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Genus Upeneus Cuvier 1829 Diagnosis. Dorsal fins VII or VIII + 9; pectoral fins 12���17; principal caudal-fin rays 7 + 8 (median 13 branched); gill rakers 4���9 + 13���24 = 18���33; lateral-line scales 28���39; lateral line complete; small scales present basally on second dorsal, anal and/or caudal fins; small teeth present on vomer and palatines; teeth on jaws multiserial and villiform; body oblong, slightly compressed or nearly rounded; barbels 15���27% SL; snout length 9.1���14% SL, subequal to or slightly shorter than postorbital length (9.7���15% SL); body frequently with one to several longitudinal stripes, varying from yellow to red, brown or black in life; oblique bars often present on caudal-fin lobes, usually retained in preservative when darkly pigmented, and their number, colour and form of diagnostic importance. Remarks. Forty-six species, including the three new species, distributed in all major oceans. Two Indo-Pacific species in the Mediterranean as Lessepsian migrants. Many species targeted by fisheries or landed as bycatch and can be found in fish markets, having moderate to high local economic value. Many different fishing methods are used including angling, beach seine, gillnetting, trapping and trawling. Most species occur in coastal waters to less than 100 m depth, maximum depth reached is 600 m. Maximum size range among species is about 89 to 300 mm SL., Published as part of Uiblein, Franz & Motomura, Hiroyuki, 2021, Three new goatfishes of the genus Upeneus from the Eastern Indian Ocean and Western Pacific, with an updated taxonomic account for U. itoui (Mullidae: japonicus-species group), pp. 298-324 in Zootaxa 4938 (3) on page 302, DOI: 10.11646/zootaxa.4938.3.2, http://zenodo.org/record/4569653

Upeneus elongatus n. sp. Elongate Goatfish Iwanaga-himeji (new standard Japanese name) (Figures 1B, C, 2; Tables 2, 3) Upeneus sp.: Tashiro et al. 2014 Material examined. Holotype. KAUM���I. 58746, 89 mm SL, Philippine Sea, S Japan, Kagoshima Prefecture, Osumi Islands, Tanega-shima Island, Kumage-gun, Nakatane, off Kumano, 30��28.13��� N, 130��58.32��� E, 25 m, set net, Mayumi Takayama, 20-01-2012 (photo of fresh holotype, Figure 1B, C) Unvouchered In situ photos of Upeneus cf. elongatus, Central Philippines, Visayas (most probably all adults): Panay, Caridad, Culasi, Antique, 11��26.89��� N, 122��03.55��� E, 7 m depth (Figure 1G); Leyte, Albuera, 10��53.45��� N, 124��42.36��� E, 20 m depth (Figure 1H); Negros, Bonbonon, Siaton, 9��03.53��� N, 123��05.97��� E, 10 m depth (Figure 1I); same locality, 7 m depth (Figure 1J). Diagnosis. Dorsal fins VII + 9; pectoral fins 13; gill rakers 8 + 19���20 = 27���28; body elongate and laterally compressed; measurements as % SL: depth at first dorsal-fin origin 21; body depth at anal-fin origin 18; caudalpeduncle depth 8.3; caudal-peduncle width 3.3; maximum head depth 18; head depth through eye 14; head length 27; snout length 10; orbit length 6.7; upper-jaw length 11; barbel length 17; caudal-fin length 29; anal-fin height 16; pelvic-fin length 23; pectoral-fin length 21; first dorsal-fin height 20, dorsal-fin spines proportionally decreasing in height; second dorsal-fin height 15; total oblique bars on caudal fin 12, upper caudal-fin lobe with 5 red bars; lower caudal-fin lobe with 7 red-brown blotch-like bars which are most conspicuous along middle of lobe and dorsally with a faint yellowish-beige stripe of less than orbit diameter in width that connects to caudal-fin base; bars mostly retained after preservation; first dorsal fin with large yellowish-beige triangular patch close to fin base covering central part of ventral half of fin; a similarly coloured, but much smaller patch at middle of second dorsal-fin base; barbels white when fresh; body and head colour in freshly deceased fish pale greyish dorsally and mostly red below eye level except for pale-grey snout region; ventral margin of head and body silvery white; a vermillion mid-lateral body stripe from snout tip through eye to middle of caudal-fin base; body and head pale brown, mid-body darkened when preserved. Description. Measurements in % SL and counts are given in Table 3; morphometric data as ratios of SL for holotype: body elongate and laterally compressed, body depth at first dorsal-fin origin 4.9; body depth at anal-fin origin 5.5; head depth through eye 6.9; head length 3.7, much larger than maximum body depth and subequal to caudal-fin length (3.5); first dorsal-fin height 4.9, larger than barbel length (6.0) and second dorsal-fin height (6.6); pelvic-fin length 4.4, larger than pectoral-fin length (4.8) and body depth at first dorsal-fin origin; caudal-peduncle depth 12, distinctly larger than orbit length (15); and caudal-peduncle width 31, much smaller than pectoral-fin width (23). Colour. Fresh HT (Figure 1B, C). Body with all scales missing, pale greyish dorsally above eye level with tiny dark-brown dots scattered along dorsal margin from below second dorsal fin to caudal-fin base; also, with a few tiny, similar-coloured dots further ventrally between first dorsal-fin origin and middle of second dorsal fin base; broad vermillion stripe of pupil diameter from behind eye to middle of caudal-fin base a; series of about 20 oblique red bands separated by thin silvery lines ventrally of stripe behind pectoral-fin base; few red blotches along ventral margin of posterior half of body partly overlapping with the oblique bands and white-silvery ventral side of body; head pale grey anterior to eye, with a weak red stripe from snout to eye of about half pupil width, stripe continues behind eye as mid-lateral body stripe; cheek below eye red and opercle region silvery white with red blotches; iris red; barbels white; caudal fin with five oblique, mostly straight brown bars of about half pupil width on upper lobe, with white hyaline interspaces between bars increasing in width from being narrower than bars proximally to more than double-bar width distally; tip of upper caudal-fin lobe white hyaline; lower caudal-fin lobe with seven redbrown blotch-like bars which are most conspicuous along middle of lobe becoming mostly hyaline towards lobe margins, with narrow hyaline-white interspaces; the two proximal bars only very weakly indicated by fine streaks on two ventral-most fin rays fusing dorsally into a single dark brown bar, followed distally by a weakly indicated red spot composed by three short streaks on two adjacent fin rays along middle of lobe; four more conspicuous red spots further distally, two as wide as pupil, positioned mainly along middle of fin lobe and tip; dorsal part of lobe with a yellowish-beige stripe of less than orbit diameter, extending from central caudal-fin base to tip of lower caudal-fin lobe, covering nearly entire lobe distally and surrounding the two distal-most red blotches; first dorsal fin with a triangle-shaped patch of yellowish-beige skin, of about snout length in its maximum extension, patch connecting fin base and second to sixth spine, covering central part of ventral half of fin, except for the first spine; first dorsalfin spine with four or five weakly indicated brown dots, spine tip unpigmented; second to third spine with brown pigmentation close to tip (second spine however broken), spines otherwise pale; second dorsal fin with four weakly indicated pale brown stripes and a yellowish-beige patch, similarly coloured as patch on first dorsal fin but much smaller, at middle of fin base between fourth and fifth spine; pectoral, pelvic and anal fins hyaline whitish, pelvic fins pale yellowish white at base. Preserved HT. Head and body mostly pale brown, head in front of eyes, belly and underside of body pale beige; a brownish mid-lateral band wider than orbit diameter along body from behind opercle to caudal-fin base, becoming thinner behind second dorsal-fin base and then widening again close to caudal-fin base; tiny scattered dark-brown dots along dorsal body margin from below second dorsal fin to caudal-fin base and, just above dark mid-body band between dorsal fins; lower half of opercle silvery white; bar pigmentation on caudal fin mostly retained; stripe on lower caudal-fin lobe lost as well as most of dorsal-fin pigmentation except for a few traces distally on first dorsal fin and at mid and further distally on second dorsal fin; other fins and barbels pale. Etymology. The name ��� elongatus ��� refers to the elongate, very shallow body and head of this species. The standard Japanese name Iwanaga-himeji has two meanings: (1) ���elongate goatfish���; (2) ���goatfish of goddess Iwanaga���, sister of the myth goddess Sakuya, whose grandchild is considered to be the first Emperor of Japan. Accordingly, the standard Japanese name for the rather similar U. itoui is Sakuya-himeji. Distribution, habitat, size. Known only from a single locality in S Japan, Tangea-shima Island, Kagoshima Prefecture; occurs on shallow sandy bottoms (25 m depth); attains at least 89 mm SL. This species may also occur in the central Philippines (see also Remarks below). Remarks. Upeneus elongatus most likely also occurs in the Philippines as indicated by unvouchered in situ photos of shallow-bodied goatfish specimens with very similar colour patterns on caudal and dorsal fins (Figure 1 G���H). However, the body colour in those specimens differs considerably from the fresh HT from S Japan, which may be due to the absence of scales in the HT. Until preserved specimens with scales intact associated with colour photographs of fish in fresh condition become available for comparative studies, we refer to this form as U. cf. elongatus., Published as part of Uiblein, Franz & Motomura, Hiroyuki, 2021, Three new goatfishes of the genus Upeneus from the Eastern Indian Ocean and Western Pacific, with an updated taxonomic account for U. itoui (Mullidae: japonicus-species group), pp. 298-324 in Zootaxa 4938 (3) on pages 304-310, DOI: 10.11646/zootaxa.4938.3.2, http://zenodo.org/record/4569653, {"references":["Tashiro, S., Takayama, M. & Motomura, H. (2014) Distributional range extension of Upeneus itoui (Perciformes; Mullidae) in East Asia, with the first records from Tanega-shima Island in the Osumi Islands, Kagoshima, Japan, and notes on an unidentified specimen of Upeneus from Tanega-shima Island. Nature of Kagoshima, 40, 69 - 74."]}

Upeneus dimipavlov n. sp. Pavlov���s Goatfish (Figures 1A, 2���6; Tables 2, 3) Upeneus guttatus Day, 1868, in part: Uiblein et al. 2017a Material examined. Holotype. VNMN-I 2056 (formerly HIFIRE F 58334), 120 mm SL, South China Sea, S-central Vietnam, Khan Hoa Province, Nha Trang, Bai Tre Bay, N of Hon Tre Island, 12��13.35��� N, 109��18.50��� E, obtained from hookah divers, ca. 15 m depth, Dimitri A. Pavlov, 19-01-2012 (photo of fresh HT, Figure 1A). Paratypes (n = 10, 111��� 134 mm SL). South China Sea, S-central Vietnam, Khan Hoa Province, Nha Trang, Xom Moi Market: HIFIRE F 58183, 129 mm SL; HIFIRE F 58184, 117 mm SL; NHMO J7223 (formerly HIFIRE F 58185), 124 mm SL; MNHN 2020-0474 (formerly HIFIRE F 58186), 122 mm SL; NMMBP 34705 (formerly HIFIRE F 58187), 111 mm SL; KAUM���I. 146834 (formerly HIFIRE F 58188), 123 mm SL; AMS I.49516-001 (formerly HIFIRE F 58189), 122 mm SL; CAS-ICH 247243 (formerly HIFIRE F 58190), 122 mm SL; Nha Trang fish market: VNMN-I 2057 (formerly HIFIRE F 58174), 134 mm SL; VNMN-I 2058 (formerly HIFIRE F 58177), 128 mm SL. Diagnosis. Dorsal fins VII + 9; pectoral fins 14���15; gill rakers 6���7 + 16���18 = 22���25; body nearly rounded and moderately elongate; measurements as % SL: body depth at first dorsal-fin origin 23���25; body depth at anal-fin origin 20���22; caudal-peduncle depth 9.7���10; caudal-peduncle width 4.8���5.4; maximum head depth 19���20; head depth through eye 16���17; head length 28���30; snout length 11���13; orbit length 5.7���6.8; upper-jaw length 11���12; barbel length 19���21; caudal-fin length 27���29; anal-fin height 16���18; pelvic-fin length 20���22; pectoral-fin length 18���20; first dorsal-fin height 19���21; dorsal-fin spines proportionally decreasing in height; second dorsal-fin height 16���18; total oblique bars on caudal fin 12 ��� 14, upper caudal-fin lobe with 5���6 red mostly straight bars; lower caudal-fin lobe with 7���8 short red bars along ventral margin which connect dorsally to a red stripe, the latter slightly wider than orbit diameter, and with about 3 short red or reddish-grey bars along dorsal margin of fin lobe that do not correspond with bars on ventral-lobe margin; caudal-fin colour patterns partly or completely lost after preservation; dorsal fins in fresh fish with 3���4 thin red or brown stripes, may be weakly retained in preservative; barbels rose whitish or white when fresh; body colour in freshly deceased specimens dorsally pale reddish orange, ventral half creamy white, sometimes with red patches or blotches, ventral margin pale creamy white; no conspicuous mid-lateral body stripe; head dorsally pale red and silvery white ventrally, with red patches below and behind eyes; body and head dorsally pale brown or brown and ventrally white when preserved. Description. Measurements in % SL and counts are given in Table 3; morphometric data as ratios of SL for holotype, data for paratypes in brackets: body nearly rounded, rather robust and moderately elongate, body depth at first dorsal-fin origin 4.2 [4.1���4.4]; body depth at anal-fin origin 4.9 [4.6���5.0]; head depth through eye 6.3 [6.0��� 6.4]; head length 3.4 [3.4���3.5], larger than maximum body depth and subequal to caudal-fin length (3.6 [3.4���3.8]); first dorsal-fin height 5.0 [4.8���5.4], subequal to barbel length (5.0 [4.7���5.4]) and larger than second dorsal-fin height (6.2 [5.7���6.2]); pelvic-fin length 4.9 [4.5���5.0], larger than pectoral-fin length (5.1 [5.1���5.5]) and subequal to body depth at anal-fin origin; caudal-peduncle depth 10 [9.8���10], clearly larger than orbit length (15 [15���17]); and caudal-peduncle width 19 [19���21], mostly larger than pectoral-fin width (22 [21���23]). Colour. Fresh HT (Figure 1A). Body pale red orange above lateral line and creamy white below, ventral side pale creamy white; several red sports and blotches from pelvic-fin base and behind opercle at eye level to level of second dorsal-fin origin; lateral line indistinct, no mid-lateral body stripe; dorsal margin of head from snout tip to above eye pale red; silvery-pale beige mouth region followed by a large bright-red patch below eye and another one posteriorly on bony opercle with three bright-red blotches just above; iris bright red; barbels whitish rose; caudal fin with five oblique, mostly straight red bars of approximately half pupil width on upper lobe, with white hyaline interspaces between bars increasing in width from being narrower than bar width proximally to double-bar width distally; tip of upper caudal-fin lobe white hyaline; lower caudal-fin lobe with seven very short red bars along ventral margin covering up to four ventral-most fin rays; creamy-hyaline interspaces between bars of less than bar width proximally with increasing width distally, continued as a pale creamy stripe behind distal-most seventh bar, not reaching lobe tip; broad red stripe along middle of lower caudal-fin lobe slightly wider than orbit diameter, stripe extending from caudal-fin base to lower lobe tip; four red bars with dark-grey tips dorsally on lower caudal-fin lobe along lobe margin being slightly wider than bars on ventral side of lobe and with white-hyaline interspaces of similar width as bars, the interspaces connecting to red stripe; first dorsal fin with hyaline-red spots on and around spines forming a dotted pattern of four horizontal stripes; first dorsal-fin tip weakly pale-whitish pigmented; second dorsal fin whitish hyaline with four red horizontal stripes, one at fin base and one at fin tip; pectoral, pelvic and anal fins hyaline creamy; pelvic fins with pale-red pigmentation on spine tip and anterior four rays. Preserved HT. Dark brown dorsally from mouth and below eye to pectoral-fin origin and distal end of lateral line near caudal-fin base; creamy white ventrally from cheeks and below pelvic-fin origin to ventral half of caudalfin base; caudal-fin bars and stripe and dorsal-fin pigmentation only faintly retained; other fins and barbels pale. Etymology. The name ��� dimipavlov ��� is used as a composed noun in apposition. It honours the ichthyologist Dr. Dimitri Alexandrovich ���Dimi��� Pavlov, Lomonosov University, Moscow, Russia, for collecting, photographing and donating mullid specimens from Vietnam (including the holotype of Upeneus dimipavlov) for taxonomic research. Distribution, habitat, size. Currently only known from Nha Trang, S-central Vietnam; possibly occurs on sandy bottoms near reefs; depth ca. 15 m; attains 134 mm SL. Remarks. The nearly rounded, robust body is a typical characteristic of this species and rather unique among japonicus -group species. The photograph of the HT (Figure 1A) is the only high-quality colour documentation of fresh fish available so far. In photos of several PT���s made during or after collection at fish markets the body colour is less intense and in two specimens barbel colour is white instead of whitish rose. After specimens were kept in a refrigerator over several days or thawed, the body and head became almost entirely red which is most likely an artefact and does not reflect natural colouration. This species has been encountered at fish markets near the type locality, indicating local commercial value., Published as part of Uiblein, Franz & Motomura, Hiroyuki, 2021, Three new goatfishes of the genus Upeneus from the Eastern Indian Ocean and Western Pacific, with an updated taxonomic account for U. itoui (Mullidae: japonicus-species group), pp. 298-324 in Zootaxa 4938 (3) on pages 302-304, DOI: 10.11646/zootaxa.4938.3.2, http://zenodo.org/record/4569653, {"references":["Uiblein, F., Gledhill, D. C. & Peristiwady T. (2017 a) Two new goatfishes of the genus Upeneus (Mullidae) from Australia and Indonesia. Zootaxa, 4318 (2), 295 - 311. https: // doi. org / 10.11646 / zootaxa. 4318.2.4"]}

Upeneus willwhite n. sp. White���s Goatfish (Figures 1F, 2���6; Tables 2, 3) Upeneus guttatus Day, 1868: White et al. 2013; Uiblein et al. 2017a (in part) Material examined. Holotype. CSIRO 7217-07, 90 mm SL, EIO, S Indonesia, Lombok, Tanjung Luar, fish market, William White (photo of fresh HT, Figure 1F) Paratype. MZB 23014, 92 mm SL, same sampling data as holotype. Diagnosis. Dorsal fins VII + 9; pectoral fins 14; gill rakers 6 + 15���17 = 21���23; body moderately elongate and laterally slightly compressed; measurements as % SL: body depth at first dorsal-fin origin 23���24; body depth at anal-fin origin 20���21; caudal-peduncle depth 9.6���9.9, caudal-peduncle width 4.2���4.3; maximum head depth 20���21; head depth through eye 16; head length 29; snout length 10���11; orbit length 7.1���7.5; upper-jaw length 12; barbel length 20; caudal-fin length 28���30; anal-fin height 16���17; pelvic-fin length 20���21; pectoral-fin length 19���20; first dorsal-fin height 20, dorsal-fin spines proportionally decreasing in height; second dorsal-fin height 17; total oblique bars on caudal fin 11 ��� 14, upper caudal-fin lobe with 5���6 brown mostly straight bars; lower caudal-fin lobe with 6���8 short brown bars ventrally, connecting dorsally to a brown stripe of about orbit diameter in its maximum width, and with about 3 short red, brownish-ochre bars along dorsal margin that do not correspond with bars on ventral-lobe margin; caudal-fin stripe and bars retained after preservation; dorsal fins in fresh fish with 3���4 thin brown stripes, retained in preservative; barbels reddish white when fresh; body colour in freshly deceased fish dark brown dorsally and pale brown ventrally, ventral margin white; mid-lateral body stripe dark brown, rather inconspicuous on snout and posterior half of body; head entirely grey brown; body and head dark brown dorsally and pale brown ventrally when preserved. Description. Measurements in % SL and counts are given in Table 3; morphometric data as ratios of SL for holotype, data for paratype in brackets: body moderately elongate and laterally slightly compressed, body depth at first dorsal-fin origin 4.3 [4.3]; body depth at anal-fin origin 4.9 [4.9]; head depth through eye 5.1 [4.8]; head length 3.4 [3.5], larger than maximum body depth and subequal to caudal-fin length (3.6 [3.4]); first dorsal-fin height 5.1 [4.9], subequal to barbel length (5.1 [5.1]) and larger than second dorsal-fin height (6.0 [5.8]); pelvic-fin length 5.0 [4.7], slightly larger than pectoral-fin length (5.2 [4.9]) and subequal to body depth at anal-fin origin; caudal-peduncle depth 10 [10], distinctly larger than orbit length (14 [13]); and caudal-peduncle width 23 [24], subequal to pectoral-fin width (22 [23]). Colour. Fresh HT. Dorsal part of head and body grey brown from snout to above eye and above lateral line; dorsal body colour appearing marbled due to grey scales with dark grey-brown edges; dark grey-brown band of about pupil diameter in width along entire dorsal body margin, becoming more conspicuous behind first dorsal fin and narrows behind second dorsal fin to a dark grey, partly blackish stripe; a similarly coloured dark grey, partly blackish mid-lateral body stripe of pupil diameter in width from behind eye to caudal-fin base crossing the faint white-greyish lateral line below second dorsal fin; below of mid-lateral body stripe scales pale grey or white with brown-greenish edges forming four oblique bands that become weaker and more ochre-coloured distally, before ending on lower part of caudal-fin base or above posterioventral half of body; a fifth faint-yellowish band from below pectoral-fin base to above anal-fin origin; head below eye from behind snout pale grey with grey-brown dots or streaks and two faint-reddish blotches on ventral margin below and behind eye; iris red to rose whitish; ventral part of head and body white creamy; caudal fin with five oblique brown-ochre bars of about half pupil width on upper lobe, the most anterior bar at lobe base bent, the remaining bars more straight; white-hyaline interspaces between bars increasing in width from being narrower than bars proximally to double-bar width distally; tip of upper caudal-fin lobe white hyaline; lower caudal-fin lobe with eight short brown bars, along ventral margin of lobe brown streaks on ventral-most fin rays; white-hyaline interspaces between bars increasing distally to more than double-bar width; dark brown stripe along middle of lower caudal-fin lobe of about orbit diameter in its maximum width, stripe originating from a large, vertically extended dark-brown patch on caudal-fin base, connecting there with mid-lateral body stripe; lower caudal-fin lobe stripe changes from dark brown to brown red distally, ending as a dark-brown lobe tip; dorsal part of lower caudal-fin lobe with three brownish-ochre bars of about twice the width of bars on ventral side of lobe and with white-hyaline interspaces as wide as these bars; first dorsal fin hyaline with rather irregularly arranged pale-brown pigmentation spots, streaks or small blotches on or between individual spines giving a weak indication of a horizontal stripe close to fin base and three more oblique stripes further above; first dorsal-fin tip unpigmented; second dorsal fin hyaline with pale-brown spots arranged into three horizontal stripes, fin tip pale-brown; pectoral, pelvic and anal fins hyaline, weakly pale reddish-brown along several fin rays; barbels rose whitish, pale red on base. Preserved HT. Dark brown dorsally at level of snout to termination of lateral line at caudal-fin base; pale brown to beige ventrally below eyes and level of pectoral-fin base; behind pectoral-fin base to second dorsal-fin base two weakly indicated brown bands intermingled with beige; caudal-fin bars and stripe and dorsal-fin pigmentation well retained; other fins pale hyaline; barbels pale. Etymology. The name ��� willwhite ��� is used as a composed noun in apposition. It honours the ichthyologist Dr. William T. White, Australian National Fish Collection, CSIRO National Research Collections Australia, Hobart, Australia, for collecting and photographing mullid specimens from S Indonesia and making them available for taxonomic research (including the types of Upeneus willwhite). Distribution, habitat, size. Only known from Lombok, S Indonesia; occurs most probably on shallow sandy bottoms; attains at least 92 mm SL. Remarks. This species was obtained at a fish market and hence may have local commercial value., Published as part of Uiblein, Franz & Motomura, Hiroyuki, 2021, Three new goatfishes of the genus Upeneus from the Eastern Indian Ocean and Western Pacific, with an updated taxonomic account for U. itoui (Mullidae: japonicus-species group), pp. 298-324 in Zootaxa 4938 (3) on pages 313-314, DOI: 10.11646/zootaxa.4938.3.2, http://zenodo.org/record/4569653, {"references":["Uiblein, F., Gledhill, D. C. & Peristiwady T. (2017 a) Two new goatfishes of the genus Upeneus (Mullidae) from Australia and Indonesia. Zootaxa, 4318 (2), 295 - 311. https: // doi. org / 10.11646 / zootaxa. 4318.2.4"]}

Upeneus itoui Yamashita, Golani & Motomura, 2011 Itou’s Goatfish Sakuya-himeji (standard Japanese name) (Figures 1D, E, 2–6; Tables 1–3) Upeneus itoui Yamashita, Golani & Motomura, 2011: Uiblein & Lisher 2013; Tashiro et al. 2014; Uiblein et al. 2017a, 2020a. Material examined. Paratypes (n = 6, 87– 134 mm SL). S Japan, Kagoshima Prefecture: KAUM–I. 10384, 87 mm SL, Minamisatsuma, Kasasa, Kataura, off E of Sakinoyama, 31°25.44’ N, 130°11.49’ E, 27 m depth, set net; KAUM–I. 13595, 118 mm SL, same sampling data (photo of fresh PT, Figure 1 E); KAUM–I. 13591, 104 mm SL, Minamisatsuma, Kasasa, NE of Matsu-shima Island, 31°25.06’ N, 130°12.32’ E, 20 m depth, set net (photo of fresh PT, Figure 1D); KAUM–I. 13594, 108 mm SL, Minamisatsuma, Kasasa, Kataura, off Kouzaki-yama, 31°26.00’ N, 130°10.05’ E, 36 m depth, set net; KAUM–I. 13715, 134 mm SL, Ibusuki, Kaimonkawajiri, 1 km SW of Kawajiri Fishing Port, 31°10’ N 130°32’ E, 40 m depth, set net; MNHN 2010-1050, 114 mm SL, Minamisatsuma, Kasasa, Kataura, off E of Sakinoyama, 31°25.44’ N, 130°11.49’ E, 27 m depth, set net. Non-types (n = 14, 88– 139 mm SL). KAUM–I. 14661, 111 mm SL, Ibusuki, Kaimonkawajiri, 1 km SW of Kawajiri Fishing Port, 31°10’ N 130°32’ E, 40 m depth, set net; KAUM–I. 27549, 111 mm SL, same sampling data; KAUM–I. 35617, 139 mm SL, Minamisatsuma, Kasasa, Kataura, off E of Sakinoyama, 31°25.44’ N, 130 11.49’ E, 27 m depth, set net; KAUM–I. 56303, 115 mm SL, same sampling data; KAUM–I. 97832, 98 mm SL, same locality and catch data; KAUM–I. 131500, 133 mm SL, same sampling data; KAUM–I. 35944, 133 mm SL, Kimotsuki, Uchinoura Bay, 31°17’ N, 131°05’ E, set net; KAUM–I. 123984, 117 mm SL, same sampling data; KAUM–I. 54077, 99 mm SL, Osumi Islands, Tanega-shima Island, Kumage-gun, Nakatane, off Kumano, 30°28.13’ N, 130°58.32’ E, 25 m depth, set net; KAUM–I. 59466, 101 mm SL, same sampling data; KAUM–I. 56749, 115 mm SL, Kimotsuki, Uchinoura Bay, 31°24.28’ N 131°02.55’ E, 40-45 m depth, set net; KAUM–I. 73580, 88 mm SL, Minamisatsuma, Kasasa, NE of Matsu-shima Island, 31°25.06’ N, 130°12.32’ E, 20 m depth, set net; KAUM–I. 110071, 131 mm SL, same sampling data; KAUM–I. 86533, 134 mm SL, Kimotsuki, Kishira, off Funama Fishing Port, 31°10’ N, 130°59’ E, set net. Diagnosis. Dorsal fins VII + 9; pectoral fins 13–14(15); gill rakers 6–7 + 16–18 = 22–25; body elongate and laterally slightly compressed; measurements as % SL: depth at first dorsal-fin origin 21–24; body depth at anal-fin origin 19–22; caudal-peduncle depth 9.0–10; caudal-peduncle width 3.9–4.6; maximum head depth 17–19; head depth through eye 13–15; head length 27–29; snout length 10–12; orbit length 5.4–6.4; upper-jaw length 9.8–11; barbel length 17–18; caudal-fin length 27–30; anal-fin height 15–17; pelvic-fin length 18–21; pectoral-fin length 19–21; first dorsal-fin height 18–21, dorsal-fin spines proportionally decreasing in height; second dorsal-fin height 15–17; total oblique bars on caudal fin 11 – 18, upper caudal-fin lobe with 5–7 mostly straight red or red-brown bars; lower caudal-fin lobe with 6–11 red or red-brown bars that are only faintly visible along ventral-lobe margin, becoming more conspicuous along central lobe; several of these bars and especially three of the most proximal ones reach dorsal-lobe margin, becoming pale reddish grey; lower caudal-fin lobe in life with a red or red-brown stripe of about orbit diameter in maximum width, covering bars only partly, that fades away post mortem; bars are mostly retained in the first years after preservation, but may be lost later; dorsal fins in fresh fish with 3–4 thin red or brown stripes, may be retained in preservative; barbels white or creamy white when fresh; body colour in freshly deceased fish dark red or dark brown dorsally and pale red or rose with red blotches or spots on lower half, ventral margin silvery white; a rather conspicuous dark brown, dark red or red mid-lateral body stripe of about pupil diameter in width at eye level from behind eye to caudal-fin base and in front of eye to nose tip, not retained in preservative; head below and behind eye silvery white, often covered with red patches or blotches; body and head dorsally grey or grey brown and ventrally often silvery white when preserved. ......continued on the next page * counts at left and right body side, respectively. Distribution and size. Known from S Japan, including Kagoshima, Miyazaki, Kochi and Ehime Prefectures and Okinawa-jima Island, Ryukyu Islands, and Taiwan; occurs on sandy bottoms; depth 4–40 m; attains 144 mm SL. Remarks. Upeneus ituoi is a rather common species which is often caught by set net in S Japan and has local commercial value. It co-occurs frequently with U. tragula Richardson, 1846 (Yamashita et al. 2011). Two specimens have been collected off Tanega-shima Island, Osumi Islands in the same area as the HT of U. elongatus (Tashiro et al. 2014).

The cuskeel genus Neobythites (Ophidiidae) is recorded for the first time from the Solomon Sea, SW Pacific, and a new species, N. solomonensis n. sp., is described based on five specimens (SL 137-166 mm) caught at 498-839 m depth in the eastern Solomon Sea. The new species is characterized by having two spines on hind margin of preoperculum, a distinct lateral body stripe and dark-brown or grey dorsal-fin margin. The most similar species is N. somaliaensis Nielsen, 1995, of which 14 specimens are compared. Neobythites solomonensis n. sp. differs from the latter in the following characters (N. somaliaensis in brackets): total vertebrae 58-61 (61-64), developed gill rakers 12-14 (9-11), orbit length 4.0-4.9 (5.0-6.1) in % SL, longest gill filament on anterior arch 4.6-5.3 (11.0-14.0) in % head length and presence (absence) of body stripe. The significance of studying colour patterns in the speciose genus Neobythites (55 valid species) is discussed.

The Madras Goatfish, Upeneus madras n. sp. (Mullidae), is described based on two specimens from Chennai, SE India, that were formerly identified as U. guttatus (Day, 1868), a widely distributed species which has the same type locality. The new species shares white barbels and oblique bars on the lower caudal-fin lobe, seven dorsal-fin spines which decrease proportionally in height and subequal pelvic and pectoral fins with the other eight species of the pori group, a newly established taxonomic species group. Upeneus madras n. sp. differs from all eight species in the lack of bars along the dorsal margin of the lower caudal-fin lobe and from the four most similar species mainly in the combination of caudal-peduncle shape and pectoral-fin width. Detailed comparisons with U. guttatus, a japonicus-group species, were made involving the study of two syntypes collected by Francis Day (the taxonomic authority of U. guttatus), from which we designate a lectotype. This species differs from U. madras n. sp. in yellow vs white barbels, shorter head, narrower pectoral fins and a mostly higher first dorsal fin with disproportionally elongated anterior spines. A review of barbel colour in the 47 Upeneus species suggests high diagnostic significance of this character for the distinction of several species and taxonomic species groups.

The so-called “posteli-species group”, a new taxonomic group of the goatfish genus Parupeneus is reviewed. This group consists of five species distributed in different areas of the Indo-Pacific, of which four species are known to extend to or occur exclusively on the deeper shelf or uppermost slope of 120-250 m: P. chrysonemus from Hawaiian Islands and Johnston Island, P. louise from French Polynesia (Central Pacific), P. moffiti from Guam (West Pacific) and P. posteli from Réunion, now also recorded from the southern Mascarene Plateau, Mauritius, South-Western Indian Ocean. One new species, P. sinai n. sp., from the northern Red Sea is here described based on a single preserved specimen collected during a French research mission about 90 years ago, of which no locality and depth information exist and the fresh colour when alive or after death is unknown. All five posteli-group species are very similar in morphometric and meristic characters, having a relatively shallow caudal peduncle, particularly long barbels, a high gill-raker number which – in combination with lack of dark body and fin pigmentation – distinguish them from all other congeners. Based on detailed examination of 62 morphometric and meristic characters, taxonomic accounts for each species and interspecific comparisons are provided, considering also size-related variation in body form in two species of which specimens of a wide size range were available for studies. Parupeneus sinai n. sp. clearly differs from the other four posteli-group species by a shorter head and snout, a lower ninth dorsal-fin ray, and in a combination of several other characters including body depth, barbel length and number of gill rakers. Comparisons among the remaining four species revealed that morphometric and meristic characters have a higher diagnostic significance than fresh colour patterns, which vary considerably among individuals. Morphological similarities of the posteli-species group with a taxonomic

The main objectives of this study were to clarify the taxonomic status of Mulloides armatus, listed as incertae sedis in current taxonomic literature, and to examine the extent of phenotypic differentiation among subspecies and populations of the yellowstripe goatfish Mulloidichthys flavolineatus (Mullidae) in the Indo- Pacific. In total, 53 quantitative morphometric and meristic characters and the visual detectability of the first dorsal-fin spine were gathered from 116 specimens split according to size into juveniles and adults. In addition, fresh colour imagery was studied. The Mulloides armatus holotype, which is broken into two pieces, could be identified as M. flavolineatus after reconstruction of standard length and detailed comparisons involving specimens from nearby the assumed type locality in the Southwestern (SW) Pacific. Then, the recently described subspecies M. f. flavicaudus of the Red Sea and populations of the nominal subspecies M. f. flavolineatus from the Indian Ocean and the Pacific, with additional subdivision into four smaller populations (SW Pacific, Wake Atoll, Hawaiian Archipelago and remaining Pacific), were studied by direct comparisons and uni- and multivariate statistical methods. For the first time, marked size differences (i.e. allometry) in morphometric characters in M. flavolineatus were documented. Mulloidichthys f. flavicaudus overlaps in all characters, singly and in combination, and under consideration of a whitish-grey vs. yellow caudal-fin colour (a diagnostic character used in the original description) with M. f. flavolineatus. These results do not support the elevation to species level in a recently published list of Red Sea fish species. Our data suggest a well-differentiated population in agreement with genetic data presented in the original description of Mulloidichthys f. flavicaudus. Statistical differences in morphometric and meristic characters were also found between the Indian Ocean and Pacific populations of M. f. flavolineatus, though at a much lower degree than between the two subspecies. Interestingly, considerable variation occurs within the Pacific populations with three specimens from Wake Atoll being distinct from nearly all other conspecifics, in having larger heads and eyes and longer barbels and pectoral fins. Therefore, we stress the significance of intraspecific differentiation in populations from remote oceanic island and atoll areas and the need to collect more data. Important is also to use results on intraspecific regional or local differentiation in widely distributed species such as M. flavolineatus in fisheries management and conservation efforts at biologically relevant scales, as well as adopting appropriate common names to facilitate the information exchange with local stakeholders.

Upeneus heterospinus n. sp. Uiblein & Pavlov Varied-spine goatfish (Figures 1, 9���12; Tables 2, 4���11) Upeneus australiae Kim & Nakaya, 2002: Uiblein & Heemstra 2010 (Nha Trang, South-central Vietnam, South China Sea). U. bensasi (Temminck & Schlegel, 1843): Provincial Keelung Girls��� High School, Keelung, Taiwan 1985 (fresh-colour photo, no locality, but size information (92 mm SL) and collecting date (03/12/1979). U. guttatus (Day, 1868): Motomura et al. 2017 (vouchered fresh-colour photo, central Philippines, W Pacific). U. margarethae: Pavlov & Emel���yanova 2016 (fresh-colour photos, South-central Vietnam, South China Sea). U. cf. margarethae: Emel���yanova & Pavlov 2014; White et al. 2013 (vouchered fresh-colour photo, S Indonesia). U. spottocaudalis Uiblein & Gledhill, 2017: Bandai et al. 2018 (vouchered fresh-colour photo and comparative data, S Japan, W Pacific). U. tragula: Froese & Pauly 2019 (fresh colour photo by A. Cornell from off Hong Kong, South China; https://www.fishbase. de/photos/PicturesSummary.php?StartRow=5&ID=5443&what=species&TotRec=14). Upeneus sp.: Kimura et al. 2018 (vouchered fresh colour photo, N Vietnam, Ha Long Bay). Upeneus sp.1: Uiblein & Lisher 2013 (Nha Trang, South-central Vietnam, South China Sea). Holotype. VNMN-I 2015, adult, 91 mm SL, W Pacific, South China Sea, South-central Vietnam, Khanh Hoa province, Nha Trang, N of Hon Tre, 12�� 13���21������ N, 109�� 18���30������ E, obtained from hookah divers, 23 Feb. 2010, collector: Dimitri Pavlov (fresh-colour photo) Paratypes. (28 adults, 1 subadult: 56���152 mm SL, 15 fresh-colour photos). W Pacific, South China Sea, Southcentral Vietnam: Phu Yen province, Vung Ro Bay, 12�� 52.11��� N, 109�� 24.66��� E, obtained from hookah divers: VNMN-I 2016, 70 mm SL, (fresh-colour photo); VNMN-I 2017, 101 mm SL (fresh-colour photo); VNMN-I 2018, 96 mm SL (fresh-colour photo); VNMN-I 2019, 56 mm SL (fresh-colour photo); Khanh Hoa province, Nha Trang Bay: N of Hon Tre, 12�� 13���21������ N, 109�� 18���30������ E, obtained from hookah divers: VNMN-I 2020, 97 mm SL (fresh-co- lour photo); VNMN-I 2021, 112 mm SL; VNMN-I 2022, 106 mm SL (fresh-colour photo); VNMN-I 2023, 113 mm SL (fresh-colour photo); VNMN-I 2024, 105 mm SL (fresh-colour photo); VNMN-I 2025, 109 mm SL; VNMN-I 2026, 100 mm SL (fresh-colour photo); VNMN-I 2027, 112 mm SL; VNMN-I 2028, 104 mm SL; VNMN-I 2029, 104 mm SL; VNMN-I 2030, 103 mm SL; VNMN-I 2031, 96 mm SL; VNMN-I 2032, 103 mm SL; S of Hon Tre, obtained from hookah divers: HIFIRE F 58110, 120 mm SL; HIFIRE F 58115, 109 mm SL; VNMN-I 2033, 123 mm SL; Cua Be: CSIRO H 8427-01, 90 mm SL (fresh-colour photo), landing site, 12�� 12.09��� N, 109�� 12.12��� E; HIFIRE F 58203, 105 mm SL (fresh-colour photo), fish market, 12�� 12.10��� N, 109�� 12.05��� E; Nha Trang city, fish market: CSIRO H 8428-01, 82 mm SL (fresh-colour photo); CSIRO H 8428-02, 82 mm SL (fresh-colour photo); HIFIRE F 58178, 152 mm SL; NHMO J 7217, 114 mm SL (fresh-colour photo); NHMO J 7218, 103 mm SL (fresh-colour photo); Nha Trang, no further locality details: MNHN 1965-272, 90 mm SL; ZMUC P49483, 84 mm SL. Non-types. (72 adults, 6 subadults: 43���127 mm SL, 30 fresh colour photos). W Pacific, South China Sea, N Vietnam, Qu���ng Ninh province, Ha Long Bay, Van Don landing site, 21�� 03.61��� N, 107��25.81��� E: HIFIRE F 58382, 97 mm SL (fresh-colour photo); NHMO J 7219, 104 mm SL (fresh-colour photo); VNMN-I 2034, 109 mm SL (fresh-colour photo); VNMN-I 2035, 113 mm SL (fresh-colour photo); VNMN-I 2036, 104 mm SL (fresh-colour photo); VNMN-I 2037, 108 mm SL; VNMN-I 2038, 108 mm SL (fresh-colour photo); Ha Long: VNMN-I 2039, 83 mm SL (fresh-colour photo); VNMN-I 2040, 83 mm SL (fresh-colour photo); VNMN-I 2041, 122 mm SL (fresh- colour photo); VNMN-I 2042, 93 mm SL (fresh-colour photo); VNMN-I 2043, 108 mm SL (fresh-colour photo); VNMN-I 2044, 94 mm SL (fresh-colour photo); VNMN-I 2045, 117 mm SL (fresh-colour photo); VNMN-I 2046, 87 mm SL (fresh-colour photo); VNMN-I 2047, 72 mm SL (fresh-colour photo); VNMN-I 2048, 82 mm SL (freshcolour photo); VNMN-I 2049, 80 mm SL (fresh-colour photo); VNMN-I 2050, 77 mm SL (fresh-colour photo); VNMN-I 2051, 81 mm SL (fresh-colour photo); VNMN-I 2052, 64 mm SL (fresh-colour photo); VNMN-I 2053, 60 mm SL (fresh-colour photo); South-central Vietnam, Khanh Hoa province, Nha Trang Bay: HIFIRE F 58231, 123 mm SL, N of Hon Tre Island, 12��14���18���N 109��15���54���E, obtained from hookah divers (fresh-colour photo); Gulf of Thailand, S Vietnam, Phu Quoc, An Thoi market, 10�� 0.90��� N, 104�� 0.86��� E: HIFIRE F 58287, 93 mm SL (freshcolour photo); NHMO J 7220, 88 mm SL (fresh-colour photo); VNMN-I 2054, 83 mm SL (fresh-colour photo); VNMN-I 2055, 114 mm SL (fresh-colour photo); Gulf of Thailand, Thailand, Ko-Sichang Island: CAS 17679, 41 (of 141): 56���94 mm SL, 13��8��� N, 100��51��� E, trawl, 18���24 m depth; Singapore: NHMO J 2134, 2, 78���81 mm SL; Philippine Sea, Central Philippines, Panay Island: UPVMI 155, 96 mm SL (fresh-colour photo); Indonesia, Bali Sea, E Java: CSIRO H 7360-04, 3: 43���66 mm SL, Banguwangi, 08��09��� S 114��23��� E; Lombok (border of W Pacfic to Indian Ocean): MZB, unregistered (field code LM308 (large)), 54 mm SL; CSIRO H 8409-02, 65 mm SL, and CSIRO H 7364-02, 127 mm SL, Tanjung Luar, 8��45��� S, 116��35��� E (both with fresh-colour photo); BPBM 29896, 96 mm SL, no locality information (fresh-colour photo). Fresh-colour photos of specimens not retained (n =79): W Pacific, Vietnam (D.A. Pavlov & F. Uiblein), South China Sea: 20 adults, Ha Long Bay, N Vietnam; 10 adults, Nha Trang Bay, South-central Vietnam; Gulf of Thailand, S Vietnam, Phu Quoc: 37 adults and 1 subadult or adult (in-situ photo); South China Sea, other areas: 1 adult, Hong Kong, China (A. Cornell); 1 adult, Taiwan (Provincial Keelung Girls��� High School, Keelung, Taiwan, 1985); Philippine Sea, Philippines: 7 adults, 1 subadult, Central Philippines (various localities, in-situ photos; Philippe & Guido Poppe���www.poppe-images.com); Japan: adult, 96 mm SL, Satsuma Peninsula, Kagoshima, S Japan (Bandai et al. 2018). Diagnosis. Dorsal fins VII or VIII + 9, the first spine minute in 8-spined fish; pectoral fins 13���15; gill rakers 4���6 + 15���18 = 21���24; lateral-line scales 28���30; measurements in % SL, adults: body depth at first dorsal-fin origin 22���26; body depth at anus 18���22; caudal-peduncle depth 9.2���11; caudal-peduncle width 3.5���5.0; maximum head depth 19���23; head depth through eye 15���18; suborbital depth 8.3���11; interorbital length 7.2���9.2; head length 27���31; snout length 9.7���13; postorbital length 10���13; orbit length 5.9���8.3; upper-jaw length 9.4���13; barbel length 16���20; interdorsal distance 13���17; caudal-peduncle length 22���26; caudal-fin length 27���32; anal-fin height 15���19; pelvic-fin length 19���23; pectoral-fin length 19���22; pectoral-fin width 3.4���4.7; first dorsal-fin height 18���23; second dorsal-fin height 16���20; measurements in % SL, subadults: body depth at first dorsal-fin origin 22���24; body depth at anus 18���21; caudal-peduncle depth 9.1���10; caudal-peduncle width 3.6���3.8; maximum head depth 19���21; head depth through eye 15���18; suborbital depth 9.0���10; interorbital length 7.6���8.1; head length 29���30; snout length 11���12; postorbital length 11���13; orbit length 7.8���7.9; upper-jaw length 9.9���12; barbel length 18���19; interdorsal distance 12���14; caudal-peduncle length 22���25; caudal-fin length 29���31; anal-fin height 16���19; pelvic-fin length 21���22; pectoral-fin length 21; pectoral-fin width 3.8���4.2; first dorsal-fin height 21���22; second dorsal-fin height 17��� 20; fresh colour: head and body dorsally red, brown or dark-grey mottled, ventrally white, often with pale beige or grey dots or red blotches (inactive, live fish with large red patches and broad vertical or oblique red bands on body); upper lobe of caudal fin with 4���6 red or brown bars (3 or 4 bars in subadults), lower caudal-fin lobe with 5���7 (3���5) red, brown or dark-brown bars, often crossed, entirely or ony dorsally, by a yellow, beige, pale red or pale brown stripe; bars on both lobes of pupil width or less in adults, interrupted by pale, partly hyaline interspaces of similar width or slightly narrower; lower caudal-lobe tip sometimes black; barbels entirely yellow; a single yellow, beige, or pale brown mid-lateral body stripe of pupil width from snout tip through eye to caudal-fin base; stripe sometimes covered by 1���4 sections with 1���4 dark dots, the posteriormost group of dots behind second dorsal-fin base, often connecting to a dark saddle behind second dorsal fin; first dorsal fin with 3 or 4 often fused, pale-red, pale-brown or beige stripes, the fin-tip area not darker pigmented; second dorsal fin with 3 or 4 red or brown well-separated stripes with hyaline interspaces; pectoral fins hyaline, pelvic and anal fins weakly pigmented and partly hyaline; preserved fish pale brown, often with remains of dark pigmentation deriving from caudal-fin bars, mid-lateral dots, saddle and/or second dorsal-fin distal stripe. Description. Measurements in % SL and counts for types are given in Table 8; morphometric data as ratios of SL for holotype, data for paratypes in brackets: body elongate, body depth at first dorsal-fin origin 4.2 [3.8���4.6], body depth at anal-fin origin 5.0 [4.5���5.4], head length 3.5 [3.3���3.7], larger than maximum body depth and subequal to caudal-fin length (3.3 [3.2���3.7]), second dorsal-fin height 6.1 [5.3���6.4], slightly shallower than first dorsal-fin height (4.7 [4.5���5.6]), barbel length (5.5 [5.0���5.9]), pelvic-fin length 5.0 [4.5���5.3], similar to length of pectoral fins (5.2 [4.7���5.3]) and body depth at first dorsal-fin origin; caudal-peduncle depth 10 [8.9���10], clearly larger than orbit length (14 [12���17]); and caudal-peduncle width 21 [20���28], subequal to pectoral-fin width (23 [22���27]). Colour. Freshly collected fish. (Figure 9). Head and body ventrally white and ventro-laterally white or palegrey whitish with numerous red, brown or dark-grey dots of half pupil size or less, placed mostly on individual scales (Figure 9 A, B, E, F); in all six specimens shown in Figure 9 (least in holotype, Figure 9 A), red blotches of different form and size or larger red pigmentation patches below or behind eye on gill cover and/or on body ventrally or ventro-laterally either covering larger areas (Figure 9 B, C) or more restricted to smaller areas on or behind belly (Figure 9 E, F); some of these blotches and patches may have catch/treatment-related context; head from above snout and eye and body above lateral line red-brown or dark-grey mottled, bordered below by a mid-lateral stripe of pupil width which runs rather straight or only slightly bent from caudal-fin base to eye in yellow, beige, pale red or pale brown colour and continues from eyes (with red or pale-red iris) to snout either in similar colour and intensity (Figure 9 B���D), or slightly darker, but weaker (Figure 9 E, F), or rather faint and indistinct (Figure 9 A); on the mid-lateral body stripe from behind head to behind second-dorsal fin up to 4 sections with 1���3 red, brown or black dots of less than pupil width; behind head until below base of second dorsal fin in adults, a thin white or pale creamy stripe separates the dorsal body pigmentation from the mid-lateral body stripe (Figure 9 A, B, E); lateral line visible especially in subadult (Figure 9 C) and small adult (Figure 9 F) as a series of mostly red, dark-grey or black dots placed well above mid-lateral stripe in anterior half of body, then starting to cross the stripe just anterior of second dorsal-fin base, continuing behind dorsal fin just below stripe towards caudal-fin base; barbels entirely yellow; caudal fin with 9���13 red, dark brown or black oblique bars crossing mostly both lobes entirely, if not placed on fin base or fin tips; upper lobe covered by 4���6 (3���4 in subadults) red, red brown, or dark brown bars of pupil or slightly narrower width, the distal-most bar sometimes covering fin tip (Figure 9 A, F); bars separated by hyaline interspaces of similar width or slightly wider; in large adult (Figure 9 E) dark-brown blotches on three intermediate bars; lower caudal-fin lobe covered by 5���7 (3���5 in subadults) red, red brown or dark brown bars and hyaline interspaces of similar width or slightly narrower than on upper fin lobe in adults, whereas more variable in subadults; in four specimens (Figure 9 A���D), the lower caudal-fin lobe ends in a black tip; a red or brown stripe stretches entirely or only dorsally along the lower caudal-fin lobe covering partly the bars in both aduts and subadults; first dorsal fin almost entirely covered with pale-red, beige or pale-brown pigmentation; second dorsal fin with 3 red stripes of orbit width or less, one stripe at or close to fin base, one at or close to fin tip, the latter and in some fish also the middle stripe slightly darker; pelvic fins hyaline with up to 5 narrow red or brown stripes; anal fin with 1���3 red or brown stripes and hyaline or whitish interspaces; pectoral fins hyaline. Live fish in situ. Active fish. (Figure 10 A���D). Head and body ventrally white, with numerous tiny red or beige dots associated with 4 scale rows and, in three of four fish, with additional, larger rounded and partly connected red blotches, ca. 25 of the latter visible in fish from the type locality which was kept in a tank (Figure10 A); fewer and more ventrally positioned blotches in two fish photographed in situ in Central Philippines (Figure 10 C, D); head from above snout and eye and body above lateral line pale-grey or rose mottled, bordered below by a mid-lateral stripe in red colour (that appears brown when photographed without flash, see Figure 10 B) of pupil width which runs rather straight from caudal-fin base to eye, continuing through eyes (with iris red-coloured in stripe shape) and then bending ventrally towards snout tip, becoming slightly narrower and weaker; 5 well separated, rectangular dark-red sections on mid-lateral body stripe (not well visible in Figure 10 B), one shorter behind gill cover, two longer ones below dorsal fins, one shorter behind second dorsal fin, and one less conspicuous, rather short section close to caudal-fin base; each section except for the posterior-most with a horizontal series of 2���4 black dots of a little less than pupil diameter; the dotted section behind the second dorsal fin connecting closely to red saddle in tank fish (Figure 10A), the latter reaching from dorsal body margin behind second dorsal fin to stripe in V-form; lateral line, as also observed in freshly collected fish, proceeds parallel above mid-lateral body stripe during first half of body, then crossing and continuing just below stripe to caudal-fin base; lateral line marked by several small red, beige or grey dots that become more conspicuous in positions of dots on mid-lateral body stripe below; barbels entirely yellow; caudal fin with 8���12 red-brown oblique bars which appear dark-brown in the small possibly adult specimen photographed at distance without flash (Figure 10 B); dorsal fin lobe crossed by 3���5 bars of pupil width, the distal-most bar covering fin tip in two specimens (Figure 10 A, D); in three specimens the bars are reduced to dark red or dark brown blotches (Figure 10 B, C, D) leaving the remaining parts of bars as pale, mostly hyaline traces (Figure 10 C, D); bars on upper lobe separated by hyaline interspaces of similar width; lower caudal-fin lobe crossed in adults by 5���7 dark red (dark brown in Figure 10 B) bars which are similarly shaped and sized as bars on upper lobe; while in the tank specimen the bars are of varying intensity but typically shaped, the bars in the other three active fish show a blotch-like concentration of intense colour in the middle of the lower lobe, leaving the ventral and dorsal areas of each bar rather inconspicuous (however still visible, e.g. Figure 10 C, D); in addition, a red stripe is formed on the lower lobe of the tank specimen that covers the lobe almost entirely (apart from ventral margin) and a pale red stripe is also weakly visible along the dorsal half of lobe in two of the three other specimens (Figure 10 C, D); the first dorsal fin (only well visible in Figure 10 A) is mostly weakly pale red, leaving the tip almost completely unpigmented hyaline; the second dorsal fin with 3-4 red to brown stripes of about pupil width or less, at least one of the distal two stripes more intensely coloured than the stripes (or single stripe) closer to fin base which appear rather pale red and partly hyaline; interspaces between stripes in part with white pigmented fin rays; pelvic and anal fins partly hyaline with weak pale grey pigmentation indicating stripes; pectoral fins hyaline, only very faintly visible. Resting fish. (Figure 10 E, only differences from the above colour description indicated). Large areas on head and body covered with red, in particular below eye, on gill cover, above pectoral-fin base, two large red bands running from dorsal fins down to ventral margin, the anterior band rather straight from first dorsal fin, the posterior band obliqueand wider, covering also the area of and below saddle; the two bands connecting on ventral body margin with each other and covering most of ventral underside except for an inverted V-shaped pale area behind pelvicfin base and caudal peduncle; caudal peduncle mostly white with three round orbit-sized red blotches on ventral margin and one more elongated red blotch just above the last ventral-margin blotch; mid-lateral body stripe only visible in areas without red band markings and almost indistinct whitish on caudal peduncle, then orange anterior to second dorsal fin between the large red bands and behind head, turning into red on gill cover, then crossing eye (with iris red-colour in stripe shape) and bending towards snout tip, still in red colour; caudal fin with 9 oblique dark grey bars, on dorsal lobe 4 bars (including one on tip) which are dark pigmented only on dorsal margin of fin, while becoming ventrally pale grey and rather indistinct; the 5 bars on lower fin lobe entirely dark grey, slightly wider than whitish interspaces, with faint appearance of a grey stripe only close to caudal-fin base; area of red pigmentation on first dorsal fin concentrated centrally and in part of fin, leaving the tip entirely pale hyaline. Preserved fish. Head and body mostly uniformly pale brown, pale greyish, greyish brown or brown, sometimes slightly darker dorsally, gill cover silvery or pale and partly transparent, lateral line, when intact, well visible in entire range from behind head to caudal-fin base; barbels pale brown or pale creamy; mid-lateral body stripe completely lost, but often remains of dark mi, Published as part of Uiblein, Franz, Gledhill, Daniel C., Pavlov, Dimitri A., Hoang, Tuan Anh & Shaheen, Shaker, 2019, Three new goatfishes of the genus Upeneus (Mullidae) from the Indo-Pacific, with a redescription of colour patterns in U. margarethae, pp. 151-196 in Zootaxa 4683 (2) on pages 180-185, DOI: 10.11646/zootaxa.4683.2.1, http://zenodo.org/record/3772664, {"references":["Kim, B. J. & Nakaya, K. (2002) Upeneus australiae, a new goatfish (Mullidae: Perciformes) from Australia. Ichthyological Research, 49, 128 - 32. https: // doi. org / 10.1007 / s 102280200016","Uiblein, F. & Heemstra, P. C. (2010) A taxonomic review of the Western Indian Ocean goatfishes of the genus Upeneus (family Mullidae) with descriptions of four new species. Smithiana Bulletin, 11, 35 - 71.","Motomura, H., Alama, U. B., Muto, N., Babaran, R. P. & Ishikawa, S. (2017) Commercial and Bycatch Market Fishes of Panay Island, Republic of the Philippines. The Kagoshima University Museum, Kagoshima, University of the Philippines Visayas, Iloilo, and Research Institute for Humanity and Nature, Kyoto, 246 pp.","Pavlov, D. A. & Emel'yanova, N. G. (2016) Reproductive features of Upeneus margarethae (Mullidae), a species recorded in the coastal zone of Vietnam for the first time. Journal of Ichthyology, 56 (4), 600 - 612. https: // doi. org / 10.1134 / S 0032945216040093","Emel'yanova, N. G. & Pavlov, D. A. (2014) Gamete ultrastructure in two species of the genus Upeneus (Mullidae) from the South China Sea. Journal of Ichthyology, 54 (4), 286 - 292. https: // doi. org / 10.1134 / S 0032945214030023","White, W. T., Last, P. R., Dharmadi, Faizah, R., Chodrijah, U., Prisantoso, B. I., Pogonoski, J. J., Puckridge, M. & Blaber, S. J. M. (2013) Market Fishes of Indonesia. ACIAR Monograph No. 155. Australian Centre for International Agricultural Research, Canberra, 438 pp.","Uiblein, F., Gledhill, D. C. & Peristiwady, T. (2017) Two new goatfishes of the genus Upeneus (Mullidae) from Australia and Indonesia. Zootaxa, 4318 (2), 295 - 311. https: // doi. org / 10.11646 / zootaxa. 4318.2.4","Bandai, A., Itou, M. & Motomura, H. (2018) First Northern Hemisphere record of the tailspot Goatfish Upeneus spottocaudalis (Perciformes: Mullidae) from Kagoshima, Japan. Japan. Journal of Ichthyology, 65, 35 - 39. https: // doi. org / 10.11369 / jji. 17 - 056","Froese, R. & Pauly, D. (Eds.) (2019) FishBase. World Wide Web electronic publication. Availabe from: https: // www. fishbase. in / search. php (accessed 2 Feburary 2019)","Kimura, S., Imamura, H., Nguyen, V. Q. & Pham, T. D. (2018) Fishes of Ha Long Bay, the Natural World Heritage Site in Northern Vietnam. Fisheries Research Laboratory, Mie University, Shima, 314 pp.","Uiblein, F. & Lisher, M. (2013) A new goatfish of the genus Upeneus (Mullidae) from Angoche, northern Mozambique. Zootaxa, 3717 (1), 85 - 95. https: // doi. org / 10.11646 / zootaxa. 3717.1.7"]}

Upeneus caudofasciatus n. sp. Uiblein & Gledhill Tailbar goatfish (Figures 1, 6���8; Tables 2, 4���9) Holotype. CSIRO H 6717-02, adult, 116 mm SL, W Pacific, NE Australia, Queensland, Great Barrier Reef, N of Rockingham Bay, 17�� 38.57��� S, 146�� 22.79��� E, FRV Gwendoline May, otter trawl, 26 m depth; collector Daniel Gledhill, 25 Nov 2003 (fresh-colour photo) Paratypes. (30 adults, 4 subadults: 53���124 mm SL, 1 fresh-colour photo). W Pacific, NE Australia, Queensland: Torres Strait: CSIRO H 6441-03, 90 mm SL, Torres Strait, S of Bristow Island, 09�� 15.65��� S, 143�� 20.72��� E, FRV Gwendoline May, otter trawl, 28 m depth; QM I.15833, 83 mm SL, W of Adolphus Passage, 10�� 38��� S, 142�� 28��� E, 16���18 m depth, trawl; QM I. 16503, 124 mm SL, Aureed Island area, 9�� 57��� S, 143�� 17��� E, trawl; Great Barrier Reef: AMS I.20753-004, 14: 53���99 mm SL, Lizard Island area, 2 nm NW of Nymph Island, 14�� 36��� S, 145�� 14��� E, prawn trawl, 15 m depth; AMS I.34398-034, 89 mm SL, South Arm Channel, Port Clinton, adjacent to West Flat, 22�� 33��� 29��� S, 150�� 45��� 19��� E, 11 m depth; CSIRO H 5957-22, 2: 83-86 mm SL, near Cape Flattery, 14��46.8���S, 145��15.7���E, FRV Gwendoline May, otter trawl, 17 m depth; CSIRO H 6519-21, 2: 78-98 mm SL, NE of Cooktown, 15�� 01.91��� S, 145�� 29.40��� E, FRV Gwendoline May, trawl, 39 m depth; CSIRO H 7212-01, 95 mm SL, SE of Cairns, 17�� 08.90��� S, 146�� 12.11��� E, FRV Gwendoline May, trawl, 35 m depth (fresh-colour photo); CSIRO H 7660-01, 2: 110-112 mm SL, E of Northumberland Islands, 21�� 37.38��� S, 150�� 07.79��� E, FRV Gwendoline May, 34 m depth, trawl; CSIRO H 7664-01, 105 mm SL, NE of Gladstone, 23�� 37.47��� S, 151�� 41.48��� E, FRV Gwendoline May, otter trawl, 38 m depth; QM I.15628, 89 mm SL (largest of 11), 2 nm off NW of Nymph Island, 14�� 36��� S, 145�� 14��� E, trawl, 15 m depth; QM I.15677, 83 mm SL, 5 nm WNW of Lizard Island, 15�� 30��� S, 145�� 22��� E, trawl, 20 m depth; QM I.16213, 83 mm SL, (1 of 6 (marked)), 3 ml NW of Lizard Island, 14�� 38��� S, 145�� 24��� E, trawl, 26 m depth; QM I.17982, 93 mm SL, SE of Cooktown, 15�� 46��� S, 145�� 35��� E, 27 m depth; QM I.18044, 86 mm SL, Lizard Island, W of, 14�� 36��� S, 145�� 3��� E, 13 m depth; QM I. 39294, 105 mm SL, NE of Yeppoon, 22�� 44.16��� S, 151�� 26.56��� E, FRV Gwendoline May, trawl, 60 m depth; QM I.39316, 2, 94-111 mm SL, Flora Passage, 17�� 3��� S, 146�� 14��� E, trawl, 37-42 m depth. * p���0.01; ** pNon-types. (11 adults, 9 subadults: 50���107 mm SL, 1 fresh-colour photo). W Pacific, NE Australia, Queensland: Torres Strait: AMS I.20827-027, 3: 80���107 mm SL, Cape York, 2 nm NE of Hannibal Island, 11�� 33��� S, 142�� 57��� E, prawn trawl, 23 m depth; AMS I.20923-001, 5: 63���97 mm SL, Cape York, 10�� 39��� S, 142�� 30��� E, prawn trawl; CSIRO H 3637-10, 69 mm SL, W of Thursday Island, 10�� 34��� S, 141�� 59��� E, FRV Southern Surveyor, trawl, 12 m depth (fresh-colour photo); CSIRO H 3637-11, 50 mm SL, W of Thursday Island, 10�� 34��� S, 141�� 59��� E, FRV Southern Surveyor, trawl, 12 m depth; CSIRO H 7661-01, 2: 60 mm SL (both), S of Banks Island, 10�� 25.20��� S, 142�� 18.60��� E, FRV Gwendoline May, trawl, 14 m depth; CSIRO H 7689-02 3: 56���62 mm SL, NE of Dungeness Island, 9�� 39.49��� S, 142�� 47.61��� E, FRV Gwendoline May, trawl, 10 m depth; QM I. 16482, 104 mm SL, sand cay SE of Coconut Island, 10�� 11��� S, 143�� 14��� E, trawl; QM I.16483, 97 mm SL, Aureed Island area, 9�� 57��� S, 143�� 17��� E, Trawl; Great Barrier Reef: AMS I.20752-013, 3 (of 5): 54���92 mm SL, 5 nm WNW of Lizard Island, 14�� 30��� S, 145�� 22��� E, prawn trawl, 20 m depth. Diagnosis. Dorsal fins VIII + 9, the first spine minute; pectoral fins 12���14; gill rakers 5���7 + 16���19 = 22���25; lateral-line scales 28���30; measurements in % SL, adults: body depth at first dorsal-fin origin 24���28; body depth at anus 19���24; caudal-peduncle depth 9.9���11; caudal-peduncle width 2.6���4.3; maximum head depth 20���24; head depth through eye 16���20; suborbital depth 9.2���13; interorbital length 7.5���8.8; head length 28���31; snout length 9.9���12; postorbital length 10���13; orbit length 6.7���8.7; upper-jaw length 9.9���13; barbel length 18���23; interdorsal distance 12���17; caudal-peduncle length 21���25; caudal-fin length 27���32; anal-fin height 16���20; pelvic-fin length 20���24; pectoral-fin length 19���23; pectoral-fin width 3.6���4.8; first dorsal-fin height 19���23; second dorsal-fin height 18���21; measurements in % SL, subadults: body depth at first dorsal-fin origin 24���27; body depth at anus 19���22; caudal-peduncle depth 9.6���11; caudal-peduncle width 2.7���3.8; maximum head depth 20���23; head depth through eye 17���20; suborbital depth 9.4���11; interorbital length 7.6���8.6; head length 30���32; snout length 11���12; postorbital length 11���13; orbit length 8.1���8.8; upper-jaw length 11���13; barbel length 19���23; interdorsal distance 12���15; caudalpeduncle length 22���24; caudal-fin length 28���32; anal-fin height 16���19; pelvic-fin length 22���25; pectoral-fin length 21���23; pectoral-fin width 3.8���4.6; first dorsal-fin height 20���23; second dorsal-fin height 18���21; fresh colour: head and body dorsally red-brown or dark grey mottled, ventrally white, often with pale beige dots; upper lobe of caudal fin with 4 or 5 red or brown bars (3 or 4 bars in subadults), lower caudal-fin lobe with 4 to 9 red, brown or darkbrown bars, sometimes dorsally covered by a red, brown or dark-brown stripe; bars on both lobes of pupil width or less, interrupted by pale, partly hyaline interspaces of less or similar width; lower caudal-lobe tip sometimes black; barbels white, pale rose whitish, or white with beige tip region; a single yellow, beige or pale brown mid-lateral body stripe of pupil width from snout tip through eye to caudal-fin base, following lateral line in anterior two thirds of body; stripe covered by 1���4 sections with horizontal series of 2���4 dark dots, the posteriormost dots behind second dorsal-fin base; first dorsal fin with 3 or 4 red or brown, often fused stripes and the tip region pale or not darker than rest of fin; second dorsal fin with 3 or 4 red or brown well-separated stripes with hyaline interspaces; pectoral fins hyaline, pelvic and anal fins weakly pigmented and partly hyaline; preserved fish pale, often with remains of dark pigmentation deriving from caudal-fin bars, mid-lateral dots, saddle and/or second dorsal-fin distal stripe. Description. Measurements in % SL and counts for types are given in Table 8; morphometric data as ratios of SL for holotype, data for paratypes in brackets: body elongate, body depth at first dorsal-fin origin 3.9 [3.5���4.2], body depth at anal-fin origin 4.7 [4.2���5.2], head length 3.4 [3.2���3.6] subequal to caudal-fin length (3.6 [3.1���3.7]), second dorsal-fin height 5.5 [4.7���5.7], only slightly shallower than first dorsal fin (5.1 [4.3���5.1]) and similar to barbel length (5.1 [4.4���5.6]), pelvic-fin length 4.7 [4.0���5.0], similar to length of pectoral fins (5.0 [4.3���5.2]) and body depth at first dorsal-fin origin; caudal-peduncle depth 9.8 [8.9���10], larger than orbit length (15 [12���15]); and caudal-peduncle width 30 [24���38], narrower than pectoral-fin width (22 [21���27]). Colour. Freshly collected fish. (Figure 7 A���C). All three specimens and especially holotype and paratype (Figure 7 A, B) show obvious signs of lesions (most probably due to trawling damage) with large red, skin-lacking patches on head below eye, gill cover or mid of body. Otherwise head and body are ventrally white, overlain by small beige to beige-rose dots associated with individual scales in paratype (Figure 7 B) and non-type (Figure 7 C); belly and ventral part of caudal peduncle white; head from above snout and eye and body above lateral line pale brown-red to dark-grey mottled, bordered below by a mid-lateral stripe of pupil width which runs rather straight or only slightly bent from caudal-fin base to eye in yellow or greenish beige colour, becoming reddish on head and bending ventrally towards snout tip (stripe on head not visible in holotype); on the mid-lateral body stripe from behind head to behind second-dorsal fin up to 4 well-separated sections just behind head, below each dorsal, and behind second dorsal fin, each with 1���3 red, brown or black dots of less than pupil width; from behind head until below base of second dorsal fin, a thin white stripe separates the dorsal body pigmentation from the mid-lateral body stripe; lateral line weakly visible as a series of mostly red dots placed well above mid-lateral stripe in anterior half of body starting to cross the stripe just in front of anterior second dorsal-fin base, continuing behind dorsal fin just below stripe towards caudal-fin base; barbels entirely white (paratype), white, intermingled with pale rose (holotype) or white with a beige tip region (non-type) (Figure 7 A���C); caudal fin with 9���14 oblique red bars crossing both lobes entirely, or placed on fin tips; upper caudal-fin lobe with 4���5 bars of pupil width or slightly narrower, the distal-most bar covering fin tip in holotype and non-type; bars separated by hyaline interspaces of similar width or slightly wider; in holotype and non-type blotches of dark red or black pigmentation on some of the bars; lower caudal-fin lobe covered by 5���9 red bars of similar width or slightly narrower (holotype) than bars on dorsal fin lobe, with hyaline interspaces being of similar width or slightly narrower; in all three specimens the ventral lobe ends in a black tip and some of the other bars contain dark-red or black blotches; the lower fin lobe is in addition crossed longitudinally by a pale-red stripe that reaches from base of fin to fin tip, crossing nearly the entire fin lobe apart from ventral margin (types) or the dorsal half of lobe (non-type); first dorsal fin almost entirely covered with pale red or pale orange pigmentation intermingled with hyaline areas; second dorsal fin with 3 red stripes of orbit width or less, one stripe at fin base, one intermediate and one at or close to fin tip, the latter being darker than the other stripes; pelvic fins hyaline with up to five narrow red stripes; anal fin with or without stripes, pectoral fins hyaline. Preserved fish. Head and body mostly uniformly pale brown, greyish brown or brown, sometimes slightly darker dorsally, gill cover silvery or pale and partly transparent, lateral line, when intact, well visible in entire range from behind head to caudal-fin base; barbels pale brown or pale creamy; mid-lateral body stripe completely lost, but often remains of dark dots mid-laterally in the area of stripe below and posterior to dorsal fins (51 % of 41 adults, 60 % of 10 subadults); dark pigmentation remains occur also in other areas: saddle behind second dorsal fin (50 % in adults vs. 100 % in juveniles), caudal fin (88 % vs. 100 %), and second dorsal-fin distal stripe (76 % vs. 100 %); on caudal fin with retained pigmentation remains of up to 6 oblique bars visible on each lobe; in adults 71 % (of 42 fish) show remains of bars on both lobes, in subadults 100 % (= all 13 fish); of all adults only 7 % show no dark pigmentation remains (pigmentation degree 0) and 34 % have dark pigmentation in all body and fin regions (pigmentation degree 4) vs. 60 % in adults. Unpigmented areas of caudal and second dorsal fins and other fins pale hyaline. Etymology. The name refers to the occurrence of oblique conspicuous bars which cross both caudal-fin lobes entirely. Distribution, habitat and size. SW Pacific, NE Australia, Queensland, from W of Thursday Island and S of Daru Island, Torres Strait to NE of Gladstone, Great Barrier Reef; maximum depth 60 m, commonly trawled above soft bottoms at between 15 and 40 m; maximum size 12.5 cm SL. Intraspecific comparisons. Subadults differ from adults in having a slightly longer head, shallower pelvic-fin depth and longer pelvic fins (Tables 2, 9; Figure 8); they differ in having fewer bars on the caudal fin (only preserved fish could be compared; Tables 2, 9); dark pigmentation is more often found in preserved subadults than in preserved adults (Tables 2, 9; see also preserved colour description above). Remarks. Because of the relatively small distribution area of the species and low numbers of specimens available from the most distant localities, no quantitative population comparisons could be conducted. In two of the 55 studied specimens the tiny first-dorsal fin spine is overgrown with skin or very small and hence difficult to detect. Though this species appears to be rather common on shallow soft bottoms near coral reefs, we could not find any in-situ fresh-colour photo documentation., Published as part of Uiblein, Franz, Gledhill, Daniel C., Pavlov, Dimitri A., Hoang, Tuan Anh & Shaheen, Shaker, 2019, Three new goatfishes of the genus Upeneus (Mullidae) from the Indo-Pacific, with a redescription of colour patterns in U. margarethae, pp. 151-196 in Zootaxa 4683 (2) on pages 168-171, DOI: 10.11646/zootaxa.4683.2.1, http://zenodo.org/record/3772664

Upeneus gubal n. sp. Uiblein Gubal goatfish (Figures 1, 5, 7; Tables 1, 5, 6) Holotype. MNHN 2011-1515, adult, 86 mm SL, WIO, Red Sea, Gulf of Suez, Strait of Gubal, N of Gubal Island, 27° 49’ 59” N, 33° 43’ 1” E, 70 m depth, 28 Dec. 1928, collector Robert P. Dollfus. Diagnosis. Dorsal fins VIII + 9, the first spine minute; pectoral fins 15; gill rakers 6 + 19 = 25; lateral-line scales 28–29; measurements in % SL: body depth at first dorsal-fin origin 24; body depth at anus 21; caudal-peduncle depth 11; caudal-peduncle width 3.3; maximum head depth 20; head depth through eye 16; suborbital depth 9.0; interorbital length 8.2; head length 29; snout length 11; postorbital length 11; orbit length 7.4; upper-jaw length 11; barbel length 19; interdorsal distance 16; caudal-peduncle length 24; caudal-fin length 31; anal-fin height 19; pelvic-fin length 22; pectoral-fin length 22; pectoral-fin width 4.8; first dorsal-fin height 22; second dorsal-fin height 19; preserved colour pale brown. Description. Measurements in % SL and counts are given in Table 1; morphometric data as ratios of SL for holotype: body elongate, body depth at first dorsal-fin origin 4.2, body depth at anal-fin origin 4.8, head length 3.4, larger than maximum body depth and subequal to caudal-fin length (3.2); second dorsal-fin height 5.2, shallower than first dorsal fin (4.4) and similar to barbel length (5.4); pelvic-fin length 4.5, equal to length of pectoral fins (4.5) and similar to body depth at first dorsal-fin origin; caudal-peduncle depth 9.2, much larger than orbit length (14); and caudal-peduncle width 30, much narrower than pectoral-fin width (21). Colour. Preserved fish. (Figure 7 D). Body and head entirely pale brown, with brown patches or blotches on head below and behind eyes and on anterior ventral side of body; barbels pale creamy; no remains of dark pigmentation deriving from caudal-fin bars, dark mid-lateral dots, saddle behind second dorsal fin, or second dorsal-fin distal stripe; fins pale brown, partly hyaline. ......continued on the next page ......continued on the next page ......continued on the next page Etymology. Named after the type locality which is situated close to Gubal Island and in the Strait of Gubal at the southern entrance to the Gulf of Suez, Northern Red Sea. The name “ gubal ” is used as a noun in apposition. Distribution, habitat and size. Single locality off Gubal Island, S Gulf of Suez, Northern Red Sea; fish caught by trawling at 70 m depth, most probably on sandy or muddy bottom; size of the specimen is 8.7 cm SL. Remarks. Fresh colour of Upeneus gubal n. sp. is yet unknown, but the available morphological evidence strongly suggests its inclusion in the margarethae group (see also the Interspecific comparisons section further below). The single type specimen was collected together with a specimen of Upeneus moluccensis (Bleeker, 1855) (registered as MNHN 2011-0093). The latter did not show any remains of dark pigmentation e.g. of first dorsal-fin tip and oblique bars on upper caudal-fin lobe as indicated to be diagnostic for preserved conspecifics (Uiblein & Heemstra 2010). The single known locality of U. gubal n. sp. in the S Gulf of Suez is about 1100 km north of the northernmost currently known occurrence of U. margarethae off Ibn Abbas Island, S coast of Sudan.

Upeneus randalli Uiblein & Heemstra, 2011 Randall���s goatfish (Figures 1, 2, 5; Tables 2, 5, 6, 9) Upeneus randalli Uiblein & Heemstra, 2011. Type locality off S Kuwait, Arabian/Persian Gulf, Kuwait. Holotype. Arabian / Persian Gulf: BPBM 33180, HT, adult, 101 mm SL, Arabian / Persian Gulf, Kuwait, off S Kuwait, 29˚00��� N, 48˚25��� E, RV Bahith, trawl, 15���20 m depth (fresh-colour photo) Paratypes (7 adults: 66���106 mm SL; 1 fresh colour photo, 1 fresh-colour drawing): Arabian / Persian Gulf: BPBM 21201, 6: 66-88 mm SL, Bahrain, fish market (fresh-colour photo of 80 mm SL fish); Gulf of Oman: ZMUC P 49161, 106 mm SL, S Iran, Chahabar (fresh-colour drawing in Blegvad & L��ppenthin (1944)) Non-type: Arabian/Persian Gulf: BPBM 29498, subadult, 60 mm SL, Bahrain (fresh-colour photo) Diagnosis. Dorsal fins VIII + 9, the first spine minute; pectoral fins 13���14; gill rakers 6���7 + 17���19 = 23���25; lateral-line scales 28���30; measurements in % SL, adults: body depth at first dorsal-fin origin 23���24; body depth at anus 19���22; caudal-peduncle depth 9.9���11; caudal-peduncle width 2.9���3.8; maximum head depth 19���21; head depth through eye 15���16; suborbital depth 8.0���10; interorbital length 7.2���8.8; head length 27���30; snout length 11���12; postorbital length 10���12; orbit length 6.2���7.4; upper-jaw length 10���11; barbel length 16���20; interdorsal distance 13���15; caudal-peduncle length 23���25; caudal-fin length 27���30; anal-fin height 15���19; pelvic-fin length 20���22; pectoral-fin length 20���22; pectoral-fin width 3.8���4.4; first dorsal-fin height 19���22; second dorsal-fin height 16���20; measurements in % SL, subadults (based on single specimen): body depth at first dorsal-fin origin 22; body depth at anus 19; caudal-peduncle depth 9.0; caudal-peduncle width 3.5; maximum head depth 18; head depth through eye 16; suborbital depth 7.0; interorbital length 7.7; head length 28; snout length 12; postorbital length 10; orbit length 8.2; upper-jaw length 11; barbel length 19; interdorsal distance 14; caudal-peduncle length 23; caudalfin length 29; anal-fin height 16; pelvic-fin length 21; pectoral-fin length 20; pectoral-fin width 3.7; first dorsal-fin height 21; second dorsal-fin height 19; fresh colour: head and body dorsally red-brown red or grey mottled, ventrally white or rose, in some fish with tiny red or brown dots; caudal fin with 11���13 (9 in subadult) brown or darkgrey oblique bars crossing both lobes; on upper caudal-fin lobe 4 or 5 bars, on lower lobe 5���8 bars; lower caudal lobe may be longitudinally crossed by a dark grey stripe (only observed in single subadult); caudal-fin bars and hyaline interspaces of about pupil width in adults; in subadult hyaline interspaces on upper lobe and bars on lower lobe wider (based on single subadults); lower caudal-fin lobe tip sometimes black; barbels white; a single yellow, pale beige or pale red mid-lateral body stripe of less than pupil width from behind head to caudal peduncle, the stripe covered by 3 sections with horizontal series of 2���4 dark brown or black dots, the posteriormost series of 2 dots behind second dorsal-fin base connecting to a dark saddle placed behind second dorsal fin; first dorsal fin with closely fused pale-red brown or pale-grey stripes; the tip region of first dorsal fin without dark pigmentation; second dorsal fin with 2���3 well-separated, brown or black stripes with hyaline interspaces, the dorsal-most stripe the largest and darkest; preserved fish brown or pale brown, mid-lateral body stripe completely lost, often with remains of dark saddle and���more rarely���other dark pigmentation patterns. Distribution, habitat and size. Arabian/Persian Gulf and inner Gulf of Oman: depth 15���20 m; maximum size 11 cm SL. Sources: 1 Uiblein & Heemstra (2010); 2 Uiblein & Causse (2013); 3 Uiblein & Gouws (2015); 4 Uiblein et al. (2016); 5 Uiblein et al. (2017); 6 current study; 7 Upeneus sanctaehelenae Bauchot, 1966 is here treated tentatively as a junior synonym of U. sulphureus. Intraspecific comparison. The single subadult differs from adults in shallower maximum body and head depth, shallower caudal peduncle, shallower suborbital depth, larger eyes, longer anal-fin base, slightly narrower pectoral fins and wider bars on caudal fin, especially on lower fin lobe which is crossed by a dark stripe at mid of lobe (Tables 2, 9; Figure 2 G, H)., Published as part of Uiblein, Franz, Gledhill, Daniel C., Pavlov, Dimitri A., Hoang, Tuan Anh & Shaheen, Shaker, 2019, Three new goatfishes of the genus Upeneus (Mullidae) from the Indo-Pacific, with a redescription of colour patterns in U. margarethae, pp. 151-196 in Zootaxa 4683 (2) on pages 188-190, DOI: 10.11646/zootaxa.4683.2.1, http://zenodo.org/record/3772664, {"references":["Uiblein, F. & Heemstra, P. C. 2011. Description of a new goatfish species, Upeneus randalli sp. nov (Mullidae), from the Persian Gulf, with remarks on and keys for Western Indian Ocean Upeneus species. Scientia Marina, 75, 585 - 94. https: // doi. org / 10.3989 / scimar. 2011.75 n 3585","Uiblein, F. & Heemstra, P. C. (2010) A taxonomic review of the Western Indian Ocean goatfishes of the genus Upeneus (family Mullidae) with descriptions of four new species. Smithiana Bulletin, 11, 35 - 71.","Randall, J. E. & Kulbicki, M. (2006) A review of the goatfishes of the genus Upeneus (Perciformes: Mullidae) from New Caledonia and the Chesterfield Bank, with a new species, and four new records. Zoological Studies, 45, 298 - 307.","Lachner, E. A. (1954) A revision of the goatfish genus Upeneus with descriptions of two new species. Proceedings of the United States National Museum, 103, 497 - 532. https: // doi. org / 10.5479 / si. 00963801.103 - 3330.497","Kim, B. J. & Nakaya, K. (2002) Upeneus australiae, a new goatfish (Mullidae: Perciformes) from Australia. Ichthyological Research, 49, 128 - 32. https: // doi. org / 10.1007 / s 102280200016","Uiblein, F., Gledhill, D. C. & Peristiwady, T. (2017) Two new goatfishes of the genus Upeneus (Mullidae) from Australia and Indonesia. Zootaxa, 4318 (2), 295 - 311. https: // doi. org / 10.11646 / zootaxa. 4318.2.4","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","Uiblein, F. & White, W. T. (2015) A new goatfish of the genus Upeneus (Mullidae) from Lombok, Indonesia and first verified record of U. asymmetricus for the Indian Ocean. Zootaxa, 3980 (1), 51 - 66. https: // doi. org / 10.11646 / zootaxa. 3980.1.3","Uiblein, F. & Lisher, M. (2013) A new goatfish of the genus Upeneus (Mullidae) from Angoche, northern Mozambique. Zootaxa, 3717 (1), 85 - 95. https: // doi. org / 10.11646 / zootaxa. 3717.1.7","Uiblein, F. & Gledhill, D. C. (2015) A new goatfish of the genus Upeneus (Mullidae) from Australia and Vanuatu, with inter- and intraspecific comparisons. Marine Biology Research, 11, 475 - 491. https: // doi. org / 10.1080 / 17451000.2014.958088","Uiblein, F. & Gouws, G. (2015) Distinction and relatedness - taxonomic and genetic studies reveal a new species group of goatfishes (Upeneus; Mullidae). Marine Biology Research, 11, 1021 - 1042. https: // doi. org / 10.1080 / 17451000.2015.1064963","Uiblein, F., Gouws G., Gledhill, D. C. & Stone, K. (2016). Just off the beach: intrageneric distinctiveness of the bandtail goatfish Upeneus taeniopterus (Mullidae) based on a comprehensive alpha taxonomy and barcoding approach. Marine Biology Research, 12, 675 - 694. https: // doi. org / 10.1080 / 17451000.2016.1190458"]}

Upeneus margarethae Uiblein & Heemstra, 2010 Margaretha’s goatfish (Figures 1–6; Tables 1–7) Upeneus margarethae Uiblein & Heemstra, 2010. Type locality off Beira, Mozambique, SW Indian Ocean. Upeneus margarethae: Uiblein & Heemstra 2011 (fresh colour photo of holotype); Bogorodsky et al. 2014 (S Saudi Arabia, Red Sea). Upeneus luzonius non Jordan & Seale, 1907: Ray et al. 2016 (fresh-colour photo from the N Bay of Bengal, E India, EIO). Upeneus tragula Richardson, 1846: Sainsbury et al. 1985 (vouchered fresh-colour photo from NW Australia, EIO). Upeneus tragula non Richardson, 1846: Ratmuangkhwang 2018 (fresh-colour photo from W Thailand, Andaman Sea, EIO). Holotype. SAIAB 82217, male, 82 mm SL, WIO, Mozambique, off Beira, 19˚56.17’ S, 35˚47.16’ E, RV Dr. F. Nansen, bottom trawl, 47 m depth; collectors Phil & Elaine Heemstra, 21 Oct 2007 (fresh-colour photo). Paratypes. (20 adults: 67–124 mm SL; 1 fresh-colour photo). WIO, Somalia: USNM 396092, PT, 124 mm SL, off Bargaal, 11°14’ N, 51°08’ E; Kenya: SAIAB 82817, 5: 89–110 mm SL, off Kipini, 02°38’ S, 40°28’ E, RV Dr. F. Nansen, bottom trawl, 11–12 m depth; Mozambique: SAIAB 82209, 89 mm SL, off Beira, 19°47.45’ S, 35°30.73’ E, RV Dr. F. Nansen, bottom trawl, 28 m depth; SAIAB 82814, 2: 89–97 mm SL, same collecting data as holotype (fresh-colour photo); Madagascar: SAIAB 82815, 11: 67–94 mm SL, Tsimipaika Bay, 8–12 m depth. Non-types. (66 adults, 11 subadults: 40–129 mm SL, 14 fresh-colour photos): WIO proper: Kenya: SAIAB 82825, 12: 40-95 mm SL, same collection data as PT SAIAB 82817; Tanzania: BPBM 33441, 100 mm SL, Mafia Island, Mafia Channel, 10–50 m depth; SAIAB 87108, 112 mm SL, Zanzibar, Mazizini, landing site adjacent to Zanzibar Beach Resort; Mtwara, Mikindani landing site (each with fresh-colour photo): HIFIRE F 58449, 79 mm SL; HIFIRE F 58450, 70 mm SL; SAIAB 104431, 77 mm SL; Zanzibar, Stone Town, landing site and fish market (each with fresh-colour photo): HIFIRE F 58451, 90 mm SL; HIFIRE F 58452, 87 mm SL; HIFIRE F 5453, 84 mm SL; HIFIRE F 58454, 85 mm SL, HIFIRE F 58455, 95 mm SL; SAIAB 104432, 3: 90–114 mm SL; Mozambique: SAIAB 81741, 128 mm SL, off Machangulo, 26°10’ S, 32°59’ E, 45 m depth; SAIAB 86466, 2: 75-84 mm SL, off Beira, 19°47.44’S, 35°30.72’E, RV Dr. F. Nansen, bottom trawl, 28-29 m depth; Madagascar: MNHN 1966-881, 90 mm SL, no locality information; SW-India, Kerala State: BPBM27694, 2, 71-82 mm SL, Vizhinjam fishing harbor; Sri Lanka: USNM 396094, 78– 79 mm SL, Colombo fish market, landed at Eravur; WIO, Red Sea: Sudan: BMNH 1960.3.15.841, 63 mm SL, Ibn Abbas Island, 5 km S of island, FRV Manihine; Saudi Arabia, off Jizan, FV Attiat Alrahman 2, bottom trawl: KAUMM 27, 90 mm SL, 16° 44.32’ N, 42° 26.941’ E, 42 m depth; KAUMM 49, 5: 96–113 mm SL, 16° 53.621’ N, 42° 23.633’ E, 25.5-28 m depth; KAUMM 59, 1 of 5: 109 mm SL, 16° 44.508’ N, 42° 29.497’ E, 18.5-21.5 m depth; KAUMM 109, 2 of 11: 67–91 mm SL, 16° 54.869’ N, 42° 26.044’ E, 21 m depth; SMF 34972, 4: 93–111 mm SL, 16° 53.621’ N, 42° 23.633’ E, 25.5-28 m depth; SMF 35011, 3: 90–116 mm SL, 16° 44.508’ N, 42° 29.497’ E, 18.5-21.5 m depth; SMF 35030, 90 mm SL, 16° 57.873’ N, 42° 25.523’ E, 16–19 m depth (fresh-colour photo); SMF 35059, 91 mm SL, 16° 54.869’ N, 42° 26.044’ E, 21 m depth; SMF 35060, 91 mm SL, 16° 44.32’ N, 42° 26.941’ E, 42 m depth; Eritrea: MNHN1984-455, 2: 93–94 mm SL, RV Dr. F. Nansen, bottom trawl (possibly off Assab, 13°21’ N, 42°25’ E, 30 m depth); EIO: Andaman Sea: Myanmar (new record): SAIAB 203672, 110 mm SL, S of Buda Island, 10°19.43’ N, 98°14.69’ E, RV Dr. F. Nansen, bottom trawl, 104 m (freshcolour photo); SAIAB 203480, 95 mm SL, NW of Bashuhino Island, 14°40.29’ N, 97°15.93’ E, RV Dr. F. Nansen, bottom trawl, 57 m depth; Thailand, Phuket: ZMUC P49560, 85 mm SL; NW Australian shelf: AMS 22831 -021, 3 (of 6): 81-97 mm SL, 140 km W of Port Hedland, 20°00’S, 117°16’E, 50 m depth; BMNH 1983 5.5.28-30, 3: 108–117 mm SL, off Port Hedland, 20°10’ S, 118°25’ E; CSIRO CA 3052, 98 mm SL, off Port Hedland, 20°03.8’S, 117°57.2’E, 38–39 m depth (fresh-colour photo); WAM 25397.004, 3 (of 7): 94–102 mm SL, Rowley Shoals, 17°29’S, 121°52’E, 42 m depth; Exmouth Gulf: WAM 23785 - 6, 100 mm SL; WAM 32680 -002, 93 mm SL, Exmouth Gulf, Bundegi Reef, 21°50.879’S, 114°15.530’E, 24 m depth; W Pacific: N Australia, Northern Territory, Arafura Sea: AMS I.21849-009, 6: 95–101 mm SL, off Arnhem Land, 11°29’ S, 134°23’ E, RV Soela, Engel trawl, 105 m depth; Queensland, Gulf of Carpentaria (new record): CSIRO A 2881, 95 mm SL, E of Mornington Island, 16°35.7’ S, 140°41.6’ E, FV Rama, prawn trawl, 18 m depth. Fresh-colour photos of specimens not retained (n =16): WIO proper: subadult or adult, Pemba Bay, near Pemba harbour, N Mozambique (R. Koch); subadult or adult, Pomene, South-central Mozambique (M. & V. Fraser); adult, Lunene Island, off Vilanculos, South-central Mozambique (A. Lund); adult and subadult, Tanzania, Mafia Channel (J.E. Randall); adult, Tuticorin, S India (K.K. Bineesh); Red Sea: 3 adults, Jizan, Saudi Arabia (S. Bogorodsky); EIO: adult, Digha Morona, West Bengal, E India (Ray et al. 2016); 4 adults, Puri, Odisha, E India (S. K. Roul); subadult, Kampuan mangrove forest, Amphoe Suksamran, Ranong, Thailand (S. Ratmuangkhwang); adult, off NW Australia, unknown locality, RV Soelae (CSIRO staff). Diagnosis. Dorsal fins VIII + 9, the first spine minute; pectoral fins 13–15; gill rakers 5–7 + 15–18 = 21–24; lateral-line scales 28–30; measurements in % SL, adults: body depth at first dorsal-fin origin 22–27; body depth at anus 20–24; caudal-peduncle depth 9.3–12; caudal-peduncle width 3.5–5.7; maximum head depth 18–23; head depth through eye 15–18; suborbital depth 8.3–12; interorbital length 7.3–9.1; head length 27–31; snout length 9.7–12; postorbital length 10–13; orbit length 6.2–9.1; upper-jaw length 9.8–12; barbel length 15–20; interdorsal distance 12–16; caudal-peduncle length 22–26; caudal-fin length 27–31; anal-fin height 14–18; pelvic-fin length 20–24; pectoral-fin length 19–24; pectoral-fin width 4.1–5.3; first dorsal-fin height 18–23; second dorsal-fin height 15–20; subadults: body depth at first dorsal-fin origin 24–26; body depth at anus 21–23; caudal-peduncle depth 10–11; caudal-peduncle width 3.4–4.4; maximum head depth 19–22; head depth through eye 15–17; suborbital depth 7.6–9.4; interorbital length 7.6–8.1; head length 28–32; snout length 9.9–12; postorbital length 11–13; orbit length 7.5–9.0; upper-jaw length 11–12; barbel length 17–18; interdorsal distance 14–17; caudal-peduncle length 24–26; caudal-fin length 29–33; anal-fin height 17–19; pelvic-fin length 22–24; pectoral-fin length 21–23; pectoral-fin width 4.3–5.1; first dorsal-fin height 21–24; second dorsal-fin height 19–21; fresh colour: head and body dorsally red, brown or pale red-brown mottled, ventrally white, often with pale red brown dots or larger red blotches (the latter together with dark vertical bands in inactive, live fish); caudal fin with maximally 15 red, brown or darkgrey oblique bars on both lobes; on upper caudal-fin lobe 4 or 5 (rarely 6) bars (3 or 4 bars in subadults), on lower lobe 6–9 bars (3–6 bars in subadults) which are often crossed or covered by a red, brown or dark-grey stripe that may cover the dorsal half of lobe or almost the entire lobe leaving only the ventral margin free; caudal-fin bars and hyaline interspaces of about pupil width in adults; lower caudal lobe tip sometimes black; barbels entirely white or mostly white, intermingled with pale rose; a single yellow, beige, or orange mid-lateral body stripe of pupil width from snout tip through eye to caudal-fin base; stripe sometimes covered by 1–4 sections with horizontal series of 2–4 dark brown or black dots, the posteriormost series of dots behind second dorsal-fin base, often connecting to a dark saddle placed behind second dorsal fin; first dorsal fin with often closely fused pale-red or pale-grey brown stripes; the tip region of first dorsal fin not darker than remainder of fin and often without any pigmentation; the tiny first dorsal-fin spine sometimes dark pigmented, also in preserved fish; second dorsal fin with 3 or 4 red or brown, well-separated stripes with hyaline interspaces; preserved fish entirely brown, pale brown or pale grey, the mid-lateral stripe completely lost, but often with remains of pigmentation deriving from caudal-fin bars, dark mid-lateral dots, a saddle and/or second dorsal-fin distal stripes. ......continued on the next page ......continued on the next page Colour Description. Freshly collected fish. (Figure 2 A-F). Head and body ventrally white or pale brassy, in adult fish (Figure 2 A–E) with numerous tiny red or red-brown dots placed on individual scales and covering up to five vertical scale rows at mid body; in some fish, as in holotype (Figure 2 A) and subadult (Figure 2 F) larger red blotches are formed on ventral side of head and body; head from above snout and eye and body above lateral line red or brown-greyish mottled, bordered below by a mid-lateral stripe of pupil width which runs rather straight or only slightly bent from caudal-fin base to eye in yellow, beige or orange colour, becoming more reddish when crossing pupil (iris entirely or partly red) and bending ventrally towards snout tip; on the mid-lateral body stripe in several adult or subadult specimens of the entire distribution area (Figure 2 B–F) 1–3 sections with horizontal series of 2–4 black, red or brown dots of about half-pupil diameter, the anterior-most section below first dorsal-fin base, the second section below second dorsal-fin base, and the third series of dots just behind second dorsal fin; the latter section is the most frequently encountered (also in preserved specimens) and often connected to a dark saddle of about half of second dorsal-fin base length which reaches from behind second dorsal fin to mid-lateral body stripe; from head behind eye until below base of second dorsal fin, a thin white stripe separates the dorsal body pigmentation from the mid-lateral body stripe; lateral line weakly visible as a series of red or brownish-grey dots placed well above midlateral stripe in anterior half of body starting to cross the stripe just anterior to second dorsal-fin base, continuing behind second dorsal fin just below stripe towards caudal-fin base; barbels entirely white or mostly white, partly intermingled with pale rose pigmentation proximally (Figure 2 E) or distally (Figure 2 C, D); caudal fin of adults entirely crossed by 10–15 red, red-brown or partly darker oblique bars; upper caudal-fin lobe covered by 4–5 (6 in a single large specimen) bars of pupil or slightly narrower width, the distal-most bar covering fin tip in some specimens (Figure 2 B, C–F); bars separated by hyaline interspaces of similar width or slightly wider; lower caudal-fin lobe covered by 6–9 bars of similar width as on upper fin lobe, with hyaline interspaces sometimes slightly narrower; in addition, the lower fin lobe is frequently covered by a red stripe that reaches from base of fin to fin tip, covering the dorsal half of lobe or almost the entire lobe, leaving at least the ventral lobe margin free where the bars and interspaces can be well recognized (Figure 2 A, C, D); in most fish with the stripe pigmentation on lower caudalfin lobe, bars and interspaces can be still seen shining through the entire fin lobe (Figure 2 B, E); furthermore, dark markings of bars can be also sometimes found at dorsal margin of lower caudal-fin lobe, especially distally, where the bars may turn into black close to or on lobe tip (Figure 2 A–E); subadults with 3 brown bars of more irregular form and size on each caudal-fin lobe, the 2 proximal bars on lower lobe nearly fused; first dorsal fin with irregular red or pale red (adults, Figure 2 A–E) or pale brown (subadult, Figure 2 F) pigmentation which does not become darker (adults), or becomes lighter (subadult), towards tip; the tiny first dorsal-fin spine is often dark pigmented; second dorsal fin with 3–4 red stripes of less than orbit width in adults, one stripe at fin base, one at or close to fin tip, the latter being darker than the other stripes in some specimens (Figure 2 C, E); 2 brown second dorsal-fin stripes in subadult, the distal stripe darker (Figure 2 F); pelvic fins hyaline with up to four narrow red stripes; anal fin with or without stripes, pectoral fins hyaline. Live fish in situ. Active fish. (Figure 3 A, B). Head and body ventrally white, with numerous tiny red or grey dots associated with scale rows; head from above snout and eye and body above lateral line pale-grey mottled, bordered ventrally by a mid-lateral body stripe of pupil width which runs rather straight from caudal-fin base to eye in yellow or orange colour, becoming red or brown when crossing pupil (iris pigmented in same colour and in shape of stripe) and bending ventrally towards snout tip; five dark, well separated, rectangular, brown or dark-grey rectangular sections on mid-body stripe, 2 shorter ones behind eye and gill cover, three longer ones each covered by a horizontal series of 2–4 black or dark brown dots of approximate pupil size (or slightly smaller); four dots on the section below first dorsal fin, two dots on the section below anterior part of second dorsal fin and three dots on posterior-most section behind second dorsal-fin base; of the latter three dots the last one is slightly elevated connecting closely to a grey saddle that reaches from dorsal body margin behind second dorsal fin to stripe; lateral line, as also observed in recently deceased fish (see further above), well visible, proceeding parallel above lateral mid-body stripe within first half of body, then crossing and continuing just below stripe to caudal-fin base; barbels white; caudal fin with 6–10 red-brown (Figure 3 B) or black (Figure 3 A) oblique bars; upper caudal-fin lobe covered by 3–4 bars of pupil or slightly narrower width, the distal-most bar covering fin tip; bars separated by hyaline interspaces of similar width or slightly wider, overlain with white pigmentation in middle parts; lower caudal-fin lobe covered by 3–6 bars of either variable (Figure 3 A) or similar (Figure 3 B) size and form; in both photographed specimens a stripe is formed along dorsal two thirds of lower caudal-fin lobe which ends with second bar in one specimen (Figure 3 A) and extends to black tip in the other fish (Figure 3 B); first dorsal fin only weakly pale grey or pale brown pigmented, without dark-pigmented tip region (in contrast to U. heemstra; Figure 3 B); second dorsal fin with three pale grey or pale brown to brown or dark-brown stripes, the distal two stripes more intensely coloured than the rather weak stripe at or close to fin base; interspaces between stripes in part white pigmented; pelvic and anal fins partly hyaline with weak pale-grey or pale-brown stripes and whitish interspaces and margin, pectoral fins hyaline, only faintly visible. Inactive, “resting” fish (Figure 3C; only differences from the above colour description are indicated). Head and body ventrally with at least 25 irregularly formed and distributed, mostly rounded red to dark-red blotches of about orbit size or smaller, extending from below eye to caudal peduncle; head from above snout and eye and body above lateral line pale brown, the latter with 5 oblique brown or dark-brown bands overlapping with the dark-brown rectangular sections on orange mid-lateral body stripe (see also above); the second to fourth band connect in part to dark-red blotches on ventral body side; behind, one weaker coloured and more isolated brown band is placed at posterior end of mid-lateral body stripe, connecting to the posterior-most red blotch ventrally; caudal fin with 5 pale red to pale brown oblique bars on dorsal lobe and at least 5 red bars on ventral lobe (lobe tip appears to be broken); no stripe apparent on lower caudal-fin lobe; dark pigmentation of first minute spine on first dorsal fin well visible. Preserved fish. Head and body mostly uniformly pale brown greyish brown or brown, sometimes slightly darker dorsally, gill cover silvery or pale and partly transparent, lateral line, when intact, well visible in entire range from behind head to caudal-fin base; barbels pale brown or pale creamy; mid-lateral body stripe completely lacking, but often remains of dark dots in the area of stripe below and posterior to dorsal fins (in 24 % of 87 adults vs. 73 % of 11 subadults); dark pigmentation remains occur also in other areas: saddle behind second dorsal fin (60 % adults vs. 82 % subadults), caudal fin (66 % vs. 100 %), and second dorsal-fin distal stripe (55 % vs. 90 %); on caudal fin with retained pigmentation remains of up to 6 oblique bars visible on upper and up to 8 bars on lower lobe visible; in adults 55 % (of 87 fish) show remains of bars on both caudal-fin lobes, in subadults 82 % (of 11 fish); of all adults 19 % (= 16 of 86 fish) show no pigmentation remains (pigmentation degree 0), while a pigmentation degree of 4 (remains of all four colour characters retained) is found in 21 % (= 18 of 86 fish). All subadults do show dark pigmentation, with pigmentation degree 4 occuring in 70 % (7 of 10 individuals). Unpigmented areas of caudal and second dorsal fins and other fins entirely pale and partly hyaline. Distribution, habitat and size. WIO: East African coast from S Mozambique and Madagascar to Central Red Sea (northernmost record from off Ibn Abas Island, S Sudan), W and S coast of India from Kerala and Tamil Nadu to Sri Lanka; EIO: E coast of India, Odisha and West Bengal, Andaman Sea off S Myanmar (new record) and W Thailand; NW Australian coast and shelf from Exmouth Gulf to Rowley Shoals; Arafura Sea, Northern Territory, and inner Gulf of Carpentaria, Queensland (new record) (Figure 1); depth from 8 m (or possibly shallower) to 105 m, commonly trawled above soft bottoms at between 20 and 30 m depth, encountered during diving on shallow sandy bottoms (Figure 3); maximum size 13 cm SL. Intraspecific Comparisons. Among the two size classes, main differences occur in both morphology and colour patterns. Subadults differ from adults in having slightly fewer rudimentary gill rakers on lower limb (Figure 4), while total gill-raker numbers on both limbs do not change, a slightly shallower suborbital depth, longer caudal fin, higher anal fin and higher second do

Neobythites kenyaensis, a cuskeel (Ophidiidae) with two ocelli on the dorsal fin, is redescribed based on 33 meristic, body shape, colour, and otolith characters obtained from 15 specimens, 12 of which were studied for the first time. Nine of the 12 new specimens were collected during a research cruise off Mozambique together with vouchered colour photographs made on board when the fish were still fresh. Two specimens were collected off Natal, South Africa, one of them only available as a fresh colour photograph and one specimen off Dar es Salaam, Tanzania. Accordingly, while N. kenyaensis was only known from Kenya and northernmost Tanzania (Zanzibar) since its description in 1995, we proved now its continuous distribution along the entire SE African coast from off Ras Ngomeni, Kenya, to off Durban, South Africa, at depths of 238-457 m. Neobythites kenyaensis is rather unique among the 54 species of its genus in possessing an ocellus positioned near the dorsal-fin origin (anterior ocellus) which it shares only with the West-Pacific N. longiventralis and West-Atlantic N. ocellatus. It differs from the latter in the following characteristics: second, central ocellus more posteriorly placed on dorsal fin, more precaudal vertebrae (13 vs. 12), less developed gill rakers (8-10 vs. 14-16), shorter head (19.5-20.5 vs. 21.5-23.5% SL), and shorter pelvic fins (9.9-12.0 vs. 16.0-21.5% SL). Both species differ clearly from N. longiventralis in having a much shorter pelvin fin (9.9-21.5 vs. 32.0-34.% SL). The central ocellus-spot size, determined by the number of rays covered by the ocellus spot, varies considerably in N. kenyaensis and reaches the largest size reported for any ocellus-bearing Neobythites species. This character is strongly and positively correlated with both latitude and depth, suggesting gradual differentiation processes among adjacent populations or subpopulations. Another finding reported for the first time in Ophidiidae is the significant correlation of three otolith measurements with fish size (SL) in N. kenyaensis.

Neobythites malhaensis Nielsen, 1995 (Figures 1���2, Tables 1, 3) Neobythites malhaensis Nielsen, 1995: 6, fig. 5a. Holotype. ZM MGU P-18915 (male, 123 mm SL), Saya de Malha Bank, Western Indian Ocean, 11��02' S, 62��15' E, 250 m depth (probably same cruise as paratypes). Paratypes (n = 3; 117���135 mm SL). Saya de Malha Bank, Western Indian Ocean: ZM MGU P-18916 (female, 117 mm SL), 11��08' S, 62��16' E, RV Professor Mesiatzev, trawl 476, 235 ��� 239 m depth, 7 Oct 1977; ZM MGU P- 18917 (female, 132 mm SL), 11��06' S, 62��19' E, RV Professor Mesiatzev, trawl 478, 240 m depth, 7 Oct 1977; ZMUC P77840 (female, 135 mm SL), RV Fiolent, 1973 (no other data). Diagnosis. Indistinct or flat spine on hind margin of preopercle; dorsal fin-rays 99���103; anal-fin rays 78���82; pectoral fin-rays 30; precaudal vertebrae 13; total vertebrae 57���59; pseudobranchial filaments 3���4; long gill rakers on anterior arch 12���13; head length 21.0���21.5 % SL; pelvic-fin length 12.5���13.0 % SL, pelvic fins not reaching anus; orbit length 4.1���4.9 % SL, 19.5���23.0 % head length, and 2.2���2.4 times in upper-jaw length; longest gill filament 1.6���1.9 % SL and 7.5���8.9 % head length; ocellus spot placed behind a vertical line through anus, the ocellus-spot distance being 42.5���45.5 % SL, and the spot covering 13 dorsal-fin rays; dorsal and anal fins not pigmented; one vertical, dark bar on body below ocellus spot; otolith length 5.2 % SL, sulcus length 3.8 % SL, and ostium height 21.5 % sulcus length. Distribution. Saya de Malha Bank, off SE Seychelles, Western Indian Ocean, at 235���250 m depth., Published as part of Uiblein, Franz & Nielsen, J��rgen G., 2018, Review of the steatiticus - species group of the cuskeel genus Neobythites (Ophidiidae) from the Indo-Pacific, with description of two new species, pp. 157-173 in Zootaxa 4387 (1) on pages 164-165, DOI: 10.11646/zootaxa.4387.1.7, http://zenodo.org/record/1186736, {"references":["Nielsen, J. G. (1995) A review of the species of the genus Neobythites (Pisces, Ophidiidae) from the western Indian Ocean with description of seven new species. Ichthyological Bulletin, 62, 1 - 19."]}

Neobythites stefanovi Nielsen & Uiblein, 1993 (Figures 1���3, Tables 1���3) Neobythites stefanovi Nielsen & Uiblein, 1993: 110. Neobythites stefanovi: Nielsen 1995: page 15; Nielsen 2002: 86 (in part); Kannan et al. 2013: 1 (here tentatively referred to N. steatiticus). Holotype. ZM MGU P18923 (male, 166 mm SL), Gulf of Aden, NW Indian Ocean, 14��48.8' N, 51��16.1' E, RV Dmitry Stefanov cr. 3, trawl 91, 245��� 335 m depth; 5 Jan. 1989. Paratypes (n = 25, 42���195+ mm SL). BMHN 1939.5. 24.1437 (female, 195+ mm SL), Gulf of Aden, 13��14.4' N, 46��14.2' E, RV John Murray, st. 35, otter trawl, 457���549 m depth, 16 Oct. 1933; SMF 26427 ���29 (2 females, 2 males, and 4 unsexed; 42���122 mm SL), Gulf of Aden, 12��16' N, 44��09.5' E, RV Meteor, MeS-287, beam-trawl; 472���479 m depth, 16 Mar. 1987; USNM 302930, female, 179 + mm SL, Gulf of Aden, 11��10.6'N, 48��11.2'E, RV Beinta, cr. 19, haul 32, 351 ��� 384 m depth, 19 Feb. 1987; ZM MGU P-18924 (female, 139 mm SL), Gulf of Aden, 13��15.5' N, 46��14.3' E, RV Akademik Petrovsky cr. 14, st. 34, 250 ��� 300 m depth, 26 Sep. 1983; ZM MGU P-18925 (male, 130 mm SL), Gulf of Aden, 15��59.8' N, 52��23.9' E, RV Dmitry Stefanov cr.3, trawl 9, 80 ��� 110 m depth, 19 Nov. 1988; ZM MGU P-18926 (female, 162+ mm SL), Gulf of Aden 14��49.2' N, 50��35.1' E, RV Dmitry Stefanov, trawl 78, 375 m depth, 3 Feb. 1989; ZM MGU P-18927 (female, 168 mm SL), same data as holotype; ZM MGU P- 18928 (female, 180 mm SL), Gulf of Aden, 13��13.7' N, 45��41.8' E, RV Dmitry Stefanov, cr.5, trawl 48, 355 ��� 365 m depth, 22 Nov. 1989; ZM MGU P-18929 (female, 190 mm SL), and ZMUC P77841 (female, 198 mm SL), Gulf of Aden, 13��09' N, 45��41' E, RV Dmitry Stefanov, trawl 51; 460 m; 24 Nov. 1989; ZM MGU P-18930 (female and male, 165���179 mm SL), and ZMUC P77842 (male, 168 mm SL), Gulf of Aden, 13��25.1' N 47��14.S' E, RV Dmitry Stefanov, trawl 68, 367 m depth, 28 Nov. 1989; ZM MGU P 18931 (female, male, and unsexed, 130���172 mm SL), and ZMUC P77843 ���44 (male and female, 163���166 mm SL), Gulf of Aden, RV Dmitry Stefanov, 1989. Non-types (n = 22, 29��� 122 mm SL). SMF 26430 (female, 122 mm SL), Red Sea, 18��35.3' N, 39��03.5' E, RV Meteor, MeS-171, beam trawl, 434���469 m depth, 23 Feb. 1987; SMF 26431 and 26434 (3 unsexed, 45���94 mm SL), Red Sea, 19��18.2' N, 38��15.5' E, RV Meteor, MeS-194, beam trawl, 537���681 m depth, 28 Feb. 1987; SMF 26432 (2 unsexed, 75���84 mm SL), Red Sea, 22��05' N, 37��09.3' E, RV Meteor, MeS-96, beam trawl, 600 m depth, 9 Feb. 1987; SMF 26433 (2 unsexed, 44���99 mm SL), Red Sea, 19��44.5' N, 37��40.2' E, RV Meteor, MeS-148, beam trawl, 517���583 m depth, 20 Feb. 1987; SMF 26435 and 26440 (2 unsexed, 49��� 31 mm SL), Red Sea, 21��28.97' N, 38��15.55' E, RV Valdivia, st. 111 TA, closing trawl, 740���785 m depth, 12 Apr. 1979; SMF 26436 and 26441 (3 unsexed, 29���46 mm SL), Red Sea, 21��26.5' N, 38��38.3' E, RV Valdivia, st. 121 TA, closing trawl, 779���801 m depth, 15 Apr. 1979; SMF 26437 and 26785 (2 unsexed, 63���115 mm SL), Red Sea, 20��94.9' N, 37��26.1' E; RV Sonne 35 TA, c1osing trawl, 490���588 m, 17 Oct. 1977; SMF 26438 (42 mm SL), Red Sea, 21��10.8' N, 37��34.0' E, RV Sonne 27 TA, closing trawl, 733���757 m depth, 16 Oct. 1977; SMF 26439 (38 mm SL), Red Sea, 21��12.0' N, 37��26.8' E, RV Sonne, 25 TA, closing trawl, 724���747 m depth, 15 Oct. 1977; SMF 26442 (5 unsexed, 29���42 mm SL), Red Sea, 21��33' N, 38��21' E, RV Valdivia, 99 TA, closing trawl, 753���804 m depth, 9 Apr. 1979. Diagnosis. No spines on hind margin of preopercle; dorsal fin-rays 89���95; anal-fin rays 73���78; pectoral finrays 24���26; precaudal vertebrae 12; total vertebrae 52���56; pseudobranchial filaments 2���6; long gill rakers on anterior arch 10���13; head length 23.0���27.5 % SL; pelvic-fin length 14.5���20.5 % SL, pelvic fins not reaching anus; orbit length 5.4���6.9 % SL, 21.5���27.5 % head length, and 1.7���2.3 times in upper-jaw length; longest gill filament 1.4���3.7 % SL and 5.4���16.0 % head length; ocellus spot placed well behind a vertical line through anus, the ocellusspot distance being 45.5���56 % SL, and the spot covering 8���12 dorsal-fin rays; dorsal and anal fins with dark margins throughout; no vertical bars on body; otolith length 4.6���5.2 % SL, sulcus length 3.4���4.1 % SL, and ostium height 20.0���28.5 % sulcus length. Distribution. Gulf of Aden, NW Indian Ocean, at 80-549 m depth and Red Sea, 434���804 m depth. Remarks. Kannan et al. (2013) reported on two specimens of N. stefanovi from Gulf of Manar, Tamil Nadu (NW Indian Ocean). After inspection of the photo and some of the data provided by Kannan et al. (2013), we assume that these specimens should be treated as tentatively referred to N. steatiticus. Uiblein (1995) reported considerable differences between the populations of the Red Sea and the Gulf of Aden based on quantitative studies of meristic, morphometric and ocellus characters. Those differences were however only statistically detectable and did not support the erection of a distinct taxon in the Red Sea. We have reconsidered these earlier findings when examining the differentiation degree among the three rather similar species N. gloriae n. sp., N. steatiticus, and N. stefanovi. While these species can be well separated by five diagnostic characters (singly or in combination), the two N. stefanovi populations clearly overlap with each other in those as well as in all other characters studied (Figure 3, Tables 2���3)., Published as part of Uiblein, Franz & Nielsen, J��rgen G., 2018, Review of the steatiticus - species group of the cuskeel genus Neobythites (Ophidiidae) from the Indo-Pacific, with description of two new species, pp. 157-173 in Zootaxa 4387 (1) on pages 169-170, DOI: 10.11646/zootaxa.4387.1.7, http://zenodo.org/record/1186736, {"references":["Nielsen, J. G. & Uiblein F. (1993) Tiefenwasser- und Tiefseefische aus dem Roten Meer. XVI. A new species of Neobythites from the NW Indian Ocean and the Red Sea. Senckenbergiana maritima, 23, 109 - 113.","Nielsen, J. G. (1995) A review of the species of the genus Neobythites (Pisces, Ophidiidae) from the western Indian Ocean with description of seven new species. Ichthyological Bulletin, 62, 1 - 19.","Nielsen, J. G. (2002) Revision of the Indo-Pacific species of Neobythites (Teleostei, Ophidiidae) with 15 new species. Galathea Report, 19, 5 - 105.","Kannan, K., John, S., Joshi, K. K. & Zacharia, P. U. (2013) First record of Neobythites stefanovi (Teleostei: Ophidiidae) from Indian waters. Marine Biodiversity Records, 6, e 71. https: // doi. org / 10.1017 / S 1755267213000523","Uiblein, F. (1995) Morphological variability between populations of Neobythites stefanovi (Pisces: Ophidiidae) from the deep Red Sea and the Gulf of Aden. Marine Ecology Progress Series, 124, 23 - 29. https: // doi. org / 10.3354 / meps 124023"]}

Uiblein, Franz, Nielsen, Jørgen G. (2018): Review of the steatiticus - species group of the cuskeel genus Neobythites (Ophidiidae) from the Indo-Pacific, with description of two new species. Zootaxa 4387 (1): 157-173, DOI: https://doi.org/10.11646/zootaxa.4387.1.7

Neobythites lombokensis n. sp. (Figures 1���2, Tables 1, 3) Neobythites meteori: Nielsen 2002: 57, fig. 47. Neobythites sp. 2: Gloerfelt-Tarp & Kailola 1984: 88 ���89 (colour photo). Holotype. ZMUC P77744 (93 mm SL), SE of Lombok Island, Indonesia, Eastern Indian Ocean, 8��58���S, 116��34��� E, JETINDOFISH, sta. TGT 1717, bottom trawl, 150���280 m depth, July 1981. Diagnosis. No spines on hind margin of preopercle; dorsal fin-rays 91; anal-fin rays 72; pectoral fin-rays 29; precaudal vertebrae 13; total vertebrae 53; pseudobranchial filaments 3; long gill rakers on anterior arch 7; head length 23.0 % SL; pelvic-fin length 13.0 % SL, pelvic fins not reaching beyond anus; orbit length 5.2 % SL, 22.5 % head length, and 53 % upper-jaw length; longest gill filament 2.2 % SL and 9.4 % head length; ocellus spot originating just behind at a vertical line through anus, the ocellus-spot distance being 43.0 % SL, and the spot covering 14 dorsal-fin rays; dorsal and anal fins not or only patchily pigmented; no vertical bars on body; otolith length 4.8 % SL, sulcus length 3.2 % SL, and ostium height 26.0 % sulcus length. Description. Body rather elongate; complete lateral line running close to dorsal edge; head and body covered with cycloid scales; origin of dorsal fin well behind opercle; origin of anal fin slightly anterior to midpoint of fish; tip of pectoral fin well in front of anal fin origin; pelvic fins reaching halfway to origin of anal fin; blunt snout shorter than diameter of eye; upper jaw ends below posterior half of eye; anterior nostril with low rim and larger posterior nostril a mere hole; spine absent on posterior margin of preopercle; opercular spine strong and straight; anterior gill arch with 2���3 short and 1 long raker on upper branch, 1 long raker in angle and lower branch with 5 long and 6 short rakers; gill filaments twice the length of the long rakers; three distinct pseudobranchial filaments. Otolith (Nielsen 2002, fig. 48B). Otolith oval, its height 1.5 times in its length; sulcus large, 1.5 times in otolith length; ostium length 1.7 times in sulcus length; ostium height 2.3 in ostium length. Dentition. Premaxillaries and dentaries with irregular tooth rows, teeth on outer rows needle-like. Blunt teeth on subtriangular vomer and palatines. Two median basibranchial tooth patches, the anterior oblong and posterior smaller and circular. Axial skeleton (from radiograph). Precaudal 13 and caudal vertebrae 40; tip of all neural and haemal spines thin and pointed; first neural spine half length of second spine; vertebrae 3���5 with slightly depressed neural spines; bases of vertebral spines 5���8 enlarged; parapophyses present on posterior seven precaudal vertebrae; pleural ribs on vertebrae 3���13; epipleural ribs not observed. Colour (fresh fish; Figure 1B 2). Body mottled pale-brown to brown, becoming paler posteriorly, abdomen whitish blue, lateral line indistinct; head brown dorsally and pale brown below and behind eye, gill cover pale; ocellus with large black spot, its horizontal diameter more than twice of orbit length, placed above anterior part of anal fin; posterior parts of dorsal fin and nearly entire anal fin patchily covered with pale-brown or brown pigment; no margin or band on anal fin. Colour (preserved fish; holotype after 36 years of preservation, Figure 1B1). Body and head mottled brown, abdomen and eyes bluish, lateral line indistinct; gill cover transparent, inside of peritoneum with many black spots of various sizes; ocellus spot dark-brown; patchily distributed traces of brown pigmentation on dorsal and anal fins. Etymology. Named after the type locality, SE off Lombok Island, southern Indonesia. Distribution. Single type specimen trawled SE of Lombok Island, Indonesia, Eastern Indian Ocean, at 150��� 280 m depth. Biology. The holotype is an unripe female caught between the lower shelf and upper continental slope. Its intestinal contents were unidentifiable except for parts of a small bivalve, suggesting a benthic lifestyle. Comparisons. Neobythites lombokensis n. sp. differs from all congeners in the following combination of characters: a single ocellus placed on dorsal fin at mid-body, no preopercular spines, 7 developed gill rakers, head length 23.0 % SL, gill filament length 2.2 % SL, orbit length 22.5 % head length and 1.9 times in upper-jaw length, and otolith sulcus length 3.2 % SL. Neobythites lombokensis seems most similar to N. meteori, sharing the following characters: dorsal fin with one ocellus, preopercular spine absent, pelvic fins ending anterior to anus, teeth needle-like and less than eight long rakers on anterior gill arch. Neobythites lombokensis differs from N. meteori in having a larger ocellus spot (spot covering 14 vs. 9 dorsal-fin rays), the ocellus placed above anterior part of anal fin (vs. above anus), and shorter gill filaments (9.4 vs. 13.0 % head length), respectively. Remarks. In a review of the genus Neobythites from the Western Indian Ocean, Nielsen (1995) described seven new species. One of these, N. meteori, was based on a specimen from off Socotra Is. Furthermore, a specimen from off Lombok was referred to N. meteori but not designated as a paratype because of a number of differences from the Socotra specimen. Subsequent examination of the Lombok specimen showed that it represented an undescribed species of Neobythites, which is herein described., Published as part of Uiblein, Franz & Nielsen, J��rgen G., 2018, Review of the steatiticus - species group of the cuskeel genus Neobythites (Ophidiidae) from the Indo-Pacific, with description of two new species, pp. 157-173 in Zootaxa 4387 (1) on pages 160-161, DOI: 10.11646/zootaxa.4387.1.7, http://zenodo.org/record/1186736, {"references":["Nielsen, J. G. (2002) Revision of the Indo-Pacific species of Neobythites (Teleostei, Ophidiidae) with 15 new species. Galathea Report, 19, 5 - 105.","Gloerfelt-Tarp, T. & Kailola, P. J. (1984) Trawled fishes of southern Indonesia and northwestern Australia. Australian Development Assistance Bureau, Australia, Directorate General of Fishes, Indonesia, and German Agency for Technical Cooperation, Federal Republic of Germany, 407 pp.","Nielsen, J. G. (1995) A review of the species of the genus Neobythites (Pisces, Ophidiidae) from the western Indian Ocean with description of seven new species. Ichthyological Bulletin, 62, 1 - 19."]}

Neobythites steatiticus Alcock, 1894 (Figures 1���3, Tables 1���3) Neobythites steatiticus Alcock, 1894: 181, pl. IX Neobythites steatiticus: Norman 1939: 76 (non steatiticus); Nielsen 1995: 11 and 2002: 84 (in part); Jawad 2014: 1 (non steatiticus). Holotype. ZSI 13435 (female, 129 mm SL), Bay of Bengal, NE Indian Ocean, 15��4'7'' N, 80��25'7'' E, RV Investigator, st. 137, Agassiz trawl, 234 m depth, 3 Mar. 1893. Non-types (n = 5, 113��� 162 mm SL). BMNH 1895.1.3.3 (former ZSI 13477), (female, 143 mm SL), Bay of Bengal, 13��1'6'' N, 80��36'56'' E, RV Investigator, st. 170, Agassiz trawl, 193 m depth, 16 Feb. 1894; USNM 46756 (former ZSI 13475), (female, 113 mm SL), USNM 256238 (former ZSI 13474), (female, 127 mm SL), and BMNH 1895.1.3.4 (female, 114 mm SL), Bay of Bengal, 13��51'12'' N, 80��28'12'' E, RV Investiga tor, st. 162, Agassiz trawl, 261���450 m depth, 30 Jan. 1894; ZMUC P 771752 (unripe, 162 mm SL), off Myanmar, 16��27' N 93��57' E, RV Dr. Fridtjof Nansen, st. 42, bottom trawl, 274���277 m depth, 6 May 2015. ������continued on the next page Tentatively referred. Tuticorin Research Center, Tamil Nadu, India, 2, 178��� 180 mm SL, Gulf of Manar, 8��44'46'' N, 78��37'24'' E, bottom trawl, 200���250 m depth, 22 Feb. 2010. Diagnosis. (Tentatively referred specimens from Gulf of Mannar not included). 0���1 weak spine on hind margin of preopercle; dorsal fin-rays 88���94; anal-fin rays 73���77; pectoral fin-rays 24���27; precaudal vertebrae 12; total vertebrae 53���57; pseudobranchial filaments 2���3; long gill rakers on anterior arch 11���14; head length 28.0��� 31.0 % SL; pelvic-fin length 14.0���15.0 % SL, pelvic fins not reaching anus; orbit length 4.8���5.2 % SL, 16.0���18.5 % head length, and 2.3���2.8 times in upper-jaw length; longest gill filament 4.4���5.9 % SL and 15.0���19.0 % head length; ocellus spot placed well behind a vertical line through anus, the ocellus-spot distance being 46.0���51.5 % SL, and the spot covering 7���10 dorsal-fin rays; median part of anal-fin dark and distal and proximal parts light; dorsal fin mostly unpigmented; up to 6 broad, rather distinct dark vertical bars on body; otolith length 5.0���5.2 % SL, sulcus length 3.6���3.7 % SL, and ostium height 21.0���24.5 % sulcus length. Distribution. Known from both sides of the Bay of Bengal, NE Indian Ocean, at 193���450 m depth. Remarks. According to Alcock (1899) ZSI held the holotype and eight additional specimens, five of which are here examined. Kannan et al. (2013:1) published the first record in Indian waters of Neobythites stefanovi. However, the illustration shows more resemblance to N. steatiticus so we treat it here as tentatively referred to N. steatiticus. Some specimens that in earlier literature were referred to N. steatiticus are now transferred to other Neobythites species: Norman (1939: 76) referred three specimens to N. steatiticus. Two of these from the Gulf of Aden have been transferred to N. stefanovi and one from off Zanzibar to N. kenyaensis Nielsen, 1995. Nielsen (1995: 11 and 2002: 84) mentioned six specimens caught in the Gulf of Oman (all from BMNH) which now are referred to N. gloriae n. sp. The specimen collected during the RV Dr. Fridtjov Nansen cruise off Myanmar represents the first record from the eastern Bay of Bengal. The photo taken on board immediately after capture is the first to document fresh colour of the species (Figure 1G 2). It reveals considerable similarity with ocellus, fin and body colour patterns shown in two different drawings of N. steatiticus provided by Alcock (1893, 1898) based on shortly preserved specimens. The slightly larger ocellus of the specimen from off Myanmar (10 vs. 7���9 dorsal-fin rays covered by the spot) contributes to the distinction of N. steatiticus from N. gloriae n. sp. (Figure 3, Tables 2���3). Jawad (2014) reported N. steatiticus from the Arabian Gulf off Iraq. However, judging from the dark blotch near the origin of the dorsal fin (vs. no anterior blotch), the position of the mid-body ocellus above or in front of anus (vs. well behind anus), and head length (21.5 vs. 28.0���31.0 % SL) this is neither a N. steatiticus nor a member of the steatiticus species group., Published as part of Uiblein, Franz & Nielsen, J��rgen G., 2018, Review of the steatiticus - species group of the cuskeel genus Neobythites (Ophidiidae) from the Indo-Pacific, with description of two new species, pp. 157-173 in Zootaxa 4387 (1) on pages 165-169, DOI: 10.11646/zootaxa.4387.1.7, http://zenodo.org/record/1186736, {"references":["Norman, J. R. (1939) Fishes. Scientific Report of the John Murray Expedition (1933 - 34), 7 (1), 1 - 116.","Nielsen, J. G. (1995) A review of the species of the genus Neobythites (Pisces, Ophidiidae) from the western Indian Ocean with description of seven new species. Ichthyological Bulletin, 62, 1 - 19.","Jawad, L. A. (2014) New record of Neobythites steatiticus Alcock, 1894 (Actinopterygii: Ophidiidae) from the marine waters of Iraq. International Journal of Marine Science, 4 (32), 1 - 3.","Kannan, K., John, S., Joshi, K. K. & Zacharia, P. U. (2013) First record of Neobythites stefanovi (Teleostei: Ophidiidae) from Indian waters. Marine Biodiversity Records, 6, e 71. https: // doi. org / 10.1017 / S 1755267213000523","Alcock, A. (1893) New species of Lophius, Neobythites, Odontostomus and Congromuraena. Journal of Asiatic Society Bengal, 62, 177 - 184.","Alcock, A. (1898) Illustrations of the zoology of H. M. Indian marine surveying steamer \" Investigator \". Royal Indian Marine, Calcutta, part V, plates 18 - 24."]}

Neobythites longipes Smith & Radcliffe, 1913 (Figures 1–2, Tables 1, 3) Neobythites longipes: Nielsen 2002: 38, fig. 9. Holotype. USNM 74126 (female, 276 mm SL), off Jolo Island, Philippines, Western Pacific, 06°02'00" N, 120°44'40" E, RV Albatross, st. 5550, bottom trawl, 472 m depth, 17 Sep. 1909. Paratypes (n = 4, 90– 207 mm SL). USNM 99070 (2 females, 165–242 mm SL), Philippines, 05°50' N, 120°31' E, RV Albatross, st. 5564, dredge, 432 m depth, 21 Sep. 1909; USNM 99073 (female, 90 mm SL), off Borneo, 04°52'45" N, 119°06'45" E, RV Albatros s, st. 5580, trawl, 296 m depth, 25 Sep. 1909; USNM 99091 (male, 207 mm SL), off Jolo, Philippines, 06°01'15" N, 120°44'20" E, RV Albatross st. 5549, trawl, 481 m depth, 17 Sep. 1909. Non-types (n = 18, 98– 301 mm SL). AMS I.22808-019 (female, 204 mm SL), 220 km N of Port Hedland, Western Australia, 17°59' S 118°17' E, RV Soela, st. SO 2/82/17–18, Engel trawl, 404–420 m depth, 3 Apr. 1982; AMS 1.22821-027 (female and male, 165–180 mm SL), 190 km northwest of Port Hedland, 18°16' S 118°12' E, RV Soela, st. SO 2/82/36–38, Engel trawl, 298–320 m depth, 10 Apr. 1982; AMS 1.23425-007 (2 males, 240–242 mm SL), northwest shelf, Western Australia, 18°46' S 117°00" E, RV Soela, st. SO 4/82/leg l, bottom trawl, 400 m depth, 1 Aug. 1982; CSIRO CA3916–19 (2 females and 2 males, 212–301 mm SL), off Port Hedland, Western Australia, 17°45.1' S 118°30.4' E, RV Soela, st. SO 183170, 442– 460 m depth, 5 Feb. 1983; NTM S.12588-020 (female, 194 mm SL), off Rowley Shoals, Western Australia, 17°22' S 118°38' E, RV Comac Endeavour, st. WH 85-15, 430 m depth, 2 Nov. 1985; NTM S.12631-005 (female and 2 males, SL 226–264 mm SL), off Rowley Shoals, Western Australia, 17°37' S 118°40' E, RV Comac Endeavour, st. YW 85-21, 400 m depth, 4 Nov. 1985; NTM S.14378-001 (male, 245 mm SL), off Cartier Reef, Timor Sea, 13°7.89' S 123°12.65' E, RV Lady Basten, st. RW 96-30, 420 m, 19 June 1996; WAM P.28081-001 (female, 230 mm SL), 45 km northeast of Rowley Shoals, Western Australia, 17°44' S 120°5' E, 431–433 m depth, 20 Aug. 1983; WAM P.29125-001 (female, 163 mm SL), Rowley Shoals, Western Australia, 17°27' S 119°44' E, RV Umitaka Maru, trawl, 20 Dec. 1969; ZMUC P779 (female, 195 mm SL), off Jolo, Philippines, trawl, 463 m depth, 27 Mar. 1914; ZMUC P77740 (male, 98 mm SL), SE of Lombok, Indonesia, 8°58’ S 116°34’ E, JETINDOFISH TGT 1718, trawl, 150–280 m depth, July 1981. Diagnosis. No spines on hind margin of preopercle; Dorsal fin-rays 96–103; anal-fin rays 72–76; pectoral finrays 27–30; precaudal vertebrae 13–14 (mostly 14); total vertebrae 57–60; pseudobranchial filaments 5–10; long gill rakers on anterior arch 8–10; head length 22.5–25.5 % SL; pelvic-fin length 28.5–54 % SL, pelvic fins reaching well beyond anus; orbit length 3.9–5.0 % SL, 17.0–21.5 % head length, and 2.4–3.2 times in upper-jaw length; longest gill filament 1.3–2.7 % SL and 5.7–11.0 % head length; ocellus spot placed behind a vertical line through anus, the ocellus-spot distance being 42.0–53 % SL, and the spot covering 9–13 dorsal-fin rays; margins of posterior third of dorsal fin and posterior two thirds of anal fin dark pigmented; no vertical bars on body; otolith length 4.8–6.2 % SL, sulcus length 4.3–4.9 % SL, and ostium height 12.5–15.0 % sulcus length. Distribution. From NW Australia to Philippines, at 150–481 m depth. Remarks. The photograph of the fresh specimen (Figure 1C 2) shows that the anterior two thirds of the lateral line and the entire margin of the anal fin to be black pigmented. The pigmentation of the dorsal fin as revealed by examination of the preserved specimen is however not evident in the fresh photograph. The overall bluish colour of the fresh specimen appears to us as artifact.

Neobythites gloriae n. sp. (Figures 1–3, Tables 1–3) Neobythites steatiticus: Nielsen 1995 (in part, Arabian Gulf, Gulf of Oman). Neobythites stefanovi: Nielsen & Uiblein 1993 and Nielsen 1995, 2002 (in part, Gulf of Oman). Holotype. USNM 309008 (male, 158 mm SL), Gulf of Oman, NW Indian Ocean, 25°02' N, 56°52' E, RV Anton Bruun, cruise 4B, st. 264A, bottom trawl, 272–291 m depth, 2 Dec 1963. Paratypes (n=8, 130– 150 mm SL). BMNH 1904.5.25.1 (1 male, 150 mm SL), Gulf of Oman, 25°32' N, 57°47' E, 320 m depth; BMHN 1910.1.31.7–10 (3 females and 1 male, 132–145 mm SL), Gulf of Oman, 25°35' N, 57°47' E, 311 m depth; BMHN 1910.1.31.23 (1 female, 130 mm SL), Arabian Gulf, 10' W of Dubai, United Arab Emirates, 26 m depth; USNM 440378 (1 male, 140 mm SL), and ZMUC P771754 (1 female, 131 mm SL), same data as for the holotype. Diagnosis. No spines on hind margin of preopercle; dorsal fin-rays 90–93; anal-fin rays 72–76; pectoral finrays 24–27; precaudal vertebrae 12; total vertebrae 54–56; pseudobranchial filaments 3–4; long gill rakers on anterior arch 11–13; head length 25.5–28.0 % SL; pelvic-fin length 13.5–17.0 % SL, pelvic fins not reaching beyond anus; orbit length 5.3–5.9 % SL, 19.5–23.5 % head length, and 2.0–2.2 times in upper-jaw length; longest gill filament 3.9–4.9 % SL and 15.5–19.0 % head length; ocellus spot placed well posterior to a vertical line through anus, the ocellus-spot distance being 49.0–51 % SL, and the spot covering 6–8 dorsal-fin rays; margins of posterior fifth to a quarter of dorsal fin and posterior half to two thirds of anal fin dark pigmented; no vertical bars on body; otolith length 4.8–5.0 % SL, sulcus length 3.4–3.7 % SL, and ostium height 21.0–26.5 % sulcus length. Description (based on holotype, differences in paratypes mentioned in round brackets). Rather elongate fish with complete lateral line in preserved specimens; on the anterior half of body, the lateral line runs with 8–10 scale rows to dorsal edge, decreasing posteriorad; head and body fully covered with cycloid scales; origin of dorsal fin above hind margin of opercle; origin of anal fin slightly in front of midpoint of fish; tip of pectoral fins ending just above origin of anal fin; pelvic fins reaching almost to origin of anal fin; blunt snout slightly shorter than diameter of eye; upper jaw ending well behind eye; anterior nostril ending in a short tube and the larger posterior nostril in a mere hole; no spines on preopercle; strong opercular spine; anterior gill raker with 1–2 (2–3) blunt and 3–4 (2–3) long rakers on upper branch; one long raker in angle and lower branch with eight long and 6 (7) blunt rakers; gill filaments 1.5–3.0 (1–4) times as long as long rakers; distinct pseudobranchial filaments 3 (3–4). Otolith. Otolith oval, its height 1.6 (1.5–1.7) times in its length; sulcus large, 1.3 (1.3–1.4) times in otolith length; ostium 1.5 times in sulcus length; ostium height 3.3 (2.6–3.3) times in ostium length. Dentition. Premaxillaries and dentaries with small, pointed teeth in 3–4 irregular rows anteriorly, decreasing to 1–2 rows posteriorad. Teeth in outer rows somewhat longer. Vomer triangular with many small, close-set teeth. Palatines with several rows of small, close-set teeth. Two median basibranchial tooth patches, the anterior elongate and the posterior much smaller and circular. Axial skeleton (from radiographs). Precaudal vertebrae 12 and caudal vertebrae 42 (42–45) and all neural and haemal spines pointed; first neural spine one third length of second spine, that is longer than the following spines; bases of neural spines more or less enlarged; parapophyses present on posterior 7–8 precaudal vertebrae; pleural ribs on vertebrae 3–11 (3–4 to 10–12); epipleural ribs on vertebrae 3–8. Colour. Preserved fish (holotype [Figure 1A] and paratypes). Body mottled brown, abdomen silvery bluish, lateral line pale brown, rather indistinct; head pale brown with brown preopercular area behind eye, gill cover transparent, eyes bluish; ocellus spot dark brown, its horizontal diameter about orbit length, placed well behind a vertical line through anal-fin origin; dorsal and anal fins with pale- to dark-brown distal margins along posterior fifth to quarter of dorsal fin and posterior half to two-thirds of anal fin; margins becoming darker towards dark caudal-fin tip. Etymology. The new species name honors the wife of the first author, Gloria Jansen Echevarria. Distribution. Inner Gulf of Oman, NW Indian Ocean, at 272–320 m depth, and southern Arabian Gulf, United Arab Emirates, at 26 m depth. Comparisons. Neobythites gloriae n. sp. differs from all congeners by the following combination of characters: a single ocellus placed on dorsal fin at mid-body, preopercular spines lacking, 11–13 developed gill rakers, head length 25.5–28.0 % SL, gill filament length 3.9–4.9 % SL, orbit length 19.5–23.5 % head length and 45–50 % upper-jaw length, and otolith sulcus length 3.4–3.7 % SL. Neobythites gloriae n. sp. is most similar to N. stefanovi, sharing the following characters: dorsal fin with one ocellus, preopercular spine absent, pelvic fins ending anterior to anus, similar number of long rakers on anterior gill arch, eyes five times or less in head length, and similar otolith sulcus length. Neobythites gloriae differs from N. stefanovi having longer gill filaments (3.9–4.9 vs. 1.4–3.7 % SL) and a slightly smaller ocellus spot (spot covering 6–8 vs. 8–12 dorsal-fin rays). These two characters in combination with head length, orbit length, and ocellus-spot distance best distinguish N. gloriae from N. steatiticus and N. steatiticus from N. stefanovi (Figure 3; Tables 1–3). Remarks. The description of Neobythites gloriae n. sp. is based on three former N. stefanovi paratypes from the inner Gulf of Oman (Nielsen & Uiblein 1993) and six specimens previously identified as N. steatiticus (Nielsen 1995). Apart from this earlier misidentified material, no additional specimens and no photographs of fresh fish were available for our study. The depth of 26 m reported for the single Arabian Gulf specimen (BMHN 1910.1.31.23), indicated as questionable by Nielsen (1995), is here confirmed based on original data recordings available from NHM.

Goatfishes of the genus Parupeneus (Mullidae), from several areas and countries of the Indo-West Pacific, were studied, focusing on species characterized by a symmetrically rounded rear end of the maxilla, which belong to the so-called heptacanthus group. We examined 62 meristic and morphometric characters and colour patterns from 180 specimens from 11 Parupeneus species, using a comprehensive alpha-taxonomy approach. When possible, the data for species represented by wider size ranges were split into two size classes (small-sized fish, < 110 mm SL, vs. large-sized fish ≥ 110 mm SL) to account for allometric changes in body form. A new species, P. inayatae Uiblein & Fahmi, is described, based on a specimen collected in SW Lombok, Indonesia, and new records for four species are reported from the Indian Ocean: Parupeneus fraserorum new for Mozambique, P. jansenii new for Myanmar and the NE Indian Ocean, P. nansen new for Tanzania, and P. procerigena new for the Nazareth Bank and Mauritus (Cargados Carajos Bank), Western Indian Ocean (WIO). The record of P. jansenii for Myanmar was compared in detail to 37 conspecific specimens from the entire distribution range (western Indonesia to Japan and Vietnam to NE Australia). The latter revealed two new Pacific records for the Central Philippines and for Shikoku Island, Japan (northernmost record of the species). For each of the eight heptacanthus-group species, diagnoses are provided along with remarks on intraspecific size- and/or population-related variation and/or other noteworthy observations. The new species can be distinguished from all other 32 congeners as follows: dorsal head margin in front of eyes concave shaped; posterior maxilla margin evenly, symmetrically rounded; body, head, and caudal peduncle moderately deep; pectoral and pelvic fins subequal in length; and caudal peduncle clearly deeper than 9.5 % SL; in life a dusky spot on lateral line below interdorsal space, followed posteriorly by a bright orange blotch. Detailed comparisons among all eight heptacanthus-group species and with three other similar and/or co-occurring congeners revealed considerable similarity between two species pairs, P. jansenii (Eastern Indian Ocean and Western Pacific) and P. nansen (WIO), and P. fraserorum and P. procerigena (both from the WIO). The need to further intensify taxonomic studies and accompanying fisheries-biology and ecological studies of goatfishes in the large realm of the Indo-West Pacific is emphasized.

Parupeneus jansenii (Bleeker, 1856) Jansen’s goatfish (Figs. 1, 3–5; Tables 2, 3) Upeneus jansenii Bleeker, 1856 (type locality Manado, Sulawesi). Parupeneus jansenii: Bleeker 1875; Randall 2004; Allen & Erdmann 2012 Syntypes. RMNH 5748, 2 specimens, 76–91 mm SL, Indonesia, Sulawesi, Manado. Non-type material (n = 27, 68– 159 mm SL). Vietnam, South-central coast: MUL.PA. CHR.04, Hai Phong Fisheries Research Institute Reference Collection, 125 mm SL; Vietnam, Khanh Hoa province, Nha Trang: HIFIRE 58155, 5 specimens, 143–152 mm SL, VNMN-I 54, 147 mm SL, and VNMN-I 64, 157 mm SL, fish market; VNMN-I 50, 143 mm SL, VNMN-I 51, 135 mm SL, and VNMN-I 52, 142 mm SL, Binh Tan fish market, 12°12.793' N; 109°11.825' E; VNMN-I 4, 159 mm SL, Hon Ro landing site, 12°12.07' N, 109°11.66' E; VNMN-I 134, 74 mm SL, and VNMN-I 135, 68 mm SL, Hon Tre Island, 12°10'– 12°14' N; 109°14'– 109°21'E, obtained from fishermen; Indonesia: RMNH 25006, 102 mm SL, and RMNH 25008, 1 (of 2), 107 mm SL; southern Bali: CSIRO H 5972-19, 146 mm SL, Jimbaran Bay, Kedonganan fish market, 8°45' S 115°10' E; southern Lombok: NTM S 10740 -006, 95 mm SL, 9°0' S 116°12' E, 100 m depth; NE Australia, Queensland, Torres Strait: Torres Strait: CSIRO H 6148-03, 118 mm SL, NE of Cape York Peninsula, 10°19.87' S 143°33.42' E, 37 m depth; CSIRO H 6926-10, 4, 94–114 mm SL, NE of Cape York Peninsula, 10°21.84' S 143°40.26' E, 40 m depth; S of Torres Strait: CSIRO H 3446-04, 114 mm SL, E of Cape York Peninsula, Blackwood Channel, 11°43.4' S, 143°42.6' E, 23 m depth; CSIRO H 7462-03, 131 mm SL, E of Newcastle Bay, 10°55.1' S 143°54.6' E, 32 m depth; Great Barrier Reef: CSIRO H 7460-04, 104 mm SL, NE of Townsville, 18°31.48' S 147°34.9' E, 76 m depth; CSIRO H 8154-01, 81 mm SL, NE of Lizard Island, 14°35.38' S 145°33.98' E, 26 m depth. Diagnosis. Dorsal fins VIII + 9; pectoral fins 14–16; gill rakers 6–7 + 2 0–23 = 26–29; measurements in % SL, large-sized fish: body depth at first dorsal-fin origin 23–26; body depth at anus 19–21; caudal-peduncle depth 8.2– 9.0; maximum head depth 22–24; head depth through eye 19–21; interorbital length 8.1–10; head length 31–34; snout length 14–16; postorbital length 11–12; orbit length 5.7–7.7; upper jaw length 12–15; barbel length 22–25; caudal-peduncle length 21–24; caudal-fin length 25–27; anal-fin height 12–14; pelvic-fin length 20–22; pectoralfin length 21–25; first dorsal-fin height 17–20; second dorsal-fin height 11–14; small-sized fish: body depth at first dorsal-fin origin 22–26; body depth at anus 18–21; caudal-peduncle depth 7.8–8.8; maximum head depth 20–24; head depth through eye 18–21; interorbital length 8.1–9.7; head length 32–34; snout length 13–15; postorbital length 11–13; orbit length 6.8–7.8; upper jaw length 11–14; barbel length 22–26; caudal-peduncle length 21–24; caudal-fin length 25–29; anal-fin height 12–14; pelvic-fin length 21–23; pectoral-fin length 22–23; first dorsal-fin height 17–19; second dorsal-fin height 11–14; maxilla posterior margin evenly, symmetrically rounded; body and head rose, pink, or vermillion, colour may become paler below lateral line and eye; scale edges darker; body silvery white ventrally; one yellow-orange mid-lateral stripe sometimes visible from behind operculum along lateral line; caudal fin red or dark orange along proximal exterior sides of lobes, the remaining parts orange-yellowish to hyaline; dorsal fins and anterior part of pelvic fins pale red or pale orange; anal fin crossed by weakly indicated red-greenish stripes; barbels white or rose to red at basis, tips white; preserved fish pale or pale brown to pale grey, head or dorsal body and head margin sometimes darker. Description. Morphometric data as ratios of SL for large-sized fish, data for small-sized fish (including syntypes) in squared brackets: body elongated, its depth at first dorsal-fin origin 3.8–4.3 [3.9–4.5]; body depth at anal-fin origin 4.8–5.3 [4.8–5.4]; caudal peduncle depth 11–12 [11–13], subequal to interorbital length (10—12 [10–12]); maximum head depth 4.1–4.5 [4.2–4.9], subequal to barbel length (4.0–4.6 [3.9–4.6]); head depth through eye 4.7–5.2 [4.9–5.6]; head length 2.9–3.2 [3.0–3.2], larger than maximum depth of body and caudal-fin length (3.7–4.0 [3.5–4.0]); snout length 6.2–7.0 [6.6–8.0], shorter than postorbital length (8.0–9.4 [7.5–8.9]) in large-sized fish; orbit length 13–18 [13–15]; anal-fin height 7.2–8.4 [7.0–8.1]; second dorsal-fin height 7.3–9.3 [7.0–8.8]; pectoral-fin length 4.0–4.7 [4.3–4.6], subequal to barbel length and subequal to pelvic-fin length (4.6– 5.0 [4.3–4.9]) in small-sized fish. Fresh colour: freshly landed large-sized fish (Figure 3A, VNMN-I 4, 159 mm SL): body and head vermillion dorsally, becoming paler ventrally, starting from below eye and from scale row above lateral line; head with four slightly bending, thin, pink stripes: the uppermost stripe starting from behind dorsal third of eye, interrupted by orbit, to head margin in front of mid-eye, paralleled by a stripe of similar length right below; a longer pink stripe starting anterior to lower eye margin and ending close to nose tip; and the lowermost pink stripe from behind eye to below eye and ending anteriorly at half distance of snout length; jaws pink; further below eye the specimen has a large vertically oriented vermillion broad band that touches the lowermost rose stripe (see above) dorsally and ends well above ventral head margin; the preopercle is pale rose with vermillion margin; the black pupil of the eye is surrounded by an almost entirely red iris; all scales on body, apart from the ventral-most scale row, have dark reddish margins that contrast with the lighter scale colour; the lateral-line scales along the anterior half of the body show small red, mid-posteriorly placed spots; a broad, pale orange-yellow stripe in width of about scale vertical length runs along lateral line from behind preopercle to caudal-fin base; belly silvery white; caudal fin pale pink at base, with vermillion bands reaching along exterior margins of lobes to tips, covering in the anterior halves of lobes 5 to 6 of the distalmost rays; the remaining inner part of the caudal fin is covered by brighter yellow-orange colour reaching from distal-most part of fin base to close to lobe tips; dorsal fins vermillion, pelvic fins with some rose traces, and pectoral fins hyaline; anal-fin colour not identified, because folded in photograph; barbels white to pale rose, tips white. Large-sized fish encountered at fish market (Figure 3C, CSIRO H 5972-19, 146 mm SL): body and head mostly entirely vermillion, dorsally darker; ventral underside of head white; eye with black pupil surrounded by red iris that contrasts weakly with the darker vermillion head colour; body colour changing laterally towards ventral side from dark to bright vermillion to beige and rose and pale pink below lateral line; belly pale pink; scales with dark red or reddish margins that contrast with the lighter colour of scale surface, apart from the scales adjacent to the anterior dorsal body margin where body colour is darkest and the contrast is diminished; caudal fin entirely vermillion at base, the colour spreading along exterior margins of lobes to tips, covering at anterior halves of lobes 5 and 6 of the distalmost rays, then continuing along the two distalmost rays until tips, becoming much weaker close to tip, especially on upper lobe; the remaining posterior part of the caudal fin is yellow-orange to beige in colour; dorsal fins are mostly vermillion or rose, spines and anterior base of first dorsal fin most intensely coloured, rays of second dorsal fin proximally vermillion and pale on tips, the beige interspaces partly covered by one to three rounded, partly connected, vertically arranged vermillion patches; pelvic fins hyaline with pale rose spine and all rays except for innermost ray with three to four beige to pale brown patches proximally; pectoral fins hyaline; anal fin hyaline with three thin pale green to pale brown stripes covering entire length of fin, one close to base, the distalmost stripe thinnest, positioned at about two thirds of fin height. Barbels vermillion at base, distal two thirds white including tip. Freshly caught small-sized fish (Figure 4A, VNMN-I 135, 68 mm SL): body and head dark orange dorsally, becoming ventrally paler, starting from below eye and lateral line; head yellowish-orange below and behind ventral half of eye, with two weakly indicated pink dashes anterior to lower half of eye; snout tip and jaws dark orange; the black pupil of the eye is surrounded by a red iris; nearly all scales on body have reddish, orange or yellowish margins that contrast with the lighter scale-surface colour; the lateral-line scales form a thin bright orange to yellow stripe from posterior of the preopercle to behind second-dorsal fin base; this stripe is surrounded by a broader yellow pigmentation starting below first dorsal-fin base that widens from ca. mid-body to caudal fin base, almost entirely covering the dorsal part of the caudal peduncle; belly silvery white; caudal fin and dorsal fins dark orange; pelvic fins hyaline with pale orange spine and first three rays; pectoral fins hyaline with orange basis; anal fin hyaline; barbels pale rose along proximal two thirds and white distally including tip. Preserved colour (syntype, RMNH 5748, 91 mm SL): body dorsally dark brown grey, lighter from lateral line, with weakly pale brown ventral half of body; head mostly dark grey brown, except for tiny pale-brown area in front of eye and pale brown region behind jaws, pale brown opercle and preopercle; fins pale, partly hyaline; other specimens often entirely pale brown or with darker, grey brown dorsal margin on head and body; Distribution, depth range, and size. Parupeneus jansenii occurs in the Western Pacific and Eastern Indian Ocean in the area ranging from south-central Vietnam and northern Philippines to southwestern Indonesia and NE Australia, with verified records from Nha Trang Bay (this study), Luzon (Randall 2004), northern Sulawesi (type locality Manado), Sunda Strait off southern Sumatra (Randall 2004), southern Bali and southern Lombok (this study), and Torres Strait to Great Barrier Reef (this study). Depth range is 24–100 m, maximum size is 159 mm SL., Published as part of Uiblein, Franz, Hoang, Tuan Anh & Gledhill, Daniel, 2017, Redescription and new records of Jansen's goatfish, Parupeneus jansenii (Mullidae), from the Western Pacific and Eastern Indian Ocean, pp. 541-559 in Zootaxa 4344 (3) on pages 546-548, DOI: 10.11646/zootaxa.4344.3.6, http://zenodo.org/record/1043749, {"references":["Bleeker, P. (1875) Revision des especes insulindiennes de la famille des Mulloides. Ferhandelingen der Koninklijke Akademie van Wetenschappen te Amsterdam, 15, 1 - 40.","Randall, J. E. (2004) Revision of the goatfish genus Parupeneus (Perciformes: Mullidae), with descriptions of two new species. Indo-Pacific Fishes, 36, 1 - 64.","Allen, G. R. & Erdmann, M. V. (2012) Reef fishes of the East Indies. Fols. I - III. Tropical Reef Research, Perth, 1292 pp."]}

Parupeneus heptacanthus (Lacepède, 1802) Cinnabar goatfish (Figs. 2–5; Table 2) Sciaena heptacantha Lacepède, 1802 (type locality not given) Upeneus cinnabarinus Cuvier in Cuvier and Valenciennes, 1829 Upeneus pleurospilos Bleeker, 1853 (type locality Ambon, Moluccas, Indonesia) Parupeneus heptacanthus Kim & Amaoka 2001, Randall 2004, Randall & Heemstra 2009, Allen & Erdman 2012 Holotype. MNHN 5438, 153 mm SL, dried specimen, type locality not given (probably Indonesia). Syntype. Parupeneus pleurospilus, RMNH 5743, 106 mm SL, Indonesia, Moluccas, Ambon. Non-type material (n = 51, 59– 193 mm SL). Vietnam: MNHN 1967-547, 185 mm SL; South-central coast: MUL.PA.HEP.02, Hai Phong Fisheries Research Institute Reference Collection, 171 mm SL; Da Nang province, Da Nang: VNMN-I 5, 121 mm SL, and VNMN-I 6, 119 mm SL, Tho Quang, Hoang Sa street landing site, 16°06.01' N 108°15.65' E; Khanh Hoa province, Nha Trang: VNMN-I 44, 186 mm SL, and VNMN-I 45, 192 mm SL, Vinh Luong landing site, 12°20.110' N, 109°12.228' E; VNMN-I 48, 139 mm SL, and VNMN-I 47, 155 mm SL, Xom Moi market, 12°14.566' N, 109°11.426' E; VNMN-I 40, 181 mm SL, VNMN-I 41, 164 mm SL, VNMN- I- 42, 161 mm SL, VNMN-I 46, 184 mm SL, VNMN-I 49, 170 mm SL, VNMN-I 66, 2 specimens: 131–138 mm SL, and VNMN-I 128, 191 mm SL, Binh Tan fish market, 12°12.793' N; 109°11.825' E; VNMN-I 137, 5 specimens: 81–106 mm SL, VNMN-I 138, 3 specimens: 64–72 mm SL, and VNMN-I 139, 2 specimens, 136–158 mm SL, Vinh Truong district, Cua Be landing site, 12°12.09' N; 109°12.122' E; VNMN-I 1, 167 mm SL, VNMN-I 2 152 mm SL, and VNMN-I 3, 128 mm SL, Hon Ro landing site, 12°12.07' N 109°11.66' E; VNMN-I 136, 59 mm SL, Hon Tre Island, 12°10'–12°14' N 109°14'–109°21' E, obtained from fishermen; Philippines: MNHN 1979-90, 1 (of 3), 149 mm SL; Indonesia: RMNH 5737, 3(of 6), 160–166 mm SL; North Sulawesi, Manado: NCIP 5033, 162 mm SL, Manado fish market, 1°29.85' N 124°50.45' E; Java Sea: RMNH 13339, 84 mm SL; southern Bali: CSIRO H 5972-17, 150 mm SL, CSIRO H 5972-18, 132 mm SL, CSIRO H 7695-03, 137 mm SL, and CSIRO H 8157-01, 184 mm SL, Jimbaran Bay, Kedonganan fish market, 8°45' S 115°10' E; Australia, Western Australia: CSIRO CA 2881, 181 mm SL, off Port Hedland, 20°00' S 117°49' E, 37–38 m depth; CSIRO CA 3859, 143 mm SL, CSIRO CA 3860, 155 mm SL, and CSIRO CA 3861, 186 mm SL, NW of Port Hedland, 19°16' S 117°55' E, 82-84 m depth; Northern Territory: CSIRO CA 2037, 193 mm SL, NW of Bathurst Island, 11°12' S 128°35' E, 38 m depth; CSIRO CA 2061, 155 mm SL, NE of N Goulburn-Island, 10°43' S 134°00' E, 62 m depth; Queensland, South of Torres Strait: CSIRO H 3894-01, 152 mm SL, and CSIRO H 3894-02, 150 mm SL, NE of Cape Grenville, 11°38' S 143°35' E, 18 m depth; Great Barrier Reef: CSIRO H 7379-07, 122 mm SL, NE of Cairns, 16°32' S 146°02' E, 54 m depth; CSIRO H 8155-01, 138 mm SL, NE of Yeppoon, 22°44' S 151°27' E, 60 m depth; CSIRO H 8156-01, 193 mm SL, N of North Reef, Capricorn Group, 23°02' S 151°55' E, 67 m depth. Diagnosis. Dorsal fins VIII + 9; pectoral fins 15–17; gill rakers 5–7 + 2 0–23 = 26–29; measurements in % SL, large-sized fish: body depth at first dorsal-fin origin 29–35; body depth at anus 23–29; caudal-peduncle depth 10– 12; maximum head depth 26–31; head depth through eye 21–25; interorbital length 7.7–9.2; head length 31–34; snout length 14–17; postorbital length 10–13; orbit length 5.8–8.1; upper jaw length 12–14; barbel length 25–29; caudal-peduncle length 21–25; caudal-fin length 29–33; anal-fin height 13–16; pelvic-fin length 22–25; pectoralfin length 23–28; first dorsal-fin height 20–23; second dorsal-fin height 11–15; small-sized fish: body depth at first dorsal-fin origin 28–30; body depth at anus 23–26; caudal-peduncle depth 10–11; maximum head depth 24–27; head depth through eye 19–22; interorbital length 7.7–8.4; head length 30–34; snout length 13–15; postorbital length 10–13; orbit length 6.6–8.3; upper jaw length 12–13; barbel length 26–28; caudal-peduncle length 22–25; caudal-fin length 29–32; anal-fin height 14–16; pelvic-fin length 23–24; pectoral-fin length 24–26; first dorsal-fin height 19–21; second dorsal-fin height 13–15; maxilla posterior margin evenly, symmetrically rounded; body and head purple red or vermillion, colour may become paler below lateral line and eye; scale edges darker; scales often with pale violet or pearly iridescent spot; body silvery white ventrally; one or two indistinct narrow yellow-orange stripes sometimes visible, one of them just above or along lateral line and more conspicuous, the other further below; a small dark red or reddish-brown spot on or just below the eighth lateral-line scale in both large- and smallsized fish; caudal fin red at base and ventrally of lower lobe, the remaining parts yellowish to pale orange; dorsal and anterior parts of anal and pelvic fins red or orange; barbels white or pale orange except for the white tips; preserved fish pale or pale brown to pale grey, head or dorsal body and head margin sometimes darker. Comparisons (Figures 1, 3–5, Tables 2, 3) Parupeneus jansenii differs from P. heptacanthus - independently of size - in shallower body and caudal peduncle, smaller maximum head depth, shorter barbels, caudal and pelvic fins, shallower first dorsal fin, shallower first ray of second dorsal fin, and lack of small dark red or reddish-brown spot on or just below the eighth lateral-line scale; furthermore, small-sized P. jansenii differs in shorter second dorsal-fin base, shallower anal fin, shorter pelvic-fin spine, and shorter pectoral fins; and large-sized P. jansenii differs in shallower head depth through eye and longer anal-fin base. Regarding meristic characters, Parupeneus jansenii differs from P. heptacanthus statistically with signifcantly fewer pectoral-fin rays, more gill rakers on upper limb, and constantly 27 lateral-line scales in contrast to 27–28 scales in the latter species (Figure 5). The currently known maximum size of P. jansenii is 159 mm SL (this study), while P. heptacanthus has been reported to reach a much larger size of 290 mm SL (Randall 2004). Parupeneus jansenii P. heptacanthus P. P. P. P. P. P. angulatus fraserorum minys nansen procerigena seychellensis Small-size&dstrok; fish Large-size&dstrok; fish Small-size&dstrok; fish Large-size&dstrok; fish Min Mean Max n Min Mean Max n Min Mean Max n Min Mean Max n HT PT PT HT PT HT ……continued on the next page Parupeneus jansenii P. heptacanthus P. P. P. P. P. P. angulatus fraserorum minys nansen procerigena seychellensis Small-size&dstrok; fish Large-size&dstrok; fish Small-size&dstrok; fish Large-size&dstrok; fish Min Mean Max n Min Mean Max n Min Mean Max n Min Mean Max n HT PT PT HT PT HT Small-size&dstrok; fish Large-size&dstrok; fish In&dstrok;onesia Vietnam Australia All In&dstrok;onesia Vietnam Australia All ……continued on the next page Small-sized fish Large-sized fish Indonesia Vietnam Australia All Indonesia Vietnam Australia All Syntypes Min Mean Max n n =2 Min Mean Max n Min Mean Max n n =1 Min Mean Max n Min Mean Max n Min Mean Max n Parupeneus jansenii differs from all other similar species (heptacanthus group and P. angulatus) as follows: from P. fraserorum, P. procigerena, and P. seychellensis in shallower body, shallower caudal peduncle, shallower head, shorter snout, shorter anal-fin base, shorter pelvic fin, and shallower eighth dorsal-fin ray; additional important differences from P. fraserorum are: fewer pectoral-fin rays, shorter barbels, shorter second dorsal-fin base, higher anal fin, shallower first ray of second dorsal fin, and wider pectoral fin; from P. procigerena: shorter postorbital length, wider interdorsal distance, shorter barbels, longer caudal peduncle, higher anal fin, shorter pectoral fins, shallower first dorsal fin, and shallower ninth ray of second dorsal fin; and from P. seychellensis: fewer gill rakers, shorter and narrower snout, and longer anal-fin base. Parupeneus jansenii differs from P. angulatus and P. nansen in shorter caudal fin, shallower third dorsal-fin spine, and fewer lateral-line scales; furthermore, it differs from P. angulatus in round vs. angular rear end of maxilla, in shorter postorbital, longer caudal peduncle, shorter anal-fin base, slightly higher first anal-fin ray and first ray on second dorsal fin; and from P. nansen it differs in shallower maximum body depth, wider barbel, and shorter pelvic fin. Parupeneus jansenii is morphologically rather similar to P. minys, especially, when comparing the 104 mm SL paratype of P. minys with P. jansenii small-sized fish. Accordingly, similar-sized P. jansenii differs in larger eyes, wider barbels, longer and wider pectoral fins, and shallower first dorsal-fin spine. Parupeneus jansenii reaches at least 159 mm SL (this study), while for P. minys a maximum size of only 106.5 mm SL has been reported (Randall & Heemstra 2009). Parupeneus jansenii shows intraspecific differences among size classes and populations (Figure 1; Table 3). The single large-sized fish from Indonesia differs from co-occurring small-sized fish in several morphometric characters. Small-sized fish of the NE Australian population differ from the other populations in having a higher eighth ray on the second dorsal fin. This difference applies also for the large-sized fish of the NE Australian population when compared with similar-sized specimens from Vietnam. The values for the geographically intermediate population from Indonesia largely overlap with those of the two other populations., Published as part of Uiblein, Franz, Hoang, Tuan Anh & Gledhill, Daniel, 2017, Redescription and new records of Jansen's goatfish, Parupeneus jansenii (Mullidae), from the Western Pacific and Eastern Indian Ocean, pp. 541-559 in Zootaxa 4344 (3) on pages 548-556, DOI: 10.11646/zootaxa.4344.3.6, http://zenodo.org/record/1043749, {"references":["Kim, B. - J. & Amaoka, K. (2001) A new species, Parupeneus procerigena, from the Saya de Malha Bank in the western Indian Ocean (Perciformes: Mullidae). Ichthyological Research, 48, 48 - 50. https: // doi. org / 10.1007 / s 10228 - 001 - 8115 - 8","Randall, J. E. (2004) Revision of the goatfish genus Parupeneus (Perciformes: Mullidae), with descriptions of two new species. Indo-Pacific Fishes, 36, 1 - 64.","Randall, J. E. & Heemstra, E. (2009) Three new goatfishes of the genus Parupeneus from the Western Indian Ocean, with resurrection of P. seychellensis. Smithiana Bulletin, 10, 37 - 50.","Allen, G. R. & Erdmann, M. V. (2012) Reef fishes of the East Indies. Fols. I - III. Tropical Reef Research, Perth, 1292 pp."]}