Pareques lineatus ( Cuvier, 1830 )

Carvalho-Filho, Alfredo, Oliveira, Claudio, Maximiano, Loriane, Tavera, Jose, Acero, Arturo & Marceniuk, Alexandre Pires, 2022, Review of the Pareques acuminatus (Bloch & Schneider, 1801) species complex with revalidation of Pareques lineatus (Cuvier, 1830) from the Western Atlantic (Percomorphacea: Sciaenidae), Zootaxa 5195 (5), pp. 401-418 : 409-414

Pareques lineatus ( Cuvier, 1830)

Figures 1 and 2 , Tables 2 and 3

Type specimens: Syntypes: MNHN 0043 (1), Brazil ; MNHN 7475 (1), Brazil .

Non-type specimens: CIRUV 017067 Colombia, Banco Animas ; UVTP 2557 Colombia, Taganga ; UF 214350 (1, 131 mm SL), 17.9119370 -67.0477090, Puerto Rico, La Parguera , fish pots ; AZUSC 5848 (7, 97–129 mm SL), Brazil, Paraíba , João Pessoa, Penha beach; NPM 3257 (1, 83 mm SL), Brazil, Pernambuco , off Pernambuco; CIUFES 1699 (1, 97 mm SL), Brazil, Espírito Santo , Vitória, Ilha do Boi ; CIUFES 1964 (1, 116 mm SL), Brazil, Espírito Santo , Itapemirim, Ilha dos Franceses ; NPM 2915 (1, 103 mm SL), Brazil, Rio de Janeiro , Macaé, Arquipélago de Santana ; AZUSC 4484 (3, 100–116 mm SL), Brazil, São Paulo , Ubatuba, Ilha das Couves ; MZUSP 70555 (1, 114 mm SL), Brazil, São Paulo , Ubatuba, Ubatuba bay ; AZUSC 2955 (1, 144 mm SL), Brazil, São Paulo , São Sebastião, Toque Toque Pequeno; AZUSC 4526 (1, 161 mm SL), Brazil, São Paulo , São Sebastião, Toque Toque Pequeno; MZUSP 46228 (2, 70–110 mm SL), Brazil, São Paulo , São Sebastião, Baraqueçaba beach; MZUSP 2460 (1, 143 mm SL), Brazil, São Paulo , Ilha da Vitória ; MZUSP 45594 (1, 125 mm SL), Brazil, São Paulo , Arquipélago dos Alcatrazes; MZUSP 44647 (1, 120 mm SL), Brazil, São Paulo , Santos, Laje de Santos ; AZUSC 3905 (1, 151 mm SL), Brazil, São Paulo , Cananéia, Ilha do Bom Abrigo ; MZUSP 55386 (1, 135 mm SL), Brazil, Santa Catarina , Florianópolis, Ilha do Arvoredo ; MZUSP 46641 (1, 143 mm SL), Brazil, Santa Catarina , Porto Belo; MZUSP 49098 (2, 115– 134 mm SL), Brazil, Santa Catarina , Porto Belo.

Diagnosis: Pareques lineatus , is distinguished from its congeners in the western Atlantic as follows: from P. acuminatus by having 50–55 lateral-line scales, excluding specimens from Colombia (vs. 44–49; Table 3 ), 8–9 scales from lateral line to origin of dorsal fin (vs. 6; Table 2 ) and 9–10 scales from lateral line to first anal fin spine (vs. 8; Table 2 ) and coloration of adults, brownish with 5–7 longitudinal, irregular white stripes thinner than dark ones (vs. 8 straight white stripes wider than dark ones; Fig. 2h ) and by absence of submarginal white stripe on soft dorsal fin (vs. presence of whitish submarginal stripe); from P. iwamotoi by absence of dark bar between eyes in adults (vs. presence of dark bar between eye), and absence of wide, transverse dark bar from dorsal-fin origin almost to pelvic-fin base (vs. presence of transverse bar); and from P. umbrosus by 5 pores on chin (vs. 6 pores), and color, brownish with white horizontal stripes (vs. brownish with thin horizontal black lines).

Pareques lineatus can also be distinguished from its congeners from the eastern Pacific as follows: from P. fuscovittatus by first dorsal without yellow rear edge (vs. with yellow rear edge), 10 dorsal-fin spines (vs. 12), 12–15 gill rakers in first arch (vs. 16–20); from P. lanfeari by head with horizontal dark stripes (vs. without horizontal dark stripes), body with 3–5 broad dark stripes and thin pale stripes between those (vs. body with 4 thin black horizontal stripes) and iris brown (vs. iris yellow-brown); from P. perissa , by having 9–11 dorsal-fin spines (vs. 12), 37–40 dorsal-fin soft rays (vs. 32–33) and seven anal-fin soft rays (vs. 8); from P. sp. A, by having 12–15 gill rakers on first arch (vs. 19–23), 10 dorsal-fin spines (vs. 8–9), and 37–40 dorsal-fin soft rays (vs. 42–45); from P. viola , by having 12–15 gill rakers on first arch (vs. 17).

Molecular diagnosis: The DNA barcode of Pareques lineatus shows that the specimens analyzed form a distinct, strongly supported cluster, with a bootstrap value of 92% ( Fig. 3 ), with genetic distances between this species and all the congeners analyzed ranging from 6.2% to 14.3% ( Table 4 ). Pareques lineatus presents 29 positions in COI with different nucleotides when compared directly with Pareques acuminatus ( Table 5 ). Pareques lineatus presents 6 unique haplotypes and differs from P. acuminatus by 29 mutational steps in COI ( Fig. 4 ).

Description: Counts and measurements in Tables 2 and 3 . Body oblong, compressed, moderately deep, dorsum elevated, tapering posteriorly to slender caudal peduncle. Head upper profile moderately steep, slightly concave from snout tip to dorsal-fin origin; lower profile of head almost straight from mandibular tip to pelvic-fin origin; snout blunt, rounded, overhangs upper jaw; posterior nostril elongate, larger than oval anterior nostril; mouth small, inferior, almost horizontal, lower jaw included; teeth villiform in both jaws, those of upper-jaw outer row enlarged, recurved; vomer and palatines toothless; upper jaw barely reaches middle of orbit; three marginal pores on snout and three smaller pores on rostrum; five pores on chin, one medial and two mental pairs; eye diameter moderate, larger than interorbital, similar to snout length; preopercular border smooth, membranous.

First dorsal fin higher than second, reaching 4 th ray when adpressed in adults, farther in younger or immature, and even caudal-fin distal end in juveniles; second dorsal fin long, posterior border somewhat rounded; pectoralfin insertion below about middle of first dorsal fin, tip pointed; pelvic-fin length slightly shorter than pectoral fin, its origin below or just after posterior pectoral-fin base, tip rounded to pointed, reaching beyond pectoral-fin tip in adults, much longer in juveniles; anal-fin base short, distal tip usually pointed, second spine rather stout; caudal fin convex to almost straight in adults, lanceolate to filamentous in juveniles.

Body and much of head covered with ctenoid scales; small ctenoid scales covering soft dorsal- and anal-fin bases. Swim bladder simple, carrot-shaped, without distinct diverticula or anterior horns.

Live colors. Adults ( Fig. 2h ). Generally dark brown to brownish grey head, body, and fins, often with golden tones on face and belly. A series of whitish longitudinal, somewhat irregular stripes on upper head, rarely on lower, and anterior body: the first a thin one extending from the insertion of the first dorsal fin along each side of the base of the second dorsal fin to about mid-fin; the second and third start together on the front of the nape then divide and pass along the upper back on each side of the body, enclosing a dark elongate triangle, then join again under the middle of the second dorsal and continue for a short distance further along the base of the soft dorsal; the fourth starts on the forehead above the eye and the fifth at the top edge of the eye, then both run parallel to near the rear of the soft dorsal, then join and run along the upper edge of the caudal peduncle; the sixth runs from the lower edge of the eye along the top edge of the pectoral base and along the lower flank to above the anal fin, where it fades out as it continues along the center of the caudal peduncle; the seventh runs parallel to the sixth, which is much more faded, and joins it above the anal fin; fins greyish brown, the border of soft dorsal with a thin black line and the posterior margin of the spinous dorsal fin usually dark, sometimes white; anterior portion of snout and lips bluish black; forehead often with a broad whitish blotch that reaches the dorsal-fin origin; iris golden brownish, pupil black.

Juveniles: Recently recruited individuals ( Fig. 2e ): yellowish white; a black transverse bar crossing upper head, a black stripe in the interorbital and two small conspicuous black dots, one on snout tip, the other just below it; a black stripe from anterior border of eye, crossing orbit to the end of the middle caudal fin; a black stripe from tip of the white spinous dorsal fin that curves back on the shoulder and merges with the lateral one below the center of the soft dorsal; soft dorsal and anal fins translucent; pectoral fins translucent, with a black basal section; pelvic fin white with a black bar down its center.

Changes with growth ( Fig. 2f, g ): As the fish grows, the existing black stripes shrink, and two additional stripes form, one running from upper head to middle of the soft dorsal and another one from below eye to lower caudal peduncle; pectoral and pelvic fins became black with outer borders white; tip of snout with large black spot and a bar from upper jaw to eye and another in the interorbital; the first dorsal is black with white outer borders and a black-centered ocellus at about the middle of the anterior part of the fin. With further growth, the black body stripes widen, become brown, and increase in number to 5–7, the first dorsal fin becomes dark with a white band along its rear edge and the ocellus reduces its size to just a dark area at the tip of the fin; the pectoral, pelvic, anal, and caudal fins are greyish with black outer borders. As the fish grows further, the overall color becomes brown, and the white color is restricted to the stripes described for adults and the spiny dorsal-fin ocellus disappears completely.

Nocturnal coloration: At night, juveniles and adults sometimes display a large black bar that runs from the tip of the spinous dorsal fin to the tip of the pelvic fin and another smaller bar from the middle of the second dorsal fin to the middle of the anal fin.

Color of preserved specimens: adults and immature specimens brown color and white stripes becomes more grayish and faded. The juveniles maintain their basic coloration, the black marks eventually slightly faded.

Distribution and habitat: Pareques lineatus apparently occur from Puerto Rico eastern through the Lesser Antilles south to Santa Catarina, Brazil ( Fig. 4 ). The northern limit of the range of P. lineatus is not clear: meristics of samples from Trinidad & Tobago (USNM 413238 and 413240) agree with P. acuminatus and the coloration with P. lineatus (pictures of live specimens can be found at https://biogeodb.stri.si.edu/caribbean/en/gallery/specie/3802), also meristics of a voucher from French Guiana (UF 242486) agree with P. acuminatus whereas its coloration is the same as P. lineatus . It thrives in rocky and coral reefs from one to about 100 meters deep, usually between three and 30 meters. Recently recruited juveniles can be observed in very shallow water under ledges or near sand pockets close to reef crevices, often in groups of 3–7 of about the same age. Young are usually found in small groups up to three individuals in depths between five and 20 meters, Adults are observed in groups of three to at least 25 individuals, among rocks, under ledges, inside caves and, eventually, at daylight, in the water column, mixing schools with fishes of similar size, such as haemulids ( Fig. 5a ). Their habits are mainly nocturnal, spending the day sheltered and feeding at night on sand, grass beds, mud, and rubble bottoms close to the reef, feeding on crustaceans, polychaetes, small bony fishes, zooplankton, and algae. A group of about 20 adults was observed and photographed by Dr. Cláudio L.S. Sampaio ( Fig. 5b ) cleaning and picking mucus or parasites off a large stingray, Hypanus berthalutzae Petean, Naylor & Lima, 2020 , that stayed quietly at the bottom, in Aracaju, Sergipe, Brazil.

Remarks: The original description of Cuvier (1830) is based in several specimens collected from Brazil, two of which become syntypes (MNHN 0043 and 7475; Fig. 1 ). He compared theses fishes only with Eques balteatus [= Eques lanceolatus ] and Pareques punctatus [= Eques punctatus ], stating correctly that the first dorsal fin was not so high, the number of rays was smaller, the teeth were stronger, and the longitudinal, grayish, stripes do not turn up to the back of neck. Also, he described their coloration as dark brown with 6–7 thin grayish longitudinal stripes. Unfortunately, he gave no scale counts, some of which are diagnostic characters for these species. Examining the pictures of the syntypes, we were able to count the lateral-line pores, the series of transverse scales, and the number of scales from the lateral line to the origin of the dorsal fin, and those from the lateral line to the origin of the anal fin. Thus, we concluded that Pareques lineatus is a valid species based on the color pattern and scale counts of the types ( Fig. 2 ; Table 3 ). Eques pulcher , as described by Steindachner (1867) is here considered as junior synonym of P. lineatus based in the lateral-line scale count (50).

Figures

FIGURE 1. Pareques acuminatus (A, B) and Pareques lineatus (C, D), body in lateral view. (A) illustration, reproduced from Seba (1759: pl. 26, fig. 33); (B) neotype, UF 210054, Florida, USA; (C) syntype, MNHN 7475, Brazil; (D) syntype, MNHN 0043, Brazil.

FIGURE 2. Color patterns of Pareques acuminatus (left column) and P. lineatus (right column). A, B: recently recruited individuals; C, D: juveniles, about 10–35 mm SL; E, F: immatures, about 35–80 mm SL; G, H: adults. Images A, C, E, and G, Carlos and Allison Estape, Florida, USA; B, Alfredo Carvalho-Filho, Praia do Forte, Bahia, Brazil; D and H: Carlos E.L. Ferreira, Arraial do Cabo, Rio de Janeiro, Brazil; F: Athila Bertoncini, Taipus de Fora, Bahia, Brazil.

FIGURE 3. Molecular phylogeny constructed by maximum likelihood method based on K2P+G distance with partial sequences of the mitochondrial gene cytochrome c oxidase subunit I. The tree with the highest log likelihood (ln L = -2351.51) is shown. Numbers on each branch indicate bootstrap values; values below 50% are not shown.

FIGURE 4. Sampling localities of examined material and haplotype network. Squares: specimens only with DNA sequenced; triangles: specimens only with morphological data; circles: specimens with sequenced DNA and morphological data. Pareques acuminatus (green) and Pareques lineatus (yellow). Question marks are of specimens with intermediate characters. Haplotype network of COI gene, constructed using the maximum likelihood approach, with the evolutionary models GTR+I+G. In the haplotype network, samples in red represent fishes from Colombia or Panama and samples in pink represent fishes from Virgin Islands.

FIGURE 5. A. A daylight school of Pareques lineatus (black individuals) and Haemulidae in open water. B. A group of P. lineatus picking mucus or parasites off a large stingray, Hypanus berthalutzae. Pictures from Aracaju, Sergipe, Brazil, by Cláudio L.S. Sampaio/UFAL.

Tables

TABLE 5. Positions in the COI gene that differ between each species.

	61	67	70	71	97	199	202	232	241	262	283	289	328	334	400
P. acuminatus	T	G	G	C	G	A	C	C	A	T	G	A	G	G	A
P. lineatus	C	A	A	T	A	G	G	T	G	G	A	G	A	A	G
	445	454	460	469	484	490	526	535	565	577	589	595	613	619
P. acuminatus	T	T	C	T	G	G	C	C	C	T	C	A	C	T
P. lineatus	C	C	T	C	A	A	T	A	T	C	A	G	T	C