taxonID	type	description	language	source
B829390BFFCCFFE3FF27FDA33BFF44AE.taxon	description	(Figures 1 – 6; Tables 1 – 7)	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFCCFFE3FF27FDA33BFF44AE.taxon	description	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).	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFCCFFE3FF27FDA33BFF44AE.taxon	materials_examined	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: BPBM 27694, 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: MNHN 1984 - 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 P 49560, 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).	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFCCFFE3FF27FDA33BFF44AE.taxon	diagnosis	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. 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 3 C; 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.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFCCFFE3FF27FDA33BFF44AE.taxon	distribution	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 dorsal fin (Tables 1, 2; Figure 4); they differ in colour in having fewer bars on the caudal fins and the bars especially on the lower caudal-fin lobe are more varied in form and size (Tables 1, 2; Figures 2,3); in preserved subadults dark pigmentation patterns are more often retained than in adults (Tables 1, 2; see also preserved-colour description above). Among the three populations of Upeneus margarethae from the WIO proper (WIOP), the Red Sea, and the EIO, Arafura Sea and Gulf of Carpentaria (N Australia, E Pacific) (EIO / NA), there is no clear distinction in any single or combination of morphological or colour characters that would support separation into different taxa (Table 1, Figures 2 – 6). However, each population can be distinguished statistically (Table 3). For instance, the EIO / NA population has significantly longer snout-pelvic-fin distance, shorter second dorsal-fin base and shallower dorsal fins than the other two populations; the Red Sea population has a significantly shallower head, smaller eyes, longer caudal fin, and narrower pectoral fins than the other two populations; the main distinction of the WIOP population is a significantly shallower body at anal-fin origin. Furthermore, several additional significant differences in morphometric characters occur among pairs of the three populations (Table 3). The considerable number of significant differences among populations contrasts however with a much higher degree of differentiation among species, as revealed by the statistical comparisons of U. margarethae with two of the other four species (Table 4, see also section on Interspecific comparisons at end of taxonomic account) Regarding meristic and colour characters, the EIO / NA population has a significantly higher number of lateralline scales than the WIOP population (Table 5) and significantly less dark pigmentation in preserved conditon than the other two populations (Table 6), with significantly fewer remains of bars on upper caudal-fin lobe and second dorsal-fin distal stripe and no occurrence of mid-lateral dots (Table 7). Addtional significant differences in preserved pigmeentation pattern concern the WIOP and the Red-Sea populations which differ in two and three color characters from the other two populations, respectively (Table 7). Regarding fresh colour patterns, no obvious population differences could be found based on the available photo documentation.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFCCFFE3FF27FDA33BFF44AE.taxon	discussion	Remarks. The tiny first dorsal-fin spine is an important diagnostic character for this species (Uiblein & Heemstra 2010) that can be best identified by using a binocular microscope and a needle, sometimes after removing scales attached to the anterior dorsal-fin base. In four of 98 studied specimens the spine was found to be reduced to a flap or little knob, making detection more difficult.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFDEFFE7FF27FB7D3B3444B0.taxon	description	(Figures 1, 6 – 8; Tables 2, 4 – 9)	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFDEFFE7FF27FB7D3B3444B0.taxon	materials_examined	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. Non-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.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFDEFFE7FF27FB7D3B3444B0.taxon	diagnosis	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.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFDEFFE7FF27FB7D3B3444B0.taxon	description	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.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFDEFFE7FF27FB7D3B3444B0.taxon	etymology	Etymology. The name refers to the occurrence of oblique conspicuous bars which cross both caudal-fin lobes entirely.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFDEFFE7FF27FB7D3B3444B0.taxon	distribution	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).	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFDEFFE7FF27FB7D3B3444B0.taxon	discussion	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.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFDAFFEEFF27FA0B3C5E410B.taxon	description	(Figures 1, 5, 7; Tables 1, 5, 6)	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFDAFFEEFF27FA0B3C5E410B.taxon	materials_examined	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.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFDAFFEEFF27FA0B3C5E410B.taxon	diagnosis	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.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFDAFFEEFF27FA0B3C5E410B.taxon	description	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.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFDAFFEEFF27FA0B3C5E410B.taxon	etymology	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.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFDAFFEEFF27FA0B3C5E410B.taxon	distribution	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.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFDAFFEEFF27FA0B3C5E410B.taxon	discussion	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.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFD2FFD1FF27FDEB3DF34572.taxon	description	(Figures 1, 9 – 12; Tables 2, 4 – 11)	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFD2FFD1FF27FDEB3DF34572.taxon	description	U. guttatus (Day, 1868): Motomura et al. 2017 (vouchered fresh-colour photo, central Philippines, W Pacific).	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFD2FFD1FF27FDEB3DF34572.taxon	description	Upeneus sp. 1: Uiblein & Lisher 2013 (Nha Trang, South-central Vietnam, South China Sea).	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFD2FFD1FF27FDEB3DF34572.taxon	materials_examined	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 º 13 ′ 21 ′′ N, 109 º 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 º 52.11 ′ N, 109 º 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 º 13 ′ 21 ′′ N, 109 º 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 º 12.09 ′ N, 109 º 12.12 ′ E; HIFIRE F 58203, 105 mm SL (fresh-colour photo), fish market, 12 º 12.10 ′ N, 109 º 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 P 49483, 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 ảng 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 LM 308 (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).	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFD2FFD1FF27FDEB3DF34572.taxon	diagnosis	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.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFD2FFD1FF27FDEB3DF34572.taxon	description	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 (Figure 10 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 10 A), 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 mid-lateral dots in the area of stripe below and posterior to dorsal fins (in 35 % of 100 adults vs. 33 % of 6 subadults); dark pigmentation remains also in other areas: saddle behind second dorsal fin (65 % vs. 67 %), caudal fin (98 % vs. 100 %) and second dorsal-fin distal stripe (92 % vs. 100 %); on caudal fin with retained pigmentation remains of up to 6 oblique bars visible on each lobe; in adults 96 % (of 101 fish) and all subadults show remains of bars on both lobes; of all adults only 2 % show no pigmentation remains (pigmentation degree 0) and 36 % have pigmentation in all body and fin regions (pigmentation degree 4); all studied subadults show pigmentation remains with pigmentation degree 4 occurring in 33 %. Unpigmented areas of caudal, second dorsal fins and other fins entirely pale and partly hyaline.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFD2FFD1FF27FDEB3DF34572.taxon	etymology	Etymology. The name refers to the variable number of 7 or 8 dorsal-fin spines in this species, which is in contrast to most other goatfish species and all other Indo-Pacific congeners which have either 7 or 8 dorsal-fin spines.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFD2FFD1FF27FDEB3DF34572.taxon	distribution	Distribution, habitat and size. W Pacific, from border to EIO in S Indonesia to Singapore, Gulf of Thailand, Vietnam, Central Philippines, South China, Taiwan, and S Japan; 6 – 24 m depth, sandy or muddy bottoms; maximum size 16 cm SL. Intraspecific comparisons. To ensure that the dimorphic expression of seven and eight dorsal-fin spines in U. heterospinus n. sp. is indeed a single-species characteristic, as is unusual in goatfishes, the two morphs were compared using the entire set of studied characters. As was expected, no differences could be discovered in any single character (Table 8), nor in any combinations of characters or in statistical comparisons of morphometric, meristic and colour characters. Subadults of U. heterospinus n. sp. differ from adults in slightly shallower body at anal-fin origin (especially when analyzing populations separately; Figure 11), fewer rudimentary gill rakers on lower limb (Figure 11; while total gill rakers do not differ among size classes), fewer oblique bars on caudal fin (Tables 2, 9), and the bars slightly wider, almost interconnected in fresh subadults (Figure 9 C) vs. narrower and more clearly separated in adults (Figure 9 A, B, D – F; Figure 10). Among the three populations of Upeneus heterospinus n. sp. from N Vietnam, South-central Vietnam and the Gulf of Thailand no clear distinction was found in any single or combination of morphological or colour characters occurs that would support separation into different taxa. However, each population can be distinguished statistically (Table 11). For instance, the Gulf of Thailand population has a significantly higher maximum body depth than the other two populations. The N Vietnam population differs significantly from the two other populations in having a shallower body at anal-fin origin, shallower caudal peduncle, shallower maximum head and suborbital depth, shorter interorbital length, longer pectoral fins and higher second dorsal fin. The South-central Vietnam population has a significantly wider snout and longer barbels than the other two populations and differs in addition in pairwise comparisons with the northern population. As stated for U. margarethae, the considerable amount of significant differences among populations contrasts however with much greater differentiation among species, as revealed by the statistical comparisons among three of the five species (Table 4, see also section on Interspecific comparisons further below). Regarding meristic characters, the Gulf of Thailand population has a significantly lower gill-raker count than the other two populations (Table 5). With respect to preserved colour patterns, the three populations differ significantly from each other with the South-central population showing the highest degree of dark pigmentation, followed by the N Vietnam and the Gulf of Thailand populations (Table 6). In none of the three populations, individuals that completely lack dark pigmentation remains (pigmentation degree 0) occur. When comparing the four examined pigmentation patterns individually, the only significant differences exist in the presence / absence of a saddle and mid-lateral dots (Table 7). The latter are completely lacking in the Gulf of Thailand population. Regarding fresh colour patterns, no clear population differences can be observed when comparing the available images (see also Figures 9, 10).	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFD2FFD1FF27FDEB3DF34572.taxon	discussion	Remarks. Upeneus heterospinus n. sp. has been reported from many areas using various names and most recently as U. spottocaudalis from Japan (Bandai et al. 2018). From Vietnam, where it is a common species that is frequently encountered in fish markets, biological data have been recently collected and published using preliminary identification information (Emel’yanova & Pavlov 2014; Pavlov & Emel’yanova 2016).	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFEDFFD6FF27F9213DAF461E.taxon	description	(Figures 1, 7, 12; Tables 2, 5, 6, 9)	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFEDFFD6FF27F9213DAF461E.taxon	materials_examined	Holotype. (only fresh-colour photo examined). BPBM 33858, adult, 70 mm SL, Coral Sea, New Caledonia, Chesterfield Islands, Chesterfield Bank, 20 ° 59 ’ 48 ’’ S, 158 ° 47 ’ 18 ’’ E, RV Alis, beam trawl, 71 m depth. Paratypes. (4 adults: 73 – 94 mm SL). Coral Sea, New Caledonia, Chesterfield Islands: BPBM 33855, 94 mm SL, Chesterfield Bank, 20 ° 51 ’ 0 ’’ S, 158 ° 45 ’ 00 ’’ E, RV Alis, shrimp trawl, 71 m depth; BPBM 39467, 88 mm SL, Chesterfield Bank, Bellona Reefs, 21 ° 24 ’ 54 ’’ S, 159 ° 09 ’ 18 ’’ E, RV Coriolis, beam trawl, 60 m depth; MNHN 2004 - 1571, 73 mm SL, and USNM 378143, 81 mm SL, Chesterfield Bank, north side, 19 ° 12 ’ 23 ’’ S, 158 ° 42 ’ 02 ’’ E, RV Coriolis, beam trawl, 68 m depth. Non-types. (4 subadults: 46 – 55 mm SL, 2 fresh-colour photos): Vanuatu, NO Alis, chalut à perche: MNHN 2008 - 1459, 46 mm SL, Espirito Santo Island, 15 ° 31.68 ′ S, 167 ° 10.80 ′ E, 36 – 43 m depth (fresh-colour photo); MNHN 2010 - 0616, 51 mm SL, NW Malo Island, 15 ° 39.90 ′ S, 167 ° 03.78 ′ E, 114 – 132 m depth; MNHN 2019 - 0026, 1 of 3, 55 mm SL, 15 ° 37.98 ′ S, 167 ° 03 ′ E, 140 – 175 m depth (wrongly referred to as MNHN 2002 - 0070 in Uiblein et al. 2016); MNHN 2019 - 0027, 1 of 4, 49 mm SL, Malo Island, Bruat Channel, 15 ° 37.32 ′ S, 167 ° 09.60 ′ E, 52 – 66 m depth (fresh-colour photo).	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFEDFFD6FF27F9213DAF461E.taxon	diagnosis	Diagnosis. Dorsal fins VIII + 9, the first spine minute; pectoral fins 12 – 13; gill rakers 6 – 7 + 17 – 18 = 23 – 25; lateral-line scales 29 – 30; measurements in % SL, adults: body depth at first dorsal-fin origin 21 – 24; body depth at anus 19 – 21; caudal-peduncle depth 9.1 – 10; caudal-peduncle width 3.2 – 3.9; maximum head depth 20 – 21; head depth through eye 17 – 18; suborbital depth 9.0 – 11; interorbital length 7.6 – 8.1; head length 29 – 30; snout length 11 – 12; postorbital length 11 – 12; orbit length 7.8 – 8.8; upper-jaw length 9.3 – 11; barbel length 20 – 22; interdorsal distance 12 – 16; caudal-peduncle length 21 – 24; caudal-fin length 27 – 29; anal-fin height 17 – 19; pelvic-fin length 22 – 23; pectoral-fin length 22 – 23; pectoral-fin width 4.0 – 4.3; first dorsal-fin height 20 – 21; second dorsal-fin height 17 – 19; measurements in % SL, subadults: body depth at first dorsal-fin origin 21 – 23; body depth at anus 18 – 20; caudal-peduncle depth 9.1 – 10; caudal-peduncle width 3.0 – 3.5; maximum head depth 19 – 21; head depth through eye 15 – 16; suborbital depth 7.5 – 9.2; interorbital length 7.0 – 7.6; head length 30 – 32; snout length 10 – 12; postorbital length 11 – 13; orbit length 8.4 – 9.1; upper-jaw length 9.7 – 12; barbel length 21 – 23; interdorsal distance 13 – 14; caudal-peduncle length 23 – 26; caudal-fin length 30 – 31; anal-fin height 16 – 17; pelvic-fin length 22 – 23; pectoral-fin length 21 – 22; pectoral-fin width 3.5 – 3.8; first dorsal-fin height 19 – 22; second dorsal-fin height 19 – 20; fresh colour: head and body dorsally orange red or grey mottled, ventrally white rose or silvery, in adults covered with red blotches along ventral margin; caudal fin with 6 dark brown or black oblique bars on both lobes; on upper caudalfin lobe 3 dark brown bars, on lower lobe 3 black bars, in adults the distal-most bar twice as wide as other bars; on both lobes hyaline interspaces mostly larger than bars; mid-laterally on caudal-fin base a brown spot both in adults and subadults; barbels yellow; a single yellow mid-lateral body stripe of pupil width from behind head to caudal peduncle; no dark dots on stripe, dark saddle behind second dorsal fin lacking or only weakly expressed; first dorsal fin with pale green pigmentation, tip pale white, often with closely fused pale-red or pale-grey brown stripes; second dorsal fin with up to 3 brown-red or grey stripes with whitish hyaline interspaces; preserved fish pale, mid-lateral stripe completely lost, yellow barbel colour sometimes retained (Figure 7 G), with remains of dark pigmentation deriving from caudal-fin bars and second dorsal-fin distal stripe or, more rarely, from saddle.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFEDFFD6FF27F9213DAF461E.taxon	distribution	Distribution, habitat and size. Coral Sea, New Caledonia, Chesterfield Islands and Vanuatu off Malo and Spiritu Santo Islands; 36 – 175 m depth, muddy bottoms, adults occur shallower than juveniles; maximum size 9.5 cm SL. Intraspecific comparisons. Subadults differ from adults in shallower head, narrower snout, slightly longer caudal peduncle and first dorsal-fin base, longer caudal fin and narrower pectoral fins (Tables 2, 9).	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFEAFFD4FF27FAE73823420E.taxon	description	(Figures 1, 2, 5; Tables 2, 5, 6, 9)	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFEAFFD4FF27FAE73823420E.taxon	materials_examined	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 & Løppenthin (1944 )) Non-type: Arabian / Persian Gulf: BPBM 29498, subadult, 60 mm SL, Bahrain (fresh-colour photo)	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFEAFFD4FF27FAE73823420E.taxon	diagnosis	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.	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
B829390BFFEAFFD4FF27FAE73823420E.taxon	distribution	Distribution, habitat and size. Arabian / Persian Gulf and inner Gulf of Oman: depth 15 – 20 m; maximum size 11 cm SL. 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).	en	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. Zootaxa 4683 (2): 151-196, DOI: https://doi.org/10.11646/zootaxa.4683.2.1
