International Journal of Food Microbiology 403 (2023) 110340

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International Journal of Food Microbiology

journal homepage: www.elsevier.com/locate/ijfoodmicro

Infection levels of Gnathostomatidae (Nematoda) larvae in commercial

fishes in north-eastern Australian waters and related food safety concerns
Shokoofeh Shamsi a, *, Jaydipbhai Suthar a, Xiaocheng Zhu a, b, Diane P. Barton a
a
School of Agricultural, Environmental and Veterinary Sciences, Gulbali Institute, Charles Sturt University, Wagga Wagga, New South Wales 2650, Australia
b
Wagga Wagga Agricultural Institute, NSW Department of Primary Industries, Wagga Wagga, New South Wales 2650, Australia

A R T I C L E I N F O A B S T R A C T

Keywords: The majority of research on the safety of marine edible fish has primarily focused on anisakid nematodes,
Zoonose neglecting the potential risks posed by other parasites, including those belonging to the family Gnathostoma­
Seafood-borne parasites tidae. In Australia, there have been few reported cases of human infections with gnathostomatid parasites since
Gnathostomiasis
2011. However, due to the absence of a standardized diagnostic test in the country, it is believed that the actual
Seafood safety
number of infections is higher than reported. This study aimed to assess the occurrence and prevalence of in­
fectious gnathostomatid parasites in selected commercial fish species in Australia. A total of 1947 marine fish
from northern Australia, representing 9 families, 16 genera, and 30 species, were examined for gnathostomatid
nematode infections. Overall, 12.3 % of the fish were found to be infected with at least one gnathostomatid larva.
Among the species examined, the yellow-dabbled flounder (Branchypleura novaezeelandiae) exhibited the highest
prevalence (83.3 %; n = 6) and the largest number of gnathostomatid larvae. The identification of the gna­
thostomatid larvae was confirmed as belonging to the genus Echinocephalus based on both morphological
characteristics and sequence data. No significant correlation was observed between the prevalence, mean
abundance, and mean intensity of infection with the length or weight of the examined fish species. Notably,
several of the infected fish species are considered popular choices in the Australian market. Hence, it is
imperative to raise awareness among relevant food safety authorities regarding the occurrence of these parasites.
The findings from this study should be taken into consideration for the revision of current seafood safety pro­
tocols in the country.

1. Introduction parasites are nematodes belonging to the family Gnathostomatidae.

The nutritional value and potential health benefits of fish con­
sumption are widely recognized. Eating fish at least two times per week
has been recommended by health professionals as part of a healthy diet
(Grieger et al., 2013). Consequently, over the past 50 years, annual
global consumption of seafood products per capita has more than
doubled (Guillen et al., 2019). Australia, being a large multicultural
country surrounded by water, offers a diverse range of seafood. How­
ever, research on seafood safety in the country has received less atten­
tion compared to other areas of food safety research (Shamsi, 2019,
2020, 2021; Sumner et al., 2015; Ziarati et al., 2022).
While significant research on the safety of marine edible fish,
particularly has focused on anisakid nematodes (Liu et al., 2015; Shamsi,
2016; Smigic et al., 2016; Sumner et al., 2015), there are several other
parasites that pose risk to human health (Shamsi, 2019). One of these
These nematodes are among the parasites that exhibit an infectious stage
during their third-stage larval development, which occurs in fish and
other marine organisms, including molluscs (Gómez-Valdez et al., 2019;
Morsy et al., 2015; Zhang et al., 2021). Human infection occurs
following the consumption of uncooked or raw infected seafood
(Gómez-Valdez et al., 2019). The resulting disease caused by gnathos­
tomatid nematodes (Gnathostoma spp. and Echinocephalus spp.) is known
as gnathostomiasis, which can be a severe and potentially life-
threatening condition (Herman and Chiodini, 2009; Liu et al., 2020).
Gnathostoma larvae can migrate through various organs and tissues in
the human body, causing tissue damage and triggering inflammatory
responses. The resulting disease, gnathostomiasis, can manifest with a
wide range of symptoms depending on the location of the parasites. The
severity of the infection can vary, ranging from mild skin inflammation
to life-threatening complications affecting the central nervous system,

* Corresponding author.
E-mail address: sshamsi@csu.edu.au (S. Shamsi).

https://doi.org/10.1016/j.ijfoodmicro.2023.110340
Received 6 May 2023; Received in revised form 15 July 2023; Accepted 22 July 2023
Available online 28 July 2023
0168-1605/© 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/by-
nc-nd/4.0/).
S. Shamsi et al.
Fig. 1. Locations of the examined fish including Locker Point (LP), Cape Voltaire (CV), Bynoe Harbour (BH), Outer Darwin Harbour (ODH), Elcho Island (EI), Groote
Eylandt (GE), Arafura Sea East (ASE), Gulf of Carpentaria (GoC), and Townsville (TSV).
eyes, and other vital organs. The diagnosis of gnathostomiasis can be
challenging due to its resemblance to other diseases and the limitations
of available diagnostic tools. Prevention strategies focus on public
health measures, health education, and the promotion of safe food
preparation practices to reduce the risk of infection.
In Australia, adult gnathostomatid (i.e. Echinocephalus spp.) are
found in elasmobranchs (Beveridge, 1987) and the infectious stage of
these parasites has been reported from both fish (Shamsi et al., 2021)
and bivalves (Beveridge, 1987) and also humans (Jeremiah et al., 2011;
Lin and Anstey, 2017), little is known about their prevalence in com­
mercial marine fish and little attention has been given to the risk caused
by them, in country's seafood safety protocols. The aim of the present
study is to determine the occurrence, and prevalence of these parasites
in selected commercial fish in Australia which then will set foundation
for establishing seafood safety protocols.
2. Materials and methods
Fish were collected across a number of projects determining parasite
faunas of marine fish in northern Australia from 1998 until 2017
(Fig. 1). Fish within the families Bothidae, Citharidae, Cynoglossidae,
Lethrinidae, Lutjanidae, Paralichthyidae, Psettodidae, Sciaenidae and
Soleidae were collected as described previously (Barton and Smales,
2015; Barton et al., 2018; Taillebois et al., 2021). Fish were measured
2
International Journal of Food Microbiology 403 (2023) 110340
with Total Length recorded in mm. Independent of the source of the fish,
fish were frozen prior to processing; at the time of processing, all in­
ternal organs encased in their mesenteries were removed and examined
for parasites. A total of 1947 fish belonging to 9 families, 16 genera and
30 species (Table 1) were examined for the infection with gnathosto­
matid nematodes according to standard protocols (Bron et al., 2021;
Fernando et al., 1972).
Parasites collected through the abovementioned projects, were sent
to the Shamsi's Parasitology Laboratory at Charles Sturt University,
Wagga Wagga, New South Wales in 2017. They underwent morpho­
logical and where possible molecular characterisation (Shamsi et al.,
2021) to determine their identification.
The prevalence (P), mean intensity (MI), and mean abundance (MA)
of nematode larvae were calculated as below (Bush et al., 1997):
P = (number of infected fish/total number of examined fish) × 100;
MI = (number of parasites/number of infected fish);
MA = number of parasites/total number of examined fish.
The data was entered into an Excel spreadsheet and transferred into
Stata version 11 (StataCorp.; College Station, United States of America).
Correlation between total length of fish with the number of gnathosto­
matid larvae was performed using a Pearson's coefficient. Fish families
S. Shamsi et al. International Journal of Food Microbiology 403 (2023) 110340

Table 1
Details of infection with gnathostomid parasites found in the examined fish in the present study. The asterisk refers to the Protonibea diacanthus, for which the total
number of fish examined is 289 but only the length of 241 was measured.
Family Fish common Fish scientific name Year/s of Length average No of examined Prevalence Total no. of MI MA
name collection (range) fish (no. of parasites in all
infected fish) fish (range)

Bothidae Weak lefteye Arnoglossus debilis 2002 107 (− ) 1 (0) 0.00 0 (0–0) 0.00 0.00
flounder
Bothidae Waite's left Arnoglossus waitei 1998, 2002 95 (64–128) 31 (2) 6.45 2 (1–1) 1.00 0.03
eye flounder
Bothidae Large scale Engyprosopon grandisquama 1998 100.4 (78–116) 18 (7) 38.89 14 (1–3) 2.00 0.78
flounder
Bothidae Flounder Engyprosopon sp. 1 2002 96.5 (63–137) 31 (0) 0.00 0 (0–0) 0.00 0.00
Bothidae Pennant Grammatobothus pennatus 2002 188.3 11 (0) 0.00 0 (0–0) 0.00 0.00
flounder (129–216)
Bothidae Three spot Grammatobothus 1998, 2002 130.9 (84–216) 39 (2) 5.13 2 (1–1) 1.00 0.05
flounder polyophthalmus
Citharidae Yellow Brachypleura 1998, 2002 123.7 6 (5) 83.33 145 (25–30) 29.00 24.17
–dabbled novaezeelandiae (115–147)
flounder
Cynoglossidae Short headed Cynoglossus kopsi 1998 109.4 (87–138) 18 (1) 5.56 1 (1–1) 1.00 0.06
tongue sole
Cynoglossidae Big –eyed Cynoglossus 1998 104.3 3 (0) 0.00 0 (0–0) 0.00 0.00
tongue –sole macrophthalmus (100− 111)
Cynoglossidae Spotfin Cynoglossus maculipinnus 1998 109.7 (71–145) 23 (0) 0.00 0 (0–0) 0.00 0.00
tongue sole
Cynoglossidae Sole Cynoglossus sp. 1 1998 99.1 (60–122) 8 (0) 0.00 0 (0–0) 0.00 0.00
Cynoglossidae Sole Cynoglossus sp. 2 1998 254 (240–267) 3 (0) 0.00 0 (0–0) 0.00 0.00
Lethrinidae Grass Lethrinus laticaudis 2013–2015 333.2 341 (4) 1.17 7 (1–2) 1.75 0.02
emperor (175–547)
Lutjanidae Saddletail Lutjanus malabaricus 2015–2016 471.3 151 (1) 0.66 1 (1–1) 1.00 0.01
snapper (310–690)
Lutjanidae Golden Lutjanus johnii 2013–2015 418.4 341 (11) 3.23 33 (1–8) 3.00 0.10
snapper (150–765)
Lutjanidae Red Emperor Lutjanus sebae 2015–2016 402.2 120 (1) 0.83 1 (1–1) 1.00 0.01
(185–600)
Lutjanidae Goldband Pristipomoides multidens 2015–2016 392.4 137 (0) 0.00 0 (0–0) 0.00 0.00
snapper (240–600)
Paralichthyidae Peacock Pseudorhombus argus 1998 172.3 (80–210) 28 (1) 3.57 1 (1–1) 1.00 0.01
flounder
Paralichthyidae Large tooth Pseudorhombus arsius 1998 196.2 38 (18) 47.37 42 (1− 10) 2.33 1.11
flounder (150–240)
Paralichthyidae Four twin Pseudorhombus diplospilus 1998 181.1 (90–330) 2 (1) 50.00 25 (25–25) 25.00 12.50
–spot flounder
Paralichthyidae Deep flounder Pseudorhombus elevatus 1998, 2002 125.6 (69–295) 63 (14) 22.22 246 (1–2) 17.57 3.90
Paralichthyidae Small tooth Pseudorhombus jenynsii 1998, 2002 180.6 14 (0) 0.00 0 (0–0) 0.00 0.00
flounder (155–210)
Paralichthyidae Spiny Pseudorhombus spinosus 1998, 2002 179.3 (89–271) 61 (33) 54.10 255 (1− 30) 7.73 4.18
flounder
Psettodidae Indian halibut Psettodes erumei 1998, 2002 223.4 (48–386) 99 (10) 10.10 20 (1–3) 2.00 0.20
Sciaenidae Black-spotted Protonibea diacanthus 2014–2015 805.7 289 (51) 17.65 129 (1–18) 2.53 0.45
croaker (387.0–1300) *
Soleidae Sole Aesopia sp. 1998 119.5 (94–147) 12 (0) 0.00 0 (0–0) 0.00
Soleidae Sole Aseraggodes sp. 1998 88.5 (76–111) 22 (3) 13.64 3 (1–1) 1.00 0.14
Soleidae Tufted sole Dexillus muelleri (previously 1998 174.2 (95–260) 26 (1) 3.85 1 (1–1) 1.00 0.04
known as Dexillichthyes
muelleri)
Soleidae Peacock sole Pardachirus pavoninus 1998 191.1 8 (0) 0.00 0 (0–0) 0.00 0.00
(169–214)
Soleidae Fringefin Zebrias quagga 1998 101.7 (64–134) 3 (0) 0.00 0 (0–0) 0.00 0.00
zebra sole
Total 1947 (166) 12.26 925 (1–30) 5.52 1.59

with >10 infected fish were tested individually and then all infected fish,
irrespective of family, were tested. p values below 0.05 were considered
as significant.
A small portion of representative specimens (up to 3 representative
worms from each fish host and each locality) were used for molecular
analysis. Genomic DNA was isolated using DNeasy Blood and Tissue Kits
(Qiagen, Australia) following the modified protocol of manufacturer's
instruction (Hossen and Shamsi, 2019). We were targeted the internal
transcribed spacer and 18S rRNA regions using multiple primer com­
binations. However, we were only able to obtain 18S rRNA sequence
from a single specimen using primer SSU_F04 (5′-GCTTGTCTCAAA­
GATTAAGCC-3′) and SSU_R26 (5′-CATTCTTGGCAAATGCTTTCG-3′)
(Blaxter et al., 1998). The sequence chromatogram was checked using
SeqMan V 8.1.0 (DNASTAR, Inc.) and primer sequences were removed
for downstream analysis. Our sequence (GenBank accession: OR088504)
was aligned with 18SrRNA sequences of the closely related species in
GenBank. Cosmocercoides pulcher (LC018444), another member of sub­
order Spirurina but Cosmocercidae family, was used as an outgroup.
Pairwise genetic distances were calculated using MEGA 11 (Kumar et al.,
2018), with align gaps ignored in the analysis. The phylogenetic tree was
calculated using MrBayes V 3.2 (Ronquist and Huelsenbeck, 2003) with
K80 + G model as suggested by JmodelTest 2.0 (Darriba et al., 2012).
The analysis was run for 2,000,000 generations until the standard de­
viation of split frequencies lower than 0.01.

3
S. Shamsi et al. International Journal of Food Microbiology 403 (2023) 110340

Fig. 2. Prevalence of gnathostomid larvae found in the present study by fish families. Dark bars indicate fish families, from left to right: Bothidae, Citharidae,
Cynoglossidae, Lethrinidae, Lutjanidae, Paralichthyidae, Psettodidae, Sciaenidae and Soleidae.

Fig. 3. Taxonomically important features for the identification of the third larval stage of the gnathostomatid larvae found in the present study. A) anterior body, B)
cephalic bulb, C) tail, D) cephalic bulb, D) tail.

3. Results prevalence and intensity of gnathostomatid larva infections. The results

revealed that 12.3 % of the examined fish were infected with at least one
In our study, we examined a total of 1947 fish to assess the gnathostomatid larva (as shown in Table 1). Notably, the greatest

4
S. Shamsi et al.
Fig. 4. Phylogenetic tree based on 18S rRNA sequences, showing the placement of our specimen (isolate number 1053-1, i.e. GenBank accession number OR088504).
Cosmocercoides pulcher, a species from the same suborder of Spirurina but a different family of Cosmocercidae, was used as an outgroup. Bayesian posterior prob­
abilities values over 90 % are indicated above the branches.
prevalence of infection, reaching 83.3 % (n = 6) was observed in Yellow-
dabbled flounder (Brachypleura novaezeelandiae), a species belonging to
the family Citharidae (Fig. 2). This was followed by Pseudorhombus spp.,
which belongs to the family Paralichthydae (as illustrated in Fig. 2 and
summarized in Table 1). Regarding the relationship between fish char­
acteristics and gnathostomatid infection, our analysis did not reveal any
significant correlation between the intensity of infection with the length
of the fish species within individual families. However, there was an
overall significant relationship between fish length and intensity of
infection (r = 0.203, df = 171, p < 0.05) if fish family was not taken into
consideration.
To accurately identify the gnathostomatid larvae, we conducted
morphological examinations and attempted to utilize sequence data.
Based on the morphology (Fig. 3), the gnathostomatid larvae in this
study were identified as belonging to the genus Echinocephalus. How­
ever, it is important to note that due to limitations in sample conditions,
genetic analysis was not feasible for the majority of the specimens. There
was a considerable time gap, ranging from six to 25 years, between the
collection date, preservation, and DNA extraction. Consequently, we
were only able to obtain a PCR product from one specimen. While our
obtained sequence (GenBank accession: OR088504) shares a high
5
International Journal of Food Microbiology 403 (2023) 110340
similarity with Echinocephalus species (GenBank accession: KY972321,
KY911549, and KF934729), with a pairwise genetic distance of <3.23
%, an alignment gap of 385 bp was observed between our sequence and
some other sequences belonging to Echinocephalus spp. in the GenBank.
This raise concerns that the sequence obtained in this study may
represent a pseudogene of the 18S rRNA gene. Although a phylogenetic
analysis has been performed (Fig. 4), it is crucial to exercise caution
when interpreting these results until more sequences become available
(Table 2). Additional sequences would provide further insights into the
phylogeny of Echinocephalus spp.
4. Discussion
It is generally known that larvae of Echinocephalus spp. inhabit in a
variety of fish, sea urchin and molluscs which serve as their intermediate
or paratenic hosts (Moravec, 2007), However, the adult stage of Echi­
nocephalus typically infects marine and freshwater stingrays (Hoberg
et al., 1998). Elasmobranchs, primarily rays but also some sharks, are
currently recognized as the only definitive hosts (Moravec and Justine,
2021). Currently, the genus Echinocephalus contains 12 recognized valid
species (Moravec and Justine, 2021). The larval stage of Echinocephalus
S. Shamsi et al. International Journal of Food Microbiology 403 (2023) 110340

Table 2
Pairwise genetic distance among 18S rRNA sequences of our sample and closely related species from the family of Gnathostomatidae, shown as p-difference (%, below
the diagonal) and number of differences (above the diagonal). Asterisk denotes specimens obtained in the present study.
1 2 3 4 5 6 7 8 9 10 11 12 13 14

1 OR088504* 24 24 19 106 112 112 112 114 112 111 110 110 106
2 KY972321 Echinocephalus 3.23 % 0 5 82 88 88 88 94 92 91 90 90 85
sp.
3 KY911549 Echinocephalus 3.23 % 0.00 % 5 82 88 88 88 94 92 91 90 90 85
sp.
4 JF934729 E. overstreeti 2.56 % 0.67 % 0.67 % 87 93 93 93 97 95 94 93 93 87
5 JF934728 Tanqua tiara 14.27 11.04 11.04 11.71 19 19 19 65 66 65 64 64 57
% % % %
6 OL830839 Tanqua sp. 15.07 11.84 11.84 12.52 2.56 0 0 70 71 70 73 73 66
% % % % %
7 OL830840 Tanqua sp. 15.07 11.84 11.84 12.52 2.56 0.00 0 70 71 70 73 73 66
% % % % % %
8 OL830841 Tanqua sp. 15.07 11.84 11.84 12.52 2.56 0.00 0.00 70 71 70 73 73 66
% % % % % % %
9 Z96946 Gnathostoma 15.34 12.65 12.65 13.06 8.75 9.42 9.42 9.42 3 4 7 7 79
binucleatum % % % % % % % %
10 Z96947 G. neoprocyonis 15.07 12.38 12.38 12.79 8.88 9.56 9.56 9.56 0.40 % 3 6 6 78
% % % % % % % %
11 Z96948 G. turgidum 14.94 12.25 12.25 12.65 8.75 9.42 9.42 9.42 0.54 % 0.40 % 5 5 77
% % % % % % % %
12 MN752132 Gnathostoma 14.80 12.11 12.11 12.52 8.61 9.83 9.83 9.83 0.94 % 0.81 % 0.67 % 0 76
sp. % % % % % % % %
13 MN752133 Gnathostoma 14.80 12.11 12.11 12.52 8.61 9.83 9.83 9.83 0.94 % 0.81 % 0.67 % 0.00 % 76
sp. % % % % % % % %
14 LC018444 14.27 11.44 11.44 11.71 7.67 8.88 8.88 8.88 10.63 10.50 10.36 10.23 10.23
Cosmocercoides pulcher % % % % % % % % % % % % %

spp. is the infective stage of the parasite, capable of infecting other
animals that consume infected intermediate and paratenic hosts.
Importantly, in their recent study, Moravec and Justine (2021) under­
scored the significant challenge in accurately identifying larval stages to
the species level. As a result, the current understanding of larval stages
often remains at broader taxonomic levels, such as family or genus,
rather than reaching the species-specific resolution. This limitation
hampers comprehensive assessments of species distributions, life cycle
dynamics, ecological interactions in various ecosystems, and also
assessing their zoonotic significance and pathogenicity.
Invasive migrations within different mammalian hosts have been
documented in Echinocephalus sinensis, as highlighted in studies by Ko
et al. (1975) and Ko (1976). Similar to other gnathostomatids, such as
Gnathostoma spinigerum, larvae of Echinocephalus are known to present a
potential health risk to humans (Hoberg et al., 1998). This poses a sig­
nificant concern, considering that many intermediate hosts, which are
frequently consumed raw or undercooked, are popular seafood choices
(Diaz, 2015; Leroy et al., 2017).
In Australia, Echinocephalus larvae have been reported in numerous
fish and invertebrate hosts (See Table 1 in Karagiorgis et al., 2022;
Shamsi et al., 2021); however, their life cycle is not completely under­
stood, primarily due to the lack of reliable specific larval identification
methods. Human cases of gnathostomiasis have been reported in
Australia (Jeremiah et al., 2011; Lin and Anstey, 2017); Nonetheless,
Shamsi and Sheorey (2018) suggest that these cases are highly likely
caused by Echinocephalus larvae, as the diagnosis relied on tests devel­
oped overseas and there have been no reports of Gnathostoma in
Australia to date.
The infective stage of the parasite that causes gnathostomiasis is
small and usually follows ingestion of only one infective, L3 larvae
(Miyazaki, 1960). Larval penetration of the intestines and portal venous
migration to the liver leads to symptoms including epigastric pain,
nausea, and vomiting beginning shortly thereafter, lasting 2 to 3 weeks
(Diaz, 2015). The prodrome is often dismissed as food poisoning, mis­
diagnosed as acute appendicitis or mesenteric adenitis, and usually
underreported (Diaz, 2015). Although observing larvae in skin, urine,
sputum, cerebrospinal fluid, and ocular specimens can lead to a defini­
tive diagnosis identification of the responsible species by microscopic
analysis of sections of larvae (Akahane et al., 1998), most reported cases
are based on serological tests that never been tested for cross reaction
with Echinocephalus spp. and most diagnoses of gnathostomiasis are
indeed presumptive.
In our study fish belonging to the families Citharidae, Para­
lichthyidae and Bothidae were among the most infected fish, with up to
30 parasites in one individual fish among the families Citharidae and
Paralichthyidae. It is important to note that caution should be exercised
when interpreting the estimated prevalence at the species level, as the
number of examined fish was relatively low for some species. For
example, only 1, 2, and 3 individuals of the following fish species were
examined: Arnoglossus debilis, Pseudorhombus diplospilus, and Zebrias
quagga, respectively.
Many of the fish species found to be infected with Echinocephalus
larvae in our study are considered as marketable edible fish (Froese
et al., 2019), and some are caught incidentally as bycatch due to their
small size. However, it is important to recognize that the parasite can be
transmitted through food chain to other edible fish (Anderson, 2000).
Therefore, it is crucial for the relevant food safety authorities to be
aware of the occurrence of these parasites and take appropriate mea­
sures to ensure the safety of the seafood consumed by the public. It
should be noted that in the present study, we did not examine the flesh of
the fish. Thus, future studies are necessary to specifically examine the
most commonly consumed edible parts of the fish. This will provide a
more comprehensive understanding of the potential risks associated
with the presence of these parasites and allow for targeted food safety
measures to be implemented.
Interestingly, our study did not find a statistically significant corre­
lation between the size of the fish and the number of parasites present in
them. However, it was observed that infected fish were primarily
benthic species that feed on bottom-living animals. This suggests that
the infection with Echinocephalus sp. is likely influenced by the biology
and feeding habits of the intermediate hosts, in this case, the fish. It is
plausible that the benthic nature of these fish exposes them to a higher
risk of acquiring the parasite from their prey or their immediate
environment.
By understanding the factors influencing the infection patterns of
Echinocephalus in fish populations, we can gain insights into the

6
S. Shamsi et al.
transmission dynamics of the parasite and potentially develop strategies
to mitigate its spread. Moreover, raising awareness among both the
fishing industry and consumers about the potential risks associated with
consuming raw or undercooked seafood can contribute to the prevention
of gnathostomiasis and ensure the overall safety of seafood products.
Declaration of competing interest
All authors declare there is no conflict of interest to report.
Data availability
Data will be made available on request.
Acknowledgements
Fish were collected as part of various projects as listed in the relevant
publications and the authors are grateful to all vessel crews and fishers
involved.
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