Brachypterois serrulifer Fowler 1938

Matsunuma, Mizuki, Sakurai, Makoto & Motomura, Hiroyuki, 2013, Revision of the Indo-West Pacific genus Brachypterois (Scorpaenidae: Pteroinae), with description of a new species from northeastern Australia, Zootaxa 3693 (4), pp. 401-440 : 426-432

Brachypterois serrulifer Fowler 1938

[New English name: Sawmaxilla Scorpionfish]

[New Standard Japanese name: Yokoba-nokogiri-kasago] ( Figures 1 , 6 C, 7, 8G–I, 9G–I, 10–11, 14B, 16–17, 19, 20C; Tables 1 , 4 )

Sebastes serrulatus (not of Richardson): Alcock 1889: 298, pl. 22, fig. 2 (off Ganjam, Bay of Bengal, India; description); Alcock 1896: 312 (off Ganjam, Bay of Bengal, India; based on Alcock 1889)

Brachypterois serrulifer Fowler 1938: 79 , fig. 35 [type locality: 8.4 miles (ca. 13.5 km) off San Fernando Point Light, west coast of Luzon Island, Philippines (16°39′36″N, 120°11′06″E), 45 fathoms (ca. 82.3 m) depth]; Herre 1945: 401 (Bay of Bengal, India; comments on distribution in Bay of Bengal; based on 17 specimens, 7 of which examined herein); Herre 1952: 432 (Manila Bay and Ragay Gulf, Luzon, Philippines and Bay of Bengal, India; description; based on 14 and 15 specimens from Philippines and India, respectively, 14 and 7 of which examined herein); Herre 1953: 563 ( Philippines and Bay of Bengal, India; listed; based on Herre 1945, 1952); Aprieto & Villoso 1977: 43 (Visayan Sea, Philippines; listed); Fourmanoir 1981: 97 (south off Lubang Island, Philippines; listed); Fourmanoir 1985: 52 ( Philippines; listed; based on Fourmanoir 1981)

Ranipterois serrulifer: Whitley 1950: 407 (new combination; see Remarks under Brachypterois )

Brachypterois serrulatus (not of Richardson): Kotthaus 1979: 25, fig. 476 (off Bombay, India; short description; 3 specimens examined herein); Gloerfelt-Tarp & Kailola 1984: 108, unnumbered fig. on top left ( Indonesia; short description; 2 specimens examined herein); Blaber et al. 1994: 392 (Gulf of Carpentaria, Australia; listed); Sujatha 1995: 57 (Bay of Bengal, India; listed); Martin et al. 1995: 917 (Gulf of Carpentaria, Australia; listed); Manilo & Bogorodsky 2003: S102 (Arabian Sea; listed)

Brachypterois serrulata (not of Richardson): Dor 1984: 85 (Red Sea; listed); Russell & Houston 1989: 81 (Arafura Sea; listed); Randall 1995: 106, fig. 227 (Gulf of Oman; short description); Krupp et al. 2000: 325, fig. 1 (off Iran, Arabian Gulf; short description; 4 specimens examined herein); Allen et al. 2003: 372, unnumbered fig. on top left [in part; Indo-West Pacific; diagnosis and ecological note; underwater photograph from North Sulawesi (G. Allen, pers. comm.)]; Allen et al. 2006: 877 (Timor and Arafura seas, Australia; review of synonymy and distribution in Australia); Kuiter & Debelius 2006: 187, unnumbered fig. on top left (in part; Indo-West Pacific; listed); Bijukumar & Deepthi 2009: 149 (off Kerala, India; listed); Al-Jufaili et al. 2010: 21 ( Oman; listed); Golani & Bogorodsky 2010: 22 (Red Sea; listed); Allen & Erdmann 2012: 217, unnumbered fig. on top ( Indonesia; short description and ecological note); Rainboth et al. 2012: 80, pl. 55, fig. 1136 (Gulf of Thailand; listed; figure taken from Gloerfelt-Tarp & Kailola 1984)

? Brachypterois serrulifer: Fourmanoir & Nhu-Nhung 1965: 94 (Nha Trang Bay, Viet Nam; short description)

Holotype. USNM 98886, 78.7 mm SL, 8.4 miles (ca. 13.5 km) off San Fernando Point Light, west coast of Luzon, Philippines (16°39′36″N, 120°11′06″E), 45 fathoms (ca. 82.3 m), R/V Albatross, 10 May 1909.

Non-type specimens. 109 specimens, 40.4–94.5 mm SL— JAPAN: HUMZ 36733 (X), 82.3 mm SL, Tosa Bay, Kochi (obtained at Mimase Port), K. Amaoka, 1956. TAIWAN: KAUM–I. 39187 (X), female, 61.3 mm SL, off Kaohsiung, H.-C. Ho, trawl, 27 June 2011; NMMB-P 2808 (X), 70.9 mm SL, Tong Gang, Pingtung, 100 m, J.- H. Wu, 8 Nov. 2001; NMMB-P 12647 (3 specimens; X), 61.2–85.1 mm SL, Tong Gang, Pingtung, M.-C. Yu, 16 Feb. 1960. PHILIPPINES: CAS 29406 (4), 59.2–64.8 mm SL, Nasipit, Agusan del Norte, Mindanao, F. B. Steiner, 15 Apr. 1973; CAS 49547 (14), 51.0– 79.3 mm SL, Manila Bay, Luzon, A. W. Herre, 25 May 1948; NSMT- P 109949, 74.7 mm SL, E. Fujii; USNM 168172, 40.4 mm SL, Manila Bay to Lingayen Gulf, west coast of Luzon, 1–4 m, R/V Albatross, 11 May 1909. MALAYSIA: KAUM–I. 22033 (X), male, 83.0 mm SL, KAUM–I. 22035 (X), female, 63.9 mm SL, off Kota Kinabalu, Sabah, Borneo, G. Ogihara, 11 Aug. 2009; KAUM–I. 49286, 76.0 mm SL, off Kota Kinabalu, Sabah, Borneo, H. Nishiyama, 2 Aug. 2012. THAILAND (Andaman Sea coast): KAUM–I. 47580, 73.3 mm SL, KAUM–I. 47581, 64.1 mm SL, off Ranong Province (obtained at Kuraburi Fishing Port), T. Yoshida & H. Nishiyama, 27 June 2012. INDONESIA: NTM S. 11016-003 (2), 64.0– 68.7 mm SL, Mentawai Strait, Sumatra (0°04′S, 99°01′E), 50 m, T. Gloerfelt-Tarp, 6 May 1983. AUSTRALIA: AMS I. 21842- 0 0 9 (2; X), 68.3–85.6 mm SL, Arafura Sea (10°35′S, 133°48′E), R/V Soela; AMS I. 26288-001, 79.8 mm SL, south east corner of Gulf of Carpentaria, 64 m, R/V Laakanuki, bottom trawl, 29 Apr. 1964; CSIRO A2507 (X), 80.6 mm SL, Arafura Sea (13°33′S, 136°57′E), 34.2 m, 4 Sept. 1963; NTM S. 10031-075 (4), 63.1–84.9 mm SL, north of Smith Point, Cobourg Peninsula (12°58′S, 132°10′E), 27 m, H. Larson, 18 Oct. 1981; NTM S. 11656-001, 92.1 mm SL, north west of Cape Wessel (10°25′S, 136°55′E), 64 m, R. Williams, 1 Feb. 1985; NTM S. 11659-006 (2), 84.6–93.9 mm SL, northern Cape Wessel (10°25′S, 136°40′E), 60 m, R. Williams; NTM S. 11895-010, 88.7 mm SL, northern Cape Wessel (10°15′S, 136°40′E), 55–60 m, M. Baron, 20 Feb. 1986; NTM S. 11896-004 (X), 92.0 mm SL, northern Cape Wessel (10°14′S, 136°34′E), 50–55 m, NT Fisheries, 20 Feb. 1986; NTM S. 12262- 0 0 1, 53.2 mm SL, north east of Goulburn Islands (10°20′S, 134°29′E), 59 m, NT Fisheries, 11 June 1987; NTM S. 12474-003, 89.2 mm SL, north of Groote Eylandt (13°16′S, 136°51′E), 37 m, R. Williams, 24 Aug. 1987; NTM S. 13038-001 (2), 70.6–77.0 mm SL, south east of Cape Grey, Gulf of Carpentaria (13°05′S, 136°45′E), 39 m, NT Fisheries, 23 Aug. 1987; NTM S. 13253-001 (X), 84.6 mm SL, northern Cape Wessel (10°14′S, 136°53′E), 53 m, R. Williams, 21 Nov. 1991; NTM S. 13260-002 (2), 55.4–78.8 mm SL, Gulf of Carpentaria (13°02′S, 139°22′E), 58 m, R. Williams, 25 Nov. 1991; NTM S. 13270-002 (3), 54.3–75.2 mm SL, Gulf of Carpentaria (11°04′S, 139°56′E), 57 m, R. Williams, 29 Nov. 1991. MYANMAR: CAS 64830 (4; 1 female, 61.2 mm SL), 52.7–78.0 mm SL, Bay of Bengal (15°08′N, 94°54′E), 35 m, R/V Anton Bruun, shrimp trawl, 1 Apr. 1963; CAS 211494 (4; 1 female, 57.3 mm SL), 54.5–70.1 mm SL, Bay of Bengal (15°08′N, 94°54′E), R/V Anton Bruun, 1 Apr. 1963. INDIA: CAS 207661 (5; 1 female, 60.8 mm SL), 60.8–66.6 mm SL, Neendakara, Kerala (8°56′N, 76°30′E), bottom trawl, 8 Oct. 1966; CAS 29590, 92.2 mm SL, 10 miles off Bombay, 5–10 m, F. B. Steiner, trawl, 4–10 Nov. 1973; SU 14649 (7; 1 female, 55.0 mm SL), 51.8–77.7 mm SL, mouth of Hugli River, Bay of Bengal, A. Herre, R/ V Lady Fraser; ZMH 5602 (3), 77.9– 66.5 mm SL, approximately 200 m off Mumbai, 70–84 m, H. Kotthaus, R/V Meteor, bottom trawl, 27 Feb. 1965. PERSIAN GULF: SMF 26000, 45.9 mm SL, 28°56.494′N, 49°43.812′E, 45 m, H. Zetzsche, bottom trawl, 14 Dec. 1991; SMF 26001 (3), 56.1–73.7 mm SL, 29°13.587′N, 49°53.955′E, 41 m, H. Zetzsche, bottom trawl, 10 Nov. 1991. OMAN: BPBM 21416, male, 76.0 mm SL, 5.6 km north of Khalil, Gulf of Oman (23°38′N, 58°21.7′E), 27.4 m, G. R. Allen, 10 Mar. 1977; CAS 60396 (19; 3 females, 65.9–67.9 mm SL), 58.7–86.4 mm SL, Gulf of Oman (26°10–13′N, 57°02′E), 64–65 m, R/V Anton Bruun, trawl, 30 Nov. 1963; WAM P. 25996.001, 72.5 mm SL, 10 km north of Khalil, Gulf of Oman (23°46′N, 58°20′E), 30–66 m, G. R. Allen, 10 Mar. 1977. ERITREA: CAS 209665 (2), 76.9–88.0 mm SL, Massawa Bay (15°40′N, 39°36′E), 79 m, L. W. Knapp, otter trawl, 20 Sept. 1971; CAS 64825, 63.6 mm SL, Massawa Bay (15°40′N, 40°23′E), 79 m, L. W. Knapp, otter trawl, 21 Sept. 1971. MADAGASCAR: MNHN 2004-0028, 94.5 mm SL, 23°36′0″S, 43°31′59″E, 340–360 m, 27 Feb. 1973.

Diagnosis. A species of Brachypterois with the following combination of characters: pectoral-fin rays 15–17 (modally 16); scale rows in longitudinal series 43–51 (47); scales above lateral line 4–7 (5); scale rows between last dorsal spine base and lateral line 4–7 (5); pre-dorsal scale rows 3–6 (4); suborbital scale rows 0–2 (1) ( Fig. 6 C); total gill rakers 15–17 (16); head length 38.8–44.6 (mean 41.7) %SL ( Fig. 7 ); orbit diameter 11.8–15.1 (13.0) %SL; upper-jaw length 19.1–22.2 (20.7) %SL; longest dorsal-fin soft ray length 20.5–26.2 (23.4) %SL; longest anal-fin soft ray length 23.3–29.0 (26.2) %SL; caudal-fin length 36.1–45.1 (40.1) %SL; posterior lacrimal spine usually directed ventrally ( Fig. 8 G–I); spine(s) on corner of outer angular ridge not especially developed, size similar to that of other spines on ridge, directed ventrally ( Fig. 9 G–I); 0–22 (mean 3.9) spines on median lateral ridge on maxilla in specimens> 58 mm SL ( Fig. 14 B); relatively few large spots on caudal fin, 2–10 (mean 5.2) spots on longest caudal-fin ray ( Figs. 10 , 16–17 ).

Color of fresh specimens. Head and body pinkish-brown, darker dorsally, paler ventrally; a poorly defined large black blotch (subequal to orbit diameter) on opercle. Five reddish-brown to brown bands on head; anteriormost band from anterior margin of eye running over snout to tip of lower jaw; second extending from anteroventral margin of eye onto lacrimal and both jaws; third from just below eye to posterodorsal corner of maxilla; fourth from posteroventral margin of eye to posterior portion of opercle; fifth from posterior margin of eye to opercle, more or less continuous with blotch on opercle. Five poorly defined reddish-brown to brown vertical bands on body; anterior-most band extending from between first and third dorsal-fin spine bases; second from between fourth and sixth spine bases; third from between eighth and penultimate spine bases; fourth Y-shaped, upper margin of anterior branch below last spine and fourth soft ray bases, upper margin of posterior branch below seventh and penultimate soft ray bases, both branches connected at level of lateral line or just below dorsal-fin base; fifth on caudal peduncle just in front of caudal-fin base. Ground color of fins similar to that of head and body. Dorsal fin with a large dark red to brown blotch on spinous portion between about sixth and ninth spines; about 4– 5 more or less dotted dark red lines on soft-rayed portion. Anal fin with about 5 more or less dotted dark red lines. Pectoral fin membrane dark brown, with about 6–8 poorly defined narrow dark red bands. Pelvic fin darker marginally, with about 5–8 poorly defined narrow dark red bands. Caudal fin membrane translucent with dark red narrow band marginally; about 20–60 small dark red to brown spots [maximum diameter 2.1–6.0 (mean 3.4) times in caudal-fin ray width at position where spot present on ray; based on 39 preserved specimens] on rays. Eye vivid yellow; iris dark blue. Color photographs of live individuals from Indonesia available in Allen et al. (2003), Kuiter & Debelius (2006), and Allen & Erdmann (2012) as B. serrulata .

Color of preserved specimens. Head and body creamy-white to brown, darker dorsally, paler ventrally; formerly reddish markings brownish and indistinct. Eye dark blue to gray; iris white to dark blue.

Distribution. Brachypterois serrulifer is widely distributed in the Indo-west Pacific region, including Madagascar, Red Sea and Persian Gulf, east to the Timor Sea and the Philippines, South and East China seas, and north to Tosa Bay, Kochi, southern Japan ( Fig. 11 ). The specimens of the 18 lots examined were from depths of 1– 360 m (mostly deeper than 50 m). According to underwater photographs of B. serrulifer (as B. serrulata ), included in Allen et al. (2003) and Allen & Erdmann (2012), the species also occurs off Bali and North Sulawesi, Indonesia. Brachypterois serrulifer and B. serrulata occur sympatrically off Borneo, southern Taiwan and Tosa Bay, southern Japan. However, the occurrence of B. serrulifer in Japan is rare, with only a single specimen recorded (vs. 51 specimens of B. serrulata recorded).

Brachypterois serrulifer is apparently not distributed off the northwestern coast of Western Australia, despite occurring off the northern coast. Allen (1997) and Allen et al. (2006) recorded B. serrulata from northwestern and northern coasts of Western Australia and northern coast of Northern Territory. However, according to distributional records of B. serrulifer we examined ( Fig. 11 ), the records of B. serrulata from the northern coasts of both states are believed to be misidentifications of B. serrulifer . Among the specimens examined only a single putative example of Brachypterois (NMV A24244, 24.4 mm SL), collected from northwest of Dampier, WA, and identified and registered as B. serrulata , was collected from Western Australia. However, that specimen is now re-identified here as Dendrochirus brachypterus (Cuvier in Cuvier & Valenciennes 1829) ( Scorpaenidae ).

Remarks. Meristics and morphometrics of specimens of B. serrulifer examined are shown in Tables 1 , 4 . The species was originally described by Fowler (1938) on the basis of a single specimen (USNM 98886, 78.7 mm SL; Fig 16 D) collected off San Fernando Point Light, west coast of Luzon, Philippines (16°39′36″N, 120°11′06″E) at a depth of 45 fathoms (ca. 82.3 m). It has been regarded by subsequent researchers (e.g., Chu 1962; Kanayama & Amaoka 1981; and Allen et al. 2006) as a junior synonym of S. serrulatus Richardson 1846 . However, examination of the holotype of B. serrulifer shows it to have the same diagnostic characters as specimens included herein under that name (see Comparisons). Brachypterois serrulifer is therefore regarded as valid. Fourmanoir & Nhu-Nhung (1965), Fourmanoir (1981, 1985) and several other authors also referred to B. serrulifer as a valid name (see synonym list), although there is no evidence that they recognized both nominal species, B. serrulata and B. serrulifer . Therefore, the record of B. serrulifer from Vietnam given by Fourmanoir & Nhu-Nhung (1965) is doubtful and probably applicable to B. serrulata .

Alcock (1889) first recorded B. serrulifer from the Bay of Bengal (as B. serrulata ), on the basis of a single specimen not examined in this study. However, the detailed description and figure of the specimen given by Alcock (1889) clearly shows it to be B. serrulifer . Brachypterois sp. reported by Oyugi (2005) from Kilifi, Kenya, but lacking information for positive identification, is most likely to represent B. serrulifer , as there are no other congeners distributed in the western Indian Ocean.

Biological notes. A female of 63.9 mm SL (KAUM–I. 22035) collected off Borneo in August 2009 had developed eggs approaching the terminal stage (about 0.5 mm in diameter). A stomach content analysis (1 specimen) showed that B. serrulifer feeds on small shrimps and fishes.

Geographical variation. A difference in the number of caudal fin spots in B. serrulifer was noted between specimens collected from the Indian ( Figs. 16 B, 17B, D) and Pacific oceans, including northern Australia ( Figs. 16 A, C–E, 17A, C). The former specimens had relatively fewer and larger spots on the caudal fin, compared with their Pacific Ocean counterparts ( Fig. 10 ) [maximum diameter of a basal spot on a middle caudal-fin ray 2.8–4.0 (mean 3.5), 2.1–6.0 (3.5), 2.6–6.0 (3.6), 2.3–3.8 (2.9) and 2.7–4.7 (3.3) times the caudal-fin ray width at the position where spot present on ray in specimens from the Persian Gulf (3 specimens), Arabian Sea (13), Bay of Bengal and Andaman Sea (8), northern Australia (8) and Philippines (7), respectively]. Since no other significant differences were found between the two groups, the difference is regarded herein as representing geographical variation in B. serrulifer .

a Upper unbranched + branched + lower unbranched rays = total rays on left side of body/ total rays on right side. b Rakers on upper limb + rakers on lower limb = total rakers.

c At vertical midline of eye.

d At posterior end of preocular spine base.

DS and LL indicate dorsal-fin spine base and lateral line, respectively.

Figures

FIGURE 1. Lateral (top) and dorsal (bottom) views of maxilla of Brachypterois serrulifer, KAUM – I. 22033, 82.1 mm SL. 1 Spines on anterior portion of articular head; 2 Spines on lateral portion of articular head; 3 Spines on median lateral ridge; 4 Dorsal ridge. Left to anterior. Bar indicates 3 mm.

FIGURE 6. Relationships of number of suborbital scale rows to standard length in (A) Brachypterois curvispina sp. nov., (B) B. serrulata and (C) B. serrulifer.

FIGURE 7. Relationships of: (A) head length; (B) orbit diameter; (C) upper jaw length; (D) longest dorsal-fin soft ray length; (E) longest anal-fin soft ray length; and (F) caudal-fin length to standard length in Brachypterois curvispina sp. nov. (stars, solid lines), B. serrulata (circles, dotted lines) and B. serrulifer (triangles, chain lines). H 1 holotype of B. curvispina; H 2 holotype of B. serrulifer; N neotype of Sebastes serrulatus.

FIGURE 8. Spines on lacrimal bones of Brachypterois curvispina sp. nov. (A – C), B. serrulata (D – F) and B. serrulifer (G – I). (A) CSIRO H 7204 - 02, male, 78.5 mm SL, holotype of B. curvispina; (B) CSIRO H 6784 - 02, female, 75.2 mm SL, paratype of B. curvispina; (C) CSIRO H 6750 - 01, sex undetermined, 57.4 mm SL; (D) KAUM – I. 24056, male, 97.9 mm SL; (E) KAUM – I. 24054, female, 76.5 mm SL; (F) KAUM – I. 24050, female, 63.9 mm SL; (G) KAUM – I. 22033, male, 83.0 mm SL; (H) CAS 60396 (1 of 19 specimens), female, 67.9 mm SL; (I) CAS 49547 (1 of 14), sex undetermined, 57.6 mm SL. LLS lateral lacrimal spine; ALS anterior lacrimal spine; PLS posterior lacrimal spine. Left to anterior. Bars indicate 5 mm.

FIGURE 9. Spines on angular bones of Brachypterois curvispina sp. nov. (A – C), B. serrulata (D – F) and B. serrulifer (G – I). (A) CSIRO H 7204 - 02, male, 78.5 mm SL, holotype of B. curvispina; (B) CSIRO H 6784 - 02, female, 75.2 mm SL, paratype of B. curvispina; (C) CSIRO H 6750 - 01, sex undetermined, 57.4 mm SL; (D) KAUM – I. 24056, male, 97.9 mm SL; (E) KAUM – I. 24054, female, 76.5 mm SL; (F) KAUM – I. 24050, female, 63.9 mm SL; (G) KAUM – I. 22033, male, 83.0 mm SL; (H) CAS 60396 (1 of 19 specimens), female, 67.9 mm SL; (I) CAS 49547 (1 of 14), sex undetermined, 57.6 mm SL. Arrows indicate spine (s) on posterior corner of outer angular ridge. Left to anterior. Bars indicate 1 mm.

FIGURE 10. Relationships of number of spots on longest caudal-fin ray to standard length in Brachypterois curvispina sp. nov. (stars), B. serrulata (circles) and B. serrulifer (triangles; dotted symbols indicate specimens from the Indian Ocean, remaining specimens from the Pacific Ocean). H holotype of B. serrulifer. Note: counts unable to be made in holotype of B. curvispina and neotype of Sebastes serrulatus.

FIGURE 11. Distributional records of Brachypterois curvispina sp. nov. (stars), B. serrulata (circles) and B. serrulifer (triangles) based on collected specimens examined in this study. Closed symbols indicate type localities.

FIGURE 14. Relationships of number of spines on median lateral ridge of maxilla to standard length in (A) Brachypterois serrulata and (B) B. serrulifer. H holotype of B. serrulifer. Note: all specimens examined, including type series, of B. curvispina sp. nov. and neotype of Sebastes serrulatus lacked spines.

FIGURE 16. Preserved specimens of Brachypterois serrulifer. (A) USNM 168172, 40.4 mm SL; (B) CAS 64830 (1 of 4 specimens), 62.3 mm SL; (C) CAS 29406 (1 of 4), 64.8 mm SL; (D) USNM 98886, 78.7 mm SL, holotype of B. serrulifer; (E) NTM S. 10031 - 075 (1 of 4), 84.9 mm SL.

FIGURE 17. Fresh specimens of Brachypterois serrulifer. (A) KAUM – I. 39187, female, 61.3 mm SL; (B) KAUM – I. 22033, male, 83.0 mm SL; (C) KAUM – I. 47580, 73.3 mm SL; (D) size unknown, off Sonmiani, Pakistan, Oct. 2009 (photo: H. B. Osmany).

Tables

TABLE 4. Meristics and morphometrics, expressed as percentages of standard length, of Brachypterois serrulifer.

SL (mm)	Holotype USNM 98886 78.7	Non-types (n = 109) 40.4–94.5
Meristics Dorsal-fin rays Anal-fin rays	XIII, 10 III, 5	Modes XII–XIII, 9–11 XIII, 10 III, 4–5 III, 5
Pectoral-fin rays a	ii + 8 + vi = 16/16	i–iii + 6–8 + vi–viii = ii + 7 + vii = 16/16 15–17/15–17
Scale rows in longitudinal series Pored lateral-line scales Scales above lateral line	48 25 6	43–51 47 23–26 24 4–7 5
Scales below lateral line Scale rows between 6th DS and LL Scale rows between last DS and LL	11 5 5	9–12 10 4–6 5 4–7 5
Pre-dorsal scale rows Suborbital scale rows Gill rakers b Morphometrics (% SL) Body depth	5 1 5 + 11 = 16 35.5	3–6 4 0–2 1 4–6 + 10–12 = 15–18 5 + 11 = 16 Means 32.0–40.0 36.6
Body depth at anal-fin origin Body width	30.1 22.2	25.8–33.1 29.4 18.1–27.4 22.1
Head length Head width Snout length Orbit diameter	42.7 14.7 8.1 13.5	38.8–44.6 41.7 13.7–16.1 14.7 7.2–9.2 8.2 11.8–15.1 13.0
Interorbital width c Interorbital width d	5.0 4.6	4.7–6.9 5.8 4.4–6.5 5.3
Upper-jaw length Maxillary depth Suborbital depth Postorbital length	21.0 7.9 1.5 20.6	19.1–22.2 20.7 6.1–8.4 7.5 0.3–1.9 0.8 19.1–23.1 20.9
Predorsal-fin length Preanal-fin length	34.3 72.7	31.3–36.1 33.6 68.2–77.5 72.8
Prepelvic-fin length 1st dorsal-fin spine length 2nd dorsal-fin spine length 3rd dorsal-fin spine length	38.6 — — —	35.3–41.3 38.4 7.4–13.6 10.3 12.4–24.5 18.2 14.8–26.4 19.4
4th dorsal-fin spine length 5th dorsal-fin spine length	— —	15.1–23.4 18.9 14.3–23.5 18.7
6th dorsal-fin spine length 7th dorsal-fin spine length 8th dorsal-fin spine length 9th dorsal-fin spine length	— — — —	15.3–22.6 18.7 14.5–22.5 18.6 15.1–21.7 17.9 12.7–18.0 15.4
10th dorsal-fin spine length	—	8.8–15.7 11.6 ......continued on the next page