Menticirrhus gracilis ( Cuvier, 1830 )

Marceniuk, Alexandre Pires, Caires, Rodrigo Antunes, Rotundo, Matheus Marcos, Cerqueira, Najila Nolie Catarine Dantas, Siccha-Ramirez, Raquel, Wosiacki, Wolmar Benjamin & Oliveira, Claudio, 2020, Taxonomic revision of the Menticirrhus americanus (Linnaeus, 1758) and M littoralis (Holbrook, 1847) (Percomorphacea: Sciaenidae) species complexes from the western Atlantic, Zootaxa 4822 (3), pp. 301-333 : 317-320

Menticirrhus gracilis ( Cuvier, 1830)

Figure 6 , Tables 3, 4

Umbrina gracilis Cuvier in Cuvier & Valenciennes 1830: 189 ( Brazil. Lectotype drawn by Chao, 1978: MNHN 44; paralectotype: MNHN 9622).— Günther 1860: 277 (catalog of fishes of the British Museum; short description).— Bauchot & Desoutter 1987:19 (type catalog)

Menticirrhus martinicensis (not of Cuvier 1830).— Jordan 1887: 539 (type specimens of the species described by Cuvier & Valenciennes; synonymy).— Jordan & Eigenmann 1899: 429 (review of the American sciaenids; synonymy).— Jordan & Evermann 1896: 1474 (fishes of North America ; description; synonymy)

Menticirrhus americanus (not of Linnaeus, 1758).—Miranda Ribeiro 1915: 422 (Fauna Brasiliense; description, synonymy).— Carvalho 1941: 65-66 (in part; fauna of São Paulo; listed).— Vasconcelos 1945:115 (in part; account of the sciaenids from Brazil; listed).— Fowler 1951: 26 ( Brazil, Rio de Janeiro; listed).— Travassos & Paiva 1957: 142 (Brazilian sciaenids; identification key; listed).— Benvegnú-Lé 1978: 49 (fish fauna of Rio Grande do Sul, Brazil; listed).— Chao 1977: 29 (in part; FAO fact sheets; description).— Chao 1978: 31 (in part; basis for classifying western Atlantic sciaenids; description).— Menezes & Figueiredo 1980: 44 (in part; guide to the fishes from southeastern Brazilian coast; short description).— Jardim 1988: 182 (in part; synopsis of Menticirrhus ).—Casatti & Menezes in Menezes et al. 2003: 87 (in part; catalog of the marine fishes of Brazil; listed).

Menticirrhus littoralis (not of Holbrook, 1847).— Menezes & Figueiredo 1980: 45 (in part; guide to the fishes from the southeastern Brazilian coast; short description).— Cassano & Levy 1990: 87-91 (biochemistry; Rio Grande do Sul; Brazil; as Menticirrhus littoralis ).

Doubtful reference to this species

Menticirrhus littoralis (not of Holbrook 1847).— Nión et al. 2016: 49 (fishes of Uruguay, common names; listed).

Material Examined. Lectotype: MNHN 44 (1), Brazil . Paralectotypes: MNHN 9622 (1), Brazil . Non-type specimens: MZUSP 69842 (1, 186 mm SL), Ubatuba, São Paulo, Brazil ; MZUSP 121488 (1, 166 mm SL), Praia do Puruba , Ubatuba, São Paulo, Brazil ; MZUSP 69832 (4, 86.6–104.4 mm SL), Ubatuba , São Paulo, Brazil ; AZUSC 723 (3, 161– 222 mm SL), Bertioga , São Paulo, Brazil ; AZUSC 1400 (1, 157 mm SL), Ilha do Arvoredo, Guarujá, São Paulo, Brazil ; MZUSP 69834 (1, 136 mm SL), Guarujá, São Paulo, Brazil ; MZUSP 69975 (1, 118 mm SL), Santos, São Paulo, Brazil ; AZUSC 3171 (1, 153 mm SL), Forte de Itaipú , Praia Grande, São Paulo, Brazil ; AZUSC 2587 (1, 140 mm SL), Cibratel, Itanhaém , São Paulo, Brazil ; AZUSC 866 (2, 245- 300 mm SL), Praia dos Pescadores , Itahaém, São Paulo, Brazil ; AZUSC 989 (2, 224- 228 mm SL), Praia dos Pescadores , Itahaém, São Paulo, Brazil ; AZUSC 5716 (31, 64- 114 mm SL), Peruibe , São Paulo ; MZUSP 69835 (3, 109- 122 mm SL), Tramandaí, Rio Grande do Sul, Brazil ; MZUSP 7961 (1, 82 mm SL), 29°09’S 50°13’W, Tramandaí , Rio Grande do Sul, Brazil ; MZUSP 70175 (1, 355 mm SL), 32°09’S 52°50’W, Rio Grande do Sul, Brazil ; MZUSP 70176 (1, 338 mm SL), 31°45’S 51°08’W, Rio Grande do Sul, Brazil ; MZUSP 7959 (1, 101 mm SL), Praia do Cassino , Rio Grande, Rio Grande do Sul, Brazil .

Diagnosis. Menticirrhus gracilis , found on the southeastern and southern coasts of Brazil, can be differentiated from all its other western Atlantic congeners by having 18–21 dorsal-fin rays (vs. 22–27, rarely 21, Table 3). The species can further be differentiated from M. americanus , found in the United States and the Gulf of Mexico, by pectoral-fin tip barely reaching tip of depressed pelvic fin (vs. surpassing tip of pelvic fin, Fig. 6 ), body without dark bars (vs. with irregular dark bars, Fig. 6 ); from M. martinicensis , found in the Caribbean and South America, by pectoral-fin tip barely reaching tip of depressed pelvic fin (vs. surpassing tip of pelvic fin, Fig. 6 ), body without dark bars (vs. with irregular dark bars, Fig. 6 ); from M. littoralis , found in the United States and the Gulf of Mexico, by the pale caudal fin, without a distinctive dark spot on the dorsal lobe (vs. with a dark spot on upper lobe, Fig. 6 ), and from M. saxatilis , found in the United States and the Gulf of Mexico, by pectoral-fin tip barely reaching tip of depressed pelvic fin (vs. surpassing tip of pelvic fin, Fig. 6 ), body without dark bars (vs. with irregular dark bars, Fig. 6 ).

Menticirrhus gracilis can also be distinguished from its congeners in the eastern Pacific as follows: from M. elongatus , ranging from the Gulf of California to Peru, by having 18–21 dorsal-fin rays (vs. 22–24, Table 3), pectoral-fin tip barely reaching tip of depressed pelvic fin (vs. surpassing tip of pelvic fin, Fig. 6 ); from M. nasus , ranging from the Gulf of California to Peru, by having six or seven anal-fin rays (vs. eight), scales present along base of soft dorsal fin (vs. soft dorsal fin naked), pectoral-fin tip barely reaching tip of depressed pelvic fin (vs. surpassing tip of pelvic fin, Fig. 6 ); from M. ophicephalus , which occurs from Ecuador to Chile, by having 48–54 scales with pores along lateral line to caudal-fin base (vs. 63–66), six or seven anal-fin rays (vs. nine), S-shaped posterior margin of the caudal fin (vs. concave), scales present along base of soft dorsal fin (vs. soft dorsal fin naked); from M. paitensis , ranging from the Gulf of California to Peru, by 48–54 scales with pores along lateral line to caudal-fin base (vs. 76–98), 18–21 dorsal-fin rays (vs. 22–25, Table 3), 16–20 pectoral-fin rays (vs. 21–23, Table 3); scales present along base of soft dorsal fin (vs. soft dorsal fin naked), pectoral-fin tip barely reaching tip of depressed pelvic fin (vs. surpassing tip of pelvic fin, Fig. 6 ); from M. panamensis , ranging from the Gulf of California to Chile, by six or seven anal-fin rays (vs. eight or nine), scales present along base of soft dorsal fin (vs. soft dorsal fin naked), pectoral-fin tip barely reaching tip of depressed pelvic fin (vs. surpassing tip of pelvic fin, Fig. 6 ); from M. undulatus , found in California, United States, by 18–21 dorsal-fin rays (vs. 23–27, Table 3), and six or seven anal-fin rays (vs. eight or nine); scales present along base of soft dorsal fin (vs. soft dorsal fin naked).

Description: D. IX-XII+18–21; A. I+6-8; P. 16–19; C. 17; GR. 10–15; LL. 48–55; SA. 6–11; SB. 12–19 ( Tables 3, 4). Body slender in lateral view, not compressed, ventral profile flat; maximum depth at first dorsal-fin origin. Dorsal profile convex, ascending, from snout to first dorsal fin, sometimes with small concavity near snout tip, nearly horizontal along first dorsal-fin base, straight, slightly descending along second dorsal-fin base, concave on caudal peduncle. Ventral profile convex, descending at snout, slightly concave on head to pelvic fin base, nearly horizontal from pelvic fin to anal-fin origin, ascending, nearly concave, along anal-fin base to caudal-fin base. Snout long, deep, with blunt to pug nose tip, triangular in lateral view. Mouth subterminal, upper jaw slender, rear tip on vertical passing through middle of eye. Premaxilla with 4–6 rows of acicular teeth, external most slightly larger, 4–7 tooth rows on dentary. Eye slightly oval, nearly two times snout length, not adjoining dorsal profile. Interorbital space smaller than orbital diameter, slightly convex, covered with ctenoid scales. Nostrils small, anterior nostril one nearly round, posterior larger, oval, nearly on horizontal line through ventral pupil border. Head with five sensory pores on snout, in a semicircle around mouth angle, followed by larger central pore, lower jaw with four pores, two on each side encircling mental barbel, pores otherwise absent; mental barbel short, rigid, with blunt tip without pore. Lateral line slightly arched to second dorsal-fin origin, then descending straight to caudal peduncle, nearly horizontal elsewhere to caudal-fin tip. Preopercle margin rough, with about 10 subtle spines. Opercle tip fleshy, blunt, well behind vertical line that passes through pectoral-fin base. Gill rakers short, two or three in upper limb rudimentary. Ctenoid scales on trunk, belly, pectoral-fin base, opercle, preopercle, infraorbital, and interorbital region to snout, cycloid scales in gular area and encircling nostrils. Dorsal fin without evident scales in interradial membranes, second dorsal fin with basal sheath of two scales; pectoral-fin base covered by small ctenoid scales, rows of cycloid scales present along pectoral-fin rays (outer third and tip naked); caudal-fin base covered with large ctenoid scales, rows of cycloid scales along rays (tips naked). Spinous dorsal fin short, first spine shortest, second and third spines longest; second dorsal fin adjoining first one. Origin of second dorsal fin well behind vertical line through pectoralfin tip, second dorsal soft rays much shorter than longest first dorsal spines. Anal-fin origin in vertical line passing through sixth-seventh ray of second dorsal fin; first spine very slender. Pectoral fin falcate, not reaching vertical line passing through last spine of first dorsal fin and pelvic-fin tip. Pelvic-fin origin behind pectoral-fin base, as long as pectoral fin. Caudal peduncle short, deep, depth greater than orbital diameter. Caudal fin short, emarginated, lower lobe slightly longer and much stouter than upper lobe.

Color in alcohol. Type material with straw-beige pigmentation, mostly faded due to the age of the specimens. Other specimens examined: upper half light brown, lower creamy white, sometimes with a silvery hue. Yellow hue may be present below eye, in operculum, pelvic and caudal fins. Dorsal and anal fins straw-beige to light brown, darkening slightly along the rays; pectoral fin light brown, somewhat darker in the upper rays, but not forming a dark hue. Caudal-fin rays dusky, slightly darker at the base ( Fig. 6B ).

Color of fresh material. Silver gray dorsally to flanks, whitish ventrally; silvery hue below eye; first dorsal fin dusky, longest spines dark-tipped; second dorsal fin dusky at base, beige elsewhere; anal fin beige; ventral fin beige with black line along spine; pectoral fin dusky, darkened in upper 3–4 rays; caudal fin dusky, darker along upper and lower caudal lobes ( Fig. 6C ).

Distribution and habitat. Western South Atlantic along the southern and southeastern coasts of Brazilian, from Rio de Janeiro to Rio Grande do Sul ( Fig. 4 ).

Fishing. Targeted by recreational fishermen.

Remarks. Cuvier described Umbrina gracilis based on specimens collected in Brazil by Auguste de Saint-Hilaire and his colleagues from the Freycinet Expedition.As Freycinet was engaged in a circumnavigation the globe, it seems reasonable to assume that Saint-Hillaire met Freycinet’s naturalists in Rio de Janeiro, where Saint-Hillaire set out on his expeditions, and that Rio de Janeiro is the type locality of U. gracilis . Cuvier confirmed that the specimen he used to describe U. gracilis had 20 dorsal-fin rays and 18 pectoral-fin rays. Günther (1860) also included this species in his catalog of fishes of the British Museum and reported that it has 75 scales on lateral line and a uniform color. Günther’s counts of the lateral scales need to be treated with caution, however, given that he may have confused lateral line counts with those of the oblique series of scales found above the lateral line. Similar discrepancies were detected by Jordan & Evermann (1898) in their account of U. martinicensis .

Umbrina gracilis was synonymized with Menticirrhus martinicensis by Jordan & Eigenmann (1899), although once again, with no explicit justification for their decision, although Jordan & Evermann (1898) confirmed that the description of this nominal species was based on “the dried skin of a young specimen, distorted and varnished”. This convinced all subsequent authors to doubt the validity of the species.

In his unpublished thesis on the taxonomy of Menticirrhus, Irwin (1971 , Fig. 58) upheld the synonymy of U. gracilis with M. americanus , following previous authors, such as Miranda Ribeiro (1915) and Fowler (1951), although he noted the existence of considerable variation in the counts of the soft dorsal fin between populations from the western North Atlantic and western South Atlantic. Irwin did not conclude this analysis, however, because that he was unable to examine an adequate sample of the material from the western South Atlantic.

In his extensive review of the classification of sciaenids of the western Atlantic, Chao (1978) reassessed Irwin’s data, and concluded that he had recognized one of the U. gracilis syntypes (MNHN 9037) as M. littoralis , but unexpectedly claimed that Irwin had disregarded this specimen as part of the type material because the locality and the collector of the specimen are not consistent with the original description of Cuvier’s (Bauchot, MNHN, Paris, pers. commun.; see also Bauchot & Desoutter 1987). Chao disregarded the validity of U. gracilis , contrarily to Irwin (1971), including it in synonymy of M. americanus . However, Chao wrongly stated specimen MNHN 44 ( Fig. 6A ) as a lectotype.

We disagree with Chao’s viewpoint on the status of U. gracilis , and consider it to be a valid member of the genus Menticirrhus for the following reasons: (i) we observed several specimens with 18–21 dorsal-fin rays and 16–19 pectoral-fin rays, which are quite similar to the original description of U. gracilis , (ii) all the specimens with 18–21 dorsal-fin rays and 16–19 pectoral-fin rays are morphologically distinct from M. americanus , presenting a slender body with a long snout, color more or less uniformly silvery gray on the back, without irregular dark bars, and pectoral fin not reaching or barely reaching the pelvic-fin tip. In addition, other Menticirrhus species, such as M. americanus , M. cuiaranensis , M. martinicensis , and M. littoralis have 22 rays in the soft dorsal fin, and (iii) the genetic evidence restricted M. gracilis to southern-southeastern Brazil, with no evidence of gene flow with populations from other areas.

Following Chao (1978), we retained specimen MNHN 44 as the lectotype of M. gracilis in our present designation, with MNHN 9622 thus becoming a paralectotype of this species ( Fig. 6B ). We independently confirmed that paralectotype has 19 pectoral-fin rays.

Figures

FIGURE 4. The Atlantic coast of North, Central, and South America showing geographic distribution. A. Menticirrhus americanus (red) and M. martinicensis (green). B. M. littoralis (yellow), M. cuiaranensis (light blue) and M. gracilis (dark blue). Some symbols represent more than one locality or a large number of specimens.

FIGURE 6. Menticirrhus gracilis in lateral view. A. Lectotype, MNHN 44, Brazil. B. Paralectotype of M. gracilis, Brazil (MNHN 9622). C. Fresh specimen, São Paulo state, Brazil, photographed by M.M. Rotundo. Scale bar equals 10 mm.