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 brevirostris ( Ribeiro, 1915 ) ( Fig. 1 ) Querimana brevirostris Ribeiro, 1915 (unpaginated), Brazil [ type locality: Brazil (no precise locality)]; holotype (probably lost); Thomson, 1997 : 494 (as a probable synonym of Mugil trichodon Poey, 1835 ). Mugil hospes (not of Jordan & Culver, 1895), Thomson, 1997 : 490 (only specimens originating from Rio de Janeiro, Brazil ); Harrison, 2003 : 1082 (only specimens originating from Brazilian coast). Mugil gaimardianus (not of Desmarest, 1831: 129), Menezes, 1983 : 4 (only specimens originating from Brazilian coast); 1985: 22 (only specimens originating from Maceió, Alagoas, Cananéia, São Paulo and Lagoa dos Patos, Rio Grande do Sul, Brazil ). Mugil sp., Menezes et al ., 2003 : 65 (only specimens originating from Brazilian coast). Material examined. Neotype—MZUSP 115087, SL 106 mm , Cananéia, São Paulo, 25°00’45”S , 47°56’07”W . Other material. MZUSP 30333, 31446, 4, SL 71–101.5 mm , Brazil , Amapá: approximately 1°26’N , 52°01’W ; MZUSP 77667, 20, SL 24.5–52 mm , Pará: Salvaterra, Praia do Jubim, 0°45’31.6”S , 48°30’51.6”W ; MZUSP 67402-03, 3, SL 135–156 mm , Maranhão: Ilha de São Luís, Rio Curuçá, approximately 2°31’S , 44°16’W ; MZUSP 67409, 67408, 67404, 4, SL 142–171 mm , Alagoas: Lagoa Mundaú, 9°38’28”S , 35°46’44”W , MZUSP 51159, 2, SL 144 and 183 mm , Coqueiro Seco, 09°38’14”S , 35°47’07”W ; MZUSP 115075,1, SL 209 mm , Maceió, 9°24’S , 35°31’W ; MZUSP 67336, 30, SL 78–100 mm , Bahia: mouth of Rio Paraguaçu, near Maragogipe, 12°7.5’S , 38°09’W ; MZUSP 115076, 1, SL 100 mm , Rio de Janeiro: Baía da Ilha Grande, 23°09’07.5”S , 44°13’44”W ; MZUSP 15076, 2, SL 64 and 66 mm , Atafona, 21°36’S , 41°01’W ; MZUSP 67319, 27, SL 22–88 mm , Atafona, Ilha da Convivência, 21°36’S , 41°02’W ; MZUSP 43717, 67383, 67393, 67406, MZUSP 115078) 10, SL 52–193 mm , São Paulo: Cananéia, 25°00’45”S , 47°56’07”W ; MZUSP 67407, 2, SL 172 and 182 mm , Cananéia, Ilha do Cardoso, 25°07’54”S , 47°57’59”W ; MZUSP 67322, 1, SL 156 mm , Iguape, Rio Ribeira de Iguape, 24°30’29”S , 47°31’39”W ; MZUSP 67317, 3, SL 155–200 mm , Santos, Praia do Embaré, 23°58’23S, 46°19’26”W ; MZUSP 15079,1, SL 32 mm , Praia Grande, 24°00’31.8”S 46°24’44”W ; MZUSP 67526, 3, SL 88–107 mm , Santa Catarina: Ibiraquera, Barra da Lagoa de Ibiraquera, 28°07’44”S , 48°39’50.2”W ; MZUSP 67411-12, 20, SL 27–64 mm , Rio Grande do Sul: Laguna dos Patos, approximately 31°09’S , 51°25’W . Diagnosis . Mugil brevirostris is the only species from the study area having the pectoral fin reaching to, or extending slightly beyond, the vertical through the dorsal-fin origin ( vs. tip of the pectoral fin not reaching the vertical through the dorsal-fin origin). Mugil brevirostris is additionally distinguished from M . liza , M . curvidens and M . trichodon in having 3 spines and 9 branched anal fin rays in adults or 2 spines, 1 unbranched and 9 branched anal fin rays in juveniles ( vs. 3 spines and 8 branched anal fin rays in adults or 2 spines, 1 unbranched and 8 branched anal fin rays in juveniles) and from M . incilis in having the dorsal-fin origin located midway between the snout tip and the caudal-fin base and 35–38 oblique scale rows between the dorsal limit of the pectoral-fin base to the caudal-fin base ( vs. the dorsal-fin origin closer to the snout tip than to the caudal-fin base and 41–44 oblique scale rows from the dorsal origin of the pectoral fin base to the caudal-fin base). Mugil brevirostris additionally differs from M . margaritae in having 35–38 oblique scale rows from the dorsal limit of the pectoral-fin base to the caudal base ( vs. 40–44 oblique scale rows from the dorsal limit of the pectoral-fin base to the caudal-fin base), and from M . curema and M . rubrioculus by having the pectoral fin with 2 unbranched and 12–14, usually 13 branched rays ( vs. 2 unbranched and 14–16, usually 15 branched rays). FIGURE 1. Mugil brevirostris , MZUSP 31446. A—Detail of mouth showing part of dentition. B—Detail of scales located on dorsal region of body below dorsal-fin base. Description. Morphometric data presented in Table 1 . Maximum examined body length 209 mm SL. Body elongate, compressed and moderately 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, straight from vertical through anterior border of orbit to vertical through spinous dorsal-fin origin, slightly convex from that point to caudal peduncle, concave along caudal peduncle. Ventral profile of head and body convex from tip of lower jaw to caudal peduncle, concave along caudal peduncle. Orbital diameter greater than snout length. Eye covered with adipose tissue, except for narrow oval-shaped central area in adults. Adipose tissue small to almost absent in specimens shorter than 30–35 mm SL. Anterior spinous dorsal fin with 4 slender spines connected by membrane, n = 138. Spinous dorsal fin-origin about midway between snout tip and caudal-fin base. Posterior soft dorsal-fin rays in specimens smaller than 30–35 mm SL ii,7, n = 45; i, 8 in adult specimens, n = 93. Unbranched pectoral-fin rays ii; first ray much smaller than second; branched rays 12–14, 13, n = 138. Tip of pectoral fins reaching to, or extending slightly beyond, vertical through spinous dorsal-fin origin. Pelvic fin I, 5. Tip of pelvic fins reaching vertical through base of third dorsal-fin spine. Anal fin in specimens smaller than 30–35 mm SL II, i, 9; III, 9 in adult specimens, n =138. Mouth subterminal. Tip of maxilla falling short of vertical through anterior border of orbit. Teeth unicuspid, spatulate with slightly curved tips ( Fig. 1 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. TABLE 1. Morphometrics of Mugil brevirostris . 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 Scales spinoid; spines well-developed and projecting on surface of scales ( Fig. 1 B). Transverse scale rows from dorsal limit of pectoral fin base to caudal-fin base 33–39, 36.7, n = 113. Horizontal scale rows from spinous dorsal-fin origin to pelvic-fin origin 12–14, 12.6, n = 112. Horizontal scale rows around caudal peduncle 18–19,18.5, n = 122. 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 3 times as long as pectoral fin in 218 mm SL specimen (MZUSP 115075). Modified axilla scale dorsal to pelvic fin about twice as long as pelvic fin in same specimen.

Standard length	138 22.0–209.0	78.6
Body depth	133 21.1–029.1	25.5	1.9
Snout to dorsal-fin origin	138 70.3–079.5	74.4	1.7
Snout to pectoral-fin origin	138 26.0–033.4	29.6	1.9
Snout to pelvic-fin origin	138 36.2–044.9	40.9	2.0
Snout to anal-fin origin	138 62.2–072.4	67.7	2.3
Caudal peduncle depth	138 09.6–011.1	12.5	0.5
Caudal peduncle length	138 15.4–022.2	18.3	1.4
Pectoral-fin length	135 19.2–024.4	22.1	1.0
Pelvic-fin length	138 15.1–020.4	17.2	1.1
Head length	138 25.0–032.7	29.3	2.0
Head width	137 50.0–058.6	53.6	2.4
Head depth	138 60.0–070.0	65.0	2.6
Lip thickness	136 03.0–007.0	0 4.8	0.7
Mouth width	138 24.0–029.6	26.7	1.2
Mouth depth	138 22.2–027.2	24.6	1.1
Horizontal orbital diameter	138 24.0–030.8	27.8	1.7
Snout length	138 22.0–025.5	23.7	0.9
Upper jaw length	138 23.2–031.2	27.0	1.6

Gill rakers close set, 21–75 on ceratobranchial portion of first arch; rakers increasing in number ontogenetically ( Fig. 2 ). Color in alcohol . Body dark dorsally, with dark color fading ventrally towards midlateral region, whitish on abdominal region. Spinous dorsal, pelvic and anal fins pale, with few scattered dark chromatophores. Soft dorsal, pectorals and caudal fins with numerous scattered dark chromatophores. Anterodorsal basal portion of pectoral fin with small dark spot covering basal portions of unbranched rays and of five dorsalmost branched rays. Cytogenetic and molecular data. Cytogenetic data unavailable. Molecular analyzes showed that the number of nucleotide differences between Mugil brevirostris and the remaining analyzed species ranges from 10.0 to 64.0 (16S) and 62.3 to 109.9 (COI) (Tables 2 to 5). The genetic distance between this species and the remaining analyzed species ranged from 0.017 to 0.127 (16S) and 0.104 to 0.215 (COI) (tables 6 and 7). The dendrogram in Figure 3 shows that this species is genetically most similar to Mugil sp. TABLE 2. Nucleotide differences observed in COI gene among the samples analyzed. Data of Mugil liza and M. cephalus introduced for comparisons 4 13 22 28 31 37 40 43 47 49 55 58 64 65 67 71 76 79 82 85 91 92 94 97 100 103 109 112 Mugil cephalus I - - - - - - - - - - A C A G U C U A U C U C A A C C U U Mugil cephalus II - - - - - - - - - -. U..... A/G U/C......... Mugil cephalus III - - - - - - - - - -... G/A.... C......... Mugil cephalus IV - - - - - - - - - -. U... C/U. A/G........... liza C G C C A A U U C A. U...... U/C.......... trichodon G A U U G A A C C C.. G. C U C...... G U U C.. rubrioculus A G C A U/C G U C C A C U.... C G.... G...... brevirostris A G C A U G U C U A C/U U A/G. C. C. C G C. G C U... Mugil sp. G A U U G G A C C A U U G. C. C. C G.. G C.. C.. incilis G/A A/G U A G/A A G C C U.. G... C.. U C U G... C C. curema A G U A G A G C C A. U G. C. C... C U... U C C. margaritae G A U A G A G C C A. U G... C......... C C. curvidens C G C G A A A C C A.. G. C. C... C. G. U... 118 121 127 130 133 139 142 145 148 151 160 163 166 169 172 175 178 181 184 187 190 193 194 196 197 199 203 205 Mugil cephalus I A C A A U U A A C U A A A C G U/C G C C A U A C A G U C A Mugil cephalus II... G......... U. C.. U..... A... Mugil cephalus III... G......... U. C........ A... Mugil cephalus IV... G......... U. C............. liza ... G......... U............... trichodon C U G C C C.. U C. G. U. C C G U U C... A C... rubrioculus . U. C. C.... G.... C.. U. C. U. A.... brevirostris ... C. C U.. C. G. U. C.... C G U. A... Mugil sp. ... C. C U.. C. G. U. C.... C. U. A C... incilis ... C C C. G U C G G U.. C.... C.. G. C U G. curema .. G C C C. G U C G G.. A C.... C. U.. C U G. margaritae .. G C C C.. U C G G U.. C.... C. U.. C U G. curvidens ... C C C... C.. C. A..... C.... C U. ......continued on the next page TABLE 2 . (Continued) 208 209 211 214 220 223 226 229 232 235 241 244 247 250 256 262 263 265 268 271 277 280 283 286 287 289 292 298 Mugil cephalus I A A C A U U A U U C A U C A C U C U C A C U C U C A U G Mugil cephalus II.. U G. C...... U G.... U......... Mugil cephalus III..... C.. C... U G.... U......... Mugil cephalus IV.. U G. C...... U G.... U....... C. liza .. U G U/C... C.... G.... U........ trichodon G. U.... A...... U... U C.. U. C/U G A C rubrioculus G C G. C. G A C U. C.. C/U U/C. G. C/U.. U. U. A A brevirostris G C. G C C. A C U G C... C. G U C. C.. U. A. Mugil sp. G C G G C C. A C U..... C. G U C. C.. U G A A incilis G. U.. C. A C U. C... C. A. C... C. G C A curema G. U. C C. A C U. C..... A U C... C.. C A margaritae G. U.. C. A C U. C U... U A U U.. C/U C.. C A curvidens G. U.... A C U. C U... U A U C U.. C.. A A 301 304 307 308 310 313 316 319 325 328 331 334 337 340 343 344 346 349 350 352 355 356 358 361 364 367 370 373 Mugil cephalus I A A G G U G A A A G U U U C A U A C A C C C G C C C A G Mugil cephalus II.. A.................... U.... Mugil cephalus III.. A.. G/A. G..... A......... U.... Mugil cephalus IV.. A.... G. A............. U... A liza .. A.... G............... U.... trichodon .... A. G. G. C. C....... U U A. U.. A rubrioculus G. A U A. C.. A.. C U C C. A G... U. U U. A brevirostris G. A U A. C.. A C C C U C C G A G... U. U.. A Mugil sp. G. A U A. C. G A C C C U U C G G G U U. U. U U. A incilis . U.. A. U G. A.. C U. C G. G... A.. U G A curema . U.. A. U A/G. A.. C G.. G. G.... U. U G A margaritae . U.. G A U.. A.. C G. C G U G...... U. A curvidens .... A A C.. A.. C G. C G A G U.. G/A.. U G A ......continued on the next page TABLE 2 . (Continued) 376 379 385 388 391 394 400 403 404 406 409 412 415 418 424 425 430 433 436 439 442 445 451 454 455 457 460 463 Mugil cephalus I U U C C U C C U/C C U G G U C U U C U U C U U A C A C U A Mugil cephalus II.. U.... C.. A U............... G Mugil cephalus III....... C.. A U............... G Mugil cephalus IV... U... C.. A U.... U...... C/U... G. liza ....... C.. A U......... U/C..... G. trichodon . C G U C U A C U A A U. U.. U A. U... U. U C G. rubrioculus ...... U. U A G/A.. U. C U C C.... U. U. G. brevirostris C. U.. U. C U G A. C U. C U..... U U C U. G Mugil sp. .. U... U C U G A U. U C C U.... C C U C U... incilis . A.. C. A C. A A U. U.. G C.. C. C C/U C. C G. curema .... C. A C. A A U. U. C G/A... U/C. C U C. C G. margaritae ...... A C. A A U. U.. G... C. C U C U C G. curvidens .. U U C. G.. G A U. U..... U C C C U C U C G 469 472 475 476 477 478 481 484 487 490 493 496 499 502 505 508 511 514 515 517 520 523 526 529 532 535 538 539 Mugil cephalus I U A U A C U U A U A/G A C U C G A U C C A U C U A C U U U Mugil cephalus II............................ Mugil cephalus III.. U/C........... A........ G.... Mugil cephalus IV C........ R/G/A.... A.............. liza ......... G.... A.............. trichodon A C.. U. C.... U.. U.. U U G... U.. C C. rubrioculus C G. C U C C. C G U. C. C.. U U.... C.. C C. brevirostris .. C C U C C.. G.... U..... C U. C U C C C Mugil sp. .. C C U C C....... C...... U. C. C C C. incilis . C C. U C G... G U C U C. C U... U C U.. C.. curema . U.. U C A/G G. G. U C U U. C U... U C U.. C.. margaritae . U.. U C G G. G G U C. U G C U... U C U.. C.. curvidens C C C. U C. G C.. U.. U.. U. G.. C C.... ......continued on the next page TABLE 3 . (Continued)

TABLE 2 . (Continued)
541	544	545	547	550	553	554	556	559	562	568	571	574	577	578	580	586	589	592	593	595	598	601	604	610	613	619	622
Mugil cephalus I A	A	U	A	A	C	U	A	U	U	U	U	A	C	C	G	U	A	C	C	A	U	U	C	U	C	A	G
Mugil cephalus II.	.	C	.	.	.	.	.	.	.	.	C	G	.	.	A	.	.	.	.	.	.	.	.	C	.	.	.
Mugil cephalus III.	.	.	.	.	.	.	.	.	.	.	C	.	.	.	A	.	.	.	.	.	.	.	.	C	.	.	.
Mugil cephalus IV.	.	.	.	.	.	.	.	.	.	.	C	.	.	.	A	.	.	.	.	.	.	.	.	C	.	.	.
M. liza .	.	U/C	.	.	.	.	.	.	.	.	C	.	.	.	.	.	.	.	.	.	.	.	.	C	.	.	.
M. trichodon C	.	C	.	.	U	C	.	.	.	.	.	G	U	.	U	.	G	.	.	.	C	C	U	C	.	G	A
M. rubrioculus A/G	C	C	C	U	.	C	.	C	C	.	.	.	U	U	A	.	U	.	.	.	C	C	U	C	.	.	A
M. brevirostris G	C	C	C	U	.	C	G	C	C	.	.	G	.	U	A	.	A/U	.	.	.	C	C	U	C	.	.	A
Mugil sp. G	C	C	C	U	A	.	.	C	C	C	C	G	.	U	A	C	G	.	.	U	C	C	U	C	.	.	A
M. incilis .	C	C	U/C	.	.	C	G	.	.	C	.	G	.	.	A	C	.	.	U	.	.	.	.	C	.	.	A
M. curema .	U	C	C/U	.	.	C	.	C	.	C	.	G	.	.	A	C	.	U	U	.	.	.	.	C	.	.	A
M. margaritae .	C/U	C	C	.	U	C	.	C	.	.	.	G	.	.	A	C	.	U	U	.	.	.	.	C	.	.	A
M. curvidens G	C	C	C	U	.	C	.	C	.	C	C	G	.	.	A	C	.	.	U	.	C	C	U	.	U	C	A
625	628	631	634	637	640	643	646	649	650	652	653	654	655
Mugil cephalus I A	G	C	A	U	G	U	A	C	C	A	C/U	A/U	U/A
Mugil cephalus II.	.	.	.	.	.	C	.	.	.	.	.	U	A
Mugil cephalus III.	.	.	.	.	.	.	.	.	.	.	.	U	A
Mugil cephalus IV.	.	.	.	.	.	.	.	.	.	.	.	A/U	U/A
M. liza .	.	.	.	.	.	.	.	.	.	G	U	U	C
M. trichodon .	.	U	C	.	U	C	G	U	.	.	U	U	C
M. rubrioculus G	A	.	.	C	.	C	.	.	U	.	U	U	C
M. brevirostris A/G Mugil sp. .	A A	..	..	C C	. A	C C	A/G G	C/U U	C/U.	..	U U	U U	C C
M. incilis .	A	.	.	.	A	.	A/G	.	.	.	U	U	C
M. curema .	.	.	.	.	.	.	.	.	U	.	U	U	C
M. margaritae .	G/A	.	.	.	.	.	.	.	U	.	U	U	C
M. curvidens .	U	U	.	C	.	C	.	.	.	.	U	U	C

TABLE 3. Nucleotide differences observed in the 16S gene among the samples analyzed. Data of Mugil liza and M . cephalus are introduced for comparisons.

12	13	15	16	17	20	32	41	47	50	51	55	70	71	73	78	81	91 101 102 114 115
Mugil cephalus I Mugil cephalus II Mugil cephalus III Mugil cephalus IV M liza M trichodon M rubrioculus M brevirostris Mugil sp M incilis M curema M margaritae M. curvidens	- - - - A U C C U U U U U 118	- - - - A U A A A A A A A 130	- - - - C C C C C U C C C 131	- - - - C U A A A C C C C 132	- - - - U U C C C C C C C 144	- - - - C C U U U C C C C 156	C U U U......... 160	A.... U U U U. U A/U U 161	C........ U... 177	U..... A A A.... 178	- - - - - G A A A G G G G 183	C.... U U U U A U U U 195	U..... C C C C C C C 200	U..... C C C C C C C 201	A.... G G G G G G G G 222	C.... U U U U U U U U 223	A........ G G G G 236	C U C G A.. U A... U A.... A.... A.. C.. G............... U.. G/A................ 238 241 242 247 250
Mugil cephalus I M ugil cephalus II Mugil cephalus III Mugil cephalus IV M liza M trichodon M rubrioculus M brevirostris Mugi l sp M incilis M curema M margaritae M. curvidens	A.... C U U U C U U U	U.... A A A A A A A A	A.... C U U U C C C U	G.... A A A A.. A U	A........ G...	U..... C C C C C C C	A........ G G G G	A..... G G G G G G G	U. C/U..........	U. C/U U/C........ C	U/C............	A........... G	U.... C. C C....	A........ G...	G.... A A A A A A A A	C.... U U U U U U U U	G.... A A A A.. G/A A	C A A U C....................... C U U G. C U U.. C U U.. C U... C U.. G C U... C U... C U .... ..continued on the next page

251 252 253 254 255 256 258 261 262 263 264 265 266 268 270 271 272 275 276 277 279 280 Mugil cephalus I G U C A A A A U C C C U U A C A G A A U A C Mugil cephalus II......... U.. C...... C.. Mugil cephalus III............ C...... C.. Mugil cephalus IV......... U.. C...... C.. Mugil lza ......... U.. C...... C.. trichodon .. U G. C. C.. U A A - U...... U rubrioculus .... G C. C.. A C A - G. A. G.. U brevirostris .... G C. C.. A C A - A. A. G. G U Mugil sp.... G C. C. U A. A - A. A. G.. U incilis .. -.. U.. U.. C A U U G A.. A.. curema ..... C.... U C A C U. A A/G G C. U margaritae ..... C.... U C A C U. A. G C. U curvidens A C... C G A.. U. A C.. A. G... 281 283 287 288 289 290 295 296 297 298 299 300 306 307 324 328 330 331 341 342 348 349 Mugil cephalus I A A - A A C U U C C A C A G/U U C G A G C C U Mugil cephalus II.. -................... Mugil cephalus III.. -................... Mugil cephalus IV.. -.... C........ A..... liza .. -................... trichodon U U -....... C. U. G. A G A... rubrioculus . U A. C.... U C. C. G U A. A. U C brevirostris U U A. C.... U C. C. G U A. A U. C Mugil sp U U A. C.... U C. C. G U A U A U. C incilis . U G. U U. C U. C. U. G. A G A U. C curema . U A G C U C... C. U. G. A G A.. C margaritae . U A G C..... C. U. G. A G A U U C curvidens U U A. U.. C.. C U U. G U A G A.. C .... ..continued on the next page Distribution . Specimens of Mugil brevirostris that have been examined in this study were found at area from the northern (Amapá) to the southern Brazilian coast (Rio Grande do Sul). This species is captured together with M . curvidens , M. curema , M . incilis , and M . rubrioculus along all its distributional range ( Fig. 4 ).

TABLE 3. (Continued)
351	354	355	357	363	364	366	367	373	374	388	389	391	395	396	397	402	405	412	414	415	422
Mugil cephalus I A	G	C	C	A	C	U	U	C	A	U	C	C	C	A	U	U	A	A	U	A	C
M ugil cephalus II.	.	.	U	.	U	.	C	U	.	.	.	.	.	.	.	.	.	.	.	.	.
Mugil cephalus III.	.	.	.	.	U	.	C	U	.	.	.	.	.	.	.	.	.	.	.	.	.
Mugil cephalus IV.	.	.	.	.	.	.	C	U	C	.	.	.	.	.	.	.	.	.	.	.	.
M liza .	.	.	.	.	.	.	C	U	.	.	.	.	.	.	.	.	.	.	.	.	.
M trichodon .	.	.	.	-	-	C	C	-	-	C	U	.	U	.	C	C	.	.	C	U	U
M rubrioculus G	.	U	.	-	-	C	C	-	-	C	U	U	.	.	C	C	C	.	.	.	U
M brevirostris G	.	.	.	-	-	C	C	-	-	C	U	A	.	.	C	C	C	.	.	.	U
Mugi l sp.	.	.	.	-	-	C	C	-	-	C	.	A	.	.	C	C	C	.	.	.	U
M incilis G	.	.	.	U	U	.	G	-	-	C	U	G	.	.	.	A	C	.	.	U	U
M curema G	.	.	.	-	-	C	.	-	-	C	U	G	.	G	C	A	C	.	.	U	.
M margaritae G	.	.	.	-	-	C	.	-	-	C	U	G	.	.	C	A	C	.	.	U	U
M. curvidens G	A	.	.	-	-	.	.	-	-	C	U	A	.	.	C	A	C	G	.	U	.
435	449	456	458	461	463	465	474	498	555	563	570
Mugil cephalus I A	U	U	A	G	C	A	C	U	A	A	U
M ugil cephalus II. Mugil cephalus III.	C.	..	. A/G	. A/G	..	..	..	..	..	..	..
Mugil cephalus IV.	.	.	.	.	.	.	.	.	.	.	.
M liza .	C	.	.	.	.	.	.	.	.	.	.
M trichodon .	.	C	.	.	.	C	U	.	.	.	.
M rubrioculus .	C	C	.	.	.	C	U	C	.	G	.
M brevirostris .	C	C	.	.	.	C	U	C	.	G	.
Mugi l sp.	C	C	.	.	.	C	U	C	.	G	.
M incilis .	C	C	.	A	U	C	U	C	G	G	C
M curema .	C	C	.	A	U	C	U	C	G	G	C
M margaritae .	C	C	.	A	U	C	U	C	G	G	C
M. curvidens U	C	C	.	.	.	C	U	C	G	G	.

TABLE 4. Number of nucleotide differences observed in 16S gene among the samples analyzed. Mean and stardard error below and under the main diagonal respectively. Data of Mugil liza and M . cephalus are introduced for comparisons.

1	2	3	4	5	6	7	8	9	10	11	12	13
M. cephalus I	3.1	2.5	3.1	2.5	6.2	7.1	7.1	7.1	7.1	7.0	7.3	7.3
M. cephalus IV	10.8	2.4	2.4	2.2	6.3	7.3	7.3	7.1	7.0	7.2	7.4	7.4
M. cephalus III	9.1	7.7	1.9	2.3	6.4	7.3	7.3	7.2	7.2	7.3	7.5	7.5
M. cephalus II	11.7	7.1	4.6	1.9	6.5	7.2	7.3	7.1	7.1	7.2	7.4	7.5
M. liza	7.7	5.1	6.6	4.0	6.6	7.4	7.3	7.2	7.1	7.2	7.5	7.5
M. trichodon	46.7	49.1	50.0	52.0	52.0	5.9	5.7	5.6	6.3	6.1	5.8	6.4
M. rubrioculus	58.7	61.1	62.0	62.0	64.0	45.0	3.0	3.8	6.7	5.9	5.6	6.2
M. brevirostris	59.7	62.1	63.0	63.0	64.0	41.0	10.0	2.5	6.5	5.7	5.4	5.7
Mugil sp.	57.7	58.1	61.0	59.0	60.0	40.0	15.0	7.0	6.6	5.9	5.6	5.7
M. incilis	61.4	61.7	62.8	63.6	63.2	50.4	49.8	48.3	50.8	5.2	4.9	6.3
M. curema	61.0	63.5	63.5	64.3	63.3	45.3	38.3	38.3	40.3	28.9	2.7	5.3
M. margaritae	60.7	63.1	63.2	64.0	63.0	41.0	32.0	32.0	34.0	26.6	8.7	5.3
M. curvidens	63.7	65.9	68.6	69.0	68.0	50.0	42.0	39.0	39.0	43.2	32.3	31.0

TABLE 5. Number of nucleotide diferences observed in COI gene among the samples analyzed. Mean and stardard error below and under the main diagonal respectively. Data of Mugil liza and M . cephalus are introduced for comparisons.

1	2	3	4	5	6	7	8	9	10	11	12	13
1 M. cephalus I	5.1	4.7	4.7	4.2	8.9	8.8	9.1	9.6	9.1	8.9	8.8	9.2
2 M. cephalus IV	28.0	3.8	3.7	3.8	8.8	8.9	8.8	9.1	8.9	8.6	8.7	9.1
3 M. cephalus III	24.5	16.6	3.5	3.6	9.0	8.8	8.7	9.1	9.1	8.6	8.6	9.3
4 M. cephalus II	26.3	16.3	14.6	4.0	8.7	8.8	8.6	9.0	9.2	8.7	8.6	9.2
5 M. liza	22.4	15.7	15.3	17.4	9.0	8.9	9.1	9.4	9.1	8.8	9.0	9.4
6 M. trichodon	98.2	98.3	101.7	94.5	108.4	9.3	9.5	9.2	9.4	9.1	9.3	9.1
7 M. rubrioculus	97.2	98.3	102.3	99.7	105.0	108.0	7.2	7.8	8.9	8.9	8.9	8.9
8 M. brevirostris	109.0	104.4	104.4	100.4	109.9	114.2	62.3	6.2	9.1	8.8	9.0	8.8
9 Mugil sp.	115.7	109.0	108.4	103.4	116.9	101.5	75.4	47.0	8.8	9.0	9.0	9.0
10 M. incilis	97.7	98.2	98.4	102.0	106.9	105.0	105.9	110.2	101.5	5.7	6.0	8.1
11 M. curema	96.6	90.6	95.4	95.6	98.1	104.1	101.7	104.8	104.8	39.8	5.1	7.9
12 M. margaritae	93.4	88.8	93.2	92.4	99.3	102.5	97.2	107.2	102.2	46.5	29.4	7.3
13 M. curvidens	101.1	100.8	102.8	100.8	105.7	101.7	97.1	98.7	95.0	78.9	74.8	73.2

TABLE 6. Genetic distances (using the Kimura-2-parameter model) observed in 16S gene among the samples analyzed. Mean and stardard error below and under the main diagonal respectively. Data of Mugil liza and M . cephalus are introduced for comparisons.

1	2	3	4	5	6	7	8	9	10	11	12	13
1 M. cephalus I	0.006	0.005	0.006	0.005	0.014	0.016	0.016	0.015	0.018	0.015	0.016	0.016
2 M. cephalus IV	0.019	0.004	0.004	0.004	0.014	0.016	0.016	0.015	0.018	0.016	0.016	0.017
3 M. cephalus III	0.016	0.014	0.003	0.004	0.014	0.016	0.016	0.016	0.018	0.016	0.017	0.017
4 M. cephalus II	0.021	0.013	0.008	0.003	0.015	0.016	0.016	0.015	0.018	0.016	0.016	0.017
5 M. liza	0.014	0.009	0.012	0.007	0.014	0.015	0.015	0.014	0.017	0.015	0.015	0.016
6 M. trichodon	0.092	0.097	0.099	0.103	0.096	0.012	0.011	0.011	0.015	0.012	0.011	0.013
7 M. rubrioculus	0.118	0.123	0.125	0.125	0.121	0.083	0.005	0.007	0.016	0.011	0.010	0.012
8 M. brevirostris	0.120	0.125	0.127	0.127	0.120	0.075	0.017	0.004	0.015	0.011	0.010	0.011
9 Mugil sp.	0.115	0.116	0.123	0.118	0.112	0.073	0.026	0.012	0.015	0.011	0.010	0.011
10 M. incilis	0.142	0.143	0.146	0.148	0.141	0.112	0.110	0.107	0.113	0.011	0.011	0.015
11 M. curema	0.123	0.128	0.128	0.130	0.119	0.084	0.070	0.070	0.073	0.062	0.005	0.010
12 M. margaritae	0.122	0.127	0.128	0.129	0.118	0.075	0.057	0.057	0.061	0.057	0.015	0.010
13 M. curvidens	0.128	0.133	0.139	0.140	0.128	0.093	0.076	0.071	0.071	0.095	0.058	0.056

TABLE 7. Genetic distances (using the Kimura-2-parameter model) observed in COI gene among the samples analyzed. Mean and stardard error below and under the main diagonal respectively. Data of Mugil liza and M. cephalus are introduced for comaprisons.

1	2	3	4	5	6	7	8	9	10	11	12	13
1 M. cephalus I	0.009	0.009	0.009	0.008	0.020	0.020	0.022	0.023	0.021	0.020	0.020	0.021
2 M. cephalus IV	0.049	0.006	0.006	0.006	0.020	0.020	0.021	0.022	0.020	0.019	0.019	0.020
3 M. cephalus III	0.042	0.028	0.006	0.006	0.020	0.020	0.020	0.022	0.021	0.020	0.019	0.021
4 M. cephalus II	0.046	0.028	0.025	0.007	0.019	0.020	0.020	0.021	0.021	0.020	0.019	0.021
5 M. liza	0.039	0.027	0.026	0.030	0.019	0.019	0.020	0.021	0.019	0.019	0.019	0.020
6 M. trichodon	0.189	0.190	0.198	0.181	0.193	0.019	0.020	0.019	0.020	0.019	0.019	0.018
7 M. rubrioculus	0.187	0.189	0.199	0.193	0.185	0.190	0.013	0.015	0.019	0.019	0.018	0.019
8 M. brevirostris	0.215	0.203	0.203	0.194	0.195	0.203	0.104	0.011	0.020	0.019	0.020	0.019
9 Mugil sp.	0.230	0.214	0.212	0.200	0.210	0.176	0.128	0.077	0.019	0.019	0.019	0.019
10 M. incilis	0.189	0.190	0.190	0.199	0.190	0.186	0.188	0.198	0.179	0.010	0.011	0.016
11 M. curema	0.186	0.173	0.184	0.184	0.172	0.184	0.179	0.186	0.186	0.064	0.009	0.015
12 M. margaritae	0.179	0.169	0.179	0.177	0.174	0.180	0.170	0.191	0.180	0.076	0.047	0.014
13 M. curvidens	0.196	0.196	0.201	0.196	0.188	0.178	0.170	0.174	0.166	0.135	0.128	0.125

FIGURE 2 . Regression of number of gill rakers versus standard length for the examined species of Mugil . Data from M . trichodon unavailable. FIGURE 3 . Dendrograms obtained by the neighbour-joining method using the Kimura-2-parameters model of nucleotide substitution for the data of genes COI (a) and 16S (b). The numbers represent the groups frequencies observed using 1,000 bootstrap pseudoreplicas. Values below 50% are not represented. Remarks. No specimen that could be identified as type-material of Querimana brevirostris was found in the collection of the Museu Nacional, Rio de Janeiro, despite a careful, approximately six months search by Marcelo Britto, ichthyologist and curator of that institution. One specimen (MNRJ 2021) in very poor condition which had been identified by Alípio de Miranda Ribeiro as Querimana brevirostris , but not cited as a type , is smaller (total length = 30 mm ) than specimens used by Ribeiro, 1915 in his original description (several specimens from an unknown locality in Brazil measuring 53 mm in length). Nine specimens (MNRJ 2977) from Baía da Guanabara, Rio de Janeiro, SL 32–37.5 mm , also identified as Querimana bevirostris are rather Mugil curvidens Valenciennes. In Ribeiro’s original description there is a clear reference to the tip of the pectoral fins reaching the vertical through the origin of the first dorsal-fin spine and the presence of 33 “lateral line” scales. The only species of Mugil that has long pectoral fins, reaching to, or extending slightly beyond, the vertical through the base of the first dorsal-fin spine is Mugil hospes Jordan & Culver, described from Sinaloa, Mexico , Pacific Ocean. However, two examined paratypes of this species (SU 28290, SL 166 mm and USNM 47446, SL 160 mm ), have respectively 43 and 45 lateral series scales along the sides of the body. Given that the specimens from Brazil identified as Mugil gaimardianus by Menezes & Figueiredo (1985) and Mugil sp. by Menezes et al . (2003) and all specimens herein examined characteristically have the tip of the pectoral fins reaching to, and sometimes extending slightly beyond, the vertical through the origin of the first dorsal-fin spine and 33–39 lateral series scales, they should be considered as Mugil brevirostris ( Ribeiro, 1915 ) . Since the the type series of this species is lost, one of the M. brevirostris specimens we have examined in the present study (MZUSP 115087, SL 106 mm ) from Cananéia, São Paulo is herein designated as neotype in order to resolve the uncertainty about the identity of the species.