was not recognized and ‘Octopus membranaceus’ was
thought to be synonymous with a number of other species
that are now recognized as distinct, though related. Among
these is Amphioctopus fangsiao (cf. Gleadall 1997, 2003),
which is a highly exploited East Asian species (Sauer et al.
2021). Both Debenedetti et al. (2014) and Guardone et al.
(2017) used sequences deposited in FISH-BOL, GenBank
and BOLD to identify DNA sequences from samples that
had been labelled as O. membranaceus. Some of these
sequences were identical with a database sequence identified
by the submitter as A. fangsiao ‘from Vietnam’ (a species
with a reported distribution only as far south as Taiwan: no
taxonomic studies have confirmed the presence of A. fang-
siao in the South China Sea). Other samples were identified
as belonging to the genus Cistopus from Thailand and India.
Debenedetti et al. (2014) and Guardone et al. (2017) sug-
gested that both these labelling and identification conflicts
were due to intentional substitutions (deliberate mislabel-
ling; see Sect. “Deliberate mislabelling”, below), consider-
ing that O. membranaceus was a species under heavy exploi-
tation and that the substituting species have a lower market
value. Debenedetti et al. (2014) also reported that a speci-
men labelled ‘Octopus membranaceus’ was identified as
O. aegina, but 21 other samples of ‘Octopus membranaceus’
Page 21 of 46 44
could not be identified using barcode analysis. This high-
lights the need to enlarge on the information available in the
reference databases, since for some octopus species, there
are no reference sequences available, either in GenBank or
BOLD (Debenedetti et al. 2014). It is clearly important to
tighten up the identifications recorded in the reference data-
bases and regular reviewing is necessary to weed out any
misidentifications. Also, it is important that the database ref-
erence sequences used are named (by their accession num-
bers) in identification studies. Unfortunately, the studies by
Debenedetti et al. (2014) and Guardone et al. (2017) did not
record the reference accession numbers (confirmed by pers
com to author IGG), so it is not possible to comment further
on the identifications used in those studies. The conclusion
is that many of the samples reported as O. membranaceus
were probably honest misidentifications (not deliberate but
based on the erroneous information available at the time).
However, specimens labelled as ‘Octopus membranaceus’
but identified as Cistopus spp. may well have been exam-
ples of deliberate mislabelling, since Cistopus is a genus of
medium-large species of inferior palatability (in both taste
and consistency) and, consequently, of inferior value.
Deliberate, fraudulent misidentifications fall into two
major categories: deliberate mislabelling (Helyar et al. 2014;
Naaum et al. 2016); and bogus identification and labelling of
catches from illegal, unregulated and unreported (IUU) fish-
ing (Macfadyen et al. 2019; Nodzenski et al. 2019; Donlan
et al. 2020). Among both these categories is included the
practice of ‘re-exporting’ as a strategy to mislead concern-
ing the true provenance of the product (Asche et al. 2022).
Deliberate mislabelling
Ensuring that seafood is accurately labelled is of great
importance to avoid negative impacts on marine popula-
tions and to achieve sustainability goals but problems with
mislabelling are significant. For example, surveys estimate
that 13% to 37% of seafood sold in the USA is incorrectly
labelled (Lewis and Boyle 2017). It is known that fishers
often mislabel seafood, especially if they caught it illegally
(IUU fishing; Warner et al. 2014; see Sect. “IUU fishing
and a summary of measures in place to combat it”). Inten-
tional species substitution can aid the introduction of ille-
gally caught fish to international markets, thereby supporting
consumption of products from poorly managed fisheries. For
instance, 55% of 44 samples labelled as ‘shark’ in Brazil
were identified as largetooth sawfish, an IUCN Critically
Endangered species the trading of which is prohibited (e.g.,
Faria et al. 2013). Such practices may give consumers the
false impression that supply is keeping up with demand,
while declines in the populations of threatened species go
unnoticed. Moreover, the fact that certain species names
appear commonly in restaurant menus makes it harder to
13
 44 Page 22 of 46
argue for increased protections for them when consumers
think that these populations are healthy and abundant given
their apparent ‘availability’.
Some species of fish and invertebrates that are used as
substitutes carry the risk of negative human health impacts
such as allergies and poisoning (Warner et al. 2016; Fer-
nandes et al. 2020; Kroetz et al. 2020). For example, samples
from Chinese shops in Italy labelled as ‘squid’ have been
identified by DNA sequencing as puffer fish in the genus
Lagocephalus, which is banned from European markets due
to the presence of life-threatening tetrodotoxin in certain tis-
sues (Armani et al. 2015). Some ‘squid balls’ marketed in
the Philippines are known to be made from pork or chicken
(Sarmiento et al. 2018) and ‘cuttlefish balls’ from Singa-
pore may include pork, mislabellings with the potential to
cause stress and conflict in a multi-religious market (Ho
et al. 2020).
In addition to species substitution, there are many other
kinds of seafood mislabelling to hide the correct identity,
origin or weight of the seafood. Examples include: misrep-
resentation of provenance (Forbes and Alexander 2015);
sourcing of farmed products subsequently labelled as wild
(Aursand et al. 2009); frozen and thawed products labelled
as fresh (see, for example, Pennisi et al. 2021); and short-
weighting or over-glazing (Silva and Hellberg 2021). The
objective of all such mislabelling is economic fraud: to
obtain more profit by assigning the name of a higher-quality
(tastier and more valuable) provenance, species or product
to seafood or other product considered much less palatable
or of lower quality, which would normally trade at a much
lower price. Those guilty of mislabelling are most often
the distributors or final seafood retailers (e.g., fishmongers,
restaurants and supermarkets; Jacquet and Pauly 2008).
Customers then lose out by purchasing low-cost seafood at
prices corresponding to high-value products. For instance,
when canned low-quality squid is mislabelled as ‘octo-
pus’, the retail price can quadruple (R. González-Gómez
pers obs), as does the corresponding loss to the consumer.
Fishermen and businesses abiding by the regulations then
struggle to compete with such cheaper products that move
more easily through the value chain when they are being
mislabelled and fraudulently sold on as more-expensive,
high-demand products.
The statistics of mislabelling reflects the fact that cepha-
lopod species tends to have a similar appearance, especially
to an untrained consumer, so it is relatively easy to take
advantage of difficulties of visual ID and distinction. From
an increasing number of market surveys (often through DNA
sequence testing and other approaches), it is known that
the incidence of (mostly deliberate) seafood mislabelling
is high. A recent study by Guardone et al. (2017), analyz-
ing 277 products belonging to different seafood categories
(fish, cephalopods, crustaceans, bivalves and amphibians)
13
Marine Biology (2024) 171:44
imported at an Italian port, showed that 23% of the analyzed
products were mislabelled. The highest percentage of misla-
belling was observed for cephalopod-based products (44%),
followed by crustaceans (17%) and fish (14%). A higher rate
was found in products imported from Asia. Percentages of
mislabelling in cephalopod products as determined by DNA
sequencing methods greatly vary between studies, although
mislabelling rates of 30–50% are not uncommon. Typi-
cally, species of higher commercial value are substituted by
another of lower value (FIS 2018). Longfin squid (Loligo)
are commonly substituted by shortfin squids of the family
Ommastrephidae such as genera Illex and Todarodes (Pardo
et al. 2018).
Legal gaps in the commercial naming of cephalo-
pod species have been underlined as factors which lead
to widespread mislabelling in the market (Herrero et al.
2012). Lack of mandatory requirements to report cepha-
lopod species for some products also encourages species
substitution. Labels on most fresh products and prepared
dishes, as well as canned cephalopods, fail to provide
cephalopod species information (Espiñeira and Vieites
2012). Labelling without providing scientific names rep-
resents from a third to two thirds of all cephalopod prod-
ucts in supermarkets (Espiñeira and Vieites 2012; Herrero
et al. 2012; Fernández et al. 2020; Velasco et al. 2021).
Mislabelling also extends to the mass-catering sector
(Pardo et al. 2018). The fact that each country has its own
legislation and that also inside the same country there are
different valid regulations for a given species or taxonomic
group, depending on the format in which it is marketed,
makes the standardization of cephalopod legislation more
difficult and cephalopod commercial naming ambiguous
(Herrero et al. 2012).
Octopus and squid commercial landings data are sub-
ject to the usual issues of under- and mis-reporting (see
Sect. “IUU fishing and a summary of measures in place
to combat it”) and, in many cases, the taxonomic identity
of the targeted species remains unknown or is taken for
granted because official fishery statistics do not attempt to
distinguish different species. Uses of umbrella or generic
common names, such as ‘octopus’ or ‘squid’, leads to
species substitution (Espiñeira and Vieites 2012; Velasco
et al. 2021), thereby confusing consumers and regula-
tors (Fernández et al. 2020). Squid landings are typically
classified as ‘long-finned’ (loliginid) or ‘short-finned’
(ommastrephid), while octopus catches are often referred
to as ‘unidentified octopus’ or ‘common octopus’, Octo-
pus vulgaris (Pierce et al. 2008; Norman and Finn 2016;
Domínguez-Contreras et al. 2018).
The generic name ‘baby octopus’ (probably implying
‘baby Octopus vulgaris’) is often applied to the adults of
several species from the small-sized genus Amphiocto-
pus marketed widely around the world (Jereb et al. 2014).
Marine Biology (2024) 171:44
Amphioctopus aegina from Indonesia and Vietnam is often
sold as ‘baby octopus’ in Australian (Martino et al. 2022)
and Hawaiian markets (Wallstrom et al. 2020), and A. aegina
and A. marginatus are sold as ‘baby octopus’ or ‘little octo-
pus’ in California (Martin 2015). Amphioctopus marginatus
was the only octopus recorded in Chinese markets by Wen
et al. (2017). Half of the Octopus vulgaris sold in Seoul
markets was substituted by A. marginatus (EJF 2019), which
has been found also as a substitute for O. vulgaris in the Ital-
ian market (Nicolè et al. 2012). In addition, possible cryptic
diversity in cephalopod species from Chinese waters (Xu
et al. 2022) suggests that many other Amphioctopus species
might be involved in the international market.
Frozen octopus sold in Spain may include American spe-
cies such as O. maya and O. mimus (Espiñeira and Vieites
2012; Fernández et al. 2020; Maggioni et al. 2020; Velasco
et al. 2021). Espiñeira and Vieites (2012) conducted a study
to identify fraud related to the labelling of food products in
Spain and found that approximately 67% of the products
(including fresh and canned products, prepared dishes, and
frozen products) had no species name. According to those
authors, this is a consequence of the current labelling regu-
lations in Spain, which do not require the particular species
to be declared in certain food formats (whole, tentacles, in
pieces, etc.). The study revealed that in prepared and canned
products, even when the species name was not included, the
commercial name was indicated as well as the commercial
format (terms for the latter being currently under review;
Secretaria General de Pesca 2020).
Instances of mislabelling are especially difficult to detect
in processed foods such as breaded or canned products
because morphological characteristics of the species are
entirely lost due to the different treatments to which they
have been subjected, including peeling, cutting, cook-
ing and canning. In this regard, it is noted that the jumbo
squid D. gigas has suckers that contain an interior rigid
ring equipped with formidable triangular teeth and its flesh
contains ammonia (see Sect. “Dosidicus gigas, the South-
east Pacific Humboldt or jumbo flying squid”). To make it
a palatable product, some companies subject it to drastic
physical and chemical processing in which the sucker rings
are removed and the ammonia off-flavours are corrected fol-
lowing a chemical treatment with hydrogen peroxide and
a cocktail of phosphates. As a result, the arms of a squid,
the most captured cephalopod in the world, chopped and
cooked, have an octopus-like appearance. Several European
companies take advantage of this fact (and the existence of
legal gaps related to labelling) and sell processed jumbo
squid as ‘octopus-like’ canned products in what is a clear
example of economic fraud.
The common octopus of the eastern Atlantic, O. vul-
garis, has worldwide socioeconomic importance and is
the most valuable cephalopod on the market (Pardo et al.
Page 23 of 46 44
2018), related to its superior taste and the fact that until
relatively recently it was regarded as a cosmopolitan species
(see Sect. “Octopus vulgaris, the common octopus of the
Eastern Central Atlantic”). Many octopus products in Spain
including prepared dishes, which do not have any species
label contain O. vulgaris (Espiñeira and Vieites 2012). It is
probably the most appreciated cephalopod on the Spanish
market due to the traditional way of preparing it as ‘Gali-
cian octopus’. Even though it has no official certificate of
designation of origin, such ‘brand names’ are used to legally
protect certain foods that are produced in a given area, thus
adding value to the product and providing enhanced access
to national and international markets.
Such brand features of O. vulgaris in association with the
high prices it can attain in the market generally, due also to
limited supplies from the main exporting countries, often
result in fraud in which the name Octopus vulgaris is used to
label other cephalopod species of lower value such as other
octopuses (e.g., Eledone sp., the ‘lesser octopus’) or even
species of squid, particularly Dosidicus gigas (Santaclara
et al. 2007; Espiñeira et al. 2010; see Sect. “Dosidicus gigas,
the Southeast Pacific Humboldt or jumbo flying squid”):
one of the most common frauds of seafood species substi-
tution involves squid labelled as octopus (Lawrence et al.
2022). The most common substitutes for O. vulgaris in the
literature are D. gigas and Amphioctopus spp. (Velasco et al.
2021). Some cans show misleading advertisements on the
main package label such as ‘octopus style’, but are labelled
elsewhere in small font as Dosidicus gigas or with generic
names such ‘cephalopod’ (Espiñeira and Vieites 2012; Her-
rero et al. 2012; Velasco et al. 2021). Mislabelling has been
reduced somewhat with the recent practice of at least includ-
ing the true species name somewhere on the package.
At the opposite end of the quality range to O. vulgaris, D.
gigas is currently the leading squid in terms of total catch
landings worldwide and it is the most common cephalopod
used as a substitute, due to its low price. It is the most fre-
quently represented cephalopod (from 48 to 100% of com-
mercial samples) in Chinese markets and the only species
retrieved in shredded and grilled, sliced and dried ‘octo-
pus’ and ‘cuttlefish’ in Zhangzhou and Qingdao markets
(Wen et al. 2017; Ye et al. 2016, 2017; Shi et al. 2020). It
is also found in ‘squid balls’ and ‘fish balls’ imported from
some countries in Southeast Asia (Sarmiento et al. 2018).
Breaded precooked slices labelled as Illex argentinus from
Spain to Bulgaria are also substituted by D. gigas (Tinacci
et al. 2018a).
Fernández et al. (2020) conducted a seafood labelling
study in three major supermarkets located in Spain and
found that 68% of cephalopod commercial products were
improperly labelled (either incorrectly, not fully informa-
tive, or incomplete) including species mixes and substitu-
tions (31%). Among other findings, the authors highlight the
13
 44 Page 24 of 46
following: (a) a product labelled as the ommastrephid squid
Illex argentinus but identified by DNA sequencing as cut-
tlefish, Sepia sp.; (b) a product labelled as a loliginid squid,
Loligo sp., but identified as a mixture of the ommastrephids
I. argentinus and D. gigas; and (c) a product labelled as
Octopus vulgaris but identified as O. maya from the Gulf of
Mexico and Caribbean.
For the present review, one of the authors (RGG) inves-
tigated a number of products available in supermarkets in
southeastern Spain (in November, 2022) and found that,
although labelled with phrases such as ‘Octopus in Galician
sauce’ (implying O. vulgaris), the packs all contained jumbo
squid, D. gigas (Fig. 7). Despite being correctly labelled in
the ingredient list as D. gigas, the ambiguity between the
commercial name of the product (implying that it is octo-
pus) and the labelled and identified squid species is clearly
intended to mislead consumers into purchasing something
quite different from what they expected. Similar mislabel-
ling examples include D. gigas deliberately mislabelled as
‘octopus in oil’, ‘octopus in Galician sauce’, ‘octopus pieces
in garlic sauce’, ‘octopus-squid in garlic sauce’ or ‘octopus-
squid in marinara sauce’ (Fig. 7), suggesting that this prac-
tice is pervasive.
These findings raise serious concerns about the transpar-
ency of the seafood supply chain and highlight the need for
traceability studies and continuous monitoring by the regu-
latory agencies. This particular fraud appears to be exten-
sive, since annual Spanish imports of D. gigas from Peru
have averaged around 60,000 t in the last three years (FAO
2021b). According to Harte et al. (2018), not only has Spain
remained Peru’s largest trade partner over the past decade,
with 24–38% of squid export by value going to Spain, but
also some companies are currently processing 6000 t per
year and are also doubling their cold storage capacity. More-
over, lower octopus catch quotas imposed in Morocco will
continue to tighten the supply of genuine O. vulgaris, and
consequently, prices can be expected to continue rising, a
situation that seems likely to boost fraudulent practices (and
profits) related to mislabelling.
The misrepresentation on these labels (obvious to some-
one familiar with the scientific names of Cephalopoda) vio-
lates the general principles of the Codex Alimentarius stand-
ard for labelling, as well as many of the mandatory labelling
requirements (FAO and WHO 2018). However, despite this
misrepresentation, it is not illegal per se, because the Codex
guidelines are recommendations for voluntary application by
member countries, serving as a basis for national legislation.
Enforcement is therefore the responsibility of each coun-
try. There is also a loophole in the Codex regarding country
of origin: ‘When a food undergoes processing in a second
country which changes its nature, the country in which the
processing is performed shall be considered to be the coun-
try of origin for the purposes of labelling.’ (FAO and WHO
13
Marine Biology (2024) 171:44
2018). Therefore, after processing, currently, these Pacific
D. gigas products can be classified, according to the Codex,
as a product of Spain. Some cans of Spanish origin have
reached the international market to the US (Lingwall 2017),
where two New York food processing and distribution com-
panies imported, processed, marketed, sold, and distributed
more than 50 t of D. gigas from Peru worth $1.1 million
that had been falsely labelled as octopus (US Department
of Justice 2019).
To establish a common organization of the markets
regarding fishery and aquaculture products, current EU
regulations concerning seafood dictate that, in order for
these to be sold, appropriate labelling should indicate: (a)
the commercial designation of the species and its scientific
name; (b) the production method; (c) the area where the
product was caught or farmed, and the category of fishing
gear used; (d) whether the product has been defrosted; and
(e) the date of minimum durability [Regulation (EU) No
1379/2013]. According to these regulations, this informa-
tion is mandatory regarding cephalopods should they be
sold live, fresh, chilled, frozen, dried, salted or in brine.
However, for canned, breaded or composite products,
using the scientific name is not mandatory. Instead, the
legal name, a customary name or a descriptive, one should
be used (European Commission 2014). In Spain, the com-
mercial name is regarded as the official name through-
out the nation (BOE 2019). In this regard, several species
names can be traded under the same ‘legal name’. For
instance, the name ‘pota’ can refer to species from any
of five different squid genera: Berryteuthis, Illex, Noto-
todarus, Ommastrephes or Todarodes. This can clearly
mislead consumers. To tackle this issue, Spanish authori-
ties have disclosed a plan to update the current national
regulations (Velasco et al. 2021) including a new list of
accepted commercial names for canned seafood where
each Spanish commercial name would be related to a sin-
gle scientific name included in the FAO ASFIS database
(CWP on Fishery Statistics Secretariat 2020).
Since correct labelling is important for protecting nature
as well as the health of, and access to fair pricing for con-
sumers, an effective labelling strategy must use the ‘one spe-
cies, one name’ rule (Golden and Warner 2014). Accord-
ingly, using the scientific name of the species in every single
food format including canned and breaded products, would
be the best approach to achieve these goals as it would
enhance the traceability of seafood, making it easier for gov-
ernments to improve food safety, protect endangered species,
enforce fisheries laws, and curb seafood fraud.
Indicating the country of origin on the label should also
be mandatory for all seafood formats. With D. gigas, as
previously indicated, Spanish companies buy large quan-
tities from other countries such as Peru and process them
afterwards in several factories based in Spain. Labelling the
Marine Biology (2024) 171:44
Fig. 7  Examples of processed seafood sold in Spain as octopus
(implying Spanish-caught Mediterranean common octopus, Octopus
vulgaris) but containing only jumbo squid (the low-value, low-quality
squid, Dosidicus gigas, of SE Pacific origin; see text). This labelling
is blatantly fraudulent, designed to mislead the consumer into believ-
ing that the contents are octopus, even though the named species is a
squid. a, b ‘Marinated octopus’. a The main package label bears the
common name ‘Pulpo’ (octopus) and then the scientific name, Dosid-
icus gigas accompanied by the Spanish flag and the words ‘Product of
Spain’. b In the small-font list of ingredients, the rear label states only
‘jumbo squid tentacles’ but also shows the word ‘Pulpo’ (octopus) in
Page 25 of 46 44
the largest font, and also capitalized, beneath which (in smaller font)
is the scientific name, Dosidicus gigas. c, d ‘Octopus, garlic style’. c
The front label reads ‘Octopus, garlic style’ and ‘Product of Spain’.
The rear label (d) lists the ingredients and begins with ‘Pulpo (Dosid-
icus gigas)’. e, f ‘Boiled tentacles’ (strongly implying that the pack
contains chopped arms of the octopus, O. vulgaris). e The front label
has an illustration of an octopus and states ‘boiled tentacles’, but the
rear label (f) states the content as ‘Dosidicus gigas’, while a recipe is
included for cooking the contents as ‘octopus Galician style’. (Image:
RGG. Commercial trademarks have been blanked out with grey rec-
tangles)
13
 44 Page 26 of 46
final canned product as a ‘product of Spain’ is legal accord-
ing to the current international regulations, but this is also
undoubtedly misleading for consumers and all concerned.
This is a significant loophole requiring rectification, at the
international level, such that the Codex should require label-
ling with the true geographic site of origin (country) of the
organism, in place of, or at least in addition to, the country
of processing. Without such legislative changes to labelling
requirements, it will continue to be difficult to trace the true
source, and affected resource, to inform true sustainability.
Attention to rigorous traceability has the potential to reduce
both the incidence and impact of mislabelling.
IUU fishing and a summary of measures in place to combat
it
Illegal, unreported and unregulated (IUU) fishing is a broad
expression covering various fishing activities that contra-
vene national or international law for fisheries management
(Agnew et al. 2009; Sumaila et al. 2020). Illegal fishing is
where any foreign or domestic vessel operates in waters
under the jurisdiction of a State without the permission of
that State, or in contravention of its laws and regulations
including those of any relevant regional fisheries manage-
ment organization (RFMO) to which that State is a party.
Unreported fishing is any activity resulting in failure to
honour obligations to report fishing activities accurately to
the relevant national authority or RFMO. Unregulated fish-
ing refers to the fishing of fish stocks to which there are no
applicable conservation or management measures, and also
to activities carried out in a RFMO area by vessels without
nationality, or by those flying the flag of a State not party to
that RFMO, in a manner that contravenes the conservation
and management measures of that State or RFMO.
IUU fishing remains one of the greatest threats to marine
ecosystems due to its potent ability to undermine efforts
to manage fisheries sustainably and to conserve marine
biodiversity. It is difficult to monitor, but IUU fishing has
been estimated to affect 26 Mt valued at US$11 to 23 bil-
lion annually (Agnew et al. 2009); and more recently, 8–14
Mt, with an estimated loss in annual economic impact (due
to the diversion of fish from the legitimate trade system)
at US$26 to 50 billion (Sumaila et al. 2020). IUU fishing
takes advantage of corrupt administrations and exploits
weak management regimes, in particular those of develop-
ing countries lacking the capacity and resources for effective
monitoring, control, and surveillance (MCS). It is ubiquitous
and insidious: found in all types and dimensions of fisheries;
on the high seas and in areas within national jurisdiction;
concerns all aspects and stages of the capture and utilisa-
tion of fish; and may be associated with organized crime. It
greatly disadvantages and discriminates against those fishers
who act responsibly in accordance with the terms of their
13
Marine Biology (2024) 171:44
fishing authorizations. Fisheries resources available to bona
fide fishers are removed by IUU fishing, which can lead to
the collapse of local fisheries, with small-scale fisheries in
developing countries proving particularly vulnerable. Prod-
ucts derived from IUU fishing can enter overseas trade mar-
kets which thereby undermines and threatens livelihoods,
exacerbates poverty, and augments food insecurity.
Octopus and squid commercial landings data, too, are
subject to issues of under- and mis-reporting and, in many
cases, the taxonomic identity of the targeted species remains
unknown or is taken for granted because official fishery sta-
tistics do not attempt to distinguish different species. How-
ever, the stigma of IUU fishing has its effects, too, making
it difficult to find IUU estimates for cephalopods: Ospina-
Alvarez et al. (2022) clearly avoided the issue when they
stated, ‘our research does not include illegal, unreported,
and unregulated (IUU) trade and discards data’.
In the USA, estimates suggest that 20–32% by weight of
all imported seafood originates from IUU sources. These
illegal imports were valued in 2011 at between $1.3 and
$2.1 billion, out of a total of $16.5 billion for the 2.3 Mt of
edible seafood imports. This trade represented between 4
and 16% of the value of the global IUU fish catch and reveals
the unintentional role of the USA, one of the largest seafood
markets in the world, in funding the profits of illegal fishing
(Pramod et al. 2014).
Combating IUU fishing has been the focus of govern-
ments, RFMOs, and intergovernmental and non-governmen-
tal organizations worldwide. This has led to the develop-
ment of an international framework to support three types
of measures to combat IUU fishing (summarized in Table 4).
Since 2015, national and international efforts to combat
IUU fishing have been driven by targets SDG 14.4 and 14.6
of the UN 2030 Agenda for Sustainable Development, which
aim to (respectively) end IUU fishing and eliminate subsidies
that contribute to IUU fishing by 2030. Of all the measures
being implemented, the most relevant to IUU fishing is the
CDS (measure outlined in Table 4, Sect. 3A), which is a sys-
tem to determine throughout the supply chain whether fish
originates from catches consistent with applicable national,
regional and international conservation and management
measures. CDS could function most effectively in synergy
with other international instruments including the PSMA and
the Global Record of Fishing Vessels, Refrigerated Transport
Vessels and Supply Vessels. Trade policy can play a critical
part in combating IUU fishing by implementing market states
measures (Table 4, Sect. 3A, B). TDS documents enable the
ID of exports of seafood sourced from fishing vessels flying
the flags of RFMO non-member states, who are barred from
harvesting species covered by the RFMO.
Both the US and the EU have procedures for trade-
restrictive measures to be taken against countries perceived
as doing too little to address IUU fishing. The EU uses Trade
Marine Biology (2024) 171:44 Page 27 of 46 44

Table 4  International regulatory framework for sustainable fisheries management and combating IUU. Measures deployed following interna-
tional agreements according to their functional relevance to fishing activities for different countries as port states, flag states or market states

(1) Port State Measures Agreement (PSMA, in force since 2016)

Aim: to prevent, deter and eliminate IUU fishing; prevent marketing of fish caught through IUU fishing by prohibiting port entry to foreign
vessels engaged in, or suspected to be engaged in, IUU fishing
(2) Voluntary Guidelines for Flag State Performance (VGFSP, in force since 2014):
Aim: to strengthen and monitor flag state compliance with obligations to ensure that vessels registered under their flags do not conduct IUU
fishing
Measures include monitoring, control and surveillance (MCS) activities such as vessel monitoring systems (VMS) and observers, which aim
to promote information exchange and cooperation among countries so that flag states refuse to register vessels that are ‘flag-hopping’ by
attempting to register with another flag state and refuse to allow the landing of vessels that have been reported for IUU fishing. Included in
VGFSP are recommendations on how countries can encourage compliance and take action against non-compliance by vessels, as well as on
how to enhance international cooperation to assist developing countries to fulfil their flag state responsibilities
(3A) Voluntary Guidelines for Catch Documentation Schemes (VGCDS, adopted by FAO in 2017). Aim: to provide assistance to States,
RFMOs, and other organisations when developing and implementing new CDS, or harmonising or reviewing existing CDS. (CDSs and TDSs
were developed in combating IUU fishing of tuna and have been implemented with success since the early 1990s)
(3B) Trade Restrictive Measures (TREMs). Along with CDSs and trade documentation schemes (TDSs), TREMs are key trade-related measures
to combat IUU fishing

Restrictive Measures (TREMs) in the form of yellow cards
(identification of non-cooperating countries) and red cards
(ban on imports). States can only become the object of EU
trade measures in their capacity as flag states; port or mar-
ket states that actively participate in the laundering of IUU
products cannot be targeted. These trade restrictions are
applied broadly to all fish imports from a particular country
regardless of the IUU fishing that triggered the identifica-
tion. While industrial operators generally have the option
of reflagging their vessels to avoid flag state-related restric-
tions, small-scale fisheries are inherently unable to escape
embargoes on their flag state. This means that embargoes
on an entire country’s fish exports are more likely to have
disproportionate impacts on small-scale fisheries.
In the USA, NOAA oversees fisheries management.
NOAA’s Office of Law Enforcement (NOAA OLE) enforces
regulations pertaining to the conservation and protection of
US living marine resources and natural habitats, of which
the Magnuson-Stevens Fishery Conservation and Manage-
ment Act and the Lacey Act Amendments of 1981 are of
high relevance to combating IUU fishing. NOAA releases a
biennial report to Congress outlining the agency’s actions to
improve international monitoring and enforcement of fisher-
ies policy. It also provides to Congress a list of nations iden-
tified as engaging in IUU fishing practices, illegal by-catch,
and other prohibited activities. Although NOAA OLE has
been tracking and identifying countries involved in IUU fish-
ing since 2009, these identifications do not lead to a ‘nega-
tive certification’, the equivalent of an EU red card. Instead,
the US legislation allows trade measures to target only fleets,
species, and product types directly tied to the IUU fishing
that has given rise to the identification.
In 2014, US President Obama issued an Executive Order
calling for the creation of The Presidential Task Force for
Combating IUU Fishing and Seafood Fraud. In March 2015,
a multi-agency committee released an action plan for com-
bating IUU fishing and seafood fraud within the US market
and containing recommendations ranging from expanding
collaboration with other nations on the issue of IUU fish-
ing, developing better technology for tracking imports, and
ratifying the UN’s Port State Measures Agreement. As a
result, NOAA Fisheries developed a national seafood trace-
ability program (known as the Seafood Import Monitoring
Program, SIMP) which applies to selected at-risk seafood
species entering US commerce. SIMP includes mandatory
collection of data from importers pertaining to the harvest,
landing, and chain of custody of products via an electronic
portal and verified via random audits. The SIMP homepage
links to a range of resources to aid industry compliance
with the new regulation, including model catch certificates,
a compliance guide, and an implementation guide. A volun-
tary ‘trusted trader’ program is also being developed which
seeks to reduce the compliance costs for eligible importers
who purchase from trustworthy supply chains with robust
traceability processes already in place.
Many other countries have ratified the PSMA and devel-
oped national rules and market access requirements to com-
bat IUU fishing. However, many countries are experiencing
challenges with implementation. Whether enacted by port,
flag or market state, any measure to combat IUU fishing
should be reliable and transparent, particularly in interna-
tional seafood trade. This requires the use of reliable and
cost-effective systems and tools to support enforcement of
market access rules that do not create unnecessary technical
barriers to trade.

13
 44 Page 28 of 46
Cephalopod traceability
Numerous confounding variables render traditional marine
population management approaches incapable of monitor-
ing cephalopod bioresources efficiently, leaving loopholes
that result in accidental mislabelling or intentional exploi-
tation which compromise sustainability. For importers and
processors in particular, the complications are significant:
Mexico, for example, represents a bare 7.8% of US ceph-
alopod imports; but Spain, which supplies half of these
imports, is itself a major importer of octopus harvested in
Mexico, Morocco, Mauritania and Argentina, which it then
re-exports to the US market (Coronado et al. 2020; CWP
2020). Seafood products of different origins are often also
mixed during processing and export.
To accurately identify cephalopod seafood products
requires a reliable, affordable, and widely accessible sys-
tem capable of tracking seafood from its point of harvest
through its distribution. Traceability technology this effec-
tive would, in turn, underwrite improved stock manage-
ment and resource sustainability, while also allowing eco-
labelling and certification to enhance product value through
premium pricing.
Ensuring that seafood supply chains are fully traceable,
and that products are legal and accurately labelled is a large
undertaking, but it is a challenge that must be met head-
on if companies are to achieve their sustainability goals.
As government regulations and international media focus
more on food safety, eliminating IUU fishing, and protect-
ing human rights in the seafood industry, it will be impor-
tant that companies are also proactive on these topics. By
better understanding and managing their supply chains and
developing the ability to trace specific products back to their
source, companies can mitigate or eliminate high risk items
while maximizing inventory control. Investing resources
into solutions now will help to protect brand value, build
consumer trust, and greatly reduce these environmental and
social issues.
In the last decade, much promising work has been done to
tackle seafood traceability issues. There are many resources,
from traceability software solutions, auditors and consult-
ants to academia, government institutions and NGOs that
can provide assistance to businesses looking to improve their
supply-chain traceability (FishWise 2018). There are also
steps companies can take now to begin to improve supply
chain traceability, including clearly communicating expecta-
tions to their suppliers, improving internal tracking systems,
conducting risk assessments and auditing high-risk items.
13
Marine Biology (2024) 171:44
Government‑instigated approaches
An example of a government-based approach at providing
the impetus for improved traceability, with significance for
cephalopod traceability, is Morocco where quota control has
been established for more than 2 decades and the authori-
ties enforce a biological rest along the Moroccan coast to
fight against illegal fishing. Owners and operators of fishing
establishments and premises are required by law (established
by the Moroccan Ministry of Agriculture, Maritime Fish-
eries, Rural Development and Water and Forests; MAPM-
DREF) to keep records of the origins of products, as well
as the quantities received, processed, shipped and stored to
contribute to eradicating illegal fishing and informal circuits
and to safeguard the resource. In practice, this is also called a
‘catch certification’, a tracking system documenting the fish
from its capture to the consumer. This certification system
has contributed to better monitoring of fishing and post-
harvest activities through the traceability of products from
the ship to export, passing through the intermediate opera-
tions of product transformation.
DNA barcoding for species identification
DNA barcoding has become established as a powerful tech-
nique to support policing of international legal frameworks
and/or local regulations and codes of conduct, both for the
protection of endangered species (Hobbs et al. 2019) and the
management of exploited populations (Lu and Luo 2020).
Barcoding has been particularly successful, and is now
widely applied in testing the accuracy of labelling of species
in seafood products (Luque and Donlan 2019), contributing
to the fight against IUU fishing (Agnew et al. 2009).
DNA-based approaches are particularly useful where tra-
ditional methods of ID are difficult or compromised, such
as where clear species delimitation is not established or
cryptic species exist, or when material exists as processed
products and distinctive morphological characters are lost.
In cephalopods, both factors are an issue. Although many
species occupy wide geographical ranges with little over-
lap among similar species from the same genus or family,
facilitating easier region-by-taxon species ID, there are also
many examples of incomplete taxonomic information com-
plicated by the presence of cryptic species (e.g., Avendaňo
et al. 2020; Fernández-Álvarez et al. 2020), as exemplified
by the recently much-improved elucidation of the global
distribution of ‘common octopus’ species comprising the
‘Octopus vulgaris complex’ (see Sect. “Identification”).
A more immediate practical issue is that cephalopods
display highly flexible morphologies with few hard body
parts on which to base clear species ID, and so quick and
reliable ID is difficult for all but experienced taxonomists,
Marine Biology (2024) 171:44
and particularly where individuals are processed into sea-
food products. Due to improved DNA sequence coverage
tied to taxonomic and geographical range descriptions for
many cephalopod species, and improved coverage of spe-
cies in more reliable DNA sequence databases such as the
Barcode of Life Data System (Hebert et al. 2003; BOLD
2022), routine confirmation of species ID and therefore
general geographical origin is available through standard
DNA barcoding methods using Polymerase Chain Reaction
(PCR)-based sequencing protocols. Such methods are now
being used to confirm the identity of cephalopod products in
the supply chain, at least to species level (Debenedetti et al.
2014; Guardone et al. 2017; Maggioni et al. 2020). However,
this system works well only as long as the reference database
DNA sequences themselves have been identified correctly
(see Sect. “Identification”) and if the reference specimens
were not obtained at a distance unreasonably far from the
type locality for the species (Vecchione et al. 2022).
Full traceability in the seafood supply chain, however,
will only be achieved if products can be traced to a particular
area inhabited by a given wild population or fished stock.
Genetic methods have shown potential to extend traceability
to the population level, as long as suitable genetic differen-
tiation between stocks exists and suitably powerful marker
systems are applicable. Recent Next-Generation Sequenc-
ing (NGS) approaches have allowed the development and
application of genome-wide genetic marker methods such
as the use of single-nucleotide polymorphisms (SNPs),
which show much greater power than previous methods to
distinguish within-species population structures (Bernatchez
et al. 2017; Barth et al. 2019). RADseq-based SNP screens
are now proven technology in enhancing the resolution of
fishery management boundaries (Benestan et al. 2015; Mul-
lins et al. 2018; McKeown et al. 2020), and have shown the
potential for traceability in abundant and widespread pelagic
finfish species (Bekkevold et al. 2015; Barth et al. 2019)
with life history and biogeography characteristics similar to
those displayed by cephalopods.
Traditional methods of population estimation are often
unsuitable for cephalopods (Young et al. 2006), because
species are not amenable to physical tagging methods
(Arkhipkin 2005) and have pronounced phenotypic plastic-
ity (van der Vyver et al. 2016), but genetic methods have
revealed sub-structuring within a number of fished species
(Shaw et al. 1999; McKeown et al. 2019). SNP analyses have
recently been applied to cephalopods (Morse et al. 2018;
Timm et al. 2020), so the potential for identifying geographi-
cal stock-of-origin is being realized.
It would, however, be naïve to think that any one method
could resolve the complex issues involved in identifying and
tracing cephalopod products through the supply chain back
to their fishery location of origin. No one method provides
answers in all situations, so an operational framework, as
Page 29 of 46 44
suggested by Cusa et al. (2022), should be applied to decide
on the most appropriate tools to apply. For example, stable
isotope profiling of seafood products and areas of origin has
shown potential for some fish species in some areas (Carrera
and Gallardo 2017; Kim et al. 2015a), and stable isotope
and multi-elemental analysis has recently been applied to
cephalopods (Martino et al. 2022).
eDNA metabarcoding for rapid species
identification
Environmental DNA (eDNA) is free DNA (or extra-organis-
mal DNA) that persists in the environment providing an indi-
cation of which organisms are currently, or were recently,
in the vicinity (Pilliod et al. 2013; Ames et al. 2021). All
living organisms shed genetic material as skin, scales, hair,
metabolic waste, etc., into their surrounding terrestrial or
aquatic environment and, much like traditional DNA barcod-
ing methods, this genetic material can be used to identify a
particular species. A principal advantage of eDNA metabar-
coding over the traditional DNA barcoding is that, by sam-
pling from the environment, an investigator no longer needs
to capture or even observe the targeted species (Bohmann
et al. 2014; Kelly et al. 2014; Deiner et al. 2016). The utility
of eDNA as a ‘forensics’ tool comes in part from its capac-
ity to utilize either passively sampled genetic material on a
filter (Bessey et al. 2021) or through active sampling using
handheld or battery-operated filtration pumps (Kelly et al.
2014). Furthermore, with the advent of more accessible and
lower-cost molecular and NGS technologies (Willette et al.
2014), eDNA metabarcoding also permits the ID of multiple
species present simultaneously from a single environmental
sample (Deiner et al. 2017). As with traditional DNA meth-
ods, eDNA metabarcoding benefits from the comprehensive
and dynamic open-access databases of genetic information
such as the Tree of Life and Barcode of Life projects (Hebert
et al. 2003; Janzen et al. 2005).
eDNA methods have garnered increasing attention as a
tool for detecting and monitoring biodiversity. Although
eDNA metabarcoding methods are used in a diverse range
of settings including soil (Hassan et al. 2022), air (Johnson
et al. 2021), ancient sedimentary rocks (Marchesini et al.
2022), and in screening for human and animal pathogens
(Farrell et al. 2021), these techniques have their origin in,
and are most extensively vetted and implemented in, aquatic
(Ficetola et al. 2008; Minamoto et al. 2012; Takahara et al.
2012) and marine (Rourke et al. 2021) environments. For
example, early trials of eDNA metabarcoding methods were
conducted in large artificial mesocosms (Kelly et al. 2014)
and yielded high levels of species ID accuracy (Miya et al.
2015). eDNA methods have also been shown to be effective
at detecting the presence of rare and low-abundance species
13
 44 Page 30 of 46
that may be overlooked using the traditional monitoring
techniques (Thomsen et al. 2014, Pikitch 2018).
The implementation of eDNA metabarcoding methods has
tremendous potential to aid in the re-envisioning and mod-
ernization of marine fisheries and the seafood supply chain.
Traditional DNA barcoding has frequently been used at the
end of the supply chain in restaurants, markets, and grocers,
where it has been effective in exposing seafood fraud including
species substitution (Naaum et al. 2016; Willette et al. 2017,
2021a; Luque and Donlan 2019). However, traditional DNA
barcoding is limited because of the need to individually sam-
ple each specimen, which is not practical early in the supply
chain where thousands of fish may be transported and pro-
cessed simultaneously. eDNA metabarcoding has the promise
to overcome this challenge (Willette et al. 2021b). Beyond spe-
cies ID en masse, applications of eDNA metabarcoding can
aid in estimating fish population size and biomass assessments
(Andres et al. 2021; Rourke et al. 2021; Stoeckle et al. 2021),
detecting threatened species in fisheries by-catch (Burgess
et al. 2018), cataloguing gut content in diet analyses (Leray
et al. 2013), and discovering instances of, and the impact of,
farm fish escapes (Aguirre-Pabón et al. 2015). After more
than a decade of robust development of eDNA metabarcod-
ing methods for monitoring biodiversity, and much more, the
timing is right for the use of this tool to shift from exploratory
trials into broader implementation (Bernatchez et al. 2017),
including routine marine assessments by fisheries agencies
(see Aylagas et al. 2018; Laroche et al. 2018). Importantly,
national and international policies are the next step in acceler-
ating this transition from novel to routine (Lodge 2022).
With regard to cephalopod fisheries, initial trials have
been performed in several countries including Japan, Mexico
and the USA, where importers and processors have shown
great interest in incorporating eDNA techniques into their
monitoring programmes, especially in view of the new SIMP
regulations for mandatory ID data collection and verifica-
tion (see Sect. “IUU fishing and a summary of measures in
place to combat it”). eDNA sampling is also one of the pro-
cedures used by the Seafood Tracer app (see Sect. “Trace-
ability apps”).
Whole genomes
Having the genome of different species available extends
the utility of both barcoding and eDNA metabarcoding for
combating illegal fishing and mislabelling by opening up
the possibilities of using other reference sequences such as
SNPs (Carvalho et al. 2010; Crawford and Oleksiak 2016;
Valenzuela-Quiñonez 2016; van Oppen and Coleman 2022).
As of December, 2022, the number of cephalopod
genomes available stood at 17, including two species
of squid and 13 species of octopus, for which whole-
genome sequencing and assembly have been completed.
13
Marine Biology (2024) 171:44
The Association for Cephalopod Research (‘CephRes’)
maintains a web page dedicated to cephalopod genomes
(CephRes 2022). Typically, DNA extraction is performed
using the Qiagen DNeasy genomic extraction kit following
the standard protocol for tissues. A paired-end sequencing
library is constructed using the Illumina TruSeq kit accord-
ing to the manufacturer’s instructions and sequenced on an
Illumina Hi-Seq platform in paired-end, 2 × 150 bp format.
The resulting fastq files are trimmed of adapter/primer
sequence and low-quality regions with Trimmomatic v0.33
(Bolger et al. 2014) and the trimmed sequence is assembled
by SPAdes v2.5 (Bankevich et al. 2012) followed by a finish-
ing step using Zanfona (Kieras et al. 2021).
Artificial intelligence, computer vision, machine
learning and deep learning
While genetic methods are improving the labelling of sea-
food, Artificial Intelligence (AI) methods can help expand
the reach and impact of genetic testing and accelerate the
rapid screening of seafood. All of the intelligence can be
accessible on a smartphone or in the Cloud. Here, AI refers
to computational systems able to perform tasks that nor-
mally require human intelligence but with increased effi-
ciency, precision, and objectivity. Computer vision is a field
of AI that deals with how computers and systems can be
used to acquire and process data from images, videos and
other digital data, and make decisions and recommendations
upon them. A subset of AI that plays a critical role in com-
puter vision is machine learning, ML, where mathemati-
cal models perform a specific task without using explicit
instructions, instead relying on patterns and inference. Deep
learning (DL) is a subset of ML that uses artificial neural
networks capable of learning from unstructured or newly
added data. The use of labelled training data can further
improve the AI predictive capability through supervised ML.
In recent years, ML and especially DL approaches are
used increasingly to automate marine species ID (Good-
win et al. 2022). DL approaches such as Artificial Neural
Network (ANN), Random Forest (RF), and Support Vector
Machine (SVM) are used increasingly to automate taxo-
nomic ID efforts (Kandimalla et al. 2022; Tan et al. 2021).
This development of ID aids was fueled by the explosion
of freely available visual data featuring living organisms
resulting from the widespread use of smart mobile devices,
digital (and analogue) cameras, as well as the digitization
of vast natural history collections. Combining the exponen-
tial growth of biological image data with cutting-edge ML
techniques such as DL opens up a world of possibilities for
automated species ID. Additionally, the widespread digitiza-
tion of natural history collections has become a top priority
for museums worldwide and has produced substantial digital
datasets. Attempts to classify fish have achieved high levels
Marine Biology (2024) 171:44
of performance and have also outperformed humans in spe-
cies recognition (Villon et al. 2018). Allken et al. (2019)
used the Inception v3 architecture to identify pelagic fish
species (blue whiting, Atlantic herring, and Atlantic mack-
erel) from trawl camera videos. Open-source AI software is
also becoming increasingly available to annotate and classify
marine species (e.g., VIAME from Kitware: https://​github.​
com/​VIAME/​VIAME).
However, only very limited ML techniques have been
applied to cephalopod classification problems. Orenstain
et al. (2016) used whole-body laboratory images of cuttle-
fish for classification of camouflaging behaviour with SVM
classifiers. Beak and statolith images were used in the ID of
three squid species including Uroteuthis duvaucelii, Loliolus
beka, and U. edulis found in the South China Sea (Jin et al.
2017). The method used to identify squid species was based
on outline geometric morphometric and stepwise discrimi-
nant analysis to evaluate the most suitable method for the
ID. The authors suggested that when the entire body of a
squid is available, a combination of statolith and beak should
be used for the ID. When only one hard tissue is available,
species ID can be subject to large errors. Another attempt to
identify squid was by Himabindu et al. (2017), who applied
an Artificial Neural Network (ANN) classifier to identify
5 squid species based on morphometric features measured
from whole-body samples.
Hu et al. (2020) developed and evaluated a rapid, low-
cost system to classify squid in industrial production. This
involved designing an easy-to-use handheld image-acquisi-
tion system combined with an automated, labour-saving, and
efficient DL model (named ‘Improved Faster Recurrent Con-
volutional Neural Network’) to identify three squid species
from the North Pacific Ocean. Three indicators (accuracy,
intersection-over-union, and average running time) were
used to evaluate the classification, and the results for the
three indicators in the test samples were means of 85.7%,
80.1%, and 0.144 s, respectively. These examples showed
the usefulness of ML methods for ID of cephalopods but
their application was limited to a single taxon group (mostly
squid). However, Tan et al. (2021) went a little further and
developed an ML ID model that can identify squid, cuttle-
fish and octopus based on beak images. The authors found
that the ANN model achieved the best testing accuracy of
91.14%, using the deep features extracted from the VGG19
model from lower beak images. The results indicated that
the deep features were more accurate than the traditional
features in highlighting morphometric differences from the
beak images of cephalopod species. In addition, the use of
lower beaks of cephalopod species provided better results
compared to the upper beaks, suggesting that the lower
beaks possess more significant morphological differences
between the studied cephalopod species (as concluded also
from morphological studies; Clarke 1986).
Page 31 of 46 44
Traceability apps
There is growing consumer awareness about the need for
seafood traceability. Sustainability commitments (such as
eco-labelling), fishery improvement projects (FIPs), con-
cerns about fraud, food safety, IUU fishing and human rights
abuses all motivate the use of traceability within the global
seafood sector (Leal et al. 2015; Lewis and Boyle 2017).
Numerous mobile applications (apps) have been devel-
oped focussed on seafood, especially in recent years. At
least 84 apps are currently available for wild-caught fisher-
ies worldwide, with different functionalities such as data
collection, information exchange among fishers, informa-
tion regarding value chains and post-harvest practices, and
for uses linked to employment, legislation and safety (Cal-
derwood 2022). Not many of these apps have traceability
functions, and even fewer have been used for cephalopods.
Still, the effective use of traceability is increasing, although
applied for different purposes in different parts of the supply
chain: from collecting and storing data, selling the catch at
the harvest level, and tracking transformation in the middle
of the supply chain, to informing consumers and to support
product quality and integrity claims at the end of the supply
chain (Lewis and Boyle 2017). Table 5 provides an overview
of tools used to improve seafood traceability which have
been used for cephalopods.
Abalobi is a suite of mobile apps, co-designed by South
African fishers and scientists with the support of telecom
companies to be an information management system for
the small-scale fisheries sector (Nthane et al. 2020). It col-
lects fisheries information and connects small-scale fish-
ers to market, facilitating the movement of fish directly to
consumers, and is backed by full supply-chain traceability.
Abalobi is focussed on a cell-phone-based modular plat-
form to facilitate the collection of information and finan-
cial transactions. The platform comprises five user-friendly
inter-connected apps tailored to the different users (Aba-
lobi-Fisher, -Monitor, -Manager, -Co-op and -Marketplace)
and an information management system (Nthane et al.
2020). For instance, ‘Abalobi Marketplace’ consists of a
digital market of fishery resources with an ecological and
social ‘story’ (https://​fishw​ithas​tory.​org) that connects buy-
ers looking for good-quality fresh, local and fully traceable
fish within a community-supported fisheries framework.
The app results in a fair, transparent and fully traceable
supply chain. Such a system is therefore able to contribute
to combating IUU, as the product is traced directly from
fisher to fork.
PescaData was developed in Mexico and is designed to
register the fishing effort of small-scale fisheries as well
as to connect fishing organisations and co-create solutions
for common problems faced by the sector. One function is
to shorten the value chain, linking fishers and consumers
13
 44 Page 32 of 46 Marine Biology (2024) 171:44

Table 5  Summary of Tools Abalobi TrazApp PescaData

characteristics of some
electronic systems with a Active since 2017 2018 2021
traceability component that are
Country of origin South Africa Peru Mexico (replicated in
in use within the cephalopod
(replicated in other other countries)
fisheries trade
countries)
Operations Geography Global National International
Language(s) Several Spanish Spanish
availability
Focussed on small-scale fisheries ✓ ✓ ✓
Supply chain actors
Producers (fishers, vessel owners) ✓ ✓ ✓
Intermediaries (middlemen, ✓ ✓ ✗
processors, wholesalers)
Fishing landing sites ✗ ✓ ✗
Consumers ✓ ✗ ✓
Functionalities
Electronic logbook* ✓ ✓ ✓
GPS ✓ ✓ ✓
Monitoring ✓ ✓ ✗
Marketplace ✓ ✗ ✓
Cephalopod species Loligo reynaudii Dosidicus gigas Dosidicus gigas
Octopus vulgaris Octopus mimus Octopus maya
Loligo gahi Octopus americanus
Compatibility with SALT standards** Yes Yes Yes
Webpage http://​abalo​bi.​org https://​traza​pp.​org https://​pesca​data.​org/

*Electronic logbook includes: records of daily catches, recording fishing effort, target species of fishing,
expenses invested in trips, capture sites, among others; **The Seafood Alliance for Legality & Traceability
(SALT) (https://​www.​saltt​racea​bility.​org/) sets principles for a comprehensive electronic Catch Documen-
tation and Traceability (eCDT) program. Note that currently the GDST global framework does not include
information on seafood traceability compliance, e.g., with regard to traceability apps (GDST 2023). How-
ever, the TrazApp web site states that the app is aligned with GDST standards (https://​www.​traza​pp.​org/#​
que; see also https://​trace​abili​ty-​dialo​gue.​org/​gdst-​capab​le-​solut​ions/)

directly. It is currently being used by around 2,000 sub-
scribers from 12 Latin American and Caribbean countries
(Villaseñor-Derbez et al. 2022). Fishers are able to identify
their catch through a species guide containing a library of
719 different commercial marine species.
TrazApp is a Peruvian electronic catch documentation
and traceability (eCDT) system developed by the World
Wildlife Foundation (WWF)-Peru with the participation of
artisanal fisheries supply-chain personnel, including fish-
ers, intermediaries, processing plants and government enti-
ties. It was designed to be user-friendly, so that each person
in the supply chain can easily generate and access reliable
real-time fishing information. Similar to Abalobi, TrazApp
comes with apps for different users: -Patrón (skipper), -Com-
erciante (intermediaries), -Desembarques (landing sites),
and -Armador (vessel owner). TrazApp is already being used
in eight different regions across Peru and includes more than
300 vessels so far, sharing information about their landings.
It was created to address the current sustainability demand
from international markets, and is designed to guarantee the
legal origin of fishing resources, increase transparency and
public access to catch information, and improve compliance
with national legislation.
Seafood Tracer is an app that was developed using
octopus species as a test group with support from the US
National Science Foundation, collating information from
eDNA, computer vision AI, and species spatial distribution,
and aiming to deliver near real-time species identification to
seafood supply chain end-users. The 1-year project began
in October 2021 and rapidly developed a working proto-
type traceability tool that (a) used images and associated
metadata, genetic data, and ecological data to validate spe-
cies identity of seafood; and (b) initiated a collaborative,
international network of scientists and fisheries industry
stakeholders to begin field testing a portable eDNA mobile
lab and purpose-built eDNA collection device in the USA,
Mexico, and Japan. The Seafood Tracer project applied
the principles of convergence research (cross-disciplinary
integration of knowledge, methods, and expertise) and was
aimed at addressing the global challenges of food security

13
Marine Biology (2024) 171:44
by boosting the transparency of seafood trading and thereby
contributing to fisheries sustainability. Although the project
ended in 2022, further scientific research is benefitting from
spin-offs of the project including the portable eDNA mobile
lab and patented eDNA collection device (patent pending).
All these apps rely on visual ID of seafood but do not
have an alternative platform to ensure that seafood products
are objectively identified correctly to species level, which
was the emphasis of the specifications and design of Seafood
Tracer (which, in addition, was based initially on cephalopod
seafood).
Conclusions and recommendations
This comprehensive review sheds light on the crucial
aspects surrounding cephalopods as seafood, emphasizing
the pressing need to address prevailing gaps and challenges
in regulations, certifications, stock assessment and fisheries
management of this species group. Shortcomings in existing
management strategies, data collection and traceability sys-
tems are contributing to transparency issues and potentially
unsustainable fishing practices. Addressing these shortcom-
ings demands the establishment of more robust traceability
systems, underpinned by stringent labelling mandates and
resilient seafood procurement policies, especially at the
importer level. Legislation and government inspection sys-
tems are pivotal, as exemplified by initiatives such as the
recently implemented SIMP in the US and Japan’s Catch
Documentation Scheme (MAFF 2022), akin to the European
Community Catch Certification Scheme introduced in 2010.
New initiatives are on the horizon such as the proposed EU
Directive on corporate sustainability and due diligence
(EUR-Lex 2022). Effectively implementing these changes
will drive a shift in management strategies towards securing
the future of cephalopod (and other) fisheries. Maintaining
momentum in urging harvesting countries and supply-chain
players to invest in science-based solutions is vital to address
the identified gaps and challenges, ensuring improved over-
all sustainability of cephalopod natural resources and sea-
food-dependent economies.
Resolving problems of (mis)identification and (mis)label-
ling is fundamental. Accurately assigned scientific names
should always be included on the labels of every food form
along with the commercial name, the fishing country and
area of origin of the species and other mandatory informa-
tion. Molecular inspection of seafood should be routinely
used as support for official monitoring to ensure regulatory
compliance (Willette et al. 2018, 2021b). Also (as suggested
by Willette et al. 2017), both end buyers (e.g., foodservice,
retailers) and consumers should be educated to become
more familiar with seafood products (e.g., appearance,
Page 33 of 46 44
flavour) and their origin to identify potential fraud related
to mislabelling.
A major scientific problem for all seafood is the perva-
sive presence of DNA sequences from misidentified speci-
mens in public databases such as GenBank and BOLD
(Sects. “Identification” and “DNA barcoding for species
identification”; see also Strohm et al. 2016). The root of
this problem is the lack of taxonomic curation (rigorous
or otherwise) of these databases. Currently, although
there have been some attempts to rectify this, there are
still no clear procedures for confirming sequence IDs that
are implemented at database level. Some attempts have
been made to flag sequences considered reliable accord-
ing to agreed standards, such as proposals for a specific
keyword ‘Barcode’ to be assigned by database curators
which would provide a subset of reference sequences
for reliable barcoding (see, for example, Hanner 2012).
However, so far, such proposals have not been taken up
(Hanner pers com to IGG). Clearly, at present it cannot be
expected that taxonomic curation will happen at the level
of large databases such as GenBank, so it will be necessary
to establish secure taxonomically curated lists specifically
for use in traceability studies. Towards this end, regarding
cephalopods initially, metadata for taxonomically curated
sequences for octopus species of the Americas are now
available as an online spreadsheet (González-Gómez et al.
2023, this issue).
With the unprecedented growth of global seafood trade,
and notably the growing importance of cephalopods in
developing nations such as around the African continent
and the Indian Ocean, it is imperative to prioritize the sus-
tainability of cephalopod (and other fisheries) harvests. It
is hoped that the insights outlined here will contribute to
the development of additional strategies aimed at address-
ing critical issues for seafood in general and for cephalopod
resources in particular. Fostering collaborative effort among
governments, stakeholders, scientific communities and an
informed public can pave the way towards a more sustain-
able and responsible seafood sector.
Acknowledgements The authors are most grateful to Athena Allen, at
Nature Bureau Ltd. UK, for her able and efficient administrative assis-
tance, particularly with collating the references and citations. Gleadall
(IGG), Moustahfid (HM), Sauer (WHHS) and Willette (DAW) grate-
fully acknowledge the efforts and support of the team during work on
the Seafood Tracer project (NSF Grant #2137582 to DAW), particu-
larly Cheryl Ames (a Co-Principal Investigator on that project), Amina
Jackson, Margarita Joaquin, Julie Kim, and Kevin Wang. Pita (CP,
University of Aveiro, Portugal) acknowledges FCT/MCTES for the
financial support to CESAM (UIDP/50017/2020 + UIDB/50017/2020
+ LA/P/0094/2020) and research contract 2020.02510.CEECIND.
WHHS also acknowledges the support of the GCRF funded One Ocean
Hub project. The authors thank Chingis Nigmatullin for suggesting the
Yuri M. Froerman dedication and providing a brief biography.
13
 44 Page 34 of 46
Author contributions IGG, HM and WHHS conceived the review and
drew up the initial content headings. All authors made substantial con-
tributions to drafting the review and revised it critically for important
intellectual content and relevance; approved the version to be pub-
lished; and agreed to be accountable for all aspects of the work in
ensuring that questions related to the accuracy or integrity of any part
of the work are appropriately investigated and resolved.
Funding This work was supported in part by US National Science
Foundation (NSF) Grant 2137582 to Demian A Willette. Paul W. Shaw
received financial support from the Aberystwyth University Impact
Leave Fund; and Warwick HH Sauer support via the GCRF funded pro-
ject, One Ocean Hub. The other authors declare that no funds, grants,
or other supports were received during the preparation of this review.
Data availability There are no new data specific to this review article.
Declarations
Conflict of interest Ian G Gleadall is Executive Director of AiCeph
LLC, Consultant to Hotland plc, Tokyo (a fast-food franchise company
specializing in octopus products), and, along with Warwick HH Sauer,
was a Consultant and Co-Principal Investigator (Co-PI) on the Seafood
Tracer project funded by NSF Award 2137582. Demian A Willette
was Senior Co-PI on the Seafood Tracer project, Hassan Moustahfid
was a Co-PI, and Isa Elegbede and Unai Markaida were Consultants.
The remaining authors have no competing interests to declare that are
relevant to the content of this article.
Ethical approval There are no requirements for ethics approval regard-
ing this review article.
Consent to participate There are no requirements for consent to par-
ticipate.
Consent to publish There are no requirements for consent to publish.
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Ian G. Gleadall1 · Hassan Moustahfid2 · Warwick H. H. Sauer3 · Lahsen Ababouch4 ·
Alexander I. Arkhipkin5,21 · Jilali Bensbai6 · Isa Elegbede7 · Abdelmalek Faraj6 · Pedro Ferreiro‑Velasco8 ·
Roberto González‑Gómez9 · Carmen González‑Vallés8 · Unai Markaida10 · Piedad S. Morillo‑Velarde9 ·
Graham J. Pierce11 · Stacy Pirro12 · Cristina Pita13 · Katina Roumbedakis14 · Yasunori Sakurai15 ·
David Scheel16 · Paul W. Shaw17 · Pedro Veiga8,18 · Demian A. Willette19 · Andreas Winter5 ·
Tadanori Yamaguchi20
* Ian G. Gleadall
enoctopus@yahoo.co.uk
Hassan Moustahfid
hassan.moustahfid@noaa.gov
Warwick H. H. Sauer
w.sauer@ru.ac.za
Lahsen Ababouch
lahsen.ababouch.la@gmail.com
Alexander I. Arkhipkin
alexander.arkhipkin@mpi.govt.nz
Jilali Bensbai
bensbaijilali@gmail.com
Page 45 of 46 44
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Young IAG, Pierce GJ, Murphy J, Daly HI, Bailey N (2006) Applica-
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Publisher's Note Springer Nature remains neutral with regard to
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Springer Nature or its licensor (e.g. a society or other partner) holds
exclusive rights to this article under a publishing agreement with the
author(s) or other rightsholder(s); author self-archiving of the accepted
manuscript version of this article is solely governed by the terms of
such publishing agreement and applicable law.
Isa Elegbede
isaelegbede@gmail.com
Abdelmalek Faraj
faraj@inrh.ma
Pedro Ferreiro‑Velasco
pedro.ferreiro@sustainablefish.org
Roberto González‑Gómez
gonzalezgomezroberto@gmail.com
Carmen González‑Vallés
carmen.gvalles@sustainablefish.org
Unai Markaida
umarkaida@ecosur.mx
13
 44 Page 46 of 46 Marine Biology (2024) 171:44

6
Piedad S. Morillo‑Velarde National Institute of Fisheries Research (INRH), 2 Bd Sidi
piedad_smv@hotmail.com Abderrahmane, 20250 Casablanca, Morocco
7
Graham J. Pierce Department of Fisheries, Lagos State University, Ojo,
g.j.pierce@iim.csic.es Nigeria
8
Stacy Pirro Sustainable Fisheries Partnership Foundation, 4348 Waianae
info@iridiangenomes.org Ave. #692, Honolulu, HI 96816, USA
9
Cristina Pita Centro Oceanográfico de Murcia (COMU-IEO), CSIC, San
c.pita@ua.pt Pedro del Pinatar, Murcia, Spain
10
Katina Roumbedakis El Colegio de la Frontera Sur, CONACyT, 24500 Lerma,
katina.roumbedakis@gmail.com Campeche, Mexico
11
Yasunori Sakurai Instituto de Investigaciones Marinas (Consejo Superior
sakurai.yasu31@gmail.com de Investigaciones Científicas), Calle Eduardo Cabello 6,
36208 Vigo, Spain
David Scheel
12
dscheel@alaskapacific.edu Iridian Genomes, Iridian Genomes, 613 Quaint Acres Dr,
Silver Spring, MD 20904, USA
Paul W. Shaw
13
pws3@aber.ac.uk CESAM ‑ Centre for Environmental and Marine Studies,
Department of Environment and Planning, University
Pedro Veiga
of Aveiro, Campus de Santiago, 3810‑193 Aveiro, Portugal
pedro.veiga@sustainablefish.org
14
Equalsea Lab, Cross‑Research in Environmental
Demian A. Willette
Technologies, CRETUS, Department of Applied
demian.willette@lmu.edu
Economics, University of Santiago de Compostela,
Andreas Winter 15782 Santiago de Compostela, Spain
awinter@naturalresources.gov.fk 15
Hakodate Research Center for Fisheries and Oceans,
Tadanori Yamaguchi Hakodate, Hokkaido, Japan
tadanoriyama@riam.kyushu-u.ac.jp 16
Institute of Culture & Environment, Alaska Pacific
1 University, Anchorage, AK 99508, USA
AiCeph LLC, Sendai, Japan
17
2 Department of Life Sciences, Aberystwyth University,
Integrated Ocean Observing System (IOOS), National
Aberystwyth, Ceredigion, UK
Oceanic and Atmospheric Administration (NOAA),
18
U.S. Department of Commerce, Silver Spring, MD, USA Centre of Marine Sciences (CCMAR), University of Algarve,
3 Campus de Gambelas, FCT Ed 7, 8005‑139 Faro, Portugal
Rhodes University, Grahamstown, South Africa
19
4 Biology Department, Loyola Marymount University,
The Food and Agriculture Organization of the United
Los Angeles, CA, USA
Nations, Rome, Italy
20
5 Kyushu University, Kasuga, Japan
Falkland Islands Government, Natural Resources – Fisheries,
21
Bypass Road, Stanley FIQQ 1ZZ, Falkland Islands Present Address: Fisheries New Zealand, 38 Bowen Street,
Wellington 6011, New Zealand

13