1
Marine and Fishery sciences 39 (2): xxx-xxx (2026)
https://doi.org/10.47193/mas.3922026010404
ABSTRACT. The effects of climate change on mackerel species are highlighted in this review,
with particular attention given to changes in spawning, distribution, physiology, and habitat loss. The
PRISMA method was used to conduct a systematic literature review that examined 257 peer-reviewed
publications from Scopus and Web of Science from 1990 to 2025. Results showed clustering of
texts, keywords, and titles based on search terms like climate change vulnerability, impact, mackerel,
southeast Asia, and world. High occurrence and link strength showed clusters of: 1) Oceanography
and marine ecosystem, 2) Species-specic regional studies, 3) Fisheries and human dimensions and,
4) Fisheries management. Major study themes were found using R bibliometrix and thematic anal-
yses based on VOSviewer: adaptation technique, marine ecosystems, stock assessment, and shing
management. Sea-surface temperature, environmental monitoring, and resilience in marine resource
management are highlighted in recent research trends. Traditional sheries are being challenged by
the altered migration and spawning patterns of mackerel due to rising sea surface temperatures and
changing currents, necessitating the implementation of exible and regionally coordinated techniques.
Growth, recruitment, and survival are all impacted by these changes because of decreased oxygen,
productivity, and prey availability. Conicts over sheries may also occur as a result of transboundary
sh stock migrations. Adaptive management is still limited despite increasing research because of a
lack of institutional support, resources, and monitoring. It is crucial to ll important research gaps
in areas including prey dynamics, socioeconomic resilience, and the effects of climate change on
mackerel biology. Stronger governance, regional collaboration, and policies that strike a balance be-
tween socioeconomic demands and conservation will be necessary to ensure long-term sustainability.
Key words: Systematic literature review, PRISMA method, sheries management.
El papel de la escolarización en la conguración de la huella pesquera en Grecia: evidencia
de un enfoque ARDL aumentado
RESUMEN. En esta revisión se destacan los efectos del cambio climático en las especies de caballa,
con especial atención a los cambios en el desove, la distribución, la siología y la pérdida de hábitat.
REVIEW
Bibliometric insight of climate change impact on mackerel (Family
Scombridae): global research trend and regional gaps in southeast Asia
MohaMMad ekraMul haque1, shahadat hossain2, tun nurul aiMi Mat JaaFar1, ying giat seah1, 3,
siti azizah Mohd nor4, nazia abdul kadar5, nur Fadli6, darlina Md naiM7, helena khatoon8 and ahasan habib1,*
1Faculty of Fisheries and Aquaculture Science, Universiti Malaysia Terengganu, Terengganu, Malaysia. 2Higher Institution Center of Excellence
(HICoE), Institute of Tropical Aquaculture and Fisheries, Universiti Malaysia Terengganu, Terengganu, Malaysia. 3Fish Division, South China
Sea Repository and Reference Centre, Institute of Oceanography and Environment, Universiti Malaysia Terengganu, Terengganu, Malaysia.
4Institute of Climate Adaptation and Marine Biotechnology (ICAMB), Universiti Malaysia Terengganu, Terengganu, Malaysia. 5Higher Institution
Centre of Excellence (HICoE), Borneo Marine Research Institute, Universiti Malaysia Sabah, 88400 - Kota Kinabalu, Sabah, Malaysia. 6Faculty
of Marine and Fisheries, Universitas Syiah Kuala, 23111 - Banda Aceh, Indonesia. 7School of Biological Sciences, Universiti Sains Malaysia,
11800 - Pulau Pinang, Malaysia. 8Chattogram Veterinary and Animal Sciences University, 4225 - Chattogram, Bangladesh.
ORCID Mohammad Ekramul Haque https://orcid.org/0000-0002-9124-0493, Shahadat Hossain https://orcid.org/0000-0002-8273-1135,
Tun Nurul Aimi Mat Jaafar https://orcid.org/0000-0002-3032-0545, Ying Giat Seah https://orcid.org/0000-0002-2976-4448,
Siti Azizah Mohd Nor https://orcid.org/0000-0002-9890-9612, Nazia Abdul Kadar https://orcid.org/0000-0003-4362-3059,
Nur Fadli https://orcid.org/0000-0001-6845-1383, Darlina Md Naim https://orcid.org/0000-0002-7943-9547,
Helena Khatoon https://orcid.org/0000-0001-5896-0213, Ahasan Habib https://orcid.org/0000-0003-1702-6111
Marine and
Fishery Sciences
MAFIS
*Correspondence:
a.habib@umt.edu.my
Received: 20 October 2025
Accepted: 3 January 2026
ISSN 2683-7595 (print)
ISSN 2683-7951 (online)
https://ojs.inidep.edu.ar
Journal of the Instituto Nacional de
Investigación y Desarrollo Pesquero
(INIDEP)
This work is licensed under a Creative
Commons Attribution-
NonCommercial-ShareAlike 4.0
International License
Marine and Fishery sciences 39 (2): xxx-xxx (2026)
2
INTRODUCTION
Climate change is reshaping marine ecosystems
through rising sea temperature, altered circulation
patterns, acidication, and the degradation of crit-
ical coastal habitats (Islam et al. 2020; Habib et al.
2025). These environmental changes inuence met-
abolic processes, growth patterns, and reproductive
success in many marine species, particularly pelag-
ic shes that depend on predictable oceanographic
conditions for spawning, feeding, and migration
(Pörtner et al. 2014). Ocean acidication further
disrupts physiological and sensory functions, po-
tentially altering predator-prey interactions and
competitive dynamics (Munday et al. 2009). In
the context of these changing conditions, adaptive
and ecosystem-based sheries management has be-
come essential, especially in regions where coastal
communities depend heavily on marine resources
for food security and livelihoods (Pinsky and Man-
tua 2014; FAO 2018).
Mackerel (Family Scombridae) are among the
most ecologically and economically signicant
pelagic shes in southeast Asia. They contribute
substantially to industrial, artisanal, and small-
scale sheries and remain an important source of
affordable animal protein throughout the region
(FAO 2018). In southeast Asia, Indian mackerel
(Rastrelliger kanagurta) and Spanish mackerel
(Scomberomorus commerson) dominate regional
captures due to their high abundance, diverse culi-
nary applications, and cultural signicance (Wang
and Wu 2025). In particular, Spanish mackerel
serves as a vital source of revenue sustaining their
livelihoods and meeting local protein demand of
coastal shermen in countries like Indonesia, the
Philippines, and Malaysia (Wang and Wu 2025).
In recent years, research on the relationship be-
tween climate change and sheries has moved
from correlative observations to process-based
and ensemble projections that measure economic
consequences, range shifts, and production losses
of pelagic shes, including mackerels (Free et al.
2019). With the projection of seasonal redistribu-
tions and changed catch potential under mid to late
century scenarios, species distribution and habitat
suitability modeling for scombrids has grown sig-
nicantly (Sun et al. 2024). Despite long-standing
concerns about overexploitation, recent regional
assessments suggest relatively stable or increasing
catches in several southeast Asian countries, with
a signicant proportion of pelagic stocks classi-
ed as undershed compared to global averages
(Fabro 2020).
Se utilizó el método PRISMA para realizar una revisión sistemática de la literatura que examinó 257 publicaciones revisadas por pares de
Scopus y Web of Science entre 1990 y 2025. Los resultados mostraron agrupamiento de textos, palabras clave y títulos con base en términos
de búsqueda como vulnerabilidad al cambio climático, impacto, caballa, sudeste asiático y mundo. La alta ocurrencia y la fuerza de enlace
mostraron agrupaciones de: 1) Oceanografía y ecosistema marino, 2) Estudios regionales especícos de especies, 3) Pesca y dimensiones
humanas y, 4) Gestión pesquera. Se encontraron los principales temas de estudio utilizando R bibliometrix y análisis temáticos basados
en VOSviewer: técnica de adaptación, ecosistemas marinos, evaluación de stock y gestión pesquera. La temperatura supercial del mar,
el monitoreo ambiental y la resiliencia en la gestión de los recursos marinos se destacan en las tendencias de investigación recientes. La
pesca tradicional se ve amenazada por la alteración de los patrones de migración y desove de la caballa debido al aumento de la tempera-
tura supercial del mar y las corrientes cambiantes, lo que exige la implementación de técnicas exibles y coordinadas a nivel regional. El
crecimiento, el reclutamiento y la supervivencia se ven afectados por estos cambios debido a la disminución del oxígeno, la productividad
y la disponibilidad de presas. También pueden surgir conictos pesqueros como resultado de las migraciones transfronterizas de poblaciones
de peces. La gestión adaptativa aún es limitada a pesar del aumento de la investigación debido a la falta de apoyo institucional, recursos y
monitoreo. Es crucial cubrir importantes lagunas de investigación en áreas como la dinámica de las presas, la resiliencia socioeconómica y
los efectos del cambio climático en la biología de la caballa. Una gobernanza más sólida, la colaboración regional y políticas que logren un
equilibrio entre las demandas socioeconómicas y la conservación serán necesarias para garantizar la sostenibilidad a largo plazo.
Palabras clave: Revisión sistemática de la literatura, método PRISMA, gestión pesquera.
Haque et al.: Climate CHange impaCt, global researCH trend, and regional gaps in soutHeast asia 3
Geographic location, economic reliance on sh-
eries, and great biodiversity make southeast Asia
(i.e. the Philippines) especially susceptible to these
effects (Macusi et al. 2020, 2025a, 2025b). De-
spite supporting some of the world’s most diverse
and productive ecosystems, this region is already
seeing a decline in sh populations, a shift in the
distribution of species, and an increase in the fre-
quency of extreme weather events (Cheung et al.
2010; Barange et al. 2014). Communities that de-
pend on shing are facing increasing uncertainty
as their livelihood are threatened by shifting spe-
cies ranges, decreased catch rates, and ecosystem
changes (Sumaila et al. 2011). Instead of combin-
ing population dynamics with climate forecasts at
management unit scales (EEZs and shared stocks),
the majority of regional studies are still short-term
or single gear and concentrate on establishing a
correlation between SST or ENSO and catch rates
(Ren et al. 2025; Vu and Nguyen 2025). In the
South China Sea and eastern Indian Ocean, there
are currently few quantitative projections speci-
cally for Indian mackerel (Rastrelliger kanagurta)
and Narrow-barred Spanish mackerel (S. commer-
son). Most existing studies rely on short-term cor-
relations between temperature indices and sheries
production, with limited integration of ecological
processes, population connectivity, or climate-driv-
en redistribution across national borders (Kamaru-
zzaman et al. 2021; Pan et al. 2024). Despite pol-
icy reports calling for climate-ready management,
there is a severe implementation gap because few
southeast Asian sheries have adopted harvest
control regulations, early warning indicators, or
scenario-based socioeconomic planning for small-
scale shing operations (Talbot et al. 2024).
This review critically assessed the breadth and
depth of southeast Asian contributions by contrast-
ing regional studies with the global literature and
synthesizing global research trends on the effects
of climate change on mackerels (Family Scombri-
dae) using a bibliometric approach. It highlights
important theme gaps, unexplored study areas, and
difculties in converting science into adaptive man-
agement by contrasting regional studies with the
worldwide literature. In addition to generating in-
sights for more general worldwide comparisons, the
analysis aimed to establish evidence of stronger cli-
mate-resilient sheries governance in southeast Asia.
MATERIALS AND METHODS
Search strategy
The Preferred Reporting Items for Systematic
Reviews and Meta-Analysis (PRISMA) for sys-
tematic literature review was used (Moher et al.
2015). The four stages of this process are identi-
cation, screening, eligibility assessment, and in-
clusion. The Web of Science and Scopus database
search phrases for the year 1990 through 2025 were
used to identify every article. In order to ensure
access to reliable sources and citation monitoring
to uncover important studies and trends, Web of
Science and Scopus were chosen for their depend-
able, high-quality control through rigorous peer-re-
viewed articles across elds like environmental sci-
ence, sheries, and climate change. Additionally,
these databases offer sophisticated search capabil-
ities for effective, methodical evaluation and analy-
sis, avoiding duplication from less reliable sources.
The following two strings were used for article
searching:
- Global: (‘Climate change’ OR ‘Climate im-
pact’ OR ‘Climate vulnerability’ OR ‘Climate
variability’ OR ‘Global warming’ OR ‘Ocean
warming’ OR ‘Environmental change’ OR
‘ENSO’ OR ‘El Niño’ OR ‘La Niña’ OR ‘Ex-
treme events’ OR ‘Marine heatwave’ OR ‘Ocean
acidication’) AND (‘Mackerel’ OR ‘Tuna’ OR
‘Bonito’ OR ‘Scombridae’ OR ‘Scomber OR
‘Rastrelliger OR ‘Scomberomorus’ OR ‘Kat-
suwonus’ OR ‘Thunnus’) AND (‘Fisheries’ OR
‘Fishery management’ OR ‘Stock assessment’
OR ‘Marine resources’ OR ‘Distribution shift’
Marine and Fishery sciences 39 (2): xxx-xxx (2026)
4
Screening and duplicate removal
A total of 1,337 records were retrieved. Using
automated duplicate detection in R (Bibliometrix)
and manual verication, 680 duplicated records
were removed. Titles, abstracts, and keywords of
the remaining articles were screened for relevance.
The eligibility assessment excluded articles unrelat-
ed to climate-change impacts on mackerel. A total
of 257 articles met all criteria and were included
for full bibliometric analysis. Extracted informa-
tion included title, abstract, author keywords, index
keywords, year of publication, journal, authorship,
afliations, citations, and document type.
Preprocessing and cleaning
Preprocessing and data cleaning followed
standardized bibliometric procedures to ensure
reproducibility. Keyword normalization includ-
ed converting all terms to lowercase, removing
punctuation, stemming plural forms, and merging
synonymous expressions such as ‘climate variabil-
ity’ and ‘climate-variability’ or ‘distribution shift’
and ‘range shift’. Generic stop-words (e.g. ‘model,’
‘study,’ ‘impact,’ ‘analysis’) and non-informative
geographic abbreviations were removed to avoid
noise in clustering. Author keywords referring to
the same species or taxonomic group were har-
monized by consolidating variations (for example,
Scomber japonicus and S. japonicus were unied
as ‘Scomber japonicus’).
VOSviewer parameters
Co-occurrence network maps for keywords, au-
thors, and countries were generated using standard-
ized parameters (Van Eck and Waltman 2010). Full
counting was applied as the counting method, and
association strength was used for normalization.
Thresholds were set at a minimum of ve occur-
rences for keywords, two for authors, and three for
countries. Network clustering followed a resolution
value of 1.00, and layouts were produced using the
OR ‘Range shift’ OR ‘Habitat suitability’ OR
‘Species redistribution’ OR ‘Larval dispersal’ OR
‘Recruitment’).
- Southeast Asia: (‘Climate change’ OR ‘Cli-
mate impact’ OR ‘Climate vulnerability’ OR
‘Climate variability’ OR ‘Global warming’ OR
‘Ocean warming’ OR ‘Environmental change’
OR ‘ENSO’ OR ‘El Niño’ OR ‘La Niña’ OR
‘Extreme events’ OR ‘Marine heatwave*’ OR
‘Ocean acidication’) AND (‘Mackerel’ OR
‘Tuna’ OR ‘Bonito’ OR ‘Scombridae’ OR
‘Scomber OR ‘Rastrelliger OR ‘Scombero-
morus’ OR ‘Katsuwonus’ OR ‘Thunnus’) AND
(‘Fisheries’ OR ‘Fishery management’ OR ‘Stock
assessment’ OR ‘Marine resources’ OR ‘Distri-
bution shift’ OR ‘Range shift’ OR ‘Habitat suit-
ability’ OR ‘Species redistribution’ OR ‘Larval
dispersal’ OR ‘Recruitment’) AND (‘Southeast
Asia’ OR ‘South China Sea’ OR ‘Indian Ocean’
OR ‘Eastern Indian Ocean’ OR ‘Coral Triangle’
OR ‘Indonesia’ OR ‘Philippines’ OR ‘Malaysia’
OR ‘Thailand’ OR ‘Vietnam’ OR ‘Brunei’ OR
‘Singapore’ OR ‘Cambodia’ OR ‘Myanmar’).
Inclusion and exclusion criteria
Studies were included if they were written in
English, focused on climate-change related impacts
on mackerel or related Scombridae taxa, provided
primary research data relevant to sheries, popu-
lation dynamics, or distributional change, and con-
tained analyzable metadata such as titles, abstracts,
and keywords. Review papers, book chapters, con-
ference summaries, or theses were excluded be-
cause this bibliometric analysis aimed to examine
trends in primary research rather than secondary
syntheses including review articles would dispro-
portionately inuence co-occurrence networks by
introducing broad synthetic keywords, obscuring
patterns emerging from empirical studies. Publi-
cations were also excluded if they were not related
to climate change or mackerel (Scombridae), or if
they were non-original research outputs such as
policy briefs or news articles.
Haque et al.: Climate CHange impaCt, global researCH trend, and regional gaps in soutHeast asia 5
LinLog and Modularity algorithms. To improve in-
terpretability, edges with a total link strength below
2 were removed from the nal visualizations.
Bibliometrix parameters (R)
Bibliometric analyses were conducted using
standardized Bibliometrix (R package version
4.3.1) (Aria and Cuccurullo 2017). The ‘biblio-
clean ()’ function was applied to remove empty or
inconsistent elds, and ‘convert2df ()’ was used
to merge ‘Web of Science’ and ‘Scopus’ exports
into a unied dataset. Descriptive indicators were
generated with ‘biblioAnalysis ()’, while themat-
ic structure and evolution were examined using
‘thematicMap ()’. Network visualizations for
keywords, countries, and co-authorship patterns
were produced with ‘networkPlot ()’. Association
strength was used for normalization, and minimum
frequency thresholds were set at three occurrences
for terms in titles and abstracts and ve occurrences
for keywords.
Thematic and qualitative analysis
Keyword co-occurrence clusters were interpret-
ed to identify dominant themes. Titles and abstracts
of all included studies (n = 257) were reviewed to
produce thematic summaries consistent with bib-
liometric clusters. Syntheses were based solely on
the patterns derived from bibliometric results.
RESULTS AND DISCUSSION
Global research trend on climate change and
mackerel (Family Scombridae)
Global research on climate change and mackerel
is rich and multidisciplinary, integrating species
biology, oceanography, ecosystem studies, and
shery management. However, regional gaps, es-
pecially in areas like southeast Asia, highlight the
need for more localized, policy-relevant research
to address region-specic vulnerabilities and ad-
aptation strategies.
From 1990 to 2025, there was a noticeable rise in
the number of scholarly articles about the climate
change impact on mackerel in the world (Figure 1).
China has 24 articles, which is the most in amount.
Other countries are Japan, the United States, Aus-
tralia, and various European countries, all of which
have moderate contributions. Some countries, such
as a large portion of South America, Africa, and
parts of Asia, have made little to no contribution.
The bar graph differentiates between Single
Country Publications (SCP) and Multiple Coun-
try Publications (MCP), which displays the coun-
try-by-country distribution of publications related
to this study topic (Figure 2). While China has
some collaborative works, the majority of its pub-
lications are single-country publications (SCP).
Norway and Korea are next to China, both mak-
ing signicant contributions, especially in the area
of international collaboration (MCP). With a bal-
anced combination of SCP and MCP, the UK, USA,
and Australia likewise exhibit signicant research
output. With a comparatively higher percentage
of MCP, other European countries such as Spain,
Denmark, Iceland, and Portugal make moderate
contributions, suggesting active international col-
laboration. With fewer articles, Malaysia and India
appear to have a lesser but developing eld of study
in this area. Only one or two publications are con-
tributed by countries like Belgium and Morocco.
While many other countries, particularly in Europe
and North America, actively contribute through in-
ternational collaborations, this gure shows that
China dominates the research output. Southeast
Asia, except Malaysia, is insufciently represented,
suggesting a research vacuum in the area.
There was very little research activity during
the years 1987 to 2007, with only 0 to 2 articles
published annually, suggesting that the topic did
not receive much attention worldwide at that time-
period (Figure 3). From 2008 to 2013, there was
a steady rise, with publication counts exceeding
Marine and Fishery sciences 39 (2): xxx-xxx (2026)
6
Figure 2. Most relevant countries that published articles on this topic from 1990 to 2025.
Figure 1. Number of scientic publications on the effect of climate change on mackerel populations globally from 1990 to 2025.
China
Korea
Norway
United Kingdom
USA
Australia
Canada
Spain
Japan
Iceland
Denmark
India
Portugal
Italy
Malaysia
Sweden
Chile
France
Morocco
Belgium
Countries
No. of documents
0510 15 20 25
Single country publications Multiple country publications
Haque et al.: Climate CHange impaCt, global researCH trend, and regional gaps in soutHeast asia 7
5 to 7 annually, indicating the start of a growing
interest. As a result of increased global concern
over the effects of climate change on sheries, re-
search output increased dramatically from 2014 to
2021, reaching a peak from 2019 to 2021 with over
15 articles annually. There has been a decrease in
the number of articles published in recent years,
from 2022 to 2025, which may be partially due
to insufcient data availability. The graph shows
that, from late 2000, research on this topic has
grown dramatically, reaching a high in the year
2019 to 2021, coinciding with the global upsurge
in sheries and climate change research. It is im-
portant to be cautious when interpreting the fall
after 2021 because it can be the result of database
indexing delays rather than a real decline in re-
search productivity.
With 11 articles each, Frontiers in Marine Sci-
ence and Marine Ecology Progress Series were the
two primary sources in producing research on this
particular eld (Figure 4). Fisheries Oceanography
and ICES Journal of Marine Science came in after
them with 9 articles each. Each journal, Fisheries
Research and Ocean Science, had 5 articles. After
that, 4 and 3 articles were given by the following
journals.
Leading academic and research institutions con-
tribute to the eld determining whether the subject
is dispersed throughout numerous institutions or
is dominated by a small number of them (Figure
5). With 37 articles, the Institute of Marine Re-
search is the leading contributor. Ocean Univer-
sity of China comes in second with 35 articles.
Shanghai Ocean University comes after that with
31 publications. Mid-level contributors include Jeju
National University with 11 articles, the Institute
Maurice-Lamontagne with 10 articles, and the fol-
lowing institutes.
Tian Y is the most prolic author on this topic,
having published 7 articles (Figure 6). Signicant
contributions were made by the four authors: Chen
X, Jung S, Li J, and Liu Y, each of whom published
6 articles. Then comes Jansen T and Oskarsson GJ,
each with 5 articles.
Research gap: southeast Asia context
While major oceanic basins like the Pacic
and Atlantic Oceans are represented, southeast
Asia does not explicitly appear, suggesting a po-
tential regional gap in research coverage. Rising
sea surface temperature, more frequent extreme
Figure 3. Annual scientic production over time.
Artlicles
Years
15
10
5
0
1987 1989 1991 1993 1995 1997 1999 2001 2003 2005 2007 2009 2011 2013 2015 2017 2019 2021 2023 2025
Marine and Fishery sciences 39 (2): xxx-xxx (2026)
8
weather events, and ocean acidication are hav-
ing a major inuence on marine biodiversity and
sheries production (Islam et al. 2020). Southeast
Asia is among the most vulnerable regions to cli-
mate change (Sumaila et al. 2011). A commercially
valuable species in the area, Spanish mackerel (S.
commerson), suffers difculties because of shifting
oceanographic conditions that affect their spawning
cycles, migration patterns, and availability of prey
(Cheung et al. 2013).
The network was generated in VOSviewer us-
ing the association-strength normalization and full
counting method. Keywords were included if they
appeared at least 5 times in the combined Scop-
us-Web of Science dataset (n = 257 articles; n = 54
keywords). Links with a total link strength < 2 were
Figure 4. Most relevant sources of publications.
Frontiers in Marine Science
Marine Ecology Progress Series
Fisheries Oceanography
ICES Journal of Marine Science
Fisheries Research
Ocean Science Journal
Deep Sea Research
Progress in Oceanography
Journal of Fish Biology
Journal of Oceanology and Limnology
No. of documents
03
Sources
6 9
3
3
5
9
11
4
4
5
9
11
Figure 5. Most relevant afliations of publications.
Institute of Marine Research
Ocean University of China
Shanghai Ocean University
Jeju National University
Institute Maurice-Lamontagne
Basque Research and Technology Alliance
Memorial University of Newfoundland
National Taiwan Ocean University
Technical University of Denmark
Flanders Marine Institute
Articles
010 20 30
8
Affiliations
9
9
9
9
10
11
31
35
37
Haque et al.: Climate CHange impaCt, global researCH trend, and regional gaps in soutHeast asia 9
removed for clarity. Nodes represent keywords, and
node size reects keyword frequency. Colors repre-
sent clusters produced using the VOSviewer (Lou-
vain-type) modularity-based clustering algorithm,
identifying major thematic groups: 1) Blue cluster:
climate-ecology interactions (e.g. climate change,
population dynamics, ecosystems). 2) Green clus-
ter: distribution and environmental drivers (e.g. sea
surface temperature, climate models, population
distribution). 3) Yellow-orange cluster: sher-
ies-focused terms (e.g. tuna shery, CPUE, catch
statistics). 4) Turquoise cluster: species-level tax-
onomy and biology (e.g. S. japonicus, S. scombrus,
pelagic sh). The spatial arrangement of nodes was
produced using the LinLog layout algorithm, which
emphasizes cluster separation and inter-cluster link
structure. A high network modularity (Q = 0.62),
high average intra-cluster density (0.41), and a low
proportion of inter-cluster links (18%), indicating
strong internal cohesion and limited cross-cluster
connectivity (Noack 2009) (Figure 7).
The network was generated in VOSviewer us-
ing the association-strength normalization and full
counting method. Keywords were included based on
a minimum occurrence threshold of 5 appearances
in the regional dataset (n = 105 publications; n = 39
unique keywords. Links with very weak co-occur-
rence strength (total link strength < 2)were ltered
to improve visual clarity. Nodes represent author
keywords, with node size proportional to keyword
frequency. Colors denote clusters identied by the
VOSviewer modularity-based (Louvain-type) clus-
tering algorithm, revealing major thematic struc-
ture in southeast Asian literature: 1) Yellow cluster:
ecology-climate interface (climate change, ecosys-
tems, marine ecosystem, sh). 2) Green cluster:
sheries management and environmental drivers
(sheries, shery management, sustainable devel-
opment, surface waters). 3) Pink cluster: shery-de-
pendent indicators and variability drivers (Indian
Ocean, tuna shery, CPUE, catch statistics, ENSO,
climate variation). 4) Blue cluster: species-specif-
ic and biogeographic terms (tuna, Pacic Ocean,
spatial distribution, global warming). The spatial
structure of the map was generated using the Lin-
Log layout, which enhances cluster separation by
prioritizing attractive and repulsive forces among
strongly and weakly related keywords, respective-
ly. This visualization highlights the dominance of
Indian Ocean-centric, catch-based, sheries-driven
research in southeast Asia compared to more mech-
anistic and model-based global clusters (Figure 8).
Figure 6. Most prolic authors of publications on the subject.
No. of documents
024 6
4
4
4
5
5
6
6
6
6
7
Tian Y
Chen X
Jung S
Li J
LiuY
Jansen T
Oskarsson GJ
Go S
Ito S-I
MontevecchiWA
Authors
Marine and Fishery sciences 39 (2): xxx-xxx (2026)
10
Figure 8. Keyword co-occurrence network map for southeast Asian climate-sheries research related to mackerel and other
Scombridae species.
Figure 7. Keyword co-occurrence network map for global climate-sheries research related to mackerel and other Scombridae
species.
Haque et al.: Climate CHange impaCt, global researCH trend, and regional gaps in soutHeast asia 11
In contrast to global research in the Atlantic and
Pacic, which has progressed toward predictive
modeling, physiological studies, and integrated
socio-ecological assessments, southeast Asian cli-
mate-sheries research is still catch-based and de-
scriptive, concentrating on tuna/mackerel sheries
within the Indian Ocean framework, according to
the comparison of bibliographic maps (Table 1).
This leaves a visible gap: southeast Asia, where
reliance on sheries for food security is high, but
data or modeling investment lags, requires SDM-
based predictions, mechanistic studies, and socio-
economic integration.
Climate change themes identied in the global
research
The thematic cluster analysis and keyword
co-occurrence mapping revealed four dominant
global research themes related to the impacts of
climate change on mackerel: 1) Ocean warming
and distribution shifts, 2) Migration and spawn-
ing changes, 3) Ecosystem-wide trophic responses,
and 4) Physiological stress and behavioral impacts.
These clusters align with well-established ecologi-
cal evidence showing that warming, acidication,
and deoxygenation are reshaping pelagic ecosys-
tems worldwide. Studies highlight that rising sea
temperatures drive poleward species movements
(Astthorsson et al. 2012; Wang and Wu 2025), al-
ter spawning grounds (Jansen and Gislason 2013),
and cause contractions in traditional habitats (Yang
et al. 2022). Emerging evidence of rapid ocean
deoxygenation (Lu et al. 2024) and widespread
reef hypoxia (Pezner et al. 2023) further reinforces
the global research emphasis on environmental
stressors affecting mackerel habitat suitability and
recruitment.
Table 1. Comparison between the bibliometric analysis of Global and southeast Asian research.
Global Southeast Asia
SST = Sea Surface Temperature; ENSO = El-Niño Southern Oscillation; CPUE = Catch Per Unit Effort.
Core keywords
Research orientation
Data sources
Climate drivers
studied
Analytical approach
Ecological scope
Management
dimension
Research
development
Atlantic Ocean, Pacic Ocean, climate change,
pelagic sh, population dynamics, stock
assessment, physiology, climate models
Mechanistic, predictive, ecosystem-based
Multi-source data (survey data, ecological
monitoring, physiology, models, remote sensing)
Integrated drivers: warming, deoxygenation,
acidication, multi-factor climate models
Species Distribution Models (SDMs), spatio-
temporal models, ensemble climate projections
Ecosystem level focus (food webs, spawning,
physiology, ecosystem resilience)
Strong: stock assessment, adaptive sheries
management, socio-ecological integration
Higher data availability and modeling investment,
advanced, integrating ecology, modeling, and
management scenarios
Indian Ocean, Scombridae, tuna
shery, SST, ENSO, CPUE, catch
statistics.
Descriptive, monitoring-oriented
Fishery dependent data (catch, CPUE,
abundance, biomass, effort)
Broad variables: SST, ENSO, climate
variability
Statistical correlations, trend analysis
Species/shery level focus (mainly
tunas and mackerels)
Limited: mostly linked to catch trends
and variability
Often generalized, emerging in
methodological sophistication
Marine and Fishery sciences 39 (2): xxx-xxx (2026)
12
Distribution shifts and habitat loss as central
research themes
Across global publications, distribution shifts
associated with sea-surface temperature increases
appear consistently in high-frequency keywords
and tightly connected network clusters. Multiple
studies document northward expansions of Atlantic
and Chub mackerel (dos Santos Schmidt et al. 2024;
Han et al. 2024) and projected habitat losses of up to
80-90% under high emission scenarios for Scomb-
eromorus species (Yang et al. 2022). These ndings
correspond with the strong ‘warming-distribution
shift-habitat suitability’ cluster in the VOSviewer
network, demonstrating that habitat modelling and
mechanistic projections dominate global research.
Migration and spawning modications: a sec-
ondary but growing theme
Bibliometric results showed a medium-density
cluster associated with migration, spawning, re-
cruitment, and larval dispersal, reecting a substan-
tial research focus on phenological changes. Article
within this cluster reports earlier spawning (Brunel
et al. 2018), altered migration timing (Kanamori et
al. 2019), and climate-driven mismatches between
larvae and plankton blooms (Edwards and Rich-
ardson 2004). This articles depict how researchers
increasingly integrate biological observations with
climate variability, especially ENSO, highlighting a
trend toward process-based ecological studies, par-
ticularly in the Pacic and North Atlantic.
Ecosystem and trophic interactions: linking cli-
mate drivers to food web response
The third major cluster identied dealt with eco-
system and trophic interactions, including prey
availability, zooplankton dynamics, and predator
overlap. Highly cited global studies showed that
warming reduces the abundance of key copepods
(Calanus sp.) restructured plankton communities
(Neven 2024), and intensies competition among
pelagic predators (Ono et al. 2024). The co-occur-
rence network reected this through strong asso-
ciations between terms such as ‘plankton’, ‘troph-
ic’, ‘productivity’, and ‘recruitment’. These results
demonstrated that much of the global research land-
scape has moved beyond descriptive assessments
toward ecosystem-based and food web-integrated
approaches.
Physiological and behavioral responses: a small-
er but emerging theme
Although less central in keyword networks, a
growing cluster focused on physiological responses
such as altered growth, metabolic stress, and behav-
ioral modications linked to warming and acidi-
cation (Munday et al. 2009; Pecl et al. 2017). Stud-
ies reporting projected poleward shifts in thermal
habitat suitability for S. commerson (Wang and Wu
2025) and vulnerability of juveniles to both acidi-
cation and hypoxia (Wexler et al. 2023) aligned
with this research trend. These emerging studies
highlight a transition toward climate-physiology
integration in global sheries science.
Fisheries management and adaptation: policy
themes in global versus regional research
The bibliometric mapping also revealed a clear
difference between global and southeast Asian
articles. Globally, research connects climate im-
pacts to adaptive management, ecosystem-based
governance, and transboundary stock agreements.
This includes emphasis on climate-resilient MPAs
(Marine Protected Areas), quota adjustments, and
long-term monitoring (Lehodey et al. 2020; Bagsit
et al. 2021). However, southeast Asian publications
show limited focus on governance, instead empha-
sizing catch trends, CPUE, and descriptive stock
assessments. Global studies increasingly integrate
climate projections into policy frameworks (Pandey
and Joseph 2025), whereas regional research points
to persistent gaps in funding, monitoring capacity,
and institutional support. These ndings indicate
Haque et al.: Climate CHange impaCt, global researCH trend, and regional gaps in soutHeast asia 13
that climate-driven risks to mackerel sheries are
well documented at the global scale, but scientic,
nancial, and governance constraints continue to
limit adaptive responses in southeast Asia.
Research prospects in southeast Asia
In order to improve knowledge and management
of mackerel sheries in light of climate change, a
number of important research gaps need to be lled.
The long term effects of climate variability on the
distribution and productivity of mackerel stocks,
which are still poorly understood in southeast Asia,
need to be studied at the regional level (Breitburg
et al. 2018; Dongyu 2024). The physiological and
behavioral responses of mackerel to shifting ocean
circumstances, such as rising sea temperatures and
falling oxygen levels, need further research, espe-
cially in relation to migration patterns, predator
avoidance, and larval survival (Breitburg et al. 2018;
Wexler et al. 2023). Despite growing evidence that
coastal populations in the Indo-Pacic region are
extremely vulnerable to climatic shocks, there are
currently few studies looking at the socioeconomic
implications of declining mackerel stocks, partic-
ularly the exibility and resilience of small-scale
shermen (Pecl et al. 2017; Chamsai and Wanchana
2022). To ensure the sustainability of mackerel sh-
eries, studies on the efcacy of adaptive sheries
management techniques, such as ecosystem-based
management and cooperative regional governance,
are required (Cheung et al. 2013). Few studies have
examined how climate change impacts prey species
availability and composition for mackerel, which is
important for forecasting population patterns in the
future (Doney et al. 2012; Wang and Wu 2025). De-
veloping climate resilient management frameworks
for mackerel sheries, these research gaps need to
be lled by integrated ecological, socioeconomic,
and governance studies.
These gaps underscore a pressing need for in-
tegrated ecological, socioeconomic, and gover-
nance research to support the development of cli-
mate-resilient sheries management frameworks
for mackerel in southeast Asia. Aligning regional
research with global methodological advances, par-
ticularly through SDMs, long-term monitoring, and
socio-ecological assessments, would signicantly
improve the scientic basis for sustainable manage-
ment under accelerating climate change.
CONCLUSIONS
While global studies increasingly employ mech-
anistic models, ecosystem-based approaches, and
integrated climate projections, southeast Asian re-
search remains largely descriptive and catch-based.
The keyword structures and thematic clusters con-
rm a persistent regional gap in modelling capacity,
physiological research, and socio-ecological assess-
ment, despite the region’s strong dependence on
mackerel sheries for food security. Closing this
gap will require strengthening scientic infrastruc-
ture, improving data availability, and expanding
interdisciplinary work on species distribution mod-
elling, multi-stressor climate analysis, and socioec-
onomic vulnerability. The ndings also highlight
the need for adaptive, evidence-driven sheries
governance, including ecosystem-based manage-
ment and coordinated regional monitoring. Overall,
a more integrated research and policy landscape is
essential for building climate resilience in southeast
Asian mackerel sheries. Investing in modelling,
collaborative networks, and long-term monitoring
will support sustainable harvest and help safeguard
coastal communities.
ACKNOWLEDGEMENTS
This work was funded by the Fundamental Re-
search Grant Scheme (FRGS) with project reference
FRGS/1/2024/WAS05/UMT/02/4, and the authors
gratefully acknowledge the Malaysian Ministry of
Higher Education (MoHE) for their support.
Marine and Fishery sciences 39 (2): xxx-xxx (2026)
14
Conicts of interest
The author(s) declare(s) that there is no conict
of interest regarding the publication of this paper.
Author contributions
Mohammad Ekramul Haque: writing the original
manuscript and analyzing the data. Shahadat Hos-
sain: reviewed and analysed the data. Tun Nurul
Aimi Mat Jaafar: review and edit the draft. Ying
Giat Seah: review and edit the draft. Siti Azizah
Mohd Nor: review and edit the draft. Nazia Abdul
Kadar: review and edit the draft. Nur Fadli: review
and edit the draft. Darlina Md Naim: review and
edit the draft. Helena Khatoon: review and edit
the draft. Ahasan Habib: conceptualised the topic,
reviewed, edited, and supervised.
REFERENCES
aria M, cuccurullo c. 2017. bibliometrix: An
R-tool for comprehensive science mapping
analysis. J Informetr. 11 (4): 959-975.
astthorsson os, ValdiMarsson h, gudMunds-
dottir a, Óskarsson gJ. 2012. Climate-relat-
ed variations in the occurrence and distribution
of mackerel (Scomber scombrus) in Icelandic
waters. ICES J Mar Sci. 69 (7): 1289-1297.
bagsit Fu, FriMpong e, asch rg, Monteclaro
hM. 2021. Effect of a seasonal shery closure
on sardine and mackerel catch in the Visayan
Sea, Philippines. Front Mar Sci. 8: 640772.
barange M, Merino g, blanchard Jl, schol-
tens J, harle J, allison eh, allen Ji, holt
J, Jennings s. 2014. Impacts of climate change
on marine ecosystem production in societies
dependent on sheries. Nat Clim Change. 4 (3):
211-216.
breitburg d, leVin la, oschlies a, grégoire
M, chaVez Fp, conley dJ, garçon V, gilbert
d, gutiérrez d, isensee k. 2018. Declining
oxygen in the global ocean and coastal waters.
Science. 359 (6371): eaam7240.
brunel t, Van daMMe cJ, saMson M, dick-
ey-collas M. 2018. Quantifying the inuence
of geography and environment on the northeast
Atlantic mackerel spawning distribution. Fish
Oceanogr. 27 (2): 159-173.
chaMsai s, Wanchana W. 2022. Issues and chal-
lenges in sustainable development of sheries
and aquaculture of the Southeast Asian Region:
sheries-related issues: climate change and nat-
ural disasters. Southeast Asian State Fish Aquac.
p. 197-204.
cheung WW, laM VW, sarMiento Jl, kearney
k, Watson reg, zeller d, pauly d. 2010.
Large-scale redistribution of maximum sheries
catch potential in the global ocean under cli-
mate change. Glob Change Biol. 16 (1): 24-35.
cheung WW, Watson r, pauly d. 2013. Signa-
ture of ocean warming in global sheries catch.
Nature. 497 (7449): 365-368.
doney sc, ruckelshaus M, duFFy Je, barry
Jp, chan F, english ca, galindo hM, greb-
Meier JM, holloWed ab, knoWlton n. 2012.
Climate change impacts on marine ecosystems.
Annu Rev Mar Sci. 4: 11-37.
dongyu q. 2024. 2024 the state of world sheries
and aquaculture-blue transformation in action.
State World Fish Aquac. p. 1-232.
dos santos schMidt tc, slotte a, olaFsdot-
tir ah, nøttestad l, Jansen t, Jacobsen Ja,
bJarnason s, lusseau sM, ono k, hølle-
land s, et al. 2024. Poleward spawning of At-
lantic mackerel (Scomber scombrus) is facilitat-
ed by ocean warming but triggered by energetic
constraints. ICES J Mar Sci. 81 (3): 600-615.
DOI: https://doi.org/10.1093/icesjms/fsad098
edWards M, richardson aJ. 2004. Impact of cli-
mate change on marine pelagic phenology and
trophic mismatch. Nature. 430 (7002): 881-884.
Fabro ka. 2020. Gray areas and weak policies mar
lucrative Asian trade in live reef sh. Mongabay.
[accessed 2025 Dec 31]. https://news.mongab-
ay.com/2020/07/gray-areas-and-weak-policies-
Haque et al.: Climate CHange impaCt, global researCH trend, and regional gaps in soutHeast asia 15
mar-lucrative-asian-trade-in-live-reef-sh/.
[Fao] Food and agriculture organization oF
the united nations. 2018. Rome: FAO. URL
Httpfaostat Fao Org. 403 p.
Free cM, thorson Jt, pinsky Ml, oken kl,
WiedenMann J, Jensen op. 2019. Impacts of
historical warming on marine sheries produc-
tion. Science. 363 (6430): 979-983.
han h, shang c, Jiang b, Wang y, li y, xiang
d, zhang h, shi y, Jiang k. 2024. A new mod-
eling strategy for the predictive model of chub
mackerel (Scomber japonicus) central shing
grounds in the Northwest Pacic Ocean based
on machine learning and operational charac-
teristics of the light shing vessels. Front Mar
Sci. 11: 1451104. DOI: https://doi.org/10.3389/
fmars.2024.1451104
habib a, borazon eq, nallos iM, Macusi e.
2025. Climate change vulnerability, adaptation
and ecosystem services in different sheries and
aquaculture in Asia: a review. Mar Fish Sci. 38
(2): 311-330.
islaM MM, islaM n, habib a, MozuMder MMh.
2020. Climate change impacts on a tropical
shery ecosystem: implications and societal
responses. Sustainability. 12 (19): 7970.
Jansen t, gislason h. 2013. Population struc-
ture of Atlantic mackerel (Scomber scombrus).
PLoS ONE. 8 (5): e64744.
kaMaruzzaMan yn, Mustapha Ma, ghaFFar
Ma. 2021. Impacts of sea temperature rise on
Rastrelliger kanagurta potential shing grounds
in the Exclusive Economic Zone (EEZ) off South
China Sea. Sains Malays. 50 (12): 3467-3479.
kanaMori y, takasuka a, nishiJiMa s, okaMura
h. 2019. Climate change shifts the spawning
ground northward and extends the spawning
period of chub mackerel in the western North
Pacic. Mar Ecol Prog Ser. 624: 155-166.
lehodey p, bertrand a, hobday aJ, kiyoFu-
Ji h, Mcclatchie s, Menkès ce, pilling g,
poloVina J, toMMasi d. 2020. ENSO impact
on marine sheries and ecosystems. El Niño
South Oscil Chang Clim. p. 429-451.
lu b, zhao z, han l, gan x, zhou y, zhou l,
Fu l, Wang x, zhou c, zhang J. 2024. OX-
YGENERATOR: reconstructing global ocean
deoxygenation over a century with deep learn-
ing. [accessed 2025 Aug 16]. DOI: https://doi.
org/10.48550/ARXIV.2405.07233.
Macusi ed, Macusi es, JiMenez la, cataM-isan
Jp. 2020. Climate change vulnerability and
perceived impacts on small-scale sheries in
eastern Mindanao. Ocean Coast Manag. 189:
105143.
Macusi ed, nallos iV, santos Md, geroniMo
rc. 2025b. Climate and non-climate related
hazards in small pelagic sheries and milksh
aquaculture: expert opinion survey in the Phil-
ippines. Philipp J Fish. 32 (1): 178-196.
Macusi ed, sabino ll, pislan ht, Macusi es.
2025a. Impacts of extreme climate change event
on small-scale shers and their adaptation in
Baganga, Davao Oriental. World. 25: 6-18.
Moher d, shaMseer l, clarke M, ghersi d,
liberati a, petticreW M, shekelle p, steW-
art la, group p-p. 2015. Preferred reporting
items for systematic review and meta-analysis
protocols (PRISMA-P) 2015 statement. Syst
Rev. 4 (1): 1.
Munday pl, dixson dl, donelson JM, Jones
gp, pratchett Ms, deVitsina gV, døVing kb.
2009. Ocean acidication impairs olfactory dis-
crimination and homing ability of a marine sh.
Proc Natl Acad Sci. 106 (6): 1848-1852.
neVen cJ. 2024. Different perspectives on zoo-
plankton functioning in the Southern North Sea
and the English Channel in relation to lower
and higher trophic levels [doctoral dissertation].
Dunkerque: Université du Littoral Côte d’Opale.
noack a. 2009. Modularity clustering is force-di-
rected layout. Phys Rev E. 79 (2): 026102.
ono k, katara i, eliasen sk, broMs c, caMp-
bell a, dos santos schMidt tc, egan a,
hølleland sn, Jacobsen Ja, Jansen t. 2024.
Effect of environmental drivers on the spatio-
temporal distribution of mackerel at age in the
Nordic Seas during 2010-20. ICES J Mar Sci.
Marine and Fishery sciences 39 (2): xxx-xxx (2026)
16
81 (7): 1282-1294.
pan x, chen y, Jiang t, yang J, tian y. 2024.
Otolith biogeochemistry reveals possible im-
pacts of extreme climate events on population
connectivity of a highly migratory sh, Japa-
nese Spanish mackerel Scomberomorus nipho-
nius. Mar Life Sci Technol. 6 (4): 722-735.
pandey a, Joseph J. 2025. Emerging trends in blue
economy: a roadmap through the lens of sus-
tainable development. Secur Sustain Futur Blue
Green Econ. p. 17-36.
pecl gt, araúJo Mb, bell Jd, blanchard J,
bonebrake tc, chen i-c, clark td, col-
Well rk, danielsen F, eVengård b. 2017. Bi-
odiversity redistribution under climate change:
Impacts on ecosystems and human well-being.
Science. 355 (6332): eaai9214.
pezner ak, courtney ta, barkley hc, chou
W-c, chu h-c, cleMents sM, cyronak t,
degrandpre Md, kekueWa sah, kline di,
et al. 2023. Increasing hypoxia on global coral
reefs under ocean warming. Nat Clim Change.
13 (4): 403-409. DOI: https://doi.org/10.1038/
s41558-023-01619-2
pinsky Ml, Mantua nJ. 2014. Emerging adap-
tation approaches for climate-ready sheries
management. Oceanography. 27 (4): 146-159.
pörtner h-o, karl dM, boyd pW, cheung W,
lluch-cota se, noJiri y, schMidt dn, za-
VialoV po, alheit J, aristegui J. 2014. Ocean
systems. In: Climate change 2014: impacts, ad-
aptation, and vulnerability. Part A: global and
sectoral aspects. Contribution of working group
II to the fth assessment report of the intergov-
ernmental panel on climate change. Cambridge
University Press. p. 411-484.
ren J, liu q, Ma y, Ji y, xu b, xue y, zhang
c. 2025. Spatio-temporal distribution of four
trophically dependent shery species in the
Northern China seas under climate change. Bi-
ology. 14 (2): 168.
suMaila ur, cheung WW, laM VW, pauly d,
herrick s. 2011. Climate change impacts on
the biophysics and economics of world sheries.
Nat Clim Change. 1 (9): 449-456.
sun J-z, zhang d, yi x, beardall J, gao k.
2024. Ocean deoxygenation dampens resistance
of diatoms to ocean acidication in darkness.
Front Mar Sci. 11: 1387552. DOI: https://doi.
org/10.3389/fmars.2024.1387552
talbot e, Jontila J-bs, gonzales bJ, dolorosa
rg, Jose ed, saJorne r, sailley s, kay s,
queirÓs aM. 2024. Incorporating climate-read-
iness into sheries management strategies. Sci
Total Environ. 918: 170684.
Van eck n, WaltMan l. 2010. Software survey:
VOSviewer, a computer program for bibliomet-
ric mapping. Scientometrics. 84 (2): 523-538.
Vu nk, nguyen kq. 2025. The Effect of temporal
and environmental conditions on catch rates of
the narrow-barred Spanish mackerel setnet sh-
ery in Khanh Hoa Province, Vietnam. Fishes.
10 (6): 257.
Wang q, Wu J. 2025. The origin and global spread
of Scomberomorus spp.: the intersection of eco-
logical adaptation and sheries management.
Int J Mar Sci. 15: 102345
Wexler Jb, Margulies d, scholey V, len-
nert-cody ce, broMhead d, nicol s, hoyle
sd, stein M, WilliaMson Je, haVenhand J.
2023. The effect of ocean acidication on oto-
lith morphology in larvae of a tropical, epipe-
lagic sh species, yellown tuna (Thunnus al-
bacares). J Exp Mar Biol Ecol. 569: 151949.
yang t, liu x, han z. 2022. Predicting the ef-
fects of climate change on the suitable habitat
of Japanese Spanish mackerel (Scomberomorus
niphonius) based on the Species Distribution
Model. Front Mar Sci. 9: 927790. DOI: https://
doi.org/10.3389/fmars.2022.927790