Search

Gymnothorax poikilospilus Chen & Huang, sp. nov. Common name: Variegated moray eel (Figs. 1, 2) urn:lsid:zoobank.org:act: 855F9025-193E-4B0E-9118-C217166CDADF Holotype. TOU-AE 0645, 768 mm TL, female, off Hsiao-men fishing harbor (23°39'N, 119°29'E), Xi-yu Township, Penghu Islands, Taiwan, ca. 10–20 m depth, trap of PVC tube, 10 June 2002, coll. Captain Ping Hong. Paratype. TOU-AE 5112, 868+ mm TL, tail tip lost, female, collected with the holotype. Diagnosis. A medium-sized moray with typical muraenid shape, anus at mid-point of body. Dorsal fin high. Eyes above mid-jaw. Jaws slightly arched and can’t completely close. Cephalic sensory pores small. Teeth uniserial, slender and pointed. Body brown or gray-brown with several rows of large dark brown patches on back of body and dorsal fin. Patches obscure during daytime and becoming obvious at night. Dorsal head dark brown and spotless. Fins with fine greyish-white margins when alive but invisible in preservation. Inner skin of nostrils and head pores whitish. Oral cavity uniform brown. Throat grooves darkish. Iris of the eye reddish-brown. Predorsal vertebrae 4, preanal vertebrae 57–59, total vertebrae 141. Description. Proportions in percent of TL for holotype: tail length 50.3, trunk length 34.8, head length 14.5, depth at gill opening 8.3, depth at anus 7.2. Proportions in percent of HL for holotype (paratype): predorsal length 60.0 (58.0), length of upper jaw 46.3 (47.8), length of lower jaw 46.8 (47.7), interorbital width 11.1 (11.7), snout length 17.9 (18.5), eye diameter 8.7 (8.3). Predorsal vertebrae 4 (4), preanal vertebrae 57 (59), total vertebrae 141 (137+) (Table 1). A medium-sized moray with typical muraenid shape, body stout, anus at mid-point of body. Dorsal fin high, originating anterior to gill opening. Anal fin shallow, its origin immediately behind anus. Gill opening slightly below lateral midline, opening as a hole, about size of eye in diameter. Eyes above mid-jaw. Jaws slightly arched and can’t completely close, teeth visible when mouth closed, jaws subequal in length. Snout acute and moderately elongate. Anterior nostril at tip of snout, as short tube shorter than eye radius in length, without folds on the anterior tip. Posterior nostril above and slightly anterior to front margin of eye, as an oval pore with raised rim. Cephalic sensory pores small, their number within range of typical muraenids (Fig. 3). Three supraorbital pores, first at tip of snout below level of anterior nostril; second immediately above base of anterior nostril; third along margin of snout above level of horizontal middle line of eye. Four infraorbital pores, first immediately below and posterior to anterior nostril base; second about at anterior one-third of distance between anterior nostril base and anterior margin of eye, slightly posterior to third supraorbital pore in horizontal; third below anterior margin of eye; fourth below posterior margin of eye. Six preoperculo-mandibular pores along lower jaw before corner of mouth; in holotype, pores arranging differently between left and right jaws but amounting the same number (Fig. 4). Two branchial pores on posterior-dorsal head, posterior to dorsal-fin origin and anterior to gill opening. Teeth slender and pointed, slightly retrorse, edge without serration (Fig. 4). Peripheral intermaxillary teeth uniserial, 6 (6–9) large canines on each side with small teeth in intervals; teeth becoming larger posteriorly. Median intermaxillary teeth uniserial, with 3 (3) tall and spaced teeth, second tooth longest. Maxillary teeth uniserial, with 18–19 (20) triangular and recurved teeth on each side, anterior most 1–2 teeth very small, following by 2–4 sharply increase-sized teeth flanked by several tiny teeth; remaining teeth gradually decreasing in size, smallest at posterior end. Vomerine teeth almost invisible, only one very small and stout tooth left in holotype, with four sockets of missing teeth. No visible vomerine teeth in paratype. Dentary teeth uniserial, with 27 (30–31) teeth on each side, anterior most 5–7 teeth obviously larger and flanked by some tiny teeth; size of remaining teeth subequal centrally and becoming much smaller at posterior end. Body brown or gray-brown in ground color, with several rows of inconspicuous large dark brown patches on back of body and dorsal fin. Ventral parts of head and trunk slightly lighter. Dorsal head dark brown without blotch. Throat grooves darkish. Inner skin of nostrils and head pores whitish. Color of oral cavity similar to head. Iris of the eye reddish-brown. Body color variable when alive. During daytime, body almost uniform brown, dark patches obscure; during nighttime, ground color turning lighter and dark patches becoming obvious (Fig. 5), fins with fine greyish-white margins (Fig. 6). Preserved color similar to fresh, pale margins of fins and dark patches nearly invisible. Distribution. Currently only known from the type locality Penghu Islands. Biology. From 10 June 2002, we had reared the holotype in our laboratory's aquarium until 4 February 2005, it accidentally crawled out of the aquarium at night and died. We dissected the holotype and found that there were many mature eggs in its abdominal cavity (Fig. 7). The mature eggs were between 0.8–1.2 mm in diameter, and the batch fecundity was about 20,000 eggs. Under microscope observation, there was the other immature egg group in its ovaries, the diameters of oocytes were about 0.15–0.26 mm. We also reared the paratype until its death on 22 September 2008. Under the microscope, its gonads belonged to immature ovaries, and there were 2 groups of oocytes with different egg diameter interval ranges: the larger group was 0.17–0.26 mm, and the smaller group was 0.07–0.13 mm. We fed these two specimens with fish and squid meat. Etymology. The specific name is from the Greek poikilo (varied, variegated) and the Greek spilos (a spot, stain), in reference to its varied body markings between daytime and nighttime. COI analyses. After alignment and trimming, 612 bp of COI sequences were left for analyses. The topology of ML tree reveals that each species is positioned in a monophyletic clade and separated from others (Fig. 8). The genus Enchelycore is not monophyletic, conforming with previous studies based on different mitochondrial genes (Tang & Fielitz 2013; Smith et al. 2019). The inter- and intra-species K2P genetic distances are 9.8–22.5% and 0.1–0.7%, respectively (Table 2). Gymnothorax poikilospilus is most closely related to E. schismatorhynchus, however, with an 11.4% K2P distance which far exceeds the general intraspecific variation of moray eels (Huang et al. 2018), confirming its validity. Remarks. Although the new species has arched jaws that can’t close completely, it is placed under the genus Gymnothorax instead of Enchelycore according to the following reasons: (1) the extents of elongation and arching of its jaws are much lower than species of the genus Enchelycore (Fig. 9); (2) its maxillary and dentary teeth are uniserial (vs. mostly biserial in Enchelycore). While hard to quantify the extent of elongated and arched jaws in Enchelycore, this characteristic could further reflect on the narrowing of the anterior part of head, including the interorbital width. According to our unpublished data, the interorbital width of Enchelycore ranges from 7.8–10.4% of HL (based on one E. bayeri, two E. lichenosa, two E. pardalis, and four E. schismatorhynchus), comparing to>10% of HL in most species of Gymnothorax. The new species has an interorbital width of 11.1–11.7% of HL which is completely within the range of Gymnothorax. In addition, arched jaws can also be found in several non- Enchelycore species, such as Enchelynassa canina and Gymnothorax eurostus, suggesting it might not be an appropriate diagnostic characteristic. On the other hand, we found that the largest examined Enchelycore schismatorhynchus (949 mm TL) has much fewer maxillary and dentary teeth compared to smaller specimens (510–642 mm TL) (maxillary teeth 15–17 vs. 29–31; dentary teeth 28–29 vs. 37–44), and all series of its teeth are arranged in single rows (vs. biserial of maxillary and dentary teeth in smaller specimens). Considering ambiguous morphological characters and the genetically non-monophyly of the genus Enchelycore, we temporarily designate the new species under the genus Gymnothorax. Further systematic research is needed to verify the validity of Enchelycore. Gymnothorax poikilospilus is morphologically most similar to Enchelycore schismatorhynchus, G. monochrous, and G. pseudothyrsoideus, three sympatric morays that possess gray-brown body color and pale margins on fins (Fig. 10). It differs from E. schismatorhynchus by the wider interorbital width (11.1–11.7% vs. 8.9–10.4% of HL), fewer predorsal (4 vs. 5–6) and preanal vertebrae (57–59 vs. 61–63), and the color of the tip of anterior nostrils (uniform brown vs. dark brown). It can be distinguished from G. monochrous by more dentary teeth (27–31 vs.16–25), and more total vertebrae (141 vs. 130–138). Lastly, G. poikilospilus is different from G. pseudothyrsoideus by having more maxillary (18–20 vs. 11–17) and dentary teeth (27–31 vs. 19–24), and more total vertebrae (141 vs. 130–135). Aside from morphometric and meristic characters, the obviously arched jaws make G. poikilospilus unique when compared with G. monochrous and G. pseudothyrsoideus; and the lack of the prominent white fin margin makes G. poikilospilus distinguishable at a glance when side by side with E. schismatorhynchus (Fig. 10). Furthermore, the pale fin margins of the three similar species are still visible in preservation while completely disappearing in G. poikilospilus. Gymnothorax poikilospilus can be excluded from the four synonyms of E. schismatorhynchus (Eurymyctera crudelis Kaup, 1856, Muraena congeroides Bleeker,1860, Muraena hemprichii Klunzinger, 1871, and Rhinamuraena eritima Jordan & Seale, 1906) by the dentition (uniserial vs. biserial maxillary and dentary teeth) and the color of fin margin (brown vs. white); and be excluded from the only synonym of G. pseudothyrsoideus, Gymnothorax makassariensis Bleeker, 1863, by more maxillary teeth (18–20 vs. 14–15) and more dentary teeth (27–31 vs. 20–23) (Böhlke & Smith 2002). Gymnothorax poikilospilus can be easily distinguished from other three similar species of Gymnothorax by having more outer maxillary teeth (18–20 vs. 10–18 in G. cinerascens, 9–16 in G. hepaticus, and 11–16 in G. longinquus), more outer dentary teeth (27–31 vs. 12–22, 12–21, and 18–22), and more total vertebrae (141 vs. 128–135, 128–132, and 128–134) (Smith et al. 2019; Huang et al. 2020). It can also be simply separated from the three Indo-Pacific jaw-arched brown morays (viz. Enchelycore bayeri, Enchelycore bikiniensis, and Enchelynassa canina) by using a single characteristic of uniserial maxillary and dentary teeth (vs. biserial in other three species) (Böhlke & Smith 2002; Loh et al. 2012; Smith et al. 2019). Comparative materials. Enchelycore schismatorhynchus. four specimens, 510–949 mm TL. Taiwan: DOS 07936 (1, 510 mm), Changbin, Taitung; DOS 08729 (1, 949 mm), Chenggong, Taitung; NMMB-P007074 (1, 642 mm), Checheng, Pingtung; NMMB-P015637 (1, 640 mm), Liuqiu, Pingtung. Gymnothorax monochrous. seven specimens, 557–891 mm TL. Philippines: DOS 06828 (4, 557– 784 mm), Bogo, Cebu. Taiwan: DOS 06376 (1, 758 mm), Donggang, Pingtung; DOS 07127 (1, 891 mm), DOS 08281 (1, 666 mm), Magong, Penghu Islands. Gymnothorax pseudothyrsoideus. five specimens, 400–876 mm TL. Taiwan: DOS 07943 (3, 564– 876 mm), Eziliao, Kaohsiung; DOS 08038 (1, 635 mm), Qianzhen, Kaohsiung; DOS 08076 (1, 400 mm), Datan, Taoyuan., Published as part of Huang, Wen-Chien, Loh, Kar-Hoe & Chen, Hong-Ming, 2022, Gymnothorax poikilospilus, a new moray eel (Teleostei: Anguilliformes: Muraenidae) from Penghu Islands, western Taiwan, pp. 87-102 in Zootaxa 5189 (1) on pages 88-97, DOI: 10.11646/zootaxa.5189.1.11, http://zenodo.org/record/7119280, {"references":["Tang, K. L. & Fielitz, C. (2013) Phylogeny of moray eels (Anguilliformes: Muraenidae), with a revised classification of true eels (Teleostei: Elopomorpha: Anguilliformes). Mitochondrial DNA, 24 (1), 55 - 66. https: // doi. org / 10.3109 / 19401736.2012.710226","Smith, D. G., Bogorodsky, S. V., Mal, A. O. & Alpermann, T. J. (2019) Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea, with description of a new species. Zootaxa, 4704 (1), 1 - 87. https: // doi. org / 10.11646 / zootaxa. 4704.1.1","Huang, W. C., Nguyen, V. Q. & Liao, T. Y. (2018) First record of the snowflake-patched moray Gymnothorax niphostigmus Chen, Shao, & Chen, 1996 (Anguilliformes; Muraenidae) in Vietnam and its validity confirmed by DNA barcoding. Journal of Applied Ichthyology, 34 (3), 687 - 690. https: // doi. org / 10.1111 / jai. 13684","Kaup, J. J. (1856) Uebersicht der Aale. Archiv fur Naturgeschichte, 22, 41 - 77. https: // doi. org / 10.5962 / bhl. part. 11240","Bleeker, P. (1860) Achtste bijdrage tot de kennis der vischfauna van Sumatra (Visschen van Benkoelen, Priaman, Tandjong, Palembang en Djambi). Acta Societatis Scientiarum Indo-Neerlandicae, 8 (art. 2), 1 - 88.","Klunzinger, C. B. (1871) Synopsis der Fische des Rothen Meeres. II. Theil. Verhandlungen der kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien, 21, 441 - 688. https: // doi. org / 10.5962 / bhl. title. 14760","Jordan, D. S. & Seale, A. (1906) The fishes of Samoa. Description of the species found in the archipelago, with a provisional check-list of the fishes of Oceania. Bulletin of the United States Bureau of Fisheries, 25 [for 1905], 173 - 455 + index 457 - 488, pls. 33 - 53.","Bleeker, P. (1863) Sur quelques especes nouvelles ou peu connues de Gymnothorax Bl. de l'Inde archipelagique. Nederlandsch Tijdschrift voor Dierkunde, 1, 167 - 171.","Bohlke, E. B. & Smith, D. G. (2002) Type catalogue of Indo-Pacific Muraenidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 152 (1), 89 - 172. https: // doi. org / 10.1635 / 0097 - 3157 (2002) 152 [0089: TCOIPM] 2.0. CO; 2","Huang, W. C., Thu, P. T. & Liao, T. Y. (2020) A new record of the long moray, Gymnothorax longinquus (Actinopterygii: Anguilliformes: Muraenidae), from southern Vietnam, supporting the uncertain record in the Gulf of Thailand. Acta Ichthyologica Et Piscatoria, 50 (2), 201 - 207. https: // doi. org / 10.3750 / AIEP / 02790","Loh, K. H., Shao, K. T. & Chen, H. M. (2012) Additions to the Taiwan marine eel fauna with first records of three rare moray eels (Anguilliformes: Muraenidae). Journal of Marine Science and Technology, 20 (2), 210 - 215. https: // doi. org / 10.51400 / 2709 - 6998.1840"]}

Five specimens of Gymnothorax longinquus (Whitley, 1948) were collected from a fish market and a fish landing site in Tac Cau and Ha Tien, southern Vietnam. These specimens represent the first record of G. longinquus in Vietnam, and further support a previously uncertain record in Thailand. The specimens collected are described and illustrated. Gymnothorax longinquus may prefer muddy shallow water from northern Australia to the Gulf of Thailand.

Fishes of family Muraenidae, popularly known as moray eels, are one of the most diverse groups among true eels. Gymnothorax is the most predominant genus in terms of diversity along Indian coastline (31 species including recent description and distributional records), but they rarely form the fishery of commercial importance. Gymnothorax dorsalis, species of elongated unpatterned moray eels characterized by eight to nine pre-dorsal vertebrae, three infraorbital pores and low dorsal and anal fin united with caudal fins is reported for the first time from the west coast of India. Gymnothorax reticularis, reticulated patterned eel with 18–19 vertical dark bars, five pre-dorsal vertebrae and 10–11 vomerine teeth is also reported for the first time from the northwest coast of India.

Gymnothorax mordax (Ayres, 1859). California Moray. To 152 cm (5 ft) TL (Miller and Lea 1972). Point Conception, California to Todos Santos (23��24.6���N, 110��13.8���W), southern Baja California (John Snow, pers. comm. to M.L.), including Isla Guadalupe, central Baja California (Reyes-Bonilla et al. 2010), and Rocas Alijos (Personal communication: Scripps Institution of Oceanography Fish Collection, La Jolla, California). A specimen reported from Clipperton Island (Personal communication: Scripps Institution of Oceanography Fish Collection, La Jolla, California) was misidentified (Katherine Dale and Arturo Ram��rez, pers. comm. to M.L.), and one from Nicaragua (Personal communication: Scripps Institution of Oceanography Fish Collection, La Jolla, California) is a leptocephalus. Early-stage larvae have been taken around Cabos San Lucas, in the lowermost part of the Gulf of California (on both sides), and down the Mexican mainland as far south as Mazatl��n (William Watson, pers. comm. to M.L.). Beltr��n-Le��n and Rios Herrera (2000) tentatively identified larvae captured off Colombia as this species. Benthic; depth: lower intertidal to 40 m (131 ft) (min.: Ruiz-Campos et al. 2010b; max.: Eschmeyer and Herald 1983)., Published as part of Love, Milton S., Bizzarro, Joseph J., Cornthwaite, Maria, Frable, Benjamin W. & Maslenikov, Katherine P., 2021, Checklist of marine and estuarine fishes from the Alaska-Yukon Border, Beaufort Sea, to Cabo San Lucas, Mexico, pp. 1-285 in Zootaxa 5053 (1) on page 33, DOI: 10.11646/zootaxa.5053.1.1, http://zenodo.org/record/5578008, {"references":["Miller, D. J. & Lea, R. N. (1972) Guide to the coastal marine fishes of California. California Department of Fish and Game Fish Bulletin, 157.","Reyes-Bonilla, H., Ayala-Bocos, A., Gonzalez-Romero, S., Sanchez-Alcantara, I., Mendoza, M. W., Bedolla-Guzman, Y. R., Ramirez-Valdez, A., Calderon-Aguilera, L. E. & Olivares-Banuelos, N. C. (2010) Checklist and biogeography of fishes from Guadalupe Island, western Mexico. CalCOFI Reports, 51, 195 - 209.","Beltran-Leon, B. S. & Rios Herrera, R. (2000) Estadios Tempranos de Peces del Pacific Colombiano. Republica de Colombia. Instituto Nacional de Pesca y Acuicultura INPA.","Ruiz-Campos, G., Gonzalez-Guzman, S., Ramirez-Valdez, A., Gonzalez-Acosta, A. F., Castro-Aguirre, J. L. & de la Cruz-Aguero, J. (2010 b) Composition, density and biogeographic affinities of the rocky intertidal fishes on the western coast of the Baja California Peninsula, Mexico. CalCOFI Reports, 51, 210 - 220.","Eschmeyer, W. N. & Herald, E. S. (1983) A Field Guide to Pacific Coast Fishes of North America from the Gulf of Alaska to Baja California. Houghton Mifflin, Boston."]}

Gymnothorax flavimarginatus (R��ppell, 1830). Yellowmargin Moray. 124 cm (48.8 in) TL (Robertson and Allen 2008). Indo-Pacific; western Pacific Ocean north to Yonaguni-jima Island, Japan (Koeda et al. 2016); Bah��a Santa Maria, southern Baja California (Jonathan et al. 2015); tip of Baja California, Costa Rica to Panama and offshore islands (Robertson and Allen 2008). Benthic; depth: 0.5���150 m (2���492 ft) (min.: Smith et al. 2019; max.: Robertson and Allen 2008)., Published as part of Love, Milton S., Bizzarro, Joseph J., Cornthwaite, Maria, Frable, Benjamin W. & Maslenikov, Katherine P., 2021, Checklist of marine and estuarine fishes from the Alaska-Yukon Border, Beaufort Sea, to Cabo San Lucas, Mexico, pp. 1-285 in Zootaxa 5053 (1) on page 33, DOI: 10.11646/zootaxa.5053.1.1, http://zenodo.org/record/5578008, {"references":["Robertson, D. R. & Allen, G. R. (2008) Shorefishes of the Tropical Eastern Pacific: an Information System. Version 1.0 (2008). Smithsonian Tropical Research Institute, Balboa.","Koeda, K., Hibino, Y., Yoshida, T., Kimura, Y., Miki, R., Kunishima, T., Sasaki, D., Furukawa, T., Sakurai, M., Eguchi, K., Suzuki, H., Inaba, T., Uejo, T., Tanaka, S., Fujisawa, M., Wada, H. & Uchiyama, T. (2016) Annotated Checklist of Fishes of Yonaguni-jima Island, the Westernmost Island in Japan. Kagoshima University Museum, Korimoto, Kagoshima.","Jonathan, M. P., Aurioles-Gamboa, D., Campos Villegas, L. E., Bohorquez-Herrera, J., Hernandez-Camacho, C. J. & Sujitha, S. B. (2015) Metal concentrations in demersal fish species from Santa Maria Bay, Baja California Sur, Mexico (Pacific coast). Marine Pollution Bulletin, 99, 356 - 361. https: // doi. org / 10.1016 / j. marpolbul. 2015.07.032","Smith, D. G., Bogorodsky, S. V., Mal, A. O. & Alpermann, T. J. (2019) Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea with description of a new species. Zootaxa, 4704, 1 - 87. https: // doi. org / 10.11646 / zootaxa. 4704.1.1"]}

Gymnothorax dovii (G��nther, 1870). Fine-spotted Moray. 170 cm (66.9 in) TL (Robertson and Allen 2008). Isla de Cedros, central Baja California (Personal communication: California Academy of Sciences Fish Collection, San Francisco, California) to Gulf of California to Ecuador, and offshore islands including Islas Gal��pagos (Robertson and Allen 2008). Benthic; depth: intertidal to 160 m (525 ft) (min.: Personal communication: Scripps Institution of Oceanography Fish Collection, La Jolla, California; max.: S��nchez-Jim��nez et al. 2018)., Published as part of Love, Milton S., Bizzarro, Joseph J., Cornthwaite, Maria, Frable, Benjamin W. & Maslenikov, Katherine P., 2021, Checklist of marine and estuarine fishes from the Alaska-Yukon Border, Beaufort Sea, to Cabo San Lucas, Mexico, pp. 1-285 in Zootaxa 5053 (1) on page 33, DOI: 10.11646/zootaxa.5053.1.1, http://zenodo.org/record/5578008, {"references":["Robertson, D. R. & Allen, G. R. (2008) Shorefishes of the Tropical Eastern Pacific: an Information System. Version 1.0 (2008). Smithsonian Tropical Research Institute, Balboa.","Sanchez-Jimenez, A., Naranjo-Elizondo, B., Rodriguez-Arrieta, A., Quesada, A., Blum, S., McCosker, J. E., Robertson, D. R., Auster, P. J. & Cortes, J. (2018) Updated catalogue of bony fishes observed in deep waters at Isla del Coco National Park and Las Gemelas Seamount, Costa Rica (Eastern Tropical Pacific). Revista de Biologia Tropical, 66, (Supplement 5), S 1 - S 113. https: // doi. org / 10.15517 / rbt. v 66 i 5.34943"]}

Gymnothorax equatorialis (Hildebrand, 1946). Equatorial Moray or Spottail Moray. To 100 cm (39.4 in) TL (G��mez et al. 2014). Bah��a San Crist��bal, southern Baja California (27��37���N, 114��55���W) (Rodr��guez-Romero et al. 2008), Gulf of California (McCosker and Rosenblatt in Fischer et al. 1995) to Iquique, northern Chile (Sielfeld et al. 2010), including southern tip of Baja California (McCosker and Rosenblatt in Fischer et al. 1995). Benthic; depth: surface (Personal communication: Scripps Institution of Oceanography Fish Collection, La Jolla, California), 5���183 m (16���600 ft) (min.: Robertson and Allen 2002; max.: Personal communication: Los Angeles County Museum of Natural History Fish Collection, Los Angeles, California)., Published as part of Love, Milton S., Bizzarro, Joseph J., Cornthwaite, Maria, Frable, Benjamin W. & Maslenikov, Katherine P., 2021, Checklist of marine and estuarine fishes from the Alaska-Yukon Border, Beaufort Sea, to Cabo San Lucas, Mexico, pp. 1-285 in Zootaxa 5053 (1) on page 33, DOI: 10.11646/zootaxa.5053.1.1, http://zenodo.org/record/5578008, {"references":["Gomez, S., Alejandro Caicedo, J. & Alonso Zapata, L. (2014) Captura de peces no comerciales con espinel experimental de fondo en Isla Gorgona, Colombia, y su area de influencia. Revista de Biologia Tropical, 62 (Supplement 1), 391 - 405. https: // doi. org / 10.15517 / rbt. v 62 i 0.16365","Rodriguez-Romero, J., Palacios-Salgado, D. S. & Gutierrez Sanchez, F. J. (2008) First record of the spottail moray Gymnothorax equatorialis (Anguilliformes: Muraenidae) off the west coast of Baja California Sur, Mexico. Marine Biodiversity Records, 1, 2 - 4. https: // doi. org / 10.1017 / s 1755267207007981","Fischer, W., Krupp, F., Schneider, W., Sommer, C., Carpenter, K. E. & Niem, V. H. (1995) Guia FAO para la identificacion para los fines de la pesca. Pacifico centro-oriental. Volume II, Vertebrados, Parte 1. Volume III, Vertebrados, Parte 2. FAO, Rome.","Sielfeld, W., Laudien, J., Vargas, M. & Villegas, M. (2010) El Nino induced changes of the coastal fish fauna off northern Chile and implications for ichthyogeography. Revista de Biologia Marina Oceanografia, 45, S 1, 705 - 722. https: // doi. org / 10.4067 / s 0718 - 19572010000400014","Robertson, D. R. & Allen, G. R. (2002) Shorefishes of the tropical eastern Pacific: an information system. CD-ROM. Smithsonian Tropical Research Institute, Balboa, Panama."]}

Gymnothorax verrilli (Jordan & Gilbert, 1883). White-edged Moray. To 43 cm (16.9 in) TL (McCosker and Rosenblatt in Fischer et al. 1995). Bah��a Magdalena, southern Baja California (Charter and Moser in Moser 1996); Chiapas, Mexico (Gonz��lez-Acosta et al. 2018) to Panama (Robertson and Allen 2015). Benthic; depth: intertidal to 18 m (60 ft) (min.: Robertson and Allen 2015; max.: Personal communication: Los Angeles County Museum of Natural History Fish Collection, Los Angeles, California)., Published as part of Love, Milton S., Bizzarro, Joseph J., Cornthwaite, Maria, Frable, Benjamin W. & Maslenikov, Katherine P., 2021, Checklist of marine and estuarine fishes from the Alaska-Yukon Border, Beaufort Sea, to Cabo San Lucas, Mexico, pp. 1-285 in Zootaxa 5053 (1) on page 34, DOI: 10.11646/zootaxa.5053.1.1, http://zenodo.org/record/5578008, {"references":["Fischer, W., Krupp, F., Schneider, W., Sommer, C., Carpenter, K. E. & Niem, V. H. (1995) Guia FAO para la identificacion para los fines de la pesca. Pacifico centro-oriental. Volume II, Vertebrados, Parte 1. Volume III, Vertebrados, Parte 2. FAO, Rome.","Moser, H. G. (Ed.). (1996) The early stages of fishes in the California Current region. CALCOFI Atlas, No. 33.","Gonzalez-Acosta, A. F., Rodiles-Hernandez, R. & Gonzalez-Diaz, A. A. (2018) Checklist of the marine and estuarine fishes of Chiapas, Mexico. Marine Biodiversity, 48, 1439 - 1454. https: // doi. org / 10.1007 / s 12526 - 016 - 0630 - y","Robertson, D. R. and Allen, G. R. (2015) Shorefishes of the Tropical Eastern Pacific: an Information System. Version 2.0 (2008). Smithsonian Tropical Research Institute, Balboa. http: // biogeodb. stri. si. edu / sftep / en / pages"]}

Gymnothorax castaneus (Jordan & Gilbert, 1883). Chestnut Moray or Panama Green Moray. To 155.2 cm (61.1 in) TL (Personal communication: Scripps Institution of Oceanography Fish Collection, La Jolla, California). Isla de Cedros, central Baja California (Robertson and Allen 2015), and Bah��a Magdalena, southern Baja California (Personal communication: Scripps Institution of Oceanography Fish Collection, La Jolla, California) to Ecuador (B��arez 1996), and perhaps to Chipana, northern Chile (Sielfeld 2010), including Gulf of California (McCosker and Rosenblatt in Fischer et al. 1995), and Islas Gal��pagos (Grove and Lavenberg 1997). Benthic; depth: intertidal to 101 m (3���331 ft) (min.: Castellanos-Galindo et al. 2014; max.: Personal communication: Los Angeles County Museum of Natural History Fish Collection, Los Angeles, California)., Published as part of Love, Milton S., Bizzarro, Joseph J., Cornthwaite, Maria, Frable, Benjamin W. & Maslenikov, Katherine P., 2021, Checklist of marine and estuarine fishes from the Alaska-Yukon Border, Beaufort Sea, to Cabo San Lucas, Mexico, pp. 1-285 in Zootaxa 5053 (1) on page 33, DOI: 10.11646/zootaxa.5053.1.1, http://zenodo.org/record/5578008, {"references":["Robertson, D. R. and Allen, G. R. (2015) Shorefishes of the Tropical Eastern Pacific: an Information System. Version 2.0 (2008). Smithsonian Tropical Research Institute, Balboa. http: // biogeodb. stri. si. edu / sftep / en / pages","Bearez, P. (1996) Lista de los peces marinos del Ecuador continental. Revista de Biologia Tropical, 44, 731 - 741.","Sielfeld, W., Laudien, J., Vargas, M. & Villegas, M. (2010) El Nino induced changes of the coastal fish fauna off northern Chile and implications for ichthyogeography. Revista de Biologia Marina Oceanografia, 45, S 1, 705 - 722. https: // doi. org / 10.4067 / s 0718 - 19572010000400014","Fischer, W., Krupp, F., Schneider, W., Sommer, C., Carpenter, K. E. & Niem, V. H. (1995) Guia FAO para la identificacion para los fines de la pesca. Pacifico centro-oriental. Volume II, Vertebrados, Parte 1. Volume III, Vertebrados, Parte 2. FAO, Rome.","Grove, J. S. & Lavenberg, R. J. (1997) The Fishes of the Galapagos Islands. Stanford University Press, Stanford.","Castellanos-Galindo, G. A., Giraldo, A. & Zapata, F. A. (2014) Tidepool fish assemblages of Gorgona Island, Colombian Pacific coast: a local and regional comparison). Revista de Biologia Tropical, 62 (Supplement 1), 373 - 390. https: // doi. org / 10.15517 / rbt. v 62 i 0.16362"]}

* Gymnothorax phalarus Bussing, 1998. Small-spotted Moray. To 93 cm (36.6 in) TL (Robertson and Allen 2002). Tip of Baja California and south-eastern Gulf of California to Peru (Robertson and Allen 2002). Benthic; depth: 25���933 m (82���3,060 ft) (min.: Robertson and Allen 2002; max.: Morera et al. 2019)., Published as part of Love, Milton S., Bizzarro, Joseph J., Cornthwaite, Maria, Frable, Benjamin W. & Maslenikov, Katherine P., 2021, Checklist of marine and estuarine fishes from the Alaska-Yukon Border, Beaufort Sea, to Cabo San Lucas, Mexico, pp. 1-285 in Zootaxa 5053 (1) on page 34, DOI: 10.11646/zootaxa.5053.1.1, http://zenodo.org/record/5578008, {"references":["Bussing, W. A. (1998) Freshwater fishes of Costa Rica. Revista Biologia Tropical, 46 (Supplement 2).","Robertson, D. R. & Allen, G. R. (2002) Shorefishes of the tropical eastern Pacific: an information system. CD-ROM. Smithsonian Tropical Research Institute, Balboa, Panama.","Morera, R. B., Calderon, F. C. & Hernandez, J. M. V. (2019) Abundancia, biomasa y estructura de la ictiofauna demersal en el oceano Pacifico de Centroamerica, basadas en datos de prospeccion pesquera realizados a bordo del B / W Miguel Oliver. Revista Ciencias Las Marinas y Costeras, 12, 27 - 47. https: // doi. org / 10.15359 / revmar. 12 - 1.2"]}

Gymnothorax aurocephalus Nashad, Mohapatra, Varghese & Bineesh sp. nov. (Figs. 1���6, Table 1) Common name: Goldenhead moray Holotype. EBRC /ZSI/ 11800, 723 mm TL, off Swaraj Dweep Island of the Andaman and Nicobar Islands (12��39.3���N, 93��03.4���E), collected on-board MFV Blue Marlin on 21 May 2018, by vertical longlining, depth 125���130 m. Paratypes. EBRC /ZSI/11801, 1 specimen (745 mm TL), collection details same as holotype; ZSI / ANRC / M/24129, 1 specimen (710 mm TL), off Interview Island, middle Andaman (12��55.3���N, 92��40.2���E), 90 m depth, hand line; FSI/PB / T/01 /2019, 1 specimen (610 mm TL, tail broken), collection details same as holotype. Diagnosis: A moderately elongated moray eel of the genus Gymnothorax with the following combination of characters: chocolate brown color overlain with small irregular white spots, having golden skin shadings close to rictus; anus slightly before midbody; pointed and serrated teeth; uniserial teeth on jaws and vomer; vertebral formula 7/61/148���149 (Fig. 2). Description: A moderately sized eel with a cylindrical body and laterally compressed tail (Fig. 3). Dorsal and anal fins are continuous with caudal fin. Dorsal fin commences above the anterior edge of gill opening and the predorsal length is 7.2���7.8 in total length (TL). Anal-fin origin immediately behind anus. Anus is located slightly before midbody and the preanal distance is 1.9���2.0 in TL. Head is broad and head length (HL) is 7.0��� 7.9 in TL. Snout blunt, snout length is 4.7���5.2 in HL. Both jaws are almost equal in size. All teeth concealed when mouth is closed. Eyes small, located almost at the midpoint between the snout and the rictus, eye diameter 11.6���12.1 in head length. Anterior nostril tubular and at lateral side of tip of snout, posterior nostril above anterior margin of eye. Gill opening a diagonal slit at midbody level. Head pores small with body colored rim. Two branchial pores located anterior to and below dorsal-fin origin well before gill opening (Fig. 4); three supraorbital pores, first (ethmoid) pore at tip of snout above lip, second above base of anterior nostril, third in between anterior and posterior nostrils; four infraorbital pores, first just below the base of anterior nostril, second in between anterior nostril and anterior margin of eye, third below the anterior margin of eye, and fourth after posterior margin of eye. There are six mandibular pores, all before rictus. Teeth pointed and serrated, uniserial in both jaws (Fig. 5). Intermaxillary teeth 5 on each side with one near symphysis, totaling 11 intermaxillary teeth, followed by 8 maxillary teeth on each side, no median intermaxillary teeth. Vomerine teeth uniserial with 8 teeth in holotype; 4���8 in paratypes. Mandibular teeth 18 on each side gradually decreasing in size posteriorly. Color. The fresh specimens were chocolate brown in color, overlain with small, irregular white spots, having golden shadings close to rictus. The spots start from the snout tip and extend to the entire body including the ventral portion, gill opening, dorsal and anal fins. Both dorsal and anal fins have chocolate brown color similar to body. Head region bears numerous, densely packed golden spots. The spots are widely spaced posteriorly and are comparatively larger than the spots in the head region (Fig. 6). Inside the mouth, the coloration is similar to the body. Tail tip is almost devoid of spots and without any white margin. Distribution. The specimen is presently known only from the holotype and three paratypes from Swaraj Dweep Island and Interview Island, Andaman Sea, India from a depth of 90���130 m depth. Etymology. Gymnothorax aurocephalus is named for the golden-colored head of the species., Published as part of Nashad, M., Mohapatra, Anil, Varghese, Sijo P., Ramalingam, L. & Mohanty, Swarup Ranjan, 2020, A new white-spotted moray eel, Gymnothorax aurocephalus sp. nov. (Muraenidae Muraeninae) from Andaman Sea, India, pp. 361-372 in Zootaxa 4877 (2) on pages 363-366, DOI: 10.11646/zootaxa.4877.2.8, http://zenodo.org/record/4424322

Gymnothorax unicolor (Delaroche 1809) ���Brown moray Taxonomy. First record from Lebanon as Gymnothorox unicolor (De la Roche 1809) by George et al. (1964: 13); subsequently recorded as Lycodontis unicolor (Delaroche) by Mouneimn�� (1977: 61); as Gymnothorax unicolor (Delaroche 1809) by Mouneimn�� (2002: 76). Material in collection: AUBM, FMNH (as Muraena unicolor) and USNM. Distribution. Mediterranean Sea, eastern Atlantic: Portugal to Azores, Madeira, Canary Islands; southern central Atlantic: Saint Helena. Conservation. IUCN: Global (LC: 20 October 2011); Med. (DD: 16 November 2007). Capture and threats: FIT, FIB, unknown. Occurrence: Common. Low priority for conservation action., Published as part of Bariche, Michel & Fricke, Ronald, 2020, The marine ichthyofauna of Lebanon: an annotated checklist, history, biogeography, and conservation status, pp. 1-157 in Zootaxa 4775 (1) on page 33, DOI: 10.11646/zootaxa.4775.1.1, http://zenodo.org/record/3983887, {"references":["George, C. J., Athanassiou, V. A. & Boulos, I. (1964) The fishes of the coastal waters of Lebanon. Miscellaneous Papers in the Natural Sciences. The American University of Beirut, 4, 1 - 24.","Mouneimne, N. (1977) Liste des poissons de la cote du Liban (Mediterranee orientale). Cybium, 3 (1), 37 - 66.","Mouneimne, N. (2002) Poissons marins du Liban et de la Mediterranee orientale. INCAM-EU / CNRS Lebanon, Beyrouth, 271 pp."]}

Gymnothorax elaineheemstrae Sithole, Smith & Gouws n. sp. Common name: Marbled leopard moray Figs. 3, 5; Table 9 Lycodontis undulatus (non Lacepède 1803): Smith 1949: 398 (Fig. 1129, Pl. 100); Smith 1953: 398 (Fig. 1129, Pl. 100); Smith 1962: 439 (Fig. E, Pl. 57); Castle & McCosker 1986: 173 (Fig. 41.24, Pl. 8). Gymnothorax cf. undulatus: King & Fraser 2014: 34, 35 (Fig. unnumbered) Holotype: SAIAB 86307 (624 mm TL), South Africa, KwaZulu-Natal, Cape Vidal, 28°9.8'S, 32°34.1'E, 6 Jan. 2009, coll. A. Paterson, G. Musson and A. Goetz. Paratypes: SAIAB 206655 (383 mm TL), same as holotype; CSIRO H 8399-01 (304 mm TL), South Africa, KwaZulu-Natal, Pennington, tide pools in front of the golf course; 30°24'S, 30°42'E, 29 Jul. 1992; AMS I.48960- 001 (665 mm TL), South Africa, Eastern Cape, Coffee Bay, 32°00'S, 29°12'E, 20 Jan. 1976; USNM 330982 (454 mm TL), South Africa, Natal, Aug. 1992. Non-type material: SAIAB 46644 (2: 285–329 mm TL), South Africa, KwaZulu-Natal, Mbibi, 27°30'S, 32°42'E, 12 Dec. 1988; SAIAB 4990 (2: 325–343 mm TL) South Africa, Eastern Cape, Mtentwana Point, 31°06'S, 30°12'E, 15 Aug. 1958; SAIAB 83082 (710 mm TL) South Africa, KwaZulu-Natal, Scottburgh 30°00'S, 31°00'E, 05 Jun. 2007; SAIAB 44497 (485 mm TL) South Africa, Eastern Cape, Algoa Bay, 34°00'S, 25°42'E, 01 Jan. 1967; SAIAB 39939 (644 mm TL) South Africa, Eastern Cape, Transkei Presley Bay, 31°54'S, 29°18'E, 01 Sep. 1974; SA- IAB 39959 (667 mm TL) South Africa, Eastern Cape, Hole-In-Hall, 32°00'S, 29°12'E, 27 Jan. 1975; SAIAB 83077 (2: 171–267 mm TL) South Africa, KwaZulu-Natal, Scottburgh, 30°17'S, 30°46'E and SAIAB 60166 (9: 85–349 mm TL), South Africa, KwaZulu-Natal, Scottburgh tide pool, 30°18'S 30°48'E, 18 Apr. 1999. Diagnosis. A moray eel with dorsal-fin origin anterior to gill opening, anal-fin origin just behind anus, with mottled and faintly reticulated color pattern on a dark brown background; three long intermaxillary canines in a median row; vomerine teeth small, in a single row; maxillary teeth biserial; vertebral count: 3–5 predorsal, 54–58 preanal, and 134–136 in total. Description. Proportions as % TL: preanal 41.1–50.8, predorsal 8.1–11.5, head length 11.3–15.0, depth at gill opening 4.2–8.4, depth at anus 3.7–6.5. As % head length: snout 15.2–18.5, eye diameter 10.0–11.8 and upper jaw 37.0–47.9. Vertebral: predorsal 3–5 (3), preanal 54–58 (56), total 134–136 (135). Body elongate, depth at gill opening 4.2–8.4 and 3.7–6.5 at anus in % TL; anus anterior to mid-body, preanal length 41.1–50.8 in % TL; head and trunk together shorter than the tail. Dorsal-fin origin anterior to gill opening, predorsal length 8.1–11.5 in % TL; anal-fin origin immediately behind anus; dorsal and anal fins continuous. Head long 11.3–15.0 in % TL, blunt anterior, concave above eye and broad behind the eye. Snout short, 15.2– 18.5 in % HL; upper jaw slightly longer than the lower jaw; anterior nostril tubular on each side of snout tip, posterior nostril round and slightly anterior to the eye. Three supraorbital pores, first pore situated just above the upper lip near tip of snout, second above the first pore and adjacent to the anterior nostril and one between anterior and posterior nostrils. Two branchial pores above and anterior to gill opening. Four infraorbital pores along the upper jaw, first between anterior nostril and anterior margin of the eye, second below anterior margin of the eye, third below the middle of the eye and the fourth below posterior edge of the eye. Five mandibular pores, all anterior to the rictus. No lateral-line pores posterior to the branchial ones. Gill opening slit-like and slightly below mid-body. Teeth slender and sharply pointed (Fig. 3); peripheral inter-maxillary teeth in single row of 6–9 (6) on each side; no intermediate inter-maxillary teeth; median inter-maxillary 2–3 (3), uniserial extremely long and widely spaced; vomerine teeth uniserial and small; inner maxillary teeth 1–4 (1), longer than outer maxillary teeth, anterior ones the largest, decreasing in size posteriorly; outer maxillary 12–16 (14) small teeth, closely spaced. Colour of fresh specimen and in preservation (Fig. 5). Head and body dark brown with mottled and faintly reticulated pattern; mottled coloration on the head extends inside the lower jaw; snout and ventral portion of the lower jaw to the beginning of anal fin paler compared to the rest of the body; dorsal- and anal-fins with dark brown margins. Distribution. Specimens of this species were collected in the Western Indian Ocean along the coast of South Africa from KwaZulu-Natal to the Eastern Cape, known to a depth of ~ 40 m in rock, intertidal and tide pools and sand bottoms. Etymology. The specific name, elaineheemstrae, is in honour of Mrs Elaine Heemstra (NRF-SAIAB Honorary Research Associate) for her tremendous support in mentoring the first author in fish taxonomy. Comparisons with other species. The dorsal-fin origin anterior to gill opening and anal-fin origin just after anus places this species in the genus Gymnothorax as currently recognized. Gymnothorax elaineheemstrae can be differentiated from other species within the genus in the Western Indian Ocean by its colour pattern and vertebral count. Gymnothorax elaineheemstrae has a mottled and faintly reticulated pattern (Fig. 5), whereas G. undulatus has irregular rectangular blotches separated by large reticulations (Fig. 4). Gymnothorax elaineheemstrae has a similar teeth arrangement to G. chilospilus Bleeker 1864, an Indo-Pacific species which G. undulatus has been confused with. However, G. chilospilus differs from G. elaineheemstrae in having fewer preanal (47–52 vs 54–58) and total vertebrae (120–129 vs 134–136); more peripheral intermaxillary teeth (6–8 vs 4–6) and clearly visible white markings on head (vs no white markings on head). The mottled pattern in G. elaineheemstrae is similar to G. flavimarginatus (Rüppell 1830), but G. elaineheemstrae has a faintly reticulated pattern and lacks the black blotch at the gill opening and the pale margin around posterior tail. Gymnothorax elaineheemstrae also differs from another closely similar species G. pictus (Ahl 1789) by the dorsal-fin origin which is anterior to gill opening (vs exactly above gill opening in G. pictus); uniserial teeth on vomer (vs. biserial); 3–5 (vs 7–11) predorsal and 54–58 (vs 56–62) preanal vertebrae. Remarks. The species retained the common name Marbled leopard moray given by King & Fraser (2014). Genetics. The COI analysis gave an aligned data set of 411 base pairs (bp) in length for 50 sequences. The neighbour-joining tree (Fig. 6) based on COI sequences revealed two distinct clades among specimens formerlyregarded as G. undulatus. Gymnothorax undulatus was made up of three lineages (Clades A, B and C) and was geographically widespread, comprising samples from the Pacific Ocean and several WIO regions. These lineages represented regions that were not distinct in the morphological analysis. Specimens from all these geographic regions had the typical G. undulatus color pattern, i.e. dark brown body with rectangular spots separated by pale reticulum. The second clade comprised Gymnothorax elaineheemstrae and was nested within the morphologically-distinct banded moray G. rueppelliae (McClelland 1844). Gymnothorax elainheemstrae is restricted to South Africa and comprises of specimens that are mottled and less reticulated. The most appropriate nucleotide substitution model for the COI data was determined to be a GTR model (Rodríguez et al. 1990), with proportion of invariable sites (I = 0.638), base frequencies of A = 0.284, C = 0.290, G = 0.157 and T = 0.269, and a rate matrix of R [A↔C] = 1.247, R [A↔ G] = 19.982, R [A↔T] = 3.994, R [C↔G] = 1.790, R [C↔T] = 18.676 and R [G↔T] = 1.000. The ML tree constructed with a loglikelihood (lnL) of -2829.731 (not shown) indicated the genetic distinction of G. undulatus and G. elaineheemstrae, similar to neighbour joining tree. Enforcing the topology to reflect the monophyly of the former G. undulatus (i.e., G. undulatus + G. elaineheemstrae as sister taxa, maintaining all internal relationships) resulted in a log-likelihood of -2840.998; this topology was significantly less likely (∆lnL = 11.266, P Gymnothorax based on COI sequences ranged from 0.073 to 0.267 (Table 10). The lowest genetic distance (0.073) was between the paintspotted moray G. pictus and lipspot moray G. chilospilus, and this was the only genetic distance G. griseus and brown spotted moray G. fuscomaculatus. The genetic distance (K2P) between G. undulatus and G. elaineheemstrae was high at 0.201. The 16S rRNA analysis involved an alignment of 404 nucleotides for 44 sequences, including three sequences of G. undulatus from Japan (Pacific Ocean). The optimal model for this dataset included a model with base frequencies of A = 0.339, C = 0.260, G = 0.196 and T = 0.205, a rate matrix of R [A↔C] = 1.277, R [A↔G] = R [C↔T] = 5.725 and R [A↔ T] = R [C↔G] = R [G↔T] = 1.000, a proportion of invariable sites (I = 0.367) and a gamma distribution of rate variation (α = 0.477). The NJ tree (Fig. 7) recovered a clade of G. undulatus that was widespread, containing samples from the Archipelago, East Africa, Seychelles and Japan, and sister to G. rueppelliae. Gymnothorax elaineheemstrae was distinct and phylogenetically divergent from the G. undulatus clade and was well-supported with 100% bootstrap. The ML tree (lnL = -2490.389; not shown) also revealed G. undulatus and G. elaineheemstrae to form distinct and phylogenetically well-separated clades. As with the COI data, enforcing the topologies to reflect the monophyly of the former G. undulatus (with G. undulatus placed as a sister taxon to G. elaineheemstrae) presented a significantly worse fit to the data (lnL = -2512.244, ∆lnL = 21.855, P G. thyrsoideus and the banded moray G. rueppelliae, and the lowest value (0.003) was between G. griseus and G. thyrsoideus, possibly indicating the misidentification of the former (see Fig. 7). The next lowest value (0.190) was between G. chilospilus and G. flavimarginatus. The genetic distance from the comparison between G. elaineheemstrae and the G. undulatus clades was 0.234. Average divergences of 0.021 to 0.081 separated potential Archipelago, widespread and Japanese (Pacific Ocean) subclades and lineages within the G. undulatus clade (Fig. 7)., Published as part of Sithole, Yonela, Smith, David G., Mwale, Monica & Gouws, Gavin, 2020, A taxonomic revision of Gymnothorax undulatus (Anguilliformes: Muraenidae) in the Western Indian Ocean, with description of a new species, pp. 161-181 in Zootaxa 4767 (1) on pages 170-174, DOI: 10.11646/zootaxa.4767.1.7, http://zenodo.org/record/3770681, {"references":["Lacepede, B. G. E. (1803) Histoire naturelle des poisons. Vol. 5. Chez F. G. Levrault, Paris, cxi + 414 pp.","Smith, J. L. B. (1949) The Sea fishes of Southern Africa. Central News Agency, LTD, Cape Town, xvi + 550 pp., 100 pls.","Smith, J. LB. (1953) The Sea fishes of Southern Africa. 3 rd Edition. Central News Agency, LTD, Cape Town, 564 pp., 107 pls.","Smith, J. L. B. (1962) The moray eels of the Western Indian Ocean and the Red Sea. Ichthyological Bulletin, 23, 421 - 444.","Castle, P. H. J. & McCosker, J. E. (1986) Family Muraenidae. In: Smith, M. M. & Heemstra, P. C. (Eds.), Smiths' Sea Fishes. MacMillan, Johannesburg, pp. 165 - 176.","King, D. & Fraser, V. (2014) The reef guide-fishes, corals, nudibranchs & other invertebrates: East & South coasts of southern Africa. Struik Nature, Cape Town, 360 pp.","Bleeker, P. (1864) Poissons inedits indo-archipelagiques de l'ordre des Murenes. Nederlandsch Tijdschrift voor de Dierkunde, 2, 38 - 54.","Ruppell, W. P. E. S. (1830) Atlas zu der Reise im nordlichen Afrika. Fische des Rothen Meers. Frankfurt am Main (Heinrich Ludwig Bronner), 95 - 141.","Ahl, J. N. (1789) Specimen ichthyologicum de Muraena et Ophichtho, quod venia exp. Fac. Med. Ups. Praeside Carol. Pet. Thunberg, etc. J. Edman, Upsaliae. 1 - 14.","McClelland, J. (1844) Apodal fishes of Bengal. Calcutta Journal of Natural History, 5, 151 - 226.","Rodriguez, F., Oliver, J. F., Marin, A. & Medina, J. R. (1990) The general stochastic model of nucleotide substitution. Journal of Theoretical Biology, 142, 485 - 501. https: // doi. org / 10.1016 / S 0022 - 5193 (05) 80104 - 3"]}

The taxonomic status of the widely distributed Indo-Pacific undulated moray eel, Gymnothorax undulatus, is revised using morphological and genetics features. Ninety-seven specimens previously identified as G. undulatus were examined and their mitochondrial COI and 16S rRNA genes were analysed. The multivariate analysis of eight morphometric characters resulted in separation with little to no overlap among some geographic regions. These groupings explained more than 90% of the total variation, with 86.6% overall classification. Two color morphs were identified, and the South African population was described as new species, Gymnothorax elaineheemstrae n. sp., distinct from G. undulatus in having mottled and faintly reticulated color pattern, 134–136 total vertebrae and further confirmed by the genetic analysis of COI and 16S rRNA with > 0.1 genetic distance. The morphological and genetics results indicate that G. undulatus, previously treated as a single species, consists of more than one species.

Gymnothorax minor is a moray eel of the family Muraenidae found in the Western Pacific Ocean. We report here its complete mitogenome as determined by Illumina next-generation sequencing and the phylogenetic relationship with its congeners and other taxa of the family Muraenidae. The whole mitogenome of G. minor had a total length of 16,574 bp, comprising 37 genes - 13 protein-coding genes (PCGs), two ribosomal ribonucleic acid (rRNA) and 22 transfer ribonucleic acid (tRNA) genes - and a control region. Excepting cox1 with GTG, the other 12 PCGs had ATG start codon. Seven of its PCGs had incomplete stop codon - five (nad2; cox1; cox2; nad3 and nad4) with T and two (atp6 and cox3) with TA. Molecular phylogeny based on 13 PCGs was concordant with 15 mitochondrial genes (13 PCGs and 2 rRNA genes). The subfamily Muraeninae as well as the subfamily Uropterygiinae were monophyletic. However, the genus Gymnothorax was paraphyletic, with G. minor forming a sister group with Rhinomuraena quaesita in the lineage containing also G. kidako and G. formosus forming a sister group with Enchelynassa canina. The phylogenetic relationship of the genus Gymnothorax and related taxa of the family Muraenidae, based on the mitochondrial cob gene, was in general similar to that based on 15 mt-genes. The mitogenome is useful for future studies on phylogenetics and systematics of eels of the family Muraenidae and other taxa of the order Anguilliformes.

Gymnothorax margaritophorus Bleeker. Indonesia: BMNH 1867.11.28.268 (1, 207, holotype). Samoa: USNM 51713 (1, 253, holotype of Gymnothorax talofa Jordan & Starks). French Polynesia, Moorea: MNHN 2008-0437 (1, 72); MNHN 2008-0438 (1, 72); MNHN 2008-0439 (1, 63); MNHN 2008-0442 (1, 290); MNHN 2008-0443 (1, 290); MNHN 2008-0444 (1, 219); MNHN 2008-0446 (1, 57). Gambier Is.: USNM 401788 [GAM-412] (1, 457). Manua’e (Scilly) I.: USNM 435145 [SCIL-256] (1, 168)., Published as part of Smith, David G., Bogorodsky, Sergey V., Mal, Ahmad O. & Alpermann, Tilman J., 2019, Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea, with description of a new species, pp. 1-87 in Zootaxa 4704 (1) on page 42, DOI: 10.11646/zootaxa.4704.1.1, http://zenodo.org/record/3563576

Gymnothorax angusticauda (Weber & de Beaufort 1916) —Shorttail Moray (Figure 8) Muraena (Priodonophis) angusticauda Weber & de Beaufort 1916: 389, fig. 388 (Near Supiori, Schouten Is., Papua New Guinea). Holotype (unique), ZMA 102162. Gymnothorax angusticauda: Randall & Golani 1995: 854, Pl. 1a; Golani & Bogorodsky 2010: 10; Bogorodsky et al. 2014: 411; Golani & Fricke 2018: 20. Red Sea material. Egypt: BPBM 19844 (2, 458–503), Nuweiba. Saudi Arabia: SMF 34962 [KAU12-731] (1, 383), Jizan. Comparative material. Taiwan: USNM 438183 (1, 486); USNM 438650 (1, 547); USNM 439078 (1, 513). Philippines: USNM 408880 (1, 560). Papua New Guinea: ZMA 102.062 (1, 474, holotype). Description. In TL: preanal length 1.9–2.2, predorsal length 11–13, head length 8.0–9.8, body depth at anus 25–34. In head length: snout length 5.3–7.1, eye diameter 8.8–11, upper-jaw length 2.8–3.5. Pores: LL 2, SO 3, IO 4, POM 6. Vertebrae: predorsal 4, preanal 58–70, total 143–152. Body moderately elongate; anus near mid-length; dorsal fin begins slightly before gill opening; tail relatively slender. Head and jaws moderate, jaws closing completely. Eye well developed, at about midpoint of upper jaw. Gill opening nearly horizontal, midlateral. Anterior nostril tubular, reaches edge of lip when depressed; posterior nostril oval, with a slightly raised rim, over anterior part of eye. Teeth conical to narrowly triangular, finely serrate on posterior edge. Upper jaw with about 5 or 6 peripheral intermaxillary teeth, narrowly triangular, retrorse; 0–3 median teeth, conical. Maxillary teeth uniserial, about 7–14, similar in shape to intermaxillary teeth. Dentary teeth uniserial, similar in shape to those in upper jaw, with about 7–19 teeth on each side. Vomer with about 3–6 teeth in a single row. Color: medium to light brown or tan, snout and lower jaw darker, abdomen sometimes white; head pores in conspicuous white spots, nostrils white; fins with narrow pale edge; iris yellow. Of the ten known specimens, the largest is 560 mm. Distribution and habitat. Known from the Red Sea, Taiwan, the Philippines, Indonesia (Bali and Sulawesi), and Papua New Guinea. The two larger specimens from the Red Sea were collected in less than 0.5 m from the fringing reef at Nuweiba. The smaller specimen was captured in a trawl at about 30 m from the southern Saudi Arabia, off Jizan, from a sandy area close to the island. Information on depth and habitat is not available for the Philippine and Indonesian specimens. Fricke (2015) reported an additional specimen collected from a depth of 15 m from Madang, Papua New Guinea. Remarks. The smallest trawled Red Sea specimen differs from all the other specimens in having the anus slightly behind midlength rather than slightly before. It also has more total vertebrae (152 vs. 143–148) and distinctly more preanal vertebrae (70 vs. 58–60). The other two Red Sea specimens do not differ in these characters from the western Pacific specimens. With the limited material available and without genetic information from other parts of the distribution area, we cannot assess the significance of these distinctions. In the COI-based phylogeny, the species is most closely related to Gymnothorax albimarginatus (Temminck & Schlegel) and an unidentified Gymnothorax species (Gymnothorax sp. 4, BOLD voucher of SBF244-11 collected from Madagascar), with which it forms a well-supported clade., Published as part of Smith, David G., Bogorodsky, Sergey V., Mal, Ahmad O. & Alpermann, Tilman J., 2019, Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea, with description of a new species, pp. 1-87 in Zootaxa 4704 (1) on pages 17-18, DOI: 10.11646/zootaxa.4704.1.1, http://zenodo.org/record/3563576, {"references":["Weber, M. & de Beaufort, L. F. (1916) The Fishes of the Indo-Australian Archipelago. III. Ostariophysi: II Cyprinoidea, Apodes, Synbranchi. E. J. Brill, Leiden, xv + 455 pp.","Randall, J. E. & Golani, D. (1995) Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea. Bulletin of Marine Science, 56 (3), 849 - 880.","Golani, D. & Bogorodsky, S. V. (2010) The fishes of the Red Sea-reappraisal and updated checklist. Zootaxa, 2463, 1 - 135. https: // doi. org / 10.11646 / zootaxa. 2463.1.1","Bogorodsky, S. V., Alpermann, T. J., Mal, A. O. & Gabr, M. H. (2014) Survey of demersal fishes from southern Saudi Arabia, with five new records for the Red Sea. Zootaxa, 3852 (4), 401 - 437. https: // doi. org / 10.11646 / zootaxa. 3852.4.1","Golani, D. & Fricke, R. (2018) Checklist of the Red Sea fishes with delineation of the Gulf of Suez, Gulf of Aqaba, endemism and Lessepsian migrants. Zootaxa, 4509 (1), 1 - 215.","Fricke, R. (2015) Twenty-one new records of fish species (Teleostei) from Madang and Papua New Guinea (western Pacific Ocean). Marine Biodiversity Records, 8, 1 - 9. https: // doi. org / 10.1017 / S 1755267215000445"]}

Gymnothorax favagineus Bloch & Schneider 1801 —Honeycomb Moray (Figure 14) Gymnothorax favagineus Bloch & Schneider 1801: 525, pl. 105 (Tranquebar, India). Holotype (unique), ZMB 7782 (stuffed).— Randall 1994: 260; Randall & Golani 1995: 858; Golani & Bogorodsky 2010: 10; Golani & Fricke 2018: 21. Red Sea material. None. Comparative material. Taiwan: USNM 312734 (1, 303); USNM 312736 (2, 182–278). Australia: USNM 176690 (2, ca 600–614); USNM 312733 (1, 304); USNM 312735 (1, 290). Description. In TL: preanal length 2.0–2.2, predorsal length 9.4–11, head length 7.6–8.8, body depth at anus 19–24. In head length: snout length 5.1–6.7, eye diameter 9.4–11, upper-jaw length 2.5–3.0. Pores: LL 2, SO 3, IO 4, POM 6. Vertebrae: predorsal 5–6, preanal 61–64, total 139–148. Body moderately elongate, anus slightly before midlength, dorsal-fin origin before gill opening. Jaws moderate, of equal length. Eye moderate, over middle of upper jaw. Anterior nostril tubular; posterior nostril in a low tube, over or just in front of anterior edge of eye. Teeth smooth, conical to narrowly triangular; intermaxillary teeth in a single peripheral series of about 4 on each side, 2 or 3 median teeth. Maxillary teeth biserial, 2–3 larger inner teeth and about 1 1–15 smaller outer teeth. Dentary with 2 or 3 large teeth anteriorly followed by about 11–14 smaller teeth, the larger teeth sometimes distinctly medial to smaller teeth. Vomerine teeth uniserial in young, biserial in adults. Color: body and head white covered with numerous, polygonal, black spots separated by narrow interspaces often forming a honeycomb-like pattern. Spots larger and more widely separated in young. Maximum size at least 2 m. Distribution and habitat. Widely distributed in the Indo-West Pacific from the southern Red Sea south to South Africa, east to Australia and Papua New Guinea. Occurs on coral and rocky reefs from depths of 1–50 m; feeds on fishes and octopuses. Remarks. The record from the Red Sea is based on a photograph taken at Hanish Island off Yemen (Randall 1994). The three specimens examined from Taiwan have fewer vertebrae (139–141) than the four specimens from Australia (146–148). This species has been confused with other dark-spotted species such as Gymnothorax isingteena (Richardson) and G. melanospilus (Bleeker). The species was not collected during the present study, and no tissue samples or COI sequence data for G. favagineus specimens from the Red Sea are available at present. The one sequence of Gymnothorax favagineus included in the phylogenetic analysis (Fig. 48) and other near identical sequences from the southwestern Indian Ocean that are deposited in BOLD (not included in the present analysis) do not differ markedly from sequences from specimens collected in distant parts of the distribution area of the species (e.g. Taiwan, Indonesia and Western Australia) indicating low levels of intra-specific genetic differentiation (not shown in the present phylogeny, Fig. 48). The closest phylogenetic relationship, although only weakly supported by bootstrapped analyses, is with Gymnothorax formosus Bleeker (represented by a specimen from the Society Islands in the present analysis, see Fig. 48)., Published as part of Smith, David G., Bogorodsky, Sergey V., Mal, Ahmad O. & Alpermann, Tilman J., 2019, Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea, with description of a new species, pp. 1-87 in Zootaxa 4704 (1) on pages 26-27, DOI: 10.11646/zootaxa.4704.1.1, http://zenodo.org/record/3563576, {"references":["Bloch, M. E. & Schneider, J. G. (1801) M. E. Blochii […] Systema ichthyologiae iconibus CX illustratum. Post obitum auctoris opus inchoatum absolvit, correxit, interpolavit Jo. Gottlob Schneider, Saxo, LX + 584 pp. https: // doi. org / 10.5962 / bhl. title. 5750","Randall, J. E. (1994) Twenty-two new records of fishes from the Red Sea. Fauna of Saudia Arabia, 14, 259 - 275.","Randall, J. E. & Golani, D. (1995) Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea. Bulletin of Marine Science, 56 (3), 849 - 880.","Golani, D. & Bogorodsky, S. V. (2010) The fishes of the Red Sea-reappraisal and updated checklist. Zootaxa, 2463, 1 - 135. https: // doi. org / 10.11646 / zootaxa. 2463.1.1","Golani, D. & Fricke, R. (2018) Checklist of the Red Sea fishes with delineation of the Gulf of Suez, Gulf of Aqaba, endemism and Lessepsian migrants. Zootaxa, 4509 (1), 1 - 215."]}

Gymnothorax randalli Smith & Böhlke 1997 —Randall’s Moray (Figure 35) Gymnothorax punctatofasciatus (non Bleeker): Randall & Golani, 1995: 865. Gymnothorax randalli Smith & Böhlke 1997: 185, figs. 1, 2, 8 (Sorongjunkong, Lombok, Indonesia). Holotype, BPBM 30138. — Golani & Bogorodsky 2010: 10; Bogorodsky et al. 2014: 411; Golani & Fricke 2018: 22. Red Sea material. Egypt: HUJ 9410 (1, 365), Nuweiba, May 1976. Saudi Arabia: SMF 34963 [KAU12-732] (1, 396), Jizan, 20–25 m, 1 Mar 2012. Comparative material. Indonesia: BPBM 30138 (1, 324, holotype); ANSP 175205 (1, 268, paratype); USNM 343860 (1, 317, paratype). Description. In TL: preanal length 2.2–2.4, predorsal length 8.9–11, head length 7.8–8.6, body depth at anus 21–29. In head length: snout length 5.7–7.6, eye diameter 7.5–10, upper-jaw length 2.8–3.3. Pores: LL 2, SO 3, IO 4, POM 6. Vertebrae: predorsal 5–9, preanal 48–52, total 124–130. Body moderately elongate; anus before midlength; dorsal-fin origin before gill opening. Snout moderately short, jaws of equal length. Eye moderate, over middle of upper jaw. Anterior nostril tubular; posterior nostril above anterior part of eye. Teeth conical to triangular, pointed, finely serrate on posterior edge. Intermaxillary teeth in a single peripheral series, 6 on each side; 0–3 median teeth. Maxillary teeth uniserial or biserial, 0–3 inner and 12–20 outer. Dentary teeth uniserial or biserial, with 0–5 larger inner teeth anteriorly, 16–24 smaller outer teeth. Vomerine teeth small, uniserial or biserial, about 6–12. Color: body pale yellowish with two longitudinal rows of 35–39 irregular dark brown spots, most notably larger than orbit and in vertical alignment with spot of other row; a third row of irregular dark brown spots midventrally, these merging with spot above to form short bars on posterior two-thirds of tail; pale interspaces between large spots with numerous small dark brown spots of variable size, except ventrally. Head with numerous dark brown spots of variable size, a few dorsally on posterior half of head larger than orbit. Iris white. Anterior nostril pale. Maximum size at least 396 mm. Distribution and habitat. A rare species known from only five specimens from southern Indonesia and the Red Sea. Smith & Böhlke (1997: 186) pointed out that the specimen from Madras, India illustrated by Day (1878, pl. 169, fig. 4) as Muraena punctatofasciata appears to be Gymnothorax randalli, indicating that the species probably occurs continuously across the northern Indian Ocean to Indonesia. Probably living in open areas on silty sand substrata. The specimen from Saudi Arabia off Jizan was trawled from at a depth of 20–25 m in open area. One individual was photographed in sand area covered with seagrasses at depth of about 15 m at Nuweiba, Egypt (Sonja Ooms, pers. comm.). Remarks. The two Red Sea specimens differ slightly in predorsal (5–7), preanal (48–49), and total (124–126) vertebrae from the Indonesian ones (7–9, 50–52, 127–130, respectively). SMF 34963 has a short row of distinctly biserial teeth on the vomer. The vomerine teeth of the other specimens are uniserial. We have no genetic data from outside the Red Sea. The sequence of the Red Sea specimen of G. randalli is part of a highly supported clade with G. minor (Temminck & Schlegel), G. cf. minor and G. mccoskeri Smith & Böhlke, the latter being phylogenetically closest to G. randalli (Fig. 48)., Published as part of Smith, David G., Bogorodsky, Sergey V., Mal, Ahmad O. & Alpermann, Tilman J., 2019, Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea, with description of a new species, pp. 1-87 in Zootaxa 4704 (1) on pages 54-55, DOI: 10.11646/zootaxa.4704.1.1, http://zenodo.org/record/3563576, {"references":["Smith, D. G. & Bohlke, E. B. (1997) A review of the Indo-Pacific banded morays of the Gymnothorax reticularis group, with descriptions of three new species (Pisces, Anguilliformes, Muraenidae). Proceedings of the Academy of Natural Sciences of Philadelphia, 148, 177 - 188.","Randall, J. E. & Golani, D. (1995) Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea. Bulletin of Marine Science, 56 (3), 849 - 880.","Golani, D. & Bogorodsky, S. V. (2010) The fishes of the Red Sea-reappraisal and updated checklist. Zootaxa, 2463, 1 - 135. https: // doi. org / 10.11646 / zootaxa. 2463.1.1","Bogorodsky, S. V., Alpermann, T. J., Mal, A. O. & Gabr, M. H. (2014) Survey of demersal fishes from southern Saudi Arabia, with five new records for the Red Sea. Zootaxa, 3852 (4), 401 - 437. https: // doi. org / 10.11646 / zootaxa. 3852.4.1","Golani, D. & Fricke, R. (2018) Checklist of the Red Sea fishes with delineation of the Gulf of Suez, Gulf of Aqaba, endemism and Lessepsian migrants. Zootaxa, 4509 (1), 1 - 215.","Day, F. (1878) The fishes of India; being a natural history of the fishes known to inhabit the seas and fresh waters of India, Burma, and Ceylon Part 4. Bernard Quaritch, London, xx + 227 pp. [pp. i-xx + 553 - 779] https: // doi. org / 10.5962 / bhl. title. 55567"]}

Gymnothorax reticularis Bloch 1795 —Reticulate Moray (Figure 36) Gymnothorax reticularis Bloch 1795: 85, pl. 416 (Coromandel coast, India). Holotype (unique), ZMB 3986.— Randall & Golani 1995: 867; Golani & Bogorodsky 2010: 10; Bogorodsky et al. 2014: 411; Golani & Fricke 2018: 23. Muraena reticulata: Rüppell 1830: 117. Red Sea material. Israel: HUJ 14667 (1, 240), Eilat. Saudi Arabia: KAUMM 415 [KAU14-155] (1, 400), Jizan, 60–65 m, 01 Nov 2014; SMF 34964 [KAU12-733] (1, 350), Jizan, 55–60 m, 29 Feb 2012. Comparative material. Mozambique: SAIAB 5052 (1, 346). Pakistan: USNM 427632 (1, 297). India: ZMB 3986 (1, 302, holotype), Coromandel (Tranquebar); ANSP 127987 (3, 207–256); USNM 343723 (1, 200). Myanmar: USNM 438261 (1, 271). Description. In TL: preanal length 2.0–2.2, predorsal length 7.1–11, head length 6.5–8.2, body depth at anus 19–29. In head length: snout length 5.2–7.8, eye diameter 8.1–13, upper-jaw length 2.9–3.5. Pores: LL 2, SO 3, IO 4, POM 6. Vertebrae: predorsal 5–7, preanal 48–53, total 115–126. Body moderately elongate; anus at or slightly before midlength; dorsal-fin origin before gill opening. Snout moderately short, jaws of equal length. Eye moderate, over middle of upper jaw. Anterior nostril tubular; posterior nostril above anterior part of eye. Teeth uniserial, conical to triangular, pointed, serrate. Intermaxillary teeth in a single peripheral series, 3–6 on each side; no median teeth. Maxillary teeth uniserial, 3–11. Dentary teeth uniserial except 1 specimen with 2 inner teeth on one side, 7–15. Vomerine teeth small and inconspicuous, uniserial but biserial in 1 specimen, about 6–12. Color: body pale gray with yellowish hue dorsally, with 16–20 dark brown bars on ventral two-thirds of body, narrower than pale interspace, obscured dorsally by numerous, irregular, small spots, some forming a reticular pattern. Dorsal and anal fins alternating with black and white or pale gray, spots on body extending onto white interspaces basally in fins. Head pale gray with dark brown spots of variable size that are longitudinally elongate posteriorly. Maximum size about 460 mm (specimen from the Mediterranean Sea). Distribution and habitat. Known from India, Pakistan, Mozambique, and the Red Sea. Both Red Sea specimens trawled off Jizan from soft substrata, from a depth of 50– 65 m. Reported also from the eastern Mediterranean Sea by Stern & Goren (2013). Remarks. This species was long confused with Gymnothorax minor. The true Gymnothorax reticularis is restricted to the Indian Ocean; records from the Pacific refer to G. minor (Smith & Böhlke 1997). Specimens from the Red Sea have more preanal (51–53) and total (123–126) vertebrae than those from elsewhere (48–50 and 115–122). The Mozambique specimen is intermediate with 122 vertebrae vs. ca 115–120 in those from India and Pakistan.Also in the Red Sea specimens, the bars are less distinct dorsally, where they tend to be lost in the general spotting. We have no genetic data from outside the Red Sea. Smith & Böhlke (1997: 187) reported that the specimen described by J.L.B. Smith (1962: 429) from Mozambique could not be located. However, the first author found it during his visit to South Africa in 2001 and took the relevant counts and measurements. Li et al. (2018) demonstrated that previous records of specimens identified as G. reticularis in the western Pacific are based on misidentification of G. minor. No COI sequences were available for G. reticularis from outside the Red Sea. Sequences of the mitochondrial COI from two Red Sea specimens that were collected during the present study were included in the multi-sequence alignment. In the resulting phylogenetic tree (Fig. 48), G. reticularis did not show a close phylogenetic affiliation with any other clade of moray eels included in the analysis., Published as part of Smith, David G., Bogorodsky, Sergey V., Mal, Ahmad O. & Alpermann, Tilman J., 2019, Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea, with description of a new species, pp. 1-87 in Zootaxa 4704 (1) on pages 55-57, DOI: 10.11646/zootaxa.4704.1.1, http://zenodo.org/record/3563576, {"references":["Bloch, M. E. (1795) Naturgeschichte der auslandischen Fische. Vol. 9. Auf Kosten des Verfassers und in Commission bei dem Buchhandler Hr. Hesse, Berlin, II + 192 pp.","Randall, J. E. & Golani, D. (1995) Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea. Bulletin of Marine Science, 56 (3), 849 - 880.","Golani, D. & Bogorodsky, S. V. (2010) The fishes of the Red Sea-reappraisal and updated checklist. Zootaxa, 2463, 1 - 135. https: // doi. org / 10.11646 / zootaxa. 2463.1.1","Bogorodsky, S. V., Alpermann, T. J., Mal, A. O. & Gabr, M. H. (2014) Survey of demersal fishes from southern Saudi Arabia, with five new records for the Red Sea. Zootaxa, 3852 (4), 401 - 437. https: // doi. org / 10.11646 / zootaxa. 3852.4.1","Golani, D. & Fricke, R. (2018) Checklist of the Red Sea fishes with delineation of the Gulf of Suez, Gulf of Aqaba, endemism and Lessepsian migrants. Zootaxa, 4509 (1), 1 - 215.","Ruppell, W. P. E. S. (1830) Atlas zu der Reise im nordlichen Afrika. Fische des Rothen Meers. Heinrich Ludwig Bronner, Frankfurt am Main, 141 pp.","Stern, N. & Goren, M. (2013) First record of the moray eel Gymnothorax reticularis Bloch, 1795 in the Mediterranean Sea, with a note on its taxonomy and distribution. Zootaxa, 3641 (2), 197 - 200. https: // doi. org / 10.11646 / zootaxa. 3641.2.8","Smith, D. G. & Bohlke, E. B. (1997) A review of the Indo-Pacific banded morays of the Gymnothorax reticularis group, with descriptions of three new species (Pisces, Anguilliformes, Muraenidae). Proceedings of the Academy of Natural Sciences of Philadelphia, 148, 177 - 188.","Smith, J. L. B. (1962) The moray eels of the western Indian Ocean and the Red Sea. Ichthyological Bulletin, Department of Ichthyology, Rhodes University, 23, 421 - 444.","Li, Y., Zhang, L., Zhao, L., Feng, J., Loh, K. H., Zheng, X. & Lin, L. (2018) New identification of the moray eel Gymnothorax minor (Temminck & Schlegel, 1846) in China (Anguilliformes, Muraenidae). ZooKeys, 752, 149 - 161. https: // doi. org / 10.3897 / zookeys. 752.24231"]}

Gymnothorax baranesi Smith, Brokovich & Einbinder 2008 —Barane’s Moray (Figure 9) Gymnothorax baranesi Smith, Brokovich & Einbinder 2008: 63, figs. 1–4 (Off Eilat, Israel). Holotype, HUJ 18976.— Golani & Bogorodsky 2010: 10; Golani & Fricke 2018: 20. Gymnothorax sp.: Khalaf & Disi 1997: 40. Red Sea material. Israel: HUJ 18976 (1, 857, holotype), Eilat; HUJ 18975 (1, 762, paratype), Eilat; USNM 388603 (1, 828, paratype), Eilat. Description. In TL: preanal length 2.0–2.1, predorsal length 7.9–8.1, head length 6.6–7.5, body depth at anus 14–19. In head length: snout length 4.8–5.7, eye diameter 11–13, upper -jaw length 2.5–2.6. Pores: LL 2, SO 3, IO 4, POM 6. Vertebrae: predorsal 6–7, preanal 52–55, total 137–142. Body moderate, anus near midlength; dorsal fin begins before gill opening. Head and jaws moderate, jaws not closing completely. Eye moderate in size, over middle of upper jaw. Gill opening midlateral.Anterior nostril tubular, short, not reaching edge of lip when depressed; posterior nostril elliptical, without raised rim, over anterior part of eye. Teeth sharp, conical to triangular, smooth. Intermaxillary with a peripheral series of 5–7 teeth on each side and 1–3 long, slender, median teeth. Maxillary teeth biserial or uniserial, the inner row with two long depressible teeth and the outer row with 14–17 smaller, triangular, retrorse teeth, continuous with peripheral intermaxillary teeth. Dentary teeth uniserial, larger anteriorly, conical to triangular. Vomer with 3–12 small teeth, in a single staggered row. Color: brown, covered with moderate-size pale spots in approximately three rows, largest around midbody, the spots rosette-like anteriorly, irregularly rounded near end of tail; spots much smaller on head, inconspicuous or absent on snout and lower jaw; spots extending onto fins; tubular anterior nostril dark brown or black; gill opening darker than surrounding area; a narrow dark blotch at angle of jaw. Maximum size at least 800–900 mm. Distribution and habitat. Apparently endemic to the Red Sea, in relatively deep water, approximately 200 m, all known records from the Gulf of Aqaba. Remarks. This species resembles Gymnothorax pharaonis n. sp. (see below) in its color pattern, but in the latter species the markings on the tail are more irregular in shape and tend to be drawn out into oblique streaks dorsally and ventrally. In addition, G. pharaonis has fewer vertebrae (122–128 vs. 137–142) and lives in shallower water. The teeth in G. baranesi are sexually dimorphic, the males having larger and fewer teeth and lacking the inner maxillary teeth. Gymnothorax baranesi was not collected during the present study and no COI sequence information was available for this species, so it could not be included in the phylogenetic analysis carried out in this study., Published as part of Smith, David G., Bogorodsky, Sergey V., Mal, Ahmad O. & Alpermann, Tilman J., 2019, Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea, with description of a new species, pp. 1-87 in Zootaxa 4704 (1) on pages 19-20, DOI: 10.11646/zootaxa.4704.1.1, http://zenodo.org/record/3563576, {"references":["Smith, D. G., Brokovich, E. & Einbinder, S. (2008) Gymnothorax baranesi, a new moray eel (Anguilliformes: Muraenidae) from the Red Sea. Zootaxa, 1678, 63 - 68. https: // doi. org / 10.11646 / zootaxa. 1678.1.4","Golani, D. & Bogorodsky, S. V. (2010) The fishes of the Red Sea-reappraisal and updated checklist. Zootaxa, 2463, 1 - 135. https: // doi. org / 10.11646 / zootaxa. 2463.1.1","Golani, D. & Fricke, R. (2018) Checklist of the Red Sea fishes with delineation of the Gulf of Suez, Gulf of Aqaba, endemism and Lessepsian migrants. Zootaxa, 4509 (1), 1 - 215.","Khalaf, M. A. & Disi, A. M. (1997) Fishes of the Gulf of Aqaba. Marine Science Station Aqaba, Jordan, 252 pp."]}

Gymnothorax pindae Smith 1962 —Pinda Moray (Figure 32) Gymnothorax pindae Smith 1962: 430, pl. 55 (fig. D) (Pinda, Mozambique). Holotype (unique), SAIAB 105.— Randall & Golani 1995: 865; Golani & Bogorodsky 2010: 10; Golani & Fricke 2018: 22. Red Sea material. Red Sea : USNM 191669 (1, 251). Egypt: USNM 312698 (4, 145–210), El Himeira, Gulf of Aqaba. Saudi Arabia: KAUMM 405 [KAU 12-1088] (1, 127), Al Lith; KAUMM 406 [KAU 13-489] (1, 222), Al Wajh; KAUMM 407 [KAU 13-596] (1, 104), Duba; KAUMM 408 (KAU 14-993), (1, 162), Al Lith; KAUMM 414 [KAU 13-692] (1, 103), Jeddah, Obhur; SMF 35169 (1, 340), Duba; SMF 35398 [KAU 13-352] (1, 323), Al Wajh; SMF 35818 [KAU 12-1027] (1, 210), Al Lith; SMF 35819 [KAU 13-447] (1, 304), Al Wajh; SMF 35820 [KAU 13- 595] (1, 198), Duba; SMF 35827 [KAU 13-693] (1, 252), Jeddah, Obhur. Comparative material. Mauritius : USNM 312725 (1, 99). Philippines: USNM 315563 (1, 144). Vanuatu: USNM 363336 (2, 50–146); USNM 363689 (1, 258). French Polynesia, Mururoa: USNM 408155 (1, 153). Manua’e (Scilly) I.: USNM 435224 [SCIL-335] (1, 114). Hawaii: BPBM 37447 (1, 285). Description. In TL: preanal length 2.2–2.5, predorsal length 6.6–11, head length 6.5–8.3, body depth at anus 15–24. In head length: snout length 4.1–6.8, eye diameter 6.8–11, upper-jaw length 2.2–3.2. Pores: LL 2, SO 3, IO 4, POM 6. Vertebrae: predorsal 5–6 (1 specimen with 10), preanal 42–44, total 118–123. Body moderately stout; anus before midlength; dorsal-fin origin before gill opening. Dorsal and anal fins high, dorsal fin height up to half body depth. Snout relatively short and tapering, jaws of equal length. Eye moderate, over middle of upper jaw. Anterior nostril long and tubular, reaching edge of lip when depressed; posterior nostril above anterior margin of eye. Intermaxillary teeth in a single peripheral series, 6–7 on each side, triangular, increasing in size posteriorly; 1–3 median teeth, long, conical. Maxillary teeth uniserial in larger specimens, biserial in smaller specimens, which have a few large inner teeth anteriorly. Dentary with 1–3 larger inner teeth anteriorly, an outer row of smaller teeth, decreasing in size posteriorly. Larger teeth anteriorly in jaws serrate. Vomerine teeth uniserial or slightly staggered, small and inconspicuous. Color: medium to dark brown, becoming nearly black posteriorly on tail and fins, with an indistinct marbled pattern of lighter brown separated by obscure darker interspaces on body and basally in the dorsal fin; often obscure horizontal lines on branchial area and anterior body. Anterior nostril dark brown. Iris yellow, margin of eye darker brown, wider posteriorly. Maximum size about 400 mm. Distribution and habitat. Throughout the Indo-Pacific from the Red Sea and east coast of Africa to the Society Islands and Hawaiian Islands. Usually found in shallow lagoon reefs and from fringing reefs in depth range 2– 43 m. Remarks. This species has been confused in the past with other plain brown morays. Schultz (1953: 113) misidentified it as Gymnothorax moluccensis and G. monochrous (see Randall & McCosker 1975: 17–18). Randall & Golani (1995: 865) reported vertebral counts of 130–135 for three specimens from Midway, but these specimens (presumably SIO 68-498 as reported by Randall & McCosker 1975: 17) are most probably Gymnothorax atolli (Pietschmann), a species that was not recognized until later. Böhlke & Randall (2000: 249) reported the range of vertebral counts as 110–124, but we have examined specimens from all corners of the Indo-Pacific and found no confirmed counts lower than 118. We suspect that the figure of 110 is either an error or based on a damaged specimen. There appears to be a considerable high level of intraspecific genetic variation. Two of the Red Sea specimens collected in this study (KAUMM 414 and SMF 34818) fall apart from the others on the COI phylogeny (Fig. 48). The majority of sequences derived from Red Sea specimens, however, fall into a subclade with specimens from the South Pacific (New Caledonia and Society Islands). The divergence between the two genetic subgroups is as prominent as that among sub-groups in G. javanicus (see above), but as in that case, we can find no morphological characters that separate them, and we cannot explain the significance of the observed genetic divergence. No closely related species can be identified from the present phylogeny for G. pindae (Fig. 48)., Published as part of Smith, David G., Bogorodsky, Sergey V., Mal, Ahmad O. & Alpermann, Tilman J., 2019, Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea, with description of a new species, pp. 1-87 in Zootaxa 4704 (1) on page 49, DOI: 10.11646/zootaxa.4704.1.1, http://zenodo.org/record/3563576, {"references":["Smith, J. L. B. (1962) The moray eels of the western Indian Ocean and the Red Sea. Ichthyological Bulletin, Department of Ichthyology, Rhodes University, 23, 421 - 444.","Randall, J. E. & Golani, D. (1995) Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea. Bulletin of Marine Science, 56 (3), 849 - 880.","Golani, D. & Bogorodsky, S. V. (2010) The fishes of the Red Sea-reappraisal and updated checklist. Zootaxa, 2463, 1 - 135. https: // doi. org / 10.11646 / zootaxa. 2463.1.1","Golani, D. & Fricke, R. (2018) Checklist of the Red Sea fishes with delineation of the Gulf of Suez, Gulf of Aqaba, endemism and Lessepsian migrants. Zootaxa, 4509 (1), 1 - 215.","Schultz, L. P., Herald, E. S., Lachner, E. A., Welander, A. D. & Woods, L. P. (1953) Fishes of the Marshall and Marianas islands. Vol. I. Families from Asymmetrontidae through Siganidae. Bulletin of the United States National Museum, 202, i-xxxii + 1 - 685. https: // doi. org / 10.5962 / bhl. part. 17831","Randall, J. E. & McCosker, J. E. (1975) The eels of Easter Island with a description of a new moray. Natural History Museum of Los Angeles County Contributions in Science, 264, 1 - 32.","Bohlke, E. B. & Randall, J. E. (2000) A review of the moray eels (Angulliformes [sic]: Muraenidae) of the Hawaiian Islands, with descriptions of two new species. Proceedings of the Academy of Natural Sciences of Philadelphia, 150, 203 - 278."]}

Gymnothorax undulatus (Lacepède 1803) —Undulated Moray (Figure 38) Muraenophis undulata Lacepède (ex Commerson) 1803: 629, 642, pl. 19 (fig. 2) (No locality). Holotype (unique), whereabouts unknown. ? Muraena undulata: Klunzinger 1871: 615 (Quseir, Egypt). ? Gymnothorax meleagris (non Shaw & Nodder): Fowler & Steinitz 1956: 270 (Eilat). Gymnothorax undulatus: Randall & Golani 1995: 868 (in part: Pl. 2F); Debelius 1998: 13; Lieske & Myers 2004: 36; Golani & Bogorodsky 2010: 10; Golani & Fricke 2018: 23. Red Sea material. Red Sea : USNM 47604 (1, 490). Egypt: USNM 312603 (1, 674), Gulf of Aqaba, Marsa Muqabila. Saudi Arabia: USNM 147430 (1, 500), Gubat Ashra. Eritrea: USNM 312608 (1, 357), Dahlak Archipelago, Delemmi. Comparative material. Arabian Gulf : BPBM 29469 (1, 693); BPBM 33383 (1, 760); BPBM 33384 (2, 432– 533). Mauritius: BPBM 20132 (1, 207). Chagos Archipelago: USNM 312615 (5, 263–407). Madagascar: MNHN B.2426 (1, 400); MNHN B.2432 (1, 317); MNHN B.2728 (1, 434); MNHN 1965-338 (1, 206); MNHN 1991-0402 (3, 49 8–595). Mozambique: SAIAB 60389 (1, 223). South Africa: SAIAB 60166 (9, 80–360); USNM 312712 (1, 570); USNM 330982 (1, 454). Wallis I. : USNM 371033 (1, 141). Description. In TL: preanal length 2.1–2.4, predorsal length 7.8–11, head length 6.6–8.2, body depth at anus 12–21. In head length: snout length 4.6–6.4, eye diameter 6.4–12, upper-jaw length 2.2–2.9. Pores: LL 2, SO 3, IO 4, POM 6. Vertebrae: predorsal 3–6, preanal 50–58, total 126–136. Body moderately elongate; anus before midlength; dorsal-fin origin before gill opening. Snout moderate, jaws of equal length. Eye moderate, over middle of upper jaw. Anterior nostril tubular; posterior nostril above anterior part of eye. Teeth conical, pointed, smooth. Intermaxillary teeth in a single peripheral series, 2–6 on each side; 2–4 median teeth, usually 3. Maxillary teeth uniserial or biserial, with 1–2 larger inner teeth anteriorly, about 9–18 smaller outer teeth. Dentary teeth biserial, with 2–4 larger inner teeth anteriorly, 16–20 smaller outer teeth. Vomerine teeth uniserial but biserial in 1 specimen, about 4–7. Color: variable over range. In Red Sea, small individuals with body and fins dark gray to black with narrow pale markings arranged in a reticular pattern. Larger individuals more grayish, pattern becoming more irregular and obscure. Top of head olive-green to yellow-green, densely spotted with small irregular dark brown spots posteriorly; snout and lower jaw yellow-green to light grey-brown without spots; corner of mouth with a white spot. Outside Red Sea most commonly dark brown to black, with narrow pale markings usually interconnected to form a reticulation; the pattern generally more obscure in larger specimens. One specimen (BPBM 33383, 760 mm, from Jana Island, Arabian Gulf), apparently of this species, dull whitish with small, irregular black spots. Maximum size about 1.5 m. Distribution and habitat. Widely distributed in the Indo-Pacific from the Red Sea and East Africa eastward to the islands off Central America. Mainly in shallow water, common on coral reefs, reported from depths of 1– 110 m. Remarks. As pointed out above, this species was confused with Gymnothorax pharaonis n. sp. described here. Gymnothorax undulatus is highly variable over its range, and further study may show that it represents a complex of species. Specimens examined from the Red Sea and Arabian Gulf have fewer vertebrae (126–129) than those from Mauritius and Chagos Archipelago (130–134) and South Africa (134–136). The specimen from Wallis Island has 131. The South African specimens differ further in coloration, showing a densely spotted, gravel-like pattern rather than reticulations (see Pl. 57E in Smith 1962). Randall & Golani (1995: 868) listed Klunzinger’s (1871) record of Muraena undulata and Fowler & Steinitz’s record of Gymnothorax meleagris in their synonymy of Gymnothorax undulatus, but as they confused G. undulatus with G. pharaonis, these may refer to the latter species instead. No Red Sea specimen was collected during this study and no sequences of the mitochondrial COI barcoding gene were available for other Red Sea specimens of G. undulatus. We therefore included COI sequences of each of two closely related clades for which COI sequences could be retrieved from various databases. One clade was largely restricted to the Indian Ocean and the other to the Pacific Ocean. However, as the type locality of the species is not documented, the whereabouts of the holotype is unknown, and we did (with the exception of one South Pacific specimen) examine only Red Sea and Indian Ocean material herein, we refrain from guessing which of the two clades might represent Gymnothorax undulatus and if the other clade represents another species. Hence, both clades are under the name G. undulatus in the presented phylogeny (Fig. 48), where they form part of a highly supported clade with G. punctatus., Published as part of Smith, David G., Bogorodsky, Sergey V., Mal, Ahmad O. & Alpermann, Tilman J., 2019, Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea, with description of a new species, pp. 1-87 in Zootaxa 4704 (1) on pages 58-60, DOI: 10.11646/zootaxa.4704.1.1, http://zenodo.org/record/3563576, {"references":["Lacepede, B. G. E. (1803) Histoire naturelle des poisons. Vo. 5. Chez Saugrain, Paris, 803 pp. [l'an VII de la Republique (1799, i. e. 1803) - an XII (1804)] https: // doi. org / 10.5962 / bhl. title. 6882","Klunzinger, C. B. (1871) Synopsis der Fische des Rothen Meeres. II. Theil. Verhandlungen der K. - K. zoologisch-botanischen Gesellschaft in Wien, 21, 441 - 688. https: // doi. org / 10.5962 / bhl. title. 1148","Fowler, H. W. & Steinitz, H. (1956) Fishes from Cyprus, Iran, Iraq, Israel and Oman. Bulletin of the Research Council of Israel, 5 B (3 - 4), 260 - 292.","Randall, J. E. & Golani, D. (1995) Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea. Bulletin of Marine Science, 56 (3), 849 - 880.","Debelius, H. (1998) Red Sea reef guide. IKAN-Unterwasserarchiv, Frankfurt, 321 pp.","Lieske, E. & Myers, R. F. (2004) Coral reef guide. Red Sea to Gulf of Aden, South Oman. Harper Collins Publishers, London, 384 pp.","Golani, D. & Bogorodsky, S. V. (2010) The fishes of the Red Sea-reappraisal and updated checklist. Zootaxa, 2463, 1 - 135. https: // doi. org / 10.11646 / zootaxa. 2463.1.1","Golani, D. & Fricke, R. (2018) Checklist of the Red Sea fishes with delineation of the Gulf of Suez, Gulf of Aqaba, endemism and Lessepsian migrants. Zootaxa, 4509 (1), 1 - 215.","Smith, J. L. B. (1962) The moray eels of the western Indian Ocean and the Red Sea. Ichthyological Bulletin, Department of Ichthyology, Rhodes University, 23, 421 - 444."]}

Gymnothorax pharaonis n. sp. —Pharaoh’s Moray (Figures 25–29) ? Muraena undulata: Klunzinger 1871: 615 (Quseir, Egypt). ? Gymnothorax meleagris (non Shaw & Nodder): Fowler & Steinitz 1956: 270 (Eilat). Gymnothorax undulatus (non Lacepède): Randall & Golani 1995 (in part, Pl. 1F, Fig. 6). Holotype. SMF 35814 [KAU13-614] (322), Red Sea, Saudi Arabia, Al Khoraybah, Yabua Island, isolated coral block on slope, 27°47’24.66” N, 35°07’48.00’’ E, 14–16 m, 23 Jun. 2013, S.V. Bogorodsky. Paratypes. Israel : BPBM 31848 (1, 475), Gulf of Aqaba, Eilat, North Beach, mooring, 7 m, 11 Nov. 1986, J.E. Randall. Egypt: BPBM 18265 (2, 270–284), S end of Sinai Peninsula, Sharm-el-Moya, reef, 15 m, 21 Sep 1974, J.E. Randall and A. Levy; BPBM 19805 (1, 331), Ras Muhammad, S tip of Sinai Peninsula, reef front in 4–6 m, 26 Oct. 1975, J.E. Randall et al.; BPBM 20825 (1, 277), Gulf of Aqaba, 7 km S of Nuweiba, A. Ben-Tuvia, 3 Aug. 1976; USNM 262775 (1, 305 mm TL), NW coast Gulf of Aqaba, Bay at El Himeira, 0–18 m, 16 July 1969, V.G. Springer et al.; USNM 312604 (6, 133–380), Gulf of Aqaba, Bay Between Marsa Mokrakh and El Himeira, NW Coast, 0–3 m, 15 July 1969, V.G. Springer et al.; USNM 312605 (4, 187–228), NW Gulf of Aqaba, Ras Burqa, 9–15 m, 21 July 1969, V.G. Springer et al.; USNM 312609 (3, 303–419), Strait of Jubal S end of Sinai Peninsula at Ras Muhammad, 0–9 m, 26 Sep 1969, V.G. Springer et al.; USNM 405385 (4, 233–260), NW coast of Gulf of Aqaba, reef near road at Marsa Muqabila, 0–2 m, 29 July 1969, V.G. Springer et al.; USNM 410185 (7, 123–227), same data as USNM 312604; USNM 410188 (4, 223–299), same data as USNM 262775; USNM 410628 (1, 310), Gulf of Aqaba, Dahab, Lighthouse, 18 m, 25 Nov. 2011, S.V. Bogorodsky. Sudan: BPBM 19733 (1, 156), 1 mile N of Port Sudan, reef flat in 0.5–1.0 m, 9 Oct 1975, J.E. Randall. Saudi Arabia: KAUMM 400 [KAU12-1059] (1, 225), Al Lith, 8 Mar. 2012, T.J. Alpermann & S.V. Bogorodsky; KAUMM 401 [KAU13-286] (1, 145), 50 km south of Al Wajh, fringing reef of seaward reef, 8–12 m, 13 Jun. 2013; KAUMM 402 [KAU13-615] (1, 224), same data as holotype; SMF 33617 (1, 49), Al Wajh, Rykhah Is, 10 Apr. 2011, S.V. Bogorodsky; SMF 33618 (1, 78), Al Lith, 30 Mar. 2011, T.J. Alpermann & S.V. Bogorodsky; SMF 35815 [KAU12-1028] (1, 182), Al Lith, 9 m, 7 Mar. 2012, T.J. Alpermann & S.V. Bogorodsky; SMF 35816 [KAU12-1060] (1, 208), Al Lith, 8 Mar. 2012, T.J. Alpermann & S.V. Bogorodsky. Non-type material (detailed counts and measurements not taken). KAUMM 403 [KAU14-818] (1, 125), Al Lith, 6–8 m, 16 Nov 2014, T.J. Alpermann & S.V. Bogorodsky; KAUMM 404 [KAU14-1011] (1, 246), Al Lith, 8–10 m, 19 Nov 2014, T.J. Alpermann & S.V. Bogorodsky; SMF 35817 [KAU14-928] (1, 213), Al Lith, 6–9 m, 18 Nov. 2014, T.J. Alpermann & S.V. Bogorodsky; USNM 312606 (4, 62–140), Egypt, NW Coast, Gulf of Aqaba, about 1 Mile North of Ras Burqa, 21 July 1969; USNM 312607 (12, 49–160), Egypt, just N of Ras Burqa, Gulf of Aqaba, NW Coast, 23 July 1969, V. G. Springer et al.; USNM 313223 (9, 64–96), Egypt, Gulf of Aqaba, Bay at El Himeira, 8 Sept. 1969, V. G. Springer et al.; USNM 410183 (28, 50–192), same data as USNM 262775; USNM 410184 (14, 56–265), same data as USNM 312604; USNM 410186 (17, 77–280), same data as USNM 312609; USNM 410187 (28, 75–225), same data as USNM 405385; USNM 410189 (1, 227), same data as USNM 262775 (cleared and stained). Diagnosis. Small to medium-size moray with slender head and jaws. Teeth sharp, slender, and smooth; intermaxillary teeth in one peripheral and one medial series; maxillary teeth in two rows, an outer row of 14–20 small teeth, and an inner row of 0–6 large, depressible teeth; dentary teeth in one row, with two large fixed teeth at anterior end, followed by a single row of 15–20 small teeth, and one large, depressible tooth just behind the large anterior teeth. Color brown with irregular dendritic pale markings, not interconnected or chain-like; oblique, conspicuous, parallel streaks present in dorsal fin (on tail). Total vertebrae 123–128. Description (data for the holotype first, for paratypes in parentheses). In TL: preanal length 2.3 (2.2–2.4), predorsal length 9.0 (7.7–10), head length 7.8 (7.0–8.4), body depth at gill opening 19 (15–28), depth at anus 23 (17–28). In head length: snout length 5.2 (4.9–7.1), eye diameter 8.7 (7.6–11), upper-jaw length 2.3 (2.3–3.1). Predorsal vertebrae 8 (5–8), preanal vertebrae 48 (47–50), total vertebrae 125 (123–128). A small to medium-sized moray eel, moderately elongate, with the anus slightly anterior to midlength. Dorsal and anal fins continuous with caudal fin, anal fin beginning immediately behind anus, dorsal fin beginning anterior to gill opening. Jaws and snout moderately slender, edges usually straight, concealing teeth when closed, but sometimes slightly arched in larger specimens; upper and lower jaws nearly equal in length. Gill opening small and pore-like, on side of head slightly below lateral midline. Anterior nostril tubular, relatively long, reaching slightly beyond edge of lip when depressed. Posterior nostril a broadly oval opening, without a conspicuous raised rim, above anterior part of eye, at a point where a horizontal line drawn from dorsal edge of eye would meet a vertical line drawn from anterior edge of eye. Lateral line with two small, inconspicuous pores at anterior end of canal, approximately under dorsal-fin origin; second pore closer to first pore than to gill opening (Fig. 25). Preoperculo-mandibular canal with six pores, all of them along lower jaw: the first and smallest located at the anterior tip of jaw, the second below and behind that, the remaining four pores extending in a line posteriorly to a point slightly anterior to rictus. Infraorbital canal with four pores: the first slightly below and behind base of anterior nostril, the second about a third of the way to eye, the third just anterior to eye, and the fourth under posterior margin of eye. Supraorbital series with three pores: the first and smallest at tip of snout just above edge of lip, the second slightly above anterior edge of base of anterior nostril, the third on top of snout directly above second infraorbital pore. No pores in supratemporal canal. Teeth slender, sharp, and smooth, without any serrations (Fig. 26). Intermaxillary teeth large, conical, sharply pointed; peripheral series with about 8–14 teeth, the anteriormost teeth smallest, increasing in size posteriorly; two or three median teeth, long, extremely sharp and depressible. Maxillary teeth in one or two rows: the inner row with 0–6 long, sharp, widely separated, depressible teeth at anterior end, the outer row with about 13–20 much smaller, fixed, triangular, recurved teeth, smallest at anterior end of row, increasing in size posteriorly to a point approximately under eye, then decreasing in size again posteriorly. Lower jaw with two large, fixed teeth at anterior end, followed by approximately 15–25 much smaller, triangular, recurved teeth; directly behind the two large anterior teeth is an even larger, depressible tooth just inside the row of smaller teeth. Approximately 3–9 very small vomerine teeth, in a single row, partly hidden in the folds of skin in roof of mouth. Color: in adults, ground color medium to dark brown with irregular, dendritic, pale markings, variable in size and form (Figs. 27 & 28). The most common form is short, broadly linear, vermicular lines or spots, sometimes expanded into snowflake-like blotches, but not interconnected or reticulated. On tail, the spots often line up to form oblique streaks extending onto dorsal fin. Markings sometimes become smaller and more closely spaced anteriorly. Fins with a narrow white edge, but this often not conspicuous. Grooves on throat as dark streaks. An inconspicuous pale stripe usually present on dorsal midline of snout. Corner of mouth dark. Posterior nostril and pores usually edged in dark brown. Juveniles uniform brown with lower jaw and throat pale (Fig. 29). Size and development. This is a relatively small species, the largest specimen examined was 475 mm TL, but only one other specimen was greater than 400 mm and only three over 300 mm. Females with large eggs were found in specimens as small as 223 mm. Males appear to mature at larger sizes than females; two that were clearly males were among the largest specimens examined, 299 and 419 mm. Females were measured at 380, 325, and 305 mm. There is some evidence of sexual dimorphism in dentition. The two males mentioned above lack the inner maxillary teeth; they also have fewer dentary teeth (14 vs. 18–26 in females). Small juveniles of this species are uniform brown with a conspicuous white lower jaw (Fig. 29A). At about 50 mm, pale spots begin to develop behind the head. As the eel grows, the spots progress posteriorly and become larger and more conspicuous, eventually assuming the dendritic pattern characteristic of adults (Fig. 29B). With growth, the pale lower jaw becomes less distinct. Variation. The specimens collected and examined were all brown with pale markings. The relative extent of pale and dark areas varies considerably among individuals, however. In most cases, the dark areas are more extensive, giving the fish a brown appearance, but occasionally the pale areas prevail. In such cases, the eel may appear pale with brown markings. In most specimens, the pale and dark markings are relatively large, but in others the markings are smaller and more scattered, giving a vermiculated appearance. In some specimens, the markings are larger posteriorly and smaller anteriorly. In larger specimens, the jaws can become arched, leaving the teeth visible when the mouth is closed. This approaches the condition seen in Enchelycore, but the dentition of Enchelycore is quite different (Smith et al. 2008: 68). Distribution and habitat. Known from the Red Sea, where it is common in shallow water, but also collected by the second author from Socotra Island outside the Gulf of Aden (Zajonz et al. 2019, listed as G. cf chilospilus Bleeker 1864). Typical habitats are crevices and shelters of fringing seaward reefs, observed from depths of 2– 30 m. May be seen out of shelter at night only. Etymology. Named for the pharaohs, the rulers of ancient Egypt, whose realm included the Red Sea. Referring also to the regal appearance of this handsomely marked fish. Remarks. This species has been confused with Gymnothorax undulatus. Like G. pharaonis, G. undulatus has pale markings on a dark background, but in G. undulatus the markings are generally interconnected in a reticulated or chain-like pattern, whereas in G. pharaonis the markings are separate. At larger sizes, G. undulatus has a distinct yellowish-green color on the head in life, which is never found in G. pharaonis. Gymnothorax undulatus is a much larger species, growing to well over 1 m in length. Mature G. pharaonis can be found at lengths less than 300 mm, a size at which G. undulatus is still immature. The two species also differ in the number of vertebrae, 122–128 in G. pharaonis vs. 126–138 in G. undulatus. Gymnothorax pharaonis also resembles G. baranesi, but in that species the pale markings on the body are more like snowflakes or rosettes. On the tail, the markings on G. baranesi are in the form of discrete spots rather than the oblique streaks found in G. pharaonis. In addition, G. baranesi has more vertebrae, 137–142. Gymnothorax pharaonis most closely resembles and is closest genetically (Fig. 48) to G. margaritophorus Bleeker, which is widely distributed in the Indo-Pacific but does not occur in the Red Sea. The latter is also a small species, brown with pale markings and a pale stripe on the top of the snout. It has horizontal dark streaks behind the eye, however, which are lacking in G. pharaonis, and it has more vertebrae (127–134). Pale individuals (e. g. from Dahab, Fig. 28B) may be confused with G. chilospilus Bleeker, but the latter species almost always has a distinctive pale spot at the corner of the lower jaw, which is lacking or not obvious in G. pharaonis. As in G. griseus and G. thyrsoideus, no reciprocal monophyly has yet evolved in the species pair G. pharaonis and G. margaritophorus. The closest relative to this pair of sibling species cannot be identified with high confidence from the present phylogeny, however, it is evident that the two species form part of a highly supported group of taxa to which another Red Sea species belongs, G. johnsoni (Fig. 48)., Published as part of Smith, David G., Bogorodsky, Sergey V., Mal, Ahmad O. & Alpermann, Tilman J., 2019, Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea, with description of a new species, pp. 1-87 in Zootaxa 4704 (1) on pages 41-45, DOI: 10.11646/zootaxa.4704.1.1, http://zenodo.org/record/3563576, {"references":["Klunzinger, C. B. (1871) Synopsis der Fische des Rothen Meeres. II. Theil. Verhandlungen der K. - K. zoologisch-botanischen Gesellschaft in Wien, 21, 441 - 688. https: // doi. org / 10.5962 / bhl. title. 1148","Fowler, H. W. & Steinitz, H. (1956) Fishes from Cyprus, Iran, Iraq, Israel and Oman. Bulletin of the Research Council of Israel, 5 B (3 - 4), 260 - 292.","Randall, J. E. & Golani, D. (1995) Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea. Bulletin of Marine Science, 56 (3), 849 - 880.","Smith, D. G., Brokovich, E. & Einbinder, S. (2008) Gymnothorax baranesi, a new moray eel (Anguilliformes: Muraenidae) from the Red Sea. Zootaxa, 1678, 63 - 68. https: // doi. org / 10.11646 / zootaxa. 1678.1.4","Zajonz, U., Lavergne, E., Bogorodsky, S., Saeed, F. N., Aideed, M. S. & Krupp, F. (2019) Coastal fish diversity of the Socotra Archipelago, Yemen. Zootaxa, 4636 (1), 1 - 108.","Bleeker, P. (1864) Poissons inedits indo-archipelagiques de l'ordre des Murenes. Nederlandsch Tijdschrift voor de Dierkunde, 2, 38 - 54."]}

Gymnothorax pseudoherrei Böhlke 2000 —Dwarf Brown Moray (Figure 33) Gymnothorax pseudoherrei Böhlke 2000: 408, figs. 2F, 3D, 7 (W side of Solino [Selinog] I., Zamboanga Del Norte, Mindanao, Philippines, 8°51’24”N, 123°24’36”E, 0–4.6 m). Holotype, USNM 357430. — Golani & Bogorodsky 2010: 10; Golani & Fricke 2018: 22. Gymnothorax herrei (non Beebe & Tee Van): Randall & Golani 1995: 860. Red Sea material. Saudi Arabia : KAUMM 409 [KAU 12-1082] (1, 112), Al Lith; KAUMM 410 [KAU 14-869] (1, 116), Al Lith; SMF 33616 (1, 107), Al Lith; SMF 35822 [KAU 12-1083] (1, 171), Al Lith; SMF 35877 [KAU 17- 245] (1, 127), Farasan Archipelago, Abkar Island. Eritrea: HUJ 15113 (2, 185-193), Dahlak Archipelago, Romia Island; USNM 312234 (8, 116–204), Sheikh el Abu; USNM 312247 (1, 150), Melita Bay. Yemen: USNM 397542 (1, 171), Hanish Island. Comparative material. Arabian Gulf : BPBM 33328 (1, 291); BPBM 33356 (3, 208–256). Gulf of Oman: BPBM 21473 (1, 208). Sri Lanka: USNM 357433 (3, 116–155, paratypes). Philippines: USNM 357430 (1, 147, holotype), Mindanao; USNM 357432 (2, 114–121, paratypes), Palawan. Indonesia: USNM 210269 (1, 148); USNM 274957 (1, 103). Papua New Guinea: USNM 357431 (2, 126–156, paratypes). Description. In TL: preanal length 2.0–2.4, predorsal length 8.1–12, head length 7.2–9.5, body depth at anus 16–25. In head length: snout length 5.7–6.9, eye diameter 7.9–11, upper-jaw length 2.7–3.4. Pores: LL 2, SO 3, IO 4, POM 6. Vertebrae: predorsal 5–8, preanal 42–50, total 111–120. Body somewhat elongate in smaller specimens, becoming moderately stout with growth; anus slightly before midlength; dorsal-fin origin before gill opening. Snout relatively short and tapering, jaws of equal length. Eye moderate, over middle of upper jaw. Anterior nostril tubular; posterior nostril above anterior part of eye. Teeth smooth, relatively short, stout and pointed. Intermaxillary teeth in a single peripheral series, 6–8 on each side, conical, increasing in size posteriorly; 1–2 median stout teeth. Maxillary teeth biserial, about 7–14 larger inner teeth and 19–21 smaller outer teeth, those in outer row obtusely pointed. Dentary with 1–4 larger inner conical teeth anteriorly and about 17–25 smaller blade-like outer teeth. Vomerine teeth stout, blunt, uniserial or staggered, about 7–13. Color: medium to dark brown with an irregular network of indistinct small, darker markings, head paler than body, lower part of head lighter, sometimes with dark lines along throat grooves, one continuing into angle of mouth. Posterior one-fourth of body and fins yellow in smaller specimens, yellow color gradually disappears with growth. Iris white with black outer ring. Body frequently covered with a gray or greenish mucus. Maximum size about 300 mm. Distribution and habitat. Northwestern Indian Ocean and western Pacific, from the Red Sea and Arabian Gulf to the Solomon Islands, in shallow water, generally less than 10 m depth. It has not been collected from the coast of Africa or the islands of the western Indian Ocean west and south of the Maldives. A cryptic species living inside coastal reefs, never seen alive. Remarks. Specimens from the northwestern Indian Ocean (Red Sea, Arabian Gulf, and Gulf of Oman) apparently grow larger than those from elsewhere. Out of more than 100 specimens reported by Böhlke (2000) from east of the Maldives, the largest was 182 mm (ANSP 144601 from Queensland, Australia). The largest of only 12 specimens reported from the northwestern Indian Ocean, by contrast, was 291 mm (BPBM 33328, from the Arabian Gulf). Several other specimens from this area exceeded 200 mm. There is a slight difference in the number of vertebrae between these two groups. Nineteen specimens from the Red Sea, Arabian Gulf, and Gulf of Oman had 113–120 total vertebrae; nine specimens from Sri Lanka, Indonesia, the Philippines, and Papua New Guinea had 111–116. The Red Sea and Arabian Gulf specimens also have dark throat grooves, which are not evident in those from the Pacific. We have no genetic data from outside the Red Sea. This species was confused in the past with the superficially similar Gymnothorax herrei Beebe & Tee-Van, from which it differs by having two branchial pores instead of one, the origin of the dorsal fin more anterior, and lacking an intermediate row of small intermaxillary teeth. In the present COI phylogeny (Fig. 48) two specimens are included that have been collected during the course of this study. As no COI sequence data from other specimens are available, we cannot infer the level of intraspecific genetic divergence. Gymnothorax pseudoherrei forms part of a well supported clade with a number of other taxa, such as G. griseus and G. thyrsoideus, but also Echidna unicolor and E. delicatula (Fig. 48)., Published as part of Smith, David G., Bogorodsky, Sergey V., Mal, Ahmad O. & Alpermann, Tilman J., 2019, Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea, with description of a new species, pp. 1-87 in Zootaxa 4704 (1) on page 51, DOI: 10.11646/zootaxa.4704.1.1, http://zenodo.org/record/3563576, {"references":["Golani, D. & Bogorodsky, S. V. (2010) The fishes of the Red Sea-reappraisal and updated checklist. Zootaxa, 2463, 1 - 135. https: // doi. org / 10.11646 / zootaxa. 2463.1.1","Golani, D. & Fricke, R. (2018) Checklist of the Red Sea fishes with delineation of the Gulf of Suez, Gulf of Aqaba, endemism and Lessepsian migrants. Zootaxa, 4509 (1), 1 - 215.","Randall, J. E. & Golani, D. (1995) Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea. Bulletin of Marine Science, 56 (3), 849 - 880."]}

Thirty-eight species of moray eels (Muraenidae) from the Red Sea are reported in an updated review. A species account is provided for each species, along with a full synonymy of all nominal species described from the Red Sea. One species is new to science, G. pharaonis, and two species are new for the Red Sea, Gymnothorax phasmatodes (Smith) and Scuticaria tigrina (Lesson). Gymnothorax pharaonis n. sp. is a common species that has long been misidentified as G. undulatus (Lacepède), although it more closely resembles G. margaritophorus Bleeker, to which it is closely related phylogenetically. It is characterized by the following combination of characters: total vertebrae 123–128; body slender, depth at anus 17–28 in TL; maxillary teeth biserial, dentary and vomerine teeth in a single row; color brown with irregular dendritic pale markings, not interconnected or chain-like, with oblique, conspicuous, parallel streaks posteriorly in dorsal fin. The new record of the distinctive whitish G. phasmatodes is based on an underwater photograph taken at Jeddah, Saudi Arabia. The new record of Scuticaria tigrina is based on a specimen collected from Jeddah, Saudi Arabia, and underwater photographs taken from the northern part of the Red Sea. Based on an integrative taxonomical approach by a combination of morphological and phylogenetic analyses and re-examination of holotype specimens, Gymnothorax cinerascens (Rüppell) is resurrected as a valid species, and it is shown that previous records of G. hepaticus (Rüppell), with which it has previously been synonymized, should refer to G. cinerascens. The true G. hepaticus is redescribed based on examination of the holotype and additional specimens collected during the course of this study. The two species are distinguished by the color of the jaws, the position of the median intermaxillary teeth, and a slight difference in the preanal length. They are also clearly distinct genetically. The Red Sea record of Gymnothorax atolli (Pietschmann) is based on an error, possibly a misidentification of G. griseus (Lacepède). A key to the species of Red Sea moray eels and a phylogenetic tree of presently known lineages of Indo-Pacific moray eels are provided including recently collected Red Sea specimens.

Gymnothorax andamanensis sp. nov. Proposed common name: Andaman brown moray (Figures 1–4 A, Table 1) Holotype. EBRC /ZSI F11227 [325 mm total length (TL)], Port Mout, Port Blair, South Andaman (11.659327°N; 92.696148°E]: 18 March 2019. Paratypes. CIARI /MF-05. [464 mm TL), date and place of collection: same as holotype. Diagnosis. This new species of moray eel belongs to the small, brown unpatterned group with the combination following of characters: dorsal-fin origin before gill opening, pre-anal length 2.1–2.2, jaw pores with black rim, two branchial pores, predorsal vertebrae 3, preanal vertebrae 57 and total vertebrae 135–136, teeth smooth, three large fang-like median intermaxillary teeth, biserial maxillary and uniserial vomerine teeth, and dentary teeth biserial, with two teeth on each side in the second row of the dentary. Description. A medium-sized, uniform brown moray with moderately elongate body, depth at gill opening 15.0–17.1 and at anus 16.0– 19.1 in TL; preanal length 2.1–2.2 in TL. Head small, 8.1 in TL; snout blunt, its length 4.4 in HL; jaws almost equal 2.2–2.4 in HL. Eye diameter of 6.7–7.6 in HL and located closure to rictus than the snout tip; interorbital space 6.7–8.8 in HL (Table 1). Anterior nostril tubular touching the tip of the snout, posterior nostril above the anterior margin of eye. Dorsal-fin origin before gill opening and closer to rictus than to gill opening. Pre-dorsal length 9.9–10.8 in total length. The dorsal fin origin is well before the 1st branchial pore. Anal fin starts from the anus. Both dorsal and anal fins high. Gill in a small diagonal slit located behind the dorsal fin origin and second branchial pore (Fig. 2). Teeth smooth, not serrated; intermaxillary teeth 11–12 on each side with alternate small and large teeth as shown in Fig. 3; 3 strong, curved median intermaxillary teeth placed almost equidistance from each other; maxillary teeth biserial, 19–20 on each side of outer row and 3 in each side of inner row; vomerine teeth uniserial with 9–10 teeth; lower jaw teeth with outer row of 26–28 teeth on each side and 2 inner teeth on each side near anterior end (Fig. 3). Head pores typical, supraorbital pores 3; infraorbital pores 4; mandibular pores 6; branchial pores 2, located above and before gill opening (Fig. 2). All head pores with black rim. Predorsal vertebrae 3, preanal vertebrae 57 and total vertebrae 135–136. Colour when fresh: body dark brown dorsally and slightly pale brown ventrally; (Figs. 1 & 4), both dorsal and anal fins body colour; eye ring whitish, jaw pores with black rim. When preserved, body uniformly brown, eye ring white. Distribution. Presently known only from the South Andaman, India. Etymology. The species is named “ andamanensis ” with reference to the Island from where it is collected.

Gymnothorax andamanensis sp. nov., a new short brown moray eel, is described here on the basis of two specimens collected from Port Mout, Port Blair, South Andaman (11.659327°N; 92.696148°E), caught using baited hand-lines in a sandy habitat at a depth of less than 2 m. The species is characterized in having the dorsal-fin origin before the gill opening, pre-anal length 2.1–2.2, jaw pores with black rim, two branchial pores, predorsal vertebrae 3, preanal vertebrae 57 and total vertebrae 135–136, teeth smooth, three large fang-like median intermaxillary teeth, biserial maxillary and uniserial vomerine teeth, and dentary teeth biserial with two teeth in each side in the second row of the dentary. The new species is compared with all 10 short brown unpatterned moray eels known from the world and two from Indian waters.

Gymnothorax smithi sp. nov. Proposed common name: Indian white spotted moray eel (Figs. 1–4, Table 1) Holotype. CMLRE IO /SS/FIS/00452 (342 mm TL), 200 m depth off Kochi, India (09°59.935’N, 75°36.086’E) collected on-board FORV Sagar Sampada during August 2011, by HSDT (High Speed Demersal Trawl). Paratypes. EBRC /ZSI/ F 11228 (362 mm TL); ZSI/ANRC-22842 (615 mm TL), collected from Neendakara fish landing centre, Kollam, Kerala, during November, 2018. Diagnosis. A moderate sized moray eel of the genus Gymnothorax with the following combination of characters: greyish brown colour overlain with small irregular white spots; anus slightly behind mid-point of body; head broad with short jaws; snout tip blunt and short; eyes small and closer to rictus than to snout tip; pointed and serrated teeth; vomerine teeth uniserial anteriorly, biserial centrally and uniserial posteriorly; 1–2 branchial pores; vertebral count 3–5 /57/130–132. Description. Morphometric data of holotype (in mm): total length 342; pre-anal length 182; pre-dorsal length 34; head length 57; snout length 9.3; eye diameter 4.5; interorbital 7.3; interbranchial 13; gill opening 5.7; depth at gill 27.6; depth at dorsal-fin origin 32; depth at anus 15; upper jaw length 19.8, and lower jaw length 19.3. A moderate sized eel with cylindrical to laterally compressed head and laterally compressed tail (Fig. 1). Dorsal and anal fins continuous with caudal fin. Dorsal fin commences before gill opening and its distance to snout tip is 8.4–10 times in TL. Anal fin-origin immediately behind anus. Anus located slightly behind mid-body, pre-anal distance is 1.83–1.88 in TL. Head broad and gently sloping towards snout, 5.5–6.0 in TL. Snout blunt and short, 6.1–6.9 in HL. Upper jaw and lower jaw are almost equal. All teeth concealed when mouth closed. Tongue well attached the floor of mouth. Eyes small, covered by membrane and closer to snout tip than rictus, eye diameter 12.7–13.1 in HL. Anterior nostril tubular at lateral side of tip of snout and posterior nostril above anterior margin of eye. Gill opening a diagonal slit at mid-body level. Head pores (Fig. 2) small but discernable. Two branchial pores located below dorsal-fin origin well before gill opening and closer to dorsal-fin origin than gill opening in holotype (both paratypes have a single branchial pore); three supra-orbital pores, first (ethmoid) pore at tip of snout above lip, second above base of anterior nostril, third between anterior and posterior nostrils; four infraorbital pores, first pore just behind base of anterior nostril, second between anterior nostril and anterior margin of eye, third below anterior margin of eye, and fourth after posterior margin of eye; six mandibular pores, first five before rictus and fifth below rictus. Teeth (Fig. 3) in jaws serrated and uniserial except in vomer and anterior dentary. Maxillary and mandibular teeth conical and sharp, vomerine teeth blunt and smaller. Intermaxillary teeth 6 on both sides, followed by 9 maxillary teeth on both sides, median intermaxillary teeth 2, inner maxillary teeth 1 on each side, just before vomerine teeth. Vomer with 4 uniserial teeth anteriorly, followed by 4 biserial teeth and 3 uniserial teeth posteriorly. Mandibular teeth 17 on each side, anterior 4 teeth a little enlarged with 4 smaller outer teeth on each side, remaining teeth decreasing in size posteriorly. Colour in formalin. Body greyish brown overlain with white spots of irregular shape. Snout, lower jaw and ventral portion of body pale. The white spots comparatively smaller in head region than body (Fig. 4). In enlarged view under microscope, there are small brown pigmentation spots inside the white spots. Margin of gill opening with similar brown pigmentation. Spots very small and more closely spaced in head region and becoming large and more widely spaced posteriorly, tail end is almost devoid of spots with dusky greyish brown colour. Dorsal fin greyish brown overlain with white spots similar to those on body. Anal fin dusky greyish brown with few isolated spots. Caudal fin ends with small white margin, visible only on close inspection. Mouth white in colour overlain with grey patches. Distribution. Presently known only from three type specimens collected by demersal trawl off Kerala coast, southeastern Arabian Sea, India at about 200 m depth (on the basis of holotype, not known for paratypes). Etymology. We are pleased to name this species as Gymnothorax smithi in honor of David G. Smith, United States National Museum, an eminent eel expert who has contributed significantly to the knowledge and understanding of Anguilliform fishes.

A new species of white-spotted moray eel Gymnothorax smithi sp. nov. is described based on three specimens collected from the southeastern Arabian Sea, India. This is the first deep water report of a moray eel from Indian waters from 200 m depth. This new species is distinguishable from its congeners by the following combination of characters: greyish brown body overlain with white spots of irregular shape, dorsal-fin origin anterior to gill opening, anus positioned slightly behind the mid-point of the body, serrated teeth and a unique vertebral count 3–5 /57/130–132. [Zoobank URL: urn:lsid:zoobank.org:act:9554CE07-E9E7-4B4F-95CD-54F2BB26FF28]

Gymnothorax undulatus (Lacepède 1803) — Undulated moray Status at New Ireland. First recorded from Drei Inseln Harbour, New Hanover by Munro (1958: 127). Distribution and habitat. New Ireland: 1.—General distribution: Red Sea, East and South Africa east to Panama, north to southern Japan and Ogasawara Islands, south to northern Australia. Found in crevices in reef flats, lagoons and seaward reef slopes, 0–50 m depth. Marine., Published as part of Andréfouët, Serge, Chen, Wei-Jen, Kinch, Jeff, Mana, Ralph, Russell, Barry C., Tully, Dean & White, William T., 2019, Checklist of the marine and estuarine fishes of New Ireland Province, Papua New Guinea, western Pacific Ocean, with 810 new records, pp. 1-360 in Zootaxa 4588 (1) on page 52, DOI: 10.11646/zootaxa.4588.1.1, http://zenodo.org/record/2988163, {"references":["Munro, I. S. R. (1958) The fishes of the New Guinea region. Fishery Bulletin, Territory of Papua and New Guinea, Port Moresby, 1, 97 - 369."]}

Gymnothorax mucifer Snyder, 1904 has been synonymized with G. kidako (Temminck & Schlegel, 1846) since 2000. Examination of material from Hawaii (type locality) and Taiwan reveal that G. mucifer is a valid species based on morphological and molecular data. It can be distinguished from G. kidako by a plainly colored lower jaw (vs. mottled), white margin of the anal fin replaced by serial pale blotches posteriorly on the tail (vs. white margin continuous to tip of tail), more dentary teeth (17–27 vs. 16–20), and fewer total vertebrae (130–141 vs. 140–143). In the present study, a redescription of G. mucifer is provided, and the distribution of G. mucifer and G. kidako is reevaluated.

Gymnothorax mucifer Snyder, 1904 (Figs. 1���5, Table 2) Diagnosis. A moderately sized moray (to at least 715 mm TL) with yellowish white spots on dark brown body and fins. Spots small and scattered evenly on top of head, grading into snowflake-like blotches on trunk and anterior part of tail, centralizing as separate rounded blotches on posterior part of tail. White margin on anal fin, replaced by serial pale blotches on posterior part of tail. Peripheral intermaxillary and median intermaxillary teeth uniserial. Maxillary teeth uniserial in larger males, biserial in immatures and females. Vomerine teeth small and stout in a staggered row, sometimes biserial centrally. Dentary teeth in one row, an additional peripheral row with small teeth present in immatures and females. Predorsal vertebrae 4���6, preanal vertebrae 51���55 and total vertebrae 130���141. Description. Measurements from Taiwanese and Hawaiian specimens. A moderately stout moray, depth at gill opening 13.2���27.0 and at anus 16.9���28.6 in TL. Anus anterior to mid-point of body, preanal length 2.1���2.4 in TL. Dorsal fin moderately high, the origin anterior to gill opening, predorsal length 9.5���11.9 in TL and 1.1���1.5 in HL. Anal fin shallow, originating just behind anus. Gill opening slightly below mid-body. Head length 7.3���8.7 in TL. Eyes above middle of jaw with diameter 8.2���12.2 in HL. Jaws subequal, 1.9���2.7 in HL, teeth not visible when mouth closed. Snout moderately elongate, 4.4���5.9 in HL. Anterior nostril at snout tip, moderately elongate and tubular, shorter than eye diameter in length; posterior nostril above the anterior margin of eye as a pore with raised rim. Head pores typical and with dark rim; three supraorbital pores, first on tip of snout, second just above anterior nostril, and third above and between first and second infraorbital pores; four infraorbital pores, first behind anterior nostril, second midway between anterior nostril and third infraorbital pore, third below anterior margin of eye, and fourth pore below posterior margin of eye; six mandibular pores along lower jaw, all before rictus; two branchial pores on postero-dorsal head before gill opening. Predorsal vertebrae 4���6 (5), preanal vertebrae 51���55 (53), total vertebrae 130���141 (137). For a and b, only specimens with intact tail were measured. a 11 specimens; b 13, 7 and 8 specimens for Taiwanese G. mucifer, G. kidako and G. niphostigmus, respectively; c count on 5 specimens, not including TOU-AE4828, TOU-AE4829, TOU-AE4830; d count on 6 specimens, not including TOU-AE0238, TOU-AE4981, TOU-AE4982, TOU-AE5582, TOU-AE5606. Dentition (Fig. 4): teeth smooth, pointed and slightly retrorse. Peripheral intermaxillary teeth uniserial, with 5��� 7 canines on each side. Median intermaxillary teeth very long, slender and uniserial, 2���3 in number, larger posteriorly. Maxillary teeth uniserial, 10���18 on each side, shorter than peripheral intermaxillary teeth, triangular, smaller on anterior and posterior sides, largest centrally; an additional inner row with 1���4 very long and pointed teeth on each side in females and immatures. Vomerine teeth small and stout, 5���20 in a staggered row, sometimes biserial centrally. Dentary teeth 17���27 on each side, anterior 4���5 pairs large, strong and retrorse, the remaining teeth smaller and subequal in size; an additional peripheral row with 3���6 pairs of small teeth present at 3 rd ���5 th anterior dentary teeth in females and immatures. Several small teeth in spaces between larger teeth on peripheral intermaxillary and anterior dentary. Color of fresh specimens (Figs. 1,3,5): dark brown in background and covered with moderate-sized yellowish white spots on top of head, body and fins. Spot arrangements variable among individuals. In general, spots aggregate to form numerous snowflake blotches on trunk and anterior part of tail, but centralize to form separate rounded blotches on posterior part of tail. Spots small and scattered evenly on top of head, scarcely perceptible on snout and lower jaw. Lower jaw, throat and ventral body lighter in color. Anal fin with prominent white margin and a dark brown submarginal band. White margin ends before tip of tail and replaced by several continuously pale blotches. Gular folds, gill opening and corner of mouth slightly dark. Iris of eye yellowish in fresh specimens. Phylogenetic analysis and comparisons. The topology of ML and Bayesian phylogenetic analyses based on the 2,438 bp of concatenated genetic markers revealed that material from Taiwan are three well recognized species. Gymnothorax mucifer is sister to G. niphostigmus while G. kidako is a sister taxon of the other two species (Fig. 6). The inter- and intra-species K2P genetic distances based on 647 bp COI fragment are 5.7���7.8% and 0.2���0.8%, respectively (Table 3). The high inter-species genetic distances which far exceed the intraspecific variation of moray eels (1.0���1.6% in average and maximum to 3.7%; Reece et al., 2011; Huang et al., 2018b) also support the recognition of three valid species. In each clade, color patterns of lower jaw and posterior tail are consistent and therefore used as diagnostic characters. The holotype (USNM 50868) of G. mucifer is well preserved and its color pattern is in good condition as originally described (Fig. 2). It has dark brown background and pale spots scattered evenly on head, grading into snowflake-like patches on trunk, and more rounded and farther apart on tail. The white margin on the anal fin is interrupted by serial pale blotches on posterior part of tail although the damaged tail may bring uncertainty (Dr. D.G. Smith, pers. comm.). Pale spots are scarcely perceptible on lower jaw and snout. Dentition and vertebral formula were not available in the original description, but recently provided by B��hlke & Smith (2002). Teeth uniserial, six in each side of peripheral intermaxillary, three in median intermaxillary, 15���16 in maxillary, 10 in vomer and 23���24 in dentary. The vertebral formula is 5���53���123+. Although there is no sequence available from Hawaii, the type locality of G. mucifer, all specimens from Hawaii identified as G. ��� kidako ��� and those from Taiwan fit the diagnosis based on the holotype of G. mucifer and can be well distinguished from G. kidako. Gymnothorax mucifer differs from G. kidako by having a plain lower jaw (vs. mottled), white margin of the anal fin with serially pale blotches on the posterior part of tail (vs. continuous to the tip of tail), more dentary teeth (17���27 vs. 16���20), and fewer total vertebrae (130���141 vs. 140���143) (Table 2, Fig. 5). Color patterns on lower jaw and posterior part of tail are reliable characters to separate G. mucifer from G. kidako even though blotches on body are quite variable in both species (Figs. 1, 5). The total vertebrae of G. mucifer and G. kidako are discrete in Taiwanese specimens (130���138 vs. 140���143) but slightly overlapped after including the Hawaiian specimens (Table 2). The Hawaiian G. mucifer tend to have slightly more vertebrae than the Taiwanese material. B��hlke & Randall (2000) noted that some Hawaiian muraenids had more vertebrae than usual depending on habitat difference or genetic isolation. Similar to the vertebral formula, the number of dentary teeth is discrete and can be used to separate the Taiwanese specimens into two species (21���27 in G. mucifer vs. 16���20 in G. kidako) (Table 2). Gymnothorax niphostigmus has similar color pattern of dark brown background with pale snowflake blotches on body, plain lower jaw and white margin on the anal fin. However, G. mucifer can be easily separated from G. niphostigmus by more numerous and larger blotches on tail (vs. blotches small, fewer and farther apart), white margin of anal fin discontinuous (vs. continuous and ends before tail tip), and fewer total vertebrae (130���141 vs. 140���146) (Table 2, Fig. 5). Gymnothorax nasuta De Buen, 1961 also has similar pale spots, white anal fin margin and overlapping vertebral formula, but is characterized by exceptionally long anterior nostrils (6���7% HL), yellowish posterior nostrils when alive, small pale spots on lower jaw and continuously white anal fin margin (De Buen, 1961; Randall and McCosker, 1975). Many Indo-Pacific congeners with similar pale spots and overlapping vertebral formula can be easily separated by lacking of the white anal fin margin, for instance, G. baranesi Smith, Brokovich & Einbinder, 2008, G. johnsoni (Smith, 1962), G. prionodon Ogilby, 1895 and G. punctatus Bloch & Schneider, 1801. Based on evidence of color pattern, morphometrics and molecular analysis, G. mucifer is clearly a valid species well separated from G. kidako and other congeners. Among our examined material, a specimen identified as G. ��� kidako ��� from Society Island (BPBM 31625, 900 mm TL) was not able to be identified to either species of Gymnothorax. This specimen is similar to G. kidako in having pale spots on body and continuously white margin of anal fin extended to tip of tail, but its dentition and plain lower jaw and snout are similar to G. mucifer. Furthermore, this specimen differs from both species by longer snout (4.2 in HL), the beginning of dorsal fin slightly anterior to gill opening, and significantly more vertebrae (VF 6���58���149) (Table 2). In addition to morphological characters, this specimen was collected from 350 m depth, a deepest record of G. ��� kidako ���. However, G. kidako is a common species abundant in shallow water according to local Japanese records and its reproduction behavior has also been reported at a shallow depth of 12 m in Sagami Bay, Japan (Moyer & Zaiser, 1982). It is not common for a muraenid to have such a wide vertical distribution. Further study is needed to confirm the identity of this specimen. Distribution and capture depth. The phylogenetic tree based on 647 bp COI gene shows that all G. kidako sequences from Taiwan are grouped with a G. kidako sequence (AP002976) from Japan, the type locality of G. kidako, and three sequences recorded as G. kidako and G. prionodon from New Caledonia (RESIC677-11) and Western Australia (FOAI571-09 and FOAO1360-18) are grouped with G. mucifer from Taiwan (Fig. 7). Furthermore, six Australian specimens are apparently G. mucifer in morphological characters and vertebral formulae, with VF 4���6, 51���55, 136���140 (QM I 5075, QM I 5141 and QM I 20041 from Queensland; CSIRO H 6465-01 and CSIRO H 8275-01, voucher specimens of FOAI571-09 and FOAO1360-18, respectively, from Western Australia; and AMS I 19103 -064 with unknown VF from New South Wales). Depending on molecular and morphological characters, the distribution ranges of G. kidako, G. niphostigmus and G. mucifer can be summarized as: (1) G. kidako is restricted to the area from Japan to Taiwan; (2) G. niphostigmus is currently known from Taiwan and Vietnam; and (3) G. mucifer is distributed over a wider range, including Hawaii, New Caledonia, Australia, and Taiwan (Fig. 8). The capture depths of G. mucifer specimens are 20���150 m in Taiwan, 12���100 m in Australia and 20���192 m in Hawaii. In general, G. mucifer inhabits a wide range of depth from 12 to 192 m. Sexual dimorphism in dentition. Sexual dimorphism in the dentition has been reported in several muraenids (Hatooka, 1986; Smith et al., 2008). Females and juveniles have more maxillary and dentary teeth than adult males and long inner maxillary teeth are lost in adult males. In our examined G. mucifer, all specimens less than 431 mm TL have inner maxillary teeth. But in specimens longer than or equal to 431 mm TL, only females have inner maxillary teeth except that one male (576 mm TL) has one on left side. There are two mid-sized male specimens (332 and 402 mm TL) with embedded inner maxillary teeth, implying a transformational stage of dentition. In addition, only immatures shorter than 339 mm and females have an additional peripheral row of small teeth outside the 3 rd ���5 th anterior dentary teeth (Fig. 4). The dentition of G. mucifer is roughly consistent with previous studies on other sexually dimorphic morays. Adult male does lose some rows of teeth, but numbers of teeth of the remaining rows in adult males are not lower than those of females and immatures. Material examined. Gymnothorax mucifer Snyder, 1904. Twenty- nine specimens, 210���666 mm TL. TAIWAN: DOS 0 6261 (552+ mm), DOS 0 6264 (592+ mm*), DOS 0 6265 (662+ mm), DOS 0 6266 (587+ mm*), DOS 0 6267 (666 mm), DOS 0 6268 (558+ mm), Hoping Island, Keelung; DOS 0 6262 (519+ mm), Aodi, New Taipei City; DOS 0 6263 (576 mm), Daxi, Yilan; TOU-AE 1949 (314 mm), TOU-AE 1950 (270 mm), TOU-AE 2350 (246 mm), TOU-AE 3471 (290 mm), TOU-AE 3665 (331 mm), TOU-AE 3692 (332 mm), TOU-AE 3776 (339 mm), TOU-AE 3782 (431 mm), TOU-AE 3783 (402 mm), TOU-AE 4163 (272+ mm), Changbin, Taitung. AUSTRALIA: AMS I 19103 -064 (274 mm), New South Wales; QM I 5075 (576+ mm), QM I 5141 (626 mm), QM I 20041 (432 mm), Queensland; CSIRO H 6465-01 (210 mm), CSIRO H 8275-01 (220 mm), Western Australia. HAWAIIAN ISLANDS: BPBM 29284 (314 mm), Hawaii; BPBM 28625 (221 mm), Molokai; BPBM 8511 (504 mm), BPBM 37046 (485 mm), BPBM 37047 (318 mm), Oahu. *Tail tip is damaged but hypural remains. Gymnothorax kidako (Temminck & Schlegel, 1846). Eight specimens, 572���676 mm TL. TAIWAN: DOS 0 3540 (658 mm), Aodi, New Taipei City; DOS 0 6258 (631 mm), DOS 0 6259 (572 mm), DOS 0 6360 (676 mm), DOS 0 6361 (626 mm), Hoping Island, Keelung; TOU-AE 4828 (633 mm), TOU-AE 4829 (595 mm), TOU-AE 4830 (666 mm), Daxi, Yilan. Gymnothorax niphostigmus Chen, Shao & Chen, 1996. Eleven specimens, 635���950 mm TL. TAIWAN: DOS 0 3536 (950 mm), Magong, Penghu; DOS 0 3537 (842 mm), TOU-AE 5582 (820 mm), TOU-AE 5606 (745+ mm), ASIZP 0 56940 (713 mm, holotype), Hoping Island, Keelung; TOU-AE 0 238 (882 mm), TOU-AE 4981 (743 mm), TOU-AE 4982 (754 mm), TFRI-TT 0 71 (635 mm, paratype), Changbin, Taitung; ASIZP 0 56941 (757+ mm, paratype), Aodi, New Taipei City; TFRI-TT 0 63 (737+ mm, paratype), Chenggong, Taitung. Gymnothorax sp. 900 mm TL. Society Islands: BPBM 31625, Moorea., Published as part of Huang, Wen-Chien, Chen, Hong-Ming & Liao, Te-Yu, 2019, Revalidation of a moray eel, Gymnothorax mucifer Snyder, 1904 (Teleostei: Anguilliformes: Muraenidae), with a revised distribution, pp. 151-165 in Zootaxa 4559 (1) on pages 154-162, DOI: 10.11646/zootaxa.4559.1.6, http://zenodo.org/record/2626781, {"references":["Snyder, J. O. (1904) A catalogue of the shore fishes collected by the steamer \" Albatross \" about the Hawaiian Islands in 1902. Bulletin of the United States Fish Commission, 22, 513 - 538.","Reece, J. S., Bowen, B. W., Smith, D. G. & Larson, A. (2011) Comparative phylogeography of four Indo-Pacific moray eel species (Muraenidae) reveals comparable ocean-wide genetic connectivity despite five-fold differences in available adult habitat. Marine Ecology Progress Series, 437, 269 - 277. https: // doi. org / 10.3354 / meps 09248","Huang, W. C., Nguyen, V. Q. & Liao, T. Y. (2018 b) First record of the snowflake? patched moray Gymnothorax niphostigmus Chen, Shao, & Chen, 1996 (Anguilliformes; Muraenidae) in Vietnam and its validity confirmed by DNA barcoding. Journal of Applied Ichthyology, 34, 687 - 690. https: // doi. org / 10.1111 / jai. 13684","Bohlke, E. B. & Smith, D. G. (2002) Type catalogue of Indo-Pacific Muraenidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 152, 89 - 172. https: // doi. org / 10.1635 / 0097 - 3157 (2002) 152 [0089: TCOIPM] 2.0. CO; 2","Bohlke, E. B. & Randall, J. E. (2000) A review of the moray eels (Angulliformes: Muraenidae) of the Hawaiian Islands, with descriptions of two new species. Proceedings of the Academy of Natural Sciences of Philadelphia, 150, 203 - 278.","De Buen, F. (1961) Peces chilenos. Familias Alepocephalidae, Muraenidae, Sciaenidae, Scorpaenidae, Liparidae y Bothidae. Revista de Biologia Marina Montemar, 1, 1 - 52.","Randall, J. E. & McCosker, J. E. (1975) The eels of Easter Island with a description of a new moray. Contributions in Science (Natural History Museum of Los Angeles County), 264, 1 - 32.","Smith, D. G., Brokovich, E. & Einbinder, S. (2008) Gymnothorax baranesi, a new moray eel (Anguilliformes: Muraenidae) from the Red Sea. Zootaxa, 1678, 63 - 68.","Smith, J. L. B. (1962) The moray eels of the western Indian Ocean and the Red Sea. Ichthyological Bulletin, 23, 421 - 444.","Ogilby, J. D. (1895) Description of a new Australian eel. Proceedings of the Linnaean Society of New South Wales (Ser. 2), 9, 720 - 721.","Bloch, M. E. & Schneider, J. G. (1801) M. E. Blochii, Systema Ichthyologiae Iconibus CX Ilustratum. Berolini. lx + 584 pp., 110 pls.","Moyer, J. T. & Zaiser, M. J. (1982) Reproductive behavior of moray eels at Miyake-jima, Japan. Japanese Journal of Ichthyology, 28, 466 - 468.","Hatooka, K. (1986) Sexual dimorphism found in teeth of three species of moray eels. Japanese Journal of Ichthyology, 32, 379 - 386.","Temminck, C. J. & Schlegel, H. (1846) Pisces. In: Siebold, P. F. de (Ed.), Fauna Japonica. Lyden Museum, Leiden, Parts 10 - 14, 173 - 269.","Chen, H. M., Shao, K. T. & Chen, C. T. (1996) A new moray eel, Gymnothorax niphostigmus, (Anguilliformes: Muraenidae) from northern and eastern Taiwan. Zoological Studies, 35, 20 - 24."]}

Gymnothorax odishi sp. nov., a new short brown moray eel, is described here on the basis of eleven specimens collected from Gopalpur, Odisha, along the east coast of India, Bay of Bengal. The species is characterized with dorsal-fin origin before gill opening, jaw pores with dark rim, small black patch just behind the eye of about eye size or more, gill opening with dark rim, two branchial pores, predorsal vertebrae 4, preanal vertebrae 55–58 and total vertebrae 133–138, three large fang like median intermaxillary teeth, uniserial maxillary and vomerine teeth. The new species is compared with all short brown unpatterned moray eels known from the world.

Gymnothorax odishi sp. nov. Proposed common name: Odisha moray (Figures 1���4, Table1) HolotypE. ZSI F 12592 /2 [690 mm total length (TL)], Gopalpur beach, Odisha, India, date of collection: 19 January 2018. ParatypES. ZSI F 12593 /2 (420 mm TL), EBRC /ZSI/F 9482 (522 mm TL), F9483 (423 mm TL), F9484 (602 mm TL), F9485 (522 mm TL), F9486 (650 mm TL), F9487 (500 mm TL), F9488 (550 mm TL), F9489 (696 mm TL), F9490 (700 mm TL), date and place of collection: same as holotype. DiagnoSiS. This new species belongs to the small brown unpatterned group of moray eels, with the following combination of characters: dorsal-fin origin before gill opening, jaw pores with dark rim, small black patch just behind eye of about eye size or larger, gill opening with dark rim, two branchial pores, predorsal vertebrae 4, preanal vertebrae 55���58 and total vertebrae 133���138, three large fang-like median intermaxillary teeth, uniserial maxillary and vomerine teeth. Dentary biserial with one tooth in each side in the second row of dentary. DESCription. A medium-sized, uniform brown moray with moderately elongate body, depth at gill opening 11.5���18.4 and at anus 17.3���24.0 in TL. Preanal length 1.9���2.1 in TL. Head small 6.7���7.6 in TL; snout short and blunt, its length 4.3���5.4 in HL; jaws almost equal, 2.2���2.8 in HL. Eye large, located slightly closure to rictus than the snout tip, eye diameter 8.6���10.8 in HL; interorbital space 6.0��� 7.9 in HL (Table 1). Anterior nostril a moderately broad tube, posterior nostril above anterior margin of eye. Pre-dorsal length 8.4���10.8 in total length. Dorsal-fin origin closer to rictus than to gill opening but before gill opening and well before and above the 1 st branchial pore. Anal fin starts from anus. Both dorsal and anal fins high. Gill opening at mid-side in a small diagonal slit. Teeth smooth, uniserial; outer intermaxillary teeth 4���8 in each side; median intermaxillary teeth 3, strong, curved, placed almost equidistance from each other; maxillary teeth uniserial, 12���14 on each side; vomerine teeth uniserial with 5���6 teeth; lower jaw with outer row of 20���24 teeth on each side and 1 inner tooth on each side near anterior end (Fig. 2). Head pores typical, supraorbital pores 3; infraorbital pores 4; mandibular pores 6���7; branchial pores 2, located above and before gill opening (Fig. 3). All head pores in dark rim. Predorsal vertebrae 4, preanal vertebrae 55���58 and total vertebrae 133���138. Colour. When fresh, body brown; upper and lower jaw and portion upto gill with yellow patches (Fig. 1 & Fig.4) which disappear immediately after preservation; small black patch just behind the eye of about eye size or more, gill opening with dark rim; rictus dark, fins dark; eye ring whitish, jaw pores with dark rim. When preserved, body uniformly brown, eye ring white. DiStribution. Presently known only from Gopalpur-on-Sea, on the southernmost part of the Odisha coast of India in the Bay of Bengal. Etymology. The species is named ��� odishi ��� on the name of the state Odisha, from where the specimens were collected., Published as part of Mohapatra, Anil, Mohanty, Swarup Ranjan, Smith, David G., Mishra, Subhrendu Sekhar & Roy, Sanmitra, 2018, Gymnothorax odishi sp. nov. (Muraenidae: Muraeninae), a short brown unpatterned moray eel from Bay of Bengal, India, pp. 123-130 in Zootaxa 4420 (1) on pages 124-126, DOI: 10.11646/zootaxa.4420.1.7, http://zenodo.org/record/1247220

Gymnothorax enigmaticus McCosker & Randall, 1982 Gymnothorax enigmaticus McCosker & Randall, 1982:18, figs. 1, 4 (type locality: off Ngatchab Beach, Angaur Island, Palau Islands, Western Caroline Islands, western Pacific). Smith, 2012:14. Huang et al., 2017:90. Remarks. Huang et al. (2017) reported the first record of this species for Taiwan, based on one specimen collected from off Jihui, eastern Taiwan. Two specimens were subsequently collected from Kenting, off southern Taiwan and Green Island (Ludao) off southeastern Taiwan by J.-F Huang. The species is widespread in the Indo-west Pacific Ocean., Published as part of Ho, Hsuan-Ching, Smith, David G., Tighe, Kenneth A., Hibino, Yusuke & Mccosker, John E., 2018, Checklist of eels of Taiwan (orders Anguilliformes and Saccopharyngiformes): An update, pp. 5-17 in Zootaxa 4454 (1) on page 12, DOI: 10.11646/zootaxa.4454.1.3, http://zenodo.org/record/1446687, {"references":["McCosker, J. E. & Randall, J. E. (1982) Synonymies of Indian Ocean eels, with the description of Gymnothorax enigmaticus, a moray previously known as G. ruppeli. Proceedings of the California Academy of Sciences (Series 4), 43 (2), 17 - 24.","Smith, D. G. (2012) A checklist of the moray eels of the world (Teleostei: Anguilliformes: Muraenidae). Zootaxa, 3474, 1 - 64.","Huang, W. - C., Shiao, J. - C. & Liao, T. - Y. (2017) New record of the enigmatic moray Gymnothorax enigmaticus McCosker and Randall, 1982 (Anguilliformes: Muraenidae) from Taiwan. Platax, 14, 89 - 95."]}

* Gymnothorax pseudoprolatus Smith, Hibino & Ho, this volume Gymnothorax pseudoprolatus Smith, Hibino & Ho, 2018a: this volume (type locality: Dong-gang, Pingtung, southwestern Taiwan). Remarks. The holotype and only known specimen was collected from southwestern Taiwan, off Dong-gang.

Two new species of elongate unpatterned moray eels related to Gymnothorax prolatus are described. Gymnothorax pseudoprolatus sp. nov. is described from one specimen collected off southwestern Taiwan and Gymnothorax vietnamensis sp. nov. is described from seven specimens collected off central Vietnam. Both differ from other species in the group in the number of vertebrae, position of anus, and dentition. Gymnothorax pseudomelanosomatus Loh et al., 2015 is reported from southern Taiwan for the first time; it was previously known from the type series collected off the northeastern tip of Taiwan. Gymnothorax angusticauda (Weber & de Beaufort 1916) is reported for the first time in Taiwan and the Philippines.

A new identification of Gymnothoraxminor (Temminck & Schlegel, 1846) is documented based on morphological characteristics and DNA barcoding. Sixty-one individuals of G.minor were collected from the East China Sea and the South China Sea. This species was previously reported as Gymnothoraxreticularis Bloch, 1795 in China because of the similarity in external shape and color. Gymnothoraxminor can be easily distinguished from G.reticularis by its color pattern of 18–20 irregular dark brown vertical bars and the body having scattered small brown spots. Additionally, the teeth are uniserial on both jaws, and the vertebrae number 137–139. By combining congener sequences of the cytochrome oxidase I (COI) gene from GenBank, two groups were detected among all the COI sequences of the currently named G.minor, which further indicated that two valid species were present based on genetic distance. A divergence also occurred on the number of vertebrae between the northern and southern populations. The phylogenetic and morphological analysis strongly supports that the northern and southern populations of G.minor are two different species. Furthermore, the distribution area of the northern G.minor has expanded southward to 5°15'N in the South China Sea. More specimens of G.minor and G.reticularis are crucial in order to define their geographical distribution boundaries and provide the correct DNA barcoding.

Gymnothorax visakhaensis sp. nov. Proposed common name: Visakha Moray (Figs. 1–4; Table-1) Holotype. ZSI F 12075 /2 (308 mm), Visakhapatnam fishing harbour (17° 42.452'N; 83° 18.823'E), Andhra Pradesh, along the south east coast of India, 0 6 December 2016. Paratypes. MARC/ZSI/ F4780 (310 mm); MARC/ZSI/ F4781 (271 mm); MARC/ZSI/ F4782 (300 mm); MARC/ZSI/ F4783 (274 mm); MARC/ZSI/ F4784 (311 mm) and ZSI F 12076/2 (288 mm), collection details same as holotype. Diagnosis. Gymnothorax visakhaensis sp. nov. shares the following combination of characters: body uniform brown, anus at about mid-point of body; snout blunt and moderate; eyes comparatively large and slightly closer to rictus than to tip of snout; dorsal-fin margin black; 2 branchial pores; 6 mandibular pores; sharply pointed and depressible teeth; intermaxillary teeth biserial, maxillary teeth uniserial; pre-dorsal vertebrae 7, pre-anal vertebrae 70–74, and total vertebrae 163–169. Description. Proportions of total length: preanal length 1.9–2.1, predorsal length 11.7–12.3, head length (HL) 8.4–8.8, depth at gill opening 27.1–30.0, depth at anus 27.4–30.8. Proportions of head length: snout 4.6–5.5, horizontal eye diameter 8.8–11.0, upper jaw length 2.3–2.8, lower jaw length 2.5–3.1. Vertebrae: predorsal 7, preanal 70–74, total 163–169. Body slender, elongate and tapering, anus at about midpoint. Dorsal and anal fins continuous with caudal fin around tail tip; dorsal-fin origin before gill opening, anal-fin origin immediately behind anus; pectoral and pelvic fins absent. Head long, snout blunt and short, eye relatively large, lower jaw slightly shorter than upper. Anterior nostril tubular and long, nearly reaching tip of snout. Posterior nostril a simple round pore, situated above anterior margin of eye. Gill opening a small slit on side below the mid-line. Branchial pores 2, anterior to gill opening, one of which is located before dorsal-fin origin; no lateral-line pores on body. Supraorbital pores 3: first (ethmoidal) near tip of snout just above lip, second above base of anterior nostril, third on snout between anterior and posterior nostrils. Infraorbital pores 4: first just behind base of anterior nostril, second on upper lip about midway between anterior nostril and anterior edge of eye, third below anterior margin of eye, fourth below posterior margin of eye. Mandibular pores 6, all before rictus. The dental pattern is shown in Fig. 3. Teeth in jaws smooth, needle-like, enlarged, sharply pointed and depressible. Intermaxillary teeth biserial, enlarged and fang-like, outer and inner series with 4 teeth on each side, outer row with one tooth near the mid-point of the two sides (1st median teeth); inner row also with one median tooth (Fig. 4). Maxillary teeth uniserial, about 14 on each side. Vomerine teeth 5–6, uniserial. Dentary teeth uniserial with 12–14 teeth in each side, however in one specimen (ZSI F 12076 /2; TL: 288 mm) the dentary is biserial with 5 teeth in the inner row. Colour. Body uniformly dark brown without spots or patterns. Ventral portion of lower jaw to gill opening slightly lighter brown than rest of body. Snout darker (almost black) in comparison to rest of body. Margin of dorsal and anal fins black. Eye rim pale white. On preservation in formalin, the colour remains the same. Distribution. The species is currently known only from the Andhra Pradesh coast of India in the Bay of Bengal. Etymology. The species is named Gymnothorax visakhaensis with reference to the collection site Visakhapatnam fishing harbour.

Gymnothorax tile (Hamilton, 1822) Muraenophis tile Hamilton, 1822: 18, 363 (Ganges River estuaries near Calcutta, India). No types known. Lycodontis literata McClelland, 1844: 186, 206, 215, pl. 7 (fig. 2) (Calcutta market, India). No types known. Strophidon maculata McClelland, 1844: 203, 215, pl. 8 (fig. 1). Replacement name for M. tile. Muraena vermiculata Richardson, 1848: 92 (India). Holotype BMNH 2008.4.21.2. Muraena gracilis Richardson, 1848: 92 (India). Lectotype BMNH 2008.4.9.5, designated by B��hlke & Smith, 2002: 113. Thyrsoidea microdon Kaup, 1856: 62 (Pondicherry, India). Holotype MNHN B-2469. Gymnothorax borneensis Bleeker, 1863: 169 (Borneo). Holotype RMNH 3797. Material examined. India: BMNH 2008.4.9.5 (1, 307 mm, lectotype of Muraena gracilis); BMNH 2008.4.9.6 (1, 255 mm, paralectotype of Muraena gracilis); BMNH 2008.4.21.2 (1, 435 mm, holotype of Muraena vermiculata); MNHN B.2469 (1, 464mm, holotype of Thyrsoidea microdon); MARC/ZSI F4530 (1, 711 mm); MARC/ZSI F4531 (1, 409 mm); MARC /ZSI F4533 (1, 645 mm); ZSI F 13843 (1,450 mm). Myanmar: USNM 372146 (1, 175 mm), at mouth of Yangon River, 29 Oct 1997. Malaysia: USNM 278309 (3,135��� 184 mm), Strait of Malacca, Parit Jawa, 4 Dec 1985; USNM 278418 (3, 93��� 232 mm), south-western Malay Peninsula, southside of Muar River, Muar, Johore, 25 Nov 1985; USNM 278436 (1,87 mm), same, 2 Dec 1985; USNM 278465 (2, 154��� 230 mm), same, 27 Nov 1985. Indonesia: RMNH 3797 (1, 217 mm, holotype of Gymnothorax borneensis, Borneo; USNM 75894 (1, 470 mm), westcoast of Borneo, Sangkulirang, 19 Dec 1913. Diagnosis. Teeth moderately stout, intermaxillary teeth in 5 rows; dorsal-fin origin over or slightly before gill opening; dorsal and anal fins high with a pale margin; predorsal vertebrae 9���11. Description. Proportions in TL: preanal 1.7���2.3, predorsal 7.6���9.8, head 6.6���8.8, depth at gill opening 14���29, depth at anus 16���26.Proportions in head length: snout 5.8���8.0, eye 9.8���17, upper jaw 3.1���3.9, lower jaw 3.4���4.4, eye 9.8���17. Pores: branchial 2, supraorbital 3, infraorbital 3, mandibular 6. Vertebrae: predorsal 9���11, preanal 55��� 59, total 119���131. Body moderate, anus near mid-length. Head moderate; snout blunt with large eye located near middle of jaw; lower jaw slightly shorter than upper jaw. Gill opening rounded, on side below midline. Dorsal and anal fins relatively high, dorsal-fin origin over or slightly before gill opening. Teeth moderately stout, not long and needlelike; intermaxillary teeth in five rows across, the outer teeth smaller than intermediate teeth; three large median teeth; maxillary teeth biserial, inner teeth fewer and larger; dentary teeth biserial anteriorly, uniserial posteriorly, inner teeth fewer and larger; vomerine teeth uniserial at anterior and posterior end, irregularly biserial in middle. Brown with moderate to small irregular white or yellow spots; dorsal and anal fins without pale margin. Distribution. Known from the northern Indian Ocean and western Pacific, from India to Malaysia, Philippines, and Indonesia. Also recorded from Papua New Guinea by Fricke et al., 2014. Occurs primarily in estuarine habitats, entering brackish and fresh water. Remarks. This species closely resembles Gymnothorax pseudotile; the distinctions are discussed above. The two specimens from Borneo have fewer vertebrae (119 and ca. 121) than those from elsewhere (125���131). Our sample size is too small to assess the significance of this, and there are no other obvious differences. The type specimens of the three synonyms described from India (Muraena vermiculata, M. gracilis, and Thyrsoidea microdon) are reported to have fewer vertebrae (122���126) than our specimens from the same area (125���129) (data from B��hlke & Smith, 2002). We have not been able to check these specimens ourselves, and we have no explanation for the discrepancy., Published as part of Mohapatra, Anil, Smith, David G., Ray, Dipanjan, Mishra, Subhrendu Sekhar & Mohanty, Swarup Ranjan, 2017, Gymnothorax pseudotile sp. nov. (Muraenidae: Muraeninae) from Bay of Bengal India, pp. 586-592 in Zootaxa 4286 (4) on pages 589-591, DOI: 10.11646/zootaxa.4286.4.11, http://zenodo.org/record/828715, {"references":["Hamilton, F. (1822) An account of the fishes found in the river Ganges and its branches. Printed for A. Constable and company, Edinburgh, vii + 405, 39 pls.","McClelland, J. (1844) Apodal fishes of Bengal. Calcutta Journal of Natural History, 5 (18), 151 - 226, pls. 5 - 14.","Bohlke, E. B. & Smith, D. G. (2002) Type Catalogue of Indo-Pacific Muraenidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 152, 89 - 172.","Kaup, J. J. (1856) Catalogue of apodal fish in the collection of the British Museum. Published by order of the Trustees, London, 163 pp., 19 pls.","Bleeker, P. (1863) Sur quelques especes nouvelles ou peu connues de Gymnothorax Bl. de l'Inde archipelagique. Nederlandsch Tijdschriftvoor de Dierkunde, 1, 167 - 171.","Fricke, R., Allen, G. R., Andrefouet, S., Chen, W. - J., Hamel, M. A., Laboute, P., Mana, R., Tan, H. H. & Uyeno, D. (2014) Checklist of the marine and estuarine fishes of Madang District, Papua New Guinea, western Pacific Ocean, with 820 new records. Zootaxa, 3832 (1), 1 - 247."]}

Cytogenetic studies in Gymnothorax funebris revealed a diploid chromosome number 2n = 42 (6 metacentrics, 4 submetacentrics, and 32 acrocentrics, FN = 52). The results obtained are novel and similar to those previously described for species belonging to Muraenidae family. The conventional karyotype is also novel and divergent from other species of the genus Gymnothorax, where a higher proportion of metacentric chromosomes predominate. The data are reported and discussed considering the cytotaxonomy of the genus. These results strongly support the current view that chromosomal alterations such as centric fusion and Robertsonian's translocations have an important role in the evolution of this group.

Two forms of muraenid leptocephali, collected from the western Pacific Ocean, were identified as Gymnothorax sagmacephalus Bohlke 1997 and Gymnothorax albimarginatus (Temminck and Schlegel 1846) on the basis of morphometric and genetic analyses. The leptocephali of each species were characterized, respectively, by counts of 172–175 and 186–191 myomeres, 43–44 and 47 predorsal myomeres, 109–113 and 127–134 preanal myomeres, and 100–104 and 118–119 last vertical blood vessel myomeres. Gymnothorax sagmacephalus leptocephali had minute melanophores over much of the head and body, closely resembling the condition in Gymnothorax minor (Temminck and Schlegel 1846), whereas those of G. albimarginatus not only had minute melanophores over much of the head and body, but also a pair of melanophore groups on the posteroventral and posterodorsal aspects of the head. Such groups are here considered to represent highly specific characters. Although a previous opinion postulated that G. sagmacephalus is a juvenile of G. albimarginatus, and the adult morphologies of the two species have a lot in common, they clearly differ in both leptocephalus morphology and genetic sequence. Therefore, G. sagmacephalus was concluded as being a valid species.

Gymnothorax indicus sp. nov. Figs. 1���3, Tables 1���3 Proposed common name: Indian unpatterned moray Holotype. ZSIF 11370 /2 (355 mm), Shankarpur, West Bengal, India (Trawling Site 21 ��06.55 'N, 87 ��58.68 'E), 22 December 2014. Paratypes. MARC/ZSI/F4412 (305 mm), MARC/ZSI/F4413 (335 mm) and ZSI F 11371/2 (310 mm), collection details same as for holotype. Diagnosis. This species belongs to the elongate, brown unpatterned group of moray eels which share the following combination of characters: anus at about mid-point of body; snout blunt and short; eyes comparatively large and slightly closer to rictus than to tip of snout; dorsal-fin margin black; 5 mandibular pores; maxillary teeth uniserial, sharply pointed; MVF: 9-79-194. Description. Proportions of TL: preanal length 2.0, predorsal length 10.1���10.6, head length 8.0���8.9, depth at gill opening 23.5���23.7, depth at anus 30.5���32.2. In head length: snout 8.1���8.7, horizontal eye diameter 6.9���9.1, upper jaw length 4.2���4.8. Vertebrae: predorsal 9, preanal 79, total 194. Body slender, elongate and tapering, anus at about midpoint. Dorsal and anal fins continuous with caudal fin around tail tip; dorsal-fin origin before gill opening, anal-fin origin immediately behind anus; pectoral and pelvic fins absent. Head long, snout blunt and short, eye relatively large, lower jaw slightly shorter than upper. Anterior nostril tubular and long, nearly reaching the tip of snout. Posterior nostril a simple round pore, situated above and behind anterior margin of eye. Gill opening a small slit on side below mid-line. Branchial pores 3, anterior to gill opening, two of which are before dorsal-fin origin; no lateral-line pores on body. Supraorbital pores 3: first (ethmoidal) near tip of snout just above lip, second above base of anterior nostril, third on snout between anterior and posterior nostrils. Infraorbital pores 4: first just behind base of anterior nostril, second on upper lip about midway between anterior nostril and anterior edge of eye, third below anterior margin of eye, fourth below posterior margin of eye. Mandibular pores 5, all before rictus. Teeth in jaws smooth, sharply pointed, not greatly enlarged, needle-like and depressible; intermaxillary teeth in a single outer series, about 6 (7) on each side, the anterior tooth on each side enlarged and fang-like; no median teeth; maxillary teeth uniserial, about 14���16 on each side; vomerine teeth 8, uniserial; dentary teeth uniserial, 19��� 20. Colour. When fresh, body uniformly pale brown without spots or patterns; margin of dorsal and anal fins dark; eye rim pale. On preservation, colour faded from brown to light brown. Distribution. The species is currently known only from the West Bengal coast of India in the northern Bay of Bengal. Etymology. The species is named with reference to the country ���India���, from within the Exclusive Economic Zone of which the specimens were captured by a trawler. Discussion. B��hlke (1997) reviewed the taxonomy of eight species of elongate unpatterned morays known from the Indo-Pacific region: Gymnothorax albimarginatus, ���G���. dorsalis, G. phasmatodes, G. prolatus, G. sagmacephalus, G. verrilli, Pseudechidna brummeri and Strophidon sathete. B��hlke & Randall (2000) described another elongate unpatterned moray, Gymnothorax polyspondylus, from Hawaii based on a single specimen. Later Loh et al. (2011) described another species Gymnothorax melanosomatus, with a key for nine species of elongate morays excluding G. verrilli. Loh et al. (2015) added another new species, Gymnothorax pseudomelanosomatus, to this group. The new species described herein brings the total to twelve species of elongate unpatterned moray eel from the Indo-Pacific region. A comparison of characters of all twelve species is given in Table 2 and their comparative meristics of those from Indian waters are given in Table 3. Characters Gymnothorax G. albimarginatus G. melanosomatus G. phasmatodes G. prolatus G. polyspondylus indicus sp. nov. Vertebrae ������continued on the next page Continued. Characters G.pseudo- G. sagmacephalus G. verrilli Strophidon dorsalis Strophidon sathete Pseudechidna melanosomatus brummeri Vertebrae Predorsal 5���7 6 ― 7 5 8 ― 9 8 ― 11 4 ― 7 anal 89 74 ― 78 71 ― 77 62 ― 65 73 ― 82 89 ― 98 Total 202���206 170 ― 176 170 ― 171 154 ― 163 186 ― 208 201 ― 214 Proportion in TL Preanal length 2.01 ― 2.02 1.9 ― 2.0 2.0 ― 2.0 2.3 ― 2.4 2.3 ― 2.9 2.0 ― 2.3 Head length 11.2 ― 11.4 8.7 ― 10.3 9.4 ― 11 8.4 ― 9.3 8.5 ― 14 11 ― 16 Pseudechidna brummeri has an extremely elongate and slender body with body depth at gill opening about 40 or more in total length, comparable only to Strophidon sathete and Gymnothorax melanosomatus. Following Loh et al. (2015), here we consider ���G���. dorsalis as Strophidon dorsalis, since the species has the anus located well before mid-body and the eye closer to the tip of the snout than to the rictus. In all other Indo-Pacific elongate, unpatterned eels of the genus Gymnothorax the preanal length lies within the range of 1.6���2.1 in total length. Further, eye diameter in Strophidon dorsalis, S. sathete and Pseudechidna brummeri is smaller in comparison to other Gymnothorax species of this group, at 12���24 in head length. The new species, Gymnothorax indicus, with preanal length 2.0 in TL, body depth at gill opening 23.4���23.7 in TL and eye diameter 6.9���9.1 in head length, is well differentiated from the other three species in the group. Gymnothorax indicus differs from all other of elongate unpatterned species in having 5 mandibular pores (cf. 6 or 7 pores). Further, the presence of 14���16 maxillary teeth and 19���20 dentary teeth in the new species differentiates it from all other Gymnothorax species in the elongate unpatterned moray group except for G. prolatus, which can be distinguished in having biserial teeth (vs uniserial in G. i n d i c u s), shorter head, longer snout and longer upper jaw (Table 2). There are more predorsal vertebrae (9) in the new species compared to other species of elongate unpatterned Gymnothorax (4���8) and Pseudechidna (4���7). Although the number of predorsal vertebrae is within the range of Strophidon species, the total vertebrae and the preanal vertebra are higher in comparison to S. dorsalis and, apart from the other differences stated earlier, S. sathete has biserial teeth in the jaws and more teeth in the maxillary and outer dentary. Further, four infraorbital pores in the new species differentiates it from G. dorsalis. Uniserial maxillary teeth differentiate the new species from Gymnothorax melanostomatus, G. prolatus, Strophidon dorsalis, S. sathete and Pseudechidna brummeri. Total vertebrae in Gymnothorax melanosomatus, G. polyspondylus, G. pseudomelanosomatus and Pseudechidna brummeri are always more than 200, while in G. phasmatodes, G. sagmacephalus, G. verrilli and Strophidon dorsalis they are considerably fewer. Gymnothorax prolatus has slightly fewer total vertebrae than the new species, G. i n d i c u s, and this character is overlapped only in G. albimarginatus and Strophidon sathete. But G. albimarginatus differs from the new species in having fewer predorsal vertebrae, more preanal vertebrae and greater preanal length (Table 2). The new species resembles G. phasmatodes in having a similar number of preanal vertebrae and proportionately similar preanal length, but the latter species has nostril and head pores with prominent white spots (white spots absent in G. indicus), margin of dorsal fin whitish (dark in G. i n d i cu s) and fewer vertebrae. Further, it has a shorter head, longer upper jaw and longer snout (Table 2). Among the long unpatterned moray eels, distribution of G. verrilli is confined to Eastern Central Pacific, between Baja California and Panama. Two recently described species, G. melanosomatus and G. pseudomelanosomatus, are known from Taiwan and G. polyspondylus from Hawaii, whereas, G. sagmacephalus occurred between Taiwan and Japan. One species, G. albimarginatus, is known to have a wider distribution in Western and central Pacific only. Only three species, i.e., P. brummeri, S. sathete and G. phasmatodes, are known to have much wider distribution in the Indo-west Pacific region. Occurrence of G. prolatus, apart from its type locality Taiwan, was recorded very recently from northern Indian Ocean (Mohapatra et al., 2015) and S. dorsalis recorded from northern Bay of Bengal (Ray & Mohapatra, 2015) extending its known range westward from western Pacific. The new species, Gymnothorax indicus sp. nov., forms the sixth long unpatterned moray from Indian Ocean., Published as part of Mohapatra, Anil, Ray, Dipanjan, Smith, David G. & Mishra, Subhrendu Sekhar, 2016, A new species of elongate unpatterned moray eel of the genus Gymnothorax (Muraenidae: Muraeninae) from the Bay of Bengal, pp. 591-598 in Zootaxa 4150 (5) on pages 592-597, DOI: 10.11646/zootaxa.4150.5.6, http://zenodo.org/record/260647, {"references":["Bohlke, E. B. (1997) Notes on the identity of elongate unpatterned Indo-Pacific morays, with description of a new species (Muraenidae, Subfamily Muraeninae). Proceedings of the Academy of Natural Sciences of Philadelphia, 147, 89 - 109.","Bohlke, E. B. & Randall, J. E. (2000) A review of the moray eels (Angulliformes: Muraenidae) of the Hawaiian Islands, with descriptions of two new species. Proceedings of the Academy of Natural Sciences of Philadelphia, 150, 203 - 278, Pls. I - IX.","Loh, K. H., Shao K. T. & Chen, H. M. (2011) Gymnothorax melanosomatus, a new moray eel (Teleostei: Anguilliformes: Muraenidae) from southeastern Taiwan. Zootaxa, 3134, 43 - 52.","Loh, K. H., Shao, K. T., Ho, H. C., Lim, P. E. & Chen, H. M. (2015) A new species of moray eel (Anguilliformes: Muraenidae) from Taiwan, with comments on related elongate unpatterned species. Zootaxa, 4060 (1), 30 - 40. http: // dx. doi. org / 10.11646 / zootaxa. 4060.1.5"]}

New species of the genera Gymnothorax (fam. Muraenidae) and Parapercis (fam. Mugiloididae) are described from coastal waters of central Vietnam. G. emmae sp. n. is a common species all along the coast from the Van Phong Bay to the Phan Thiet Bay and differs from other species of this genus in characters of coloration, dentition, and the vertebral formula. P. bicoloripes sp. n. is common in the Phan Thiet Bay and is also know from one specimen from the Nha Trang Bay; from other species of the genus, it differs in coloration and meristic characters.

Two new species of bucephalid trematodes are described from the rectum and intestine of the western moray eel Gymnothorax woodwardi McCulloch (Anguilliformes: Muraenidae: Muraeninae) off Point Peron in Western Australia. Dollfustrema gibsoni n. sp. is distinguished by body size, a pharynx that is intertesticular and level (latero-dextrally) with the anterior portion of the cirrus-sac, an ovary positioned dextrally to the testes and slightly anterior to (in part) the anterior testis, a uterus that extends anteriorly to the vitelline follicles but not to the level of the rhynchus, and vitelline follicles that form a confluent arc anterior to the gonads. Muraenicola nom. nov. is proposed as a replacement generic name for the pre-occupied Folliculovarium Gu & Shen 1983 nec Singh & Sinha 1981. Muraenicola botti n. sp. is distinguished from its congeners by body size, the size of the cirrus-sac (relative to body size), and in possessing tegumental spines, testes that are oblique (rather than in tandem) and eight ovarian lobes. It differs further in having an intestinal caecum that extends anteriorly to the level of the vitelline follicles and in the position of the pharynx and cirrus-sac relative to each other (lateral in part) as well as to the gonads. M. botti n. sp. also lacks a metraterm. These are the first reports of these genera from fishes off Australia and from the southern hemisphere.

Based on light and scanning electron microscopical studies of nematode specimens from the digestive tract of some rarely collected anguilliform and perciform fishes off New Caledonia, three new species of Cucullanus Müller, 1777 (Cucullanidae) are described: C. austropacificus n. sp. from the longfin African conger Conger cinereus (Congridae), C. gymnothoracis n. sp. from the lipspot moray Gymnothorax chilospilus (Muraenidae), and C. incognitus n. sp. from the seabream Dentex fourmanoiri (Sparidae). Cucullanus austropacificus n. sp. is characterized by the presence of cervical alae, ventral sucker, alate spicules 1.30-1.65 mm long, conspicuous outgrowths of the anterior and posterior cloacal lips and by elongate-oval eggs measuring 89-108 × 48-57 μm; C. gymnothoracis n. sp. is similar to the foregoing species, but differs from it in the absence of cervical alae and the posterior cloacal outgrowth, in the shape and size of the anterior cloacal outgrowth and somewhat shorter spicules 1.12 mm long; C. incognitus n. sp. (based on female morphology) differs from other congeneric species parasitic in the Sparidae mainly in possessing cervical alae, the postequatorial vulva, phasmids situated at the mid-length of the tail and in the size of the eggs (75-84 × 45-66 μm). A key to species of Cucullanus parasitizing anguilliform fishes is provided.Trois nouvelles espèces de Cucullanus (Nematoda: Cucullanidae) de poissons marins au large de la Nouvelle-Calédonie, avec une clé des espèces de Cucullanus d’Anguilliformes.Sur la base d’une étude en microscopie photonique et électronique à balayage de spécimens de nématodes provenant du tube digestif de quelques poissons anguilliformes et perciformes rarement pêchés en Nouvelle-Calédonie, trois nouvelles espèces de Cucullanus Müller, 1777 (Cucullanidae) sont décrites : C. austropacificus n. sp. de Conger cinereus (Congridae), C. gymnothoracis n. sp. de Gymnothorax chilospilus (Muraenidae) et C. incognitus n. sp. de Dentex fourmanoiri (Sparidae). Cucullanus austropacificus n. sp. est caractérisé par la présence d’ailes cervicales, une ventouse ventrale, des spicules ailés de 1,30-1,65 mm de long, des excroissances remarquables des lèvres cloacales antérieure et postérieure et d’oeufs ovales et allongés mesurant 89-108 × 48-57 μm. C. gymnothoracis n. sp. est semblable à l’espèce précédente, mais en diffère en l’absence d’ailes cervicales et d’excroissance cloacale postérieure, par la forme et la taille de l’excroissance cloacale antérieure et par des spicules un peu plus courts de 1,12 mm. C. incognitus n. sp. (basé sur la morphologie de la femelle) diffère des autres espèces congénères parasites chez les Sparidae principalement par la possession d’ailes cervicales, une vulve postéquatoriale, des phasmides situés à mi-longueur de la queue et par la taille des oeufs (75–84 × 45–66 μm). Une clé des espèces de Cucullanus parasitant les poissons anguilliformes est fournie.

Herein is given information on eight species of morays recorded from the soft grounds in the Bay of Nha Trang (Central Vietnam); one of these species is, obviously, new for science. Gymnothorax prolatus formerly known by two specimens from off Taiwan is recorded for the first time for Vietnam. There are doubts in the correctness of separation of the genera Strophidonand Pseudechidnafrom the genus Gymnothorax in its present understanding. Herein is given a description of a juvenile specimen of the genus Gymnothoraxfrom the coral reefs of Tre Island, which does not fit into the descriptions of all the known species of this genus. A key for identification of morays inhabiting the soft grounds of the Bay of Nha Trang is given.