Menticirrhus americanus ( Linnaeus, 1758 )

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 : 315-317

Menticirrhus americanus ( Linnaeus, 1758)

Figure 1 , Tables 3, 4

Alburnus americanus ).—Catesby 1736: 2, plate 12 (the Whiting; fishes of Carolina, United States; short description and illustration)

Cyprinus americanus Linnaeus 1758: 321 . (Nec. Linnaeus, 1766 = Notemigonus bosei Cuvier & Valenciennes , America).

Perca alburnus . — Linnaeus 1766: 482 (South Carolina).

Centropomus alburnus .— Lacépêde 1802: 249, 257 (description; without locality).

Umbrina alburnus .—Cuvier in Cuvier & Valenciennes 1830: 180-186 (in part; description; New York; cited).

Umbrina phalaena .— Girard 1858: 167 (type locality, Texas; syntypes, MCZ 482 [ex USNM 620] (1); USNM 619 (1, not found in 1993), 620 (orig. 4, now 1), 128273 [ex USNM 620] (2), USNM 154721 [ex USNM 620])

Menticirrhus alburnus .— Jordan & Gilbert 1883: 577 (synopsis; listed)

Menticirrhus americanus ).— Jordan & Eigenmann 1889: 430-431 (description; Chesapeake Bay to Texas).— Jordan & Evermann 1898: 1474 (fishes of North and Middle America; description).— Briggs, 1958: 281 (in part; formerly assigned from New York to Argentina).— Trewavas 1964: 116 (American sciaenids with single mental barbel; definition of Menticirrhus , species list).— Irwin 1971: 69-74 (revision; in part; formerly from Long Island to Argentina; unpublished data).— Chao 1977: 28 (in part; FAO identification sheet; Cape Cod to Buenos Aires).— Hoese & Moore 1977: 203-204 (in part; formerly assigned from New York to Argentina).— Chao 1978: 30, 53 (in part; basis for classifying western Atlantic sciaenids; formerly assigned from North Atlantic to Argentina).— Robins & Ray 1986:187 (Atlantic coast fishes; listed and short description).— Boschung 1992:145 (catalog of the fishes of Alabama; listed).— Murdy et al. 1997:198 (fishes of Chesapeake Bay; listed).— Castro-Aguirre et al. 1999: 372 (ichthyofauna of Mexico; listed).— Chao 2003:1626 (in part; FAO Species Identification Guide of fishes of the Western Central Atlantic; short description).— Nelson et al. 2004:147 (common names of the fishes from the United States and Canada; listed).— McEachran & Fechhelm 2005:431 (fishes of the Gulf of Mexico; list and description).— Page et al. 2013: 152 (common and scientific names of fishes from the United States, Canada and Mexico; listed).— Raz-Guzmán et al. 2018: 346 (checklist; Laguna de Tamiahua, Mexico).— Robins et al. 2018: 436 (guide to the fishes from Florida; listed).

Material Examined. Neotype: USNM 196402 (1, 159 mm SL), Gulf of Mexico , Jackson County, Ocean Springs 3 miles north of Horn Island , Mississippi . Non-type specimens: ANSP 168780 (1), Savannah River Estuary, Jasper/ Chatham Co., Georgia / South Carolina, United States ; ANSP 168781 (1), Savannah River Estuary, Jasper/ Chatham Co., Georgia / South Carolina, United States ; ANSP 168782 (1), Savannah River Estuary, Jasper/ Chatham Co., Georgia / South Carolina, United States ; ANSP 168783 (2), Savannah River Estuary, Jasper/ Chatham Co., Georgia / South Carolina, United States ; ANSP 168784 (2), Savannah River Estuary, Jasper/ Chatham Co., Georgia / South Carolina, United States ; ANSP 79853 (1), Gulf bay at Englewood, Florida, United States ; USNM 158610 (1), bay at Clearwater , Florida, United States ; MZUSP 69614 (2, 127- 140 mm SL), off Matanzas near St. Augustine, Florida, United States , USNM 22832 (1), Pensacola , Florida, United States ; USNM 93594 (1), off Corpus Christi , Texas, United States ; USNM 451060 (1, 154 mm SL) Gulf of Mexico , Jackson County, Ocean Springs 3 miles north of Horn Island, Mississippi ; ANSP 70907 (2), Galveston , Texas, United States ; USNM 155969 (2), New Orleans , Louisiana, United States ; USNM 155972 (1), New Orleans , Louisiana, United States ; ANSP 151465 (1), Puerto Rico, United States .

Diagnosis. Menticirrhus americanus , which is found in the United States and the Gulf of Mexico, can be differentiated from its congeners in the western Atlantic as follows: from M. cuiaranensis , which is found on the northern, northeast and eastern coasts of Brazil, by having 19–23, rarely 19, pectoral-fin rays (vs. 16–19, Table 2), pectoral-fin tip surpassing tip of depressed pelvic fin (vs. barely reaching tip of pelvic fin, Figs. 1 , 3 ), body with irregular dark bars (vs. without dark bars, Figs. 1 , 3 ); from M. gracilis , which is found on the southeastern and southern coast of Brazil, by having 23–25 dorsal-fin rays (vs. 18–21, Table 2), and body with irregular dark bars (vs. without dark bars, Figs. 1 , 4 ); from M. littoralis , which is found in the United States and the Gulf of Mexico, by a dusky caudal fin, with no distinctive dark spot on the dorsal lobe (vs. pale caudal fin, with a dark spot on the upper lobe, Figs. 1 , 2 ), pectoral-fin tip surpassing tip of depressed pelvic fin (vs. barely reaching tip of pelvic fin, Figs. 1 , 2 ), body with irregular dark bars (vs. without dark bars, Figs. 1 , 2 ), and from M. saxatilis , which is found in the United States and the Gulf of Mexico, by having body usually with eight or nine diffuse dark bars, the second and third forming a “V” shape below the nape and spinous dorsal fin (vs. body with seven or eight distinct oblique bars, with the second and third bars forming a “V” shape below the spinous dorsal fin), no longitudinal stripe below lateral line (vs. stripe present, extending to the tip of the caudal fin), spinous dorsal fin low, not reaching to base of the second soft ray (vs. spinous dorsal fin high, its tip reaching beyond the base of the fourth soft ray of the dorsal fin, Fig. 1 ).

The morphology of this species is virtually identical to M. martinicensis , which occurs from the Caribbean to South America, except that many M. americanus specimens present a small hump in the dorsal profile behind head ( Fig. 1 ), whereas the dorsal profile of M. martinicensis is always steep and gently convex in this region ( Fig. 5 ). These taxa are also separated by the molecular evidence (see Molecular Diagnosis).

Menticirrhus americanus 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 its body with irregular dark bars (vs. without dark bars, Fig. 1 ); from M. nasus , ranging from the Gulf of California to Peru, by having 10–11 gill rakers in the first arches (vs. 4–9, Table 2), scales present along base of soft dorsal fin (vs. soft dorsal fin naked), body with irregular dark bars (vs. without dark bars, Fig. 1 ); from M. ophicephalus , ranging from Ecuador to Chile, by 48–52 scales with pores along lateral line to caudal-fin base (vs. 63–66, Table 2), pectoral-fin tip surpassing tip of depressed pelvic fin (vs. barely reaching tip of pelvic fin, Fig. 1 ), S-shaped posterior margin of caudal fin (vs. concave), scales present along base of soft dorsal fin (vs. soft dorsal fin naked), body with irregular dark bars (vs. without dark bars, Fig. 1 ); from M. paitensis , ranging from the Gulf of California to Chile, by having 48–52 scales with pores along lateral line to caudal-fin base (vs. 76–98, Table 2), scales present along base of soft dorsal fin (vs. soft dorsal fin naked), body with irregular dark bars (vs. without dark bars, Fig. 1 ); from M. panamensis , ranging from the Gulf of California to Chile, by having 23–25 dorsal-fin rays (vs. 18–22, Table 2), 10–11 gill rakers in the first arches (vs. 3–8, Table 2), scales present along base of soft dorsal fin (vs. soft dorsal fin naked), body with irregular dark bars (vs. without dark bars, Fig. 1 ); from M. undulatus , which is found in California, in the United States, by having 20–23, rarely 19, pectoral-fin rays (vs. 17–19, Table 2), scales present along base of soft dorsal fin (vs. soft dorsal fin naked), pectoral-fin tip surpassing tip of depressed pelvic fin (vs. barely reaching tip of pelvic fin, Fig. 1 ).

Description: D. X+23–25; A. I+6–8; P. 19–22; C. 17; GR. 11; LL. 50–52; SA. 8–9; SB. 12–13 ( Tables 3, 4). Body slender in lateral view, not compressed, ventral profile flat; maximum depth at first dorsal-fin origin. Dorsal profile steeply ascending on snout to eye-level, convex over eye, ascending to first dorsal-fin base, frequently with “hump” over pectoral-fin base, convex along first dorsal-fin base, straight, descending along second dorsal-fin base, concave on caudal peduncle. Ventral profile descending from mouth to gular area, sometimes with slight concavity, convex in gular area, horizontal on belly, ascending, nearly concave, along anal-fin base to caudal-fin base. Snout much longer than eye, deep, with blunt tip. Mouth subterminal, upper jaw slender, rear tip barely reaching vertical line passing through middle of eye. Premaxilla with 4–7 rows of acicular teeth, those of external row (about 10–15 on each side) slightly larger, caniniform; dentary with 4–7 tooth rows. Eye slightly oval, nearly two times snout length, not adjoining dorsal profile, pupil slightly elongated dorsoventrally, teardrop shaped. Interorbital space smaller than orbital diameter, slightly convex, covered with ctenoid scales. Nostrils small, anterior nearly round, posterior larger, oval, nearly on horizontal line through ventral pupil border. Head with five sensory pores on snout, in semicircle around mouth corner, 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, straight, descending to caudal peduncle, nearly horizontal elsewhere to caudal-fin tip. Preopercle margin rough, with about 10 subtle spines. Opercle extremity fleshy, blunt, well behind vertical line that passes through pectoral-fin base. Gill rakers short. Ctenoid scales on trunk, belly, pectoral-fin base, opercle, preopercle, infraorbital, and interorbital region to snout, cycloid on gular area and encircling nostrils. Dorsal fin without scales on interradial membranes, second dorsal with basal scale on each interradial membranes; pectoral-fin base covered by small ctenoid scales, rows of cycloid scales along pectoral-fin rays (outer third 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 pectoral-fin tip, second dorsal soft rays much shorter than longest dorsal spines.Anal-fin origin at vertical line passing through sixth-seventh ray of second dorsal fin; first spine very slender. Pectoral fin falcate, reaching vertical line passing through two thirds of pelvic fin length. Pelvic-fin origin behind pectoral-fin base, as long as pectoral fin. Caudal peduncle short, deep, depth larger than orbital diameter. Caudal fin short, emarginated, lower lobe slightly longer and much stouter than upper lobe.

Color in alcohol. Trunk beige or light brown, darker on dorsum, sometimes with dark patches on sides. Snout light brown, opercle and infraorbital region dark brown. First dorsal fin dark brown, dusky in distal third, second dorsal fin light brown, some ray tips lighter or beige. Anal fin dark brown, slightly darkened at tip. Pectoral fin brown, distal half darkened. Caudal fin darkened at the base ( Fig. 1B ).

Color of fresh material. Silvery on head and flanks, grayish silvery dorsally; obscure dark areas on flanks and dorsum, often with irregular bars over the lateral line; nape with dusky bar; opercle and gill opening dusky. Dorsal and anal fins dusky, pectoral fin black with yellow hue on lower rays, lighter at base; pelvic fin yellow, with black line along pelvic spine. Dusky line on the ventral margin to the caudal peduncle ( Fig. 1A ).

Distribution and habitat. Western North Atlantic from the east coast of the United States to the Gulf of Mexico ( Fig. 4 ). This is a coastal species, typically found in estuaries at depths of up to 60 meters.

Fishing. Targeted by artisanal fishermen and using castnets and fishing weir. Taken as food in some areas.

Remarks. Linnaeus (1758) proposed Cyprinus americanus based on the figure and short account of Catesby (1743) in his description of the fish fauna of the “Carolinas”. Although the image presents certain inaccuracies in fin counts and other features, it clearly represents a Menticirrhus species, and based on Linnaeus’ description, it seems reasonable to conclude that this species is M. americanus from the western North Atlantic. Linnaeus (1766) provided additional information on C. americanus , but Wheeler (1985) concluded that this is different from the data provided by Catesby.

In their review of the American sciaenids, Jordan & Eigenmann (1889) indicated that Menticirrhus americanus was distributed along the east coast of the United States, from Chesapeake Bay to Texas. They also synonymized two species with M. americanus , Perca alburnus Linnaeus 1886 , based on the specimens Linnaeus received from Charleston, and Umbrina phalaena Girard 1858 , based on the description of specimens obtained in Mexico. Jordan and Eigenmann also remarked that M. martinicensis , described originally from the Caribbean by Cuvier and Valenciennes (see notes on this species below) is remarkably similar to M. americanus . Jordan & Evermann (1898) provided a short description of M. americanus and regarded this species to be “exceedingly alike [ M. martinicensis ] distinguished perhaps by a slightly smaller number of rays in the dorsal fin” and that “Were it not that the Sciaenoid fauna of South America is chiefly different from that of North America, we should scarcely hesitate to place martinicensis in the synonymy of americanus .” The close similarity between these two taxa led Miranda Ribeiro (1915) to assume that Menticirrhus americanus is present in Brazil and included M. martinicensis as its junior synonym. Although some contemporary authors still recognized the validity of M. jamaicensis (e.g., Ihering 1897; Jordan & Evermann 1898; Gilbert 1900; Fowler 1941), the decision of Miranda Ribeiro (1915) was followed in most subsequent studies (e.g. Hoese & Moore 1977; Chao 1977, 1978).

Here we consider M. americanus to be restricted to the Gulf of Mexico, based primarily on the genetic evidence, given that the meristic data broadly overlap with those of M. martinicensis from the Caribbean region and the western South Atlantic. One other trait that may be potentially useful for separation of these taxa is the tendency for the M. americanus specimens to have a small “hump” in the dorsal head profile, although this needs to be considered with caution, given that this feature is not depicted, for example, in McEachran and Fechhelm (2005).

Menticirrhus americanus was originally described without support of biological material and, until now, no type specimens had been allocated. For this reason, we selected specimen USNM 196402, 159 mm SL, collected east of the Cape Hatteras National Seashore , North Carolina, in the United States, as the neotype of this species .

Figures

FIGURE 1. Menticirrhus americanus in lateral view. A. Fresh specimen, Gulf of Mexico, photographed by Ross Robertson. B. Neotype, USNM 196402, 159 mm SL, Horn Island, Mississippi, United States.

FIGURE 2. Menticirrhus littoralis in lateral view. A. Holotype, ANSP 11576, 195 mm SL, South Carolina, United States. B. Live specimen captured in the Gulf of Mexico by Luiz Malabarba. C. Fresh specimen, Mexico, photographed by Eduardo Perusquia. Scale bar equals 10 mm.

FIGURE 3. Menticirrhus martinicensis in lateral view. A. Holotype, MNHN 7498, Martinique Island. B. Fresh specimen, Amapá state, Brazil, photographed by A. P. Marceniuk. Scale bar equals 10 mm.

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 5. Menticirrhus cuiaranensis in lateral view.A. Holotype, MPEG 38944, 222 mm SL, Cuiarana, Pará, Brazil. B. Fresh specimen, Maracanã, Pará, Brazil, photographed by A. P. Marceniuk. C. Type-locality of M. cuiaranensis, an artisanal fishermen village in Salinópolis, Pará state, at the time of the holotype collection.