Marine fish parasites of Vietnam: a comprehensive review and updated list of species, hosts, and zoogeographical distribution.

With a long coastline stretching from tropical to subtropical climate zones, and an immense exclusive economic zone with over 4000 islands, the Vietnamese marine waters support a rich and biodiverse parasite fauna. Although the first parasitological record was in 1898, systematic studies of the parasite fauna have increased during the last 50 years. This comprehensive review covers the current state of knowledge of marine fish parasites in Vietnam and lists 498 species found in 225 fish species, and their geographical distribution. In addition, 251 marine parasite species have newly been added to the already known fauna of 247 species since 2006 (more than two-fold increase). The most speciose group was the Digenea, which accounted for 43% of the total parasite species biodiversity, followed by Monogenea (23.5%), Crustacea (11.6%), Nematoda, and Acanthocephala (8.0% each). The shallow and muddy Gulf of Tonkin showed a rich parasite fauna, accounting for 66.3% of the whole marine parasite fauna of Vietnam, with Digenea accounting for 51% of the regional total parasite richness, followed by Monogenea (27%), Acanthocephala (8.8%), and Nematoda (5.8%). Only a few species belonged to Hirudinea, Myxozoa, and Cestoda, suggesting that these taxa may be understudied. Despite significant progress in studies of marine fish parasites in Vietnam since 2006, only about 12% and 13% of the total fish species have been examined for parasites in the whole country and the Gulf of Tonkin, respectively. © V.T. Truong et al., published by EDP Sciences, 2022.

Introduction

Vietnam has a long coastline of more than 3260 km and an immense exclusive economic zone (one million km2), comprising the coastlines of more than 4000 islands and thousands of square kilometers of coral reefs [120, 131]. The Vietnamese Sea stretches from tropical to subtropical climate zones (from the Namzu Islands, 9°40′ N, 104°22′ E, in the Gulf of Thailand to Daochao Island, 20°50′ N, 107°20′ E, in the Gulf of Tonkin), opening to the Pacific Ocean and being known for its biodiverse marine ecosystems supporting a rich fauna and flora, including fish parasites. Although Billet [26] recorded the first parasite, Distomum hypselobagri (Trematoda), systematic studies of the parasite fauna did not begin until the 1960s (e.g., [69, 71, 82, 97–99], etc.), and the first review was published only 16 years ago [19]. Recently, studies using both molecular [11–13, 16, 20, 24, 25, 36] and morphological methods [12, 13, 20, 24, 92] have been conducted in order to elucidate the systematics of marine parasites in Vietnam.

According to Palm and Bray [103], marine fish parasites are important in ecosystems. In open Hawaiian waters (Pacific Ocean), the authors discovered an average of 2.2 parasite species per marine fish species. In contrast, Klimpel et al. [66] discovered 1.5 metazoan parasite species per deep-sea fish species. According to Palm [102], each fish species in the world harbors up to 3–4 metazoan parasite species on average. In this sense, Arthur and Te [19] reported that the estimated number of parasite species per marine fish species in Vietnamese waters was as high as 3.0. There are currently 1876 species of marine fish reported in Vietnam’s marine waters [45]. Despite this, only 247 parasite species infecting 82 marine fish species were documented by 2006, accounting for about 4.4% of the total fish community [19]. Such data demonstrate that the number of parasites known to science is insufficient to provide credible estimates of the local parasite community.

Because of the importance of marine parasites in fisheries, aquaculture, and human health, more research has been conducted recently, focusing on commercially important fish (e.g., [88, 90, 132, 135]). Furthermore, because parasites can be used as a tool for monitoring climate change and environmental health [86, 102, 129], studies regarding these organisms are essential. However, no systematic update pertaining to parasites in marine fish from Vietnam has been published since 2006, and since that date, much has changed in the knowledge of such organisms. However, Poulin [123] asserts that, due to the absence of an adequate method for determining parasite diversity, species richness remains the most straightforward and thus most relevant indicator of diversity. Therefore, the present study is aimed at: (1) shedding more light on the marine parasite richness in Vietnam by listing, correcting, and arranging the latest information, including the host and geographical distribution of parasites of marine fish; and (2) providing a current state of knowledge on parasite research in the Vietnamese marine waters, with particular emphasis on the Gulf of Tonkin (GOT).

Methods

Study sites and data collection

Vietnam is a Southeast Asian country divided into three distinct geographical and climatic regions: North, Central, and South Vietnam. The aquaculture industry is vital to the Vietnamese economy, especially brackish and marine fish farming along the country’s coastline, particularly in the north [67].

A total of 137 references published between 1901 and 2022 were retrieved to gather knowledge on marine fish parasites and their occurrence in cultured and wild marine fish in Vietnam, focusing on the latest studies. This list was also based on information from the checklists of parasites of fishes in Vietnam [19], and it was expanded to provide a parasite-host list organized on a taxonomic basis, including information about each parasite species, sites of infection, the known geographical distribution in Vietnam, and references. More recently, reviews of individual species of parasites, including morphology, host-parasite distribution, zoogeography, and occasionally molecular analysis have become available; for instance, Amin et al. [2, 8, 14, 15] on Rhadinorhynchus trachuri Harada, 1935, Neoechinorhynchus johnii Yamaguti, 1939, Sclerocollum neorubrimaris Amin, Heckmann, Ha, 2018, respectively and Serrasentis sagittifer (Linton, 1889) Linton, 1932. The animal science databases (https://www.cabi.org/animalscience/disease-health/), MEDLINE, PubMed, Scopus, Web of Science, and Google Scholar were searched using the keywords: parasite, fish, marine, and Vietnam.

In addition, we included our most recent results from the Gulf of Tonkin, which included 40 Acanthopagrus latus, 1 Acanthocybium solandri, 7 Neotrygon kuhlii, 47 Protonibea diacanthus, 37 Trachinotus blochii, 7 Telatrygon zugei, and 2 Thynnus sp. (from 2014 to 2015). The fish were collected in the local fish markets in Ha Long (20°94′96″ N, 107°08′23″ E) and Cat Ba (20°72′75″ N, 107°04′67″ E) between 2014 and 2015, kept on ice and transferred to the Fisheries College’s laboratory in Bac Ninh (close to Ha Noi). The parasite examination was performed using a Nikon SMZ-1 stereomicroscope and the standard procedures described by Palm [101] and Palm and Bray [103]. For taxonomic identification, permanent mounts were prepared following standard methods for Digenea, Monogenea, Nematoda, Cestoda, Acanthocephala, and Crustacea (Isopoda, Copepoda) according to Riemann [124], Palm [101], Arai et al. [18], Dojiri and Ho [42] and Paladini et al. [100]. The method to identify trichodinid ciliates was modified after Klein [65]. Parasites were identified using an Olympus BX53F DIC microscope based on taxonomic keys and original descriptions. The monogenean identification followed Chisholm and Whittington [39], Dang et al. [41], and Hendrix [51]. The identification of crustaceans was based on Kabata and Margolis [59], Shultz [127], and Dojiri and Ho [42]. The digenean identification followed Bray et al. [32], Gibson et al. [47], Jones et al. [57], and Bray and Justine [30]. The identification of cestodes was based on Khalil [63] and Palm [101]. Identification of nematodes was based on Anderson et al. [17] and Gibbons [46]. The protozoan identification followed Bruno et al. [33], and Lom and Dyková [78]. Identification of acanthocephalans was based on Amin [1] and Arai et al. [18]. Other groups of invertebrates retrieved from the literature were identified using methods developed by different authors.

Furthermore, numerous scientific names for recorded hosts from the literature have been amended and corrected following the FishBase database (https://www.fishbase.in/search.php; 2021). The scientific names of existing parasites were carefully checked and corrected using various reliable sources, such as the World Register of Marine Species database (http://www.marinespecies.org/index.php), Ocean Biodiversity Information System (https://obis.org/), and Integrated Taxonomic Information System (https://www.itis.gov/).

Vietnam’s 3260 km of coastline is home to 25 coastal provinces and three cities by the sea. The following abbreviations are used to denote administrative and oceanic divisions where the parasites have been reported (Table 1)

Table 1

The abbreviated names of the municipalities, provinces, and sea or ocean parts where samples were collected.

Municipalities/provinces/sea or ocean parts		Municipalities/provinces/sea or ocean parts
Full names	Abbreviation	Full names	Abbreviation
Bac Lieu province	BL	Nam Dinh province	ND
Binh Thuan province	BT	Nghe An province	NA
Gulf of Tonkin	GOT	Quang Binh province	QB
Gulf of Thailand	GOTh	Quang Ninh province	QN
Hai Phong city	HP	South China Sea	SCS
Ha Long Bay, Quang Ninh	HL-QN	Thanh Hoa province	TH
Thua Thien Hue province	Hue	Ba ria-Vung tau province	VT
Khanh Hoa province	KH
Kien Giang province	KG	The Gulf of Tonkin (GOT) includes HP, NA, ND, QB, QN, and TH

Data analysis

The retrieved literature data, such as the reference, study period, study location, site of infection, fish host, taxa, and parasite species studied, were entered into an excel spreadsheet. This analysis only included parasites identified at the species level. In the list, the parasite taxon levels, i.e., subfamilies, genera, and species, were organized alphabetically. For the entire country of Vietnam and the Gulf of Tonkin, parasite species were classified into eight main taxa: Myxozoa (My), Ciliophora (Ci), Monogenea (Mo), Digenea (D), Cestoda (C), Nematoda (N), Acanthocephala (A), and Crustacea (Cr).

In order to have an insight into parasite richness, the total number of parasite species for Vietnam and the GOT was used to calculate the parasite species ratio per fish host. It is determined by dividing the total number of parasite species parasitizing fishes by the number of infected hosts. Microsoft Excel software was used to perform calculations and descriptive analysis of the collected data.

Results and discussion

Parasites in marine fish of Vietnam

Table 2 contains information about the parasite and host taxa, the site of infection, geographical localities, and literature sources.

Table 2

List of marine fish parasites in Vietnamese waters.

Parasite		Taxa			Hosts		Site of infection	Locality/ies		References
Phylum: Myzozoa Cavalier-Smith & Chao, 2004
Class: Conoidasida Levine, 1988
Order: Eucoccidiorida Léger, 1911
Family: Eimeriidae Minchin, 1903
Goussia sp.		My			Lates calcarifer		Sw	KH		[122]
Phylum: Cnidaria Hatschek, 1888
Class: Myxozoa Grassé, 1970
Subclass: Myxosporea Bütschli, 1881
Order: Bivavulida Shulman, 1959
Family: Ceratomyxidea Doflein, 1899
Ceratomyxa binhthuanensis Chinh, Ha, Doanh, Violettta		My			Epinephelus fasciatus		Gb	KH		[38]
Ceratomyxa sp.		My			Epinephelus bleekeri, E. coioides, E. malabaricus, E. tauvina, Lates calcarifer		Gb	KH, HP, ND		[122, 133, 135]
Family: Meglitschiidae Kovaleva, 1988
Meglitschia insolita (Meglitsch, 1960)		My			Epinephelus coioides		Gb	KH		[135]
Family: Myxobolidae Thélohan, 1892
Henneguya cerebralis Pronin, 1972		My			Lates calcarifer		Gi	KH		[122]
Henneguya lata Nguyen, Chinh, Ngo, Van Tuc, Itoh, Yoshinaga, Shirakashi & Doanh, 2021 in [Chinh NN et al. (2021)]1		My			Acanthopagrus latus		Gi	QN		[37]
Henneguya sp.		My			Acanthopagrus latus		Gi, Sw	QN		Present study
Myxobolus sp.		My			Epinephelus bleekeri, E. coioides, E. tauvina		K	KH		[135]
Phylum: Ciliophora Doflein, 1901
Class: Oligohymenophorea, Stein 1859
Subclass: Peritrichia Stein, 1859
Order: Mobilida Kahl, 1933
Family: Trichodinidae Claus, 1951
Trichodina japonica Imai, Miyazaki & Nomura, 1991		Ci			Epinephelus bleekeri, E. coioides, E. malabaricus, E. tauvina		Gi	KH		[140]
Trichodina rostrata Kulemina, 19681		Ci			Lates calcarifer		Gi, Sk	KH		[122]
Trichodina sp.		Ci			Epinephelus bruneus, E. tauvina, E. sexfasciatus, E. coioides, Lutjanus erythropterus, Rachycentron canadum, Sciaenops ocellatus		Gi, Sk	HP, QN, NA, KH, VT		[34, 121, 133, 140]
Order: Sessilida Stein 1933
Family: Epistylididae Kahl, 1935
Apiosoma sp.		Ci			Epinephelus bleekeri, E. coioides, E. malabaricus, E. tauvina		Sk	KH		[135]
Family: Scyphiddidae Kahl, 1935
Ambiphrya sp.		Ci			Epinephelus bleekeri, E. coioides, E. malabaricus, Lates calcarifer		Sk	KH		[140]
Class: Phyllopharyngea de Puytorac, Batisse, Bohatier, Corliss, Deroux, Didier, Dragesco, Fryd-Versavel, Grain, Grollere, Horasse, Mode, Laval, Roque, Savoie & Tuffrau, 1974
Order: Chlamydodontida Deroux, 1976
Family: Chilodinellidae Deroux, 1970
Chilodonella sp.		Ci			Epinephelus bleekeri, E. coioides		Sk	KH		[135]
Order: Dysteriida Deroux, 1970
Family: Hartmannulidae Poche, 1913
Brooklynella hostilis Lom & Nigrelli, 1970		Ci			Epinephelus bleekeri, E. bruneus, E. sexfasciatus, E. tauvina*		Gi, Sk	QN, KH		[34, 140]
Class: Prostomatea Schewiakoff, 1896
Order: Prorodontida Corliss, 1974
Family: Holophryidae Perty, 1852
Cryptocaryon irritans Brown, 1951		Ci			Epinephelus bleekeri, E. coioides, E. malabaricus, Lates calcarifer, Sciaenops ocellatus		Gi, Sk	KH, HP, ND		[133, 140]
Phylum: Platyhelminthes Minot, 1876
Class: Monogenea Van Beneden, 1858
Subclass: Monopisthocotylea Odhner, 1912
Order: Capsalidea Lebedev, 1988
Family: Capsalidae Baird, 1853
Allobenedenia epinepheli (Bychowsky & Nagibina, 1967) Yang, Kritsky & Sun, 2004	Mo			Epinephelus coioides			Gi	KH		[140]
Allobenedenia yamagutii (Egorova, 1994) Yang, Kritsky & Sun, 2004	Mo			Hyporthodus nigritus (Holbrook, 1855)			Gi, Sk	KH, QN		[88, 140]
Benedenia epinepheli (Yamaguti, 1937) Meserve, 1938	Mo			Epinephelus bleekeri, E. bruneus, E. coioides, E. malabaricus, E. sexfasciatus, E. tauvina*, Lutjanus argentimaculatus			Sk, Fi	KH, QN		[34, 135]
Benedenia sp.	Mo			Epinephelus bruneus, E. coioides, E. sexfasciatus, E. tauvina *, Lutjanus erythropterus, Rachycentron canadum			Sk	HP, ND, QN, KH		[34, 133, 140]
Capsala affinis (Mamaev, 1968) Chisholm & Whittington, 2007	Mo			Auxis thazard, Euthynnus affinis			Gi	SCS		[80]
Capsala notosinense (Mamaev, 1968) Chisholm & Whittington, 2007	Mo			Euthynnus affinis			Gi	SCS		[80]
Capsala paucispinosa (Mamaev, 1968) Chisholm & Whittington, 2007	Mo			Euthynnus affinis, Thunnus thynnus			Gi, Sk	SCS		[80]
Capsala sp.	Mo			Thunnus thynnus			Gi	SCS		[80]
Encotyllabe spari Yamaguti, 1934	Mo			Gymnocranius griseus, Plectorhinchus sp.			Gi	GOT		[81]
Megalocotyle lutiani Lebedev, 1970	Mo			Lutjanus lutjanus			Gi	GOT		[73]
Neobenedenia melleni (MacCallum, 1927) Yamaguti, 1963	Mo			Epinephelus bleekeri, E. coioides, E. malabaricus, Lates calcarifer, Lutjanus argentimaculatus			Sk, Fi	KH		[53, 140]
Neobenedenia sp.	Mo			Rachycentron canadum			Sk	KH		[53]
Sessilorbis limopharynx Mamaev, 1970	Mo			Platax orbicularis			Gi	GOT		[81]
Sprostoniella multitestis Bychowsky & Nagibina, 1967	Mo			Platax orbicularis			Gi	GOT		[81]
Trilobiodiscus lutiani Bychowsky & Nagibina, 1967	Mo			Lutjanus argentimaculatus			–	GOT		[43]
Trochopus antigoniae Egorova & Korotaeva, 1990		Mo			Antigonia rubescens		Gi	SCS		[44]
Order: Dactylogyridea Bychowsky, 1937
Family: Ancyrocephalidae Bychowsky, 1937
Ancyrocephalus macrogaster Yamaguti, 1953		Mo			Gerres filamentosus		Gi	GOT		[81]
Ancyrocephalus parspinicirrus Mamaev, 1970		Mo			Drepane punctata		Gi	GOT		[81]
Ancyrocephalus unicirrus Tripathi, 1959		Mo			Pomadasys argenteus		Gi	GOT		[81]
Haliotrema bilobatum (Yamaguti, 1953) Bychowsky & Nagibina, 1970		Mo			Drepane longimana, D. punctata		Gi	GOT		[81]
Haliotrema cromileptis Young, 1968		Mo			Epinephelus bleekeri, E. coioides		Gi	KH		[41, 135]
Haliotrema epinepheli Yamaguti, 1968		Mo			Epinephelus bleekeri, E. coioides		Gi	KH		[41, 135]
Haliotrema geminatohamula Bychowsky & Nagibina, 1971		Mo			Leiognathus nuchalis		Gi	QN		[88]
Haliotrema spinicirrus (Yamaguti, 1953) Bychowsky & Nagibina, 1970		Mo			Decapterus maruadsi, Drepane punctata		Gi	QN, GOT		[81, 88]
Haliotrema sp.		Mo			Acanthopagrus latus, Epinephelus bruneus, E. bleekeri, E. coioides, E. sexfasciatus, E. tauvina*		Gi	GOT, KH		[34, 41, 135] Present study
Ligophorus hamulosus Pan & Zhang in Pan 1999		Mo			Moolgarda engeli, M. seheli		Gi	QN		[88]
Metahaliotrema kulkarnii Venkatanarasaiah, 1981		Mo			Scatophagus argus		Gi	GOT		[68]
Metahaliotrema mizellei Venkatanarasaiah, 1981		Mo			Scatophagus argus		Gi	QN, GOTh		[68, 88]
Metahaliotrema scatophagi Yamaguti, 1953		Mo			Scatophagus argus		Gi	QN		[68]
Metahaliotrema simile Kritsky, Nguyen, Ha & Heckman, 2016		Mo			Scatophagus argus		Gi	QN		[68]
Metahaliotrema yamagutii Mizelle & Price, 1964		Mo			Scatophagus argus		Gi	GOTh		[68]
Metahaliotrema ypsilocleithrum Kritsky, Nguyen, Ha & Heckman, 2016		Mo			Scatophagus argus		Gi	QN, GOTh		[68]
Murraytrema pricei Bychowsky & Nagibina, 1977		Mo			Nibea albiflora		Gi	QN		[88]
Paradiplectanotrema trachuri (Kovaleva, 1970) Gerasev, Gayevskaya & Kovaleva, 1987		Mo			Argyrosomus japonicus, Johnius carouna		St	QN		[88]
Platycephalotrema platycephali (Yin & Sproston, 1948) Kritsky & Nitta, 2019		Mo			Platycephalus indicus		Gi	QN		[88]
Protogyrodactylus scapulasser (Mamaev, 1970) Gusev, 1973		Mo			Gerres filamentosus		Gi	GOT		[81]
Family: Diplectanidae Monticelli, 1903
Calydiscoides flexuosus (Yamaguti, 1953) Young, 1969		Mo			Nemipterus Japonicus		Gi	QN		[88]
Acleotrema nenue (Yamaguti, 1968) Dominques & Boeger, 2007		Mo			Lates calcarifer		Gi, Sk	KH		[122]
Diplectanum sp.		Mo			Plectorhinchus cinctus		Gi	GOT		[81]
Laticola latesi (Tripathi, 1959) Yang, Kritsky, Sun, Zhang, Shi & Agrawal, 2006		Mo			Lates calcarifer		Gi, Sk	KH		[122]
Laticola paralatesi (Nagibina, 1976) Yang, Kritsky, Sun, Zhang, Shi & Agrawal, 2006		Mo			Lates calcarifer		Gi, Sk	KH		[122]
Murraytrema pricei Bychowsky & Nagibina, 1977		Mo			Nibea albiflora		Gi	QN		[88]
Paradiplectanum blairense (Gupta & Khanna, 1974) Domingues & Boeger, 2008		Mo			Sillago japonica, S. sihama		Gi	QN		[88]
Pseudorhabdosynochus coioidesis Bu, Leong, Wong, Woo & Foo, 1999		Mo			Epinephelus bleekeri, E. coioides, E. malabaricus, E. tauvina		Gi	KH		[135]
Pseudorhabdosynochus cupatus (Young, 1969) Kritsky & Beverley-Burton, 1986		Mo			Epinephelus sexfasciatus		Gi	QN		[34]
Pseudorhabdosynochus epinepheli (Yamaguti, 1938) Kritsky & Beverley-Burton, 1986		Mo			Epinephelus bleekeri, E. bruneus, E. coioides, E. sexfasciatus, E. tauvina		Gi	QN, KH		[34, 135]
Pseudorhabdosynochus grouperi (Bu, Leong, Wong, Woo & Foo, 1999) Wu, Li, Zhu & Xie, 2005		Mo			Epinephelus bleekeri, E. coioides, E. malabaricus, E. tauvina, Lates calcarifer		Gi	KH		[140]
Pseudorhabdosynochus lantauensis (Beverley-Burton & Suriano, 1981) Kritsky & Beverley-Burton, 1986		Mo			Epinephelus bleekeri, E. coioides, E. malabaricus		Gi	KH		[135]
Pseudorhabdosynochus serrani (Yamaguti, 1953) Kritsky & Beverley-Burton, 1986		Mo			Epinephelus bleekeri, E. coioides, E. malabaricus, E. tauvina		Gi	KH		[135]
Pseudorhabdosynochus summanoides Yang, Gibson & Zeng, 2005		Mo			Epinephelus bleekeri, E. coioides, E. malabaricus		Gi	KH		[135]
Pseudorhabdosynochus summanae (Young, 1969) Kritsky & Beverley-Burton, 1986		Mo			Epinephelus coioides, E. bleekeri		Gi	KH		[135]
Pseudorhabdosynochus sp.		Mo			Epinephelus coioides		Gi	QN, HP, ND		[133]
Order: Gyrodactylidea Price, 1943
Family: Tetraonchoididae Bychowsky, 1951
Paratetraonchoides inermis Bychowsky, Gussev & Nagibina, 1965		Mo			Ichthyscopus lebeck		Gi	GOT		[35]
Pavlovskioides ichthyoscopi Bychowsky, Gussev & Nagibina, 1965		Mo			Ichthyscopus lebeck		Gi	GOT		[35]
Pavlovskioides litoralis Bychowsky, Gussev & Nagibina, 1965		Mo			Trachinocephalus myops		Gi	GOT		[35]
Pseudotetraonchoides bleekeriae Bychowsky, Gussev & Nagibina, 1965		Mo			Bleekeria viridianguilla		Gi	GOT		[35]
Subclass: Polyopisthocotylea Odhner, 1912
Order: Mazocraeidea Bychowsky, 1957
Family: Allodiscocotylidae Tripathi, 1959
Allodiscocotyla chorinemi Yamaguti, 1953		Mo			Caranx sp., Decapterus sp., Scomberoides lysan		Gi	GOT, SCS		[73, 76, 118]
Allodiscocotyla diacanthi Unnithan, 1962		Mo			Acanthopagrus pacificus, Decapterus sp.		Gi	QN, GOT, SCS		[73, 76, 93, 118]
Camopia rachycentri Lebedev, 1970		Mo			Rachycentron canadum		Gi	GOT		[74]
Metacamopia chorinemi (Yamaguti, 1953) Lebedev, 1984		Mo			Scomberoides lysan, Selar crumenophthalmus		Gi	GOT, SCS		[73, 76, 118]
Metacamopia lebedevi Nguyen, Nguyen, Ha, Ngoc, Ngoc, Le, Tatonova & Greiman, 2020		Mo			Acanthopagrus pacificus		Gi	QN		[93]
Family: Axinidae Monticelli, 1903
Allopseudaxine macrova (Unnithan, 1957) Yamaguti, 1963		Mo			Auxis thazard, Caranx sp.		Gi	SCS		[72, 80]
Loxuroides pricei Nguyen, Nguyen, Bui & Ha, 2016		Mo			Cypselurus naresii		Gi	GOT, QB		[92]
Unnithanaxine naresii Nguyen, Nguyen, Bui & Ha, 2016		Mo			Cypselurus naresii		Gi	GOT, Hue		[92]
Family: Bychowskicotylidae Lebedev, 1969
Bychowskicotyle plectorhynchi Lebedev, 1969		Mo			Plectorhinchus cinctus		Gi	GOT		[74]
Yamaguticotyla jucunda (Lebedev, 1972) Lebedev, 1984		Mo			Pomadasyidae gen. sp.		Gi	GOT		[74]
Family: Diclidophoridae Cerfontaine, 1895
Osphyobothrus bychowskyi Khoche & Chauhan, 1969		Mo			Saurida tumbil		Gi	GOT, SCS		[83, 114]
Family: Gastrocotylidae Price, 1943
Churavera triangula (Mamaev, 1967) Lebedev, 1986		Mo			Auxis thazard		Gi	SCS		[74, 80]
Engraulicola thrissocles (Tripathi, 1959) Lebedev, 1971		Mo			Anchoviella sp., Thryssa mystax		Gi	GOT		[74]
Gastrocotyle indica Subhapradha, 1951		Mo			Alepes djedaba			SCS		[74]
Gastrocotyle kurra Unnithan, 1968		Mo			Decapterus sp.		Gi	GOT		[74]
Gastrocotyle trachuri Van Beneden & Hesse, 1863		Mo			Decapterus sp., Selar crumenophthalmus, Trachurus trachurus		Gi	GOT, SCS		[73, 76, 118]
Gastrocotyle sp.		Mo			Alectis indicus, Decapterus sp.		Gi	GOT, SCS		[73, 76, 118]
Pseudaxine bivaginalis Dillon & Hargis, 1965		Mo			Selaroides leptolepis		Gi	GOT		[72]
Pseudaxine trachuri Parona & Perugia, 1890		Mo			Carangoides malabaricus, Caranx sp., Decapterus sp.		Gi	GOT, SCS		[73, 76, 118]
Pseudaxine bychowskyi (Lebedev, 1977) Bouguerche, Tazerouti, Gey & Justine, 2020		Mo			Alepes djedaba, Alepes kleinii		Gi	SCS		[74]
Pseudaxine caballeroi (Lebedev, 1977) Bouguerche, Tazerouti, Gey & Justine, 2020		Mo			Alepes djedaba, Alepes kleinii		Gi	SCS		[74]
Pseudaxine sp.		Mo			Alectis indicus, Auxis thazard, Caranx sp., D. muroadsi, Decapterus sp.		Gi	QN, GOT, SCS		[73, 76, 88, 118]
Pseudaxine vietnamensis (Lebedev, Parukhin & Roitman, 1970) Bouguerche, Tazerouti, Gey & Justine, 2020		Mo			Caranx sp., Decapterus sp., Selar crumenophthalmus, Selaroides leptolepis, Seriola dumerili		Gi	QN, GOT, SCS		[73, 76, 88, 118]
Sibitrema poonui Yamaguti, 1966		Mo			Auxis thazard, Euthynnus affinis, T. thynnus		Gi	SCS		[74, 80]
Family: Gotocotylidae Yamaguti, 1963
Cathucotyle cathuaui Lebedev, 1968		Mo			Scomberomorus commerson, S. guttatus.		Gi	GOT, SCS		[72, 73]
Gotocotyla acanthura (Parona & Perugia, 1896) Meserve, 1938		Mo			Scomberomorus guttatus		Gi	GOT		[73]
Gotocotyla bivaginalis (Ramalingam, 1961) Rohde, 1976		Mo			Scomberomorus commerson		Gi	GOT		[74]
Gotocotyla laticauda Lebedev, 1970		Mo			Scomberomorus commerson		Gi	GOT		[73]
Family: Heteraxinidae Unnithan, 1957
Bicotyle perpolita Lebedev, 1968		Mo			Pampus argenteus, Parastromateus niger		Gi	GOT, SCS		[69, 81]
Heteraxine heterocerca (Goto, 1894) Yamaguti, 1938		Mo			Caranx sp., Selar crumenophthalmus		Gi, Phc	GOT, SCS		[69, 81]
Kannaphallus virilis Unnithan, 1957		Mo			Alectis indicus, Carangoides malabaricus,		Gi	SCS, GOT		[73, 76, 118]
Karvolicola ruber Nguyen, Nguyen, Tatonova, 2020		Mo			Otolithes ruber		Gi	QB		[94]
Karvolicola tuyeti Nguyen, Nguyen, Tatonova, 2020		Mo			Nibea albiflora		Gi	QB		[94]
Lethrinaxine parva Mamaev, 1970		Mo			Gymnocranius griseus		Gi	GOT		[81]
Monaxine formionis Unnithan, 1957		Mo			Parastromateus niger		Gi	GOT		[81]
Family: Heteromicrocotylidae Unnithan, 1961
Heterapta chorinemi (Tripathi, 1956) Unnithan, 1961		Mo			Acanthopagrus pacificus		Gi	QN		[93]
Heteromicrocotyla carangis Yamaguti, 1953		Mo			Carangoides malabaricus, Seriola sp., Seriolina nigrofasciata, Carangidae gen. sp.		Gi	GOT, SCS		[69, 73, 76, 118]
Heteromicrocotyla polyorchis Unnithan, 1961		Mo			Carangidae gen. sp.		Gi	GOT, SCS		[73, 76, 118]
Heteromicrocotyla vaginispina Unnithan, 1961		Mo			Carangoides malabaricus		Gi	GOT, SCS		[73, 118]
Heteromicrocotyla sp.		Mo			Carangidae gen. sp., Carangoides malabaricus, Decapterus sp., Selaroides leptolepis		Gi	GOT, SCS		[73, 76, 118]
Family: Hexostomatidae Price, 1936
Hexostoma thynni (Delaroche, 1811) Rafinesque, 1815		Mo			Auxis thazard		Gi	SCS		[80]
Neohexostoma euthynni (Meserve, 1938) Price, 1961		Mo			Auxis thazard, Euthynnus affinis		Gi	SCS		[80]
Family: Mazocraeidae Price, 1936
Neomazocraes dorosomatis (Yamaguti, 1938) Price, 1943		Mo			Clupanodon thrissa		Gi	QN		[88]
Paramazocraes thrissocles Tripathi, 1959		Mo			Thrissocles sp.		Gi	GOT
Family: Microcotylidae Taschenberg, 1879
Caballeraxine chainanica (Lebedev, Parukhin & Roitman, 1970) Lebedev, 1972		Mo			Carangoides malabaricus		Gi	GOT, SCS		[118]
Diplostamenides sciaenae (Goto, 1894) Mamaev, 1986		Mo			Seriola sp.		Gi	GOT, SCS		[73, 118]
Incisaxine dubia Mamaev, 1970		Mo			Gerres sp.		Gi	GOT		[81]
Intracotyle orientalis Mamaev, 1970		Mo			Pomadasys argenteus		Gi	GOT		[81]
Lutianicola haifonensis Lebedev, 1970		Mo			Lutjanus russellii, L. sebae		Gi	GOT		[73]
Microcotyle sp.		Mo			Carangoides malabaricus		Gi	GOT		[73]
Polylabroides cf. guangdongensis Zhang & Yang, 2000		Mo			Acanthopagrus latus		Gi	QN		Present study
Polylabroides tienyenensis Nguyen, Nguyen, Ha, Ngoc, Ngoc, Le, Tatonova & Greiman, 2020		Mo			Acanthopagrus pacificus		Gi	QN		[93]
Polylabroides tonkinensis Nguyen, Nguyen, Ha, Ngoc, Ngoc, Le, Tatonova & Greiman, 2020		Mo			Acanthopagrus pacificus		Gi	QN		[93]
Tonkinaxine homocerca Lebedev, Parukhin & Roitman, 1970		Mo			Carangoides malabaricus, Seriola dumerili, Seriola sp., Seriolina nigrofasciata		Gi	GOT, SCS		[76, 118]
Family: Plectanocotylidae Monticelli, 1903
Triglicola tonkinensis Mamaev & Parukhin, 1972		Mo			Lepidotrigla sp.		Gi	GOT		[84, 119]
Family: Protomicrocotylidae Johnston & Tiegs, 1922
Bilaterocotyloides carangis Ramalingam, 1961		Mo			Carangidae gen. sp., Megalaspis cordyla		Gi	GOT, SCS		[73, 118]
Bilaterocotyloides madrasensis Radha, 1966		Mo			Megalaspis cordyla		Gi	GOT		[73, 76, 118]
Vallisiopsis contorta Subhapradha, 1951		Mo			Lactarius lactarius		Gi	GOT, SCS		[74]
Family: Thoracocotylidae Price, 1936
Pricea multae Chauhan, 1945		Mo			Scomberomorus commerson, S. guttatus		Gi	GOT		[73]
Pseudothoracocotyla ovalis (Tripathi, 1956) Yamaguti, 1963		Mo			Scomberomorus commerson, S. guttatus		Gi	GOT, SCS		[73, 119]
Monogenea gen. sp.		Mo			Abalistes stellaris		Gi	SCS		[119]
Class: Trematoda Rudolphi, 1808
Subclass: Digenea Carus, 1863
Order: Diplostomida Olson, Cribb, Tkach, Bray & Littlewood, 2003
Suborder: Diplostomata Olson, Cribb, Tkach, Bray & Littlewood, 2003
Superfamily: Schistosomatoidea Stiles & Hassall, 1898
Family: Aporocotylidae Odhner, 1912
Cardicola congruentus Lebedev & Mamaev, 1968		D			Euthynnus affinis		Bvg	GOT		[75]
Cardicola grandis Lebedev and Mamaev, 1968		D			Makaira sp.		Iw	GOT		[75]
Cardallagium cf. anthicum (Bullard & Overstreet, 2006) Yong, Cutmore, Jones, Gauthier & Cribb, 2017		D			Rachycentron canadum		He	KH		[141]
Order: Plagiorchiida La Rue, 1957
Suborder: Apocreadiata Olson, Cribb, Tkach, Bray & Littlewood, 2003
Superfamily: Apocreadioidea Skrjabin, 1942
Family: Apocreadiidae Skrjabin, 1942
Homalometron sp.		D			Gerres filamentosus		In	GOT		[81]
Schistorchis skrjabini Parukhin, 1963		D			Abalistes stellaris, Triacanthus biaculeatus		In	GOT		[106, 119]
Sphincteristomum acollum Oshmarin et al., 1961		D			Abalistes stellaris		In	GOT		[99, 119]
Suborder: Bivesiculata Olson, Cribb, Tkach, Bray & Littlewood, 2003
Superfamily: Bivesiculoidea Yamaguti, 1934
Family: Bivesiculidae Yamaguti, 1934
Bivesicula claviformis Yamaguti, 1934		D			Amphiprion clarckii, A. perideraion; A. polymmus		S	KH		[27]
Bivesicula sp., Metacercaria		D			Amphiprion clarckii, A. perideraion		In	KH		[142]
Paucivitellosus vietnamensis Atopkin, Besprozvannykh, Ngo, Van Ha, Van Tang, Ermolenko & Beloded, 2016		D			Liza subviridis		In	GOT		[20]
Suborder: Bucephalata La Rue, 1926
Superfamily: Bucephaloidea Poche, 1907
Family: Bucephalidae Poche, 1907
Alcicornis baylisi Nagaty, 1937		D			Carangoides malabaricus		In	SCS		[113]
Alcicornis carangis MacCallum, 1917		D			Caranx sp.		In	GOT, SCS		[71, 73]
Bucephalus fragilis Velasquez, 1959		D			Megalaspis cordyla, Scomberoides lysan		In	SCS		[113]
Bucephalus gorgon (Linton, 1905) Eckmann, 1932		D			Seriolina nigrofasciata		S	GOT		[95]
Bucephalus introversus Manter, 1940		D			Seriolina nigrofasciata		In	SCS		[118]
Bucephalus varicus Manter, 1940		D			Atropus atropos, Atule mate, Rachycentron canadum, Selar crumenophthalmus		S, In	SCS, GOT		[113, 117, 118]
Bucephalus paraheterotentaculatus Velasquez, 1959		D			Seriola dumerili, Seriolina nigrofasciata		In	SCS		[113, 118]
Bucephalus polymorphus von Baer, 1827		D			Lates calcarifer		In	KH		[140]
Bucephalus sp.		D			Gerres filamentosus		S	SCS, GOT		[81]
Prosorhynchus epinepheli Yamaguti, 1939		D			Epinephelus coioides, E. bleekeri, E. bruneus, E. malabaricus, E. sexfasciatus, E. tauvina, Lutjanus argentimaculatus		In	KH, QN		[34, 90, 140]
Prosorhynchus pacificus Manter, 1940		D			Epinephelus bleekeri, E. coioides, E. malabaricus		In	KH		[135]
Prosorhynchus luzonicus Velasquez, 1959		D			Epinephelus coioides		S, In, Py	HP, ND, QN, TH		[132]
Prosorhynchus tonkinensis Truong, Palm, Bui, Thuy Ngo & Bray, 2016		D			Epinephelus coioides		S, Py	ND		[132]
Prosorhynchus maternus Bray & Justine, 2006		D			Epinephelus coioides		S	HP		[132]
Prosorhynchus sp. A2		D			Epinephelus coioides		S	HP		[132]
Prosorhynchus sp. B3		D			Epinephelus coioides		S	HP		[132]
Prosorhynchus sp.		D			Acanthopagrus latus, Auxis thazard		In	NA, QB, QN SCS		[80, 90]; Present study
Rhipidocotyle laruei Velasquez, 1959		D			Psettodes erumei		S, In	SCS, GOT		[114, 119]
Rhipidocotyle pentagonum (Ozaki, 1924) Eckmann, 1932		D			Auxis thazard, Euthynnus affinis, Thunnus thynnus		S, In	SCS		[95]
Rhipidocotyle sp.		D			Drepane punctata, Ephippus orbis, Gerres filamentosus, Gymnocranius griseus, Leiognathus equulus, Parastromateus niger, Leiognathidae gen. sp., Sciaenidae gen. sp.		Gi, Go, In, K, Ve	GOT, SCS		[81, 95, 133]
Prosorhynchinae gen. sp.		D			Epinephelus coioides		In	QN		[133]
Superfamily: Gorgoderoidea Looss, 1901
Family: Callodistomidae Odhner, 1910
Cholepotes sp.		D			Parupeneus multifasciatus		In	KH		[27]
Superfamily: Gymnophalloidea Odhner, 1905
Family: Fellodistomidae Nicoll, 1909
Complexobursa vjetnamensis Oshmarin & Mamaev, 1963		D			Terapon theraps		In	GOT		[98]
Lintonium vibex (Linton, 1900) Stunkard & Nigrelli, 1930		D			Abalistes stellaris, Aluterus monoceros, Monacanthus chinensis, Scomberoides lysan, Scomberomorus commerson		In, L	GOT, SCS		[73, 95, 113]
Proctoeces sp.		D			Acanthopagrus latus		In	QN		Present study
Pseudosteringophorus sp.		D			Ephippus orbis		In	GOT		[81]
Tergestia laticollis (Rudolphi, 1819) Stossich, 1899		D			Alepes melanoptera, Decapterus sp., Megalaspis cordyla, Selar crumenophthalmus, Selaroides leptolepis		In	SCS		[118]
Family: Tandanicolidae Johnston, 1927
Buckleytrema indicum Gupta, 1956		D			Arius sp.		In	GOT		[98]
Monodhelmis torpedinis Dollfus, 1937		D			Arius arius		In	HP		[90]
Suborder: Haploporata Pérez-Ponce de León & Hernández-Mena, 2019
Superfamily: Haploporoidea Nicoll, 1914
Family: Haploporidae Nicoll, 1914
Parahaploporus elegantus Atopkin, Besprozvannykh, Ha, Nguyen & Nguyen, 2019		D			Osteomugil cunnesius		In	KH		[23]
Parasaccocoelium mugili Zhukov, 1971		D			Liza haematocheila		In	QN, HP		[23]
Pseudohaploporus vietnamensis Atopkin, Besprozvannykh, Ha, Tang, Nguyen, Nguyen & Chalenko, 2018		D			Moolgarda seheli, Osteomugil engeli		In	GOT		[22]
Pseudohaploporus planilizum Atopkin, Besprozvannykh, Ha, Nguyen, Nguyen & Chalenko, 2018		D			Planiliza subviridi		In	GOT		[22]
Pseudohaploporus pusitestis Atopkin, Besprozvannykh, Ha, Nguyen & Nguyen, 2019		D			Moolgarda seheli			HP		[23]
Pseudohaploporus sp.		D			Moolgarda seheli		In	GOT		[22]
Skrjabinolecithum spasskii Belous, 1954		D			Liza haematocheila, Mugil cephalus		In	HP		[90]
Suborder: Haplosplanchnata Olson, Cribb, Tkach, Bray & Littlewood, 2003
Superfamily: Haplosplanchnoidea Poche, 1925
Family: Haplosplanchnidae Poche, 1926
Haplosplanchnus pachysoma (Eysenhardt, 1829) Looss, 1902		D			Liza engeli		In	QN		[24]
Suborder: Hemiurata Skrjabin & Guschanskaja, 1954
Superfamily: Hemiuroidea Looss, 1899
Family: Accacoeliidae Odhner, 1911
Tetrochetus hansoni (Parukhin, 1964) Hafeezullah, 1982		D			Aluterus monoceros		In	GOT		[107]
Tetrochetus sp.		D			Nemipterus hexodon		S	KH		[27]
Family: Bathycotylidae
Bathycotyle sp.		D			Pampus argenteus		Gic	GOT		[73]
Family: Derogenidae Nicoll, 1910
Derogenes varicus (Müller, 1784) Looss, 1901		D			Rachycentron canadum, Triacanthus biaculeatus		S, In	GOT		[117, 119]
Gonocercella pacifica Manter, 1940		D			Drepane punctata		In	GOT		[81]
Gonocercella sp.		D			Atropus atropos, Psettodes erumei, Scomberoides lysan		In	GOT, SCS		[113, 118]
Family: Dictysarcidae Skrjabin & Guschanskaja, 1955
Elongoparorchis pneumatis Rao, 1961		D			Arius sp.		Sb	SCS		[96]
Family: Didymozoidae Monticelli, 1888
Colocyntotrema auxis Yamaguti, 1951		D			Scombridae gen. sp.		Bc, In	SCS		[80]
Didymodiclinus epinepheli (Abdul-Salam, Sreelatha & Farah, 1990) Pozdnyakov, 1994		D			Epinephelus coioides		Mu	KH		[135]
Lobatozoum multisacculatum Ishii, 1935		D			Auxis thazard		Iw, S, Ve	SCS		[80]
Metanematobothrium bivitellatum Mamaev, 1968		D			Auxis thazard, Euthynnus affinis		Bc, Bvg, L, K	SCS		[80]
Monilicaecum ventricosum Yamaguti, 1942		D			Abalistes stellaris, Psettodes erumei		Bvg, L	GOT		[119]
Multtubovarium amphibolum Mamaev, 1970		D			Platax orbicularis		Gi, Ht	GOT		[81]
Nematobothrium sp.		D			Euthynnus affinis, Thunnus thynnus		L, Phc	SCS		[80]
Neometadidymozoon polymorphis (Oschmarin & Mamaev, 1963) Yamaguti, 1971		D			Priacanthus tayenus		Bc, Gi, Mu	GOT		[98]
Neometanematobothrioides rachycentri (Parukhin, 1969) Yamaguti, 1971		D			Rachycentron canadum		Bc, Gi	GOT, SCS		[116, 118]
Oesophagocystis dissimilis (Yamaguti, 1938) Yamaguti, 1970		D			Auxis thazard, Euthynnus affinis, Thunnus thynnus		O, S, In	SCS		[80]
Torticaecum fenestratum (Linton, 1907) Yamaguti, 1942		D			Psettodes erumei, Triacanthus biaculeatus		Bvg, L	GOT		[119]
Didymozoidae gen. sp.		D			Atule mate, Auxis thazard, Carangoides malabaricus, Echeneis naucrates, Epinephelus coioides, Euthynnus affinis, Leiognathus equulus, Psettodes erumei, Rachycentron canadum, Selar crumenophthalmus, Seriolina nigrofasciata, Thunnus thynnus, Leiognathidae gen. sp.		Bc, Ve, Gi, In, K, L	GOT, SCS		[80, 81, 113, 115, 117, 133]
Family: Hemiuridae Looss, 1899
Allostomachicola secundus (Srivastava, 1939) Yamaguti, 1958		D			Chirocentrus dorab		S	GOT		[81]
Aphanurus mugilus Tang 1981		D			Moolgarda engeli		I	HP		[21]
Aphanurus stossichii (Monticelli, 1891) Looss, 1907		D			Drepane punctata, Ephippus orbis, Pampus argenteus, Thunnus thynnus		S, In	GOT, SCS		[73, 80, 81]
Aphanurus sp.		D			Mugil sp.		In	GOT		[90]
Dinurus euthynni Yamaguti, 1934		D			Auxis thazard, Euthynnus affinis, Thunnus sp.			SCS		[80]
Dinurus longisinus Looss, 1907		D			Carangoides malabaricus		S	GOT		[113, 118]
Dinurus selari Parukhin, 1966		D			Atropus atropos, Atule mate, Carangoides malabaricus, Decapterus sp., Rachycentron canadum, Selar crumenophthalmus, Selaroides leptolepis, Carangidae gen. sp.		S, In	GOT, SCS		[19, 111, 113, 118]
Dinurus sp.		D			Mene maculata		S	GOT		[81]
Ectenurus selari (Parukhin, 1966) Yamaguti, 1971		D			Atule mate, Caranx sp., Epinephelus bruneus, E. coioides, E. sexfasciatus, E. tauvina*, Megalaspis cordyla, Rachycentron canadum, Selar crumenophthalmus, Selaroides leptolepis, Carangidae gen. sp.		S, In	GOT, SCS		[34, 73, 112, 113, 118, 121]
Ectenurus theraponae Oshmarin, 1965		D			Terapon theraps		S	GOT		[95]
Ectenurus trachuri (Yamaguti, 1934) Yamaguti, 1970		D			Caranx sp., Selar crumenophthalmus, Carangidae gen. sp.		S	GOT, SCS		[71, 73]
Ectenurus sp.		D			Selar crumenophthalmus		S	GOT		[95]
Erilepturus formosae Reid, Coil & Kuntz, 1966		D			Decapterus sp., Dussumieria elopsoides		S	GOT		[73, 81]
Erilepturus hamati (Yamaguti, 1934) Manter, 1947		D			Eleutheronema tetradactylum, Epinephelus bleekeri, E. coioides, E. malabaricus, Platycephalus indicus, Nibea albiflora,		In	NA		[90, 138]
Erilepturus neopacificus (Velasquez, 1962) Gupta & Jain, 1992		D			Lates calcarifer		Gi	KH		[122]
Erilepturus sp.		D			Acanthopagrus latus, Sciaenidae gen. sp.		S	GOT, HL		[95]; Present study
Elytrophalloides sp.		D			Thryssa mystax		S	ND		[90]
Hemiurus arelisci Yamaguti, 1938		D			Scomberoides lysan		S	QN, ND		[88, 90]
Lecithochirium alectis Yamaguti, 1970		D			Nibea albiflora		In	QN, ND		[88, 90]
Lecithochirium imocavum (Looss, 1907) Skrjabin & Guschanskaja, 1955		D			Auxis thazard, Euthynnus affinis, Ilisha sp., Thunnus thynnus, Thunnus sp.		S	GOT		[80, 81]
Lecithochirium magnaporum Manter, 1940		D			Atropus atropos		S	SCS		[113]
Lecithochirium microstomum Chandler, 1935		D			Atropus atropos		S	SCS		[113]
Lecithochirium monticellii (Linton, 1898) Skrjabin & Guschanskaja, 1955		D			Amphiprion clarkia, Atropus atropos, Echeneis naucrates, Selar crumenophthalmus		S	KH, SCS		[27, 113, 115]
Lecithochirium sp.		D			Mene maculata, Seriolina nigrofasciata		S	GOT, SCS		[81, 113]
Lecithocladium apolecti Velasquez, 1962		D			Ephippus orbis, Gerres filamentosus, Parastromateus niger, Rastrelliger kanagurta, Leiognathidae gen. sp.		S, In	GOT		[73, 81]
Lecithocladium excisiforme Cohn, 1902		D			Alepes melanoptera, Caranx sp., Selaroides leptolepis		S	GOT, SCS		[118]
Lecithocladium excisum (Rudolphi, 1819) Lühe, 1901		D			Alepes melanoptera, Caranx sp., Decapterus maruadsi, Decapterus sp., Elentheronema tetradactylum, Sardinella sp., Selar crumenophthalmus, Scomberoides lysan		S	GOT, SCS		[73, 81, 88, 118]
Lecithocladium harpodontis Srivastava, 1942		D			Atropus atropos, Decapterus sp., Dussumieria elopsoides, Ilisha sp., Selar crumenophthalmus		S	GOT, SCS		[81, 113, 118]
Lecithocladium megalaspis Yamaguti, 1953		D			Megalaspis cordyla		S, In	SCS		[113]
Lecithocladium pampi Lebedev, 1968		D			Pampus argenteus		In	GOT, SCS		[70, 73]
Lecithocladium seriolellae Manter, 1954		D			Carangoides malabaricus, Selar crumenophthalmus		S, In	SCS		[113, 118]
Lecithocladium sp.		D			Mene maculata		S	GOT		[81]
Merlucciotrema praeclarum (Manter, 1934) Yamaguti, 1971		D			Platycephalus indicus, Eleutheronema tetradactylum		S, In	QN, ND		[90]
Parahemiurus clupeae Yamaguti, 1953		D			Herklotsichthys quadrimaculatus		S	KH		[64]
Parahemiurus merus (Linton, 1910) Manter, 1940		D			Atropus atropos, Decapterus sp., Sardinella sp., Scomberoides lysan		S, In	GOT, SCS		[81, 118]
Parahemiurus sp.		D			Alepes kleinii		S	QN		[88]
Plerurus digitatus (Looss, 1899) Looss, 1907		D			Auxis thazard, Euthynnus affinis, Thunnus sp., Carangidae gen. sp.			GOT, SCS		[80, 113, 118]
Stomachicola muraenesocis Yamaguti, 1934		D			Congresox talabonoides		S	QN (QB)		[88, 90]
Tubulovesicula angusticauda (Nicoll, 1915) Yamaguti, 1934		D			Epinephelus merra, Rachycentron canadum, Trachinocephalus sp.		S, In	GOT, SCS		[64, 95, 118]
Tubulovesicula lindbergi (Layman, 1930) Yamaguti, 1934		D			Echeneis naucrates, Psettodes erumei, Carangidae gen. sp.		S	GOT, SCS		[115, 118, 119]
Tubulovesicula marsupialia Oshmarin, 1965		D			Saurida tumbil		In	GOT		[95]
Tubulovesicula sp.		D			Trachinocephalus myops		S, In	KH		[27]
Hemiuridae gen. sp.		D			Euthynnus affinis, Thunnus thynnus, Thunnus sp.			SCS		[80]
Family: Hirudinellidae Dollfus, 1932
Hirudinella ventricosa (Pallas, 1774) Baird, 1853		D			Euthynnus affinis, Seriolina nigrofasciata, Thunnus thynnus		S	SCS		[80, 113]
Family: Lecithasteridae Odhner, 1905
Aponurus carangis Yamaguti, 1952		D			Decapterus sp., Rachycentron canadum, Selar crumenophthalmus		S, In	GOT, SCS		[73, 113, 118]
Aponurus laguncula Looss, 1907		D			Amphiprion polymnus, Atropus atropos, Drepane punctata, Dussumieria elopsoides, Ilisha sp., Leiognathus sp., Megalaspis cordyla, Parastromateus niger, Sardinella sp., Selar crumenophthalmus, Seriolina nigrofasciata, Thunnus sp., Carangidae gen. sp.		In, S	GOT, SCS		[80, 81, 113, 118, 142]
Aponurus pyriformis (Linton, 1910) Overstreet, 1973		D			Amphiprion clarckii, Parastromateus niger, Platax orbicularis		In, S	GOT, KH		[81, 142]
Aponurus sp.		D			Epinephelus coioides		S	GOT		[133]
Hysterolecitha nahaensis Yamaguti, 1942		D			Amphiprion clarckii, A. frenatus, A. perideraion, A. polymnus, Dascyllus trimaculatus		S	SCS		[64, 142]
Lecithaster mugilis Yamaguti, 1970		D			Moolgarda engeli, M. seheli, Liza subviridis		In	GOT		[25]
Lecithaster stellatus Looss, 1907		D			Seriolina nigrofasciata		S	SCS		[113]
Trifoliovarium triacanthi (Parukhin, 1964) Bray & Cribb, 2000		D			Triacanthus biaculeatus		In	GOT		[108]
Family: Sclerodistomidae Odhner, 1927
Prosogonotrema bilabiatum Vigueras, 1940		D			Abalistes stellaris, Ephippus orbis, Platax orbicularis		S	GOT		[81, 107, 119]
Prosogonotrema symmetricum Oshmarin, 1965		D			Lutjanus sp., Pristipomoides typus		S	GOT		[95, 118]
Prosorchis chainanensis Lebedev, 1970		D			Ephippus orbis, Pampus argenteus, Parastromateus niger		S	GOT		[73, 81]
Hemiuroidea gen. sp.		D			Alectis indicus, Gnathanodon speciosus, Megalaspis cordyla, Rachycentron canadum, Seriolina nigrofasciata, Terapon theraps, Carangidae gen. sp.		In	GOT, SCS		[95, 113, 117]
Suborder: Lepocreadiata Olson, Cribb, Tkach, Bray & Littlewood, 2003
Superfamily: Lepocreadioidea Odhner, 1905
Family: Aephnidiogenidae Yamaguti, 1934
Aephnidiogenes barbarus Nicoll, 1915		D			Pomadasys argenteus		In	GOT		[81]
Family: Gyliauchenidae Fukui, 1929
Gyliauchen tarachodes Nicoll, 1915		D			Siganus fuscescens		In	HP		[90]
Family: Lepocreadiidae Odhner, 1905
Diploproctia drepanei Mamaev, 1970		D			Drepane punctata		In	GOT		[81, 90]
Diploproctodaeum haustrum (MacCallum, 1919) La Rue, 1926		D			Aluterus monoceros		In	QB		[90]
Diploproctodaeoides longipygum (Oshmarin, Mamaev & Parukhin, 1961) Reimer, 1981		D			Abalistes stellaris		In	GOTh, GOT		[99]
Diploproctodaeum macracetabulum Oshmarin, Mamaev & Parukhin, 1961		D			Abalistes stellaris, Triacanthus biaculeatus		In	GOTh, GOT		[99]
Diploproctodaeum plataxi Mamaev, 1970		D			Platax orbicularis		In	GOT		[81]
Diploproctodaeum rutellum (Mamaev, 1970) Bray, Cribb & Barker, 1996		D			Ephippus orbis, Platax orbicularis		In	GOT		[81]
Echeneidocoelium indicum Simha & Pershad, 1964		D			Echeneis naucrates, Psettodes erumei		In, S	GOT, SCS		[112, 119]
Hypocreadium cavum Bray & Cribb, 1966		D			Abalistes stellaris		In	GOT		[95]
Hypocreadium scaphosomum (Manter, 1940) Bravo Hollis & Manter, 1957		D			Abalistes stellaris, Aluterus monoceros, Triacanthus biaculeatus		In	GOTh, GOT		[119]
Hypocreadium sp.		D			Abalistes stellaris			GOTh		[119]
Lepidapedon megalaspi Parukhin, 1966		D			Carangoides malabaricus, Decapterus sp., Megalaspis cordyla, Rachycentron canadum		In	GOT, SCS		[112, 113]
Lepocreadium sp.		D			Ephippus orbis, Triacanthus biaculeatus		In	GOT		[81, 119]
Multitestis magnacetabulum Mamaev, 1970		D			Ephippus orbis, Platax orbicularis		In	GOT		[81]
Neoallolepidapedon fistulariae (Oshmarin, 1965) Yamaguti, 1965		D			Fistularia petimba		S	GOT		[96]
Opechona formiae Oshmarin, 1965		D			Pampus argenteus, Parastromateus niger, Leiognathidae gen. sp.		S, In	GOT		[73, 95]
Trigonotrema alatum Goto & Ozaki, 1929		D			Brachiostegus japonicus, Drepane longimana, D. punctata, Ephippus orbis		In, S	GOT		[95]
Lepocreadiidae gen. sp.		D			Plectorhinchus cinctus		In	GOT		[81]
Suborder: Monorchiata Olson, Cribb, Tkach, Bray & Littlewood, 2003
Superfamily: Monorchioidea Odhner, 1911
Family: Monorchiidae Odhner, 1911
Alloinfundiburictus cacuminatus (Nicoll, 1915) Wee, Cutmore, Pérez-del-Olmo & Cribb, 2020		D			Pomadasys argenteus		In, S	GOT		[95]
Alloinfundiburictus cryptostoma (Oshmarin, 1966) Wee, Cutmore, Pérez-del-Olmo & Cribb, 2020		D			Pomadasys argenteus		In, S	GOT		[81]
Ancylocoelium tropicum (Manter, 1940) Wee, Cutmore, Pérez-del-Olmo & Cribb, 2020		D			Carangoides malabaricus, Megalaspis cordyla		In	GOT		[81]
Hurleytrematoides chaetodoni (Manter, 1942) Yamaguti, 1954		D			Chaetodon sp.		In	GOT		[81]
Huridostomum formionis Mamaev, 1970		D			Parastromateus niger		In	GOT		[81]
Infundiburictus chaetodipteri (Thomas, 1959) Wee, Cutmore, Pérez-del-Olmo & Cribb, 2020		D			Pomadasys argenteus		In, S	GOT		[81]
Leiomonorchis mamaevi Madhavi, 2008		D			Leiognathus equulus, Parastromateus niger, Leiognathidae gen. sp.		In	GOT		[81]
Monorcheides diplorchis Odhner, 1905		D			Scolopsis taeniopterus		I	KH		[27]
Monorchis diplovarium Mamaev, 1970		D			Pomadasys argenteus		In	GOT		[81]
Opisthomonorcheides decapteri Parukhin, 1966		D			Atule mate, Decapterus sp.		In	GOT, SCS		[112, 113]
Opisthomonorcheides ovacutus (Mamaev, 1970) Machida, 2011		D			Parastromateus niger		In	GOT		[81]
Opisthomonorchis carangis Yamaguti, 1952		D			Carangoides malabaricus, Carangidae gen. sp.		In	GOT, SCS		[118]
Paralasiotocus macrorchis (Yamaguti, 1934) Wee, Cutmore, Pérez-del-Olmo & Cribb, 2020		D			Plectorhinchus cinctus, Plectorhinchus sp.		In	GOT		[81]
Proctotrema sp.		D			Pomadasys argenteus, Selar crumenophthalmus		In	GOT, SCS		[95, 113, 119]
Sinistroporomonorchis lizae (Liu, 2002) Wee, Cutmore, Pérez-del-Olmo & Cribb, 2020		D			Moolgarda cunnesius, Liza engeli, L. longimanus, L. subviridis, Valamugil seheli		In	GOT		[20]
Suborder: Opisthorchiata La Rue, 1957
Superfamily: Opisthorchioidea Looss, 1899
Family: Cryptogonimidae Ward, 1917
Beluesca plectorhyncha (Mamaev, 1970) Miller & Cribb, 2007		D			Plectorhinchus cinctus		In	GOT		[81]
Metadena eurystoma Oshmarin, 1965		D			Sciaenidae gen. sp.		In	GOT		[95]
Metadena longa (Oshmarin, Mamaev & Parukhin, 1961) Miller & Cribb, 2008		D			Pristipomoides typus		In	GOT		[99]
Metadena sp.		D			Lutjanus russellii		In	QB		[90]
Neometadena ovata (Yamaguti, 1952) Miller & Cribb, 2008		D			Lutjanus russellii		In	QB		[90]
Pseudometadena celebesensis Yamaguti, 1952		D			Lates calcarifer		In	KH		[140]
Siphoderina echinostomus (Oshmarin, Mamaev & Parukhin, 1961) Miller & Cribb, 2008		D			Pristipomoides typus		In	GOT		[99]
Siphoderina morosovi (Parukhin, 1965) Miller & Cribb, 2008		D			Rachycentron canadum		In	GOT, SCS		[110, 118]
Siphoderina sp.		D			Lutjanus argentimaculatus		In	QB		[90]
Family: Heterophyidae Leiper, 1909
Centrocestus sp.		D			Epinephelus coioides		Mu, Fi	GOT		[133]
Heterophyopsis continua (Onji & Nishio, 1916) Price, 1940		D			Epinephelus bleekeri, E. coioides		Mu, Fi	KH		[130, 135, 137]
Procerovum varium Onji & Nishio, 1916		D			Epinephelus bleekeri, E. coioides		Mu, Fi	KH		[135]
Heterophyidae gen. sp.		D			Epinephelus coioides, Protonibea dicanthus, Sardinella sp.		In	GOT, SCS		[81, 133]; Present study
Suborder: Pronocephalata Skrjabin, 1955
Superfamily: Paramphistomoidea Fischoeder, 1901
Family: Cladorchiidae Fischoeder, 1901
Cleptodiscus sp.		D			Triacanthus biaculeatus		In	GOT		[119]
Suborder: Transversotremata Olson, Cribb, Tkach, Bray & Littlewood, 2003
Superfamily: Transversotrematoidea Witenberg, 1944
Family: Transversotrematidae Witenberg, 1944
Transversotrema patialense (Soparkar, 1924) Crusz & Sathananthan, 1960		D			Epinephelus bleekeri, E. coioides		Us	KH, ND		[133, 140]
Suborder: Xiphidiata Olson, Cribb, Tkach, Bray & Littlewood, 2003
Superfamily: Brachycladioidea Odhner, 1905
Family: Acanthocolpidae Lühe, 1906
Acanthocolpus liodorus Lühe, 1906		D			Chirocentrus dorab, Ilisha sp., Sardinella sp.		Ve	GOT		[82]
Pleorchis hainanensis Shen, 1983		D			Johnius carouna, Nibea albiflora, Pennahia argentata		In	QN, QB		[90]
Pleorchis sciaenae Yamaguti, 1938		D			Acanthopagrus berda, Nibea albiflora, Sciaenidae gen. sp.		In	QN, HP		[90, 95]
Stephanostomum bicoronatum (Stossich, 1883) Fuhrmann, 1928		D			Johnius carouna		In	QN, ND		[88]
Stephanostomum ditrematis (Yamaguti, 1939) Manter, 1947		D			Scomberoides lysan, Seriola dumerili, Seriolina nigrofasciata		In	SCS, QN, ND		[49, 113]
Stephanostomum fistulariae (Yamaguti, 1940) Manter & Van Cleave, 1951		D			Fistularia petimba, Harpadon nehereus		In	GOT, HP		[90, 95]
Stephanostomum hispidum (Yamaguti, 1934) Manter, 1940		D			Seriolina nigrofasciata		In	SCS		[113]
Stephanostomum imparispine (Linton, 1905) Manter, 1940		D			Abalistes stellaris, Aluterus monoceros, Echeneis naucrates, Psettodes erumei, Rachycentron canadum, Seriolina nigrofasciata, Triacanthus biaculeatus		Bc, Gi, In	GOT, GOTh, SCS		[113, 114, 119]
Stephanostomum orientale (Srivastava, 1939) Madhavi, 1976		D			Seriola dumerili, Seriolina nigrofasciata		In	SCS		[113]
Stephanostomum tenue (Linton, 1898) Linton, 1934		D			Chirocentrus dorab		S	GOT		[84]
Stephanostomum sp.		D			Ephippus orbis, Epinephelus coioides, Sciaenidae gen. sp.		In	GOT		[81, 114, 133]
Tormopsolus carangi Parukhin, 1976		D			Carangoides malabaricus		In, S	SCS		[118]
Tormopsolus filiformis Sogandares-Bernal & Hutton, 1958		D			Carangoides malabaricus, Rachycentron canadum		In	GOT, SCS		[113, 119]
Tormopsolus orientalis Yamaguti, 1934		D			Carangoides malabaricus		S	GOT, SCS		[71, 73, 113]
Superfamily: Opecoelioidea Ozaki, 1925
Family: Opecoelidae Ozaki, 1925
Allopodocotyle epinepheli (Yamaguti, 1942) Pritchard, 1966		D			Drepane punctata		In	GOT		[81]
Allopodocotyle sp.		D			Epinephelus coioides		In	QN, HP		[133]
Cainocreadium labracis (Dujardin, 1845) Nicoll, 1909		D			Scorpaenopsis cacopsis, Synodus variegatus		S	KH		[27]
Coitocaecum gymnophallum Nicoll, 1915		D			Acanthopagrus berda, A. latus		In	QN, HP		[90], Present study
Helicometra fasciata (Rudolphi, 1819) Odhner, 1902		D			Epinephelus bleekeri, E. coioides, E. malabaricus, E. quoyanus, E. sexfasciatus		In	GOT, KH		[34, 90, 139]
Helicometra pisodonophi Nguyen Van Ha, 20123		D			Pisodonophis cancrivorus		In	QN, HP		[48]
Helicometrina nimia Linton, 1910		D			Epinephelus sexfasciatus		In	QB
Helicometra sp.		D			Acanthopagrus latus, Epinephelus coioides		In	QN		[133]; Present study
Macvicaria sp.		D			Amphiprion clarckii		In	KH		[142]
Neonotoporus decapteri Parukhin, 1966		D			Decapterus sp.		In	SCS, GOT		[112, 113]
Opegaster brevifistula Ozaki, 1928		D			Sillago sihama		In	GOT		[90]
Opecoelus haduyngoi Nguyen Van Ha, 2012		D			Acanthopagrus berda		In	QN, HP		[48]
Opecoelus pteroisi Shen, 1986		D			Pennahia argentata		In	QN, HP		[88, 90]
Opecoelus sphaericus Ozaki, 1925		D			Ephippus orbis, Platax orbicularis		In	GOT		[81]
Opecoelus sp.		D			Megalaspis cordyla		In	SCS		[113]
Opecoelina vixiintestina Oshmarin, 1965		D			Terapon theraps		In, S	GOT		[95]
Opecoelina sp.		D			Epinephelus amblycephalus		In	QB		[90]
Opegaster parapristipomatis Yamaguti, 1934		D			Gerres filamentosus		In	GOT		[81]
Opistholebes amplicoelus Nicoll, 1915		D			Lagocephalus lunaris		In	HP		[90]
Podocotyloides petalophallus Yamaguti, 1934		D			Plectorhinchus cinctus, Plectorhinchus sp.		In	GOT		[81]
Pseudopecoeloides carangis (Yamaguti, 1938) Yamaguti, 1940		D			Alectis indicus, Megalaspis cordyla		In	SCS		[113]
Pseudopecoeloides sp.		D			Atropus atropus		In	HP		[90]
Phyllotrema sp.		D			Pinjalo pinjalo		In	QB		[90]
Pycnadenoides pagrosomi Yamaguti, 1938		D			Sciaenidae gen. sp.		In	GOT		[95]
Vesicocoelium solenophagum Tang, Hsu, Huang & Lu, 1975		D			Gerres limbatus		In	QB		[90]
Superfamily: Gorgoderoidea Looss, 1901
Family: Gorgoderidae Looss, 1899
Phyllodistomum carangis (MacCallum, 1913) Cutmore & Cribb, 2018		D			Scomberoides lysan		In	QN, HP, SCS		[90, 113]
Phyllodistomum lancea Mamaev, 1968		D			Auxis thazard, Euthynnus affinis		In	SCS		[80]
Phyllodistomum notosinicum Lebedev, 1970		D			Scomberomorus sp., Rastrelliger brachysoma		In	GOT, SCS		[73, 90]
Phyllodistomum parukhini Yamaguti, 1971		D			Rachycentron canadum		K, Ub	GOT, SCS		[106]
Phyllodistomum psettodi Parukhin, 1966		D			Psettodes erumei		Ub	GOT		[112]
Phyllodistomum strictum Oshmarin, 1965		D			Parastromateus niger		In	GOT		[95]
Phyllodistomum sp.		D			Nibea albiflora		Ub	QN, HP		[88, 90]
Xystretrum abalisti Parukhin, 1963		D			Triacanthus biaculeatus		Ub	GOT		[119]
Superfamily: Microphalloidea Ward, 1901
Family: Faustulidae Poche, 1926
Paradiscogaster drepanei Mamaev, 1970		D			Drepane longimana, D. punctata		In	GOT		[81]
Family: Microphallidae Ward, 1901
Microphallus sp.		D			Parupeneus multifasciatus		In	KH		[27]
Family: Zoogonidae Odhner, 1902
Plectognathotrema ovatum Parukhin, 1964		D			Aluterus monoceros		In	GOT, SCS		[107, 119]
Digenea gen. sp.		D			Alectis indicus, Alepes melanoptera, Atropus atropos, A. oreolatus, Atule mate, Carangoides chrysophrys, C. malabaricus, Caranx sp., Decapterus sp., Echeneis naucrates, Epinephelus coioides, Gnathanodon speciosus, Megalaspis cordyla, Pomadasys argenteus, Psettodes erumei, Rachycentron canadum, Scomberoides lysan, Selar crumenophthalmus, S. crumenophthalmus, Selaroides leptolepis, Seriola dumerili, Seriolina nigrofasciata, Carangidae gen. sp.		Bc, Ve, Mu, In, S	SCS, GOT		[19, 34, 133]
Class: Cestoda Rudolphi, 1808
Subclass: Eucestoda Southwell, 1930
Order: Bothriocephalidea Kuchta, Scholz, Brabec & Bray, 2008
Family: Bothriocephalidae Blanchard, 1849
Bothriocephalus manubriformis (Linton, 1889) Ariola, 1900		C			Makaira sp.		In	GOT		[73]
Order: Diphyllobothriidea Kuchta, Scholz, Brabec & Bray, 2008
Family: Diphyllobothriidae Lühe, 1910
Diphyllobothrium sp.		C			Alepes melanoptera, Atule mate		In, Mu	SCS		[113]
Order: Lecanicephalidea Hyman, 1951
Family: Cephalobothriidae Pintner, 1928
Tylocephalum sp.		C			Epinephelus bleekeri, E. coioides, E. malabaricus, Lates calcarifer		In	KH		[140]
Order: Onchoproteocephalidea Caira, Jensen, Waeschenbach, Olson & Littlewood, 2014
Family: Proteocephalidae La Rue, 1911
Proteocephalus sp.		C			Lates calcarifer		In	KH		[122]
Order: Tetraphyllidea Carus, 1863
Family: Tetraphyllidea incertae sedis
Scolex sp.		C			Auxis thazard, Thunnus sp.		Bc, L	SCS		[80]
Tetraphyllidea gen. sp. plerocercoid		C			Abalistes stellaris, Acanthocepola limbata, Carangoides malabaricus, Caranx sp., Decapterus muroadsi, Decapterus sp., Drepane punctata, Echeneis naucrates, Epinephelus coioides, Mene maculata, Parastromateus niger, Protonibea diacanthus, Selar crumenophthalmus, Carangidae gen. sp.		Bc, Gb, In, S	GOT, SCS		[19, 73, 81, 119, 133]; Present study
Order: Trypanorhyncha Diesing, 1863
Suborder: Trypanobatoida Olson, Caira, Jensen, Overstreet, Palm & Beveridge, 2010
Superfamily: Tentacularioidea Poche, 1926
Family: Tentaculariidae Poche, 1926
Nybelinia sp.		C			Alectis indicus, Carangoides malabaricus, Caranx sp., Cepola schlegelii, Decapterus muroadsi, Decapterus sp., Leiognathus sp., Megalaspis cordyla, Mene maculata, Parastromateus niger, Platax orbicularis, Pomadasys argenteus, Rastrelliger kanagurta, Scomberoides lysan, Scomberomorus commerson, S. guttatus, Scorpaenodes sp., Selar crumenophthalmus, Selaroides leptolepis, Selaroides sp., Seriola dumerili, Seriola sp., Seriolina nigrofasciata, Triacanthus biaculeatus, Carangidae gen. sp., Leiognathidae gen. sp., Scombridae gen. sp.		Bc, Go, In, K, L, S	GOT, SCS		[73, 81, 119, 143]
Tentacularia coryphaenae Bosc, 1802		C			Acanthocybium solandri		SW	HP		Present study
Superfamily: Eutetrarhynchoidea Guiart, 1927
Family: Eutetrarhynchidae Guiart, 1927
Dollfusiella sp. A4		C			Telatrygon zugei		Sv	HP		Present study
Dollfusiella sp. B		C			Neotrygon kuhlii		Sv	HP		Present study
Oncomegas wageneri (Linton, 1890) Dollfus, 1929		C			Acanthocepola limbata, Cepola schlegelii, Cepola sp.		Bc	GOT		[81]
Prochristianella sp.		C			Telatrygon zugei		Sv	HP		Present study
Eutetrarhynchidae gen. sp.		C			Chirocentrus dorab, Ilisha sp.		Bc	GOT		[81]
Family: Rhinoptericolidae Carvajal & Campbell, 1975
Shirleyrhynchus cf. butlerae Beveridge & Campbell, 1988		C			Neotrygon kuhlii, Telatrygon zugei		Sv	HP		Present study
Suborder: Trypanoselachoida Olson, Caira, Jensen, Overstreet, Palm & Beveridge, 2010
Superfamily: Lacistorhynchoidea Guiart, 1927
Family: Pterobothriidae Pintner, 1931
Pterobothrium platycephalum (Shipley & Hornell, 1906) Dollfus, 1930		C			Mene maculata, Parastromateus niger, Platax orbicularis		Bc	GOT		[81]
Family: Lacistorhynchidae Guiart, 1937
Grillotia sp.		C			Drepane punctata, Platax orbicularis		Bc	GOT		[81]
Superfamily: Otobothrioidea Dollfus, 1942
Family: Lacistorhynchidae Guiart 1937
Callitetrarhynchus gracilis (Rudolphi, 1819) Pintner, 1931		C			Auxis thazard, Epinephelus coioides, E. malabaricus, Euthynnus affinis, Thunnus thynnus, Thunnus sp.		S	SCS, KH		[80, 140]
Family: Otobothriidae Dollfus, 1942
Otobothrium sp.		C			Abalistes stellaris, Aluterus monoceros, Auxis thazard, Cepola schlegelii, Cepola sp., Drepane punctata, Ephippus orbis, Gymnocranius griseus, Leiognathus equula, Leiognathus sp., Parastromateus niger, Platax orbicularis, Pomadasys argenteus, Leiognathidae gen. sp.		Bc, Sw, In	GOT		[80, 81, 119]
Family: Pseudotobothriidae Palm, 1995
Parotobothrium balli (Southwell, 1929) Palm, 2004		C			Epinephelus bleekeri, E. coioides, E. malabaricus		S	KH		[135]
Trypanorhyncha gen. sp.		C			Abalistes stellaris, Acanthocepola limbata, Alepes melanoptera, Aluterus monoceros, Atule mate, Carangoides malabaricus, Caranx sp., Cepola schlegelii, Decapterus sp., Echeneis naucrates, Ephippus orbis, Gymnocranius griseus, Mene maculata, Parastromateus niger, Pomadasys argenteus, Psettodes erumei, Rachycentron canadum, Selaroides leptolepis, Carangidae gen. sp.		Bc, Gi, In, Mu	GOT, GOTh, SCS		[19, 81, 109, 113, 115, 117, 119]
Cestoda gen. sp.		C			Abalistes stellaris, Alectis indicus, Alepes melanoptera, Atropus atropos, A. oreolatus, Atule mate, Carangoides chrysophrys, C. malabaricus, Caranx sp., Decapterus sp., Echeneis naucrates, Epinephelus coioides, Gnathanodon speciosus, Megalaspis cordyla, Psettodes erumei, Rachycentron canadum, Selar crumenophthalmus, Selaroides leptolepis, Seriola dumerili, Seriolina nigrofasciata, Carangidae gen. sp.			GOT		[19, 109, 133]
Phylum: Nematoda Rudolphi, 1808
Class: Chromadorea Inglis, 1983
Order: Rhabditida Chitwood, 1933
Suborder: Spirurina Railliet & Henry, 1915
Infraorder: Ascaridomorpha De Ley & Blaxter, 2002
Superfamily: Ascaridoidea Baird, 1853
Family: Anisakidae Railliet & Henry, 1912
Anisakis sp. Larvae		N			Abalistes stellaris, Acanthocepola limbata, Alectis indicus, Alepes melanoptera, Atropus atropos, A. oreolatus, Atule mate, Auxis thazard, Carangoides chrysophrys, Carangoides malabaricus, Caranx sp., Cepola schlegelii, Cepola sp., Chirocentrus dorab, Decapterus muroadsi, Decapterus sp., Drepane longimana, D. punctata, Dussumieria elopsoides, Echeneis naucrates, Ephippus orbis, Epinephelus bleekeri, E. coioides, E. malabaricus, Euthynnus affinis, Gerres filamentosus, Gerres sp., Gymnocranius griseus, Ilisha sp., Leiognathus equulus, Leiognathus sp., Lutjanus russellii, L. sebae, Makaira sp., Megalaspis cordyla, Mene maculata, Pampus argenteus, Parastromateus niger, Platax orbicularis, Pomadasys argenteus, Psettodes erumei, Rachycentron canadum, Rastrelliger kanagurta, Sardinella sp., Scomberoides lysan, Scomberomorus commerson, S. guttatus, Scomberomorus sp., Selar crumenophthalmus, Selaroides leptolepis, Selaroides sp., Seriola dumerili, Seriola sp., Seriolina nigrofasciata, Thunnus thynnus, Xiphias sp., Carangidae gen. sp., Chaetodontidae gen. sp., Scombridae gen. sp., Leiognathidae gen. sp.		Bc, Sw, In,	GOT, SCS, GOTh		[19, 73, 80, 113, 118, 119, 140]
Contracaecum sp.		N			Abalistes stellaris, Alectis indicus, Alepes melanoptera, Atropus atropos, Atule mate, Auxis thazard, Carangoides malabaricus, Caranx sp., Decapterus sp., Echeneis naucrates, Euthynnus affinis, Lutjanus russellii, L. sebae, Makaira sp., Megalaspis cordyla, Mene maculata, Pampus argenteus, Parastromateus niger, Platax orbicularis, Psettodes erumei, Rachycentron canadum, Rastrelliger kanagurta, Sardinella sp., Scomberoides lysan, Scomberomorus commerson, S. guttatus, Scomberomorus sp., Selar crumenophthalmus, Selar sp., Selaroides leptolepis, Selaroides sp., Seriola dumerili, Seriola sp., Seriolina nigrofasciata, Thunnus thynnus, Xiphias sp., Carangidae gen. sp., Scombridae gen. sp.		Bc, Sw, In, K	GOT, SCS, GOTh		[19, 73, 113, 118, 119, 140]
Family: Ascarididae Baird, 1853
Porrocaecum sp.		N			Abalistes stellaris, Alectis indicus, Alepes melanoptera, Atropus atropos, Atule mate, Carangoides chrysophrys, C. malabaricus, Caranx sp., Chirocentrus dorab, Decapterus muroadsi, Decapterus sp., Dussumieria elopsoides, Echeneis naucrates, Gnathanodon speciosus, Ilisha sp., Lutjanus lutjanus, L. russellii, L. sebae, Megalaspis cordyla, Pampus argenteus, Psettodes erumei, Rastrelliger kanagurta, Sardinella sp., Scomberoides lysan, Scomberomorus commerson, S. guttatus, Scomberomorus sp., Selar crumenophthalmus, Selaroides leptolepis, Selaroides sp., Seriola dumerili, Seriola sp., Seriolina nigrofasciata, Triacanthus biaculeatus, Carangidae gen. sp., Scombridae gen. sp.		Bc, In	GOT, GOTh, SCS		[73, 81, 109, 113, 119]
Family: Raphidascarididae Hartwich, 1954
Hysterothylacium aduncum (Rudolphi, 1802)		N			Epinephelus bleekeri, E. coioides, E. malabaricus		Sw	KH		[135]
Hysterothylacium chorinemi (Parukhin, 1966) Bruce & Cannon, 1989		N			Atule mate, Scomberoides lysan		In	SCS		[113]
Hysterothylacium incurvum (Rudolphi, 1819) Deardorff & Overstreet, 1980		N			Xiphias sp.		In	GOT		[73]
Hysterothylacium saba (Yamaguti, 1941) Deardorff & Overstreet, 1980		N			Scombridae gen. sp.		In	GOT		[73]
Hysterothylacium sp.		N			Amphiprion clarckii, A. frenatus, A. polymnus		L	KH		[142]
Iheringascaris inquies (Linton, 1901) Deardorff & Overstreet, 1980		N			Rachycentron canadum		In	GOT, SCS		[118]
Raphidascaris sp. (Larva)		N			Echeneis naucrates, Epinephelus bleekeri, E. coioides, E. malabaricus			SCS		[115, 135]
Superfamily: Seuratoidea Hall, 1916
Family: Cucullanidae Cobbold, 1864
Cucullanus decapteri Parukhin, 1966		N			Decapterus sp.		In	SCS		[113]
Cucullanus heterochrous Rudolphi, 1802		N			Psettodes erumei		In	SCS		[114]
Cucullanus sp.		N			Drepane punctata, Gymnocranius griseus		In	GOT		[81]
Infraorder: Gnathostomatomorpha De Ley & Blaxter, 2002
Superfamily: Gnathostomatoidea Railliet, 1895
Family: Gnathostomatidae Railliet, 1895
Echinocephalus spinosissimus (von Linstow in Shipley et Hornell, 1905)		N			Abalistes stellaris, Echeneis naucrates		Io	SCS, GOT		[118, 119]
Echinocephalus sp.		N			Acanthopagrus latus, Echeneis naucrates, Ilisha sp., Triacanthus biaculeatus,		In	GOT. SCS		[81, 115, 119] Present study
Infraorder: Spirurina incertae sedis
Superfamily: Dracunculoidea Stiles, 1907
Family: Philometridae Baylis & Daubney, 1926
Buckleyella buckleyi Rasheed, 1963		N			Scomberoides lysan		Py	SCS		[111, 118]
Philometra balistii (Rasheed, 1963) Vidal-Martínez, Aguirre- Macedo & Moravec, 1995		N			Abalistes stellaris		Oe	GOTh, GOT		[116, 119]
Philometra spinosa Vo, 20105		N			Epinephelus coioides		Fi	KH		[135]
Philometra sp.		N			Abalistes stellaris, Carangoides malabaricus, Caranx sp., Decapterus sp., Epinephelus bleekeri, E. coioides, E. malabaricus, Megalaspis cordyla, Parastromateus niger, Psettodes erumei, Sardinella sp., Triacanthus biaculeatus, Leiognathidae gen. sp		Op, Fi	GOT, SCS, QN, KH		[19, 71, 73, 117–119]
Philometra sp. 16		N			Epinephelus coioides		Fi	ND		[133]
Philometra sp. 2		N			Epinephelus coioides		eyes	HP		[133]
Philometroides atropi (Parukhin, 1966) Moravec & Ergens, 1970		N			Atropus atropos		Bc	GOT, SCS		[112, 113]
Philometroides sp.		N			Euthynnus affinis, Rachycentron canadum		Bc	GOT		[117]
Infraorder: Spiruromorpha De Ley & Blaxter, 2002
Superfamily: Camallanoidea Travassos, 1920
Family: Camallanidae Railliet & Henry, 1915
Camallanus sp.		N			Echeneis naucrates, Psettodes erumei		In	GOT, SCS		[114, 115]
Procamallanus (Spirocamallanus) guttatusi (Andrade-Salas, Pineda-López & García-Magaña, 1994)		N			Epinephelus bleekeri, E. coioides, E. malabaricus		In	KH		[140]
Procamallanus istiblenni (Noble, 1966) Moravec & Sey, 1988		N			Amphiprion clarckii, A. frenatus, A. perideraion, A. polymnus		In	KH		[142]
Superfamily: Habronematoidea Ivaschkin, 1961
Family: Cystidicolidae Skrjabin, 1946
Ascarophis sp.		N			Echeneis naucrates, Epinephelus malabaricus, Gymnocranius griseus		In	GOT, SCS		[81, 114, 115, 140]
Ctenascarophis gastricus Mamaev, 1968		N			Auxis thazard, Euthynnus affinis,		In	SCS		[80]
Prospinitectus mollis (Mamaev, 1968) Petter, 1979		N			Auxis thazard, Euthynnus affinis, Thunnus thynnus,		In	SCS		[80]
Spinitectus echenei Parukhin, 1967		N			Echeneis naucrates		In	SCS		[115]
Spinitectus sp.		N			Auxis thazard		In	SCS		[80]
Superfamily: Physalopteroidea Railliet, 1893
Family: Physalopteridae Railliet, 1893
Cestocephalus petterae (Le-Van-Hoa, Pham-Ngoc-Khue & Nguyen-Thi-Lien, 1972) Moravec & Justine, 2018		N			Polynemus plebeius		In	SCS		[52]
Rasheedia deblocki (Le-Van-Hoa, Pham-Ngoc-Khue & Nguyen-Thi-Lien, 1972) Moravec & Justine, 2018		N			Eleutheronema tetradactylum		In	SCS		[52]
Superfamily: Thelazioidea Skrjabin, 1915
Family: Rhabdochonidae Skrjabin, 1946
Heptochona dorabi (Mamaev, 1968) Moravec, 1975		N			Chirocentrus dorab		In	GOT		[81]
Class: Enoplea Inglis, 1983
Subclass: Dorylaimia Inglis, 1983
Order: Trichinellida Hall, 1916
Superfamily: Trichinelloidea Ward, 1907
Family: Capillariidae Railliet, 1915
Capillaria ariusi (Parukhin, 1989) Arthur & Te, 2006		N			Arius sp.		In	GOTh		[119]
Capillaria sp.		N			Acanthopagrus latus, Epinephelus coioides, Gerres filamentosus		In	GOT, SCS		[81] Present study
Pseudocapillaria (Pseudocapillaria) echenei (Parukhin, 1967) Moravec, 1982		N			Echeneis naucrates		In	SCS		[115, 119]
Nematoda gen. sp.		N			Alectis indicus, Alepes melanoptera, Atropus atropos, A. oreolatus, Atule mate, Carangoides chrysophrys, C. malabaricus, Caranx sp., Chirocentrus dorab, Decapterus muroadsi, Decapterus sp., Echeneis naucrates, Gnathanodon speciosus, Megalaspis cordyla, Psettodes erumei, Rachycentron canadum, Selar crumenophthalmus, Selar sp., Selaroides leptolepis, Seriola dumerili, Seriolina nigrofasciata, Carangidae gen. sp.			GOT, SCS		[19, 114, 119]
Phylum: Acanthocephala Kohlreuther, 1771
Class: Eoacanthocephala Van Cleave, 1936
Order: Neoechinorhynchida Ward, 1917
Family: Neoechinorhynchidae Ward, 1917
Neoechinorhynchus ampullata Amin, Ha & Ha, 2011		A			Megalops cyprinoides		In	GOT		[4]
Neoechinorhynchus (Neoechinorhynchus) ascus Amin, Ha & Ha, 2011		A			Moolgarda seheli		In	GOT		[4]
Neoechinorhynchus (Neoechinorhynchus) dimorphospinus Amin & Sey, 1996		A			Liza subviridis		In	KG		[14]
Neoechinorhynchus (Neoechinorhynchus) johnii Yamaguti, 1939		A			Eleutheronema tetradactylus, Johnius carouna, Johnius sp. Otolithes ruber,		In	VT, BL, KH, QN, QB		[11]
Neoechinorhynchus (Neoechinorhynchus) longinucleatus Amin, Ha & Ha, 2011		A			Strongylura strongylura		In	GOT		[4]
Neoechinorhynchus (Neoechinorhynchus) manubrianus Amin, Ha & Ha, 2011		A			Johnius carouna, Nibea albiflora		In	GOT		[4]
Neoechinorhynchus (Neoechinorhynchus) pennahia Amin, Ha & Ha, 2011		A			Pennahia argentata		In	GOT		[4]
Neoechinorhynchus (Neoechinorhynchus) plaquensis Amin, Ha & Ha, 2011		A			Clupanodon thrissa		In	HP		[90]
Order: Gyracanthocephala Van Cleave, 1936
Family: Quadrigyridae Van Cleave, 1920
Acanthogyrus (Acanthosentis) fusiformis Amin, Chaudhary, Heckmann, Ha & Singh, 2019		A			Arius sp.		In	SCS, BL		[10]
Acanthogyrus (Acanthosentis) indicus Tripathi, 1959		A			Pomadasys argenteus,		In	GOT		[81]
Class: Palaeacanthocephala Meyer, 1931
Order: Echinorhynchida Southwell & Macfie, 1925
Family: Arhythmacanthidae Yamaguti, 1935
Heterosentis holospinus Amin, Heckman & Ha, 2011		A			Plotosus lineatus		In	QN		[5]
Heterosentis mongcai Amin, Heckmann & Ha, 2014		A			Acreichthys sp., Epinephelus sp.		In	QN		[7]
Heterosentis paraholospinus Amin, Heckmann & Ha, 2018		A			Leiognathus equulus, Megalaspis cordyla, Nuchequula flavaxilla		In	BT, KH, QN		[9]
Family: Cavisomidae Meyer, 1932
Filisoma indicum Van Cleave, 1928		A			Scatophagus argus		In	KG		[7]
Neorhadinorhynchus atypicalis Amin & Ha, 2011		A			Siganus fuscescens		In	QN		[3]
Neorhadinorhynchus nudum (Harada, 1938) Yamaguti, 1939		A			Auxis thazard, Decapterus sp., Euthynnus affinis, Thunnus thynnus		In	KH, SCS		[9, 80, 119]
Family: Echinorhynchidae Cobbold, 1879
Echinorhynchus sp.		A			Amphiprion clarckii		In	KH		[142]
Family: Paracanthocephalidae Golvan, 1960
Acanthocephalus halongensis Amin & Ha, 2011		A			Decapterus kurroides		In	NA		[3]
Family: Rhadinorhynchidae Lühe, 1912
Australorhynchus multispinosus Amin, Heckmann & Ha, 2018		A			Fistularia petimba		In	KH		[12]
Cathayacanthus spinitruncatus Amin, Heckmann & Ha, 2014		A			Leiognathus equulus, Nuchequula flavaxilla		In	HP, QN		[50]
Gorgorhynchus medius (Linton, 1908) Chandler, 1934		A			Selar crumenophthalmus		Bc, Io	GOT		[118]
Gorgorhynchus tonkinensis Amin & Ha, 2011		A			Decapterus kurroides		In	QN		[3]
Gorgorhynchus sp.		A			Abalistes stellaris			GOT		[119]
Micracanthorhynchina kuwaitensis Amin & Sey, 1996		A			Strongylura strongylura		In	QN		[3]
Rhadinorhynchus carangis Yamaguti, 1939		A			Carangoides malabaricus		In	GOT, SCS		[73, 118]
Rhadinorhynchus circumspinus Amin, Rubtsova & Nguyen, 2019		A			Triacanthus biaculeatus		In	HP		[13]
Rhadinorhynchus ditrematis Yamaguti, 1939		A			Decapterus sp.		In	GOT, GOTh, SCS		[71, 73]
Rhadinorhynchus dorsoventrospinosus Amin, Heckmann & Nguyen Van Ha, 2011		A			Decapterus maruadsi		In	QN		[6]
Rhadinorhynchus hiansi Soota & Bhattacharya, 1981		A			Sarda orientalis		–	KH		[16]
Rhadinorhynchus johnstoni Golvan, 1969		A			Cypselurus hexazona		–	QB		[7]
Rhadinorhynchus laterospinosus Amin, Heckmann & Nguyen Van Ha, 2011		A			Alectis ciliaris, Auxis rochei, A. thazard, Balistes sp., Harpadon nehereus, Leiognathus equulus, Lutjanus bitaeniatus, Megalaspis cordyla, Nuchequula flavaxilla, Tylosurus sp.		In	QN, HP, BT		[6, 15]
Rhadinorhynchus multispinosus Amin, Rubtsova & Ha, 2019		A			Decapterus maruadsi		In	HP		[13]
Rhadinorhynchus pacificus Amin, Rubtsova & Ha, 2019		A			Auxis thazard		In	HP		[13]
Rhadinorhynchus pristis (Rudolphi, 1802) Lühe, 1911		A			Carangoides malabaricus		In	GOT, SCS		[73, 118]
Rhadinorhynchus trachuri Harada, 1935		A			Auxis thazard, Megalaspis cordyla, Tylosurus sp		In	KH, BT		[2]
Family: Transvenidae Pichelin & Cribb, 2001
Pararhadinorhynchus magnus Van Ha, Amin, Ngo & Heckmann, 2018		A			Scatophagus argus		In	GOT		[50]
Sclerocollum neorubrimaris Amin, Heckmann & Ha, 2018		A			Siganus guttatus		In	KH		[12]
Family: Isthmosacanthidae Smales, 2012
Serrasentis sagittifer (Linton, 1889) Linton, 1932		A			Abalistes stellaris, Atropus atropos, Carangoides malabaricus, Echeneis naucrates, Euthynnus affinis, Gerres filamentosus, Gerres sp., Gymnocranius griseus, Lutjanus russellii, Pomadasys argenteus, Rachycentron cacadum, Sardinella sp., Scomberoides lysan, Trachurus declivis, Triacanthus biaculeatus		Py, In, Bc	GOT, GOTh, SCS		[71, 73, 80, 81, 113, 118, 119]
Acanthocephala gen. sp.		A			Echeneis naucrates, Psettodes erumei			GOT		[109]
Phylum: Annelida Lamarck, 1809
Class: Clitellata Michaelsen, 1919
Subclass: Hirudinea Savigny, 1822
Order: Rhynchobdellida Blanchard, 1894
Family: Piscicolidae Johnston, 1865
Piscicola sp.		H			Lates calcarifer		Gi, Sk	KH		[122]
Zeylanicobdella arugamensis de Silva, 1963		H			Epinephelus bleekeri, E. coioides, E. malabaricus, Lates calcarifer, Lutjanus argentimaculatus		Sk	KH		[139]
Phylum: Arthropoda von Siebold, 1848
Subphylum: Crustacea Brünnich, 1772
Class: Hexanauplia Oakley, Wolfe, Lindgren & Zaharof, 2013
Subclass: Copepoda Milne Edwards, 1840
Order: Cyclopoida Burmeister, 1834
Family: Ergasilidae Burmeister, 1835
Ergasilus sp.		Cr			Epinephelus coioides, E. malabaricus		Gi	KH		[139]
Order: Siphonostomatoida Burmeister, 1835
Family: Caligidae Burmeister, 1835
Abasia sp.			Cr			Saurida tumbil	Gi		GOT		[61]
Anuretes branchialis Rangnekar, 1953			Cr			Sarda sp.	–		–		[61]
Caligus arii Bassett-Smith, 1898			Cr			Arius sp	–		–		[60]
Caligus bonito Wilson, 1905			Cr			Euthynnus affinis	–		–		[61]
Caligus confusus Pillai, 1961			Cr			Abalistes stellatus, Decapterus sp.	Gi		GOT		[61]
Caligus constrictus Heller, 1865			Cr			Decapterus sp.	Gi		GOT		[61]
Caligus epidemicus Hewitt, 1971			Cr			Epinephelus bleekeri, E. coioides, E. malabaricus, E. tauvina, Lates calcarifer	Sk		KH		[122, 136]
Caligus fortis Kabata, 1965			Cr			Abalistes stellatus	–		–		[61]
Caligus multispinosus Shen, 1957			Cr			Pampus argenteus	–		–		[61]
Caligus pelamydis Krøyer, 1863			Cr			Sphyraena jello	–		–		[61]
Caligus robustus Bassett-Smith, 1898			Cr			Decapterus sp.	Gi		GOT		[61]
Caligus stromatei Krøyer, 1863			Cr			Epinephelus bleekeri, E. coioides, E. malabaricus, E. tauvina, Lates calcarifer	Gi		KH		[122, 136]
Caligus sp.			Cr			Acanthopagrus latus, Epinephelus bleekeri, E. coioides	Gi		KH, QN		[133, 140] Present study
Caligodes laciniatus (Krøyer, 1863)			Cr			Ablennes hians	–		–		[61]
Hermilius pyriventris Heller, 1865			Cr			Unidentified Marine catfish	–		–		[61]
Mappates plataxus Rangnekar, 1958			Cr			Platax teira, Sarda sp.	–		–		[61]
Lepeophtheirus atypicus Lin, Ho & Chen, 1996			Cr			Siganus fuscescens	–		–		[61]
Lepeophtheirus longipalpus Bassett-Smith, 1898			Cr			Arius acutirostris	–		–		[61]
Lepeophtheirus sp.			Cr			Epinephelus tauvina*	Sk		GOT		[34]
Parapetalus hirsutus (Bassett-Smith, 1898)			Cr			Eleutheronema tetradactylum	–		–		[61]
Parapetalus occidentalis Wilson, 1908			Cr			Sphyraena jello	–		–		[61]
Sinocaligus formicoides (Redkar, Rangnekar et Murti, 1949)			Cr			Dussumieria elopsoides	–		–		[61]
Synestius caliginus Steenstrup et Lutken, 1861			Cr			Parastromateus niger	–		–		[61]
Parapetalus sp.			Cr			Rachycentron canadum	Gi		KH		[53]
Family: Hatschekiidae Kabata, 1979
Hatschekia foliolata Redkar, Rangnekar & Murti, 1950			Cr			Parastromateus niger, Nemipterus peronii					[61]
Hatschekia hanguyenvani Kovaleva, Nguyen et Ngo, 2017			Cr				Gi		GOT		[62]
Hatschekia rotundigenitalis Yamaguti, 1939			Cr			Unidentified marine host	–		–		[61]
Hatschekia sp.			Cr			Amphiprion polymnus	Gi		KH		[142]
Pseudocongericola sp.			Cr			Congresox talabonoides	–		–		[61]
Family: Kroyeriidae Kabata, 1979
Kroyeria spatulata Pearse, 1948			Cr			Carcharhinus sorrah	–		–		[61]
Family: Lernaeopodidae Milne Edwards, 1840
Charopinopsis quaternia (Wilson C.B., 1935)			Cr			Scomberoides lysan	–		–		[61]
Parabrachiella trichiuri (Yamaguti, 1939)			Cr			Lutjanus erythropterus	Gi		GOT		[61]
Neobrachiella sp.			Cr			Valamugil engeli	–		–		[61]
Naobranchia sp.						Gerres filamentosus	–		–		[61]
Family: Lernanthropidae Kabata, 1979
Chauvanium chauvani Kazatchenko, Kovaleva, Nguyen and Ngo, 2017			Cr			Alepes melanoptera	Gi		GOT		[61]
Lernanthropus alatus Pillai, 1964			Cr			Alepes melanoptera, Caranx sp., Decapterus sp.	–		–		[61]
Lernanthropus carangis Pillai, 1964			Cr			Upeneus sulfureus	–		–		[61]
Lernanthropus chirocentrosus Tripathi, 1959			Cr			Chirocentrus dorab	–		–		[61]
Lernanthropus cornutus Kirtisinghe, 1937			Cr			Ablennes hians	–		–		[61]
Lernanthropus francai Nunes-Ruivo, 1962			Cr			Larimichthys croceus	–		–		[61]
Lernanthropus latis Yamaguti, 1954			Cr			Lates calcarifer	Gi		KH		[140]
Lernanthropus lappaceus Wilson, 1912			Cr			Eleutheronema tetradactylum, Arius maculatus	–		–		[61]
Lernanthropus opisthopteri Pillai, 1964			Cr			Ilisha elongata	–		–		[61]
Lernanthropus otolithi Pillai, 1963			Cr			Johnius carouna	–		–		[61]
Lernanthropus polynemi Richiardi, 1881			Cr			Unidentified host	–		–		[61]
Lernanthropus sanguineus Song, 1976.			Cr			Lutjanus johnii	–		–		[61]
Lernanthropus trifoliatus Bassett-Smith, 1898			Cr			Arius maculatus, Arius sp., Polydactylus sextarius	–		–		[61]
Lernanthropus villiersi Delamare-Deboutteville et Nunes-Ruivo, 1954			Cr			Gerres filamentosus	–		–		[61]
Lernanthropinus decapteri (Pillai, 1964) Pillai, 1967			Cr			Decapterus maruadsi, Elisha filigera	–		–		[61]
Lernanthropinus gibbosus (Pillai, 1964)			Cr			Saurida tumbil	–		–		[61]
Lernanthropinus sphyraenae (Yamaguti et Yamasu, 1959)			Cr			Mene maculata	–		–		[61]
Sagum sanguineus (Song, 1976)			Cr			Lutjanus johnii	–		–		[61]
Sagum vietnamiensis Kazatchenko, Kovaleva, Nguyen and Ngo, 2017			Cr			–	Gi		GOT		[61]
Family: Pseudocycnidae Wilson C.B., 1922
Cybicola armatus (Bassett-Smith, 1898) Kensley et Grindley, 1973			Cr			Sphyraena jello, Euthynnus alleteratus, Scomberomorus commerson	–		–		[61]
Pseudocycnus appendiculatus Heller, 1865			Cr			Auxis thazard, Euthynnus affinis, E. alleteratus	–		–		[61]
Family: Trebiidae Wilson C.B., 1905
Trebius elongatus Capart, 1953			Cr			Taeniura meyeni	–		–		[61]
Class: Malacostraca Latreille, 1802
Order: Isopoda Latreille, 1817
Family: Corallanidae Hansen, 1890
Alcirona sp.		Cr			Epinephelus coioides		Sk	QN, HP		[133]
Corallana sp.		Cr			Epinephelus tauvina*		Sk	GOT		[34]
Family: Cymothoidae Leach, 1818
Ceratothoa verrucosa (Schioedte & Meinert, 1883)		Cr			Epinephelus coioides		Gi	KH		[135]
Family: Gnathiidae Leach, 1814
Gnathia sp.		Cr			Epinephelus bleekeri, E. coioides		Gi	KH		[133, 140]
Gnathiidae gen. sp.		Cr			Epinephelus coioides		Gi	GOT		[133]

This species has not been registered in WoRMS and not confirmed by international experts.

Prosorhynchus sp. A & Prosorhynchus sp. B have not been identified, but they are distinct based on some morphological characters.

This species has not been confirmed by international experts.

Dollfusiella sp. A and B were not identified to species level, but it was possible to distinguish them based on morphology.

Temporary name: This species has not been registered in WoRMS and not confirmed by international experts.

Philometra sp. 1 and 2 were not fully identified, but it was possible to distinguish them based on morphology.

Abbreviations

Taxon names: A: Acanthocephala; C: Cestoda; Ci: Ciliophora; Cr: Crustacea; D: Digenea; H: Hirudinea; Mo: Monogenea; My: Myxozoa; N: Nematoda.

Sites of infection: Bc: Body cavity; Bvg: Blood vessel gill; Fi: Fin; Gb: Gall bladder; Gi: Gill; Gic: Gill cavity; Go: Gonad; He: Heart; Ht: Head tissues; In: Intestine; Io: Inner organs; Iw: Intestine wall; K: Kidney; L: Liver; Mu: Muscle; O: Oesophagus; Oe: Orbit of the eye; Op: Operculum; Phc: Pharyngeal cavity; Py: Pyloric caeca; S: Stomach; Sb: Swimming bladder; Sk: Skin; Sw: Stomach wall; Sv: Spiral valve; Ub: Urine bladder; Us: Under scales; Ve: Vitreous eye.

*Before 2005, Epinephelus coioides was misidentified as E. tauvina.

Parasite richness

A total of 498 parasite species have been recorded from 225 marine fish in Vietnam belonging to the following taxa: Myxozoa and Ciliophora (8 each), Monogenea (117), Digenea (214), Cestoda (17), Nematoda (37), Acanthocephala (37), Hirudinea (2), and Crustacea (58), demonstrating the high diversity of the marine fish parasites inhabiting Vietnamese waters. The current average number of parasite species per fish species was 2.2, which was lower than the previously estimated species richness in Vietnamese (3.0) and German coastal waters (3.1) [19, 104]. This lower species richness appears to be due to a higher proportion of the fish community being investigated compared to 16 years ago (225 species vs. 82 fish species reported in 2006) so that a better prediction of the species richness can be made. It could also be explained by a decline in host species richness and population density, the major universal determinants of variations in parasite species richness [123]. However, the new figure of 2.2 is comparable to that of New Caledonia (1.9), the Indo-West Pacific (1.7), and Hawaii’s open Pacific waters (2.2) ([58, 103, 126], respectively). These regions’ similar latitudinal patterns could account for their parasite richness [28].

Our findings showed that only 225 of the total 1876 species of marine fish recorded in Vietnam [21] had been investigated for parasite species, accounting for only 12% of the entire fish community; this indicates that the parasite fauna of marine fishes in Vietnam is still poorly known.

Parasite composition and proportion in marine fish of Vietnam

The present results showed that the proportion of marine fish parasites observed in Vietnam is similar to that previously described in Hawaiian waters ([103]; Fig. 2b). As mentioned, the proximity of the latitudinal range between Vietnam and Hawaii might be the reason for such similarities in the proportion of the parasite taxa.

Figure 2

Proportion (%) of (a) the recorded fish parasite species in Vietnam (498 species, present study), and (b) fish parasites species in Hawaii (652 species [103]).

A map of Vietnam with abbreviated names of municipalities, provinces, and sea or ocean areas where samples were collected.

Digenea and Monogenea were the most common taxa, accounting for 43% and 23.5% of Vietnam’s total marine fish parasite fauna, respectively. The overall taxon proportions are similar to those found in Hawaii, except for Nematoda and Acanthocephala, which were higher in Vietnam than in Hawaii (7.4% and 7.4% in Vietnam vs. 3.1% and 1.4% in Hawaii, respectively). Many factors can influence the richness of these two parasite taxa, e.g., the zooplankton (e.g., Copepoda, Amphipoda), which may act as intermediate hosts for Nematoda and Acanthocephala and were reported to be more diverse and abundant in Vietnam than in Hawaii [56, 87]. Furthermore, since acanthocephalans use amphipods or ostracods more frequently as intermediate hosts, they are abundant in shallow waters in Ha Long Bay but not on the Hawaiian coast [56, 87]; these parasites are typically found in benthic and benthopelagic fish final hosts [54]. As a result, when compared to the 2006 list, 29 additional acanthocephalans have been added to the current local list [3–13, 50], 22 of which are new parasite species that have never been recorded anywhere other than Vietnam. Those 22 species are Acanthocephalus halongensis Amin & Ha, 2011; Acanthogyrus (Acanthosentis) fusiformis Amin, Chaudhary, Heckmann, Ha & Singh, 2019; Neoechinorhynchus ampullata Amin, Ha & Ha. 2011; Neoechinorhynchus ascus Amin, Ha & Ha, 2011; Neoechinorhynchus longinucleatus Amin, Ha & Ha, 2011; Neoechinorhynchus manubrianus Amin, Ha & Ha, 2011; Neoechinorhynchus pennahia Amin, Ha & Ha, 2011; Neoechinorhynchus plaquensis Amin, Ha & Ha, 2011; Heterosentis holospinus Amin, Heckmann & Ha, 2011; Heterosentis mongcai Amin, Heckmann & Ha, 2014; Heterosentis paraholospinus Amin, Heckmann & Ha, 2018; Australorhynchus multispinosus Amin, Heckmann & Ha, 2018; Cathayacanthus spinitruncatus Amin, Heckmann & Ha, 2014; Rhadinorhynchus circumspinus Amin, Rubtsova & Nguyen, 2019; Rhadinorhynchus dorsoventrospinosus Amin, Heckmann & Ha, 2011; Rhadinorhynchus laterospinosus Amin, Heckmann & Ha, 2011; Rhadinorhynchus pacificus Amin, Rubtsova & Ha, 2019; Rhadinorhynchus multispinosus Amin, Rubtsova & Ha, 2019; Neorhadinorhynchus atypicalis Amin & Ha, 2011, Pararhadinorhynchus magnus Ha et al., 2018; Gorgorhynchus tonkinensis Amin & Ha, 2011, and Sclerocollum neorubrimaris Amin, Heckmann & Ha, 2018. These findings suggest that prior to 2006, acanthocephalans were neglected in the parasitological studies regarding marine fish from Vietnam.

To date, it is known that the Cestoda, particularly the orders Trypanorhyncha and Tetraphyllidea, are found in a wide range of marine fish species worldwide [101, 103]. They are tapeworms from elasmobranchs that have been considered excellent indicators of host ecology [105]. In Vietnam, cestodes accounted for only 3.4% of the total parasite fauna in marine fish (17 out of 498 species; Fig. 2a), a much lower proportion than in Hawaii (9.2% represented by 60 species; [103]; Fig. 2b); this could be because of a lack of definitive hosts (elasmobranchs) due to shallow waters or, more likely, a lack of research interest in these parasites in Vietnam.

The hirudinean parasites were in very low proportion compared to other parasite taxa. These annelids have one of the largest body sizes among the groups of parasites in marine fish and are well known to be pathogenic in farmed finfish [40]. Moreover, hirudineans occur mainly in freshwater [134] which may explain their low proportion of occurrence in marine fish observed here (0.4%; Fig. 2a). This low proportion was also observed by Palm and Bray [103] in Hawaiian waters (0.5%; Fig. 2b).

In comparison to data from Hawaii [103], Crustacea were found in similar proportions in Vietnamese marine fish (11.6% vs. 10.3%; Fig. 2a), but Myxozoa proportions were higher (1.6% vs. 0.9%), and no Ciliophora were found in Hawaii, while there was 1.6% in Vietnam. However, myxozoans have recently been reported in a low proportion in Vietnamese marine waters (0.09% equal to 2 species) when compared to other groups. However, Mackenzie and Kalavati [79] reported 223 myxosporean parasites from North Pacific marine fish, identifying them as one of the most common parasite groups in marine fish [79]. Thus, the true diversity of myxozoans in Vietnam is expected to be much higher than the current findings. The lack of information and data for this group could be attributed to rudimentary sampling methods and a lack of well-trained personnel to investigate this parasite group. Similarly, the Ciliophora, a worldwide distributed parasite of aquatic teleosts, was only found in small numbers. Depending on their host species, this taxon can range from harmless ectocommensals to dangerous parasites in fish aquaculture [79]. Bui [34] and Phan [121] reported Ciliophora as parasites causing disease in Vietnam’s finfish mariculture. These parasites that infected marine fish in Vietnam were most likely reported as multi-species groups (e.g., Trichodina spp.), making identification difficult. This means that the actual number of ciliates in Vietnamese marine waters may be much higher than the eight species found so far.

Parasitic crustacean species accounted for 11.6% of total marine fish parasites reported in Vietnam thus far, slightly higher than the figure found in Hawaiian marine waters (10.3%; Fig. 2). This parasite group has been identified as the most diverse and widespread subphylum of arthropods in marine fish [29, 126]. They have also been identified as critical pathogenic agents of cultured marine fish in the Asian-Pacific region [77]. Given the critical role that disease control plays in ensuring the success of aquaculture, further research on parasites is both necessary and warranted.

Parasite composition and proportion in the Gulf of Tonkin

The current study presents 330 marine fish parasite species in the GOT. They are classified into eight taxa, Myxozoa (3), Ciliophora (3), Monogenea (89), Digenea (168), Cestoda (11), Nematoda (19), Acanthocephala (29), and Crustacea (8), demonstrating a high number and rich species composition in the GOT. The parasite richness in the GOT may be influenced by several factors, including geographical latitude, the environment, and the availability of intermediate hosts [55, 85]. The GOT is in South-East Asia, a region with a diverse range of parasites and hosts [125, 126], including the first intermediate host, mollusks [91], which may be the primary factor affecting endoparasite diversity, such as the digeneans (168 species) we discovered here. However, the GOT’s shallow water depth and muddy sediment limit the distribution of open-water elasmobranchs (e.g., sharks and rays), the expected final hosts of marine Cestoda, particularly Trypanorhyncha (Fig. 3). As a result, only seven Trypanorhyncha species have been identified in the GOT so far.

Figure 3

Marine fish parasites in the Gulf of Tonkin: (a) Proportion of the total recorded fish parasite species (%), and (b) Composition (species richness) of parasite species only found in the Gulf of Tonkin.

Most of the recorded marine parasite species in Vietnam (66.3%) have been found in the GOT, indicating that the GOT is a hotspot for interested collectors and institutions supporting research due to its high parasite biodiversity. However, apart from its natural biodiversity, most parasitological research may be concentrated in the GOT due to the urgency of fish disease research serving the high-density aquaculture area in the GOT or because conducting research here is more convenient than in other regions. The composition related to the parasite fauna in the GOT was generally similar to that of all Vietnamese marine environments (Figs. 2a and 3a). Digenea were proportionally the taxon in the GOT with the highest species richness (51%; Fig. 3a). This proportion in the GOT was higher than that observed for the whole of Vietnam (this study; Fig. 2a) and similar to that reported in Hawaii ([103], Fig. 2b). A possible explanation for the previous results relates to the environmental features of the GOT, i.e., muddy and shallow waters with rocky and limestone substrates [89, 120] which support an abundant fauna of mollusks that act as intermediate hosts for digenetic trematodes. This is also consistent with the findings of Sujatha and Madhavi [128] and Bray et al. [31], which indicated that the Digenea were typically shallow-water parasites.

Other major parasite groups in the GOT reported in this study (Fig. 3a) include Monogenea (27%), Acanthocephala (8.8%), Nematoda (5.8%), Cestoda (3.3%), Crustacea (2.4%), Ciliophora (0.9%), and Myxozoa (0.9%). It is worth noting that, of the 330 species of marine fish parasites discovered in the GOT, up to 235 species (71%) are found only here and nowhere else in Vietnam (Fig. 3b). For example, the current effort revealed that up to 21/29 acanthocephalans are found only in the GOT (Fig. 3b), suggesting that they are GOT native species.

The parasite-host ratio of 2.7 was calculated based on a total of 330 parasite species that have been recorded in 122 marine fish species in the GOT. This ratio is higher than the average for Vietnamese marine water (2.2), Hawaii (2.2), New Caledonia (1.9), and the Indo–West Pacific (1.7) [58, 103, 126]. Since the GOT is thought to have around 928 marine fish species [144], only a tiny portion of the fish species has been examined for parasites (13.1%), indicating that more attention needs to be paid to exploring the region’s high diversity of marine fish parasites.

Conclusions

This is a comprehensive review to understand the diversity and richness of Vietnam’s marine parasite fauna. In 225 fish species, 498 marine fish parasite species have been identified, compared to 247 parasites in 82 fish species in 2006. Digenea (43%) and Monogenea (23.5%) had the highest levels of species richness. According to the data currently available, most parasites in Vietnamese fish come from the GOT (330 species, equivalent to 66.3%).

In Vietnam, the estimated marine parasite richness decreased from 3.0 in 2006 to 2.2 in 2022. Only 12% of the marine fish species have been studied for parasites, so a large part of the fish community still needs to be studied to evaluate and predict better parasite richness and diversity in this tropical-subtropical country. This compilation of parasite records shows significant progress in studying marine fish parasites in Vietnam over the last 16 years. However, it is still in its infancy, leaving a sizable task for the future, as species classification is the first critical step in characterizing any ecosystem. It is a challenging but fascinating task to learn about evolutionary biology and the history of nature, while discovering the true diversity of these marine ecosystems. Understanding pathogenic threats is critical for Vietnam’s growing finfish mariculture industry. Recent research shows that parasites can be used to study climate change and the environment. Thus, studying marine fish parasites in Vietnam is urgently needed in the future, especially using molecular data to characterize and classify the fauna.

The findings of this study will help create a database for marine fish parasites in Vietnam. Additionally, it promotes aquaculture success in Vietnam, reduces the risk of fish disease transmission between countries, and reduces the risk of consuming parasite-infected fishery and aquaculture products in Vietnam and elsewhere.