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Corresponding author: Hsuan-Ching Ho ( ogcoho@gmail.com ) Corresponding author: Chien-Hsiang Lin ( chlin.otolith@gmail.com ) Academic editor: Barry C Russell
© 2024 Shing-Lai Ng, Hsin-Wei Liu, Dominique P. Mediodia, Yen-Ting Lin, Chieh-Hsuan Lee, Ching-Feng Lin, Shih-Pin Huang, Siao-Man Wu, Chih-Ren Tung, Hsuan-Ching Ho, Chien-Hsiang Lin.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Ng S-L, Liu H-W, Mediodia DP, Lin Y-T, Lee C-H, Lin C-F, Huang S-P, Wu S-M, Tung C-R, Ho H-C, Lin C-H (2024) An updated checklist of fishes of Dongsha Island, Taiwan, northern South China Sea. In: Ho H-C, Russell B, Hibino Y, Lee M-Y (Eds) Biodiversity and taxonomy of fishes in Taiwan and adjacent waters. ZooKeys 1220: 175-242. https://doi.org/10.3897/zookeys.1220.131100
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Dongsha Island, situated in the northern part of the South China Sea, is surrounded by coral reefs and deep-sea habitats. The coastal areas of the atoll, a marine protected area, serve as important nursery habitats for many reef fish species. At the same time, the offshore deep-sea continental slopes are historically important fishing grounds. Although previous inventories primarily focused on coral reef fishes within the atoll listing 652 species from 73 families, comprehensive surveys of fishes from deeper waters have been incomplete. In this study, the species composition of the fish fauna around Dongsha was updated by analyzing large volumes of frozen bycatch from commercial deep-sea trawlers operating in the area for the past four years and reviewing the existing literature. The species list increased to 1087 species belonging to 167 families, including several documented as new records and potentially undescribed species. This updated checklist also includes images of each species and most of their sagittal otoliths. This will assist further taxonomic work and significantly enhance understanding of marine biodiversity in the South China Sea.
Biodiversity, ichthyofauna, ichthyology, Pratas Island, taxonomy
The South China Sea is an arm of the western Pacific Ocean that borders the Southeast Asian mainland. Renowned for its significant marine biodiversity (
Besides its coral reef ecosystems and associated fishes inhabiting Dongsha Atoll, the surrounding waters are also areas for commercial deep-sea fishing. Notably, the commercial trawl fishery, active in the deeper slope areas since the early 2000s, yields high volumes of deep-sea organisms. These catches, including a wide range of fish and invertebrate species (
The purpose of this paper is to review and compile existing data and literature to provide a comprehensive overview of the fish fauna in and around Dongsha Island. While the list of coral and seagrass-associated fishes is derived from previous surveys outside of the marine protected area (Fig.
Our study used two distinct datasets: occurrence records from the literature on coral and seagrass-associated shallow-water fishes and newly collected occurrence data from deep-water fishes sampled via commercial bottom trawlers. Both datasets represent different ecological settings with distinct sampling techniques. For the former, we reviewed existing literature (
Each fish was measured for standard length (SL), total length (TL), and weight (WT), with preanal length (PAL) recorded when necessary. Specimens were photographed before muscle tissue was subsampled from the right caudal region. These tissue samples were preserved in 95% ethyl alcohol and deposited at the National Academy of Marine Research in Kaohsiung for future reference.
For images of teleostean otoliths, the left sagittal otoliths were primarily used; right otoliths were reversed for consistency and noted with an (R) in each caption (
The occurrence dataset listed below was uploaded to the Global Biodiversity Information Facility (GBIF) and was published (
Apristurus nakayai Iglésias, 2012
Fig.
This species was originally described from New Caledonia, with one record from Papua New Guinea, and was recently recorded around Dongsha Island (
Iago garricki Fourmanoir & Rivaton, 1979
Fig.
This species is restricted to the tropical western Pacific, and is recently redescribed based on five specimens around Dongsha Island (
Etmopterus lii Ng, Liu & Joung, 2024
Fig.
This species was recently described from the northern South China Sea, where we collected our samples. Juveniles are more common around Dongsha Island, while adults are rare, suggesting that the latter may inhabit deeper waters.
Okamejei picta Ng, Ho, Joung & Liu, 2023
Fig.
This species was described around Dongsha Island, based on two specimens (
Images of fresh specimens collected around Dongsha Island A Chimaera phantasma, 511 mm TL B Hydrolagus mitsukurii, 633 mm TL C Sinobatis borneensis, 338 mm TL D Notoraja tobitukai, 448 mm TL E Dipturus gigas, 404 mm TL F Dipturus tengu, 400 mm TL G Okamejei sp., 266 mm TL H Okamejei picta, 429 mm TL I Dipturus wuhanlingi, 753 mm TL J Tetronarce tokionis, 576 mm TL K Plesiobatis daviesi, 801 mm TL. Scale bars: 5 cm.
Images of fresh specimens collected around Dongsha Island A Cirrhoscyllium formosanum, 380 mm TL B Dichichthys melanobranchus, 320 mm TL C Apristurus herklotsi, 413 mm TL D Apristurus macrostomus, 361 mm TL E Apristurus nakayai, 421 mm TL F Galeus eastmani, 342 mm TL G Galeus nipponensis, 566 mm TL H Galeus sauteri, 408 mm TL I Cephaloscyllium fasciatum, 356 mm TL J Iago garricki, 515 mm TL. Scale bar: 5 cm.
Images of fresh specimens collected around Dongsha Island A Etmopterus bigelowi, 261 mm TL B Etmopterus brachyurus, 268 mm TL C Etmopterus decacuspidatus, 270 mm TL D Etmopterus lii, 296 mm TL E Etmopterus cf. molleri, 264 mm TL F Etmopterus lucifer, 287 mm TL G Heptranchias perlo, 383 mm TL H Centrophorus tessellatus, 613 mm TL I Squalus montalbani, 371 mm TL J Squalus japonicus, 410 mm TL K Zameus squamulosus, 356 mm TL L Pristiophorus japonicus, 622 mm TL. Scale bars: 5 cm.
Ophichthus kusanagi Hibino, McCosker & Tashiro, 2019
Fig.
This species was described from Japan and reported around Dongsha Island (
Ophichthus pratasensis Ho, Ng & Lin, 2022
Figs
This species was described around Dongsha Island (
Congriscus maldivensis (Norman, 1939)
Figs
This species is widespread in the Indo-West Pacific and was recently recorded by
Lestrolepis philippina (Fowler, 1934)
Figs
This species was commonly misidentified as Lestrolepis japonica over the past few decades (e.g.,
Kuronezumia macronema (Smith & Radcliffe, 1912)
Figs
This species was originally described from the Philippines and was recorded around Dongsha Island (
Amarsipus carlsbergi Haedrich, 1969
Figs
This species is widespread in the Indo-West Pacific and was reported as a new record around Dongsha Island by
Synagrops atrumoris Mediodia & Lin, 2024
Figs
This recently described new species (
Lophiodes lugubris (Alcock, 1894)
Figs
This species is widely distributed in the Indo-West Pacific and was recently redescribed by
Lophiodes triradiatus (Lloyd, 1909)
Figs
This deepwater species was only found around Dongsha Island and was not recorded around Taiwan.
Eptatretus sheni (Kuo, Huang & Mok, 1994)
Fig.
This species was described from Taiwan. Though common around Taiwan, it was not reported in other regions. Our records around Dongsha Island show a substantial range extension. This species is rare around Dongsha Island.
Eptatretus taiwanae (Shen & Tao, 1975)
Fig.
This species was described from Taiwan and is common in the area. It was recognized as endemic. Our first record around Dongsha Island suggests a broader distribution range. This species is rare around Dongsha Island.
Rubicundus rubicundus (Kuo, Lee & Mok, 2010)
Fig.
This attractive cuskeel species was described from Taiwan and is considered endemic. The sole specimen we collected represents the first record around Dongsha Island. This species is very rare around Dongsha Island.
Centrophorus tessellatus Garman, 1906
Fig.
Some species of the genus Centrophorus are very difficult to identify, including a long-snout species group. Within this group, two species occur in the northwestern Pacific, C. tessellatus and C. isodon Chu, Meng & Liu, 1981. We tentatively identified the long-snout specimens as C. tessellatus, which is not uncommon in southwestern Taiwan and is recorded in Japan. This species is uncommon around Dongsha Island, and most specimens are juveniles.
Etmopterus bigelowi Shirai & Tachikawa, 1993
Fig.
This species is one of the circumglobal lanternshark species, yet records from the northwestern Pacific were limited to off Japan. Five specimens were collected in our study, confirming their occurrence around Dongsha Island, South China Sea.
Squalus montalbani Whitley, 1931
Fig.
One of the widespread spurdog species, having records from northeastern Taiwan to northern Australia. Yet, no records are known from the South China Sea. This species is common around Dongsha Island. Large individuals are sold separately.
Atractodenchelys brevitrunca Vo & Ho, 2020
Figs
This species was recently described in central Vietnam by
Images of fresh specimens collected around Dongsha Island A Aldrovandia affinis, 188.65 mm TL B Dysomma polycatodon, 241.19 mm TL C Dysomma anguillare, 225.71 mm TL D Dysommina orientalis, 290.99 mm TL E Dysomma dolichosomatum, 290.86 mm TL F Synaphobranchus kaupii, 330.85 mm TL G Synaphobranchus oligolepis, 252.15 mm TL H Synaphobranchus affinis, 215.86 mm TL I Atractodenchelys brevitrunca, 605.05 mm TL. Scale bar: 5 cm.
Dysomma polycatodon Karrer, 1983
Figs
This species is widely distributed on continental shelf in the Indo-West Pacific, and is common around Taiwan. Only one specimen was recorded around Dongsha Island, suggesting the shallower habitat of this species.
Otolith images of fresh specimens collected around Dongsha Island A Aldrovandia affinis, CHLOL 20601, 204.25 mm TL B Synaphobranchus oligolepis, CHLOL 23597, 275.98 mm TL C Synaphobranchus kaupii, CHLOL 23598, 330.85 mm TL (R) D Synaphobranchus affinis, CHLOL 26162, 215.86 mm TL E Dysomma anguillare, CHLOL 25422, 225.71 mm TL (R) F Dysomma dolichosomatum, CHLOL 20664, 322.95 mm TL G Dysomma polycatodon, CHLOL 9815, 241.19 mm TL H Dysommina orientalis, CHLOL 20492, 294.48 mm TL I Atractodenchelys brevitrunca, CHLOL 25213, 494.47 mm TL. Scale bar: 1 mm.
Synaphobranchus oligolepis Ho, Hong & Chen, 2018
Figs
This species was recently described around Taiwan. It is common near the type locality but rare around Dongsha Island.
Coloconger maculatus Ho & Tang, 2021
Figs
This species was recently described around Taiwan based on a single specimen. Our specimen collected around Dongsha Island is the second known specimen.
Coloconger raniceps Alcock, 1889
Figs
This species is widespread in the Indo-West Pacific, but is not well presented in museums. It is rare around Dongsha Island.
Ophichthus megalops Asano, 1987
Figs
This species was originally described from Japan. It is occasionally found in bycatches of bottom trawl around Taiwan, but it is rare around Dongsha Island.
Images of fresh specimens collected around Dongsha Island A Pisodonophis boro, 476.99 mm TL B Neenchelys sp., 311.24 mm TL C Ophichthus obtusus, 467.93 mm TL D Ophichthus sp., 448.31 mm TL E Ophichthus pratasensis, 860.23 mm TL F Ophichthus urolophus, 549.79 mm TL G Ophichthus kusanagi, 504.80 mm TL H Ophichthus megalops, 592.09 mm TL I Nettastoma solitarium, 557.13 mm TL. Scale bar: 5 cm.
Otolith images of fresh specimens collected around Dongsha Island A Ophichthus pratasensis, CHLOL 20629, 495.21 mm TL B Ophichthus megalops, CHLOL 22042, 592.09 mm TL C Ophichthus sp., CHLOL 17929, 442.76 mm TL D Ophichthus urolophus, CHLOL 9821, 577.00 mm TL E Ophichthus obtusus, CHLOL 19481, 453.22 mm TL F Pisodonophis boro, CHLOL 21208, 476.99 mm TL G Nettastoma solitarium, CHLOL 29404, 648.86 mm TL H Coloconger maculatus, CHLOL 20620, 171.06 mm TL I Coloconger raniceps, CHLOL 27101, 207.50 mm TL J Coloconger scholesi, CHLOL 27100, 191.54 mm TL. Scale bar: 1 mm.
Ophichthus obtusus McCosker, Ide & Endo, 2012
Figs
This species is restricted to the northwestern Pacific and has records from Japan, Taiwan, and Vietnam. It is common around Dongsha Island and Taiwan.
Acromycter nezumi (Asano, 1958)
Figs
This species is restricted to the northwestern Pacific. It is common around Dongsha Island but rare around Taiwan.
Images of fresh specimens collected around Dongsha Island A Congriscus megastoma, 348.09 mm TL B Congriscus maldivensis, 312.06 mm TL C Nemichthys scolopaceus, 507.37 mm TL D Macrocephenchelys brachialis, 461.81 mm TL E Macrocephenchelys brevirostris, 36.052 mm TL F Acromycter nezumi, 272.56 mm TL G Blachea xenobranchialis, 227.95 mm TL H Gnathophis heterognathos, 183.87 mm TL I Ariosoma meeki, 367.31 mm TL J Ariosoma emmae, 329.76 mm TL K Gavialiceps taiwanensis, 690.19 mm TL. Scale bar: 5 cm.
Ariosoma emmae Smith & Ho, 2018
Figs
This species was described from southwestern Taiwan. It is rare around Dongsha Island.
Otolith images of fresh specimens collected around Dongsha Island A Congriscus megastoma, CHLOL 27848, 307.35 mm TL B Congriscus maldivensis, CHLOL 27847, 217.73 mm TL C Bathycongrus wallacei, CHLOL 20479, 333.05 mm TL D Bathycongrus bleekeri, CHLOL 19487, 179.51 mm TL E Bathycongrus sp., CHLOL 20668, 315.03 mm TL F Bathycongrus retrotinctus, CHLOL 22039, 300.22 mm TL G Ariosoma meeki, CHLOL 20626, 326.77 mm TL H Macrocephenchelys brevirostris, CHLOL 9816, 360.52 mm TL I Ariosoma emmae, CHLOL 21649, 302.42 mm TL. Scale bar: 1 mm.
Bathycongrus bimaculatus Smith & Ho, 2018
Fig.
This species was described from southwestern Taiwan. The sole specimen around Dongsha Island, suggests a broader distribution in the South China Sea.
Images of fresh specimens collected around Dongsha Island A Bathycongrus retrotinctus, 254.48 mm TL B Bathycongrus wallacei, 273.51 mm TL C Bathycongrus sp., 275.39 mm TL D Bathycongrus bimaculatus, 153.61 mm TL E Bathycongrus bleekeri, 179.51 mm TL F Bathyuroconger cf. vicinus, 378.46 mm TL G Bathyuroconger parvibranchialis, 729.12 mm TL H Bathyuroconger albus, 127.79 mm TL I Bathyuroconger sp., 494.24 mm TL J Bathyuroconger fowleri, 375.93 mm TL. Scale bar: 5 cm.
Bathycongrus bleekeri Fowler, 1934
Figs
This species is restricted to the northwestern Pacific. It is rare around Dongsha Island.
Bathyuroconger parvibranchialis (Fowler, 1934)
Figs
This species was originally described from the Philippines. It was recently documented in Taiwan by
Otolith images of fresh specimens collected around Dongsha Island A Acromycter nezumi, CHLOL 20456, 208.92 mm TL B Gavialiceps taiwanensis, CHLOL 21638, 620.48 mm TL C Blachea xenobranchialis, CHLOL 22753, 257.88 mm TL D Macrocephenchelys brachialis, CHLOL 25636, 461.81 mm TL E Gnathophis heterognathos, CHLOL 9812, 183.87 mm TL F Bathyuroconger cf. vicinus, CHLOL 27098, 378.46 mm TL G Bathyuroconger fowleri, CHLOL 22668, 234.89 mm TL H Bathyuroconger albus, CHLOL 22038, 127.79 mm TL I Bathyuroconger parvibranchialis, CHLOL 22077, 729.12 mm TL J Bathyuroconger sp., CHLOL 16037, 494.24 mm TL. Scale bar: 1 mm.
Blachea xenobranchialis Karrer & Smith, 1980
Figs
This species is widespread in the Indo-West Pacific and was previously reported from Taiwan by
Congriscus megastoma (Günther, 1877)
Figs
This species is restricted to the northwestern Pacific. It is not uncommon around Dongsha Island.
Macrocephenchelys brachialis Fowler, 1934
Figs
This species is widespread in the Indo-West Pacific. It is common around Taiwan, yet only one specimen was collected around Dongsha Island.
Rouleina squamilatera (Alcock, 1898)
Figs
This species has scattered distribution in the Indo-West Pacific. It is rare around Dongsha Island.
Otolith images of fresh specimens collected around Dongsha Island A Nemichthys scolopaceus, CHLOL 23020, 641.46 mm TL B Sigmops elongatus, CHLOL 15420, 146.75 mm SL C Diplophos vicinia, CHLOL 19048, 138.84 mm SL D Rouleina squamilatera, CHLOL 15938 166.50 mm SL E Rouleina watasei, CHLOL 16466, 221.15 mm SL F Nansenia ardesiaca, CHLOL 20443, 142.76 mm SL. Scale bar: 1 mm.
Diplophos vicinia Koeda & Ho, 2019
Figs
This species was recently described from southern Taiwan and Papua New Guinea. The specimens we collected around Dongsha Island, suggest that this species is also found in the South China Sea and is sympatric with the morphologically similar congener D. taenia.
Polyipnus matsubarai Schultz, 1961
Figs
This species has been reported from Japan and Hawaii (Emperor Seamounts). The occurrence around Dongsha Island suggests its broad distribution in the northwestern Pacific to Hawaii.
Images of fresh specimens collected around Dongsha Island A Polyipnus matsubarai, 79.97 mm SL B Polyipnus spinifer, 54.97 mm SL C Polyipnus stereope, 37.93 mm SL D Sternoptyx pseudobscura, 24.90 mm SL E Sternoptyx diaphana, 24.19 mm SL F Argyripnus sp., 87.00 mm SL G Polymetme surugaensis, 115.57 mm SL H Polymetme elongata, 124.14 mm SL I Polymetme corythaeola, 214.45 mm SL J Argyropelecus affinis, 46.60 mm SL. Scale bar: 3 cm.
Otolith images of fresh specimens collected around Dongsha Island A Polyipnus matsubarai, CHLOL 16404, 79.97 mm SL B Polyipnus spinifer, CHLOL 21531, 52.83 mm SL C Polyipnus stereope, CHLOL 20639, 43.47 mm SL D Argyripnus sp., CHLOL 3446, 86.97 mm SL E Argyropelecus affinis, CHLOL 15486, 60.46 mm SL F Polymetme elongata, CHLOL 20610, 154.19 mm SL G Polymetme surugaensis, CHLOL 20580, 187.44 mm SL H Polymetme corythaeola, CHLOL 29482, 214.45 mm SL. Scale bar: 1 mm.
Polyipnus spinifer Borodulina, 1979
Figs
This species is widespread in the western Pacific. It is not uncommon around Taiwan, especially in mid-water trawlers, but rare around Dongsha Island.
Polymetme corythaeola (Alcock, 1898)
Figs
This species can be found in the western Pacific. It is very common around Dongsha Island but rare around Taiwan.
Polymetme surugaensis (Matsubara, 1943)
Figs
This species is widespread in the western Pacific. It is common in mid-water trawlers around Taiwan but rare around Dongsha Island.
Borostomias elucens (Brauer, 1906)
Figs
While having circumglobal distribution in tropical and temperate oceans, this species is rare around Dongsha Island.
Borostomias pacificus (Imai, 1941)
Figs
This species is restricted to the northwestern Pacific and is rare around Dongsha Island.
Images of fresh specimens collected around Dongsha Island A Astronesthes lucifer, 100.27 mm SL B Astronesthes indica, 84.31 mm SL C Astronesthes chrysophekadion, 96.90 mm SL D Astronesthes indopacifica, 129.70 mm SL E Astronesthes trifibulata, 123.68 mm SL F Chauliodus sloani, 176.27 mm SL G Photostomias tantillux, 105.26 mm SL H Stomias nebulosus, 76.28 mm SL I Borostomias pacificus, 113.63 mm SL J Photonectes albipennis, 191.94 mm SL. Scale bar: 3 cm.
Photostomias tantillux Kenaley, 2009
Figs
This species is widespread in the Pacific. Only one specimen was collected around Dongsha Island.
Otolith images of fresh specimens collected around Dongsha Island. Scanning electron microscope images of A Sternoptyx diaphana, CHLOL 21042, 24.19 mm SL B Photonectes albipennis, CHLOL 17271, 191.94 mm SL C Photostomias tantillux, CHLOL 20618, 105.26 mm SL D Chauliodus sloani, CHLOL 20056, 159.87 mm SL E Stomias nebulosus, CHLOL 29697 92.64 mm SL F Xiphasia setifer, CHLOL 26196, 312.83 mm TL. Scale bar: 0.5 mm.
Otolith images of fresh specimens collected around Dongsha Island A Astronesthes trifibulata, CHLOL 15414, 123.68 mm SL B Astronesthes indopacifica, CHLOL 22771, 74.26 mm SL C Astronesthes indica, CHLOL 21597, 84.31 mm SL D Astronesthes lucifer, CHLOL 26368, 118.29 mm SL E Astronesthes chrysophekadion, CHLOL 23215, 96.90 mm SL F Leptostomias robustus, CHLOL 27092, 338.66 mm SL G Borostomias elucens, CHLOL 27093, 245.61 mm SL H Borostomias pacificus, CHLOL 28799, 223.81 mm SL. Scale bar: 1 mm.
Chlorophthalmus pectoralis Okamaura & Doi, 1984
Figs
This species is found in the western Pacific. It is common around Dongsha Island but rare around Taiwan.
Otolith images of fresh specimens collected around Dongsha Island A Chlorophthalmus acutifrons, CHLOL 20303, 104.77 mm SL B Chlorophthalmus pectoralis, CHLOL 21197, 159.37 mm SL C Rosenblattichthys alatus, CHLOL 17910, 121.03 mm SL D Ateleopus japonicus, CHLOL 21595, 233.22 mm SL E Ijimaia dofleini, CHLOL 19918, 727.27 mm SL. Scale bar: 1 mm.
Dolichosudis fuliginosa Post, 1969
Figs
This species is widely distributed in the Atlantic and Pacific.
Images of fresh specimens collected around Dongsha Island A Stemonosudis cf. siliquiventer, 242.07 mm SL B Stemonosudis rothschildi, 172.22 mm SL C Lestidiops sp., 146.08 mm SL D Lestidium longilucifer, 190.84 mm TL E Lestidium prolixum, 209.71 mm SL F Lestidium orientale, 210.45 mm SL G Lestrolepis philippina, 191.77 mm SL H Sudis sp., 191.00 mm SL I Dolichosudis fuliginosa, 241.49 mm SL J Magnisudis sp., 318.06 mm SL. Scale bar: 3 cm.
Otolith images of fresh specimens collected around Dongsha Island A Dolichosudis fuliginosa, CHLOL 18720, 249.25 mm SL B Lestidiops sp., CHLOL 16047, 146.08 mm SL C Lestidium orientale, CHLOL 24778, 227.03 mm SL (R) D Lestidium prolixum, CHLOL 22028, 222.87 mm SL (R) E Lestidium longilucifer, CHLOL 20615, 199.43 mm TL (R) F Lestrolepis philippina, CHLOL 20260, 191.46 mm SL (R) G Stemonosudis cf. siliquiventer, CHLOL 23015, 227.89 mm SL H Stemonosudis rothschildi, CHLOL 23330, 154.29 mm TL I Sudis sp., CHLOL 16187, 191.00 mm SL J Magnisudis sp., CHLOL22413, 294.11 mm SL. Scale bar: 1 mm.
Lestidium longilucifer Ho, Graham & Russell, 2020
Figs
This species was recently described from southern Taiwan and Australia. It is rare around Dongsha Island and Taiwan.
Lestidium orientale Ho, Tsai & Li, 2019
Figs
This species, recently described around southwestern Taiwan, is restricted to the northwestern Pacific. It is rare around Dongsha Island, but common around Taiwan.
Lestidium prolixum Harry, 1953
Figs
This species is restricted to the northwestern Pacific. It is rare around Dongsha Island, but common around Taiwan (
Stemonosudis rothschildi Richards, 1967
Figs
This species is almost circumglobally distributed in tropical oceans, except eastern Pacific.
Dasyscopelus selenops (Tåning, 1928)
Figs
This species is circumglobally distributed through tropical and warm temperate oceans, except eastern Pacific. It is rare around Dongsha Island, and was recently recorded off southwestern Taiwan (
Images of fresh specimens collected around Dongsha Island A Neoscopelus porosus, 147.05 mm SL B Neoscopelus microchir, 87.87 mm SL C Neoscopelus macrolepidotus, 110.69 mm SL D Neoscopelus sp., 132.66 mm SL E Dasyscopelus obtusirostris, 75.20 mm SL F Dasyscopelus asper, 62.70 mm SL G Dasyscopelus selenops, 59.36 mm SL H Electrona risso, 57.32 mm SL I Bolinichthys sp., 72.95 mm SL J Lampanyctus sp., 97.39 mm SL K Ceratoscopelus sp., 63.30 mm SL L Benthosema fibulatum, 58.90 mm SL M Lampadena luminosa, 108.84 mm SL. Scale bar: 3 cm.
Diaphus adenomus Gilbert, 1905
Figs
Although widespread in the Pacific and Atlantic, this species has never been reported in the tropical northwestern Pacific. The two specimens collected in the present study represent the first record in the South China Sea.
Images of fresh specimens collected around Dongsha Island A Diaphus splendidus, 83.65 mm SL B Diaphus lucidus, 77.38 mm SL C Diaphus fragilis, 118.97 mm SL D Diaphus perspicillatus, 63.58 mm SL E Diaphus parri, 47.04 mm SL F Diaphus garmani, 51.20 mm SL G Diaphus suborbitalis, 62.73 mm SL H Diaphus watasei, 132.33 mm SL I Diaphus luetkeni, 64.93 mm SL J Diaphus problematicus, 70.60 mm SL K Diaphus adenomus, 161.15 mm SL. Scale bar: 3 cm.
Otolith images of fresh specimens collected around Dongsha Island A Neoscopelus macrolepidotus, CHLOL 22410, 107.12 mm SL B Neoscopelus microchir, CHLOL 20187, 107.30 mm SL C Neoscopelus porosus, CHLOL 20608, 129.94 mm SL D Neoscopelus sp., CHLOL 20517, 83.60 mm SL E Benthosema fibulatum, CHLOL 9210, 57.15 mm SL F Bolinichthys sp., CHLOL 29700, 60.66 mm SL G Dasyscopelus obtusirostris, CHLOL 22678, 75.20 mm SL H Dasyscopelus asper, CHLOL 24112, 62.13 mm SL I Dasyscopelus selenops, CHLOL 20183, 63.61 mm SL J Ceratoscopelus sp., CHLOL 20651 61.32 mm SL K Diaphus garmani, CHLOL 22594, 48.91 mm SL L Diaphus splendidus, CHLOL 29230, 33.81 mm SL M Diaphus luetkeni, CHLOL 17523, 64.93 mm SL N Diaphus suborbitalis, CHLOL 22680, 73.40 mm SL O Diaphus problematicus, CHLOL 22774, 81.69 mm SL P Diaphus lucidus, CHLOL 16019, 91.21 mm SL Q Diaphus parri, CHLOL 20570, 47.04 mm SL R Diaphus perspicillatus, CHLOL 22485, 56.48 mm SL S Diaphus fragilis, CHLOL 20604, 108.61 mm SL T Diaphus watasei, CHLOL 3480, 114.45 mm SL. Scale bars: 1 mm.
Zenion japonicum Kamohara, 1934
Figs
This species is widespread in the Pacific. Only one specimen was collected around Dongsha Island, yet it appears to be more common around Taiwan.
Images of fresh specimens collected around Dongsha Island A Parazen pacificus, 89.49 mm SL B Polymixia longispina, 64.33 mm SL C Polymixia berndti, 115.2 mm SL D Cyttomimus affinis, 75.63 mm SL E Xenolepidichthys cf. dalgleishi, 90.42 mm SL F Xenolepidichthys dalgleishi, 51.16 mm SL G Cyttopsis rosea, 147.47 mm SL H Zenion sp., 42.07 mm SL I Zenion japonicum, 61.92 mm SL J Zu cristatus, 216.98 mm SL. Scale bar: 3 cm.
Otolith images of fresh specimens collected around Dongsha Island A Diaphus adenomus, CHLOL 29085, 161.15 mm SL B Lampadena luminosa, CHLOL 17521, 89.71 mm SL C Zenion sp., CHLOL 30182, 82.69 mm SL (R) D Polymixia berndti, CHLOL 29480, 71.65 mm SL E Polymixia longispina, CHLOL 20312, 76.05 mm SL F Lampanyctus sp., CHLOL 18810, 97.39 mm SL (R) G Electrona risso, CHLOL 16387, 57.32 mm SL H Zenopsis nebulosa, CHLOL 5787, 378.54 mm SL I Cyttomimus affinis, CHLOL 23366, 56.18 mm SL J Grammicolepis brachiusculus, CHLOL 20638, 140.27 mm SL K Cyttopsis rosea, CHLOL 29192, 99.27 mm SL L Xenolepidichthys cf. dalgleishi, CHLOL 17498, 97.39 mm SL M Xenolepidichthys dalgleishi, CHLOL 21836, 104.22 mm SL N Zenion japonicum, CHLOL 30180, 61.92 mm SL (R) O Parazen pacificus, CHLOL 18765, 122.83 mm SL. Scale bars: 1 mm.
Coelorinchus macrorhynchus Smith & Radcliffe, 1912
Figs
This species is widespread from southwestern Taiwan to western Australia, but has never been recorded around Dongsha Island, until the present study. It is not uncommon around Dongsha Island, yet most of the specimens collected are small. The largest specimen examined was 302 mm TL.
Images of fresh specimens collected around Dongsha Island A Gadomus colletti, 114.77 mm PAL B Coelorinchus cingulatus, 88.53 mm PAL C Coelorinchus sheni, 184.66 mm PAL D Coelorinchus brevirostris, 76.92 mm PAL E Coelorinchus macrorhynchus, 80.41 mm PAL F Coelorinchus sp., 125.80 mm PAL G Coelorinchus longissimus, 95.10 mm PAL H Coelorinchus smithi, 138.37 mm PAL I Malacocephalus nipponensis, 100.03 mm PAL. Scale bar: 3 cm.
Otolith images of fresh specimens collected around Dongsha Island A Gadomus colletti, CHLOL 20823, 103.93 mm PAL B Kuronezumia macronema, CHLOL 27490, 208.00 mm TL (R) C Kuronezumia bubonis, CHLOL 27491, 51.66 mm PAL D Coelorinchus sp., CHLOL 20068, 112.86 mm PAL E Coelorinchus brevirostris, CHLOL 15969, 83.09 mm PAL F Coelorinchus cingulatus, CHLOL 17475, 79.90 mm PAL G Coelorinchus longissimus, CHLOL 15397, 105.80 mm PAL H Coelorinchus macrorhynchus, CHLOL 15763, 126.55 mm PAL I Coelorinchus smithi, CHLOL 17376, 125.91 mm PAL J Coelorinchus sheni, CHLOL 19815, 214.53 mm PAL K Malacocephalus nipponensis, CHLOL 28535, 106.91 mm PAL. Scale bar: 1 mm.
Coelorinchus sheni Chiou, Shao & Iwamoto, 2004
Figs
Only one specimen was collected from Dongsha Island. It is also very rare around Taiwan and Japan (
Coelorinchus smithi Gilbert & Hubbs, 1920
Figs
This species is widely distributed in the central Indo-West Pacific, and is rare around Dongsha Island and Taiwan.
Kuronezumia bubonis Iwamoto, 1974
Figs
This species has a scattered distribution in the western Atlantic, western Indian Ocean, and the southwestern Pacific. Only one specimen was collected around Dongsha Island, representing the first record from the northwestern Pacific.
Spicomacrurus kuronumai (Kamohara, 1938)
Figs
This species was previously known only from Japan to Taiwan. The present record around Dongsha Island suggests its broad distribution in the northwestern Pacific.
Images of fresh specimens collected around Dongsha Island A Sphagemacrurus pumiliceps, 23.47 mm PAL B Hymenocephalus lethonemus, 37.27 mm PAL C Hymenocephalus longibarbis, 199.32 mm TL D Hymenocephalus striatissimus, 39.66 mm PAL E Pseudocetonurus cf. septifer, 25.13 mm PAL F Kumba gymnorhynchus, 22.12 mm PAL G Kumba japonica, 18.44 mm PAL H Kumba punctulata, 21.27 mm PAL. Scale bar: 3 cm.
Otolith images of fresh specimens collected around Dongsha Island A Pseudocetonurus cf. septifer, CHLOL 22917, 25.13 mm PAL B Nezumia condylura, CHLOL 28936, 27.50 mm PAL C Nezumia spinosa, CHLOL 28000, 38.12 mm PAL D Sphagemacrurus pumiliceps, CHLOL 20138, 23.13 mm PAL E Kumba punctulata, CHLOL 20826, 34.22 mm PAL F Kumba japonica, CHLOL 29704, 15.49 mm PAL G Kumba gymnorhynchus, CHLOL 27679, 22.12 mm PAL H Mataeocephalus sp., CHLOL 28763, 156.82 mm TL I Hymenocephalus striatissimus, CHLOL 20135, 33.43 mm PAL J Hymenocephalus longibarbis, CHLOL 20012, 58.81 mm PAL K Hymenocephalus lethonemus, CHLOL 20137, 39.68 mm PAL L Spicomacrurus kuronumai, CHLOL 23002, 52.10 mm PAL. Scale bar: 1 mm.
Images of fresh specimens collected around Dongsha Island A Kuronezumia bubonis, 51.66 mm PAL B Kuronezumia macronema, 208 mm TL C Ventrifossa longibarbata, 41.13 mm PAL D Ventrifossa johnboborum, 35.24 mm PAL E Ventrifossa sazonovi, 166.76 mm TL F Ventrifossa divergens, 56.05 mm PAL G Ventrifossa cf. longibarbata, 52.96 mm PAL H Nezumia condylura, 30.56 mm PAL I Nezumia spinosa, 45.92 mm PAL J Mataeocephalus sp., 54.96 mm PAL K Spicomacrurus kuronumai, 37.02 mm PAL. Scale bar: 3 cm.
Ventrifossa johnboborum Iwamoto, 1982
Figs
This species is widespread in the central Indo-West Pacific and the southeastern Pacific. Only two small individuals were collected around Dongsha Island.
Otolith images of fresh specimens collected around Dongsha Island A Ventrifossa johnboborum, CHLOL 27105, 35.24 mm PAL B Ventrifossa longibarbata, CHLOL 22754, 48.88 mm PAL C Ventrifossa divergens, CHLOL 25656, 69.58 mm PAL D Ventrifossa sazonovi, CHLOL 20707, 56.86 mm PAL E Ventrifossa cf. longibarbata, CHLOL 17064, 47.15 mm PAL. Scale bar: 1 mm.
Gadella jordani (Böhlke & Mead, 1951)
Figs
This species is widespread in the Indo-West Pacific and is one the most common species in deepwater trawl fisheries around Taiwan. However, this species appears to be less common around Dongsha Island.
Otolith images of fresh specimens collected around Dongsha Island A Gadella jordani, CHLOL 27463, 205.96 mm SL (R) B Physiculus sp. 1, CHLOL 27649, 112.77 mm SL C Physiculus sp. 2, CHLOL 4203, 56.27 mm PAL D Physiculus chigodarana, CHLOL 22372, 168.71 mm SL E Physiculus japonicus, CHLOL 18348, 57.47 mm PAL F Bregmaceros japonicus, CHLOL 15755, 62.54 mm SL G Diretmus argenteus, CHLOL 22570, 52.10 mm SL. H Diretmoides veriginae, CHLOL 18692, 118.47 mm SL Scale bars: 1 mm.
Laemonema robustum Johnson, 1862
Fig.
This species has a circumglobal distribution in tropical and temperate oceans. It is known from one specimen captured by a deepsea longline fishery around Dongsha Island.
Physiculus chigodarana Paulin, 1989
Figs
This species is restricted to the northwestern Pacific. It is common around Taiwan but rare around Dongsha Island. Based on our observations, this species usually occurs in shallower waters.
Hoplostethus roseus Su, Lin & Ho, 2022
Figs
This species was recently described from Taiwan and is widespread in the central Indo-West Pacific. We found that this species is not uncommon around Dongsha Island, suggesting its broad distribution in the South China Sea.
Images of fresh specimens collected around Dongsha Island A Diretmoides veriginae, 157.03 mm SL B Diretmus argenteus, 51.84 mm SL C Bregmaceros japonicus, 82.38 mm SL D Encheliophis sp., 165.00 mm SL E Beryx mollis, 118.03 mm SL F Hoplostethus melanopus, 174.52 mm SL G Hoplostethus roseus, 110.34 mm SL H Hoplostethus sp., 107.77 mm SL I Hoplostethus robustispinus, 213.97 mm SL. Scale bar: 3 cm.
Otolith images of fresh specimens collected around Dongsha Island A Hoplostethus sp., CHLOL 29039, 140.17 mm SL B Hoplostethus melanopus, CHLOL 20387, 157.22 mm SL C Beryx mollis, CHLOL 20633, 122.49 mm SL D Hoplostethus roseus, CHLOL 22715, 110.74 mm SL E Hoplostethus robustispinus, CHLOL 22731, 213.97 mm SL. Scale bar: 1 mm.
Glyptophidium argenteum Alcock, 1889
Figs
This species has a broad distribution in the Indo-West Pacific. It is rare around Taiwan and Dongsha Island, where only one specimen was collected.
Images of fresh specimens collected around Dongsha Island A Luciobrotula bartschi, 133.17 mm PAL B Hoplobrotula armata, 81.73 mm PAL C Homostolus acer, 68.27 PAL D Monomitopus sp., 58.58 mm PAL E Neobythites bimaculatus, 50.38 mm PAL F Neobythites unimaculatus, 96.75 mm PAL G Glyptophidium japonicum, 56.51 mm PAL H Glyptophidium lucidum, 74.65 mm PAL I Glyptophidium argenteum, 65.90 mm PAL J Glyptophidium oceanium, 77.92 mm PAL K Dicrolene tristis, 51.33 PAL L Neobythites longipes, 51.33 mm PAL M Pycnocraspedum microlepis, 142.31 mm PAL N Lamprogrammus brunswigi, 440.00 mm TL. Scale bar: 3 cm.
Otolith images of fresh specimens collected around Dongsha Island A Dicrolene tristis, CHLOL 20052, 63.84 mm PAL B Encheliophis sp., CHLOL 4101, 165.00 mm TL C Hoplobrotula armata, CHLOL 24857, 196.93 mm SL D Homostolus acer, CHLOL 20168, 73.17 mm PAL E Glyptophidium japonicum, CHLOL 29128, 56.51 mm PAL F Glyptophidium lucidum, CHLOL 23426, 58.25 mm PAL G Glyptophidium argenteum, CHLOL 22281, 65.90 mm PAL H Glyptophidium oceanium, CHLOL 19990, 77.92 mm PAL. I Monomitopus sp., CHLOL 22272, 59.58 mm PAL J Luciobrotula bartschi, CHLOL 20829, 231.74 mm SL K Pycnocraspedum microlepis, CHLOL 15786, 142.31 mm PAL. L Neobythites longipes, CHLOL 22758, 68.89 mm SL M Lamprogrammus brunswigi, CHLOL 17964, 440.00 mm TL N Neobythites unimaculatus, CHLOL 6266, 88.86 mm PAL O Neobythites bimaculatus, CHLOL 9881, 57.53 mm PAL P Neobythites sivicola, CHLOL 27431, 48.80 mm PAL. Scale bars: 1 mm.
Glyptophidium japonicum Kamohara, 1936
Figs
Similar to the congeners, this species can be found in the Indo-West Pacific. It appears more common than the other sympatric congeners around Dongsha Island and Taiwan.
Glyptophidium oceanium Smith & Radcliffe in Radcliffe, 1913
Figs
As in other congeners, this species is widespread in the Indo-West Pacific. It is rare around Dongsha Island and around Taiwan.
Neobythites bimaculatus Nielsen, 1997
Figs
Being widespread in the central Indo-West Pacific, this species is very common around Dongsha Island, but less common around Taiwan. It is especially similar to N. macrops Günther, 1887 described from the Philippines. However, N. macrops has not been reported in the South China Sea before. Therefore, all specimens are identified as N. bimaculatus.
Neobythites unimaculatus Smith & Radcliffe in Radcliffe, 1913
Figs
This species is widespread in the central Indo-West Pacific. It is known only from one specimen around Dongsha Island but appears to be more common around the shallower waters of Taiwan. The depth record around Dongsha Island appears to be the lower depth limit of this species.
Cataetyx lepidogenys (Smith & Radcliffe, 1913)
Figs
This species is restricted to the northwestern Pacific. While it is common off southwestern Taiwan, we only collected a few specimens around Dongsha Island.
Images of fresh specimens collected around Dongsha Island A Bythitidae indet., 42.50 mm PAL B Barathronus maculatus, 69.18 mm PAL C Diplacanthopoma sp., 103.86 mm PAL D Pseudonus squamiceps, 41.47 mm PAL E Saccogaster horrida, 58.20 mm PAL F Cataetyx lepidogenys, 71.10 mm PAL G Saccogaster tuberculata, 59.34 mm PAL. Scale bar: 3 cm.
Otolith images of fresh specimens collected around Dongsha Island A Saccogaster horrida, CHLOL 18127, 55.20 mm PAL B Saccogaster tuberculata, CHLOL 22284, 59.34 mm PAL C Pseudonus squamiceps, CHLOL 19987, 37.55 mm PAL D Barathronus maculatus, CHLOL 25717, 69.18 mm SL E Diplacanthopoma sp., CHLOL 19402, 76.62 mm PAL F Bythitidae indet., CHLOL 19985, 42.50 mm PAL (R) G Cataetyx lepidogenys, CHLOL 18149, 71.10 mm PAL. Scale bar: 1 mm.
Pseudonus squamiceps (Lloyd, 1907)
Figs
This species is found in the central Indo-West Pacific. Although poorly presented in museums, we found that it is not uncommon around Dongsha Island.
Saccogaster horrida Nielsen, Schwarzhans & Cohen, 2012
Figs
This species was known only from western Australia. The sole specimen collected around Dongsha Island in the present study suggests a potential broad distribution in the central Indo-West Pacific.
Barathronus maculatus Shcherbachev, 1976
Figs
This species is widespread in the Indo-West Pacific. It is rare around Dongsha Island, as it usually occurs in much deeper waters (e.g.
Images of fresh specimens collected around Dongsha Island A Exocoetus monocirrhus, 144.25 mm SL B Hirundichthys oxycephalus, 211.01 mm TL C Decapterus macrosoma, 124.06 mm SL D Decapterus tabl, 91.56 mm SL E Xiphasia setifer, 312.83 mm TL F Decapterus macarellus, 174.18 mm TL G Trachurus japonicus, 125.57 mm SL. Scale bars: 3 cm.
Otolith images of fresh specimens collected around Dongsha Island A Exocoetus monocirrhus, CHLOL 19408, 144.25 mm SL B Decapterus macarellus, CHLOL 20653, 174.18 mm TL C Decapterus macrosoma, CHLOL 17765, 124.06 mm SL (R) D Hirundichthys oxycephalus, CHLOL 15698, 211.01 mm TL E Trachurus japonicus, CHLOL 24117, 104.79 mm SL F Decapterus tabl, CHLOL 22730, 251.70 mm SL. Scale bar: 1 mm.
Otolith images of fresh specimens collected around Dongsha Island A Lepidoblepharon ophthalmolepis, CHLOL 22379, 67.13 mm SL B Poecilopsetta plinthus, CHLOL 23359, 69.60 mm SL C Chascanopsetta lugubris, CHLOL 14976, 230.17 mm SL D Chascanopsetta prognatha, CHLOL 15675, 194.29 mm SL E Symphurus orientalis, CHLOL 18684, 87.69 mm SL. Scale bar: 1 mm.
Images of fresh specimens collected around Dongsha Island A Nesiarchus nasutus, 430.94 mm SL B Nealotus tripes, 143.09 mm SL C Rexea bengalensis, 241.14 mm SL D Rexea prometheoides, 235.74 mm SL E Promethichthys prometheus, 267.82 mm SL F Scombrolabrax heterolepis, 161.77 mm SL G Neoepinnula orientalis, 193.94 mm SL H Gempylus serpens, 506.87 mm SL I Ruvettus pretiosus, 211.29 mm SL J Thyrsitoides marleyi, 386.84 mm SL. Scale bar: 5 cm.
Otolith images of fresh specimens collected around Dongsha Island A Scombrolabrax heterolepis, CHLOL 21499, 161.77 mm SL B Gempylus serpens, CHLOL 21292, 506.87 mm SL C Nealotus tripes, CHLOL 18041, 143.09 mm SL D Nesiarchus nasutus CHLOL 23266, 389.39 mm SL E Promethichthys prometheus, CHLOL 21502, 292.37 mm SL F Neoepinnula orientalis, CHLOL 20027, 185.78 mm SL G Rexea bengalensis CHLOL 5395, 241.14 mm SL H Rexea prometheoides, CHLOL 5394, 235.80 mm SL I Ruvettus pretiosus, CHLOL 22035, 292.42 mm SL J Thyrsitoides marleyi, CHLOL 24722, 390.60 mm SL K Aphanopus sp., CHLOL 28066, 437.53 mm SL (R) L Benthodesmus tenuis, CHLOL 22846, 510.72 mm SL M Benthodesmus sp., CHLOL 16539, 665.63 mm SL. Scale bars: 1 mm.
Cubiceps pauciradiatus Günther, 1872
Figs
This species has circumglobal distribution in tropical and temperate oceans. It is very rare around Dongsha Island, known only from one specimen.
Images of fresh specimens collected around Dongsha Island A Champsodon snyderi, 90.06 mm SL B Champsodon longipinnis, 127.54 mm SL C Cubiceps whiteleggii, 163.50 mm SL D Cubiceps pauciradiatus, 115.74 mm SL E Amarsipus carlsbergi, 146.27 mm SL F Psenes pellucidus, 229.58 mm SL G Psenes cyanophrys, 141.89 mm SL H Psenes arafurensis, 123.44 mm SL I Cubiceps baxteri, 265.12 mm SL J Psenopsis anomala, 76.07 mm SL. Scale bar: 3 cm.
Otolith images of fresh specimens collected around Dongsha Island A Psenopsis anomala, CHLOL 26057, 76.07 mm SL B Amarsipus carlsbergi, CHLOL 29706, 146.27 mm SL C Cubiceps whiteleggii, CHLOL 21285, 112.40 mm SL D Cubiceps pauciradiatus, CHLOL 24988, 115.74 mm SL E Cubiceps baxteri, CHLOL 20390, 119.10 mm SL F Psenes arafurensis, CHLOL 24971, 132.51 mm SL G Psenes pellucidus, CHLOL 20622, 213.19 mm SL H Psenes cyanophrys, CHLOL 27368, 123.32 mm SL. I Champsodon longipinnis, CHLOL 24121, 127.54 mm SL J Champsodon snyderi, CHLOL 21520, 90.06 mm SL K Bathysphyraenops simplex, CHLOL 15774, 69.53 mm SL. Scale bars: 1 mm.
Cubiceps whiteleggii (Waite, 1894)
Figs
Although widely distributed in the Indo-West Pacific, this species has not been recorded around Dongsha Island. Here, we found that this species is very abundant in the area, outnumbering the congeners.
Champsodon longipinnis Matsubara & Amaoka, 1964
Figs
This species is widespread in the Indo-West Pacific, and is common off southwestern Taiwan. However, only one specimen was collected around Dongsha Island, which suggests that this species may prefer shallower waters.
Malakichthys elegans Matsubara & Yamaguti, 1943
Figs
This species is widespread in the central Indo-West Pacific, and is one of the most abundant species in bottom trawl fisheries in Taiwan. However, only one specimen was collected around Dongsha Island, possibly due to a much deeper sampling depth.
Images of fresh specimens collected around Dongsha Island A Ostracoberyx dorygenys, 73.53 mm SL B Parascombrops serratospinosus, 107.83 mm SL C Bathysphyraenops simplex, 69.53 mm SL D Malakichthys elegans, 170.51 mm SL E Synagrops japonicus, 220.42 mm SL F Synagrops atrumoris, 132.91 mm SL G Xenocephalus elongatus, 219.48 mm SL. Scale bar: 5 cm.
Otolith images of fresh specimens collected around Dongsha Island A Parascombrops serratospinosus, CHLOL 5589, 92.81 mm SL B Synagrops japonicus, CHLOL 24379, 203.28 mm SL C Synagrops atrumoris, CHLOL 26169, 129.07 mm SL D Epigonus pectinifer, CHLOL 22798, 97.02 mm SL E Epigonus megalops, CHLOL 22100, 102.02 mm SL (R) F Epigonus denticulatus, CHLOL 27481, 103.79 mm SL G Ostracoberyx dorygenys, CHLOL 28712, 103.95 mm SL H Malakichthys elegans, CHLOL 24712, 122.29 mm SL (R). I Neobathyclupea gracilis, CHLOL 17470, 159.61 mm SL J Neobathyclupea japanotaiwana, CHLOL 22536, 194.90 mm SL K Neobathyclupea malayana, CHLOL 22087, 187.50 mm SL. Scale bars: 1 mm.
Parascombrops serratospinosus (Smith & Radcliffe, 1912)
Figs
This species is found in the western Pacific. Although common off southwestern Taiwan, the species is very rare around Dongsha Island, which suggests that it usually occurs in shallower waters.
Epigonus denticulatus Dieuzeide, 1950
Figs
This species has a circumglobal distribution in tropical and temperate oceans. It is uncommon around Dongsha Island and Taiwan.
Images of fresh specimens collected around Dongsha Island A Neobathyclupea malayana, 229.75 mm SL B Neobathyclupea gracilis, 179.90 mm SL C Neobathyclupea japanotaiwana, 181.32 mm SL D Pentaceros japonicus, 100.59 mm SL E Priacanthus zaiserae, 124.10 mm SL F Owstonia grammodon, 183.01 mm SL G Owstonia aurora, 111.14 mm SL H Brama orcini, 102.62 mm SL I Brama dussumieri, 142.96 mm SL J Epigonus pectinifer, 94.61 mm SL K Epigonus megalops, 161.37 mm SL L Epigonus denticulatus, 169.85 mm SL. Scale bar: 3 cm.
Epigonus megalops (Smith & Radcliffe, 1912)
Figs
This species is restricted to the Philippines. The sole specimen we recognized around Dongsha Island reflects a broad distribution in the South China Sea.
Epigonus pectinifer Mayer, 1974
Figs
Although widespread circumglobally in tropical and temperate oceans, this species is rare around Dongsha Island and Taiwan.
Neobathyclupea gracilis (Fowler, 1938)
Figs
This genus was established by
Neobathyclupea japanotaiwana (Prokofiev, 2014)
Figs
This species has a narrower distribution than its congeners, known only from Japan and Taiwan. However, this species is not uncommon around Dongsha Island, suggesting a broad distribution in the northwestern Pacific.
Neobathyclupea malayana (Weber, 1913)
Figs
Although widespread in the Indo-West Pacific, this species is rare around Dongsha Island. It is also more prominent in body size than the two sympatric congeners.
Brama dussumieri Cuvier, 1831
Figs
This species can be found in tropical and warm temperate oceans. The two specimens from our collection are the first record of this species around Dongsha Island.
Brama orcini Cuvier, 1831
Figs
This species is widespread in the Indo-West Pacific. It is common around Taiwan, but only one specimen was collected around Dongsha Island.
Owstonia aurora Liao, Reyes & Shao, 2022
Figs
This species was recently described from the Philippines. Our present record, represented by one specimen around Dongsha Island, suggests its broad distribution in the South China Sea.
Owstonia grammodon (Fowler, 1934)
Figs
This species has scattered records in Indonesia and Taiwan. The two specimens collected around Dongsha Island confirm its broader distribution in the tropical Indo-West Pacific.
Lythrichthys cypho (Fowler, 1938)
Figs
This species was recently revised by
Images of fresh specimens collected around Dongsha Island A Bembrops caudimacula, 165.04 mm SL B Lioscorpius longiceps, 139.94 mm SL C Ectreposebastes imus, 56.56 mm SL D Plectrogenium kamoharai, 43.27 mm SL E Phenacoscorpius megalops, 65.03 mm SL F Setarches guentheri, 126.15 mm SL G Erisphex pottii, 62.07 mm SL H Lythrichthys cypho, 53.30 mm SL I Lythrichthys eulabes, 137.08 mm SL. Scale bar: 3 cm.
Otolith images of fresh specimens collected around Dongsha Island A Pentaceros japonicus, CHLOL 26254, 100.59 mm SL B Brama dussumieri, CHLOL 25658, 142.96 mm SL C Brama orcini, CHLOL 15403, 56.19 mm SL D Priacanthus zaiserae, CHLOL 5243, 124.10 mm SL (R) E Bembrops caudimacula, CHLOL 22142, 165.04 mm SL F Owstonia grammodon, CHLOL 24036, 183.01 mm SL G Owstonia aurora, CHLOL 20621, 111.14 mm SL. Scale bars: 1 mm.
Otolith images of fresh specimens collected around Dongsha Island A Setarches guentheri, CHLOL 24292, 131.63 mm SL B Lioscorpius longiceps, CHLOL 22373, 136.03 mm SL C Lythrichthys eulabes, CHLOL 24508, 120.40 mm SL D Phenacoscorpius megalops, CHLOL 24396, 62.60 mm SL E Ectreposebastes imus, CHLOL 28795, 47.71 mm SL F Lythrichthys cypho, CHLOL 24502, 47.83 mm SL G Plectrogenium kamoharai, CHLOL 9880, 49.42 mm SL H Erisphex pottii, CHLOL 29164, 62.07 mm SL. Scale bars: 1 mm.
Lythrichthys eulabes Jordan & Starks, 1904
Figs
Being revised together with L. cypho, this species is also widespread in the Indo-West Pacific. It is very common around Dongsha Island and Taiwan.
Setarches guentheri Johnson, 1862
Figs
This species has circumglobal distribution in tropical and subtropical oceans. It is not uncommon around Dongsha Island and Taiwan.
Lepidotrigla pectoralis Fowler, 1938
Figs
This species is possibly endemic to the northwestern Pacific, and is rare around Dongsha Island.
Images of fresh specimens collected around Dongsha Island A Pterygotrigla macrorhynchus, 179.11 mm SL B Pterygotrigla cajorarori, 225.51 mm SL C Paraheminodus murrayi, 193.98 mm SL D Heminodus philippinus, 248.83 mm SL E Lepidotrigla sp., 115.30 mm SL F Lepidotrigla pectoralis, 127.46 mm SL G Peristedion riversandersoni, 158.52 mm SL H Peristedion orientale, 128.99 mm SL I Scalicus hians, 165.47 mm SL J Hoplichthys gilberti, 288.42 mm SL K Hoplichthys fasciatus, 78.74 mm SL L Hoplichthys filamentosus, 184.54 mm SL M Scalicus orientalis, 129.56 mm SL N Satyrichthys milleri, 265.63 mm SL O Chelidonichthys spinosus, 132.61 mm. Scale bar: 3 cm.
Otolith images of fresh specimens collected around Dongsha Island A Scalicus orientalis, CHLOL 27465, 128.73 mm SL B Scalicus hians, CHLOL 29478, 165.47 mm SL C Satyrichthys milleri, CHLOL 20905, 103.55 mm SL D Paraheminodus murrayi, CHLOL 20864, 245.67 mm SL E Lepidotrigla sp., CHLOL 23056, 115.30 mm SL F Lepidotrigla pectoralis, CHLOL 29737, 124.66 mm SL G Pterygotrigla macrorhynchus, CHLOL 8397, 155.76 mm SL H Pterygotrigla cajorarori, CHLOL 23058, 171.52 mm SL I Chelidonichthys spinosus, CHLOL 29166, 212.35 mm SL (R) J Peristedion riversandersoni, CHLOL 23051, 164.13 mm SL K Peristedion orientale, CHLOL 21074, 151.57 mm SL. Scale bar: 1 mm.
Pterygotrigla cajorarori Richards & Yato, 2012
Figs
Although widespread in the Indo-West Pacific, this species appears to be uncommon around Dongsha Island.
Paraheminodus murrayi (Günther, 1880)
Figs
This species is widely distributed in the Indo-West Pacific, and is common around Dongsha Island, but rare around Taiwan.
Satyrichthys milleri Kawai, 2013
Figs
This species is widely distributed in the central Indo-West Pacific. It is common off southwestern Taiwan but rare around Dongsha Island.
Hoplichthys filamentosus Matsubara & Ochiai, 1950
Figs
This species is widespread in the central Indo-West Pacific. It is common around Taiwan, but rare around Dongsha Island.
Stlengis distoechus Bolin, 1936
Figs
This species was formerly restricted to Japan. The present record around Dongsha Island suggests its broader distribution in the northwestern Pacific.
Psychrolutes macrocephalus (Gilchrist, 1904)
Figs
This species has a widespread distribution in the Indo-West Pacific. Only a few specimens were collected around Dongsha Island.
Otolith images of fresh specimens collected around Dongsha Island A Hoplichthys gilberti, CHLOL 9890, 89.80 mm SL B Hoplichthys fasciatus, CHLOL 24449, 78.74 mm SL C Hoplichthys filamentosus, CHLOL 22098, 204.66 mm SL D Psychrolutes macrocephalus, CHLOL 25008, 172.17 mm SL E Stlengis distoechus, CHLOL 24045, 48.77 mm SL. Scale bar: 1 mm.
Lophiodes iwamotoi Ho, Séret & Shao, 2011
Figs
This species was originally described from off Society Islands. A few specimens were collected around Dongsha Island, representing a substantial range extension.
Otolith images of fresh specimens collected around Dongsha Island A Erythrocles schlegelii, CHLOL 21498, 177.46 mm SL (R) B Lophiodes iwamotoi, CHLOL 23849, 109.84 mm SL C Lophiodes mutilus, CHLOL 23149, 250.73 mm SL D Lophiodes lugubris, CHLOL 23262, 265.64 mm SL E Lophiodes naresi, CHLOL 9768, 185.84 mm SL F Lophiodes triradiatus, CHLOL 20515, 437.17 mm SL. Scale bar: 1 mm.
Chaunax apus Lloyd, 1909
Figs
This species is widespread in the Indo-West Pacific. Although not previously recorded from Dongsha Island, we found it very common in the area but rare around Taiwan.
Chaunax breviradius Le Danois, 1978
Figs
This species is common around Taiwan and is restricted to the South China Sea. It has never been recorded around Dongsha Island before, until the present study.
Halieutopsis echinoderma Ho, 2021
Figs
This species was recently described from Taiwan and the Coral Sea, and one specimen was collected around Dongsha Island, suggesting its broad distribution in the central Indo-West Pacific.
Images of fresh specimens collected around Dongsha Island A Halieutopsis echinoderma, 64.26 mm SL B Halieutopsis nasuta, 54.01 mm SL C Halieutaea coccinea, 65.84 mm SL D Halieutopsis sp., 53.07 mm SL E Malthopsis annulifera, 85.54 mm SL F Malthopsis mitrigera, 55.29 mm SL G Malthopsis kobayashii, 75.44 mm SL H Malthopsis tiarella, 66.21 mm SL I Halicmetus ruber, 86.79 mm SL J Halicmetus cf. ruber, 56.29 mm SL K Halicmetus reticulatus, 77.90 mm SL L Coelophrys micropus, 64.00 mm SL. Scale bar: 3 cm.
Halieutopsis nasuta Alcock, 1891
Figs
Although being widespread in the Indo-West Pacific, this species is very rare around Dongsha Island, known only from a few specimens. The present record confirms its appearance in the South China Sea.
Otolith images of fresh specimens collected around Dongsha Island A Chaunax penicillatus, CHLOL 20309, 127.92 mm SL B Chaunax apus, CHLOL 20308, 95.51 mm SL C Chaunax breviradius, CHLOL 5399, 131.88 mm SL D Chaunax sp., CHLOL 15746, 160.97 mm SL E Coelophrys micropus, CHLOL 27454, 64.00 mm SL F Malthopsis mitrigera, CHLOL 17434, 55.29 mm SL G Malthopsis tiarella, CHLOL 4394, 58.70 mm SL H Malthopsis kobayashii, CHLOL 6852, 61.09 mm SL I Malthopsis annulifera, CHLOL 3678, 85.54 mm SL J Halieutopsis sp., CHLOL 18748, 46.37 mm SL K Halieutopsis echinoderma, CHLOL 29043, 64.26 mm SL L Halieutopsis nasuta, CHLOL 25666, 73.65 mm SL M Halicmetus ruber, CHLOL 21065, 65.15 mm SL N Halicmetus cf. ruber, CHLOL 24204, 64.74 mm SL O Halicmetus reticulatus, CHLOL 20910, 101.23 mm SL P Halieutaea coccinea, CHLOL 27444, 65.84 mm SL. Scale bars: 1 mm.
Otolith images of fresh specimens collected around Dongsha Island A Triacanthodes anomalus, CHLOL 22706, 73.57 mm SL B Bathyphylax bombifrons, CHLOL 12732, 43.93 mm SL C Paratriacanthodes retrospinis, CHLOL 23527, 90.30 mm SL (R) D Halimochirurgus alcocki, CHLOL 22698, 143.53 mm SL (R) E Tydemania navigatoris, CHLOL 22702, 110.20 mm SL (R). Scale bar: 1 mm.
Malthopsis kobayashii Tanaka, 1916
Figs
This species was originally described from Japan and resurrected by
Okamejei sp.
Fig.
This sole neonate specimen is light yellow on the dorsal side and white on the ventral side, without any blotches or patterns. This species is under investigation by SLN.
Mobula spp.
Several large specimens were observed at the landing site but were dissected, making specific identification impossible. Nevertheless, this record shows the existence of Mobula species in the deep waters around Dongsha Island.
Neenchelys sp.
Fig.
This sole specimen belongs to Neenchelys but morphologically does not resemble any members.
Ophichthus spp.
Figs
These specimens belong to Ophichthus. More specimens without precise identification are found in the collection.
Bathycongrus sp.
Figs
This species has certain differences in morphology from other congeners. It is common around Dongsha Island.
Bathyuroconger cf. vicinus
Figs
The sole specimen resembles B. vicinus, but has a longer dorsal fin and more elongated body.
Bathyuroconger sp.
Figs
This species is most similar to B. albus, but its otoliths are distinctly different from those of the congeners.
Argyripnus sp.
Figs
This species represents the first record of the genus in the South China Sea. However, the specimen was partially damaged and was not retained.
Lestidiops sp.
Figs
One specimen with a partially damaged body was identified at a generic level. This species is the only representative of the genus from our collection.
Magnisudis sp.
Figs
This species is uncommon around Dongsha Island. Most of the individuals are severely damaged.
Stemonosudis cf. siliquiventer
Figs
This species is rare around Dongsha Island but common around Taiwan (
Sudis sp.
Figs
A single specimen was collected from Dongsha Island. Although both species under Sudis were predicted to be circumglobal, our specimen represents the first record of the genus in the South China Sea.
Neoscopelus sp.
Figs
This species is currently known only around Dongsha Island, and is abundant. It differs from other congeners by the unique photophore pattern and the pinkish coloration and is recognized as undescribed.
Bolinichthys spp.
Figs
The specimens are easily recognizable as Bolinichthys by having a whitish crescent on the posterior edge of the eye. However, almost all specimens are partially damaged and lack diagnostic photophores; thus, species-level identification is impossible. They are very common around Dongsha Island.
Ceratoscopelus sp.
Figs
Ceratoscopelus warmingii (Lütken, 1892) is the only species previously reported in the South China Sea (
Lampanyctus spp.
Figs
This genus has a complicated taxonomic history and is especially difficult to identify. All specimens are conservatively identified at a generic level. This genus is uncommon around Dongsha Island.
Zenion sp.
Figs
This common species is currently known only from Dongsha Island. It is similar to Z. japonicum, but differs in the dorsal-fin morphology and some body morphometrics. It is recognized as undescribed. and is currently under investigation by YTL in the special issue.
Xenolepidichthys cf. dalgleishi
Figs
These specimens are similar to X. dalgleishi in all aspects, but without the conspicuous black blotches on the body.
Coelorinchus sp.
Figs
This species is similar to C. smithi, but differs from the morphology of the occipital scale and body morphometrics. It is the most abundant macrourid species collected around the studied area and is possibly restricted to the South China Sea.
Mataeocephalus sp.
Figs
This species differs from the two sympatric species, M. cristatus Sazonov, Shcherbachev & Iwamoto, 2003 and M. hyostomus (Smith & Radcliffe, 1912), by having the underside of snout largely naked (
Pseudocetonurus cf. septifer
Figs
The sole specimen collected around Dongsha Island resembles the Taiwanese specimen listed in
Ventrifossa cf. longibarbata
Figs
This species differs from V. longibarbata by having fewer transverse scale rows, body morphometrics, and having distinct black blotch on the first dorsal fin. The status of the species is currently under investigation by SLN. It is common around Dongsha Island.
Physiculus sp. 1
Figs
This species resembles the Physiculus sp. 1 in
Physiculus sp. 2
Figs
This species resembles the Physiculus sp. 3 in
Hoplostethus sp.
Figs
This species is similar to H. roseus, but differs from the latter by having the gular region covered with scales and the oral cavity pale (
Encheliophis sp.
Figs
Known only from two specimens in rather poor condition around Dongsha Island. We conservatively identified the specimens to a generic level. This is the first record of this genus in the area.
Monomitopus sp.
Figs
Members of this genus are exceptionally similar to each other, thus making it difficult to identify. A comprehensive taxonomic revision of all the members is needed. This species is common around Dongsha Island.
Bythitidae indet.
Figs
This specimen resembles members of the genus Cataetys, but it has a more slender body. Due to the limited knowledge of the taxonomy of the bythihtids in this region, we conservatively assigned this specimen to the familial level.
Diplacanthopoma sp.
Figs
This genus is poorly studied, and it is uncertain how many species are valid. As a result, all specimens are identified to a generic level. This species is uncommon around Dongsha Island.
Aphanopus sp.
Figs
The sole specimen represents the first record of the genus Aphanopus around Taiwan and the South China Sea. The status of the species if under investigation by YTL.
Benthodesmus sp.
Figs
This species is uncommon around Dongsha Island. It is similar to B. tenuis in morphological aspects but with a dense black body coloration and a shorter orbital length.
Lepidotrigla sp.
Figs
This species resembles Lepidotrigla sp. sensu
Chaunax sp.
Figs
This species is morphologically distinct from other known Chaunax species. It is not uncommon around Taiwan but rare around Dongsha Island.
Halicmetus cf. ruber
Figs
This species is similar to, yet distinct from H. ruber. This is also common around Dongsha Island.
Halieutopsis sp.
Figs
This species, represented by a few specimens, is recognized as undescribed.
The earliest record of the ichthyofauna around Dongsha Island was a report by
In this study, we cataloged a total of 1087 species across 167 families, and our recent efforts alone identified 337 species from 93 families (Suppl. material
The updated checklist not only enriches our understanding of marine biodiversity but also serves as a critical resource for future research and conservation initiatives. Notably, based on the IUCN Red List of Threatened Species (
We thank all the people who helped with processing the fish specimens: Chia-Yen Lin, Shu-Ling Lee, Ying-Cheng Tseng, Tai-Yen Lin, and the members of Marine Paleontology Lab, BRCAS (https://otolithlin.biodiv.tw/). We are grateful to the following fellows for their assistance in the identification of some specimens: Yung-Chieh Chiu (Fisheries Research Institute), Chi-Ngai Tang, Man-Kwan Wong (National Taiwan University), Jian-Fu Huang (National Taiwan Ocean University), and Yo Su (National Kaohsiung University of Science and Technology). We thank Daphne Hoh from the Taiwan Biodiversity Information Facility for assisting with the data publishing. We thank Yung-Tai Li for communicating and contacting the trawling fishery companies. We thank Iizuka Yoshiyuki and Yu-Shiang Wang (Institute of Earth Sciences Academia Sinica) for help with SEM images. We thank the following captains of the vessels who provided access to the samples and detailed information on the trawling sites: Dong Yuan Jyi No. 168, Jin Roei Yih No. 33, Jin Roei Yih No. 37, Chian Chun No. 1, and Jin Yang. We also thank Barry C. Russell for his valuable insights and thoughtful review of this paper.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This study was supported by the Intramural funding of Academia Sinica, Taipei, Taiwan, and National Science and Technology Council, Taiwan (Grant No. 109-2116-M-001-022, 110-2116-M-001-009, 111-2116-M-001-033, and 112-2116-M-001-017-MY3) to C-HL.
Shing-Lai Ng – Conceptualization, Methodology, Validation, Writing - Original draft, Writing - Review and Editing. Hsin-Wei Liu – Conceptualization, Methodology, Data Curation, Validation, Writing - Review and Editing. Dominique P. Mediodia – Data Curation, Writing - Original draft, Writing - Review and Editing. Yen-Ting Lin – Methodology, Writing - Original draft, Writing - Review and Editing. Chieh-Hsuan Lee – Methodology, Data Curation, Writing - Review and Editing. Shih-Pin Huang – Data Curation, Validation, Writing - Review and Editing. Siao-Man Wu – Data Curation, Writing - Review and Editing. Ching-Feng Lin – Methodology, Writing - Original draft, Writing - Review and Editing. Chih-Ren Tung – Writing - Review and Editing. Hsuan-Ching Ho – Conceptualization, Methodology, Validation, Writing - Original draft, Writing - Review and Editing, Supervision. Chien-Hsiang Lin – Conceptualization, Methodology, Validation, Writing - Original draft, Writing - Review and Editing, Supervision.
Shing-Lai Ng https://orcid.org/0000-0001-6914-9100
Hsin-Wei Liu https://orcid.org/0009-0001-7650-0883
Dominique P. Mediodia https://orcid.org/0000-0002-7305-6588
Yen-Ting Lin https://orcid.org/0009-0002-3641-1947
Chieh-Hsuan Lee https://orcid.org/0009-0001-8492-8556
Shih-Pin Huang https://orcid.org/0000-0002-5398-5646
Siao-Man Wu https://orcid.org/0009-0004-3702-1996
Chih-Ren Tung https://orcid.org/0009-0001-7647-3627
Hsuan-Ching Ho https://orcid.org/0000-0003-1154-601X
Chien-Hsiang Lin https://orcid.org/0000-0002-9843-9729
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Checklist of fishes around Dongsha Island
Data type: docx
Explanation note: table S1. Checklist of fishes around Dongsha Island based on the literature and our collection. Species recorded and described around Dongsha Island for the first time (in this study) are in bold font.