Journal of Sea Research 193 (2023) 102386
Contents lists available at ScienceDirect
Journal of Sea Research
journal homepage: www.elsevier.com/locate/seares
A scientometric review of climate change and research on crabs
Chandra Segaran Thirukanthan a, *, Mohamad Nor Azra a, b, c, *, Nor Juneta Abu Seman b,
Suzanne Mohd Agos b, Hidir Arifin b, Hani Amir Aouissi d, e, f, Fathurrahman Lananan g,
Huan Gao h
a
Institute of Marine Biotechnology, Universiti Malaysia Terengganu, 21030 Kuala Nerus, Terengganu, Malaysia
Climate Change Adaptation Laboratory, Institute of Marine Biotechnology (IMB), Universiti Malaysia Terengganu (UMT), 21030 Kuala Nerus, Terengganu, Malaysia
Research Center for Marine and Land Bioindustry (Earth Sciences and Maritime), National Research and Innovation Agency (BRIN), Pemenang, West Nusa Tenggara,
83352, Indonesia
d
Scientific and Technical Research Center on Arid Regions (CRSTRA), Biskra 07000, Algeria
e
Laboratoire de Recherche et d’Etude en Aménagement et Urbanisme (LREAU), Université des Sciences et de la Technologie (USTHB), Algiers 16000, Algeria
f
Environmental Research Center (CRE), Badji-Mokhtar Annaba University, Annaba 23000, Algeria
g
East Coast Environmental Research Institute, Universiti Sultan Zainal Abidin, Gong Badak Campus, 21300 Kuala Nerus, Terengganu, Malaysia
h
School of Marine Science and Fisheries, Jiangsu Ocean University, Lianyungang 222005, Jiangsu, China
b
c
A R T I C L E I N F O
A B S T R A C T
Keywords:
Adaptation
Climate-induced temperature changes
Harmful algal bloom
Hypoxia
Ocean acidification
Predation
Sea level rise
Crabs categorized as cold-blooded organisms are especially at risk as climate change worsens. Their current
situation was not well documented, especially in terms of scientometric analysis. The present study aims to
investigate the relationship between research on crabs and climate change-related studies, with a focus on
identifying trends and hotspots over time. The analysis was based on a collection of over 2834 relevant docu­
ments and 107,502 cited references indexed in the Web of Science Core Collection (WOSCC) database from 1977
to 2022. The findings indicated an increase in research in recent decades, with the USA as the largest contributor,
followed by China and Brazil. Researchers from the USA and Germany were among the top published authors in
the field. The most highly cited studies in WOSCC focused on the relationship between harmful algal blooms and
crab research. Of these studies, 20 clusters were generated, with the most influential cluster identified as related
to “ocean acidification,” “blue king crab,” and “mud crab fishery.” The most frequently cited and influential
keywords in the field were “climate change” and “hypoxia,” respectively. Our conclusion is that the fields of
“research on crabs” and “climate change” are thriving and that further exploration of the adaptation strategies of
these organisms is necessary. This knowledge will benefit scientific communities, philanthropic funders or
related governments, fisheries-related industries, and NGOs towards the sustainable management of commercial
crab species in the future.
1. Introduction
The interrelated issues of climate change, fisheries, food security,
and biodiversity play a crucial role in the projected increase of the
human population by 2050 (Rice and Garcia, 2011; Béné et al., 2015;
Islam and Wong, 2017; Schnitter and Berry, 2019; Muluneh, 2021). In
recent years, climate change has become the focus of intense debate and
discussion (Hassan et al., 2023). Climate change refers to long-term al­
terations in the mean or variability of weather patterns or the Earth’s
surface temperature within the Earth’s atmosphere, typically over de­
cades or longer (WBCKP World Bank Climate Change Knowledge Portal,
2023). This change is primarily driven by the emission of gases such as
carbon dioxide, which trap heat from the sun in the Earth’s lower at­
mosphere, leading to an increase in the Earth’s temperature known as
global warming (Bolaji and Huan, 2013; Farmer and Cook, 2013; Jakada
et al., 2022; Dhaka and Kumar, 2023). These changes can result in a
variety of impacts, including heatwaves, changes in precipitation pat­
terns, heavy floods, and rising sea levels (Mohd et al., 2019; Zscheischler
et al., 2020; Ødemark et al., 2023). Climate change is projected to have
significant impacts on global ecology, particularly on poikilothermic
animals, altering the distribution, abundance, behaviour, and physi­
ology of populations and communities within these groups (Gutierrez
* Corresponding authors at: Institute of Marine Biotechnology (IMB), Universiti Malaysia Terengganu (UMT), 21030 Kuala Nerus, Terengganu, Malaysia.
E-mail addresses: thiru@umt.edu.my (C.S. Thirukanthan), azramn@umt.edu.my (M.N. Azra), fathurrahman@unisza.edu.my (F. Lananan).
https://doi.org/10.1016/j.seares.2023.102386
Received 4 April 2023; Received in revised form 7 May 2023; Accepted 8 May 2023
Available online 9 May 2023
1385-1101/© 2023 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/bync-nd/4.0/).
C.S. Thirukanthan et al.
Journal of Sea Research 193 (2023) 102386
et al., 2008; Lister and Garcia, 2018; Missionário et al., 2022; Bowman
Jr and Post, 2023).
Poikilothermy refers to the characteristic of organisms whose body
temperature is not internally regulated but is instead influenced by their
external environment. In contrast, homeothermic or warm-blooded or­
ganisms, such as mammals and birds, maintain a relatively constant
body temperature despite variations in their surroundings (Wood, 2005;
Gracey et al., 2004). Crabs, as aquatic organisms, are poikilothermic,
meaning that their physiological processes, including development,
reproduction, and metabolism, are heavily dependent on the tempera­
ture of their environment. For example, elevated water temperatures
can increase crabs’ metabolism, leading to enhanced growth and
reproduction (Robinson, 2010). However, excessive temperature in­
creases can result in stress and decreased survival. Conversely, low
water temperatures can reduce metabolism and hinder growth and
reproduction. Furthermore, water temperature fluctuations can impact
the reproduction timing and the development of eggs and larvae (Culos
and Tyson, 2014). As poikilothermic organisms, crabs are particularly
vulnerable to the impacts of climate change, as rising sea temperatures
brought on by global warming can disrupt their physiology and repro­
duction, potentially leading to population declines (Briffa et al., 2013).
Crabs are of significant importance, as they are a potential food source
and play a vital role in fisheries and ecosystem services (Azra et al.,
2021; Henriksson et al., 2021). They have been identified as one of the
groups that may be affected by climate change (Ray et al., 2019; Azra
et al., 2022).
Scientometrics is a field that studies the quantitative aspects of sci­
ence and technology, including the measurement of research produc­
tivity, impact, and trends. As a branch of the discipline known as the
“science of science,” scientometrics has been widely used across various
fields of study to identify patterns and thematic scope of selected topics
(Chen and Song, 2019; Chen, 2020; Chen, 2022). It is widely recognized
as a powerful tool for research and innovation policy (Ivancheva, 2008).
The application of these quantitative methods also allows for mapping
the disciplinary structure and inter-relations of research dynamics, as
well as identifying the characteristics of potential research areas for the
future (Kovács et al., 2015; Daradkeh et al., 2022). The CiteSpace soft­
ware can process scientometrics records as well as cited references from
sources such as the Web of Science, Scopus, Lens, and Dimension (Chen,
2022). Cited references are considered a crucial criterion in broadening
the scope of scientific analysis and providing a more comprehensive
understanding of the topic and the quality of research (Aksnes et al.,
2019).
This scientometric review seeks to elucidate the progression and
contemporary trends in the domain of crab research vis-à-vis climate
change, employing the comprehensive WOSCC database as its resource.
The overarching goal is to forecast emerging research focal points by
scrutinizing various dimensions of the existing literature, including: (i)
publication and citation metrics, (ii) highly influential articles, (iii)
authorship, institutional affiliations, preferred journals, and geograph­
ical distribution, (iv) collaborative networks and funding bodies, (v)
burgeoning research disciplines, (vi) cluster examination of scholarly
works, and (vii) keyword co-occurrence patterns, dissemination, and
burst dynamics.
Considering the pressing need for sustainable bioresource manage­
ment and biodiversity conservation amid shifting climatic conditions, a
thorough examination of diverse research avenues is paramount. The
outcomes of this scientometric review are anticipated to offer valuable
guidance for a wide array of stakeholders, encompassing crab fishers,
postgraduate scholars, funding agencies, academic institutions, and
early career investigators, thereby laying the groundwork for future
inquiries. Furthermore, this study is expected to foster multidisciplinary
synergies among various institutions and research domains, tran­
scending geographical and disciplinary boundaries. To effectively
decipher the research outcomes and inform future policy-oriented
management strategies, it is essential to engage in a multidisciplinary
approach that amalgamates the insights from fisheries, the food in­
dustry, climate science, and biodiversity research.
2. Scientometric analysis
A systematic and structured electronic search was conducted using
the WOSCC, a leading bibliographic database in scientific literature. The
search was restricted to the time period from January 1977 to December
2022 and was conducted in English. The search string was constructed
using Boolean operators (OR), an asterisk (*) symbol, and quotation
marks (“”) in the Topics search (TS) of the WOSCC database. The asterisk
symbol was used to account for variations in keywords, and the quota­
tion marks were used to ensure that the keywords were interpreted with
exact meanings. The common name of the crab species was obtained
from the FAO factsheet of FishStatJ version 4.02.08 (updated November
2022), and climate change-related elements were adapted from a pre­
vious study (Azra et al., 2022). Due to a large number of species names
for crabs, we did not consider species-specific terms for keyword anal­
ysis. To avoid confusion with unrelated research on the animal “crab,”
we excluded other keywords such as crab algorithm, etc., as described in
the subsequent section.
Crab: (“crab”) NOT (“crabtree”) NOT (“crab tree”) NOT (“crab
apple”) NOT(“Lifelike crab”) NOT (“embedded crab”) NOT (“trans­
lucent crab”) NOT (“Black cuboid crab”) NOT (“crab cros*”) NOT (“crab
waist”) NOT (“CRAB algorithm”) OR (“Antarti* stone crab”) OR
(“Atlant* rock crab”) OR (“Batw* coral crab”) OR (“black stone* crab”)
OR (“blue crab”) OR (“blue king crab”) OR (“blue* swim* crab”) OR
(“brown king crab”) OR (“Callinectes swim* crab”) OR (“channel clin*
crab”) OR (“Charybdis crab”) OR (“Chin* mitten crab”) OR (“Coconut
crab”) OR (“common spider crab”) OR (“Dana swim* crab”) OR (“deep*
sea red crab”) OR (“Dungeness crab”) OR (“edible crab”) OR (“fiddler
crab”) OR (“Gazami crab”) OR (“giant land crab”) OR (“giant stone
crab”) OR (“giant swim* crab”) OR (“globose king crab”) OR (“golden
king crab”) OR (“Green crab”) OR (“Green mud crab”) OR (“hair crab”)
OR (“Harbour spider* crab”) OR (“Henslow* swim* crab”) OR (“Indo*
Pacific swamp crab”) OR (“Jonah crab”) OR (“king crab”) OR (“knobby
swim* crab”) OR (“Maja spider crab*”) OR (“mangrove ghost crab”) OR
(“marine crab*”) OR (“Mediterranean shore crab”) OR (“Mola rock
crab”) OR (“mud crab”) OR (“orange mud crab”) OR (“Pacific rock
crab”) OR (“pelagic red crab”) OR (“port* spider crab”) OR (“Portunus
swim* crab”) OR (“Queen crab”) OR (“red crab”) OR (“red king crab”)
OR (“red snow crab”) OR (“red stone crab”) OR (“red vermillion crab”)
OR (“right* hand* hermit crab*”) OR (“rock crab”) OR (“Scylla spp”) OR
(“shamefaced crab”) OR (“snow crab”) OR (“softshell red crab”) OR
(“Southern king crab”) OR (“Southern spider crab”) OR (“Southwest
Atlantic red crab”) OR (“Spanner crab”) OR (“spider crab”) OR
(“Spinous spider crab”) OR (“stone crab”) OR (“stone king crab”) OR
(“Subantarctic stone crab”) OR (“swim* crab”) OR (“Tanner crab*”) OR
(“Velvet swim* crab”) OR (“Warty crab”) OR (“West Afric* fiddler
crab”) AND climate change: (“climat*”) OR (“climat* chang*”) OR
(“global warm*”) OR (“seasonal* variat*”) OR (“extrem* event*”) OR
(“environment* variab*”) OR (“anthropogenic effect*”) OR (“green­
house effect*”) OR (“sea level ris*”) OR (“erosio*”) OR (“agricult*
runoff”) OR (“weather* variab*”) OR (“weather* extrem*”) OR
(“extreme* climat*”) OR (“environment* impact*”) OR (“environment*
chang*”) OR (“anthropogenic stres*”) OR (“temperature ris*”) OR
(“temperature effect*”) OR (“warm* ocean”) OR (“sea surface* tem­
perat*”) OR (heatwav*) OR (acidific*) OR (hurrican*) OR (“el nino”) OR
(“el-nino”) OR (“la nina”) OR (la-nina) OR (drought*) OR (flood*) OR
(“high precipit*”) OR (“heavy rainfall*”) OR (“CO2 concentrat*”) OR
(“melt* of the glacier*”) OR (“melt* ice*”) OR (“therm* stress*”) OR
(“drought”) OR (“hypoxia”) OR (“harm* alga* bloom*”) OR
(“eutrophication”).
A total of 2834 relevant original research articles were retrieved
from the WOSCC database in January 2023. Non-original research ar­
ticles and articles written in non-English languages were excluded from
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C.S. Thirukanthan et al.
Journal of Sea Research 193 (2023) 102386
been published, indicating that research on crabs under a changing
climate is a topic of growing interest.
the analysis, resulting in a final sample of 2451 articles. The raw data
was downloaded as a text file, renamed as “download_1.txt,” and
continuously renamed with a numeric format until the analysis was
completed. The data was then imported into CiteSpace software version
6.1.R6 Advanced for Windows users in the CiteSpace-6.1.6.msi format.
CiteSpace is a widely used computational tool for visualizing patterns
and trends in scientific literature. The analysis of the data involved four
different types of scientometric methods: (i) frequency analysis and cooccurrence analysis of keywords, (ii) co-citation analysis of cited refer­
ences, (iii) clustering analysis of document titles, and (iv) burst analysis
of articles. The results of these categories, along with the quantitative
analysis of metadata, will be presented in the Results section of the
study.
Additionally, several concepts and metrics commonly used in the
CiteSpace software, such as centrality, burst detection, and sigma, will
be applied. Centrality refers to a key connector between various cita­
tions or references in the field, while burst detection refers to the process
of identifying sudden increases in the frequency of selected references.
This metric is commonly used in trend or pattern analysis studies and
involves the use of various algorithms for burst detection (Tattershall
et al., 2020). Sigma is a useful criterion for identifying potentially
important works that are attracting significant attention in the field
(Chen et al., 2012). Using CiteSpace, the average silhouette value for
each cluster was determined, with a higher value indicating a greater
similarity between cluster members (Yang et al., 2019).
3.2. Main contributor in the field
3.2.1. Global publications
Analysis of global publications on crabs associated with climate
change, as indicated in Fig. 2, revealed that 106 countries or states had
relevant contributions. The United States (933), China (248), Brazil
(182), Canada (160), England (158), Australia (158), United Kingdom
(158), Germany (153), and Argentina (97) are the main contributors in
terms of published articles, accounting for almost 85.2% of the total
publication. These nations’ increasing interest in this field of study un­
derscores the importance of conducting extensive research and devel­
opment in the area. However, the study also identified a deficiency of
research being conducted in the African region, with more than half
(59%) of the literature emanating from the North American region.
Consequently, there is a need to expand research efforts to include
understudied areas to obtain a more comprehensive understanding of
the impact of climate change on crab populations worldwide. Funding
agencies that wish to support the advancement of this field of research
may also consider directing resources towards addressing the research
gaps identified in this study.
3.2.2. Highly cited authors, major funding agencies, highly cited affiliations
and major publication sources
Table 1 displays the highly published output in terms of the number
of publications in the field of crab and climate change. The table pro­
vides information across four categories: author, organization, funding
and journal name. Half of the expertise (i.e., author) and the most
published organizations in the field mostly come from the U.S.A. Hans-O
Portner from Alfred Wegener Institute holds the record for the highest
number of publications with 25, followed by John Spicer with 24 from
Plymouth University and Sven Thatje with 21 from the University of
Southampton.
National Science Foundation (NSF), funded over 233 publications,
accounting for nearly 10% of the total crab and climate change publi­
cations. This was followed by funding from the National Natural Science
3. Trends and evolution of the literature in the field
3.1. Annual publication trends and productive journals
The trend of publications has increased notably in the past few de­
cades, with a sharp increase from 2014 to 2022, indicating that
academia has been giving increasing attention to this topic. According to
the WOSCC database, a total of 2451 publications with 52,020 total
citations and 33,186 total references (Fig. 1) were retrieved for research
on crabs under changing climates. Within the last decade (2012− 2022),
64.9% of the datasets, equivalent to 1591 papers, have been published.
In the previous five years alone (2018–2022), 881 of those papers have
Fig. 1. Total number of publications and citations on research on crabs and climate change from 1977 to 2022.
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C.S. Thirukanthan et al.
Journal of Sea Research 193 (2023) 102386
Fig. 2. Total publications per nation for research on crabs associated with climate change, with the darker brown hues, denote countries that have published a more
substantial number of works, whereas lighter shades indicate a lower volume of publications. (For interpretation of the references to colour in this figure legend, the
reader is referred to the web version of this article.)
Table 1
Top Ten Authors, Funding Bodies, Affiliations, and Journals in the Climate Change and Crab Research Domain from 1977 to 2022: A WOSCC Analysis.
Highly published author
Author name
Top funding bodies
Count
Funding Agencies
Most published institutions
Count
Affiliation
Most publication sources
Count
Sources title
Hans-Otto Pörtner
(Alfred Wegener Institute)
25
National Science Foundation (N.S.F.)
233
National Oceanic and Atmospheric
Administration, U.S.A.
145
John Spicer
(Plymouth University)
24
National Natural Science Foundation
of China (NSFC)
120
University of California, U.S.A.
108
21
National Council for Scientific and
Technological Development
75
Helmholtz Association, Germany
89
Marine Biology
78
20
European Commission
69
77
Estuarine, Coastal and
Shelf Science
67
74
Journal of Crustacean
Biology
63
PlosOne
54
Sven Thatje
(University of
Southampton)
Louis E.
Burnett
(College of Charleston)
William C. Long
(NOAA)
Richard B. Forward
(Duke Marine Lab Road
Beaufort)
Blaine D. Griffen
(Brigham Young
University)
Karen G. Burnett
(College of Charleston)
Oscar O.
Iribarne
(Instituto de
Investigaciones Marinas y
Costeras)
Stefano
Cannicci
(The University of
Florence)
20
19
National Oceanic Atmospheric Admin
(NOAA)
Coordination Foundation for the
Improvement of Higher Education
Personnel (CAPES)
Alfred Wegener Institute Helmholtz
Centre for Polar and Marine Research,
Germany
National Scientific and Technical
Research Council, Argentina
63
55
State University System of Florida, U.
S.A.
66
Marine Ecology Progress
Series
Journal of Experimental
Marine Biology and
Ecology
Count
Comparative
Biochemistry &
Physiology: Part A
Journal of Experimental
Biology
171
123
19
Natural Sciences and Engineering
Research Council of Canada (NSERC)
54
University of Washington, U.S.A.
59
16
São Paulo Research Foundation
49
University of Washington Seattle, U.S.
A.
55
16
U.K. Research Innovation (UKRI)
45
French National Centre for Scientific
Research, France
50
Frontier in Marine Science
35
15
Natural Environment Research
Council (NERC)
43
Fisheries and Oceans Canada
50
Estuaries and Coast
32
Foundation of China (NSFC), the National Council for Scientific and
Technological Development, the São Paulo Research Foundation, the
Coordination Foundation for the Improvement of Higher Education
Personnel (CAPES) from Brazil, the European Commission, National
46
41
Oceanic Atmospheric Admin (NOAA), Natural Sciences and Engineering
Research Council of Canada, UK research Innovation (UKRI), and Nat­
ural Environment Research Council (NERC) from the United Kingdom.
Ellegaard (2018) proposed a novel approach to evaluate academic
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C.S. Thirukanthan et al.
Journal of Sea Research 193 (2023) 102386
disciplines with the highest publication counts were Marine and Fresh­
water Biology, Ecology, Oceanography, Zoology, and Environmental
Sciences. The sizes of the nodes in the disciplines of Fisheries, Biology,
Biochemistry & Molecular Biology, Physiology, Multidisciplinary Sci­
ences, and Biodiversity Conservation indicate strong connections and
their relevance to this research domain. While certain disciplines such as
Toxicology, Geosciences, Evolutionary Biology, Limnology, Water Re­
sources, Genetics & Heredity, and Biotechnology & Applied Sciences
have a smaller number of published papers, their relatively high
betweenness centrality values suggest their importance in interdisci­
plinary research, predicting their potential for future growth in this
field. These results provide a comprehensive picture of the multiple
disciplinary intersections within the field of crabs and climate change,
highlighting the importance of diverse approaches to tackling complex
ecological problems. (See Fig. 5.)
support for a particular field by analyzing the network of research in­
stitutions. In this study, 948 clustering nodes were identified in the
network, and 670 collaborative links were established between them
(Fig. 3). The 2451 documented affiliations that emerged from this
network underscore the significance of the field of inquiry in academia,
highlighting the intensity and depth of the investigation. Fig. 3 show­
cases the top institutions that have contributed significantly to this
collaborative network, with the National Oceanic and Atmospheric
Administration from the U.S.A. emerging as the leading contributor. The
top ten contributing institutions produced a remarkable 775 articles,
representing 31.6% of all crab and climate change publications. This
information offers a rich foundation for future collaborative efforts in
the field, especially for early-career researchers or postgraduate students
seeking research supervision. Additionally, Marine Ecology Progress
Series, published by Inter-Research Science Center in Germany, carried
most of the papers published on crabs associated with climate change,
followed by JEMBE: Journal of Experimental Marine Biology and Ecol­
ogy (Elsevier) and Marine Biology (Springer Verlag). This information
could be useful in reducing article submission time and optimizing
research outputs.
3.4. Highly cited articles
The top ten highly cited articles in the WOSCC database were
analyzed, and their conclusions are presented in Table 2. The results
indicated that numerous elements of climate change, such as ocean
warming, hypoxia, harmful algal blooms, pollution, and acidification,
have a significant impact on crab populations, movements, physiolog­
ical status, adaptation responses, and the harvest industry worldwide
(Pihl et al., 1991; Orbea et al., 2002; Mueter and Litzow, 2008; Lelong
et al., 2012; McCabe et al., 2016). The most active area of study was
found to be harmful algal blooms, with a mean of 34 citations per year.
These blooms are caused by environmental factors such as nutrient
pollution and changes in surface water temperature and can result in the
production of toxins or other harmful compounds that harm aquatic
organisms such as fish and shellfish (Moore et al., 2019; Anderson et al.,
3.3. Important research disciplines
The present study employed CiteSpace’s “Category” node type to
generate a visual map of research disciplinary categories that reflect the
publications addressing issues related to the impact of climate change on
crabs. The centrality of a network, comprising 103 nodes and 243 links,
was calculated following the automated data simplification and merging
using the CiteSpace algorithm (Fig. 4). The resulting distribution map
demonstrated that the study of crabs is a complex research topic that
encompasses numerous fields of study. In descending order, the five
Fig. 3. A collaborative network of institutions researching crabs and climate change from 1977 to 2022. Line thickness represents the connection between in­
stitutions, while the nodes’ colours, ranging from magenta (1977) to yellow (2022), indicate the progression of research over time.
5
C.S. Thirukanthan et al.
Journal of Sea Research 193 (2023) 102386
Fig. 4. Network of linked research disciplines. Line thickness represents the connection between research disciplines, while the colours of the nodes, ranging from
magenta (1977) to yellow (2022), indicate the progression of research over time. The thickness of the lines between the two nodes is proportional to the strength of
the linkages between the two research disciplines. The sizes of the modes are proportional to the frequency of the subject category co-occurrence. (For interpretation
of the references to colour in this figure legend, the reader is referred to the web version of this article.)
2021; Karlson et al., 2021). The study by Farrell et al. (2000) did not find
a link between research on crabs and climate change-related studies, but
this study was still included in the analysis due to its inclusion as a crabrelated study based on the use of keywords such as “flood*” and “crabeating” in the abstract.
The primary cluster (#0), labelled blue king crab by LLR and ocean
acidification by LSI, is the largest, with 145 members and a high
silhouette value of 0.916, suggesting its homogeneity. This cluster is
closely associated with cluster (#1), labelled as ocean acidification by
LLR and fatty acid composition by LSI, with a silhouette value of 0.898.
The leading paper cited in the cluster (#0) and (#1) is by Bednaršek
et al. (2021), a meta-analysis exploring the biological effects of ocean
acidification on decapods. The review highlights the decreased survival,
calcification, growth, development, and abundance of decapods in
response to acidification, with sensitivity increasing in decapod larvae.
Interestingly, the review also revealed that the amplitude of these re­
actions varied significantly among taxonomic groups and multi-species
assemblages, indicating the heterogeneity of species’ responses. Other
factors, such as nutritional states such as amino acid and fatty acid
composition or source population, also appeared to affect the organisms’
responses (Ramaglia et al., 2018). Additionally, the study reported a
trend towards increased sensitivity to acidification when organisms
were exposed to rising seawater temperatures, underscoring the com­
plex interplay between multiple stressors on decapod physiology. These
findings demonstrate the potential of using cluster analysis to identify
key research themes and highlight emerging areas of investigation,
providing valuable insights into the potential impacts of global climate
change on decapod populations.
It is interesting to note that four species of crabs have branched off to
form their own research cluster, highlighting their ecological signifi­
cance and the need for continued research. Of note, the intertidal por­
celain crab Petrolisthes cinctipes is identified as a key species in the third
largest cluster (#2) with a high silhouette value of 0.943, indicating its
relevance in the context of ocean acidification research (LLR). Studies
3.5. Cluster analysis
Cluster analysis is a well-established statistical technique for data
analysis and knowledge discovery that has proven effective in identi­
fying latent semantic themes in textual data (Chen and Morris, 2003;
Zhong et al., 2019). This method enables researchers to divide a large
body of research data into multiple units based on the degree of term
correlation, facilitating the identification of research themes, trends,
and connections within a particular field of study (Olawumi and Chan,
2018; Zhong et al., 2019). The homogeneity of a cluster can be assessed
using the mean silhouette index, which ranges from − 1 to 1. In this
study, we employed the Log-Likelihood (LLR) and Latent Semantic Index
(LSI) algorithms available in CiteSpace to map the link between climate
change and crabs, resulting in the identification of 20 clusters in the
scientometric analysis (Table 3 and Fig. 5). The LLR algorithm assessed
the similarity between text content and topics, while the LSI algorithm
categorized technical terminologies. To conduct this analysis, we used
the literature cited by the citing literature from 1977 to 2022 as the data
source for the scientometric study, carried out using the co-citation
network of cited references. These findings highlight the potential of
cluster analysis in uncovering complex patterns and relationships within
a particular research domain, offering a valuable tool for identifying
emerging research areas and informing future research endeavours.
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C.S. Thirukanthan et al.
Journal of Sea Research 193 (2023) 102386
Fig. 5. The reference co-citation network from 1977 to 2022 for crab and climate change publications. The size of the nodes in the literature co-citation network
reflects how often the paper has been cited, and the network nodes’ colours, which range from dark to light, show how the research has developed from its earliest to
most recent stages. The literature co-citation network was divided into 20 clusters using the network clustering method. Line thickness represents the connection
between research clusters, while the colours of the nodes, ranging from magenta (1977) to yellow (2022), indicate the progression of research clusters over time. (For
interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)
have demonstrated that intertidal zone porcelain crabs, which are highly
adapted to heat, possess limited ability to acclimate their heat tolerance
limits. Consequently, marine intertidal porcelain crabs living in high
intertidal zones may have limited acclimation capacity (Stillman and
Somero, 2000; Stillman and Tagmount, 2009). These findings indicate
that high intertidal porcelain crabs, which already live at the upper
thermal tolerance range, may be negatively impacted by the extreme
temperature increases due to climate change. The fifth largest cluster
(#4), with shore crab Carcinus maenas and environmental hypoxia as the
primary focus, suggests a strong relationship between the species and
the impact of low oxygen conditions. Studies have investigated the shore
crab’s adaptation strategies, such as the ability of adults to feed in
hypoxia, which is closely linked to the thickness of the water-blood
barrier in the gills, allowing them to uptake oxygen even in hypoxic
conditions without the need to switch to anaerobic metabolism (Legeay
and Massabuau, 2000).
Meanwhile, the seventh largest cluster (#6) focuses on the ecology
and behaviour of the mud crab Scylla serrata, an important species
known for its adaptability to various environments due to its ability to
tolerate broad temperatures (16–35 ◦ C) and salinity (1–56 ppt) ranges
(Alberts-Hubatsch et al., 2016). Mud crabs are in high demand due to
the international markets; therefore, studying the species’ ecology and
behaviour is critical (Fazhan et al., 2017). Lastly, the Dungeness crab
Cancer magister, a valuable species, is highlighted in the 10th largest
cluster (#10), with a high silhouette value of 0.97, which indicates its
importance in the field. Since the mid-1800s, Dungeness crab has been
harvested commercially, and it now forms the backbone of one of the
most lucrative fisheries off the mainland West Coast of the United States,
bringing in over $200 million annually (Richerson et al., 2020). These
findings suggest that research on these important crab species is ongoing
and highly relevant in the field of marine ecology, with implications for
conservation and management efforts.
The 4th largest cluster (#3) has 82 members and a silhouette value of
0.992. It is labelled as adaptive significance by LLR and brachyuran crab
settlement by LSI, indicating a focus on the mechanisms of crab adap­
tation during their migration from estuaries to the coastal ocean. Christy
and Morgan’s (1998) study is the major cited article in this cluster,
which describes the adaptation mechanisms of 9 different crab larva
species. The findings from this cluster are closely associated with the 6th
largest cluster, labelled as “wind-driven estuary” by both LLR and LSI.
These two clusters highlight the importance of understanding the ad­
aptations of crab species during migration and the impact of winddriven estuaries on their settlement.
Cluster #8, with a silhouette value of 0.959 and 45 members, focuses
on episodic hypoxia, a common phenomenon in temperate estuaries that
leads to low dissolved oxygen levels (<2 mgL− 1) as a result of increased
anthropogenic nutrient loading. The spatial extent and temporal dura­
tion of seasonal hypoxia, determined by the hydrodynamics of the
estuarine system, can affect the avoidance behaviour of some crab
species, such as Callinectes sapidus (Christy and Morgan, 1998; Bell et al.,
2003). Chronic hypoxia develops slowly over several days and is limited
to the deep basins of estuaries, whereas episodic hypoxia rapidly moves
into near-shore habitats during wind-driven hypoxic upwelling events
(Bell et al., 2003). These findings have significant implications for
7
C.S. Thirukanthan et al.
Journal of Sea Research 193 (2023) 102386
Table 2
Top ten highly cited articles in research on crabs, associated with the climate
change elements based on the latest WOSCC database (until January 2023).
References
Moullac and
Haffner,
2000
Mueter and
Litzow,
2008
Journal
Aquaculture
Ecological
Applications
Total
citation
425
350
Average
per year
17.7
21.8
Pihl et al.,
1991
Marine
Biology
309
9.36
McCabe et al.,
2016
Geophysical
Research
Letters
272
34
Farrell et al.,
2000
Molecular
Ecology
272
11.33
Lelong et al.,
2012
Phycologia
261
21.75
Orbea et al.,
2002
Aquatic
Toxicology
242
11
Cumberlidge
et al., 2009
Biological
Conservation
222
14.8
Bibby et al.,
2007
Biology
Letters
180
10.59
Albecker and
McCoy,
2017
Frontiers in
Zoology
179
25.57
Table 3
Top 10 ranked clusters and labels produced by LSI and LLR on research on crabs
and climate change-related studies.
Main conclusion
Changes in
environmental
variables induced
alteration of immune
response in
Crustacean, and
tolerant to metal
stress
The warming sea in
the Arctic is the
primary cause of the
distribution pattern
in snow crabs over
the past two decades
Periodic hypoxias
affect the oxygen
tolerance, movement
and feeding habit of
blue crab
Harmful algal bloom
negatively impacted
the shellfish harvest
industry in US and
Canada’s west coast
Not included in the
topics1
Harmful algal blooms
contaminated the
shellfish animals,
such as crabs,
worldwide
Long-term
anthropogenic
sources of pollution
impacted the
environment and
further affected the
crab’s biomarker
responses
Global conservation
status of freshwater
crab under changing
climate
Ocean acidification
impact marine life,
especially gastropod,
in the presence of
crab through
chemical cues
Salinity stress in the
coastal area caused
by the flooding, storm
surges and erosion
caused the adaptation
response of crabeating frog
Cluster
Size
Silhouette
Year
Label (Latent
Semantic Indexing)
Label (Log Likelihood
Ratio)
ocean acidification
fatty acid
composition
blue king crab
0
145
0.916
2013
1
127
0.898
2017
2
85
0.943
2008
3
82
0.992
1994
4
68
0.986
1987
5
56
0.962
2002
6
47
0.944
2019
8
9
45
45
0.959
0.981
2003
2013
10
44
0.97
1991
ocean acidification
brachyuran crab
settlement
environmental
hypoxia
wind-driven
estuary
mud crab Scylla
serrata
episodic hypoxia
crab herbivory
crab Cancer
magister
ocean acidification
intertidal porcelain
crab Petrolisthes
cinctipe
adaptive
significance
shore crab Carcinus
maena
wind-driven estuary
mud crab Scylla
serrata
episodic hypoxia
crab herbivory
Dungeness crab
Cancer magister
2003; Linton and Greenaway, 2007; Alberti et al., 2011). For example,
crab’s herbivory causes zonation in marsh plants along the northern
rocky intertidal Chinese salt marsh (He et al., 2015). Massive losses of
high salt marsh grass Spartina patens have been reported in New En­
gland, and investigators have linked this phenomenon to heavy her­
bivory by the crab Sesarma reticulatum (Holdredge et al., 2009; Gedan
and Bertness, 2010). Understanding the complex dynamics of crab
herbivory is essential for managing and conserving marine ecosystems.
3.6. Keywords themes
The principle of keyword co-occurrence analysis is based on counting
the frequency with which a set of keywords appears in the same docu­
ment, clustering these words based on the number of occurrences,
reflecting their affinity, and then analyzing the structural shifts of the
disciplines and topics represented by these words (Radhakrishnan et al.,
2017; Zou et al., 2022).
Keywords, which are the primary words and phrases that describe
the central concepts of articles, can be utilized to monitor the develop­
ment of research areas and domains (Lee and Su, 2010; Wu et al., 2019).
In this study, we utilized CiteSpace to analyze keyword occurrences and
relative frequency, identifying the strongest burst keywords and cooccurrences and interpreting the mapping language. 735 keywords
and 4412 links were extracted from CiteSpace between 1977 and 2022.
The abundance of linked lines (exceeding the number of nodes - 856)
and complex linkages between keywords indicate the broad scope of
research on the impact of climate change on crabs. Analysis of highfrequency keywords revealed eight keyword clusters as research hot­
spots and frontiers in the field of climatic impact on crabs (Fig. 6): #0
snow crab, #1 subsequent recovery, #2 blue crab, #3 temperaturedependent, #4 ocean acidification, #5 fiddler crab, #6 northern Car­
olina estuaries, #7 hypoxia stress, and #8 Domoic acid. These clusters
can be analyzed independently to determine the most applicable de­
scriptors (Table 4).
The major keywords used to discuss “"Snow crab” are population,
pattern, dynamics, community, and ecosystem; “"Subsequent recovery”
are temperature, response, shore crab, hypoxia, and thermal tolerance;
“blue crab” are Callinectes sapidus, abundance, recruitment, Chesapeake
Bay, and habitat; “temperature-dependent” are crab, growth, crustacea,
Brachyura, and behaviour; “"Ocean acidification” are climate change,
impact, life history, survival, and acid-base balance; “fiddler crab” are
Carcinus maena, Decapoda, seasonal variation, evolution, and adaptive
significance; “Northern Carolina Estuaries” are salinity, size, sediment,
population dynamics, and accumulation; “Hypoxia stress” are oxidative
1
Out of the scope of the topic about research on crabs and climate changerelated studies.
understanding the distribution and behaviour of crab populations in
estuarine environments subjected to episodic hypoxia.
Crabs play a significant role in marine ecosystems, and studying their
impact is essential. Crab herbivory, the ninth-largest cluster, is a crucial
component of marine ecology that looks at how crab-eating behaviour
affects seagrass meadows, salt marshes, and other macrophytes. Crab
herbivory is ambivalent to ecosystem health, as it can enhance plant
growth and limit the spread of invasive species but also cause significant
damage to the primary producers (Alberti et al., 2008; Alberti et al.,
2011). Many studies have investigated the feeding behaviour of crabs
and their ecological impact on various plant species (Erickson et al.,
8
C.S. Thirukanthan et al.
Journal of Sea Research 193 (2023) 102386
Fig. 6. Distribution of co-cited clustered keywords in the field of crab and climate change from research articles from 1977 to 2022.
stress, gene expression, dissolved oxygen, mud crab, and Eriocheir
sinensis; “Domoic acid” are mortality, harmful algal bloom, aquaculture,
toxin, and benthic community
Table 5 shows the top 10 keywords with the strongest citation burst.
The detected keywords were “Hypoxia”, “Chesapeake Bay”, “Shore
crab”, “Predation”, “Callinectes sapidus”, “Expression”, “Vertical migra­
tion”, “Sea level rise”, “Chinese mitten crab”, and “Carcinus maenas.”
The keyword “Sea level rise” showed a remarkable increase in popu­
larity and citations from 2017 to 2022, reflecting the growing impor­
tance of climate change-related studies in recent times. Meanwhile,
keywords such as “Chesapeake Bay”, “Shore crab”, “Vertical migration”,
and “Carcinus maenas” had a comparatively prolonged citation burst of
more than ten years.
Among the three species with the highest citation burst were Calli­
nectes sapidus, Chinese mitten crab, and Carcinus maenas. Callinectes
sapidus, also known as the Atlantic blue crab, plays a crucial role in the
structure and function of coastal benthic food webs (Mancinelli et al.,
2017) and is highly valued for its high nutritional content (Sharov et al.,
2003) (Zotti et al., 2016). However, studies have indicated that blue
crab larvae are particularly susceptible to low dissolved oxygen levels
(Tomasetti et al., 2018) and ocean acidification (Giltz and Taylor, 2017),
leading to mortality. The Chinese mitten crab, Eriocheir sinensis, is one of
the most economically significant aquatic animals in the world, partic­
ularly in East Asia, with the highest consumption of all crabs in terms of
economic value (Wang et al., 2016). It is considered invasive due to its
euryoecious nature, wide range of habitats, and high biotic potential
(Chen et al., 2007; Stentiford et al., 2011). Lastly, the shore crab, Car­
cinus maenas, is commercially important in its native region of the
northeast Atlantic coastline (Young and Elliott, 2019) but is considered
one of the world’s 100 worst invaders by the International Union for the
Conservation of Nature (IUCN) (Leignel et al., 2014).
4. Emerging hotspots and general discussions
One aspect of scientometrics is meta-analyses of research trends and
the evaluation of the impact of scientific works. A valuable tool in this
regard is offered by CiteSpace, which enables researchers to conduct
timeline co-citation analysis to visualize the relationships between ref­
erences. This type of analysis provides a graphical representation of the
connections between references and can be utilized to identify patterns
and trends in the scientific literature (Chen, 2022). Through this, re­
searchers can gain a deeper understanding of the evolution of their field,
monitor the impact, and make informed decisions regarding the direc­
tion of future research (Fig. 7).
In recent years, an increase in publications on the impact of climate
change on crabs has been noted. Notably, five clusters have emerged as
the most prominent research areas, namely (#0) “blue king crab”, (#1)
“ocean acidification”, (#6) “mud crab Scylla serrata”, (#9) “crab her­
bivory”, and (#13) “crab larvae”. The significant impact of these topics
on the industry has led to their emergence as hotspots of research in
climate change and capture fisheries. It is important to note that these
clusters are interlinked, suggesting a correlation between the effects of
climate change on different aspects of crab populations. Utilizing
timeline co-citation analysis can provide a deeper understanding of the
relationships between these clusters, identify research gaps, and suggest
potential areas for future study. This approach can prove highly valuable
for researchers seeking to publish cutting-edge research in the field of
climate change and crabs in high-impact reputable journals.
Our meta-analyses have revealed that climate change’s impact on the
distribution and productivity of crabs has received extensive attention in
the literature. Among the shellfish group, crabs are among the highestvalue species and are mainly found in intertidal or bottom ocean areas
(Madduppa et al., 2021). To fully comprehend the future of crabs in the
context of global climate change, we must address several crucial
questions, including (1) the effect of rising sea temperatures and levels
on crab habitat, (2) the impact of ocean acidification, (3) the influence of
9
C.S. Thirukanthan et al.
Journal of Sea Research 193 (2023) 102386
regions globally. The climatic impact drivers recently published by the
IPCC project climate change’s impacts on land, coastal or open ocean
regions worldwide and predict that these factors will impact the earth’s
ecosystem for the next 20 to 30 years (4IPCC, 2014). Therefore, ongoing
evidence-based discussions are urgently needed to assess and analyze
the risk of these factors towards marine animals such as crabs. Such
discussions must be informed by the findings, providing a robust foun­
dation for understanding climate change’s impacts and developing
effective mitigation and adaptation strategies.
Table 4
Most frequent keyword label describing the cluster label for crab and climate
change publications (1977–2022).
Cluster
Cluster label
#0
Snow crab
#1
Subsequent
recovery
#2
Blue crab
#3
Temperature
dependent
#4
Ocean acidification
#5
Fiddler crab
#6
Northern Carolina
Estuaries
#7
Hypoxia stress
#8
Domoic acid
Keyword descriptors
Population, pattern, dynamics, community,
ecosystem, ecology, variability, marine, salt
marsh, management, diversity, bay, predation,
invasive species, community structure,
consequence, climate, biodiversity, coastal,
fishery, spatial distribution
Temperature, response, shore crab, hypoxia,
thermal tolerance, water, adaptation, oxygen
consumption, transport, tolerance, Carcinus
maenas, carbon dioxide, exposure, crayfish,
oxygen, stress, performance, hemolymph, acidbase regulation
Callinectes sapidus, abundance, recruitment,
Chesapeake Bay, habitat, Cancer magister,
Dungeness crab, behavioral response, dispersal,
estuary, vertical migration, Crustacean. Larvae,
Delaware Bay, Gulf of Mexico, California,
megalopa
Crab, growth, crustacea, fish, Brachyura,
behaviour, system, biology, shrimp, hermit crab,
conservation, invertebrate, food, reproduction,
cadmium, juvenile, island, blood, resilience,
environmental change
Climate change, impact, life history, survival, acidbase balance, larval development, embryonic
development, Chasmagnathus granulate, carbon,
estuarine crab, population structure, carbonic acid,
green crab, seawater acidification, Chionoecetes
opilio, CO2, ocean
Carcinus maena, Decapoda, seasonal variation,
evolution, decapod Crustacean, adaptive
significance, Mytilus edulis, biochemical
composition, brachyuran crab, chemical
composition, nitrogen, Uca pugilator, cycle,
thermoregulatory behaviour, sexual selection
Salinity, size, sediment, population dynamics,
accumulation, sea level rise, gulf, forest, heavy
metal, fresh water, ocypodidae, eutrophication,
density, organic matter, Spartina alterniflora
Oxidative stress, gene expression, expression,
dissolved oxygen, mud crab, Eriocheir sinensis,
identification, Chinese mitten crab, Portunus
trituberculatus, protein, metabolism, Litopenaeus
vannamei, immune response, gene, phenotypic
plasticity, Atlantic blue crab, geographic variation
Mortality, harmful algal bloom, aquaculture, toxin,
benthic community, indicator species, sea lion,
sand crab, Pseudo nitzschia Bacillariophyceae,
Antarctic krill
4.1. Climate-induce temperatures changes and sea level rise
The term “rising temperature” in the context of climate change refers
to the observed increase in the global average temperature attributed to
human activities, particularly the emission of greenhouse gases into the
atmosphere. According to the Intergovernmental Panel on Climate
Change (IPCC), the global average temperature has risen by between 0.8
and 1.2 degrees Celsius (2 degrees Fahrenheit) above pre-industrial
levels and is expected to continue to rise in the future (Allen et al.,
2019). This warming is occurring at an unprecedented rate and is pri­
marily due to human activities, such as the burning of fossil fuels that
releases significant amounts of carbon dioxide into the atmosphere
(Saklani and Khurana, 2019). Carbon dioxide and methane are two
gases that act as a blanket, trapping the sun’s heat and contributing to
global warming (Hitz and Smith, 2004; Thomton et al., 2014). The
increased frequency and intensity of heatwaves are a key indicator of
global warming and have implications for human health, agriculture,
and wildlife, particularly aquatic species (Perkins-Kirkpatrick and
Lewis, 2020).
Additionally, the rise in temperature causes sea levels to rise due to
the thermal expansion of saltwater and the melting of glaciers on land,
resulting in shifts in weather patterns, more extreme weather events,
and changes in the distribution of marine organisms (Hansen, 2007;
Allen et al., 2019). Projections for sea-level rise are not optimistic for the
future, with estimates of a rise of 0.26 to 0.77 m (about 8 in. to 2.5 ft) by
the end of the century, according to the latest reports from prestigious
international organizations such as the Intergovernmental Panel on
Climate Change (IPCC) (Esteban et al., 2020). However, local relative
sea-level change can differ from the global mean sea-level rise due to
variables such as ocean currents, wind patterns, coastline geometry, and
others (Johnston, 1993; Slangen et al., 2012).
The impacts of rising water temperatures on crab populations have
been extensively studied and have been found to have significant effects
on both their physiological and behavioral responses (Azra et al., 2020).
Elevated temperatures have been associated with changes in oxygen
consumption (Bartolini et al., 2013), ammonia excretion (da Silva
Vianna et al., 2020), and overall energy expenditure (Guerin and Stickle,
1992; Jungblut et al., 2018) in crabs. The respiratory pigment haemo­
cyanin, found in the blood of crustaceans, has been identified as playing
a role in thermal tolerance adaptation in crabs (Giomi and Pörtner,
2013). Haemocyanin can undergo conformational changes in high
temperature conditions, increasing its ability to bind oxygen and its
thermal stability (Verberk et al., 2016). Elevated temperatures have also
been observed to result in increased expression levels of haemocyanin,
potentially as a means of accommodating the increased oxygen transport
demands from increased metabolic activity (Giomi and Pörtner, 2013).
These mechanisms allow for improved thermal tolerance and survival in
crabs under varying temperature conditions (Valère-Rivet et al., 2017).
In addition to physiological effects, rising water temperatures can
also impact crab population distribution and abundance. For instance,
the fiddler crab (Leptuca uruguayensis) that inhabits vegetated areas
exhibited limitations in thermal adaptation, as evidenced by a decrease
in feeding rate, while Leptuca leptodactyla, which inhabits unvegetated
areas, was able to adjust its metabolic rate in response to temperature
increase (da Silva Vianna et al., 2020). Warmer temperatures have also
been found to shorten the winter dormancy period, increasing the
Table 5
Top 10 keywords with the strongest citation burst for crab and climate change
publications (1977–2022).
Keyword
Year
Strength
Begin
End
hypoxia
Chesapeake Bay
shore crab
predation
Callinectes sapidus
expression
vertical migration
sea level rise
Chinese mitten crab
Carcinus maenas
1991
1991
1991
2004
1993
2014
1994
2017
2015
1991
14.46
11.71
9.51
9.04
8.73
8.72
8.65
7.12
7.05
6.77
1991
1994
1997
2004
2008
2020
1994
2017
2020
1991
2000
2004
2010
2011
2014
2022
2005
2022
2022
2008
extreme weather events, and (4) the emergence of new diseases.
Moreover, climate change elements such as sea surface temperature, sea
level rise, ocean heatwave, and ocean acidity are among multiple cli­
matic impact drivers that are expected to alter in coastal and open ocean
10
C.S. Thirukanthan et al.
Journal of Sea Research 193 (2023) 102386
Fig. 7. Timeline co-citation cluster analysis. Nodes represent reference names, whereas lines represent connections between those references. Larger nodes indicate
higher frequencies of citations. References with strong citation bursts are shown with red rings, whereas references having high centrality are shown with yellow
nodes. The longer the colour line segment in the figure, the larger the time span of citations. (For interpretation of the references to colour in this figure legend, the
reader is referred to the web version of this article.)
likelihood of survival for some crab species, and thus, potentially
increasing population productivity. The survivability of blue crab (Cal­
linectes sapidus) has been projected to increase by at least 20% by 2100,
as a result of shorter winter periods (Glandon et al., 2019). However, the
expression of this increased productivity will depend on the responses of
other food web components and potential changes in fishery manage­
ment policies over the same period.
Rising sea levels have been found to result in the destruction of
intertidal habitats, which can have a negative impact on crab pop­
ulations. Coastal habitats such as marshes, beaches, and estuaries, which
are crucial for the survival and reproduction of many crab species, are
disappearing globally, as sea levels rise in regions such as the Persian
Gulf (Sharifian et al., 2021), India (Khan et al., 2021), and Bangladesh
(Sarwar, 2005). However, vertical accretion has been documented as a
means of adaptation against sea-level rise (Morris et al., 2016).
growth, and survival of marine organisms, as well as the entire marine
ecosystem, from tiny plankton to predatory fish, and potentially impact
human food security and coastal protection (Fabry et al., 2008; Kroeker
et al., 2013). Studies have shown that ocean acidification has negative
impacts on crab larvae’s survival, growth, physiology, and behaviour,
such as decreasing swimming speed and changing settlement behaviour
(Long et al., 2013; Punt et al., 2014; Punt et al., 2016). A stagestructured pre-recruit model was developed to study the impact of
ocean acidification on the survival of red king crabs. The results showed
a decline in expected yields and profits over the next 50 to 100 years
(Punt et al., 2014). Another similar simulation model was developed to
study the impact of ocean acidification on the survival and hatching
rates of southern Tanner crab (Chionoecetes bairdi) larvae, which
demonstrated that juvenile survival had the greatest impact, decreasing
by 20% over a period of 75 years (Punt et al., 2016). The altered
behaviour of marine organisms in high CO2 conditions has been docu­
mented in marine fishes, with changes in homing, predator detection,
feeding, and habitat choice observed (Roggatz et al., 2016). In another
study using shore crabs (Carcinus maenas) as a model system, changes in
pH were found to impair the functionality of peptide signalling cues
(Roggatz et al., 2016). The direction and speed of the vertical
displacement of Florida stone crab larvae (Menippe mercenaria) were also
impacted by changes in pH, with larvae swimming downwards more
quickly at lower pH levels, potentially impeding larval transport and
northward dispersal (Gravinese et al., 2020). The dispersal of crab
populations may be impacted by ocean acidification, leading to alter­
ations in community structure and ecosystem dynamics (Gravinese,
2018; Gravinese et al., 2022). Changes in the availability of prey and
habitat may drive crab migration in search of favorable conditions,
resulting in increased competition for resources (Ross et al., 2011).
One of the most pronounced effects of ocean acidification on crabs is
its impact on their calcification processes. Calcifying organisms regulate
biomineralization through passive and active ion movement into and
4.2. Ocean acidification
Ocean acidification, which is a decrease in the ocean’s pH levels, is
primarily caused by the absorption of carbon dioxide (CO2) from the
atmosphere into the ocean, as a result of human activities such as the
combustion of fossil fuels, deforestation, and other changes in land use
(Doney et al., 2009; Hönisch et al., 2012). Approximately 25–30% of the
anthropogenic CO2 released into the atmosphere is absorbed by the
ocean (Sabine et al., 2004). Since the industrial revolution, the ocean’s
pH has decreased by approximately 0.1 units, due to the rising CO2
levels in the atmosphere (Hoegh-Guldberg et al., 2007). The average pH
of seawater is naturally slightly alkaline, with a range of 7.7 to 8.3
(Kuroyanagi et al., 2009). Ocean acidification is projected to continue to
rise in the future, with a predicted decrease in pH of 0.3 to 0.4 units by
the end of the century, according to the Intergovernmental Panel on
Climate Change (IPCC) (Doney et al., 2009). Ocean acidification can
have a detrimental effect on the physiology, behaviour, reproduction,
11
C.S. Thirukanthan et al.
Journal of Sea Research 193 (2023) 102386
out of a calcification compartment isolated from the ambient seawater
(Weiner and Dove, 2003; Fabry et al., 2008). The microhardness of the
Tanner crab (Chionoecetes bairdi) claw was reduced by 38% under lower
pH conditions, altering the elemental content of the carapace (Dickinson
et al., 2021). A decrease in pinching strength was observed in the shellcrushing crab Acanthocyclus hassleri in lower pH environments, affecting
its ability to feed and create microhabitats for other species living in
mussel beds (Manríquez et al., 2021).
Ocean acidification may also hinder the reproductive success of
crabs, reducing recruitment and population growth. Increased levels of
CO2 in the water have been shown to cause developmental abnormal­
ities in crab larvae, including malformations of the exoskeleton and
slower growth rates (Kroeker et al., 2014). Research has indicated that
the Tanner crab may divert energy away from reproduction and towards
adapting to ocean acidification, due to the high energy costs associated
with this adaptation (Meseck et al., 2016). Reduced pH was found to
negatively impact the development and hatching success of stone crabs,
delaying embryonic development by 24% and hatching success by 28%,
but having no effect on embryo size (Gravinese, 2018).
diseases (Daszak et al., 2000). Disease dynamics may be differently
affected by climate change based on the host-pathogen interaction and
geographical location, and alterations in food web dynamics can lead to
an increase in pathogen exposure and resource competition, further
stressing the animals (Peeler and Taylor, 2011).
Crabs, like many marine organisms, are susceptible to disease out­
breaks due to environmental stressors associated with climate change.
The virulence of several pathogens has been linked to rising tempera­
tures. For instance, Vibrio infections have been associated with climate
change, as the bacterium thrives in warm, saline waters and can infect
numerous marine organisms, including crabs. Over 60% of mortality
from the “milky disease” of mud crab in Southern China was caused by
the pathogenic infection of V. parahaemolyticus, leading to substantial
economic losses (Li et al., 2008; Xie et al., 2014). Studies examining the
correlation between temperature and salinity in Vibriosis outbreaks in
blue crabs from seven coastal marsh locations in Louisiana showed a
higher prevalence of Vibrio infection with warmer temperatures (Sul­
livan and Neigel, 2018). A similar study conducted in Southern China
found that warmer waters enhanced the mortality rate of mud crabs by
promoting the proliferation of virulent pathogens in mud crab (Scylla
paramamosain) aquaculture ponds.
Parasites have specific temperature requirements for their vital
processes, and climate change can directly impact these requirements
(Marcogliese, 2008). However, the local host environment can also play
a role in mitigating the effects of climate change on parasites (Lohmus
and Bjorklund, 2015). Changes in host distribution, immunology,
behaviour, and physiology, all of which are sensitive to temperature, can
ultimately influence the impact of climate change on parasites (Call­
away et al., 2012). These indirect effects may be more significant than
direct effects and highlight the strong link between parasite responses
and host reactions to climate change. Previous studies (Shields, 2019)
reported that the prevalence of parasitic dinoflagellate infestations in
snow crabs and blue crabs was much higher in warmer waters.
In summary, the susceptibility of crabs to diseases increases due to
climate change due to changes in water temperature, pH, food web
dynamics, ocean currents, and population dynamics. These changes can
weaken the immune system of crabs, making them more susceptible to
infection by pathogens and increasing the likelihood of disease
outbreaks.
4.3. Extreme weather conditions
Extreme weather events, such as hurricanes, typhoons, heat waves,
droughts, and floods, are characterized by unusual and severe weather
conditions. They can cause significant impacts on habitats and the life
cycles of species, leading to disruptions in the ecosystem (Stott, 2016).
Climate change, resulting from human activities such as the burning of
fossil fuels and deforestation, can exacerbate the frequency and intensity
of extreme weather events (O’Neill et al., 2017). The populations and
habitats of crabs are among the marine organisms that may be severely
impacted by these changes.
A study conducted on the sandy beaches along the coast of Brazil
showed that the population density of the ghost crab (Ocypode quadrata)
was negatively impacted by the combined effects of storm waves and
urbanization (Machado et al., 2019). In the Mondego estuary in
Portugal, a long-term (15 years) study of the green crab (Carcinus mae­
nas) revealed a correlation between the population dynamics of the crab
and the extreme weather events brought on by climate change. The
study found that the recruitment of the green crab increased during
drought periods, and other studies also showed that environmental
variables such as precipitation rate and ocean currents had an impact on
the populations of green crabs in this region (de Rivera et al., 2011;
Yamada et al., 2015).
Moreover, communities that rely on natural resources for food and
livelihoods are particularly vulnerable to the impacts of climate change.
For example, a study of the frequent tropical cyclones in Bua Province,
Fiji, showed that 52% of mud crab fishers had stopped harvesting crabs
after the cyclone, due to the need to rebuild their homes and difficulties
in traveling to collection sites and markets (Thomas et al., 2019).
5. Conclusion
In conclusion, this scientometric review underscores the pressing
need to address the myriad consequences of climate change on crab
populations and their associated ecosystems. The findings reveal that
rising sea temperatures, ocean acidification, and alterations in precipi­
tation patterns and sea-level rise collectively pose significant challenges
to the well-being of these vital species. The implications of these envi­
ronmental stressors extend to the global crab industry, marine har­
vesters, and other stakeholders who rely on healthy crab populations for
their livelihoods and ecosystem services.
Given the importance of implementing adaptive strategies to coun­
teract the negative impacts of climate change, the present study high­
lights the necessity for continued research and the development of
mitigation measures. By fostering collaborative efforts among re­
searchers, policy-makers, and industry stakeholders, it is possible to
enhance our understanding of the intricate relationships between
climate change and crab populations, and to develop effective solutions
that will ensure the long-term sustainability of these essential marine
resources.
In acknowledging the limitations of this scientometric analysis, it is
crucial to consider the constraints associated with the use of Citespace
software. Specifically, Citespace permits the incorporation of only one
database at a time; for this study, we employed the WOSCC database.
Consequently, this singular database utilization may preclude the
comprehensive representation of the existing literature or the intricate
4.4. The vulnerability of crabs to disease outbreaks in a changing climate
The presence of diseases in invertebrate aquaculture, such as crab
farming, can result in insecurity in food production. Environmental
factors, including temperature and salinity, as well as human-caused
disturbances, are known to affect the transmission of infectious dis­
eases between pathogens and their host organisms (Mydlarz et al., 2006;
Rowley et al., 2014; Maulu et al., 2021). Several studies indicate that the
prevalence, dispersion, and severity of diseases in aquatic ecosystems
have likely increased due to anthropogenic stressors (Lafferty, 2009).
The emergence of infectious diseases is a major concern for both humans
and animals worldwide. The exchange of pathogens with wild pop­
ulations and the intentional transfer of broodstocks for cultivation have
led to the emergence of new diseases in aquaculture species (Peeler and
Taylor, 2011). Climate change, including rising temperatures and ocean
acidification, is a contributing factor to the spread of new infectious
12
C.S. Thirukanthan et al.
Journal of Sea Research 193 (2023) 102386
research networks related to crabs and climate change. The potential
exclusion of pertinent research from other databases could result in an
incomplete portrayal of the current state of knowledge in this field.
Despite this limitation, the insights derived from this review contribute
substantially to the broader understanding of the challenges confronting
crab populations amid a shifting climate. Furthermore, these findings
serve as a solid foundation for future investigations and policy devel­
opment aimed at addressing this critical area of research.
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Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Data availability
The authors do not have permission to share data.
Acknowledgments
The present review was partly supported by the Ministry of Higher
Education Malaysia (MOHE), under the Long Term Research Grant
program (LRGS/1/2020/UMT/01/1; LRGS UMT Vot No. 56040) enti­
tled “Ocean Climate Change: Potential Risk, Impact and Adaptation
Towards Marine and Coastal Ecosystem Services in Malaysia’, with a
sub-project entitled "Charting the Effects of Climate Change and Acidi­
fication through Marine Organism Physiological Responses."
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