Redescription of Orthopristis ruber and Orthopristis scapularis (Haemulidae: Perciformes), with a hybridization zone off the Atlantic coast of South America Author Marceniuk, Alexandre Pires Author Caires, Rodrigo Antunes Author Machado, Leonardo Author Cerqueira, Najila Nolie Catarine Dantas Author Serra, Rayla Roberta M. De S. Author Oliveira, Claudio text Zootaxa 2019 2019-04-01 4576 1 109 126 journal article 27520 10.11646/zootaxa.4576.1.5 f30f953f-f745-452b-8498-a53ce12e04ba 1175-5326 2624718 A5242453-3D24-43A3-8EF9-07F841C6B314 Orthopristis ruber Cuvier 1830 Figures 1 a–f, Tables 3 & 4 Pristipoma rubrum Cuvier [G.] in Cuvier & Valenciennes 1830 : 283 [type locality; Brazil ]. Syntypes : MNHN 0000–4084 (2), 7734 (2), A–0457 (1), A–7821 (1). Orthopristis ruber .— Jordan & Fesler 1893 : 499 [notes on etymology, habitat and distribution].— Miranda Ribeiro 1903 : 171 [fishing of “Annie” trawl; listed].—Miranda Ribeiro 1915: 390 [fauna Brasiliense; description].— Miranda Ribeiro 1918 : 109 [fauna Brasiliense; synonymy].— Bauchot et al . 1983 :42 [ type catalog].— Figueiredo & Menezes 1980 : 34 –35 [fishes of southeastern Brazilian coast; list and short description].— Andreata & Séret 1996 : 586 [fishes at continental shelf limits Vitoria, Trindade and Martim Vaz; listed].—Moura & Menezes in Menezes et al . 2003 :85 [in part; catalog of marine fishes of Brazil ; listed]. Doubtful references for this species Pristipoma lineatum Cuvier in Cuvier & Valenciennes 1830 : 287 [type locality: Brazil ]. Brazil . Syntypes : MNHN 0000–7740 (1, dry).— Jordan & Fesler 1893 : 499 [synonymy under O. ruber ] Material examined. ( 42 specimens , 59–217 mm SL). Brazil : MZUSP 68089 , 5 : 101–112 mm SL, Espírito Santo , 18°45' S 39°35' W ; MZUSP 68058 , 3 : 96–122 mm SL, Espírito Santo , 18°45' S 39°35' W ; MZUSP 91637 , 2 : 59– 63 mm SL, Espírito Santo , Guarapari, Morro Beach, 20°39'14" S 40°28'24" W ; MZUSP 3155 , 2 : 112–154 mm SL, Rio de Janeiro, Rio de Janeiro ; MZUSP 3172 ( 1, 128 mm SL), Rio de Janeiro, Rio de Janeiro ; MZUSP 3171 , 1 : 111 mm SL, Rio de Janeiro, Rio de Janeiro ; LBP 22153, 2 : 125–158 mm SL, Guanabara Bay , Rio de Janeiro ; AZUSC 1474 , 3 : 206–217 mm SL, ilha Grande , Rio de Janeiro ; AZUSC 1812 , 3 : 104–184 mm SL, ilha Bela , São Paulo ; LBP 8670 , 2 : 184–214 mm SL, Ubatuba , São Paulo ; AZUSC 4338 , 3 : 131–180 mm SL, Santos , São Paulo ; AZUSC 1158 , 3 : 169–191 mm SL, Itanhaém , São Paulo ; LBP 21672, 1 : 173 mm SL, Cananéia , São Paulo ; LBP 12175, 1 : 214 mm SL, Cananéia , São Paulo ; AZUSC 1252 , 4 : 149–211 mm SL, Gravatá beach, Laguna , Santa Catarina ; MZUSP 68080 , 2 : 108–180 mm SL, Rio Grande do Sul ; MZUSP 68079 , 4 : 133–174 mm SL, 30°63’S, 49°81’W, Rio Grande do Sul . Diagnosis. Orthopristis ruber can be differentiated from its congeners as follows: from Orthopristis chrysoptera , found from New York to Cuba , by having 10 rays in the anal fin (vs. 12–13); from Orthopristis cantharinus , found in the Galapagos Islands, by having 52–57 scales with pores in the lateral line (vs. 65–72, Table 4a ); 11–13 scales above the lateral line (vs. 9–10, Table 4b ), and 10 rays in the anal fin (vs. 12–13); from Orthopristis chalceus , found from the Gulf of California to Peru (including the Galapagos), by having 11–13 scales above the lateral line ( vs . 9–10, Table 4b ); from Orthopristis forbesi , found in the Galapagos Islands, by having a finely-serrated pre-operculum (vs. smooth), 52–57 scales with pores in the lateral line ( vs . 80–85, Table 4a ), 11–13 scales above the lateral line ( vs. 9–10, Table 4b ), and 10 rays in the anal fin ( vs. 11); from Orthopristis reddingi , found from the Gulf of California to southern Mexico , by the scaleless soft dorsal and anal fins ( vs. with small scales on membranes) and 11–13 scales above the lateral line ( vs. 8, Table 4b ); from Orthopristis scapularis , found in northern and northeastern South America and the northern and northeastern Brazilian coast, by having a body with conspicuous vertical dark bars ( vs . without conspicuous vertical dark bars, sometimes with inconspicuous vertical dark bars, Fig. 1 ), a large and conspicuous black humeral spot above the base of the pectoral fin base ( vs. indistinct or rarely visible, Fig. 1 ), a small black spot, present on rear border of operculum ( vs. absent, Fig. 1 ), 13– 15 rakers in the lower limb of the first gill arch ( vs. 10–12, Table 4g ), 20–23 rakers in the second gill arch ( vs. 16– 19, rarely 20, Table 4h ); and from the Orthopristis hybrid by having 20–23 rakers in the second gill arch ( vs. 17–19 in the hybrid, Table 4h ). Molecular diagnosis. The DNA barcode of O. ruber shows that the specimens analyzed form a distinct, strongly supported cluster, with a bootstrap value of 99% ( Fig. 2 ), with genetic distances between this species and all the congeners analyzed ranging from 3.6% to 10.6%. The genetic distance to Atlantic species is 3.6% to O. scapularis and 10.6% to O. chrysoptera ( Table 5 ). The O. ruber haplotypes differ from all other members of the genus analyzed in 16 to 47 sites, with 16 from O. scapularis and 47 from O. chrysoptera in the Atlantic. Description. Dorsal-fin rays XII, 14–15 (usually 15), first unbranched, remaining branched; anal-fin rays III, 10, all branched; pectoral-fin rays 17–18, first ray shorter, unbranched, second ray about three times longer, unbranched, remaining ones branched; pelvic-fin rays I, 5, all branched; principal caudal-fin rays 9 + 8 = 17, uppermost and lowermost rays unbranched; lateral-line scales 52–57; scales above lateral line to base of first dorsal-fin spine 10–13 and 15–17 scales below lateral line to first anal-fin spine; gill rakers on first arch 21–25 (8– 11 on upper); gill rakers on second arch 20–23 (rarely 18 or 19); branchiostegal rays 7. Body compressed, relatively deep and long, greatest depth at spinous dorsal origin. Head large, dorsal profile steeply inclined from snout to first dorsal fin spine, with slight concavity before nostrils, also over eye in some specimens, forming small bump. Snout convex, moderately long as wide. Eye large, ellipsoid. Nostrils small, anterior teardrop shaped, with small dorsal skin flap, above horizontal line through lower third of eye, posterior oval, crowned with tiny slender, fleshy papillae, about half the size of first, slightly above anterior one. Mouth large, tip of maxilla reaching a vertical through rear margin of posterior nostril (reaching anterior eye margin in juveniles); lips fleshy and relatively thick, upper lip protruding slightly beyond tip of lower jaw; teeth conical, outer series enlarged (more so on upper jaw), about 20 on each side; 6 tooth rows anteriorly on each jaw, one or two rows posteriorly; numerous small fleshy papilla interdigitating among teeth; vomer and palatines toothless; tongue smooth, with broadly rounded tip; lower margin of chin with 3 prominent pores, a pair anterior and one posterior, pattern resembling the holes in a bowling ball; cheeks and chin pierced by irregular series of minute pores, difficult to discern from scale sockets; gill rakers moderately long, without spines, those on lower arm longest, about half pupil length, raker on angle shortest. Opercle lacking exposed spine, covered with 6–7 vertical rows of ctenoid scales; posterior upper edge of preopercle finely serrate (more so in juveniles) covered with skin, rough ventrally, with a small bump; subopercle and interopercle borders smooth. Small, ctenoid scales (sometimes covered with skin) on flanks, belly from nearby branchial isthmus to anal fin origin, cheeks (about 15 series under eye to preopercle) and interorbital to line between nostrils; lateral line continuous, gently curved anteriorly, following dorsal contour of body, straight on caudal peduncle; bases of dorsal fins with sheath of 1–2 rows of ctenoid scales, basal sheath over anal fin base with 3 rows of ctenoid scales, inter radial membranes of dorsal and anal fins naked. Scales on pectoral fin base, also with rows of 6 scales along pectoral rays. Dorsal fin scarcely notched, its origin just anterior to vertical from posterior margin of opercle; base of soft dorsal-fin usually subequal to that of spinous portion; interspinous membranes distinctly incised; dorsal-fin spines strong, fourth spine longest; origin of anal fin below base of 2 nd –3 rd soft dorsal-fin ray; first anal-fin spine short, remaining spines stout, more than 2 times longer than 1 st ; anal-fin base short; caudal fin forked, emarginated in specimens smaller than 100 mm SL. Pectoral fin slightly falcate, not reaching or barely reaching tip of pelvic fin, fifth pectoral-fin ray longest, subequal to pelvic fins; origin of pelvic fins posterior to lower base of pectoral fins, on vertical from base of 4th to 5th dorsal-fin spine; pelvic-fins nearly reaching anus (reaching anus in juveniles and some subadults), first ray longest (second subequal to first). FIGURE 1. Body in lateral view, Orthopristis ruber (left column). A–B. Syntypes of (MNHN-IC-A-0457 and MNHN-IC-A- 7821). C. Live specimen (not preserved, São Paulo state, image of Gabriel Togni). D–E. Fresh specimens (not preserved, D. Santa Catarina state, image of Leonardo Machado, E. Espírito Santo state, image of João Gasparini). F. Fixed specimen (AZUSC 1812, São Paulo). Body in lateral view, Orthopristis scapularis (right column). G. Holotype of O. scapularis (ANSP 45084). H. Live specimen (not preserved, Ceará state). I-J-K. Fresh specimens (not preserved, I. Rio Grande do Norte state, image of Alfredo Carvalho, J. Venezuela, image of James van Tassell & Ross Robertson, K. Amapá state, image of Marceniuk). L. Fixed specimen (MPEG 34292, Alagoas state). FIGURE 2. Neighbor-joining tree of the species of Orthopristis based on partial sequences of the cytochrome oxidase c subunit I. Numers near nodes represent bootstrap support. Color in life. Light gray, lighter ventrally, dark brown dorsally on head from upper lip to origin of first dorsal fin. Black spots (dark brown ventrally) ¼ pupil length on flanks, on rear edge of scales, smaller dots behind upper jaw, on pre-operculum and operculum. Irregularly horizontal black bar from rear eye border to posterior margin of operculum, width equal to pupil length, forming small, dark black spot posteriorly. Large and conspicuous oval humeral black spot above pectoral fin base, followed by six vertical black bars, width equal to eye length, last three bars darker on caudal peduncle. First dorsal fin hyaline, with four dark bars continuous with the bars on the body, second dorsal fin dark brown; pectoral and pelvic fins hyaline, anal fin dark yellow, caudal fin dirty beige, dark on base ( Fig. 1c ). Dark bars fade soon after death. Color of preserved specimens. Light brown, lighter ventrally, with dark spot on rear border of scales, forming irregularly oblique dark brown lines on the flanks. Dark brown spots on operculum and below eyes. Small, marked black spot on rear border of operculum and a conspicuous black humeral spot above pectoral fin base. Trunk with six dark faded irregular vertical bars, width nearly equal to eye length. Dorsal and anal fins hyaline, with dark brown line along the base of the hyaline pectoral and pelvic fins, caudal fin dusky, darker in middle rays ( Fig. 1f ). Distribution and habitat. Widely distributed in Brazil from Espírito Santo state (eastern coast) to Rio Grande do Sul (southern coast). The species is abundant in estuarine and coastal waters over sandy, rocky or muddy bottoms, to depths of 70 m ( Fig. 4 ). Size Maximum TL 400 mm , commonly 300 mm . Fisheries. Orthopristis ruber is frequently caught by artisanal fishers using lines, weirs or gillnets. The species is also taken as by-catch, in particular by industrial pink shrimp fisheries, and is typically retained for sale, rather than being discarded ( Vianna & Almeida 2005 ). Remarks. Orthopristis lineatus Cuvier (1830) was based on a specimen collected together with the holotype of Orthopristis ruber , in Rio de Janeiro , Brazil . This species was subsequently synonymized with O. ruber , without comment, by Jordan & Fesler (1893) , an arrangement followed by Miranda Ribeiro (1918) and all subsequent authors. However, the correct identification of this species is difficult to ascertain. While the holotypes of O. ruber and O. lineatus have the same fin ray and scales counts, the coloration of O. lineatus is distinct from that observed in O. ruber , in particular the absence of the black vertical lines. The synonymy of Orthopristis lineatus with O. scapularis is also difficult because the bronze spots forming oblique lines on flanks, typical of O. scapularis , were not detected in the only extant syntype (dry specimen) of O. lineatus ( Bauchot et al . 1983 ) . It is possible that O. lineatus may represent a hybrid form (see description of hybrid below), but as the type specimen was stuffed, it is impossible to obtain counts of the gill rakers, so the taxon is treated herein as Orthopristis nomen dubium.