Taxonomic review of the species of Mugil (Teleostei: Perciformes: Mugilidae) from the Atlantic South Caribbean and South America, with integration of morphological, cytogenetic and molecular data Author Menezes, Naércio A. Author Nirchio, Mauro Author Oliveira, Cláudio De Author Siccharamirez, Raquel text Zootaxa 2015 3918 1 1 38 journal article 10.11646/zootaxa.3918.1.1 c79e3d9f-7b3d-4ec0-bfd8-fc1d2dd1645a 1175-5326 287820 9F5CA16E-19A9-4BAF-B951-21E6396A85BF Mugil curema Valenciennes, 1836 ( Fig. 5 ) Mugil curema Valenciennes 1836: 87 ( Type locality: Bahia, Brazil ; lectotype : MNHN A-3683, designated by Harrison, 1993: 145; paralectotype : MNHN 3653 (1), Cuba ;?A- 4641(1, dry), Maracaibo; A-4655 (1, dry), Martinique ; A-4671 (1, dry), Martinique ; Ribeiro, 1915 : [unpaginated (Bahia)]; Blanc & Hureau, 1972 ( type catalog); Menezes, 1983 :3 (only specimens originating from the Caribbean region to southern Brazil ); Menezes & Figueiredo, 1985 :21 (only specimens originating from West Indies to southern Brazil ; Cervigón, 1992 : 363 (northern coast of South America ); Cervigón, 1993: 269 ( Venezuela , diagnosis, biological and ecological data); Thomson, 1997 : 487 (only specimens originating from Caribbean region to southern Brazil ); González-Bencomo et al ., 1997 : 159 ( Venezuela ); Aguilera, 1998 : 51 ( Venezuela ); Keith et al ., 2000 : 24 ( Guyana ); Marin, 2000 : 76 ( Venezuela ); Camargo & Isaac, 2001 : 41 (northern Brazil ); Lim et al ., 2002 : 70 ( Martinique ); Collette et al ., 2003 : 102 (West Indies); Nirchio et al ., 2003 : 113 ( Venezuela ); Harrison, 2003 : 1080 (only specimens originating from West Indies to Rio de Janeiro, Brazil ; size, habitat, biology and distribution); Menezes et al ., 2003 : 65 (only specimens originating from the Caribbean region to southern Brazil ); González-Castro et al ., 2006 : 91 ( Argentina ); Harrison, 2007: 457 ( Venezuela ); González-Castro et al ., 2012 : 1517 ( Argentina ). Material examined. MZUSP 67337, 46, SL 34–61 mm , Brazil , Pará: Marapanim, 0°42’S , 47°42’W ; MZUSP 67328, 4, SL 115–130 mm , Maranhão, Ilha São Luís, Rio Curuçá, approximately 2°31’S , 44°16’W ; MZUSP 48096, 5, SL 38–62 mm , Rio Grande do Norte, Natal , Ponta do Morcego, 5°47’56.4”S , 35°12’16.6”W ; LBP 4934, 1, SL 185 mm , Pernambuco: Tamandaré, 08°40’31” S , 35°06’S , 35°06’44”W ; MZUSP 42029, 1, SL 142 mm , Alagoas, Coqueiro Seco, Canal de Cadoz, 9°24’54.7”S , 35°31’16.5”W ; MZUSP 67334, 3, SL 139–155 mm , 67333, 5, SL 130–145 mm , 67396, 4, SL 115–207 mm , Maceió: Lagoa Mundaú, 9°38’28”S , 35°46’44”W ; MZUSP 67520, 2 SL 56.5 and 63 mm , Praia da Avenida, 9°35’26” S , 35°47’58”W ; MZUSP 115073, 3, 230– 267 mm , Barra de Santo Antônio 9°24’16.8”S , 35°31’47”W ; MZUSP 67400, 67398, 8, SL 110–250 mm , Sergipe: Rio Sergipe, 10°51’49.8”S , 37°01’53”W ; MZUSP 67382, 17, SL 45–131 mm , Aracaju, 10°51’49.8”S , 37°01’55”W ; MZUSP 51724, 3, SL 113–118 mm , Bahia: Nova Viçosa, Rio Peruípe, Porto de Nova Viçosa, 17°54’S , 39°22’W ; MZUSP 60776, 5, SL 19–84, Caravelas, 17°43’S , 39°14’W ; MZUSP 67373, 1, SL 55 mm , Salvador , Rio Vermelho, 13°00’40.5”S , 38°29’19.2”W ; MZUSP 67358, 1, SL 28.5 mm , Ilha de Itaparica, Rio Penha, 9°24’57.5”S , 40°31’12.8”W ; MZUSP 67359, 16, SL 31– 29 mm , Maragogipe, Barra do Paraguaçu, 12°45’S , 38°56’W ; MZUSP 67316, 102687 4, SL 118.5–200 mm , Ilhéus, 14°48’49.2”S , 39°02’0.7”W ; MZUSP 82166, 1, SL 145 mm , between Valença and Itacaré, approximately 13°59’S , 39°02’W ; MZUSP 67374, 18, 43–52 mm , Espírito Santo: Guarapari, Praia dos Namorados, 28, SL 31–98 mm , 20°33’11.5”S , 40°29’19.1”W ; MZUSP 673853, SL 55–121 mm , Santa Leopoldina, Rio Santa Maria da Vitória, 20°4’S , 40°44’W ; MZUSP 1330, 1, SL 220 mm , Linhares, Rio Doce, 20°11’S , 40°5’W ; MZUSP 67324, 4, SL 126–167, Lagoa Nova, 19°24’S , 40°1’W ; MZUSP 60348, 45, SL 76–244 mm , Rio de Janeiro: Ilha Grande, Baía da Ilha Grande, 23°09’07.5”S , 44°13’44”W ; MZUSP 29296, 28295, 27871, 4, SL 124–250 mm , Itaguaí, Foz do Rio da Guarda, 22°52’S , 43°46’W ; MZUSP 67340, 16, SL 20–282, Rio Paraíba do Sul, 21°48’S , 41°45’W ; MZUSP 44942, 1, SL 300 mm , Nova Iguaçu, Rio Guandu, 22°46’S , 43°26’W ; MZUSP 67391, 2, SL 130 and 190 mm , Cabo Frio, Lagoa Araruama, 22°52’22.6”S , 42°00’29.7”W ; MZUSP 67386, 2, SL 116 and 148 mm , Arraial do Cabo, 22°58’56.2”S , 42°01’14”W ; MZUSP 67329, 6, SL 65–104 mm , Atafona, Pontal 21°36’S , 41°01’W ; MZUSP 67355, 3, SL 50–87.5 mm , Ilha da Convivência, 21°36’S , 41°02’W ; MZUSP 67339, 9, SL 71–88 mm , Ilha do Lima, 21°42’S , 41°01’W ; MZUSP 67360, 2, SL 64 and 69 mm , São Fidélis, Córrego Pedra d’Água, 21°04’S , 41°45’W ; MZUSP 2359, 1195, 2, SL 193 and 224 mm , São Paulo: Santos, 23°57’14.5”S , 46°20’07”W ; MZUSP 108257, 1, SL 255 mm , Praia Grande, 24°01’40.8”S , 46°32’W ; MZUSP 58719, 1 SL 232 mm , Peruibe, Rio Itinguaçu, 24°18’44.8”S , 47°00’06”W ; LBP 10004, 25, SL 48–205 mm , Bertioga, 23°51’38”S , 46°09’10.5”W ; MZUSP 67390, 67395, 67397, 67325, 67326, 13, SL 164–264 mm , Cananéia, 25°00’53.7”S , 47°56’04”W , LBP 7573, 1, SL 220 mm , Cananéia, 26°06’34”S , 47°17.2”W ; MZUSP 115074, 10, SL 21–245 mm , Ilha do Cardoso, Cananéia, 25°07’54”S , 47°57’59”W ; MZUSP 67363, 67338, 13, SL 36–255 mm , São Vicente , 23°57’48.5”S , 46°29’14.6”W ; MZUSP 67331, 11, SL 77–103.5 mm , Caraguatatuba, 23°36’47”S , 45°24’46”W ; MZUSP 107459, 23, SL 28–97 mm , Ubatuba, núcleo Picinguaba, riacho Canto da Paciência, 23°22’48.6”S , 44°50’20.3”W ; MZUSP 67345, 16, SL 37–60 mm , Ubatuba, Praia do Itaguá, 23°27’20.3”S , 45°10.3’W ; MZUSP 67323, 1, SL 220 mm , Ubatuba, 23°22’17”S , 45°01’07”W ; MZUSP 67354, 7, SL 65–190 mm , Boca do rio da Lagoa, 23° 09’7.9”S , 46°42’6.3”W ; MZUSP 67392, 674201, SL 164 and 156 mm , São Sebastião, 23°47’44”S , 45°26’53”W ; MZUSP 13343, 1, SL 212 mm , Santa Catarina: Florianopólis, Rio Biguaçu, 27°29’S , 48°39’55”W ; MZUSP 14315-14316, 14318-14321, 14323-14324 14326, 67367, 67371, 67387, 27, SL 40–212 mm , Rio Grande do Sul: Tramandaí, 29°59’S , 50°07’W . Diagnosis . Mugil curema differs from congeners from the study area except M . brevirostris , M . incilis , M . margaritae , and M . rubrioculus in having an anal fin with 3 spines and 9–10 branched rays in adults or 2 spines, 1 unbranched and 9–10 branched rays in juvenile ( vs. 3 spines and 8 branched rays in adults or 2 spines, 1 unbranched and 8 branched rays in juveniles). Mugil curema is distinguished from M . brevirostris in having the pectoral fin not reaching of the vertical through the origin of the first dorsal-fin spine ( vs. reaching to, or extending slightly beyond, the vertical through the origin of the first dorsal-fin spine), and from M . incilis in having the origin of the first dorsal-fin spine midway between the snout tip and the caudal-fin base ( vs. origin of the first dorsal-fin spine much closer to the snout tip than the caudal-fin base) and 35–39 oblique scales from the dorsal limit of the pectoral-fin base to the caudal-fin base ( vs . 43–47 oblique scale rows from the dorsal limit of the pectoral-fin base to the caudal-fin base). Mugil curema differs from M . margaritae in having 35–39 oblique scales rows from the dorsal limit of the pectoral-fin base to the caudal-fin base ( vs. 40–44 oblique scale rows from the dorsal limit of the pectoral-fin base to the caudal-fin base) and from M . rubrioculus in having the anterodorsal portion of second dorsal fin uniformly dark or just slightly darker than remainder of fin, but not forming a spot ( vs. a conspicuous black spot on the anterodorsal portion of the second dorsal fin), and a conspicuous dark spot extending over most of the basal portion of the pectoral fin ( vs. basal portion of the pectoral fin overall dark or with small inconspicuous dark spot). Description . Morphometric data presented in Table 8 . Maximum examined body length 300 mm SL. Body elongate, compressed, deep compared to congeners. Greatest body depth at vertical through spinous dorsal-fin origin. Dorsal profile of head and body slightly convex along tip of snout, convex from vertical through anterior border of orbit to caudal peduncle, slightly concave along caudal peduncle. Ventral profile of head and body strongly convex from tip of lower jaw to caudal peduncle, straight to slightly concave along caudal peduncle, to vertical through caudal-fin base. Orbital diameter greater than snout length. Eye largely covered by adipose tissue,leaving only narrow oval-shaped central area uncovered in adults. Adipose tissue almost absent in specimens smaller than 30–35 mm SL. FIGURE 5. Mugil curema , MZUSP 114037. A—Detail of mouth showing dentition. B—Detail of scales located on ventral region of body above anal-fin origin. TABLE 8 . Morphometrics of Mugil curema . Standard length expressed in mm; measurements through head length are percentages of standard length; last eight entries are percentages of head length.

Characters	n range	mean	SD
Standard length	454 19.0–300.0	87.5
Body depth	446 22.6–035.6	28.3	2.3
Snout to dorsal-fin origin	454 70.6–081.6	76.0	1.8
Snout to pectoral-fin origin	454 25.0–035.1	30.0	2.0
Snout to pelvic-fin origin	454 35.3–045.6	41.4	1.8
Snout to anal-fin origin	454 66.4–076.0	72.2	1.8
Caudal peduncle depth	453 09.1–012.8	11.3	0.6
Caudal peduncle length	417 14.0–020.5	17.0	1.3
Pectoral-fin length	446 17.0–022.7	20.4	1.1
Pelvic-fin length	452 14.3–021.0	17.1	1.1
Head length	453 24.0–033.0	30.0	2.1
Head width	453 54.0–064.0	59.8	2.2
Head depth	452 62.8–074.0	69.6	2.3
Lip thickness	428 04.0–007.5	0 6.0	0.8
Mouth width	441 22.7–033.0	28.6	1.6
Mouth depth	440 20.0–028.0	24.2	1.1
Horizontal orbital diameter	454 23.6–033.5	30.0	2.3
Snout length	453 20.0–027.0	23.0	1.0
Upper jaw length	454 21.6–031.6	27.0	1.8

Anterior spinous dorsal fin with 4 slender spines connected by membrane, n = 454. Spinous dorsal fin-origin about midway between snout tip and caudal-fin base. Posterior soft dorsal-fin rays ii,7, n = 45 in specimens smaller than 30–35 mm SL; i, 8 in adults, n = 410. Unbranched pectoral-fin rays ii, first ray much smaller than second; branched rays 14–16, 14.8, n = 454. Tip of pectoral fins not reaching vertical through spinous dorsal-fin origin, but only to vertical slightly beyond pelvic-fin base. Pelvic fin with I,5. Tip of pelvic-fin reaching vertical through base of second dorsal-fin spine. Anal fin in specimens smaller than 30–35 mm SL II,i,9–10; III, 9–10 in adults, n =454. Mouth subterminal. Tip of maxilla extending slightly beyond vertical through anterior border of orbit. Teeth unicuspid, spatulate with slightly curved tips ( Fig. 5 A); external teeth on upper lip larger than inner teeth. Single row of close set, finer teeth on lower lip smaller than upper lip teeth. Scales spinoid; spines rudimentary on surface of scales, not projecting posteriorly along margin of scales ( Fig. 5 B). Transverse scale rows from dorsal limit of pectoral-fin base to caudal-fin base 34–39, 37.1, n = 399. Horizontal scale rows from spinous dorsal-fin origin to pelvic-fin origin 12–13, 12.6, n = 392. Horizontal scale rows around caudal peduncle 17–18, 17.7, n = 379. Soft dorsal and anal fins densely scaled except for narrow scaleless distal margin. Basal portion of pectoral fin fully covered by small scales extending between interradial membranes but not reaching distal margin of fin. Modified axilla scale dorsal to pectoral-fin base about 2.5 times as long as pectoral fin in 215 mm SL specimen (MZUSP 114037). Modified axilla scale dorsal to pelvic fin half length of pelvic fin in same specimen. Gill rakers close set; 20–75 rakers on ceratobranchial portion of first arch; rakers increasing in number ontogenetically ( Fig. 2 ) Color in alcohol . Body dark above with dark color fading ventrally midlateral region, whitish on abdominal region. Spinous dorsal, pelvic and anal fins pale with few scattered dark chromatophores. Soft dorsal fin dark with anterodorsal tip darker than remainder of fin. Pectoral and caudal fins profusely covered with dark chromatophores. Anterodorsal basal portion of pectoral fin with relatively long, dark spot covering basal portions of unbranched rays and 9–10 dorsalmost branched rays, vertically elongate whitish spot under it extending through bases of remaining ventralmost branched rays. Distal margin of caudal fin darker than remainder of fin. Cytogenetic and molecular data. Chromosome data of specimens from coastal waters of the Paranaguá Bay, Paraná State, Brazil show a karyotype with chromosome complement 2n=28, composed of 20M + 4ST + 4A ( Nirchio et al ., 2005 ) that corresponds to the caryotype originally described for the species from Louisiana, USA by Le Grande & Fitzsimons (1976) . C-banding in specimens from Brazil display only pericentrometric heterochromatic blocks on AG-NORs located on the telomeric region of the short arm of the subtelocentric chromosome pair number 11. Molecular analyses showed that the number of nucleotide differences between Mugil curema and the remaining analysed species ranges from 26.6 to 64.3 (16S) and 39.8 to 104.8 (COI) (Tables 2 to 5). The genetic distance between this species and the remaining analyzed species ranged from 0.057 to 0.130 (16S) and 0.064 to 0.186 (COI) ( Tables 6 and 7 ). The dendrogram in Figure 1 shows that this species is genetically most similar to M. margaritae . Distribution . Mugil curema is known throughout the study area ( Fig. 6 ), but specimens from the southern Caribbean, Uruguay and Argentina were unavailable for study. This species has been captured with M. brevirostris , M. curvidens , M. incilis and M. rubrioculus at many sites within the study area.