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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Vet. Sci.</journal-id>
<journal-title>Frontiers in Veterinary Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Vet. Sci.</abbrev-journal-title>
<issn pub-type="epub">2297-1769</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
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<article-meta>
<article-id pub-id-type="doi">10.3389/fvets.2025.1529827</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Veterinary Science</subject>
<subj-group>
<subject>Brief Research Report</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Helminth absence and invasion success of blackchin tilapia (<italic>Sarotherodon melanotheron</italic>) in Thailand</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name><surname>Suwannarat</surname> <given-names>Nannaphat</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
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<contrib contrib-type="author" corresp="yes">
<name><surname>Ribas</surname> <given-names>Alexis</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
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<contrib contrib-type="author">
<name><surname>Miquel</surname> <given-names>Jordi</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
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<contrib contrib-type="author">
<name><surname>Poonlaphdecha</surname> <given-names>Srisupaph</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
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<aff id="aff1"><sup>1</sup><institution>Program in Fishery Science and Aquatic Resources, Department of Agricultural Technology, King Mongkut&#x2019;s Institute of Technology Ladkrabang, Prince of Chumphon Campus</institution>, <addr-line>Chumphon</addr-line>, <country>Thailand</country></aff>
<aff id="aff2"><sup>2</sup><institution>Parasitology Section, Department of Biology, Healthcare and Environment, Faculty of Pharmacy and Food Science, University of Barcelona</institution>, <addr-line>Barcelona</addr-line>, <country>Spain</country></aff>
<aff id="aff3"><sup>3</sup><institution>Institut de Recerca de la Biodiversitat (IRBio), Universitat de Barcelona</institution>, <addr-line>Barcelona</addr-line>, <country>Spain</country></aff>
<author-notes>
<fn fn-type="edited-by" id="fn0001">
<p>Edited by: Felipe A. Hern&#x00E1;ndez, Universidad Austral de Chile, Chile</p>
</fn>
<fn fn-type="edited-by" id="fn0002">
<p>Reviewed by: Pamela Mu&#x00F1;oz, Universidad Austral de Chile, Chile</p>
<p>Miguel Rubio-Godoy, Instituto de Ecolog&#x00ED;a (INECOL), Mexico</p>
</fn>
<corresp id="c001">&#x002A;Correspondence: Alexis Ribas, <email>aribas@ub.edu</email></corresp>
</author-notes>
<pub-date pub-type="epub">
<day>07</day>
<month>02</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>12</volume>
<elocation-id>1529827</elocation-id>
<history>
<date date-type="received">
<day>18</day>
<month>11</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>23</day>
<month>01</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2025 Suwannarat, Ribas, Miquel and Poonlaphdecha.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Suwannarat, Ribas, Miquel and Poonlaphdecha</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<sec id="sec1">
<title>Introduction</title>
<p>This study investigates the helminth absence in the invasive blackchin tilapia (<italic>Sarotherodon melanotheron</italic>) in Thailand, testing the Enemy Release Hypothesis (ERH). The ERH suggests that invasive species thrive in new habitats due to the lack of natural parasites that control their populations in native environments. The recent introduction of <italic>S. melanotheron</italic> in Thailand has raised concerns about its ecological and economic impacts.</p>
</sec>
<sec id="sec2">
<title>Methods</title>
<p>We surveyed 164 blackchin tilapia from six different locations in Chumphon Province, Thailand, including the sea, estuary, canal, and shrimp farms, examining them for helminths. Fishermen provided data on the first capture dates in the surveyed areas to determine how long the populations have been established.</p>
</sec>
<sec id="sec3">
<title>Results</title>
<p>No helminths were detected in any of the examined fish. The absence of parasites was consistent across all surveyed environments, suggesting a lack of parasitic burden in the population.</p>
</sec>
<sec id="sec4">
<title>Discussion</title>
<p>The absence of helminths may contribute to the successful expansion of <italic>S. melanotheron</italic> in Thailand, supporting the Enemy Release Hypothesis.</p>
</sec>
</abstract>
<kwd-group>
<kwd>helminth</kwd>
<kwd>invasive fish</kwd>
<kwd>enemy release hypothesis</kwd>
<kwd><italic>Sarotherodon melanotheron</italic></kwd>
<kwd>Thailand</kwd>
</kwd-group>
<counts>
<fig-count count="3"/>
<table-count count="0"/>
<equation-count count="0"/>
<ref-count count="36"/>
<page-count count="6"/>
<word-count count="4041"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Parasitology</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec sec-type="intro" id="sec5">
<label>1</label>
<title>Introduction</title>
<p>The cichlid fish <italic>Sarotherodon melanotheron</italic> R&#x00FC;ppell 1852 (Cichlidae) is originally distributed in Africa, where it inhabits lagoons and estuaries from Mauritania to the Democratic Republic of the Congo (<xref ref-type="bibr" rid="ref1">1</xref>). This species has since been introduced to other continents, including America (Suriname and the United States) (<xref ref-type="bibr" rid="ref1">1</xref>). In Asia, it has been intentionally introduced for aquaculture purposes, particularly in countries such as Cambodia and the Philippines (<xref ref-type="bibr" rid="ref2 ref3 ref4">2&#x2013;4</xref>). In its native Africa, its original distribution range has expanded, demonstrating its capacity to colonize new habitats, such as reservoirs (<xref ref-type="bibr" rid="ref5">5</xref>). As a euryhaline species, <italic>S. melanotheron</italic> shows high adaptability to various aquatic environments, being found in coastal and lagoon waters as well as freshwater reservoirs in places like Hawaii (<xref ref-type="bibr" rid="ref6">6</xref>). This includes physiological strategies to protect gills from environmental threats and to maintain protein integrity (<xref ref-type="bibr" rid="ref7">7</xref>). The reproductive traits of the blackchin tilapia also exhibit significant physiological plasticity, allowing it to adapt to different salinities. This adaptability is crucial for its survival and reproductive success in varying environmental conditions (<xref ref-type="bibr" rid="ref8">8</xref>).</p>
<p>Furthermore, the introduction of <italic>S. melanotheron</italic> to new regions for aquaculture has had significant economic and ecological impacts. In Asia, for instance, its use in aquaculture has provided a valuable source of protein and income for local communities. However, the introduction of non-native species can also pose risks to local ecosystems, potentially leading to competition with native species and changes in local biodiversity. Overall, the adaptability and resilience of <italic>S. melanotheron</italic> make it a species of significant interest both ecologically and economically. Its ability to thrive in diverse environments and its use in aquaculture underscore its importance, highlighting the need for careful management to mitigate potential ecological impacts.</p>
<p>Tilapias, including the genera <italic>Oreochromis</italic>, <italic>Sarotherodon,</italic> and <italic>Tilapia</italic>, host a rich variety of parasites, many of which have been translocated with their hosts to newly colonized areas (<xref ref-type="bibr" rid="ref9">9</xref>). The parasites of <italic>Sarotherodon melanotheron</italic> have been well studied in their native distribution areas in Africa, with research conducted in Nigeria (<xref ref-type="bibr" rid="ref10">10</xref>), Benin (<xref ref-type="bibr" rid="ref11">11</xref>), and the Ivory Coast (<xref ref-type="bibr" rid="ref12">12</xref>, <xref ref-type="bibr" rid="ref13">13</xref>). These studies have reported a high diversity of parasites found on the skin, gills, lamellae, opercula, and intestines, including several taxonomic groups: Myxosporidia (<italic>Myxobolus</italic>), Monogenea (<italic>Cichlodigyrus</italic>, <italic>Enterogyrus</italic> and <italic>Scutogyrus</italic>), Digenea (<italic>Clinostomum</italic> and <italic>Euclinostomum</italic>), Cestoda (Gryporhynchidae), Acanthocephala (<italic>Acanthogyrus</italic>), Nematoda (<italic>Eustrongylides</italic>), and Copepoda (<italic>Ergasilus</italic> and <italic>Lernaea</italic>) (<xref ref-type="bibr" rid="ref10 ref11 ref12 ref13 ref14">10&#x2013;14</xref>). Two of the genera reported in Africa, <italic>Clinostomum</italic> and <italic>Eustrongylides</italic>, are considered zoonotic (<xref ref-type="bibr" rid="ref15">15</xref>, <xref ref-type="bibr" rid="ref16">16</xref>). However, data on parasites in introduced populations outside Africa are very limited. For instance, there is no existing data for Thailand, which is our study area. This lack of information poses a challenge for managing the health of introduced populations and impacts of these translocated parasites in new environments. In addition to their ecological significance, wild and farmed tilapias present a potential zoonotic risk, as they can transmit parasitic infections to human populations (<xref ref-type="bibr" rid="ref17">17</xref>).</p>
<p>In Southeast Asia, <italic>S. melanotheron</italic> was first recorded on scientific literature in open lakes in the Philippines in 2011, and by 2015, it was reported in the coastal waters of Manila Bay (<xref ref-type="bibr" rid="ref2">2</xref>). The origin of <italic>S. melanotheron</italic>&#x2019;s introduction in Thailand remains unclear, but it is relatively recent. The first establishment and proliferation were reported in 2002, and by the summer of 2024, the issue was raised in both the Thai parliament and the media due to growing concerns about the negative impacts on aquatic environments and the economy, with estimated losses in the millions of dollars (<xref ref-type="bibr" rid="ref18">18</xref>). In other regions, such as Hawaii, the introduction of this fish has led to adverse ecological impacts (<xref ref-type="bibr" rid="ref6">6</xref>). In Thailand, a recent survey in Samut Songkhram Province found that blackchin tilapia was the dominant species at the Don Hoi Lot Ramsar Site (<xref ref-type="bibr" rid="ref19">19</xref>). In Chumphon Province, our survey area, <italic>S. melanotheron</italic> has spread across all estuaries. Although there is no official report on the introduction of this invasive fish in the area, it appears to be a recent occurrence, as confirmed by the Marine and Coastal Research Center (Central Gulf of Thailand), Mueang Chumphon District, Chumphon Province, through personal communication with one of the authors.</p>
<p>The Enemy Release Hypothesis (ERH) suggests that the abundance or impact of certain non-native species is influenced by the lack of natural enemies in their introduced range compared to their native range, with parasites being one of these natural enemies (<xref ref-type="bibr" rid="ref20">20</xref>). According to Shinn et al. (<xref ref-type="bibr" rid="ref9">9</xref>), the translocation of parasites in tilapia species varies based on their life cycles. Trematodes with more complex life cycles, involving multiple hosts, often result in the absence of the same species of first intermediate host (a mollusk), unlike monogeneans, which do not require intermediate hosts. Parasites significantly influence the invasive process of vertebrates by weakening the immune responses of native species, facilitating disease spread, and disrupting ecological balances. This amplifies the impact of invasive vertebrates on local ecosystems (<xref ref-type="bibr" rid="ref21 ref22 ref23 ref24">21&#x2013;24</xref>). Understanding the role of parasites in these invasions is crucial for assessing their contribution to invasion success, their potential to introduce novel diseases, and their impacts on native species and ecosystems. This knowledge can inform better management and control strategies. The aim of this study was to investigate the helminth community of this fish species to help explain its high invasive success in Thai aquatic systems. According to the ERH, we expect the blackchin tilapia to exhibit lower helminth diversity in the introduced population studied in Thailand compared to its native distribution areas in Africa.</p>
</sec>
<sec sec-type="materials|methods" id="sec6">
<label>2</label>
<title>Materials and methods</title>
<p>The collecting of blackchin tilapia (<xref ref-type="fig" rid="fig1">Figure 1</xref>) was performed from July to August 2024. A total of six points were surveyed in Chumphon Province, covering four different environments: sea, estuary, canal and shrimp farm (<xref ref-type="fig" rid="fig2">Figure 2</xref>). The specific locations and details are as follows: (1) Sea (<italic>n</italic>&#x202F;=&#x202F;30): about 3,000 m from Saphli Pier, Pathio District; coordinates of Saphli Pier: latitude 10.587698, longitude 99.282759. (2) Estuary of canal (<italic>n</italic>&#x202F;=&#x202F;23): Pak Phraek, Sawi District; coordinates: latitude 10.311691, longitude 99.150417. (3) Sea in front of estuary of canal (<italic>n</italic>&#x202F;=&#x202F;20): Thung Kha, Mueang Chumphon District; coordinates: latitude 10.347286, longitude 99.151261. (4) Shrimp farm 1 (<italic>n</italic>&#x202F;=&#x202F;25): Wisai Nuea, Mueang Chumphon District; coordinates: latitude 10.337490, longitude 99.147035. (5) Shrimp farm 2 (<italic>n</italic>&#x202F;=&#x202F;30): Wisai Nuea, Mueang Chumphon District; coordinates: latitude 10.352470, longitude 99.136356. (6) Canal (approximately 3,000 m from the mouth to the sea) (<italic>n</italic>&#x202F;=&#x202F;36): Pak Nam, Mueang Chumphon District; coordinates: latitude 10.435289, longitude 99.23701 (<xref ref-type="fig" rid="fig3">Figure 3</xref>). Local fishermen were advised to contact to the University when blackchin tilapia were caught during daily fishing activities. Upon notification, the fish were collected and transported to the laboratory for immediate dissection. The survey for helminths was conducted on the skin surface, fins, mouth, liver, body cavities and gills using a stereomicroscope. The gastrointestinal tract was examined following the method described by Justine et al. (<xref ref-type="bibr" rid="ref25">25</xref>). All individuals were photographed, weighed, and measured from head to tail fork. To assess the arrival data of the blackchin tilapia, 10 fishermen at each locality were asked, &#x201C;In which year did you first catch blackchin tilapia in this area?&#x201D; If there was no consensus among them, the oldest reported date was considered valid. Salinity was determined using a Master Refractometer (Atago, Japan) in October 2024. This project was reviewed and approved by the Animal Care and Use Committee of King Mongkut&#x2019;s Institute of Technology Ladkrabang, Thailand, under approval number ACUC&#x2013;KMITL-RES/2024/012.</p>
<fig position="float" id="fig1">
<label>Figure 1</label>
<caption>
<p><italic>Sarotherodon melanotheron</italic> from our study area, easily recognized by the distinctive black area on its chin, is readily identified by local fishermen.</p>
</caption>
<graphic xlink:href="fvets-12-1529827-g001.tif"/>
</fig>
<fig position="float" id="fig2">
<label>Figure 2</label>
<caption>
<p><bold>(A)</bold> One of the representative habitats of <italic>Sarotherodon melanotheron</italic> in the canal, <bold>(B)</bold> blackchin tilapia under a boat in the canal.</p>
</caption>
<graphic xlink:href="fvets-12-1529827-g002.tif"/>
</fig>
<fig position="float" id="fig3">
<label>Figure 3</label>
<caption>
<p>Sampled points of <italic>Sarotherodon melanotheron</italic> in Chumphon Province (province shaded in red on the map of Thailand).</p>
</caption>
<graphic xlink:href="fvets-12-1529827-g003.tif"/>
</fig>
</sec>
<sec sec-type="results" id="sec7">
<label>3</label>
<title>Results</title>
<p>A total of 164 blackchin tilapia were examined, ranging in size from 65 to 250&#x202F;mm, with an average length of 158.61&#x202F;mm. The fish weighed between 4.9 and 239.4&#x202F;g, with an average weight of 92.76&#x202F;g. Of these, 80 were female, 80 were male, and 4 were undetermined sex. No helmints were detected in any of the studied organs. The arrival of the blackchin tilapia was estimated as follows: (1) Sea: about 3,000 m from Saphli Pier, Pathio District: 2019; (2) Estuary of canal: Pak Phraek, Sawi District: 2018; (3) Sea in front of estuary of canal: Thung Kha, Mueang Chumphon District: 2018; (4) Shrimp farm 1: Wisai Nuea, Mueang Chumphon District: 2019; (5) Shrimp farm 2: Wisai Nuea, Mueang Chumphon District: 2019; (6) Canal (approximately 3,000 m from the mouth to the sea): Pak Nam, Mueang Chumphon District: 2016. Salinity varied across the different surveyed habitats as follows: (1) 32&#x2013;33&#x202F;ppt; (2) 32&#x2013;33&#x202F;ppt; (3) 32&#x2013;33&#x202F;ppt; (4) 15&#x2013;20&#x202F;ppt; (5) 15&#x2013;20&#x202F;ppt; and (6) 15&#x2013;20&#x202F;ppt.</p>
</sec>
<sec sec-type="discussion" id="sec8">
<label>4</label>
<title>Discussion</title>
<p>To our knowledge, this is the first survey of helminths in an introduced population of <italic>Sarotherodon melanotheron</italic> in Thailand, and it may be extended to Southeast Asia [as reviewed by Acosta-P&#x00E9;rez et al. (<xref ref-type="bibr" rid="ref17">17</xref>)]. Our research tested the Enemy Release Hypothesis (ERH), which suggests that alien species experience greater invasion success in new habitats because they are free from the natural enemies, such as parasites, that regulate their populations in their native environments. This absence of parasites allows invasive species to outcompete native species. Introduced species often originate from relatively small subsets of their native populations, which reduces the likelihood of introducing parasites. Additionally, if the newly colonized area lacks the necessary intermediate hosts, parasites cannot become established (<xref ref-type="bibr" rid="ref20">20</xref>, <xref ref-type="bibr" rid="ref26">26</xref>). In the present study, the absence of parasites is notable, including monogeneans of African origin that have spread to new geographical areas, such as Mexico in <italic>Oreochromis niloticus</italic> (<xref ref-type="bibr" rid="ref27">27</xref>). In China (<xref ref-type="bibr" rid="ref28">28</xref>) also reported nine monogenean species infecting cultured tilapia. Similarly, in Madagascar, a subset of dactylogyridean monogeneans arrived from continental Africa with invasive fish (<xref ref-type="bibr" rid="ref29">29</xref>). In Panama&#x2019;s Canal watershed, <italic>O. niloticus</italic> was infected by a single parasite (monogenean) species from its native range (<xref ref-type="bibr" rid="ref30">30</xref>). Our findings align with Firmat et al. (<xref ref-type="bibr" rid="ref31">31</xref>), who documented the complete loss of gill parasites in the invasive cichlid <italic>O. mossambicus</italic> in New Caledonia. A key factor to consider is the time since these populations were established. In the study by Jim&#x00E9;nez-S&#x00E1;nchez et al. (<xref ref-type="bibr" rid="ref32">32</xref>), the populations had been established for approximately 30&#x202F;years, whereas in our study, the introduction is much more recent, between four and 8 years depending on the locality. The importance of colonization time has been demonstrated in other cases as well. For example, the dataset of 32 British freshwater fish species and their helminth parasites was analyzed, including seven introduced host species. The analysis concluded that helminth species richness is correlated with the time since the fish hosts were introduced to Britain (<xref ref-type="bibr" rid="ref33">33</xref>). Also, it has been shown that the helminth parasite diversity of the Eurasian round goby (<italic>Neogobius melanostomus</italic>) introduced in Canada doubled after 15 years. The authors reached this conclusion by comparing the original study, conducted soon after the establishment of this invasive fish, with a resampling conducted 15 years later using the same sampling design (<xref ref-type="bibr" rid="ref34">34</xref>, <xref ref-type="bibr" rid="ref35">35</xref>, <xref ref-type="bibr" rid="ref36">36</xref>). These findings suggest that the recent introduction of <italic>S. melanotheron</italic> in Thailand may explain the absence of helminths in our survey conducted at the invasion front. The lack of helminths could be attributed to the short time since introduction, which may not have allowed sufficient time for the establishment and spread of parasites. Continued monitoring and further studies are necessary to understand the long-term dynamics of parasite&#x2013;host interactions in these newly introduced populations.</p>
<p>Further studies are necessary to evaluate the potential advantages that cichlids might gain in the invasive areas when infected with a low richness and diversity of helminths (<xref ref-type="bibr" rid="ref32">32</xref>). The absence of parasites in this population could be significant, and examining sympatric fish species will be relevant for further understanding. A limitation of our study is the lack of a survey for metacercariae in muscle tissue using a digestion protocol due to logistical constraints, a gap that future research should address. In addition to the ecological negative effect, fish-borne zoonotic trematodes, including heterophyids and opisthorchiids, are widespread in native and farmed fish in Southeast Asia (<xref ref-type="bibr" rid="ref35">35</xref>). Data on the susceptibility of <italic>S. melanotheron</italic> to these zoonotic trematodes in natural populations would be valuable. An experimental study by Kopolrat et al. (<xref ref-type="bibr" rid="ref35">35</xref>) under controlled conditions examined the susceptibility to <italic>Haplorchis taichui</italic> in five aquaculture fish species from Thailand, showing susceptibility variations in the success of parasitization. Highly susceptible fish species included <italic>Barbonymus gonionotus, Cyprinus carpio</italic>, and <italic>Cirrhinus mrigala</italic>, with values of 93.33 and 100% after exposure to cercariae. In contrast, cichlids such as Nile tilapia (<italic>O. niloticus</italic>) and red tilapia (<italic>Oreochromis aureus</italic> x <italic>Oreochromis mossambicus</italic>) showed no susceptibility. A study in Thailand on translocated Nile tilapia found three zoonotic species: <italic>Stellantchasmus falcatus</italic>, <italic>Haplorchis pumilio</italic>, and <italic>Procerovum</italic> var<italic>ium</italic> (<xref ref-type="bibr" rid="ref36">36</xref>). It should be noted that this fish has been present in Thailand for decades, making it an incomparable model to newly introduced species. Additionally, blackchin tilapia is not commonly valued as a food resource, so the potential transmission to these trematodes to humans should not be a major concern. However, the normalization of this fish in Thai ecosystems could eventually lead to its introduction as a food resource, potentially raising food safety issues.</p>
<p>In conclusion, the absence of helminths at the invasion front of blackchin tilapia may partly explain the successful expansion of this invasive species. This lack of parasitic burden could provide a competitive advantage, facilitating its spread and establishment in new environments.</p>
</sec>
</body>
<back>
<sec sec-type="data-availability" id="sec9">
<title>Data availability statement</title>
<p>The raw data supporting the conclusions of this article will be made available by the authors, without undue reservation.</p>
</sec>
<sec sec-type="ethics-statement" id="sec10">
<title>Ethics statement</title>
<p>The animal study was reviewed and approved by the Animal Care and Use Committee of King Mongkut&#x2019;s Institute of Technology Ladkrabang, Thailand, under approval number ACUC&#x2013;KMITL-RES/2024/012. The study was conducted in accordance with the local legislation and institutional requirements.</p>
</sec>
<sec sec-type="author-contributions" id="sec11">
<title>Author contributions</title>
<p>NS: Investigation, Writing &#x2013; original draft, Funding acquisition. AR: Investigation, Writing &#x2013; original draft, Formal analysis, Methodology, Writing &#x2013; review &#x0026; editing. JM: Investigation, Writing &#x2013; original draft. SP: Formal analysis, Investigation, Methodology, Writing &#x2013; original draft, Writing &#x2013; review &#x0026; editing.</p>
</sec>
<sec sec-type="funding-information" id="sec12">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research, authorship, and/or publication of this article. This project was financially supported by King Mongkut&#x2019;s Institute of Technology Ladkrabang, Thailand, under Research Fund number 2567-02-08-006.</p>
</sec>
<ack>
<p>We are grateful to the staff and students from the Department of Agricultural Technology at King Mongkut&#x2019;s Institute of Technology Ladkrabang, Prince of Chumphon Campus, Chumphon, Thailand, for their logistic help in the sampling collection and laboratory facilities for this project.</p>
</ack>
<sec sec-type="COI-statement" id="sec13">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec sec-type="ai-statement" id="sec14">
<title>Generative AI statement</title>
<p>The author(s) declare that no Gen AI was used in the creation of this manuscript.</p>
</sec>
<sec sec-type="disclaimer" id="sec15">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<ref-list>
<title>References</title>
<ref id="ref1"><label>1.</label><citation citation-type="other"><person-group person-group-type="author"><name><surname>Lal&#x00E8;y&#x00E8;</surname> <given-names>P</given-names></name></person-group>. <article-title>Sarotherodon melanotheron</article-title>. <comment>The IUCN Red List of Threatened Species</comment>, (<year>2020</year>). Available at: <ext-link xlink:href="https://www.iucnredlist.org/species/182038/58328597" ext-link-type="uri">https://www.iucnredlist.org/species/182038/58328597</ext-link> (Accessed October 11, 2024).</citation></ref>
<ref id="ref2"><label>2.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Ordo&#x00F1;ez</surname> <given-names>JF</given-names></name> <name><surname>Asis</surname> <given-names>AMJM</given-names></name> <name><surname>Catacutan</surname> <given-names>BJ</given-names></name> <name><surname>Dela Pena</surname> <given-names>J</given-names></name> <name><surname>Santos</surname> <given-names>MD</given-names></name></person-group>. <article-title>First report on the occurrence of invasive black-chin tilapia <italic>Sarotherodon melanotheron</italic> (Ruppell, 1852) in Manila Bay and of Mayan cichlid <italic>Cichlasoma urophthalmus</italic> (Gunther, 1892) in the Philippines</article-title>. <source>BioInvasions Rec</source>. (<year>2015</year>) <volume>4</volume>:<fpage>115</fpage>&#x2013;<lpage>24</lpage>. doi: <pub-id pub-id-type="doi">10.3391/bir.2015.4.2.08</pub-id></citation></ref>
<ref id="ref3"><label>3.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Cuvin-Aralar</surname> <given-names>MLA</given-names></name></person-group>. <article-title>Introduced aquatic species for inland aquaculture: boon or bane?</article-title> <source>Fish People</source>. (<year>2016</year>) <volume>14</volume>:<fpage>26</fpage>&#x2013;<lpage>39</lpage>.</citation></ref>
<ref id="ref4"><label>4.</label><citation citation-type="other"><person-group person-group-type="author"><name><surname>Froese</surname> <given-names>R</given-names></name> <name><surname>Pauly</surname> <given-names>D</given-names></name></person-group>. <article-title>FishBase</article-title>. <comment>Available at:</comment> <ext-link xlink:href="http://www.fishbase.org" ext-link-type="uri">http://www.fishbase.org</ext-link> (<year>2024</year>).</citation></ref>
<ref id="ref5"><label>5.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>P&#x00E8;l&#x00E8;b&#x00E8;</surname> <given-names>ROE</given-names></name> <name><surname>Imorou Toko</surname> <given-names>I</given-names></name> <name><surname>Verheyen</surname> <given-names>E</given-names></name> <name><surname>Van Steenberge</surname> <given-names>M</given-names></name></person-group>. <article-title>Molecular identification of an invasive <italic>Sarotherodon</italic> species from the atchakpa freshwater reservoir (Ou&#x00E9;m&#x00E9; river basin, Benin) and comparison within <italic>S. melanotheron</italic> using coi markers</article-title>. <source>Diversity</source>. (<year>2021</year>) <volume>13</volume>:<fpage>297</fpage>. doi: <pub-id pub-id-type="doi">10.3390/d13070297</pub-id></citation></ref>
<ref id="ref6"><label>6.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Gaither</surname> <given-names>MR</given-names></name> <name><surname>Aeby</surname> <given-names>G</given-names></name> <name><surname>Vignon</surname> <given-names>M</given-names></name> <name><surname>Meguro</surname> <given-names>Y</given-names></name> <name><surname>Rigby</surname> <given-names>M</given-names></name> <name><surname>Runyon</surname> <given-names>C</given-names></name> <etal/></person-group>. <article-title>An invasive fish and the time-lagged spread of its parasite across the Hawaiian archipelago</article-title>. <source>PLoS One</source>. (<year>2013</year>) <volume>8</volume>:<fpage>e56940</fpage>. doi: <pub-id pub-id-type="doi">10.1371/journal.pone.0056940</pub-id>, PMID: <pub-id pub-id-type="pmid">23468894</pub-id></citation></ref>
<ref id="ref7"><label>7.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Blondeau-Bidet</surname> <given-names>E</given-names></name> <name><surname>Tine</surname> <given-names>M</given-names></name> <name><surname>Gonzalez</surname> <given-names>AA</given-names></name> <name><surname>Guinand</surname> <given-names>B</given-names></name> <name><surname>Lorin-Nebel</surname> <given-names>C</given-names></name></person-group>. <article-title>Coping with salinity extremes: gill transcriptome profiling in the black-chinned tilapia (<italic>Sarotherodon melanotheron</italic>)</article-title>. <source>Sci Total Environ</source>. (<year>2024</year>) <volume>929</volume>:<fpage>172620</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.scitotenv.2024.172620</pub-id>, PMID: <pub-id pub-id-type="pmid">38642748</pub-id></citation></ref>
<ref id="ref8"><label>8.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Dugu&#x00E9;</surname> <given-names>R</given-names></name> <name><surname>Baras</surname> <given-names>E</given-names></name> <name><surname>Gueye</surname> <given-names>M</given-names></name> <name><surname>Avarre</surname> <given-names>JC</given-names></name> <name><surname>Combes</surname> <given-names>Y</given-names></name> <name><surname>Cochet</surname> <given-names>C</given-names></name> <etal/></person-group>. <article-title>Egg production in the euryhaline tilapia <italic>Sarotherodon melanotheron heudelotii</italic>, experimentally maintained in fresh, sea and hypersaline waters</article-title>. <source>Aquat Living Resour</source>. (<year>2014</year>) <volume>27</volume>:<fpage>63</fpage>&#x2013;<lpage>72</lpage>. doi: <pub-id pub-id-type="doi">10.1051/alr/2014008</pub-id></citation></ref>
<ref id="ref9"><label>9.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Shinn</surname> <given-names>AP</given-names></name> <name><surname>Avenant-Oldewage</surname> <given-names>A</given-names></name> <name><surname>Bondad-Reantaso</surname> <given-names>MG</given-names></name> <name><surname>Cruz-Laufer</surname> <given-names>AJ</given-names></name> <name><surname>Garc&#x00ED;a-V&#x00E1;squez</surname> <given-names>A</given-names></name> <name><surname>Hern&#x00E1;ndez-Orts</surname> <given-names>JS</given-names></name> <etal/></person-group>. <article-title>A global review of problematic and pathogenic parasites of farmed tilapia</article-title>. <source>Rev Aquac</source>. (<year>2023</year>) <volume>15</volume>:<fpage>92</fpage>&#x2013;<lpage>153</lpage>. doi: <pub-id pub-id-type="doi">10.1111/raq.12742</pub-id></citation></ref>
<ref id="ref10"><label>10.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Chinedu</surname> <given-names>P</given-names></name> <name><surname>Effiong</surname> <given-names>J</given-names></name> <name><surname>Chukwuemenam</surname> <given-names>F</given-names></name></person-group>. <article-title>Co-parasitism and morphometrics of three Clinostomatids (Digenea: Clinostomatidae) in <italic>Sarotherodon melanotheron</italic> from a tropical freshwater Lake</article-title>. <source>Anim Res Int</source>. (<year>2009</year>) <volume>6</volume>:<fpage>982</fpage>&#x2013;<lpage>6</lpage>. doi: <pub-id pub-id-type="doi">10.4314/ari.v6i2.48129</pub-id></citation></ref>
<ref id="ref11"><label>11.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Bouko</surname> <given-names>B</given-names></name> <name><surname>Tossavi</surname> <given-names>ND</given-names></name> <name><surname>Gand&#x00E9;</surname> <given-names>JJG</given-names></name> <name><surname>Onzo</surname> <given-names>AA</given-names></name> <name><surname>Zannou</surname> <given-names>TB</given-names></name> <name><surname>Ibikounl&#x00E9;</surname> <given-names>M</given-names></name></person-group>. <article-title>Assessing the diversity of metazoan parasites of <italic>Sarotherodon melanotheron</italic> (Cichlidae) from inlands waters in southern Benin (West Africa)</article-title>. <source>Asian J Res Zool</source>. (<year>2023</year>) <volume>6</volume>:<fpage>160</fpage>&#x2013;<lpage>72</lpage>. doi: <pub-id pub-id-type="doi">10.9734/ajriz/2023/v6i4133</pub-id></citation></ref>
<ref id="ref12"><label>12.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Blahoua</surname> <given-names>K</given-names></name> <name><surname>N&#x2019;Douba</surname> <given-names>V</given-names></name> <name><surname>Kone</surname> <given-names>T</given-names></name> <name><surname>Kouassi</surname> <given-names>J</given-names></name></person-group>. <article-title>Variations saisonni&#x00E8;res des indices &#x00E9;pid&#x00E9;miologiques de trois monog&#x00E8;nes parasites de <italic>Sarotherodon melanotheron</italic> (pisces: Cichlidae) dans le lac d&#x2019;Ayam&#x00E9; I (C&#x00F4;te d&#x2019;Ivoire)</article-title>. <source>Sci Nat</source>. (<year>2009</year>) <volume>6</volume>:<fpage>39</fpage>&#x2013;<lpage>47</lpage>. doi: <pub-id pub-id-type="doi">10.4314/scinat.v6i1.48578</pub-id></citation></ref>
<ref id="ref13"><label>13.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Adou</surname> <given-names>YE</given-names></name> <name><surname>Blahoua</surname> <given-names>KG</given-names></name> <name><surname>Yeo</surname> <given-names>K</given-names></name> <name><surname>Konate</surname> <given-names>S</given-names></name> <name><surname>Tiho</surname> <given-names>S</given-names></name></person-group>. <article-title>Parasitofauna of Blackchin tilapia <italic>Sarotherodon melanotheron</italic> (Teleostei: Cichlidae) from Ebrie lagoon, C&#x00F4;te d&#x2019;Ivoire</article-title>. <source>Int J Fish Aquat Stud</source>. (<year>2021</year>) <volume>9</volume>:<fpage>354</fpage>&#x2013;<lpage>60</lpage>. doi: <pub-id pub-id-type="doi">10.22271/fish.2021.v9.i3e.2510</pub-id></citation></ref>
<ref id="ref14"><label>14.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Pariselle</surname> <given-names>A</given-names></name> <name><surname>Lambert</surname> <given-names>A</given-names></name> <name><surname>Euzet</surname> <given-names>L</given-names></name></person-group>. <article-title>A new type of haptor in mesoparasitic monogeneans of the genus <italic>Enterogyrus</italic> Paperna, 1963, with a description of <italic>Enterogyrus foratus</italic> n. sp. and <italic>E. coronatus</italic> n. sp, stomach parasites of cichlids in West Africa</article-title>. <source>Syst Parasitol</source>. (<year>1991</year>) <volume>20</volume>:<fpage>211</fpage>&#x2013;<lpage>20</lpage>. doi: <pub-id pub-id-type="doi">10.1007/BF00009785</pub-id></citation></ref>
<ref id="ref15"><label>15.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Park</surname> <given-names>CW</given-names></name> <name><surname>Kim</surname> <given-names>JS</given-names></name> <name><surname>Joo</surname> <given-names>HS</given-names></name> <name><surname>Kim</surname> <given-names>J</given-names></name></person-group>. <article-title>A human case of <italic>Clinostomum complanatum</italic> infection in Korea</article-title>. <source>Korean J Parasitol</source>. (<year>2009</year>) <volume>47</volume>:<fpage>401</fpage>&#x2013;<lpage>4</lpage>. doi: <pub-id pub-id-type="doi">10.3347/kjp.2009.47.4.401</pub-id>, PMID: <pub-id pub-id-type="pmid">19967090</pub-id></citation></ref>
<ref id="ref16"><label>16.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Robertson</surname> <given-names>LJ</given-names></name></person-group>. <article-title>Parasites in food: from a neglected position to an emerging issue</article-title>. <source>Adv Food Nutr Res</source>. (<year>2018</year>) <volume>86</volume>:<fpage>71</fpage>&#x2013;<lpage>113</lpage>. doi: <pub-id pub-id-type="doi">10.1016/bs.afnr.2018.04.003</pub-id>, PMID: <pub-id pub-id-type="pmid">30077225</pub-id></citation></ref>
<ref id="ref17"><label>17.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Acosta-P&#x00E9;rez</surname> <given-names>V</given-names></name> <name><surname>&#x00C1;ngeles-Hern&#x00E1;ndez</surname> <given-names>J</given-names></name> <name><surname>Vega-S&#x00E1;nchez</surname> <given-names>V</given-names></name> <name><surname>Zepeda-Vel&#x00E1;zquez</surname> <given-names>AP</given-names></name> <name><surname>A&#x00F1;orve-Morga</surname> <given-names>J</given-names></name> <name><surname>Ponce-Noguez</surname> <given-names>JB</given-names></name> <etal/></person-group>. <article-title>Prevalence of parasitic infections with zoonotic potential in tilapia: a systematic review and meta-analysis</article-title>. <source>Animals (Basel)</source>. (<year>2022</year>) <volume>12</volume>:<fpage>2800</fpage>. doi: <pub-id pub-id-type="doi">10.3390/ani12202800</pub-id>, PMID: <pub-id pub-id-type="pmid">36290186</pub-id></citation></ref>
<ref id="ref18"><label>18.</label><citation citation-type="other"><person-group person-group-type="author"><name><surname>Guinto</surname> <given-names>J</given-names></name> <name><surname>Sricham</surname> <given-names>J</given-names></name></person-group>. <article-title>Thailand wages war against &#x201C;alien&#x201D; tilapia fish</article-title>. <comment>Available at:</comment> <ext-link xlink:href="https://www.bbc.com/news/articles/cjjw9e077d8o" ext-link-type="uri">https://www.bbc.com/news/articles/cjjw9e077d8o</ext-link> (<year>2024</year>).</citation></ref>
<ref id="ref19"><label>19.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Siriwattanarat</surname> <given-names>R</given-names></name> <name><surname>Wongroj</surname> <given-names>W</given-names></name> <name><surname>Ngamniyom</surname> <given-names>A</given-names></name> <name><surname>Vilasri</surname> <given-names>V</given-names></name> <name><surname>Chungthanawong</surname> <given-names>S</given-names></name> <name><surname>Punnarak</surname> <given-names>P</given-names></name></person-group>. <article-title>Fish diversity and environmental factors at the Don hoi lot Ramsar site</article-title>. <source>Thailand Biodiversitas J Biol Divers</source>. (<year>2024</year>) <volume>25</volume>:<fpage>1394</fpage>&#x2013;<lpage>403</lpage>. doi: <pub-id pub-id-type="doi">10.13057/biodiv/d250407</pub-id>, PMID: <pub-id pub-id-type="pmid">39771294</pub-id></citation></ref>
<ref id="ref20"><label>20.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Colautti</surname> <given-names>RI</given-names></name> <name><surname>Ricciardi</surname> <given-names>A</given-names></name> <name><surname>Grigorovich</surname> <given-names>IA</given-names></name> <name><surname>MacIsaac</surname> <given-names>HJ</given-names></name></person-group>. <article-title>Is invasion success explained by the enemy release hypothesis?</article-title> <source>Ecol Lett</source>. (<year>2004</year>) <volume>7</volume>:<fpage>721</fpage>&#x2013;<lpage>33</lpage>. doi: <pub-id pub-id-type="doi">10.1111/j.1461-0248.2004.00616.x</pub-id></citation></ref>
<ref id="ref21"><label>21.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Dunn</surname> <given-names>AM</given-names></name> <name><surname>Hatcher</surname> <given-names>MJ</given-names></name></person-group>. <article-title>Parasites and biological invasions: parallels, interactions, and control</article-title>. <source>Trends Parasitol</source>. (<year>2015</year>) <volume>31</volume>:<fpage>189</fpage>&#x2013;<lpage>99</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.pt.2014.12.003</pub-id>, PMID: <pub-id pub-id-type="pmid">25613560</pub-id></citation></ref>
<ref id="ref22"><label>22.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Prenter</surname> <given-names>J</given-names></name> <name><surname>MacNeil</surname> <given-names>C</given-names></name> <name><surname>Dick</surname> <given-names>JTA</given-names></name> <name><surname>Dunn</surname> <given-names>AM</given-names></name></person-group>. <article-title>Roles of parasites in animal invasions</article-title>. <source>Trends Ecol Evol</source>. (<year>2004</year>) <volume>19</volume>:<fpage>385</fpage>&#x2013;<lpage>90</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.tree.2004.05.002</pub-id>, PMID: <pub-id pub-id-type="pmid">16701290</pub-id></citation></ref>
<ref id="ref23"><label>23.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Poulin</surname> <given-names>R</given-names></name></person-group>. <article-title>Invasion ecology meets parasitology: advances and challenges</article-title>. <source>Int J Parasitol Parasites Wildl</source>. (<year>2017</year>) <volume>6</volume>:<fpage>361</fpage>&#x2013;<lpage>3</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.ijppaw.2017.03.006</pub-id>, PMID: <pub-id pub-id-type="pmid">30951572</pub-id></citation></ref>
<ref id="ref24"><label>24.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Brown</surname> <given-names>GP</given-names></name> <name><surname>Shine</surname> <given-names>R</given-names></name> <name><surname>Rollins</surname> <given-names>LA</given-names></name></person-group>. <article-title>A biological invasion modifies the dynamics of a host&#x2013;parasite arms race</article-title>. <source>Proc R Soc B</source>. (<year>2024</year>) <volume>291</volume>:<fpage>20232403</fpage>. doi: <pub-id pub-id-type="doi">10.1098/rspb.2023.2403</pub-id>, PMID: <pub-id pub-id-type="pmid">38351807</pub-id></citation></ref>
<ref id="ref25"><label>25.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Justine</surname> <given-names>JL</given-names></name> <name><surname>Briand</surname> <given-names>MJ</given-names></name> <name><surname>Bray</surname> <given-names>RA</given-names></name></person-group>. <article-title>A quick and simple method, usable in the field, for collecting parasites in suitable condition for both morphological and molecular studies</article-title>. <source>Parasitol Res</source>. (<year>2012</year>) <volume>111</volume>:<fpage>341</fpage>&#x2013;<lpage>51</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s00436-012-2845-6</pub-id>, PMID: <pub-id pub-id-type="pmid">22327319</pub-id></citation></ref>
<ref id="ref26"><label>26.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Torchin</surname> <given-names>ME</given-names></name> <name><surname>Lafferty</surname> <given-names>KD</given-names></name> <name><surname>Dobson</surname> <given-names>AP</given-names></name> <name><surname>McKenzie</surname> <given-names>VJ</given-names></name> <name><surname>Kuris</surname> <given-names>AM</given-names></name></person-group>. <article-title>Introduced species and their missing parasites</article-title>. <source>Nature</source>. (<year>2003</year>) <volume>421</volume>:<fpage>628</fpage>&#x2013;<lpage>30</lpage>. doi: <pub-id pub-id-type="doi">10.1038/nature01346</pub-id>, PMID: <pub-id pub-id-type="pmid">12571595</pub-id></citation></ref>
<ref id="ref27"><label>27.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Pinacho-Pinacho</surname> <given-names>CD</given-names></name> <name><surname>S&#x00E1;nchez-Angeles</surname> <given-names>J</given-names></name> <name><surname>Mart&#x00ED;nez-Ram&#x00ED;rez</surname> <given-names>E</given-names></name> <name><surname>Calixto-Rojas</surname> <given-names>M</given-names></name> <name><surname>Rubio-Godoy</surname> <given-names>M</given-names></name></person-group>. <article-title>Feral parasites: African monogeneans recorded in ferine Nile tilapia in Oaxaca, Mexico</article-title>. <source>Aquaculture</source>. (<year>2023</year>) <volume>577</volume>:<fpage>739974</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.aquaculture.2023.739974</pub-id>, PMID: <pub-id pub-id-type="pmid">39843627</pub-id></citation></ref>
<ref id="ref28"><label>28.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Zhang</surname> <given-names>S</given-names></name> <name><surname>Zhi</surname> <given-names>T</given-names></name> <name><surname>Xu</surname> <given-names>X</given-names></name> <name><surname>Zheng</surname> <given-names>Y</given-names></name> <name><surname>Bilong</surname> <given-names>CFB</given-names></name> <name><surname>Pariselle</surname> <given-names>A</given-names></name> <etal/></person-group>. <article-title>Monogenean fauna of alien tilapias (Cichlidae) in South China</article-title>. <source>Parasite</source>. (<year>2019</year>) <volume>26</volume>:<fpage>4</fpage>. doi: <pub-id pub-id-type="doi">10.1051/parasite/2019003</pub-id>, PMID: <pub-id pub-id-type="pmid">30714897</pub-id></citation></ref>
<ref id="ref29"><label>29.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>&#x0160;imkov&#x00E1;</surname> <given-names>A</given-names></name> <name><surname>&#x0158;ehulkov&#x00E1;</surname> <given-names>E</given-names></name> <name><surname>Rasoloariniaina</surname> <given-names>JR</given-names></name> <name><surname>Jorissen</surname> <given-names>MWP</given-names></name> <name><surname>Scholz</surname> <given-names>T</given-names></name> <name><surname>Falt&#x00FD;nkov&#x00E1;</surname> <given-names>A</given-names></name> <etal/></person-group>. <article-title>Transmission of parasites from introduced tilapias: a new threat to endemic Malagasy ichthyofauna</article-title>. <source>Biol Invasions</source>. (<year>2019</year>) <volume>21</volume>:<fpage>803</fpage>&#x2013;<lpage>19</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s10530-018-1859-0</pub-id></citation></ref>
<ref id="ref30"><label>30.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Roche</surname> <given-names>DG</given-names></name> <name><surname>Leung</surname> <given-names>B</given-names></name> <name><surname>Mendoza Franco</surname> <given-names>EF</given-names></name> <name><surname>Torchin</surname> <given-names>ME</given-names></name></person-group>. <article-title>Higher parasite richness, abundance and impact in native versus introduced cichlid fishes</article-title>. <source>Int J Parasitol</source>. (<year>2010</year>) <volume>40</volume>:<fpage>1525</fpage>&#x2013;<lpage>30</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.ijpara.2010.05.007</pub-id>, PMID: <pub-id pub-id-type="pmid">20600073</pub-id></citation></ref>
<ref id="ref31"><label>31.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Firmat</surname> <given-names>C</given-names></name> <name><surname>Alibert</surname> <given-names>P</given-names></name> <name><surname>Mutin</surname> <given-names>G</given-names></name> <name><surname>Losseau</surname> <given-names>M</given-names></name> <name><surname>Pariselle</surname> <given-names>A</given-names></name> <name><surname>Sasal</surname> <given-names>P</given-names></name></person-group>. <article-title>A case of complete loss of gill parasites in the invasive cichlid <italic>Oreochromis mossambicus</italic></article-title>. <source>Parasitol Res</source>. (<year>2016</year>) <volume>115</volume>:<fpage>3657</fpage>&#x2013;<lpage>61</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s00436-016-5168-1</pub-id>, PMID: <pub-id pub-id-type="pmid">27334451</pub-id></citation></ref>
<ref id="ref32"><label>32.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Jim&#x00E9;nez-S&#x00E1;nchez</surname> <given-names>A</given-names></name> <name><surname>S&#x00E1;nchez-Nava</surname> <given-names>P</given-names></name> <name><surname>de Jes&#x00FA;s</surname> <given-names>R-RF</given-names></name> <name><surname>Flores-Nava</surname> <given-names>B</given-names></name></person-group>. <article-title>Testing the enemy release hypothesis in the invasive fish <italic>Amatitlania nigrofasciata</italic> (Perciformes: Cichlidae) in Mexico</article-title>. <source>Russ J Biol Invasions</source>. (<year>2022</year>) <volume>13</volume>:<fpage>385</fpage>&#x2013;<lpage>91</lpage>. doi: <pub-id pub-id-type="doi">10.1134/S2075111722030079</pub-id></citation></ref>
<ref id="ref33"><label>33.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Gu&#x00E9;gan</surname> <given-names>JF</given-names></name> <name><surname>Kennedy</surname> <given-names>CR</given-names></name></person-group>. <article-title>Maximum local helminth parasite community richness in British freshwater fish: a test of the colonization time hypothesis</article-title>. <source>Parasitology</source>. (<year>1993</year>) <volume>106</volume>:<fpage>91</fpage>&#x2013;<lpage>100</lpage>. doi: <pub-id pub-id-type="doi">10.1017/S0031182000074862</pub-id>, PMID: <pub-id pub-id-type="pmid">8479807</pub-id></citation></ref>
<ref id="ref34"><label>34.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Gendron</surname> <given-names>AD</given-names></name> <name><surname>Marcogliese</surname> <given-names>DJ</given-names></name> <name><surname>Thomas</surname> <given-names>M</given-names></name></person-group>. <article-title>Invasive species are less parasitized than native competitors, but for how long? The case of the round goby in the Great Lakes-St. Lawrence Basin</article-title>. <source>Biol Invasions</source>. (<year>2012</year>) <volume>14</volume>:<fpage>367</fpage>&#x2013;<lpage>84</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s10530-011-0083-y</pub-id></citation></ref>
<ref id="ref35"><label>35.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Kopolrat</surname> <given-names>K</given-names></name> <name><surname>Sithithaworn</surname> <given-names>P</given-names></name> <name><surname>Tesana</surname> <given-names>S</given-names></name> <name><surname>Andrews</surname> <given-names>RH</given-names></name> <name><surname>Petney</surname> <given-names>TN</given-names></name></person-group>. <article-title>Susceptibility, metacercarial burden, and mortality of juvenile silver barb, common carp, mrigal, and tilapia following exposure to Haplorchis taichui</article-title>. <source>Parasitol Res</source>. (<year>2015</year>) <volume>114</volume>:<fpage>433</fpage>&#x2013;<lpage>42</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s00436-015-4326-1</pub-id></citation></ref>
<ref id="ref36"><label>36.</label><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wiriya</surname> <given-names>B</given-names></name> <name><surname>Hedegaard Clausen</surname> <given-names>J</given-names></name> <name><surname>Inpankaew</surname> <given-names>T</given-names></name> <name><surname>Thaenkham</surname> <given-names>U</given-names></name> <name><surname>Jittapalapong</surname> <given-names>S</given-names></name> <name><surname>Satapornvanit</surname> <given-names>K</given-names></name> <etal/></person-group>. <article-title>Fish-borne trematodes in cultured Nile tilapia (Oreochromis niloticus) and wild caught fish from Thailand</article-title>. <source>Vet Parasitol</source>. (<year>2013</year>) <volume>198</volume>:<fpage>230</fpage>&#x2013;<lpage>4</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.vetpar.2013.08.008</pub-id></citation></ref>
</ref-list>
</back>
</article>