Sphyraena stellata, anew barracudafromthe Indo-Pacific, with redescriptionsofS. helleri Jenkins, 1901 and S. novaehollandiae Günther, 1860 (Perciformes: Sphyraenidae) Author Morishita, Satoshi The Kagoshima University Museum, 1 - 21 - 30 Korimoto, Kagoshima 890 - 0065, Japan. & satopool 05060323 @ yahoo. co. jp; https: // orcid. org / 0000 - 0003 - 2349 - 3349 Author Motomura, Hiroyuki The Kagoshima University Museum, 1 - 21 - 30 Korimoto, Kagoshima 890 - 0065, Japan. & motomura @ kaum. kagoshima-u. ac. jp; http: // orcid. org / 0000 - 0002 - 7448 - 2482 text Zootaxa 2020 2020-05-11 4772 3 545 566 journal article 22257 10.11646/zootaxa.4772.3.6 0754157d-d8d3-4076-b8bf-0d932c7dc5ec 1175-5326 3819738 7092DA88-22A8-4606-9A77-235DA17263C4 Sphyraena stellata n. sp. [New English name: Yellow-striped Barracuda; Japanese name: Hoso-kamasu] ( Figures 2C , 4 , 7–10 , 11 C–F, 12; Tables 1 , 2 ) Sphyraena africana (not of Gilchrist & Thompson): Senou 2014: 577 , unnumbered fig. [Yoron-jima island, Amami Islands, Japan (based on KAUM–I. 55091)]. Sphyraena novaehollandiae (not of Günther): Smith 1956: 40 , fig. 1; Talwar & Kacker 1984: 86 ; Yoshino 1984: 121 , pl. 106, fig. J. ( Japan ); Senou 1997: 654 , unnumbered figs. (Iriomote-jima island, Yaeyama Islands, Japan ). Sphyraena helleri (not of Jenkins): Thinker 1978: 186 , unnumbered figs. (Hawaiian Islands); Hoover 1993: 18 , unnumbered fig. (Puako, Hawai’i Island, Hawaiian Islands); Randall et al . 1997a: 292 (Great Barrier Reef); Randall et al . 1997b: 60 , pl. 19C (Ogasawara Islands, Japan ); Senou 2001: 3693 , unnumbered fig. (western Central Pacific); Senou et al . 2002: 230 [Hachijo-jima island, Izu Islands, Japan (based on 3 photographs, KPM-NR 8888, 9268, 11531); Randall 2005: 607 , unnumbered fig. ( Marshall Islands ); Randall 2007: 440 , unnumbered figs. (Maui Island, Hawaii island, Hawaii Islands); Senou et al . 2007: 71 [Miyako Islands, Japan (based on 2 photographs, KPM-NR 68291, 68292); Randall & Cea 2011: 121 , unnumbered fig. (Easter Island); Allen & Erdman 2012: 1038 , unnumbered fig. ( Phoenix Islands ); Senou 2013: 1639 , unnumbered fig. (Pacific Ocean); Ikeda & Nakabo 2015: 222 , PL 221-1 [Susami, Wakayama , Japan (based on WW 10902 (1))]; Koeda et al . 2016: 105 , fig. 488. (Yonaguni-jima island, Yaeyama Islands, Japan ); Jeong 2017: 57 , unnumbered fig. [ Kagoshima Bay, Kagoshima Pref. , Japan (based on KAUM–I. 14726)]; Kimura et al . 2017: 170 , fig. 4 [Kuchinoerabujima island, Osumi Islands, Japan (based on KYUM-PI 4959)]; Nakae et al . 2018: 334 [Amami-oshima island, Amami Islands, Japan (based on 2 specimens , KAUM–I. 108200, NSMT-P 131265 )]. Senou 2019: 394 , unnumbered figs. [Amami Islands, Japan (based on 3 specimens , KAUM–I. 55091, 108200, 122320 )]. Holotype . KAUM–I . 122319, 437.0 mm SL, off Okidomari Port , Okinoerabu-jima island , Amami Islands , Japan , 27°24′16′′N , 128°32′57′′E , 20 m , line-fishing, J. Nakamura , 23 Oct. 2018 . FIGURE 6. Cyanine Blue stained heads of Sphyraena novaehollandiae showing sensory canal pores. (A) entire head of AMS I 44768-004, 423.8 mm SL; (B and C) suborbital regions of AMS IA 659, 429.9 mm SL and AMS I 44768-001, 403.7 mm SL, respectively. Paratypes . 40 specimens , 98.0–587.0 mm SL. MALDIVES : BPBM 34729 , 415.0 mm SL, Maldives Islands . JAPAN : Honshu : FRLM 11241, 243.4 mm SL, Ago Bay , Shima , Mie Pref. , 34°28′23′′– 34°28′38′′N , 136°75′43′′– 136°77′2′′ E, S. Kimura , 22 Aug. 1991 ; FRLM 28489, 293.6 mm SL, Ago Bay , Shima , Mie Pref. , H. Nakamura , 28 Sep. 2001 ; FRLM 34109, 286.3 mm SL, Kumano-nada , Owase , Mie Pref. , K. Tsumoto , 30Aug. 2007 ; FRLM 39546, 315.1 mm SL, Ago Bay , Shima , Mie Pref. , 34°28′23′′– 34°28′38′′N , 136°75′43′′–136°77′2′′ E, D. Sasaki et al. , 28 Oct. 2011 ; FRLM 40372, 283.6 mm SL, Ago Bay , Shima , Mie Pref. , 34°28′23′′– 34°28′38′′N , 136°75′43′′–136°77′2′′ E, D. Sasaki et al. , 6 Jan. 2012 ; FRLM 42172, 216.5 mm SL, Ago Bay , Shima , Mie Pref. , 34°28′23′′– 34°28′38′′N , 136°75′43′′–136°77′2′′ E, S. Kimura et al. , 12 Sep. 2012 ; FRLM 46606 , 316.0 mm SL, Kumano-nada , Owase , Mie Pref. , Y. Hibino , 15 Dec. 2013 . Izu Islands : NSMT-P 131265 , 414.0 mm SL, Tori-shima island , 28°28′N , 129°28′E , 7 May 2017 . Kyushu : KAUM–I . 14726, 334.0 mm SL, off Izashiki Port , Kagoshima Bay , Kagoshima Pref. , 31°05′N , 130°41′E , 30–40 m , set net , M. Yamada , 20 Jan. 2009 ; KAUM–I . 22798, 126.5 mm SL, Kasasa , Minamisatsuma , Kagoshima Pref. , 31°25′44′′N , 130°11′49E , 27 m , set net , M. Itou , 11 Jun. 2009 ; KAUM–I . 120245, 275.4 mm SL, Uchinoura Bay , Kimotsuki , Kagoshima Pref. , 31°17′29′′N , 131°06′59′′E , 40 m , set net , H. Hata et al. , 13 Sep. 2018 ; Ryukyu Islands : KAUM–I . 107721, 307.5 mm SL, off Osaki , Tanega-shima island, Osumi Islands , 30°39′N , 130°54′E , 10–15 m , roll net , FV An’ei-maru , 31 Aug. 2017 ; KAUM–I . 115064, 393.1 mm SL, KAUM–I . 115088, 412.1 mm SL, KAUM–I . 115846, 431.8 mm SL, KAUM–I . 115847, 466.3 mm SL, KAUM–I . 115848, 446.8 mm SL, KAUM–I . 115849, 430.0 mm SL, KAUM–I . 115850, 380.7 mm SL, off Kiriishi Port , Suwanose-jima island , Tokara Islands , 29°36′32′′N , 129°42′53′′E , 20 m , line-fishing, B. Jeong et al. , 26 Apr. 2018 ; KAUM–I . 55091, 123.0 mm SL, Chabana Port , Yoron-jima island , Amami Islands , 27°02′57′′N , 128°24′19′′E , 0.5 m , night light, H. Hata , 30 Jun. 2013 ; KAUM–I . 65293, 421.0 mm SL, KAUM–I . 65294, 434.5 mm SL, off Okinoerabu-jima island , Amami Islands , 27°22′N , 128°34′E , Y. Sakurai , 28 Jun. 2014 ; KAUM–I . 108200, 280.4 mm SL, off Amami-oshima island , Amami Islands , 28°28′N , 129°28′E , T . Maekawa , 7 May 2016 ; KAUM–I . 122320, 411.0 mm SL, KAUM–I . 122321, 433.5 mm SL, off Okidomari Port , Okinoerabu-jima island , Amami Islands , 27°24′16′′N , 128°32′57′′E , 20 m , line-fishing, K. Uehara , 23 Oct. 2018 ; NSMT-P 103279 , 451.0 mm SL, Amami-oshima island , Amami Islands , 28°28′N , 129°28′E , 19 Jul. 2010 , Purchased at Naze Fish Market ; KAUM–I . 132237, 412.4 mm SL, KAUM–I . 132243, 405.9 mm SL, Tomari , Naha , Okinawa , Y. Sakurai , 6 Jul. 2019 . JOHANSTON ATOLL : BPBM 3837, 449.5 mm SL, no further locality data, 1923. MARSHALL ISLAND: USNM 140958 , 2 specimens , 99.8–100.7 mm SL, Rigili Island , Eniwetok Atoll , nightlight, L.P. Schultz & F. Cali , 25 May 1946 . PHOENIX ISLANDS : BPBM 25533 , 98.4 mm SL, Hull Island , nightlight, 5 Aug. 1950 . FIJI : BMNH 1930.8 . 5.15, 437.3 mm SL, no further locality data. FRENCH POLYNESIA : MNHN 1980-0088 , 440.0 mm SL, Rangiroa Island , Tuamotu Islands ; BPBM 6384 , 2 specimens , 383.0–436.0 mm SL, Tahiti Island , Society Islands , J.E. Randall , 3 Sep. 1967 , fish market. PITCAIRN ISLANDS : BPBM 16600 , 587.0 mm SL, north side of Oeno Atoll , 13 m , spear, J.E. Randall , 18 Dec. 1970 ; BPBM 36427 (body in two pieces) , Henderson Island , T . Brooke , 7 Dec. 1991 . EASTER ISLAND: BPBM 39335 (head and tail only) , Motu Nui , 30 m , spear, M. Garcia , 7 Feb. 1986 . Diagnosis. A species of Sphyraena ( Fig. 7 ) characterized by the following combination of characters: one gill raker on first gill arch; pelvic-fin insertion located slightly before vertical through first dorsal-fin origin; pored lateral-line scales 134–141 (modally 137), total lateral-line scales 139–148 (146); scales above and below lateral line 15–17 (15) and 14–16 (15), respectively; snout comparatively short, its length 13.6–15.8 (mean 14.4) % SL; upper jaw short, its posterior tip not reaching to below anterior nostril ( Fig. 2C ), its length 10.0–12.2 (10.8) % SL; eye small, orbit diameter and depth 4.3–7.0 (4.9) % and 3.9–5.6 (4.5) % SL, respectively; anal-fin base shortish, its length 6.9–8.1 (7.5) % SL; last dorsal- and anal-fin ray lengths 4.0–5.6 (4.7) % and 3.6–5.6 (4.6) % SL, respectively; anus not close to anal-fin origin, anterior and posterior margins of anus to anal-fin origin 7.5–11.9 (9.9) % and 5.2–8.3 (6.8) % HL, respectively; head sensory canal pores on suborbital area simple or poorly branched, their lowermost parts not close to lacrimal bone margin, large smooth area lacking canal pores present on mid-margin of lacrimal bone ( Figs. 2C , 6 ); two yellow stripes on lateral body surface (remaining as black stripes in preserved specimens); and caudal fin gray. Description. Counts and proportional measurements as percentages of SL are given in Table 1 . Body elongate, cylindrical, moderately compressed anteriorly, progressively more compressed posteriorly. Dorsal profile rising slightly from snout tip to first dorsal-fin origin, horizontal to second dorsal-fin origin, thereafter gently lowering to caudal-fin base. Ventral profile of head and body lowering from lower-jaw tip to pelvic-fin insertion, almost parallel to body axis from pelvic-fin insertion to anal-fin origin, thereafter rising to lowermost point of caudal-fin base. Second dorsal-fin origin just above anal-fin origin. Head large, conical. Eye and pupil round, lateral on head; orbit elliptical. Nostrils close to each other, located anterior to orbit. Anterior nostril rounded, located slightly below uppermost point of orbit. Posterior nostril semicircular, short flap on anterior edge. Lower jaw longer than upper jaw. Upper jaw short, posterior tip not reaching to below anterior nostril (reaching to anterior nostril in specimens less than ca. 200 mm SL). Upper jaw with a single row of numerous small teeth; 2 pairs of large, recurved, caniniform teeth on symphysis of premaxillary, anterior pair shorter than posterior pair. Lower jaw with a single row of spaced teeth, several anterior teeth smaller; a single large, slightly recurved, caniniform tooth on symphysis of dentary; all jaw teeth ankylotic. Palatine with a single row of irregularly sized caniniform teeth. Vomer toothless. Posterior opercular margin rounded, located anterior to uppermost point of pectoral-fin insertion. Branchial membrane on each side joined distally. Pectoral-fin insertion slightly below lateral-median line of body; posterior tip pointed, failing to reach vertical line through pelvic-fin insertion; uppermost ray short, rudimentary. First spine of first dorsal fin longest (second spine in some specimens), last spine shortest and joined to dorsum by fin membrane. Second dorsal fin somewhat falcate, anterior margin rising from fin origin to posterior tip of second fin ray, distal margin concave; first ray unbranched, other rays branched; last ray elongated. Posterior tip of depressed pelvic fin extending to vertical through last spine origin of first dorsal fin; all pelvic-fin rays branched, first ray longest. Anus oval, long longitudinally, positioned anterior to anal-fin origin. Anal fin somewhat falcate, anterior margin lowering from fin origin to posterior tip of second fin ray, distal margin concave; first spine of anal fin short, rudimentary; first ray unbranched, other rays branched; last ray elongated. Caudal peduncle compressed. Caudal fin forked, without a pair of small lobes on central margin. Lateral line continuous, slightly downward from uppermost point of opercle to point just above anal-fin base end, thereafter parallel to body axis. Body, including ventral surface from isthmus to anal-fin origin and pectoral-fin base, covered with cycloid scales. Anterior half of head without scales, except for opercular area covered with small cycloid scales and mandible with row of 4–6 embedded scales. Pelvic and anal fins and axillary region of pectoral fin lacking scales. Upper and lower gill arches with rough platelets; single gill raker long, slender and rough. Opercular gills present. Swimbladder present. FIGURE 7. Fresh (A–E) and preserved (F) specimens of Sphyraena stellata n. sp. (A, F) KAUM–I. 122319, holotype, 430.7 mm SL, Okinoerabu-jima island, Amami Islands, Japan; (B–D) KAUM–I. 120245, paratype, 275.4 mm SL, Uchinoura Bay, Kagoshima Pref., Japan; and (E) KAUM–I. 55091, paratype, 123.0 mm SL, Yoron-jima island, Amami Islands, Japan. (A, B, E, F), lateral views; (C); dorsal view; and (D) ventral view. FIGURE 8. Underwater photograph of a school of Sphyraena stellata n. sp. (Okinoerabu-jima island, Amami Islands, Japan: Photo by K. Uehara). Specimens not collected. Coloration in life and fresh ( Figs. 7 A–E, 8): Coloration of holotype (KAUM–I. 122319 ; Fig. 4A ) presented first, followed by paratypes (KAUM specimens) (if different). Head and body dorsally yellowish-greenish-silver (blackish), ventrally whitish-silver. Two narrow yellow stripes on lateral body surface; upper line from upper margin of opercle to center of caudal-fin base, on lateral-line scales; lower line from uppermost point of pectoral-fin insertion to lower caudal-fin base. Jaws and lips whitish-silver. Pupil black. Pectoral fin darkish transparent, a small black blotch on uppermost point of pectoral-fin base. Spines of first dorsal fin whitish-gray. Spines and rays of second dorsal fin darkish gray; membrane darkish transparent. Spines and rays of pelvic fin white, membrane whitish translucent (rarely with a small black blotch on pelvic-fin insertion). Anal-fin spines whitish; membrane whitish translucent. Caudal fin gray. Coloration in alcohol ( Fig. 7F ): Body dark brown dorsally, elsewhere pale brown. Two blackish lateral stripes (somewhat indistinct). A black blotch on uppermost point of pectoral-fin base. (A small indistinct black blotch on pelvic-fin insertion in some specimens). All spines and rays milky white. FIGURE 9. Cyanine Blue stained (A–C) and living individual (D) heads of Sphyraena stellata n. sp. showing sensory canal pores. (A) entire head of KAUM–I. 122320, 411.0 mm SL; (B and C) suborbital regions of KAUM–I. 14726, 334.0 mm SL and FRLM 42172, 216.5 mm SL, respectively; and (D) underwater photograph taken from Okinoerabu-jima island, Amami Islands, Japan by K. Uehara (specimen not collected). Areas indicated by broken red lines indicate absence of sensory canals. Distribution. The new species is widely distributed in the Indo-Pacific Ocean ( Fig. 4 ), examined voucher specimens having been collected from the Maldives , southern Japan , including the Ryukyu Islands, and Fiji and oceanic islands in the Pacific Ocean ( Johnston Atoll , Marshall Island, Phoenix Islands , French Polynesia , Pitcairn Islands , and Easter Island). The species has also been recorded from Madagascar , northeastern Australia , and the Hawaiian Islands ( Smith 1956 ; Thinker 1978 ; Hoover 1993 ; Randall et al. 1997a ; Randall 2007 ). Etymology. The specific name stellata is derived from Latin meaning starry, referring to the two yellow lateral stripes on the body. Comparisons. The species group within Sphyraena characterized by a single gill raker on the first gill arch, currently includes the following six valid species: Sphyraena acutipinnis Day, 1876 , S. africana Gilchrist & Thompson, 1909 , S. helleri , S. novaehollandiae , S. stellata and S. waitii Ogilby, 1908 ( Senou 2001 , 2013 ; Fricke et al. 2019 ; this study). The species treated here ( S. helleri , S. novaehollandiae , and S. stellata ) are clearly separated from the other species in having the pelvic-fin insertion located slightly before a vertical through the first dorsal-fin origin [vs. located behind such a vertical or (rarely) just below the first dorsal-fin origin], and the posterior tip of the upper jaw not reaching to below the posterior nostril (vs. reaching beyond the posterior nostril). Scale counts are also useful for identification of species of Sphyraena ( Doiuchi & Nakabo 2005 ; Morishita et al. 2020a , b). Compared with S. helleri and S. novaehollandiae , S. stellata exhibits the highest numbers of pored and total lateral-line scales [ viz., 134–141 (modally 137) and 139–148 (146), respectively in S. stellata vs. 121–127 and 132–134, respectively in S. helleri and 120–138 and 127–146, respectively in S. novaehollandiae : Tables 1 , 2 ]. Scale count above the lateral line are also an important diagnostic character for the aforementioned species [ viz., scales above lateral-line 14–16 (15) in S. stellata vs. 12–13 in S. helleri , and 9–11 (10) in S. novaehollandiae : Tables 1 , 2 ]. FIGURE 10. Relationships of (A) snout length, (B and C) orbit diameter and depth, respectively, and (D) upper-jaw length (as % of SL) to SL (mm) in Sphyraena helleri (yellow stars), S. novaehollandiae (blue squares), and S. stellata n. sp. (red circles). Arrowheads indicate holotypes. The diagnostic importance of head sensory canal pores is well established in many fish families ( e.g., Gobiidae and Gobiescoidae) ( Akihito et al. 2013 ; Fujiwara et al. 2018 ) and we found it helpful for distinguishing between species of Sphyraena . Among S. helleri , S. novaehollandiae , and S. stellata , only S. novaehollandiae has finely branched canals on the suborbital region, whereas the other species have simple canals (sometimes weakly branched) ( Figs. 2 , 3 , 6 , 9 ). Although the lowermost parts of such canals on the suborbital region are very close to the lacrimal bone margin in S. helleri and S. novaehollandiae , those of S. stellata showed greater separation from the margin ( Figs. 2 , 3 , 6 , 9 ). Additionally, S. stellata is unique in having a large smooth area lacking canal pores on the mid-margin of the lacrimal bone ( Figs. 2C , 9 ). FIGURE 11. Ventral views of anus and anal-fin origin of (A) Sphyraena helleri , BPBM 8787, 545.8 mm SL, Oahu Island, Hawaiian Islands; (B) S. novaehollandiae , NSMT-P 112435, 598.0 mm SL, south coast of Australia; and (C) S. stellata n. sp. , BPBM 16600, 587.0 mm SL, Pitcairn Islands. Red, blue, and black arrows indicate anterior and posterior margins of anus, and anal-fin origin, respectively. D, E and F show relationships of anal-fin base length (as % of SL), and AMA and PMA to anal-fin origin, respectively (as % of HL) to SL (mm), in S. helleri (yellow stars), S. novaehollandiae (blue squares), and S. stellata n. sp. (red circles). Arrowheads indicate holotypes. Sphyraena helleri differs from S. novaehollandiae and S. stellata by having a larger eye, and longer snout and upper jaw in similarly sized individuals [ viz., orbit diameter and depth 4.8–5.9% and 4.8–5.6% SL, respectively, in S. helleri (vs. 3.4–5.4% and 3.3–4.9% SL, respectively, in S. novaehollandiae and S. stellata > 200 mm SL: Fig. 10B, C ); snout length 15.0–16.4% SL (vs. 12.2–14.9% SL: Fig. 10A ), upper-jaw length 12.8–12.9% SL (vs. 9.2–11.3% SL: Fig. 10D ), and posterior tip of upper jaw reaching to or beyond anterior nostril (vs. not reaching to below anterior nostril: Fig. 2 )]. FIGURE 12. Relationships of (A) length of last ray of second dorsal fin and (B) length of last ray of anal fin (as % of SL) to SL (mm) in Sphyraena helleri (yellow stars), S. novaehollandiae (blue squares), and S. stellata n. sp. (red circles).Arrowheads indicate holotype. TABLE 1. Counts and measurements of examined specimens of S. helleri , S. novaehollandiae , and S. stellata n. sp.

Sphyraena helleri		Sphyraena novaehollandiae		Sphyraena stellata n. sp.
Holotype	Non-types	Holotype	Non-types	Holotype	Paratypes
USNM 49692	n = 3	BMNH 1847.6.17.37	n = 14	KAUM–I. 122319	n = 40
Standard length (SL, mm)	315.4	243.3–545.8	340.2	270.8–598.0	437.0	98.4–587.0
Counts	Modes	Modes
Dorsal-fin rays	V–I, 9	V–I, 9	V–I, 9	V–I, 9	V–I, 9	V–I, 9	V–I, 9	V–I, 9
Anal-fin rays	II, 8	II, 8–9	II, 8	II, 8	II, 8	II, 8	II, 8	II, 8
Pectoral-fin rays (left)	ii, 12	ii, 12	ii, 12	ii, 10–12	ii, 11	ii, 12	ii, 11–12	ii, 12
Pectoral-fin rays (right)	–	ii, 12	–	ii, 11–12	ii, 11	ii, 11–12	ii, 12
Pelvic-fin rays	I, 5	I, 5	I, 5	I, 5	I, 5	I, 5	I, 5	I, 5
Gill rakers	1	1	1	1	1	1	1	1
Pored lateral-line scales	121	126–127	124	120–138	129	140	134–141	137
Total lateral-line scales	132	133–134	132	129–146	132	144	139–148	146
Scales above lateral line	13	12–13	broken	9–11	10	16	15–17	15
Scales below lateral line	14	14–15	broken	11–13	12	15	14–16	15
Measurements (%SL)	Means	Means
Body depth at pelvic-fin origin	broken	10.2–13.6	11.7	8.1–11.9	10.0	12.3	8.5–14.7	12.0
Body depth at 2nd dorsal-fin origin	broken	9.9–11.0	–	8.8–11.4	10.2	11.3	8.3–11.8	10.7
Body width	broken	7.8–9.9	–	6.3–8.2	6.9	8.7	7.4–9.5	8.4
Head length	31.1	31.1–33.8	27.6	24.3–29.8	28.1	31.3	29.1–35.9	31.4
Snout length	15.0	15.2–16.4	12.7	12.2–14.0	13.2	14.3	13.6–15.8	14.4
Orbit diameter	5.9	4.8–5.8	4.5	3.4–4.5	4.0	4.9	4.3–7.0	4.9
Orbit depth	–	4.8–5.6	–	3.3–4.2	3.7	4.6	3.9–5.6	4.5
Postorbital length	10.5	10.8–11.5	10.5	10.1–11.8	11.0	12.4	11.0–13.6	12.1
Interorbital width	4.3	4.5–6.3	–	3.7–4.9	4.2	5.9	4.4–6.0	5.4
Upper-jaw length	11.9	12.8–12.9	10.0	9.2–10.7	10.0	10.5	10.2–12.2	10.8
Maxillary depth	2.5	2.6–2.8	2.0	1.9–2.4	2.1	2.0	1.9–2.5	2.2
Pre-dorsal-fin length	46.3	46.0–46.8	44.8	43.7–47.1	45.4	46.5	43.9–47.3	45.6
Pre-2nd dorsal-fin length	74.8	73.0–75.4	73.6	71.5–74.9	73.2	74.8	72.4–76.0	73.9
Pre-anal-fin length	76.4	74.5–75.7	74.9	71.7–75.7	73.9	75.1	71.9–77.9	74.6
Pre-pelvic-fin length	46.1	46.3–46.9	42.5	41.9–46.7	44.0	45.8	43.5–47.2	45.0

......continued on the next page TABLE 1. (Continued)

Sphyraena helleri		Sphyraena novaehollandiae		Sphyraena stellata n. sp.
Holotype	Non-types	Holotype	Non-types	Holotype	Paratypes
USNM 49692	n = 3	BMNH 1847.6.17.37	n = 14	KAUM–I. 122319	n = 40
Pre-pectoral-fin length	31.8	31.4–33.6	28.9	27.3–30.3	28.9	31.3	29.6–35.0	31.4
D1-D2	28.6	26.7–28.8	28.4	25.8–30.2	28.0	28.3	25.8–29.8	28.3
P2-A	30.2	28.8–29.4	32.4	28.7–31.7	30.4	29.8	26.1–32.2	29.8
D1-P2	–	10.3–13.8	–	8.1–12.5	10.2	12.7	9.4–14.8	12.2
D2-A	–	10.0–11.0	–	8.8–11.6	10.2	11.3	8.8–12.0	10.8
P1-P2	14.7	13.8–15.4	13.8	14.6–18.6	15.7	15.1	10.5–16.6	14.4
D1-P1	16.8	15.7–16.9	17.9	16.6–20.5	18.0	17.5	13.8–18.6	16.6
Pectoral-fin length	broken	11.0	broken	9.5–11.4	10.5	10.4	7.9–10.9	9.7
Pelvic-fin length	broken	8.7–9.0	broken	7.7–8.8	8.1	8.8	6.5–9.7	8.4
1st dorsal-fin base length	6.3	6.0–6.5	5.6	5.5–6.5	6.0	5.5	5.0–7.1	5.9
2nd dorsal-fin base length	7.7	7.3–7.8	7.6	7.4–8.2	7.9	7.6	6.7–8.4	7.7
Length of 1st dorsal-fin spine	broken	7.3	7.0	6.8–8.4	7.4	8.2	5.4–9.2	8.2
Length of 2nd dorsal-fin spine	broken	8.7	6.9	6.7–8.2	7.5	8.1	5.9–9.3	8.2
Length of 1st ray of 2nd dorsal fin	broken	8.6	4.5	7.6–9.3	8.2	7.8	6.4–8.4	7.7
Length of 2nd ray of 2nd dorsal fin	broken	7.7	broken	7.5–8.8	8.3	7.7	6.6–8.3	7.5
Length of penultimate ray of 2nd dorsal fin	broken	3.2–3.5	broken	3.4–4.1	3.8	3.0	2.3–4.1	3.5
Length of last ray of 2nd dorsal fin	broken	4.2–4.3	broken	3.6–4.5	4.0	4.3	4.0–5.6	4.7
Anal-fin base length	7.5	6.8–7.4	8.2	7.6–9.2	8.5	7.4	6.9–8.1	7.5
Length of 1st ray of anal fin	broken	8.3	broken	6.9–8.9	7.9	7.8	6.3–8.7	7.5
Length of 2nd ray of anal fin	broken	7.8–8.0	broken	6.9–8.2	7.6	7.5	6.4–8.1	7.4
Length of penultimate ray of anal fin	broken	3.0–3.6	broken	3.3–4.4	3.8	3.1	3.0–4.3	3.6
Length of last ray of anal fin	broken	3.6–4.7	broken	3.3–4.3	3.9	4.5	3.6–5.6	4.6
Caudal-peduncle length	17.7	16.9–17.5	17.1	17.1–20.7	18.8	19.1	17.4–20.8	18.8
Caudal-peduncle depth	5.3	4.7–5.6	5.4	5.0–6.3	5.4	5.1	4.1–5.7	5.1
Upper caudal-fin length	broken	19.0	broken	15.9–17.1	16.8	17.9	15.0–19.3	17.0
Lower caudal-fin length	broken	19.1	broken	14.5–17.2	15.9	17.8	14.4–18.5	16.9
AMA to anal-fin origin (%HL)	broken	7.1–8.2	broken	5.8–8.0	6.4	8.6	7.5–11.9	9.9
PMA to anal-fin origin (%HL)	6.5	4.1–4.9	broken	3.1–5.1	4.0	5.8	5.2–8.3	6.8

TABLE 2. Frequency distribution of pored lateral-line scales, total lateral-line scales, and scales above and below lateral line in Sphyraena helleri , S. novaehollandiae , and S. stellata n. sp.

Pored lateral-line scales
120	121	122	123	124	125	126	127	128 129	130	131	132	133	134	135	136 137		138	139	140 141
S. helleri S. novaehollandiae S. stellata n. sp.	2	1H	1	2H	1	1 1	2 1	3	1	1	1 1	8	5	8	1 3	5	4H	5
Total lateral-line scales
127	128	129	130	131	132	133	134	135 136	137	138	139	140	141	142	143 144		145	146	147 148
S. helleri S. novaehollandiae S. stellata n. sp.	1	2	2	1H 2H	2 1	1 1	2	1	1	1	2	1	7	1 7	4H	6	1 8	2	1
Scales above lateral line				Scales below lateral line
9	10		11	12	13	14	15	16	17		11	12	13	14	15	16
S. helleri S. novaehollandiae S. stellata n. sp.	1	9	4	2H	2	24	12H	1	3	7	4 2	3H 14	1 22H 1

H indicates holotype Sphyraena novaehollandiae can be distinguished from S. helleri and S. stellata in having two additional unique characters: longer anal-fin base, its length 7.6–9.2 (mean 8.5) % SL [vs. 6.8–7.5 % SL in S. helleri and 6.9–8.1 (7.5) % SL in S. stellata : Fig. 11 A–D) and yellowish-gray caudal fin (vs. gray in S. helleri and S. stellata ). In addition, S. novaehollandiae has slightly shorter posteriormost dorsal and anal fin rays than S. stellata , with lengths 3.6–4.5 (4.0) % and 3.3–4.3 (3.9) % SL, respectively vs. 4.0–5.6 (4.7) % and 3.6–5.6 (4.7) % SL, respectively, in the latter ( Fig. 12 ). Additionally, S. stellata has two other distinguishing characters: longer distance from anterior and posterior margins of anus to anal-fin origin 7.5–11.9 (9.9) % and 5.2–8.3 (6.8) % HL, respectively [vs. 6.5–8.2 % and 4.1–4.9 % HL, respectively, in S. helleri and 5.8–8.0 (6.9) % and 3.1–5.1 (4.0) % HL, respectively, in S. novaehollandiae : Fig. 11 A–C, E–F], and two yellow stripes on the lateral body surface (remaining as black stripes in preserved specimens) (vs. lateral body surface without stripes or bands in S. helleri and S. novaehollandiae : Figs. 1 , 5 , 7 ). Comparative material examined. Sphyraena acutipinnis : ZSI-F 1599, holotype , 452.0 mm SL, Sind , Pakistan . Sphyraena africana : SAM 10149, holotype , 187.6 mm SL, Durban , KwaZulu-Natal , South Africa . Sphyraena waitii : CSIRO H 7281-08 , 262.1 mm SL, Queensland , Australia .