Orthopristis scapularis Fowler 1915

Marceniuk, Alexandre Pires, Caires, Rodrigo Antunes, Machado, Leonardo, Cerqueira, Najila Nolie Catarine Dantas, Serra, Rayla Roberta M. De S. & Oliveira, Claudio, 2019, Redescription of Orthopristis ruber and Orthopristis scapularis (Haemulidae: Perciformes), with a hybridization zone off the Atlantic coast of South America, Zootaxa 4576 (1), pp. 109-126 : 117-119

Orthopristis scapularis Fowler 1915

( Figures 1 g–l, Tables 3 & 4)

Orthopristis scapularis Fowler 1915: 536 , Fig. 4 [type locality: Port of Spain, Trinidad Island, West Indies]. Holotype (unique): ANSP 45084.— Fowler 1916: [fishes Costa Rica; listed].— Fowler 1944: 467 [Vanderblit Expedition; Costa Rica; Colon; listed].— Böhlke 1984: 148 [type catalog]

Orthopristis ruber (not of Cuvier, 1830).— Uyeno et al. 1983:351 [fishes trawled off Suriname and Guyana; listed.— Cervigón 1992:336 [fishes septentional coast of South America; listed].— Cervigón 1993:213 [fishes of Venezuela; listed].— Aguilera 1998:50 [fishes of Venezuela, western portion; listed].— Rocha et al. 1998: 564 [fishes of coast of Paraíba, Brazil; listed].– Camargo & Isaac 2001:142 [fishes of northern Brazilian coast; listed].— Lindeman & Toxey 2003:1548 [fishes of western Central Atlantic; list and short description].—Moura & Menezes in Menezes et al. 2003:85 [in part; catalog of marine fishes of Brazil; listed].— Tavera et al. 2012:6 [phylogeny of Orthopristis ].— Marceniuk et al. 2017:41 [fishes of Caeté; northern Brazilian coast; listed].–Tavera et al. 2018:214, 221 [phylogeny of Orthopristis ].

Material examined. (52 specimens, 122–224 mm SL). Venezuela: USNM 361933, 2: 122–136 mm SL, Gulf of Venezuela, Venezuela. French Guiana: USNM 400901 , 4 : 172–197 mm SL, off coast of French Guiana, French Guiana. Brazil: MPEG 35856 , 3 : 192–200 mm SL, 3°44.277'N– 50°12.030'W, Amapá, Brasil; MPEG 35713 , 1 : 220 mm SL, 3°41.411'N– 50°20139'W, Amapá, Brasil; MPEG 35601 , 1 : 131 mm SL, 2°19.122'N– 48°36.465'W, Amapá, Brasil; MPEG 35633 , 2 : 134–148 mm SL), 2°07.783'N– 48°30.1'W, Amapá, Brasil; MPEG 35637 , 1 : 153 mm SL, 2°03.645'N– 48°30.390'W, Amapá, Brasil; MPEG 35647 , 1 : 149 mm SL, 1°51.812'N– 48°17.320'W, Amapá, Brasil; MPEG 35109 , 1 : 219 mm SL, 1°04´77´´N– 47°54´06´´W, Amapá, Brasil; MPEG 35085 , 1 : 224 mm SL, 1°04´32´´N– 47°54´00´´W, Amapá, Brasil; MPEG 3564 , 1 : 161 mm SL), Amazon estuary, Pará, Brazil; MPEG

33154, 2: 182–190 mm SL, Cuiarãna, Pará, Brazil; MPEG 33155 , 1 : 195 mm SL, Cuiarãna, Pará , Brazil ; MPEG 33153 , 1 : 214 mm SL, Ajuruteua, Bragança, Pará , Brazil ; MPEG 34407 , 1 : 196 mm SL, Ajuruteua, Bragança, Pará , Brazil ; MPEG 34420 , 3 : 174–191 mm SL, Ajuruteua, Bragança, Pará , Brazil ; LBP 21413, 5 : 167–184 mm SL, Bragança, Pará ; MPEG 34576 , 5 : 127–195 mm SL, estuary of the River Timonha, Chaval , Piaui ; MPEG 34639 , 5 : 143–155 mm SL, Morgado beach, Acaraú , Ceará ; MPEG 34576 , 5 : 127–159 mm SL, Chaval , Piaui ; MPEG 34247 , 3 : 123–136 mm SL, Paripuera, Sauaçuhy , Alagoas ; MPEG 34292 , 1 : 127 mm SL, Jatiúca beach, Maceió , Alagoas ; MPEG 34264 , 3 : 132–148 mm SL, Jaraguá, Maceió , Alagoas ; MPEG 34252 , 1 : 144 mm SL, Jaraguá, Maceió , Alagoas .

Diagnosis. Orthopristis scapularis can be differentiated from its congeners as follows: from Orthopristis chrysoptera , found from New York to Cuba, by having 10–11 rays in the anal fin (vs. 12–13); from Orthopristis cantharinus , found in the Galapagos Islands, by having 51–57 scales with pores in the lateral line (vs. 65–72, Table 4a), and 10–11 rays in the anal fin (vs. 12–13); from Orthopristis chalceus , found from the Gulf of California to Peru (including the Galapagos), by having an inconspicuous humeral spot (vs. humeral spot absent); from Orthopristis forbesi , found in the Galapagos Islands, by having a finely-serrated pre-operculum (vs. smooth preoperculum), 5–57 scales with pores in the lateral line (vs. 80–85, Table 4a) 10–12 rakers in the lower limb of the first gill arch (vs. 13–14, Table 4g); from Orthopristis reddingi , found from the Gulf of California to southern Mexico, by the absence of scales on the soft dorsal and anal fins (vs. small scales on the membranes) and 9–11 scales above the lateral line (vs. 8, Table 4b); from Orthopristis ruber , found off the southern and southeastern coast of Brazil, by the lack of the conspicuous vertical dark bars, sometimes with inconspicuous vertical dark bars (vs. with conspicuous vertical dark bars, Fig. 1 ), humeral black spot above pectoral fin base indistinct or rarely visible (vs. very distinct, Fig. 1 ), small black spot on rear border of operculum absent (vs. present, Fig. 1 ), 10–12 rakers on inferior limb of the first gill arch (vs. 13–14, Table 4g), 16–29, rarely 20 rakers in the second gill arch (vs. 20–23, rarely 20, Table 4h); and from the Orthopristis hybrid by having 13–15 rakers on the lower limb of the first gill arch, vs. 10–12 ( Table 4g).

Molecular diagnosis. The DNA barcode of O. scapularis shows that the specimens analyzed form a distinct, strongly supported cluster, with a bootstrap value of 99% ( Fig. 2 ) and a K2P divergence of 3.6–10.9% from all other Orthopristis sequences analyzed. The genetic distance to Atlantic species ranged from 3.6% for Orthopristis ruber to 10.9% for O. chrysoptera ( Table 5). The haplotypes of O. scapularis differed from all other members of the genus analyzed at between 16 and 51 sites, ranging from 16 in the case of O. ruber , and to 51 for the O. chrysopterain Atlantic.

Description. Dorsal-fin rays XII, 14–15, first unbranched, remaining branched; anal-fin rays III, 10, all branched; pectoral-fin rays 17–19 (rarely 19), 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 51–57; scales above lateral line to base of first dorsal-fin spine 9–12 and scales below lateral line to first anal-fin spine 13–16 (usually 15); gill rakers on first arch 18–23 (8–11 on upper); gill rakers on second arch 15–20 (rarely 20); 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 digitating 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 with 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 diameter, raker at 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 an 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).

Color in life. Light beige, whitish ventrally, brown dorsally on head from upper lip to origin of first dorsal fin. Bronze spots about half pupil length on flanks, over rear border of scales, forming wavy lines ventrally, and corresponding yellow spots below eye, pre-operculum, and operculum. Irregular horizontal bronze bar from rear eye border to posterior margin of operculum, width equal to pupil diameter. Dark irregular humeral spot faded, nearly equal to eye length; black vertical bars missing. First dorsal fin hyaline, second dorsal fin orange bronze; pectoral and pelvic fins hyaline, anal fin dark dusky, caudal fin dirty orange, with dark brown hue on base of upper rays ( Fig. 1h ).

Color of preserved specimens. Light gray to brown, lighter ventrally, with dark spot on rear border of scales, forming irregular oblique dark brown lines on flanks. Dark brown areas on operculum and below eyes; black humeral spot and dark bars on flanks missing. Dorsal and anal fins hyaline, with dark brown line along bases pectoral and pelvic fins hyaline, caudal fin dusky, darker in middle rays ( Fig. 1l ).

Distribution and habitat. Widely distributed from Santa Marta, Colombia to the Bahia state in northeast coast of Brazil. The species is abundant in estuarine and coastal waters, in depths to 70 meters over sandy, rocky or muddy bottoms ( Fig. 4 ).

Size. Maximum TL 400 mm, commonly 300 mm.

Fisheries. Orthopristis scapularis is an important local fishery resource and is sold both fresh and salted ( Anderson et al., 2015). It is caught by artisanal fishers using lines, weirs or gillnets. In Pará, the species is taken as by-catch by the industrial pink shrimp fleet, and is not retained for sale (Marceniuk et al. in press).

Remarks. Orthopristis scapularis was described by Fowler (1915), based on one specimen from Port of Spain, Trinidad, in the British West Indies ( Fig. 4 ), with the specific epithet alluding to the presence of a spot in the humeral region. Fowler concluded that this new species was related closely to O. chrysoptera , known from the waters of the northern Gulf of Mexico, eastern United States, Cuba and Bermuda. Orthopristis scapularis differs in having fewer anal fin rays (10–11 vs. 12–13). Since this publication, the species was cited by Fowler (1944) in inventories in the Caribbean Sea ( Costa Rica) by the Vanderbilt Expedition and other vessels, but it was never treated in subsequent publications ( Courtenay & Salman 1978; Böhlke 1984; Lindeman & Toxey 2003). In Fowler’s original description of O. scapularis , the recorded 12 rakers on the lower limb of the first branchial arch has 12 rakers, which is consistent with the value recorded by Uyeno et al. (1983), i.e., 11 rakers, in Suriname, and lower than the raker counts recorded for O. ruber specimens from the southeastern coast of Brazil. Even so, the results of our survey indicate that populations from the central western Atlantic, as far as northeastern Brazil are distinct from those of southern and southeastern coast of Brazil, based on the lack of the conspicuous vertical dark bars, sometimes with inconspicuous vertical dark bars ( Fig. 1 ), humeral black spot above pectoral fin base indistinct or rarely visible ( Fig. 1 ), small black spot on rear border of operculum absent ( Fig. 1 ), and gill rakers counts ( Table 4 f-h), which supports the validity of O. scapularis , a position reinforced conclusively by the molecular evidence ( Fig. 2 ).

Figures

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.

FIGURE 4. The Atlantic coast of South America, showing the geographic distribution of Orthopristis scapularis (green), Orthopristis ruber (yellow), Orthopristis hybrid (red), locations with material, in zoological collections, not examined (dark blue), and localities with record in the literature (light blue). Some symbols represent more than one locality or a large number of specimens. Non-type specimens are represented with circles, and type specimens are indicated by square