Squalius namak, Khaefi, Roozbehan, Esmaeili, Hamid Reza, Sayyadzadeh, Golnaz, Geiger, Matthias F. & Freyhof, Jörg, 2016

Khaefi, Roozbehan, Esmaeili, Hamid Reza, Sayyadzadeh, Golnaz, Geiger, Matthias F. & Freyhof, Jörg, 2016, Squalius namak, a new chub from Lake Namak basin in Iran (Teleostei: Cyprinidae), Zootaxa 4169 (1), pp. 145-159 : 148-158

Squalius namak , new species

( Figs. 2–4 )

Holotype. ZM-CBSU G 121, 136 mm SL; Iran: Markazi prov.: spring Bolagh (Cheshmeh Bolagh) at Shazand, east of Anjirak , 34°00'38"N 49°50'51"E; H.R. Esmaeili, A. Gholamifard & A. Teimori, 22 June 2009.

Paratypes. All from Iran: Markazi prov.: ZM-CBSU G111, 10, 170– 202 mm SL; ZM-CBSU E769, 28, 57– 188 mm SL; ZM-CBSU G433, 5, 144- 168 mm SL; FSJF 3521, 5, 105– 146 mm SL; same data as holotype.—ZM- CBSU G1001, 2, 155– 189 mm SL; spring Bolagh (Cheshmeh Bolagh) at Shazand, east of Anjirak, 34 ° 0 0 ' 38 " N 49 ° 50 ' 51 " E.—ZM-CBSU D871, 22, 37–110 mm SL; FSJF 3522, 5, 57–122 mm SL; Poledoab River at Shazand, 34 ° 0 2 ' 36 " N 49 ° 21 ' 0 9 " E.

Material for molecular genetic analysis. All from Iran: ZM-CBSU M271; M272; M273, Markazi prov.: spring Bolagh (Cheshmeh Bolagh) at Shazand , east of Anjirak , 34°00'38"N 49°50'51"E (GenBank accession numbers: KX516770 , KX516771 , KX516772 ) . —ZM-CBSU M361; M362, Markazi prov.: Poledoab River at Shazand , 34°02'36"N 49°21'09"E (GenBank accession numbers: KX516773 , KX516774 ) . —ZM-CBSU M132; M134, Hamedan prov.: Saleh Abad River between Hamedan and Sanandaj , 34°35'23.3"N 48°19'48.1"E (GenBank accession numbers: KX516768 , KX516769 ) .

Diagnosis. Squalius namak is distinguished from the Squalius species found in the adjacent Caspian Sea and Persian Gulf basins by having an unique pigmentation pattern of the flank scales (a bold, grey or brown, roundish or crescent shaped blotch at the tip of each flank scale vs. no bold blotch at the tip of each flank scale: Figs. 3 , 5 ). Squalius namak is further distinguished from the species of Squalius in the Caspian Sea and Persian Gulf basins by a combination of characters. It differs from S. berak and S. turcicus by having a wide and thick symphysial knob on the lower jaw ( Fig. 9 ) (vs. no knob or knob thin and small, Figs. 10–11 ). It is distinguished from S. lepidus from the Euphrates and Tigris drainages by having fewer scales along the lateral line (39–43+1–3 vs. 45–50+1–3), a convex (vs. straight) posterior anal-fin margin, and orange caudal-, anal- and pelvic-fin rays ( v. hyaline or with a yellow hue). Further, the head of S. namak is blunt (vs. pointed) and deeper (head depth 50–63% HL vs. 44–53) and the caudal peduncle shorter than in S. lepidus (15–20% SL vs. 19–23). Squalius namak is distinguished from S. berak from the Qweik, Euphrates and Tigris drainages by having poorly-pigmented scale pockets of the flank scales above the lateral line (vs. scale pockets with a dark-grey, crescent-shaped mark at least on midlateral flank) and orange caudal-, anal- and pelvic-fin rays (vs. hyaline or with a yellow hue). Squalius namak is distinguished from S. turcicus from the Caspian Sea basin by the upper lip clearly projecting beyond the lower lip ( Fig. 6 ) (vs. tip of upper lip level with that of lower lip, Fig. 8 ). Squalius namak is also characterized by four fixed, diagnostic nucleotide substitutions in the mtDNA COI barcode region.

Description. See Figures 2–4 for general appearance and Table 1 for morphometric and Table 2 for meristic data. Body elongated, compressed; dorsal and ventral body profile slightly convex; upper head profile concave or straight. Interorbital area concave. Belly between anus and posterior pelvic-fin base compressed. Snout pointed, conical. A shallow hump at nape. Mouth subterminal, upper lip projecting beyond lower lip. Mouth cleft straight, oblique. Snout obtuse, markedly rounded. Head length greater than body depth at dorsal-fin origin. Uppermost point of mouth cleft between level of lower margin of pupil and lower margin of eye. Lower jaw-quadrate junction about at vertical through anterior margin of eye. A very wide, thick symphysial knob on lower jaw ( Fig. 9 ). Upper lip thick, anterior depth 1.6–2.0 times in its lateral length. Eye diameter 1.3–2.5 times in interorbital distance, 15– 32% HL. Caudal-peduncle depth 1.2–1.9 times in its length. Largest recorded individual 189 mm SL.

Holotype Range Mean SD

Standard length (mm) 135.9 32.1–189 91.7

In percent of standard length

Dorsal fin with 3–4 (mode 3) simple and 7½–10½ (mode 8½ or 9½) branched rays, distal margin straight or slightly convex. Dorsal-fin origin behind vertical through pelvic-fin origin. Tip of dorsal fin reaching vertical through anal-fin origin when adpressed. Anal fin with 3–4 (mode 3) simple and 7½–10½ (mode 8½) branched rays; distal margin convex. Posteriormost point of anal fin at tip of 3rd or 4th branched ray. Pectoral fin with 15–18 (usually 16–17), pelvic fin with 9–11 (mode 9) rays. A large axillary pelvic lobe present. Caudal fin forked, with 9+8 branched rays. Lateral line complete, with 39–43 scales on body, 1–3 on caudal-fin base. Scale rows between dorsal-fin origin and lateral line 6½–8½ (mode 7½), scale rows between pelvic-fin origin and lateral line 3½–4½ (mode 3½); 14–18 (mode 16) circumpeduncular scales, 19–24 predorsal scales. Gill rakers on first gill arch 9–12 (mode 10). No external sexual dimorphism observed.

Total lateral-line scales

N 3 9 4 0 4 1 4 2 4 3 4 4 4 5 4 6 4 7 4 8 4 9 5 0 5 1

S. berak * 1 6 6 7 3

S. lepidus 1 5 1 3 5 4 1 1 S . namak 56 6 14 17 14 3 2

S. turcicus 4 4 3 1 1 1 4 1 2 4

S. turcicus * 24 1 6 11 5 1

Scales above lateral line Scales below lateral line

N 6 7 8 9 1 0 N 3 4 5

S. berak * 1 6 1 6 S. berak * 16 4 12

S. lepidus 15 14 1 S. lepidus 15 15

S. namak 56 2 53 1 S. namak 56 52 4 S . turcicus 44 1 28 14 1 S . turcicus 44 42 2 S . turcicus * 2 4 1 2 3 S . turcicus * 2 4 2 2 2 Predorsal scale

N 1 6 1 7 1 8 1 9 2 0 2 1 2 2 2 3

S. berak 3 2 1

S. lepidus 1 5 1 2 3 4 3 S. namak 5 6 1 5 1 5 1 7 1 2 6 S . turcicus 44 1 1 10 17 6 5 4 Branched anal-fin rays Branched dorsal-fin rays

N 6½ 7½ 8½ 9½ N 6½ 7½ 8½ 9½

S. berak * 1 6 1 4 2 S . berak * 1 6 1 6 S. lepidus 15 6 9 S . lepidus 15 2 13 S . namak 56 1 15 36 4 S . namak 56 11 43 2 S. turcicus 44 10 32 2 S. turcicus 44 1 4 39 S. turcicus * 24 19 5 S . turcicus * 24 3 21 Coloration. In life: head and body silvery brown, darker on back. Belly white. A faint black bar from uppermost gill opening to pectoral-fin base. Peritoneum black. Each flank-scale margin with dark brown or black pigments forming a reticulate pattern. Scale pocket poorly pigmented above lateral line, with a bold, brown or black, crescent-shaped, vertically elongated or roundish anterior scale mark on scales below lateral line. Free margins of scales above lateral line pale brown or grey, each with a bold, brown or black, crescent-shaped, vertically elongated or roundish blotch on posterior tip ( Figs. 3 , 5 ). Caudal-, anal- pelvic- and pectoral-fin rays orange, caudal-fin rays with black pigments. Dorsal-fin membranes hyaline with dark-grey rays.

In preservation: Head and body pale brown, darker on back. Dorsal and caudal fins with blackish rays and hyaline membranes; rays whitish and membranes yellowish in pectoral, pelvic and anal fins.

Distribution. Squalius namak is known from the springs Bolagh ( Fig. 12 ) and Eskan in the Poledoab River drainage and the spring Emamzadeh Abdolah in the Qom River drainage, all in the Namak Lake basin. It also occurs in the Hableh River in the Kavir basin.

Etymology. The species is named for the Namak Lake basin. A noun in apposition.

Remarks. In the Euphrates and Tigris drainage, S. lepidus is very widespread and usually inhabits larger rivers and streams in lowland habitats (own observations). In headwater streams and rivers in mountainous habitats, a second Squalius species is found in the Euphrates and Tigris drainage (own observations) usually identified as S. cephalus ( Abdoli 2000) . A syntopic occurrence of both species was detected by JF in the Little Zab River drainage (Tabon River) in Iraqi Kurdistan. Here we identify this chub as S. berak , a species described from the Qweik River in Syria, a small endorheic drainage west to the Euphrates. Squalius berak from the Sünnep, a headwater stream of the Qweik in Turkey, shares the COI barcode sequence with superficially similar, round-headed Squalius found in the Euphrates and Tigris drainage. However, the lineage sorting between S. lepidus and S. berak is not complete and both species seem occasionally to hybridize. This might lead to the false classifications of a few individuals and populations of both species based on their COI barcode sequence as seen in Figure 1.

Naseka & Bogutskaya (2009) identified the Squalius populations of the southern Caspian Sea basin as S. orientalis . Squalius orientalis was described from Abkhazia ( Berg 1949), which is situated between Georgia and Russia, in the easternmost Black Sea basin. Squalius orientalis is treated as a valid species by Stoumboudi et al. (2006), Doadrio & Carmona (2004) and Turan et al. (2009, 2013), without discussing in detail how it is distinguished from S. cephalus . Özuluğ & Freyhof (2011) discussed the case and kept S. orientalis as a synonym of S. cephalus based on the lack of studies of the morphological characters distinguishing S. orientalis from S. cephalus . The present analysis, however, includes COI sequences of S. orientalis from Georgia and Turkey. These fishes are very close to S. turcicus , a species described from the upper Arax River in Turkey by De Filippi (1865). Özuluğ & Freyhof (2011) suggested that S. turcicus might be a valid species occurring in the southern Caspian Sea basin and Turan et al. (2013) supported this view and provided some morphological data distinguishing this species from S. orientalis . Our molecular data suggest that S. orientalis and S. turcicus are very closely related and might represent just one species ( S. orientalis ). Squalius turcicus might be more widespread and Squalius populations from the Lake Urmia basin, as well as those from the Iranian Sefid River and from the Iranian Talar River (flowing to the Caspian Sea at Bahnamir), might belong to this species. More and geographically focused studies are needed to better understand the distribution of S. turcicus in the rivers of the southern Caspian Sea basin.

Comparative materials. Squalius berak : NMW 48915, 6 , syntypes, 64–304 mm SL ; Syria: Aleppo.— ZM- CBSU J1725, J1740, J1741, 3, 133– 137 mm SL ; Iran: West Azerbaijan prov.: Little Zab River at road between Piranshahr and Sardasht, 36°28'36"N 45°19'54"E.—ZM-CBSU G1723, 2, 43-52 mm SL ; Iran: Kordistan prov.: Boeen River at road between Baneh and Boeen, 35°56'35.5"N 45°56'36.5"E.— IUSHM 2010-984, 4 , 112– 143 mm SL; FSJF 2912, 4 , 113– 146 mm SL ; Turkey: Kilis prov.: stream Sünnep 10 km east of Kilis, 36°45'50"N 37°15'14"E.— FSJF 3262, 8 , 83–135 mm SL ; Iraq: Nalparez River , 35°34.24'N 45°51.78'E.— FSJF 3349, 9 , 45– 158 mm SL ; Iraq: Kuna Massi stream in Sevanja , 35°47.35'N 45°24.18'E.

Squalius lepidus : FSJF 2893, 15, 43–190 mm SL; Turkey: Gaziantep prov.: tributary to Merziman stream south of Yavuzeli , 37°16'36"N 37°31'56"E.— FSJF 2996, 6 , 109– 133 mm SL , Turkey: Gaziantep prov.: Merziman stream south of Yavuzeli , 37°17'32"N 37°43'23"E.— IUSHM 2010-991, 2 , 174– 188 mm SL; FSJF 2862, 2 , 183– 213 mm SL ; Turkey: Batman prov.: Tigris River 5 km west of Hasankeyf , 37°43'25''N 41°21'37''E.—ZM-CBSU G794, 3, 145– 125 mm SL ; Iran: Kermanshah prov.: Seymareh River west of Charu Dareh , 33°39'40"N 47°03'29"E.—ZM-CBSU G797A, 1, 127 mm SL ; Iran: Kermanshah prov.: Seymareh River south of Mahizan , 33°40'54"N 47°03'22"E.

Squalius turcicus : IUSHM 2013-1057, 6 , 87–178 mm SL; Turkey: Kars prov.: stream between Sarikamis and Handere, about 4 km southwest of Sarikamis , 40°19'10"N 42°32'41"E.—ZM-CBSU D864, 4, 137– 65 mm SL ; Iran: Ardabil prov.: spring Chay Almas east of Sarein , 38°09'31"N 48°11'37"E.—ZM-CBSU D504, 22, 38–145 mm SL ; Iran: East Azarbaeijan prov.: stream Chay near Vardin , 38°26'49"N 46°59'14"E.

Materials used for molecular analysis. Squalius berak : FSJF DNA- 1383 , Turkey: Kilis prov.: stream Sünnep 10 km east of Kilis, 36°45'50"N 37°15'14"E (GenBank accession numbers: KX516744 , KX516745 , KX516746 ) .—FSJF DNA-1416; FSJF DNA- 2355 , Turkey: Sivas prov.: Kangal stream under railway bridge at Çetinkaya , 39°15'4.644"N 37°37'7.464"E (GenBank accession numbers: KX516734 , KX516735 , KX516736 , KX516737 , KX516741 ) .— FSJF DNA- 1585 , Turkey: Erzurum prov.: Karasu stream at Kandilli , 39°91'24''N 40°85'40''E (GenBank accession numbers: KX516738 , KX516739 , KX516740 ) .—FSJF DNA-2257; FSJF DNA- 2275; FSJF DNA- 2290 ; Iraq: Tabin River west of Zarbi , 35°48'06''N 44°58'47''E (GenBank accession number: KX516752 ) .— FSJF DNA- 2259 , Iraq: stream Zalm at Khurmal , 35°18.38'N 45°58.26'E (GenBank accession number: KX516750 ) .— FSJF DNA- 2389 , Turkey: Erzurum prov.: stream Arkaçayırlar at Paşayurdu close to the road from Ilıca to Aşkal , 39°58'59.952"N 40°59'31.308"E (GenBank accession number: KX516743 ) . —ZM-CBSU M464; M465, Iran: West Azerbaeijan prov.: Badin Abad River at Piranshahr , 36°28'36''N 45°19'54''E (GenBank accession numbers: KX516747 , KX516748 ) .

Squalius lepidus : FSJF DNA- 1445 , Turkey: Gaziantep prov.: tributary to Merziman stream south of Yavuzeli , 37°16'36"N 37°31'56"E (GenBank accession numbers: KX516753 , KX516754 , KX516755 ). —FSJF DNA-1441; FSJF DNA-1442; FSJF DNA-1447; FSJF DNA- 1462 ; Turkey: Batman prov.: Tigris River 5 km west of Hasankeyf , 37°43'25''N 41°21'37''E (GenBank accession numbers: KX516764 , KX516765 , KX516766 , KX516767 ). —FSJF DNA-2248; FSJF DNA-2271; FSJF DNA- 2287 ; Iraq: Tabin River west of Zarbi , 35°48'06''N 44°58'47''E (GenBank accession numbers: KX516761 , KX516762 , KX516763 ).—ZM-CBSU M751; M752; M755; M756 , Iran: Ilam prov.: Seymareh River at Seymareh bridge, at Darehshar , 33°39'19''N 47°02'15''E (GenBank accession numbers: KX516756 , KX516757 , KX516758 , KX516759 ).—ZM-CBSU M1116 , Iran: Kohkiluye and Boyerahmad prov.: Bashar River at Talegah , 30°38'55''N 51°37'06''E (GenBank accession number: KX516760 ).

Squalius orientalis : FSJF DNA- 260 , Georgia: River Galidzga about 5 km upstream from coast, 42°42'28'' N 41°29'06''E (GenBank accession numbers: KX516776 , KX516777 ) .— FSJF DNA- 1589 , Turkey: Çoruh River at Borçka , 41°36'54''N 41°67'77''E (GenBank accession number: KX516775 ) .— FSJF DNA- 1780 , Georgia: Chakhota River at Natanebi , 42º55'59''N 41º50'10''E (GenBank accession number: KX516778 ) .— ZFMK:ICH:TIS:NB203; Georgia: Chakhvata stream ca. 30 km west of Guria, 41°55'59''N 41°50'10''E (GenBank accession numbers: KX516779 , KX516780 ) .

Squalius turcicus : FSJF DNA- 2344 ; Turkey: Kars prov.: Güllü stream at Bölükbaşı village 3 km west of Selim , 40°28'15.924"N 42°44'11.76"E (GenBank accession numbers: KX516786 , KX516787 ) .—FSJF DNA-2357; FSJF DNA 2372 ; Turkey: Ardahan prov.: Lake Çıldır at Akçakale village , 41°4'53.292"N 43°17'42.432"E (GenBank accession numbers: KX516788 , KX516789 ) .— FSJF DNA- 2393 , Turkey: Kars prov.: stream crossing the road between Sarıkamış and Handere, 4 km southwest of Sarıkamış , 40°19'9.588"N 42°32'41.46"E (GenBank accession number: KX516790 ) . —ZM-CBSU M461, Iran: Ardabil prov.: Aras River at Pasrabad Moghan , 38°35'05"N 47°48'09"E (GenBank accession number: KX516783 ) . —ZM-CBSU M152; M153, Iran: Zanjan prov.: Qezel Owzan River south west of Balobin village , 36°19'26"N 48°09'13"E (GenBank accession numbers: KX516798 , KX516799 ) . —ZM-CBSU M359; M360, Iran: Ardabil prov.: Gharehsu River at Kangavar , 38°35'05"N 47°48'07"E (GenBank accession numbers: KX516781 , KX516782 ) . —ZM-CBSU M462, M463, Iran: Ardabil prov.: Balakhluchay River at Nir , 38°02'02"N 47°58'30"E (GenBank accession numbers: KX516784 , KX516785 ) .— ZM- CBSU S928F, Iran: Kurdistan prov.: Serudan River at road between Marivan and Saghez, 35°44'34''N 46°26'40''E (GenBank accession number: KX516803 ) . —ZM-CBSU M154, Iran: Kurdistan prov.: Divan Darreh at Nesar Olya , 35°52'22''N 47°04'17''E (GenBank accession number: KX516800 ) . —ZM-CBSU M285, Iran: Mazandaran prov.: Azar River at Shirgah , 36°18'15''N 52°53'07''E (GenBank accession number: KX516802 ) .— ZFMK:ICH:TIS:NB80; Russia: stream Shuraozen about 2 km west of Korkmaskala , 42°59'55''N 47°12'06''E (GenBank accession number: KX516801 ) .—ZFMK:ICH:TIS:66542; ZFMK:ICH:TIS:66543; Turkey: Digor stream at Digor , 40°22'32.48"N 43°24'57.57"E (GenBank accession numbers: KX516791 , KX516792 ) .— ZFMK:ICH:TIS:66572; ZFMK:ICH:TIS:66573; Turkey: stream Selim in Selim , 40°28'9.68"N 42°46'57.64"E (GenBank accession numbers: KX516793 , KX516794 ) .—ZFMK:ICH:TIS:66601; ZFMK:ICH:TIS:66602; IUSHM 2012-1052 ; Turkey: Çıldır Lake , 41°3'3.44"N 43°13'46.64"E (GenBank accession numbers: KU729274 , KX516795 , KX516796 ) .

Petroleuciscus borysthenicus : ZFMK:ICH:TIS:NB187; Georgia: Gubistskali stream about 8 km east of Samtredia, 42°10'10''N 42°25'53''E (GenBank accession numbers: KX516732 , KX516733 ).

Figures

FIGURE 2. Squalius namak, holotype, ZM-CBSU G 121, 136 mm SL; Iran: spring Bolagh.

FIGURE 3. Squalius namak, paratypes, ZM-CBSU G 119, 113 mm SL, ZM-CBSU E 780, 96 mm SL; Iran: spring Bolagh.

FIGURE 4. Squalius namak, ZM-CBSU G 1001, paratype, 189 mm SL; Iran: spring Bolagh.

FIGURE 5. Pattern of flank-scale pigmentation below dorsal fin: a, S. berak, ZM-CBSU J 1725, 143 mm SL; Iran: Little Zab River between Piranshahr and Sardasht; b, S. lepidus ZM-CBSU G 794, 145 mm SL; Iran: Seymareh River west of Charu Dareh; c, S. namak ZM-CBSU G 437, 145 mm SL; Iran: Bolagh Spring at Shazand; d, S. turcicus IUSHM 1059, 148 mm SL; Turkey: stream between Sarikamis and Handere, about 4 km south of Sarikmis.

FIGURE 6. Shape of head in Squalius namak. a, ZM-CBSU E 780, 98 mm SL; b, ZM-CBSU E 783, 99 mm SL; c, ZM-CBSU G 119, 115 mm SL; d, ZM-CBSU G 120, 129 mm SL; e, ZM-CBSU G 121, 138 mm SL; f, ZM-CBSU G 437, 145 mm SL; Iran: spring Bolagh at Shazand.

FIGURE 8. Shape of head in Squalius turcicus. a, ZM-CBSU D 522, 71 mm SL; Iran: stream Chay near Vardin; b, IUSHM 1057, 119 mm SL; c, IUSHM 1058, 134 mm SL; d, IUSHM 1059, 148 mm SL; e, IUSHM 1060, 152 mm SL; f, IUSHM 1061, 153 mm SL; Turkey: stream between Sarikamis and Handere, about 4 km southwest of Sarikamis.

FIGURE 9. Shape of knob in Squalius namak. a, ZM-CBSU E 780, 98 mm SL; b, ZM-CBSU E 783, 99 mm SL; c, ZM-CBSU G 119, 115 mm SL; d, ZM-CBSU G 120, 129 mm SL; e, ZM-CBSU G 121, 138 mm SL; f, ZM-CBSU G 437, 145 mm SL; Iran: spring Bolagh at Shazand.

FIGURE 10. Shape of knob in Squalius berak. a, ZM-CBSU G 1723, 43 mm SL; b, ZM-CBSU G 1724, 52 mm SL; Iran: Boein River at road at between Baneh and Boeen; c, ZM-CBSU J 1740, 137 mm SL; d, ZM-CBSU J 1725, 143 mm; Iran: Little Zab River between Piranshahr and Sardasht.

FIGURE 11. Shape of knob in Squalius turcicus. a, ZM-CBSU D 522, 70 mm SL; Iran: stream Chay near Vardin; b, IUSHM 1057, 119 mm SL; c, IUSHM 1058, 134 mm SL; d, IUSHM 1059, 148 mm SL; e, IUSHM 1060, 152 mm; f, IUSHM 1061, 153 mm SL; Turkey: stream between Sarikamis and Handere, about 4 km southwest of Sarikamis.

FIGURE 12. Type locality of S. namak; Iran: spring Bolagh.

Tables

TABLE 1. Morphometric data of Squalius namak (holotype ZM-CBSU G 121, paratypes ZM-CBSU E 100, n = 10; ZM- CBSU E 769, n = 28; ZM-CBSU G 1001, n = 2; ZM-CBSU D 871, n = 22).

Head length 28.4	24.6–32.5	28.4	1.5
Snout length 9.3	6.8–9.7	8.2	0.5
Postorbital distance 15.3	12.4–17.7	15.3	0.9
Interorbital distance 10.3	8.8–12.3	10.3	0.7
Predorsal length 56.4	50.6–59.3	56.0	1.5
Postdorsal length 52.9	45.4–61.9	54.2	2.8
Dorsal-fin base length 18.7	15.7–24.7	20.0	2.0
Dorsal-fin depth 11.7	9.1–15.5	11.1	0.8
Anal-fin base length 13.8	13.1–21.2	16.8	1.7
Anal-fin depth 10.3	7.6–13.5	10.3	0.8
Preanal length 74.5	65.7–75.8	72.9	1.5
Pectoral-fin length 16.9	13.3–20.0	17.5	1.2
Pelvic-fin length 14.8	12.9–17.8	15.4	0.9
Depth of caudal peduncle 12.2	9.2–12.8	11.7	0.6
Body depth at dorsal-fin origin 24.8	22.5–28.4	25.4	1.3
Distance between pectoral and anal-fin origins 46.3	39.4–48.5	44.7	2.2
Distance between pectoral and pelvic-fin origins 25.3	20.4–27.0	23.6	1.5
Distance between pelvic and anal-fin origins 21.6	17.4–24.2	21.3	1.4
Length of caudal fin 20.6	16.8–29.4	22.4	2.7
Length of caudal peduncle 15.1	14.0–20.5	17.3	1.3
In percent of head length
Head depth 56	50–63	56	3.0
Snout length 33	25–33	29	1.9
Postorbital distance 54	44–61	54	2.4
Interorbital width 36	32–44	36	2.1
Eye diameter 20	15–32	22	3.2
Length of caudal fin 73	48–73	61	5.6
Mouth width 31	18–35	25	3.5