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While kelp forests are some of the best-surveyed ecosystems in California, information on cryptic inhabitants and their role within the community are lacking. Kelp itself provides overall structure to the habitat; however the rocky reef to which the kelp attaches is known to provide additional structure for cryptic species. Gymnothorax mordax, the California moray, is an elusive predatory species that is considered abundant in the waters around Catalina Island. However, no life history data exists for this species. We examined habitat composition, relative abundance, size pattern distributions, and biomass of G. mordax within Two Harbors, Catalina Island. Habitats were sampled using a combination of baited trap collection and transect surveys using SCUBA. A total of 462 G. mordax were captured, primarily in shallow (< 10 m) waters. Individuals of G. mordax were associated with mostly boulder and cobble substrates. Measurements of relative abundance and density indicate that G. mordax is more prevalent than reported in previous studies. We also discovered that the 6 trapping sites from which all morays were collected, differed in size structuring and density while the relatively high biomass did not change across sites. In general, southern facing sites exhibited higher densities of morays, while northern facing sites showed more size structuring. We show how the structural complexity of the rocky reef habitat in an already diverse kelp forest ecosystem, can support a high biomass of a cryptic elongate predatory fish.

Moray eels comprise a large radiation of elongate marine predators that are thought to swallow large prey whole but also circumvent gape constraints by manipulating prey into more manageable pieces. Prey manipulation behaviors include shaking, rotation, knotting, and ramming prey against another object to assist in swallowing. Most morays feed on a wide variety of prey that vary in mechanical properties such as stiffness and toughness, which could potentially affect feeding behaviors. There is little diet data informing us of the maximum prey size morays can swallow whole and whether maximum prey size differs between prey types. Our study examines feeding behaviors for the California moray (Gymnothorax mordax) in the laboratory. We recorded morays feeding on freshly thawed fish and cephalopods of varying size. We found that prey size had a strong effect on total feeding time and manipulation duration for both fish and cephalopods. While morays were observed using a diversity of prey manipulation behaviors and the durations for each of these behaviors increased with prey size, prey type had no effect on manipulation behaviors employed. Total manipulation duration, however, comprised a greater proportion of total feeding time for fish compared to cephalopods. As relative prey mass (RPM) increased for cephalopods, morays spent a greater proportion of their total feeding time transporting prey. Transport rate was higher for cephalopod prey but the relationship between RPM and transport rate was negative for both prey types. Despite this decrease in transport rate, we attribute the lower total feeding times for larger cephalopod prey compared to fish to behavioral tactics of morays. Morays used the corners of the aquaria to aid in the transport of larger cephalopod prey. We hypothesize that the deformable tissues of cephalopods and the presumably low coefficient of friction of their thawed mantles and tentacles may be difficult for the recurved teeth on the pharyngeal jaws to pierce and grip during transport.

Scaling patterns of tooth morphology can provide insights on prey capture strategy and dietary patterns as species grow through ontogeny. We report the scaling of dentition and diet and how it relates to body size in the California moray, Gymnothorax mordax. We sampled lengths, widths, and curvature for teeth lining five distinct regions of the oral jaws across 21 G. mordax individuals ranging from 383 to 1110 mm total length. Absolute tooth length in relation to moray size shows positive allometry only for the outer maxillary teeth, while teeth lining the inner maxilla display positive allometry in tooth base width. All other regions exhibit isometric growth in both length and width relative to moray size. Similar to previous descriptions of other moray species, the longest teeth in the oral jaws are the median intermaxillary teeth. This series of three teeth are depressible and rooted in the center of the ethmovomer, the bone that forms the roof of the rostrum. We hypothesize that caudal mobility of the median intermaxillary teeth aids in prey transport by enabling the pharyngeal jaws to remove pierced prey without requiring full abduction of the oral jaws. The predominantly isometric tooth growth in G. mordax suggests that the oral teeth grow proportionately as individuals increase in size. Stomach contents from the field suggest that G. mordax is highly piscivorous. While a strong positively allometric relationship between vertical gape and head length supports the expectation that moray increase relative prey size over ontogeny, we found no relationship between prey standard length and moray size. This suggests that while larger individuals are capable of consuming larger prey, individual G. mordax are opportunistic predators that do not specialize on prey of a specific size over ontogeny.

Lionfish (Pterois miles) were observed avoiding coral pinnacles inhabited by the moray eels Gymnothorax flavimarginatus and G. javanicus in the northern Red Sea, Egypt. Release of lionfish (Standard Length 93–104 mm) in such coral pinnacles in November 2016 resulted in almost immediate predation by large moray eels (Total Length > 1 m). Predation by moray eels may be the key control mechanism of population growth in the native biogeographical range of Pterois spp. and may indirectly explain the success of the invasive populations. This is the first video-documented record of moray eels feeding on the lionfish P. miles.

This study describes the length‐weight and length‐length relationships for four species of Garra, viz., G. amirhosseini, G. gymnothorax, G. mondica and G. rosica in Iranian basins. The specimens were collected using hand net, cast net and also electrofishing in 27 different sites from different basins from November 2017 to June 2018. Significantly different b values from the expected value in length–weight relationships were observed for all the species but one. The value of the parameter b varied between 2.70 and 3.21 among the species.

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."]}

Diet has a strong influence on the morphology and physiology of vertebrates. Members of numerous fish clades have provided excellent comparative models for examining the effects of diet, primarily carnivory vs. herbivory, on organ systems. In this study, we comparatively examined the effects of carnivory, specifically independent evolutions of piscivory and durophagy, on body dimensions, organ topology, and organ proportions between nine moray species (Muraenidae). We found that organ placement via the anteroposterior organ positions differed between members in the two diet categories, although general organ topology is conserved. The stomach (31 - 55%) and intestine (68 - 76%) consistently occupied the majority of the body cavity irrespective of diet. Diet was shown to influence overall body dimensions and relative organ proportions across all moray species. Durophagous morays have shorter head lengths and body depths, and shorter heart, liver, gall bladder, and spleen lengths compared to piscivorous species. There was also greater variation in organ lengths among durophagous species than within piscivorous species. We attribute this greater variation in organ lengths to the multiple independent origins of durophagy within Muraenidae. Our single intraspecific comparison of the California moray, Gymnothorax mordax, a piscivorous species, revealed that all organs scale isometrically with body cavity length over ontogeny. The stomach grows proportionally with body cavity length and is the second longest organ in G. mordax at all ontogenetic stages, with the longest being the intestine. This suggests that morays can consume large quantities of prey or relatively large individual prey throughout their life history. In addition to scaling patterns of G. mordax, our study is the first to investigate the effects of diet and the intraspecific scaling patterns on the viscera of members of the Muraenidae, a clade of obligate carnivores that inhabit most marine ecosystems.

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."]}

On the basis of traditional morphological classification, Gymnothorax minor has been misidentified as Gymnothorax reticularis in China. Because of the special ecological habits and aggressiveness of this species, little research has been performed on its population genetics. Given the general decline in offshore fishery resources, it is necessary to study the genetic characteristics of G. minor in order to protect this resource. In the present study, six different geographical populations of G. minor were collected from their distribution areas in China, and their genetic signatures were investigated for the first time based on the control region fragments. The results showed that G. minor had a very high degree of genetic diversity, and there was no genetic differentiation between these populations. However, the genetic analysis failed to reject panmixia among geographic populations of this species. The life-history characteristics of G. minor and the heterogeneity of its habitat promote the rapid growth of its populations, resulting in gene mutation accumulation and rich genetic diversity. G. minor experienced a population expansion event during the late Pleistocene, which shaped its various genetic signatures. The prolonged pelagic larval duration (PLD) of this species ensures that its leptocephali effectively avoid various physical obstacles and have the opportunity to follow ocean currents to various coastal areas of China. Finally, this moray eel exchanges genetic material with recruitment populations from other locations. All the current genetic signature results for G. minor will serve as a reference for genetic research on moray eels with similar life-history characteristics.

Being able to tie their body into a knot is beneficial to some moray eels and other cylindrical vertebrates for various reasons, although not all morays’ tie knots. Knot tying can generate leverage for feeding activity by moving the knot up the body, and this leverage may be affected by knot type. Three types of knots are known to have been used by morays’ and this study further elucidates this behaviour and ability. Baited Remote Underwater Videos (BRUVs) were used to compare knot tying between the moray eels Gymnothorax prasinus and Gymnothorax prionodon in the Solitary Islands region, eastern Australia. Gymnothorax prasinus tied more knots overall and more types of knot than G. prionodon (five types vs one type, respectively). Both tied the overhand knot and this was the most common knot used by G. prasinus. The figure of eight knot was used two times more than the double overhand knot, being relatively wider and likely easier to move up the body. Gymnothorax prasinus also tied two undescribed knots, now called the ‘moray knot’ and the ‘morays banana knot’, which have more crossings and are bulkier, increasing potential for generating leverage. However, their uncommon use indicates costs may outweigh benefits of these more complex knots. Lower habitat complexity in deeper waters may influence differences in knot tying as G. prasinus were only recorded shallower than 40 m and G. prionodon mainly deeper. These findings further reveal the utility and complexity of knot tying in moray species and cylindrical vertebrates generally.

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 minor has been often confused with G. reticularis because of the similar morphological characteristics. The complete mitochondrial genome sequence of G. minor is 16,575 bp in length, in...

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."]}

Moray eels (Gymnothorax species) from tropical waters have long been known to be high-risk species, and the consumption of particularly the viscera or ungutted eels can result in severe ciguatera (known as Gymnothorax or moray eel poisoning), characterized by prominent neurological features. In this review, the main objective was to describe the risk and severity of ciguatera caused by eating moray eels in different parts of the world. Moray eels can accumulate very high ciguatoxin (CTX) levels in the flesh and particularly the liver. Therefore, even the smaller ones can be toxic and the consumption of an average portion (particularly liver) can result in severe or fatal ciguatera. Moray eels (particularly when ungutted) must never be served in gatherings since they can cause mass poisoning because of their large sizes and high CTX levels. Apart from regulatory measures restricting or excluding access, the public should be repeatedly warned to avoid eating moray eels.