Menticirrhus martinicensis ( 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 : 321-325

Menticirrhus martinicensis ( Cuvier, 1830)

Figure 3 , Tables 3, 4

Umbrina martinicensis Cuvier in Cuvier & Valenciennes 1830: 186 (original description; type locality: Martinique Island. Syntypes: MNHN 7498 (1)).— Günther, 1860: 277 (catalog of fishes from the British Museum; Martinique; short description).— Bauchot & Desoutter 1987:20 (catalog of types of MNHN).— González Bencomo et al. 1997:159 (Ichthyofauna of San Carlos, Maracaibo Lake, Venezuela; listed).

Umbrina januaria Steindachner 1876:170 (type locality: Rio de Janeiro, Brazil. Type material not yet found at NMW).— Jordan & Eigenmann 1899: 429 (synonymy with Menticirrhus martinicensis ).— Jordan & Evermann 1896: 1474 (fishes of North America; synonymy with Menticirrhus martinicensis ).

Menticirrhus martinicensis .— Jordan 1887: 539 (type specimens of the species described by Cuvier & Valenciennes; two specimens from Martinique; short description).— Jordan & Eigenmann 1889: 429 (review of American Sciaenidae ; West Indies to Patagonia).—Berg 1895: 56 ( Argentina; listed).— Jordan & Evermann 1896: 1474 (fishes of North America; description; comparison with Menticirrhus americanus ).— Ihering 1897: 40 (fishes of Rio Grande do Sul, Brazil; listed).— Gilbert 1900: 172 (results of Branner-Agassiz expedition; Maceió, Brazil; listed).— Devincenzi 1924: 240 ( Uruguay; listed).—Gliesch 1925: 33 (fish fauna; Torres, Rio Grande do Sul, Brazil; listed).— Fowler 1941:168 (fishes of Brazil; listed).— Chiesa 1945: 73 (fishes of Buenos Aires; listed).— Buen 1950: 115 (fish fauna of Uruguay; listed); Santos 1952: 149 (marine fishes of Brazil; listed).—Pozzi & Bordale 1953: 168 (systematic summary of Argentinean fishes; listed).— Travassos & Paiva 1957: 141 (Brazilian sciaenids; identification key; listed).— Ringuelet & Aramburu 1960:66 (Argentinean fishes; list and description).— Barcellos 1962: 12 (fishes soled in Rio Grande do Sul, Brazil; listed).— Travassos & Rêgo-Barros 1971: 66 (Brazilian sciaenids; listed).

Menticirrhus americanus (not of Linnaeus 1758).—Miranda Ribeiro 1915: 422 (Fauna Brasiliense; description; synonymy).— Carvalho 1941:65-66 (fish fauna; São Paulo, Brazil; listed).— Carvalho & Ramos 1941: 24 (fish fauna of Rio Ribeira de Iguape, Brazil; listed).— Vasconcelos 1945:115 (Brazilian sciaenids; listed).— Fowler,1951: 26 (fish collected during Wilkes Expedition; Rio de Janeiro, Brazil; listed).— Travassos & Paiva 1957: 141 (Brazilian sciaenids; identification key; listed).— Franco 1959: 59 (diet of Brazilian fishes; listed).— Nomura & Menezes, 1964: 369 (marine fishes; listed).—Iheing 1968: 502 (Dictionary of Animals; listed).— Menezes 1969: 55 (expeditions to Torres and Chuí, Rio Grande do Sul, Brazil; listed).— Vazzoler 1969: 14 (fish fauna of Santos, São Paulo, Brazil; listed).— Irwin 1971: 69-74 (in part; unpublished thesis on Menticirrhus ).— Travassos & Rêgo-Barros 1971: 66 (Brazilian sciaenids; listed).— Vazzoler & Motonaga 1971: 56 (exploratory fishing; Rio Grande do Sul, Brazil; listed).— Jardim 1973: 20-21 (Brazilian sciaenids; listed).— Benvegnú 1973: 496 ( Brazil).— 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 of the fishes from southeastern Brazilian coast; short description).— Gomes et al. 1983: 187-191 (Cananeia, São Paulo, Brazil; karyotype).—Uyeno & Sato in Uyeno et al. 1983: 366 (fishes trawled off Suriname and French Guyana; short description, image).—Casatti & Menezes in Menezes et al. 2003: 87 (in part; catalog of marine fishes of Brazil; listed).

Doubtful references to this species

Umbrina arenata Cuvier in Cuvier & Valenciennes 1830: 190 (type locality: Brazil. Syntypes: Syntypes: MNHN 7472 (1, dry), 7500 (1)).— Jordan & Eigenmann 1899: 429 (synonymy with Menticirrhus martinicensis ).— Jordan & Evermann, 1896: 1474 (fishes of North America; synonymy with Menticirrhus martinicensis ).— Bauchot & Desoutter 1987:20 (catalog of types of MNHN).

Material examined. Holotype: MNHN 7498 , Martinique Island, West Indies . Non-type specimens: USNM 300466 (3, 152– 174 mm SL), 2.5 miles east of Dangriga , Belize ; USNM 389992 (1), Bocas del Toro , Laguna de Chiriqui, Panama ; MPEG 35218 (1, 254 mm SL), 01° 03’ N 47° 49’ W, Amapá , Brazil ; MPEG 35090 (1, 301 mm SL), 01° 13’ N 48° 06’ W, Amapá, Brazil ; MPEG 35100 (1, 269 mm SL), 01° 15’ N 48° 00’ W, Amapá, Brazil ; MPEG 35206 (1, 258 mm SL), 01° 31’ N 48° 03’ W, Amapá, Brazil ; AZUSC 5499 (1, 270 mm SL), continental shelf, Amapá, Brazil ; AZUSC 5302 (2, 167– 305 mm SL), continental shelf, Pará , Brazil ; MPEG 35093 (2, 232– 249 mm SL), 00° 59’ N 47° 49’ W, Pará , Brazil ; MPEG 35241 (4, 123– 132 mm SL), Ajuruteua , Bragança, Pará, Brazil ; MPEG 34419 (1, 276 mm SL), Ajuruteua , Bragança, Pará, Brazil ; MPEG 34513 (1, 232 mm SL), Bragança , Pará, Brazil ; MPEG 34527 (3, 284– 287 mm SL), Bragança , Pará, Brazil ; MPEG 32957 (1, 175 mm SL), Furo da Ostra , Bragança, Pará, Brazil ; MPEG 34649 (8), Canárias , Piauí, Brazil ; LBP 20079 (2, 175– 176 mm SL), Fortaleza , Ceará, Brazil ; LBP 20084 (4, 152– 175 mm SL), Fortaleza , Ceará, Brazil ; MPEG 34285 (3, 126– 259 mm SL), Barra de Santo Antônio , Alagoas, Brazil ; MPEG 34245 (1, 199 mm SL), Paripueira , Alagoas, Brazil ; MPEG 34339 (1, 203 mm SL), Praia de Jatiúca , Maceió, Alagoas, Brazil ; MPEG 34352 (2, 132- 167 mm SL), Jaraguá , Maceió, Alagoas, Brazil ; MPEG 34362 (4, 119– 141 mm SL), Jaraguá , Maceió, Alagoas, Brazil ; MPEG 34398 (2, 124 mm SL), Jaraguá , Maceió, Alagoas, Brazil ; AZUSC 5934 (10, 129– 165 mm SL), mouth of the Rio Sergipe, Aracajú, Sergipe, Brazil ; MZUSP 7956 (1, 204 mm SL), Aracajú , Sergipe, Brazil ; LBP 22929 (3, 149– 193 mm SL), Porto Seguro , Bahia, Brazil ; MZUSP 82215 (4, 118– 210 mm SL), Bahia, Brazil ; MZUSP 61326 (1, 111 mm SL), Rio Caravelas , Caravelas, Bahia, Brazil ; LBP 24080 (1, 169 mm SL), Guarapari , Espírito Santo, Brazil ; AZUSC 4549 (8, 172– 214 mm SL), Vitória , Espirito Santo ; LBP 10544 (3, 132– 146 mm SL), Macaé , Rio de Janeiro, Brazil ; LBP 21416 (2, 154– 204 mm SL), Rio de Janeiro, Rio de Janeiro, Brazil ; MZUSP 7958 (1, 189 mm SL), Ubatuba , São Paulo, Brazil ; AZUSC 421 (1, 189 mm SL), Bertioga , São Paulo, Brazil ; LBP 10073 (1, 145 mm SL), Bertioga Channel , Bertioga, São Paulo, Brazil ; MZUSP 47525 (1, 118 mm SL), Guarujá , São Paulo, Brazil ; MZUSP 69620 (3, 156– 189 mm SL), Cananeia , São Paulo, Brazil ; LBP 20752 (1, 170 mm SL), Peruibe , São Paulo, Brazil ; AZUSC 1150 (4, 176– 276 mm SL), Florianópolis , Santa Catarina, Brazil ; MZUSP 69617 (1, 140 mm SL), 31°15’S 50°07’ W, Rio Grande do Sul, Brazil ; MZUSP 69671 (1, 102 mm SL), 31°45’S 49°58’ W, Rio Grande do Sul, Brazil ; MZUSP 69674 (4, 136– 154 mm SL), 32°06’S 51°49’ W, Rio Grande do Sul, Brazil ; MZUSP 69796 (3, 170– 184 mm SL), 19°08’S 39°58’ W, Espírito Santo, Brazil ; MZUSP 69799 (1, 127 mm SL), 28°07’S 48°09’ W, Santa Catarina, Brazil .

Diagnosis. Menticirrhus martinicensis , which is found in the Caribbean and Atlantic South America, 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 19–24, rarely 19, pectoral-fin rays (vs. 16–19, Table 3), pectoralfin tip surpassing tip of depressed pelvic fin (vs. barely reaching tip of pelvic fin, Fig. 5 ), body with irregular dark bars (vs. without dark bars, Fig. 5 ); from M. gracilis , which is found on the southern and southeastern coasts of Brazil, by having 22–25 dorsal-fin rays (vs. 18–21, Table 3), body with irregular dark bars (vs. without dark bars, Fig. 5 ); from M. littoralis , which is found in the United States and the Gulf of Mexico, by a dusky caudal fin, without a distinctive dark spot on the dorsal lobe (vs. pale caudal fin, with a dark spot on the upper lobe, Fig. 5 ), pectoral-fin tip surpassing tip of depressed pelvic fin (vs. barely reaching tip of pelvic fin, Fig. 5 ), body with irregular dark bars (vs. without dark bars, Fig. 5 ), and from M. saxatilis , ranging from the United States to the Gulf of Mexico, by body with eight or nine diffused bars, with the second and third bars forming a faint V-shape below the nape and spinous dorsal fin (vs. body with seven or eight distinct oblique bars, second and third bars forming a V-shape below the spinous dorsal fin, and a longitudinal stripe below the lateral line, extending to the tip of the caudal fin, Fig. 5 ), spinous dorsal-fin lower, not reaching to base of the second soft ray (vs. spinous dorsal fin high, its tip reaching beyond the base of the fourth soft dorsal-fin ray).

This species is virtually identical, morphologically, to M. americanus , which occurs from the western Atlantic, United States and Gulf of Mexico, except that many of the M. americanus specimens have a small hump in the dorsal profile ( Fig. 1 ), behind head, whereas the dorsal profile of M. martinicensis is always steep and gently convex in this area ( Fig. 5 ). These taxa were also differentiated conclusively by the molecular evidence (see Molecular Diagnosis).

Menticirrhus martinicensis can also be distinguished from its congeners in the eastern Pacific as follows: from M. elongatus , which occurs from the Gulf of California to Peru, by body with irregular dark bars (vs. without dark bars, Fig. 5 ); from M. nasus , which occurs from the Gulf of California to Peru, by having scales present along base of soft dorsal fin (vs. soft dorsal fin naked), body with irregular dark bars (vs. without dark bars, Fig. 5 ); from M. ophicephalus , ranging from Ecuador to Chile, by 47–58 scales with pores along lateral line to caudal-fin base (vs. 63–66), pectoral-fin tip surpassing tip of depressed pelvic fin (vs. barely reaching tip of pelvic fin, Fig. 5 ), Sshaped posterior margin of the 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. 5 ); from M. paitensis , ranging from the Gulf of California to Chile, by having 47–58 scales with pores along lateral line to caudal-fin base (vs. 76–98), scales present along base of soft dorsal fin (vs. soft dorsal fin naked), body with irregular dark bars (vs. without dark bars, Fig. 5 ); from M. panamensis , ranging from the Gulf of California to Chile, by having 23–25, rarely 22, dorsal-fin rays (vs. 18–22, Table 3), scales present along base of soft dorsal fin (vs. soft dorsal fin naked), body with irregular dark bars (vs. without dark bars, Fig. 5 ); from M. undulatus , which is found in California, United States, by having 20–24, rarely 19, pectoral-fin rays (vs. 17–19, Table 3), 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. 5 ).

Description: D. IX-XII+22–25; A. I+6–8; P. 19–23; C. 17; GR. 7–15; LL. 48–56; SA. 7–11; SB. 12–18 ( Tables 3, 4). Body slender in lateral view, not compressed, ventral profile flat; maximum depth at first dorsal fin origin. Dorsal profile convex at snout, nearly straight, ascending over eye-level, slightly convex to occipital region, convex to first dorsal fin, slightly convex from first dorsal-fin base to second dorsal fin, straight, descending to second dorsal-fin base, concave on caudal peduncle. Ventral profile nearly horizontal from mouth to gular region, convex to pelvic-fin origin, concave from pelvic fin base to vent, ascending, slightly concave at anal-fin base, nearly straight on caudal peduncle. Head moderately large, deep, stocky. Snout moderately long, very deep, with blunt tip. Mouth subterminal, upper jaw slender, rear tip on vertical line slightly behind vertical line passing through middle of eye. Teeth conical. Eye nearly 1.5 times snout, slightly oval, almost adjoining dorsal profile. Interorbital space smaller than orbital diameter, slightly convex, covered with ctenoid scales. Nostrils small, close together, anterior oval, posterior larger, slit-like, below horizontal line through border of ventral pupil. Lateral sensory canal on head visible in infraorbital, dentary, and preopercle; lower jaw with four pores, two on each side encircling mental barbel; mental barbel short, rigid, with blunt tip and no pore. Lateral line quite arched to second dorsal-fin origin, straight elsewhere to caudal-fin tip. Preopercle margin rough to serrated, with about 8 spines. Opercle tip blunt, not very evident, on vertical line that passes through pectoral-fin base. Gill rakers short, two or three in upper limb rudimentary. Scales present on trunk, belly, pectoral-fin base, opercle, preopercle, infraorbital to middle of eye (naked elsewhere), and interorbital region to snout; opercle with about eight vertical rows of scales. First dorsal fin without scales, membranes of second dorsal fin and anal fin with one or two series of 5–7 small cycloid scales. Pectoral fin unscaled in smaller specimens, with cycloid scales on basal fourth in larger ones. Caudal-fin base covered with large cycloid scales to one third its length. Spinous dorsal fin short, first spine shortest, second and third spines longest; notch between first and second dorsal fin. Origin of second dorsal fin behind vertical line through pectoral-fin tip, second dorsal soft rays much shorter than longest first dorsal spines. Anal-fin origin on vertical line passing through sixth-seventh ray of second dorsal fin; first spine very slender. Pectoral fin large, wide, reaching vertical line passing through last first dorsal spine and pelvic-fin tip. Pelvic fin origin slightly behind pectoral-fin base, shorter than pectoral fin. Caudal peduncle moderately long, deep, depth larger than orbital diameter. Caudal fin short, emarginated, rays on upper lobe longer than those on lower lobe.

Color in alcohol. Background color light brown, darker on head, flanks sometimes with subtle dark brown blotches. Most fins dark brown, longest dorsal spines with black tips; second dorsal fin yellow, lighter along base; pectoral fin with extensive dark hue on lowest rays; pelvic fin yellow, with dense dark hue along pelvic spine. Caudal fin lighter to tip ( Fig. 3B ).

Color of fresh material. Head dark brown, flanks gray with dark brown blotches dorsally, typically forming irregular oblique streaks. First dorsal fin dusky, second dorsal fin gray with two dark areas along base; anal fin dark, pectoral fin black, caudal fin rusty gray, with dark central area, more so at the tip of median rays ( Fig. 3B ).

Distribution and habitat. Western Atlantic from the Caribbean region ( Martinique and the West Indies) to Argentina ( Fig. 4 ). Commonly found in coastal waters at depths of up to 60 meters, over muddy or sandy bottoms.

Fishing. Targeted by artisanal fisheries and taken as bycatch in otter trawls and by other industrial fisheries.

Remarks. Cuvier ( Cuvier & Valenciennes, 1830) described Umbrina martinicensis based on a single specimen (MNHN 7498, Fig. 5A ), collected from Martinique Island, in the West Indies. In their identification key for Brazilian sciaenids, Travassos & Paiva (1957) considered both Menticirrhus americanus and M. martinicensis to be valid species, contradicting the position of most contemporary authors, but supported their arrangement by showing that the two species can be differentiated by a number of features, such as the second dorsal-fin counts (24–25 rays in M. americanus vs. 22–23 in M. martinicensis ), the shape of the mental barbel (the tip is rounded in M. americanus and pointed in M. martinicensis ), and orbital diameter (greater in M. americanus ). Travassos & Rêgo-Barros 1971, probably based on Travassos’ earlier work (and probably also unaware of Irwin’s work), upheld the validity of the two taxa, but assumed that they occur in practically the same geographic area ( M. americanus from Chesapeake Bay to Patagonia and M. martinicensis from the West Indies to Patagonia). Many of the differences identified by Travassos & Paiva (1957) were not recovered in the present study. The meristic data on M. americanus and M. martinicensis broadly overlap, and the orbital diameter also varies with size.

Umbrina arenata , which was also described by Cuvier (1830), was proposed based on specimens brought by Delalande from Brazil ( Fig. 8A ). Although meristic data give in the original description for this taxon are consistent with those of M. cuiaranensis in some features, such as the pectoral (18 rays) and second dorsal-fin counts (23–24 rays, based on Cuvier’s original description), the coloration is more similar to that observed in M. americanus and M. martinicensis , with six oblique dark bands dorsally, and the length of the pectoral fin, which extends beyond a vertical line passing through the tip of the pelvic fin. Jenyns (1842) provided further information on the specimens collected by Charles Darwin in Bahia Blanca and Maldonado ( Argentina), which he identified as U. arenata . However, these specimens have more pectoral-fin rays (21) than described in Cuvier’s original description. Günther (1860) also presented data he attributed to U. arenata , based on two specimens he obtained from the Gulf of Mexico and Jamaica, which had 22–25 rays in the second dorsal fin, and 73–78 scales in the lateral line. This species was later synonymyzed with Menticirrhus martinicensis by Jordan & Eigenmann (1889), without details, and this decision was subsequently substantiated in Jordan & Evermann (1898). The authors concluded that the values provided by Günther on the number of scales in the lateral line were probably wrong, and concluded “We see no reason, therefore, for not placing arenata in the synonymy of martinicensis .”

We also examined images of two syntypes of U. arenata: MNHN 7472 ( Fig. 8B ), which is a poorly-preserved stuffed specimen, and MNHN 7500 ( Fig. 8A ), which is quite small and in a poor condition. We nevertheless noted that MNHN 7472 has 21 rays in the pectoral-fin rays and that the pectoral tip reaches well beyond the tip of the pelvic fin, whereas MNHN 7500 appeared to have 18 rays (counts were very difficult to determine in this specimen), and the pectoral-fin tip does not reach the tip of the pelvic fin. Regardless of the possible influence of the size and condition of these specimens, the sum of the evidence indicates that these Menticirrhus arenata syntypes may represent other Menticirrhus species. We tentatively identify specimen MNHN 7472 as M. martinicensis , although we were unable to identify MNHN 7590 conclusively.

Given the substantial differences in data on U. arenata by the different authors, the overall poor condition of the type material, and the fact that we cannot confidently determine which of the known syntypes was actually depicted in Cuvier’s original description, we declare U. arenata to be a nomen dubium.

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 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.

FIGURE 8. Syntypes of Umbrina arenata in lateral view. A. MNHN 7500, Brazil. B. MNHN 7472, Brazil. Scale bar equals 10 mm.