Upeneus margarethae Uiblein & Heemstra, 2010

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), pp. 151-196 : 154-167

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.

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 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.

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.

Figures

FIGURE 1. Map showing the distribution of the six species of the margarethae group. Three populations of Upeneus margarethae are indicated by different symbols. Numbers indicate multiple sampling localities.

FIGURE 2. (A–F) Upeneus margarethae; (A) HT, SAIAB 82217, 82 mm SL, WIO, Mozambique, off Beira (O. Alvheim); (B) SMF 35030, 90 mm SL, Red Sea, Saudi Arabia, off Jizan, (S.V. Bogorodsky); (C) 86 mm SL, Tuticorin, S India (K.K. Bineesh); (D) SAIAB 203480, 95 mm SL, EIO, Myanmar, NW od Basuhino Island (P. Psomadakis; side-reversed image); (E) CSIRO CA 3052, 98 mm SL, EIO, NW Australia, off Port Hedland (CSIRO staff); (F) subadult, 47 mm SL, EIO, Thailand, Kampuan Mangrove forest, Suksamran, Ranong (S. Ratmuangkhwang); (G, H) U. randalli: (G) HT, BPBM 33180, 101 mm SL, Arabian/ Persian Gulf, off S Kuwait, (J.E. Randall); (H) BPBM29498, 60 mm SL, Bahrain (J.E. Randall).

FIGURE 3. Upeneus margarethae photographed in situ in Mozambique, WIO; (A) subadult or small adult, Pemba Bay, N Mozambique, near Pemba harbour (R. Koch); (B) at right: subadult or small adult, with subadult U. heemstra (at left), Pomene, South-central Mozambique (M. & V. Fraser); (C) adult, resting on bottom, Lunene Island, off Vilanculos, South-central Mozambique (A. Lund).

FIGURE 4. Three selected morphological characters against SL in Upeneus margarethae. The 65 mm demarcation between subadults and adults is indicated by a dotted black dash. The three populations are indicated by different symbols.

FIGURE 5. Four morphometric characters in adults of three Upeneus species of the margarethae group against SL and each other. For Upeneus margarethae, the three populations are indicated by different symbols.

FIGURE 6. Two morphometric characters against SL and each other and second dorsal-fin height against total number of gill rakers in adults of two margarethae-group species. For Upeneus margarethae, the three populations are indicated by different symbols. The distinction between U. caudofasciatus n. sp. and the EIO/N Australian population of U. margarethae is indicated by dashed and dotted outlines, respectively.