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Serranus cabrilla (Linnaeus, 1758) English name: Comber Figures 3���4; Tables 1���5 Perca cabrilla: Linnaeus 1758: 294 (no locality given [Mediterranean Sea]). Paracentropristis cabrilla: Fowler 1936: 1291 Serranus cabrilla: Poll 1954:69���72, Fig. 19 (Gabon to s. Angola); Robins and Starck 1961: 261���262 (figs. 1a, 7c, 8a, 8c; + Spain and Italy); Smith & Smith 1966: 88 (in part, not painting; Atlantic species from Mediterranean southwards, colonising South Africa to Natal); Heemstra & Randall 1986: 536 (in part, not Figure or Plate; Mediterranean, eastern Atlantic, English Channel to South Africa, and round coast to Durban). Diagnosis. Dorsal-fin X, 13���14; pectoral-fin rays 14���16; gill rakers 7���8+14���16 (22���24 total); circumpeduncular scales 34���38; lateral-line scales 70���77; scales from dorsal fin to lateral line 9���12; scales from the anal-fin origin to lateral line 22���25. Caudal fin emarginate to slightly forked; cheek, nape, opercle, pectoral-fin base and chest scaly. Description. Body elongate and slightly compressed; body depth less than head length, about 3.4���5.2 in SL; head length 2.6���2.8 in SL. Maxilla extending to below middle of the eye; lower jaw projecting beyond upper jaw; opercle with three flat spines, upper two spines well developed. Snout length greater than eye diameter, snout length 8.8���11.4 in SL, eye diameter 10.0��� 13.9 in SL. Snout, interorbital and maxilla naked. Small scales present at bases of dorsal, anal and pectoral fins; caudal and pelvic fins with or without some small scales basally. Preopercle margin coarsely serrate with developed spines near angle. Premaxillary teeth in several irregular rows, outer teeth enlarged, with large canines anteriorly. Vomer more or less V-shaped, with small teeth. Palatines with patch of small teeth. The first two dorsal-fin spines shortest, the sixth spine usually longest. No distinct notch between dorsal spines and soft rays but soft rays slightly elevated. Pectoral-fin origin slightly behind pelvic-fin origin, distal tip of pectoral fin reaching anus, slightly behind anal-fin origin. Fresh Color (Fig. 4). Body light brown to reddish with eight or nine dark vertical bars dorsolaterally; two or three white horizontal bands from head to tail; ventral part of head and belly mostly white; head with two or three orange-brownish stripes below and behind eye; caudal and soft dorsal fins with small bluish dots; tips of caudal fin reddish or dark brown; pelvic and pectoral fins pale. Juveniles with white midlateral stripe bordered by two thick black stripes, the upper stripe running from tip of snout through middle of eye to upper margin of operculum onto trunk above midlateral line to upper half of caudal peduncle, the lower stripe from base of pectoral fin to lower half of caudal peduncle; the dark stripes often partially broken into dark blotches (Iwamoto & Wirtz 2018; no juvenile specimens available for examination). Color in preservation. Specimens lose all color in preservation. Distribution. From the British Isles to Angola, including Azores, Madeira, Canary Islands, S��o Tom�� and Pr��ncipe, and Cape Verde Islands, also through Mediterranean and Black Sea, and in the Red Sea (Heemstra & Anderson 2016; Iwamoto & Wirtz 2018)., Published as part of Sithole, Yonela, Heemstra, Elaine & Mwale, Monica, 2021, Revalidation and redescription of Serranus knysnaensis Gilchrist, 1904 (Perciformes: Serranidae) with a new distribution record, pp. 99-113 in Zootaxa 5057 (1) on pages 107-108, DOI: 10.11646/zootaxa.5057.1.6, http://zenodo.org/record/5705482, {"references":["Linnaeus, C. (1758) Systema Naturae. 10 th Edition. Impensis Direct. Laurentii Salvii, Holmiae, ii + 824 pp.","Fowler, H. W. (1936) The marine fishes of West Africa based on the collection of the American Museum Congo expedition, 1909 - 1915. Part II. Bulletin of the American Museum of Natural History, 70, 607 - 1493.","Poll, M. (1954) Poissons IV. Teleosteens Acanthopterygiens (premiere partie). Resultats Scientifiques. Expedition Oceanographique Belge dans les Eaux Cotieres Africaines de l'Atlantique Sud (1948 - 1949), Bruxelles, 4, 1 - 390.","Robins, C. R. &. Starck II, W. A. (1961) Materials for a revision of Serranus and related fish genera. Proceedings of the Academy of Natural Sciences of Philadelphia, 113, 259 - 314.","Smith, J. L. B. & Smith, M. M. (1966) Fishes of the Tsitsikama Coastal National Park. Swan Press Ltd, Johannesburg, 161 pp.","Heemstra, P. C. & Randall, J. E. (1986) Family Serranidae. In: Smith, M. M. & Heemstra, P. C. (eds), Smiths' Sea Fishes. MacMillan, Johannesburg, pp. 509 - 537.","Iwamoto, T. & Wirtz, P. (2018) A synopsis of the Eastern and Central Atlantic combers of the genus Serranus (Teleostei: Perciformes: Serranidae). Proceedings of the California Academy of Sciences, 65, 1 - 39.","Heemstra, P. C. & Anderson, W. D. (2016) Serranidae. Groupers (seabasses, hinds, creolefish, combers, anthiines, soapfish). In: Carpenter, K. E. & De Angelis, N. (Eds.), The Living Marine Resources of the Eastern Central Atlantic. 4. Bony Fishes. Part 2 (Perciformes to Tetraodontiformes) and sea turtles. FAO species Identification Guide for Fishery Purposes. FAO, Rome, pp. 2365 - 2413."]}

Sithole, Yonela, Heemstra, Elaine, Mwale, Monica (2021): Revalidation and redescription of Serranus knysnaensis Gilchrist, 1904 (Perciformes: Serranidae) with a new distribution record. Zootaxa 5057 (1): 99-113, DOI: https://doi.org/10.11646/zootaxa.5057.1.6

Genus Serranus Cuvier, 1816 Type species Perca cabrilla Linnaeus, 1758 Diagnosis. Dorsal-fin X, 12���15; anal-fin rays III, 7; pectoral-fin rays 12���17; pelvic-fin rays I, 5, pectoral fin above, or slightly in front of, pelvic fin; gill rakers 6���12+12���23 (18���35 total); circumpeduncular scales 26���38; lateral-line scales 67���77; scales from dorsal fin to lateral line 9���12; scales from anal-fin origin to lateral line 20���26. Caudal fin truncate to emarginate or moderately forked, 17 principal rays with 15 branched; scales ctenoid, some species with cycloid scales on head or trunk; supramaxilla absent; maxilla, snout and chin naked; teeth on dentary, premaxillary, vomer and palatines; mouth moderately protractile; opercle with usually three flat spines (two opercular spines in S. drewesi, Iwamoto & Wirtz, 2018), central spine largest and directed horizontally; preopercle serrate; lateral line complete with pored scales. Branchiostegal rays 7. Vertebrae 10 + 14 = 24., Published as part of Sithole, Yonela, Heemstra, Elaine & Mwale, Monica, 2021, Revalidation and redescription of Serranus knysnaensis Gilchrist, 1904 (Perciformes: Serranidae) with a new distribution record, pp. 99-113 in Zootaxa 5057 (1) on page 100, DOI: 10.11646/zootaxa.5057.1.6, http://zenodo.org/record/5705482, {"references":["Cuvier, G. (1816) Le Regne Animal distribue d'apres son organisation pour servir de base a l'histoire naturelle des animaux et d'introduction a l'anatomie comparee. Les reptiles, les poissons, les mollusques et les annelides, 2, i - xviii + 1 - 532.","Linnaeus, C. (1758) Systema Naturae. 10 th Edition. Impensis Direct. Laurentii Salvii, Holmiae, ii + 824 pp.","Iwamoto, T. & Wirtz, P. (2018) A synopsis of the Eastern and Central Atlantic combers of the genus Serranus (Teleostei: Perciformes: Serranidae). Proceedings of the California Academy of Sciences, 65, 1 - 39."]}

Serranus knysnaensis Gilchrist, 1904 English name: African seabass Figures 1���3; Tables 1���5 Serranus knysnaensis Gilchrist 1904: 2, Pl. 19 (Knysna, Cape Province, South Africa); Smith 1949: 193, Fig. 427 (South Africa); Heemstra & Heemstra 2004: 174 (unnumb. painting; South Africa); King & Fraser 2014: (unnumb. photograph p. 69; endemic, from False Bay in Western Cape to northern KwaZulu-Natal). Serranus cabrilla (in part): Smith & Smith 1966: 88 (painting 429; Atlantic species from Mediterranean southwards, colonising South Africa to Natal); Heemstra & Randall 1986: 536 (Fig. 166.76, Pl. 45; Mediterranean, eastern Atlantic, English Channel to South Africa, and round coast to Durban) Diagnosis. Dorsal-fin X, 14���15; pectoral-fin rays 15���16; gill rakers 6���8+12���15 (18���22 total); circumpeduncular scales 26���34; lateral-line scales 67���76; scales from dorsal fin to lateral line 9���12; scales from the anal-fin origin to lateral line 22���26. Caudal fin emarginate. Scales small, ctenoid and deciduous. Body elongate, not much compressed; body depth less than head length, about 3.3���4.2 in SL; head length 2.5��� 2.9 in SL. Maxilla extending to below middle of the eye; jaws subequal; lower jaw projecting; opercle with three flat spines. Eye diameter greater than interorbital length and less than snout length, eye diameter contained 9.6���13.1 in SL. Snout, interorbital and maxilla naked. Pectoral fin originates before dorsal fin and slightly before the pelvic fin. Distal tip of pectoral fin before anal-fin origin. Description. Proportional measurements are given in Table 3. Body elongate; dorsal fin continuous, with spinous membrane moderately incised between spines, becoming less incised posteriorly; dorsal-fin spines graduated, but with first two spines noticeably shorter, and ninth and tenth spines subequal; soft-rayed portion of fin higher than spinous part. First anal-fin spine shortest; second anal-fin spine thicker than others. Anterior lateral-line tubes with ascending branch. Peduncle longer than deep. Patch of small teeth on vomer and palate. Preopercle margin serrate with more developed spines near the angle. Snout and interorbital naked; preopercle, opercle, operculum and postorbital scaly; chest and belly scaly; snout to eye naked. Lateral line complete, arching slightly from upper end of gill cavity and following body outline to caudal peduncle where it becomes horizontal; lateral line with pores. Fresh color (Fig. 1a). Head and body light (pale) to yellowish brown dorsally, pale pinkish to white ventrally. Head with three yellow to bronze stripes, two below and one behind eye, the lowest stripe from just behind maxilla to preopercle, the second stripe from below anterior of eye to subopercle and occasionally, the pectoral-fin base, the upper stripe from rear ventral part of eye to opercular margin near upper pectoral-fin base; posterior bony ridge of the orbit yellow. Body with two irregular usually darker brown to yellowish-brown longitudinal bands of subequal width (~2/3 eye diameter), or with lower band slightly narrower, the upper band from either side joining on premaxilla and running from snout tip through eye to upper caudal-fin base, with narrow white border (Color in preservation (Fig. 1b). Specimens often lose color almost completely in preservation. Two irregular dark, longitudinal bands from head to caudal-fin base, and dark stripes below and behind the eye visible in some specimens. Holotype uniformly dark brown, no visible pattern. Distribution. Serranus knysnaensis occurs in the southwestern Indian Ocean from South Africa (Knysna, Western Cape, rarely southwards) and northwards to Bazaruto, southern Mozambique. New records from north of Durban (Fig. 2a, Umhlanga; SAIAB 193652, uThukela [formerly Tugela] Banks; Fig. 2b, Sodwana Bay by C. van Jaarsveld sourced from Fishwise Professional: https://www.fishwisepro.com/pictures/default?Sid=34346&Info=%27 Serranus %20knysnaensis%27 As well as from Mozambique and Madagascar (see Material examined, also Figs. 2c and 2d)), are reported in this study. Species delimitation with DNA barcoding. The pairwise genetic distances (Table 4) of the COI mtDNA sequences for all Serranus species under the K2P model ranged from 1.60% to 25.30%. The pairwise genetic distance (Table 4) between COI mtDNA sequences for S. knysnaensis from South Africa and S. cabrilla from the Atlantic Ocean and Mediterranean Sea was low at 1.60%. Although, this was among the lowest genetic distances between the analysed Serranus species, it was comparable to that between two distinct species, S. novemcinctus to S. cabrilla (1.61%), and lower than the genetic distance between S. knysnaensis and S. novemcinctus (1.99%). These divergence estimates were also comparable to the COI sequence divergence reported between other serranid species (e.g Victor 2012; Vella & Vella 2016; Iswarya et al. 2018) and other marine fish species (e.g. Uiblein & Gouws 2014). Furthermore, K2P genetic distance more than 1% for the COI gene is considered as a threshold of species delimitation for marine fishes (Avise 2000; Zhang & Hanner 2011). The maximum likelihood tree (Fig. 3) based on 92 COI mtDNA sequences of 18 species including the outgroup (Cephalopholis nigripinnis) recovered monophyletic clusters corresponding to all species including the WIO species (S. knysnaensis and S. novemcinctus). All these three clades were strongly supported as monophyletic groups with bootstrap>75%. The S. knysnaensis and S. cabrilla clusters (Table 5) had the highest Intra/Inter ratios (0.28 and 0.38 respectively), suggesting that the divergence within these lineages was high relative to the divergence with their closest species. The small values (S. knysnaensis. Remarks: Serranus knysnaensis can be distinguished from the other Serranus species in the SWIO, S. novemcinctus, by having 12���15 lower limb gill rakers (versus 21���23) and 26���34 circumpeduncular scales (versus 34���36) (Table 2). Serranus knysnaensis also differs from S. novemcinctus by two dark prominent longitudinal bands and is endemic to southern Africa while S. novemcinctus has nine dark vertical bars dorsally on the body and it is distributed in Amsterdam and St. Paul Islands, and Madagascar. Serranus knysnaensis differs from S. cabrilla by having 26���34 circumpeduncular scales (versus 34���38) and 18���22 total gill rakers (versus 22���24) (Table 2). The color pattern of S. knysnaensis is very similar to that of S. cabrilla, however, S. knysnaensis can be distinguished by two dark prominent longitudinal bands, whereas S. cabrilla has two or three longitudinal bands often partially broken into dark blotches. The two species also have different distribution range; S. knysnaensis is a southern African endemic from Knysna to Bazaruto, southern Mozambique and Madagascar, with rare waifs reported south of Knysna to False Bay. Serranus cabrilla is known from the British Isles to Angola, including Azores, Madeira, Canary Islands, S��o Tom�� and Pr��ncipe, and Cape Verde Islands, also through Mediterranean and Black Sea, and in the Red Sea (Heemstra & Heemstra 2004; Heemstra & Anderson 2016; Iwamoto & Wirtz 2018). Heemstra & Heemstra (2004) described S. knysnaensis as a mainly rocky bottom species occurring in depths of 1���200 m, it is also often found at the mouth of estuaries (Smith & Smith 1966). Whereas S. cabrilla is reported over rocky as well as soft bottom, from shore to 450 m (Heemstra & Anderson 2016; Iwamoto & Wirtz 2018). Furthermore, the S. knysnaensis clade differs from S. novemcinctus and S. cabrilla by 1.99 to 1.60% genetic distance, respectively. Based on differences in the morphological characters, distribution and genetic data, S. knysnaensis is a valid species distinct from S. cabrilla and South African records of S. cabrilla are misidentifications of S. knysnaensis. Available genetic data, from the two WIO species (S. knysnaensis and S. novemcinctus) shows that they are closely related to each other, as well as S. cabrilla. The low genetic diversity (approx. 1.60%) was significant, suggesting that these lineages could represent ecomorphs or lineages that have only recently diverged (speciated) in this genus., Published as part of Sithole, Yonela, Heemstra, Elaine & Mwale, Monica, 2021, Revalidation and redescription of Serranus knysnaensis Gilchrist, 1904 (Perciformes: Serranidae) with a new distribution record, pp. 99-113 in Zootaxa 5057 (1) on pages 102-105, DOI: 10.11646/zootaxa.5057.1.6, http://zenodo.org/record/5705482, {"references":["Gilchrist, J. D. F. (1904) Description of new South African fishes. Marine investigation in South Africa, 3, 1 - 16.","Smith, J. L. B. (1949) The Sea fishes of Southern Africa. Central News Agency, Johannesburg, 550 pp.","Heemstra, P. C. & Heemstra, E. (2004) Coastal Fishes of Southern Africa. SAIAB and NISC (Pty) Ltd, Grahamstown, xxiv + 488 pp.","King, D. & Fraser, V. (2014) The Reef Guide: Fishes, Corals, Nudibranchs and other Vertebrates of East and South Coasts of Southern Africa. Struik Nature, Cape Town, 360 pp.","Smith, J. L. B. & Smith, M. M. (1966) Fishes of the Tsitsikama Coastal National Park. Swan Press Ltd, Johannesburg, 161 pp.","Heemstra, P. C. & Randall, J. E. (1986) Family Serranidae. In: Smith, M. M. & Heemstra, P. C. (eds), Smiths' Sea Fishes. MacMillan, Johannesburg, pp. 509 - 537.","Victor, B. C. (2012) Hypoplectrus floridae n. sp. and Hypoplectrus ecosur n. sp., two new Barred Hamlets from the Gulf of Mexico (Pisces: Serranidae): more than 3 % different in COI mtDNA sequence from the Caribbean Hypoplectrus species flock. Journal of the Ocean Science Foundation, 5, 1 - 19.","Vella, A. & Vella, N. (2016) Genetic barcoding and preliminary phylogenetic analysis of Serranidae species from Maltese coastal waters, with a perspective on their Mediterranean phylogeography. Natural and Engineering Sciences, 1, 66 - 77. https: // doi. org / 10.28978 / nesciences. 286369","Iswarya, D. V., Kandula, S. & Khedkar, G. D. (2018) DNA barcoding of five species of groupers (Pisces: Serranidae) off Visakhapatnam, central eastern coast of India. Mitochondrial DNA Part A, 29, 659 - 663. https: // doi. org / 10.1080 / 24701394.2017.1339188","Uiblein, F. & Gouws, G. (2014) A new goatfish species of the genus Upeneus (Mullidae) based on molecular and morphological screening and subsequent taxonomic analysis. Marine Biology Research, 10, 655 - 681. https: // doi. org / 10.1080 / 17451000.2013.850515","Avise, J. C. (2000) Phylogeography: the History and Formation of Species. Harvard University Press, Cambridge, 447 pp. https: // doi. org / 10.2307 / j. ctv 1 nzfgj 7","Zhang, J. & Hanner, R. (2011) DNA barcoding is a useful tool for the identification of marine fishes from Japan. Biochemical Systematics and Ecology, 39, 31 - 42. https: // doi. org / 10.1016 / j. bse. 2010.12.017","Heemstra, P. C. & Anderson, W. D. (2016) Serranidae. Groupers (seabasses, hinds, creolefish, combers, anthiines, soapfish). In: Carpenter, K. E. & De Angelis, N. (Eds.), The Living Marine Resources of the Eastern Central Atlantic. 4. Bony Fishes. Part 2 (Perciformes to Tetraodontiformes) and sea turtles. FAO species Identification Guide for Fishery Purposes. FAO, Rome, pp. 2365 - 2413.","Iwamoto, T. & Wirtz, P. (2018) A synopsis of the Eastern and Central Atlantic combers of the genus Serranus (Teleostei: Perciformes: Serranidae). Proceedings of the California Academy of Sciences, 65, 1 - 39."]}

Serranus novemcintus Kner, 1864 English name: Barred seabass Figures 3, 5; Tables 1���5 Serranus novemcinctus Kner 1864: 483, Pl. 2, Fig. 1 (St. Paul Island and Cape of Good Hope, southern Indian Ocean); Heemstra & Randall 1986: 536 (Fig. 166.77; Amsterdam and St. Paul Islands). Diagnosis. Dorsal-fin X, 14; pectoral-fin rays 16���17; gill rakers 9���12+21���24 (31���35 total); circumpeduncular scales 34���36; lateral-line scales 67���74; scales from dorsal fin to lateral line 9���10; scales from the anal-fin origin to lateral line 20���23. Caudal fin emarginated; scales ctenoid; maxilla naked; teeth on vomer and palatines; opercle with three flat spines, central spine largest and directed horizontally; cheek, pectoral-fin base and chest scaly. Description. Body elongate; body depth less than head length, about 3.2���4.9 in SL; head length 2.4���3.6 in SL. Maxilla extending to below middle of the eye; lower jaw projecting beyond upper jaw; opercle with three flat spines, the upper two spines well developed. Scales ctenoid; small scales at the bases of dorsal, anal and pectoral fins. Preopercle margin coarsely serrate, developed spines near angle. The first two dorsal-fin spines shortest. No distinct notch between dorsal spines and soft rays but soft rays slightly elevated. Color (Fig. 5). Brownish with nine dark vertical bars dorsally on body, the first four below the spinous dorsal fin; the next three under the soft dorsal rays, the eighth on the caudal peduncle and the last at the caudal-fin base; diagonal bar on the preopercle, originating behind the eye to margin of opercle at mid-pectoral fin position. Ventral body white. No spots on the fins but dorsal fin membranes may have some irregular bluish lines on basal third of fin membrane. Color in preservation. Specimens lose color in preservation but the pattern, i.e. nine vertical bars on body, and diagonal bar on the preopercle, remains visible. Distribution. Amsterdam and St. Paul Islands, and at Walters Shoals south of Madagascar (Heemstra & Randall 1986; Collette & Parin 1991)., Published as part of Sithole, Yonela, Heemstra, Elaine & Mwale, Monica, 2021, Revalidation and redescription of Serranus knysnaensis Gilchrist, 1904 (Perciformes: Serranidae) with a new distribution record, pp. 99-113 in Zootaxa 5057 (1) on pages 108-109, DOI: 10.11646/zootaxa.5057.1.6, http://zenodo.org/record/5705482, {"references":["Kner, R. (1864) Specielles Verzeichniss der wahrend der Reise der kaiserlichen Fregatte \" Novara \" gesammelten Fische. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften. Mathematisch-Naturwissenschaftliche Classe, 49, 481 - 486.","Heemstra, P. C. & Randall, J. E. (1986) Family Serranidae. In: Smith, M. M. & Heemstra, P. C. (eds), Smiths' Sea Fishes. MacMillan, Johannesburg, pp. 509 - 537.","Collette, B. B. & Parin, N. V. (1991) Shallow-water fishes of Walters Shoals, Madagascar ridge. Bulletin of Marine Science, 48, 1 - 22."]}

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