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Accepted by M.R. de Carvalho: 10 Jun. 2010; published: 28 Jun. 2010 59
ZOOTAXA
ISSN 1175-5326 (print edition)
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Zootaxa 2519: 59–68 (2010)
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An old taxonomic dilemma: the identity of the western south Atlantic lebranche
mullet (Teleostei: Perciformes: Mugilidae)
NAÉRCIO A. MENEZES1 , CLÁUDIO DE OLIVEIRA2 & MAURO NIRCHIO3
1Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42494, 04218-970, São Paulo,SP, Brazil. E-mail: naercio@usp.br
2Departamento de Morfologia, Instituto de Biociências, Universidade Estadual Paulista, 18618-000, Botucatu, SP, Brazil.
E-mail: cláudio@ibb.unesp.br
3Escuela de Ciências Aplicadas del Mar, Universidad de Oriente, Apartado Postal 147, Porlamar, Isla de Margarita, Venzuela.
E-mail: nirchio@cantv.net
Abstract
The identification of the lebranche mullet in the western south Atlantic has long been problematical. In most recent
works either Mugil liza Valenciennes and M. platanus Günther, 1880 or M. liza and M. cephalus Linnaeus, 1758 were
recognized from the region and more rarely the occurrence of only one species has been proposed but without sufficient
morphological, biochemical or molecular data to allow the designation of the taxonomically appropriate name. Analysis
of meristic and morphometric data taken from samples collected from Venezuela to Argentina, clearly indicates that there
is only one species of lebranche mullet in the Caribbean Sea region and the Atlantic coast of South America and that
Mugil liza is the appropriate name. The comparison of the combined data from all the samples of M. liza with the data
taken from one sample of M. cephalus that originated in the Mediterranean, the possible locality from which type
specimens were collected (Eschmeyer and Fricke, 2009), revealed significant differences indicating that they are
different species. It is also suggested that individuals from the western north Atlantic identified as M. cephalus might
represent a population of M. liza in this region.
Key words: Mugilid species, Taxonomy, Misidentification, Caribbean and South American Atlantic waters
Resumo
A identificação da tainha que ocorre no Atlântico sul ocidental tem sido exaustivamente discutida e é
problemática. Nos trabalhos mais recentes Mugil liza Valenciennes, 1836 e M. platanus Günther, 1880 ou M.
liza e M. cephalus Linnaeus, 1758 foram reconhecidas e mais raramente a ocorrência de apenas uma espécie
foi sugerida sem dados suficientes, tanto morfológicos como bioquímicos ou moleculares para possibilitar a
designação do nome taxonomicamente mais apropriado. A análise de dados merísticos e morfométricos
obtidos de amostras coletadas da Venezuela à Argentina, indica claramente a existência de uma só espécie de
tainha na região do Caribe e na costa Atlântica da América do Sul e Mugil liza é o nome que deve ser a ela
atribuído. A comparação dos dados de todas as amostras combinados com uma amostra de M. cephalus obtida
do Mediterrâneo, considerado como a provável localidade de onde os exemplares-tipo foram coletados
(Eschmeyer and Fricke, 2009), revelou diferenças significantes indicando que elas são espécies distintas.
Também é sugerido que indivíduos do Atlântico norte ocidental identificados como M. cephalus podem
representar apenas uma população de M. liza nesta região.
Introduction
The identification and distribution of the lebranche or striped mullet in the Atlantic waters of South America
has long been controversial. Thomson (1963) considered Mugil cephalus Linnaeus, 1758 to be widely
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distributed in tropical and subtropical zones of all seas, but did not confirm its presence along the Atlantic
coast of South America. Menezes (1983, 2003) and Menezes & Figueiredo (1985) did not consider Mugil
cephalus the appropriate name for the striped mullet found in coastal waters from southeastern Brazil to
Argentina. Rather they proposed that Mugil liza Valenciennes, 1836, and M. platanus Günther, 1880, were
two allopatric species distributed respectively north and south of Rio de Janeiro that should be recognized in
the western south Atlantic. In his worldwide revision of the Mugilidae Thomson (1997) identified specimens
collected in coastal waters of Brazil and Argentina as Mugil cephalus and considered M. platanus to be its
junior synonym. He also considered M. liza Valenciennes, 1836, as occurring from the West Indies to Brazil
without precisely defining the southern limit of its distribution. Harrison (2003) subsequently extended the
southern distribution of this species to Rio de Janeiro and also pointed out that M. cephalus was present from
the West Indies to Argentina. In a more recent study Cousseau (2005) demonstrated that the species found in
southern Brazilian and Argentinean coastal waters is Mugil platanus which she recognized as different from
M. liza. This conclusion was confirmed by Castro et al. (2008) who additionally indicated that M. platanus
differs from M. cephalus based on mitochondrial DNA, morphometrics and meristics. Heras et al. (2009)
suggested that Mugil platanus and M. liza “represent a continuum of a single species closely related but
distinct from M. cephalus”. Those authors did not analyze all of the taxonomic implications involved in the
recognition of a single species in the western south Atlantic, a task beyond the use of molecular data alone.
In this paper we provide evidence from morphological data indicating that Mugil liza Valenciennes is
different from M. cephalus Linnaeus and is the only lebranche mullet found along the Atlantic coast of South
America.
Methods and material
Counts and measurements were made on the left side of the specimens. Measurements of small specimens
were taken with calipers and of large specimens with a ruler and recorded to tenths of a millimeter.
Measurements of standard length and those referring to the origin of the fins were all made orthogonal to the
main body axis and the remaining ones were made point to point. The following measurements were taken:
body depth, measured vertically down from the origin of the first (spinous) dorsal-fin origin; head length,
from tip of premaxillary to posteriormost tip of opercle including its membranous margin; head depth, its
deepest posterior portion; head width, its widest posterior portion; upper jaw length, from the tip of the
premaxillary to the posteriormost tip the of the maxilla; orbital diameter, the distance from the anterior to the
posterior margins of the orbit with the adipose eyelid excluded; snout length, from the tip of the premaxillary
to the anterior margin of the orbit; interorbital distance, the least distance between the borders of the frontal
bones; caudal peduncle depth, the least depth measured vertically; pectoral-fin length, from the base of the
uppermost ray to the posteriormost tip of the longest ray; pelvic-fin length, from the base of the spine to the tip
of the longest ray; snout to second (soft) dorsal-fin origin; snout to pectoral-fin origin; snout to pelvic-fin
origin; snout to anal-fin origin.
Meristic counts are as follows: Circumpeduncular scale rows, the number of horizontal rows around the
caudal peduncle, counted just anterior to the point of caudal flexure; transverse scale rows, the number of
oblique rows counted from the scale immediately above the insertion of the pectoral fin to the caudal flexure
(equivalent to lateral scale rows of various authors); horizontal scale rows, counted from the origin of the first
(spinous) dorsal-fin origin vertically to row passing through the pelvic-fin origin; and gill rakers, counted on
ceratobranquial portion of the first gill arch. Other counts and measurements follow Thomson (1997).
Meristic data were treated through comparative box plot since the data sets from each sample are
considered non-parametric. The Mann-Whitney test (BioEstat 5.0, in Ayres et al., 2007) was then used
because we think it expresses better differences among samples from populations geographically arranged in a
linear fashion and is useful to detect clinal variation. A difference was considered statistically significant
when p 0.05. Gill rakers increase in number according to increase in standard length and were compared
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THE IDENTITY OF THE LEBRANCHE MULLET
through regression analysis (Fig.2). The regression trend proved to be logarithmic.
For comparative purposes the samples are represented by specimens from limited latitudinal ranges as
follows: Venezuela (10º57’N – 11º10’N); Northeastern Brazil (02º31’S – 10º53’S); Southeastern Brazil
(21º31’S – 25º31’S); South Brazil (30º00’S – 32º8’S); and Uruguay and Argentina (34º53’S – 36°10’S).
Specimens examined are from the following institutions: Museu de Zoologia, Universidade de São Paulo,
Brazil (MZUSP), Laboratório de Biologia de Peixes, Departamento de Morfologia, Instituto de Biociências,
Universidade Estadual Paulista, Botucatu, São Paulo, Brazil (LBP), and Museo Nacional de Ciencias
Naturales (MNCN-CSIC), Madrid, Spain.
Synonymies are by no means complete and include primarily the most relevant species geographical
distribution data from the literature in order to be consistent with the results of this study.
A full species description was not considered necessary and only diagnostic characters are presented to
allow the recognition of the species from its congeners.
Mugil liza Valenciennes, 1836
Fig. 1
Mugil liza Valenciennes in Cuvier & Valenciennes 1836: 83 (Brazil, Puerto Rico, Maracaibo, Cuba, Martinique,
Surinam, Cayenne, Buenos Aires; lectotype: MNHN A-4659, Martinique Island, West Indies, designated by
Harrison 1993: 146; different lectotype (A-1051) erroneously designated by Thomson, 1997: 472; paralectotypes:
MNHN 6307 (1), Buenos Aires, A-1050 (1, dry), Cayenne, A-3668 (1), Suriname, A-4642 (1, dry), Maracaibo, A-
4656 (1, dry), Maracaibo, A-4657 (1, dry), Maracaibo, A-5769 (1) Puerto Rico (not Martinique); Steindachner,
1879: 26 (only specimens from Río Magdalena, Colombia); Thomson, 1978 [(unpaginated), only specimens
originating from the West Indies to Natal, Brazil)]; 1997: 492 (West Indies to Brazil); Menezes, 1983: 4 (only
specimens originating from Caribbean Islands to Rio de Janeiro, Brazil; 2003: 65 (only specimens originating from
the West Indies to Rio de Janeiro, Brazil); Menezes & Figueiredo, 1985: 23 (only specimens originating from the
West Indies to Rio de Janeiro, Brazil); Cervigón, 1992: 363 (northern coast of South America); Bencomo, Borjas &
Castillo, 1997: 159 (Venezuela); Aguilera, 1998: 51 (Venezuela); Marin, 2000: 76 (Venezuela); Eiras-Stofella,
Chavert-Almeida, Fanta & Vianna, 2001: 122 (southeastern Brazil; ultrastructure of gills); Lopez, Morgan &
Montenegro, 2002: 64 (Argentina, listed); Lopez, Miquelarena & Menni, 2003: 78 (Argentina); Harrison, 2003:
1084 (only specimens originating from the West Indies to Rio de Janeiro, Brazil); Araújo, Teixeira & Oliveira, 2004:
78 (northeastern Brazil).
Mugil brasiliensis Spix & Agassiz, 1831: 134 [(misprinted 234), pl. 72; type locality: Atlantic off Brazil]. Name
suppressed and placed on Official Index Opinion 1787. Steindachner, 1876: 88 (only specimens from Brazil); Jordan
& Evermann, 1896: 810 (Cuba to Patagonia); Evermann & Marsh, 1902: 112 (Puerto Rico); Fowler, 1903: 743
(Brazil, Surinam, St. Martin); 1936: 586 (West Indies, Brazil, Surinam); Meek & Hildebrand, 1923: 274 (West
Indies to Brazil); Schultz, 1949: 111 (Venezuela); Ringuelet & Aramburu, 1960: 57 (Argentina, listed); Cervigón,
1993: 371 (Venezuela).
Mugil lebranchus Poey, 1860: 251, pl. 18 (Cuba to Patagonia)
Mugil lisa Ribeiro, 1915 [(unpaginated), Brazil].
Mugil platanus Günther, 1880: 9 (type locality: Río de la Plata, Buenos Aires, Argentina); Jordan & Swain, 1884: 266
(Argentina); Ribeiro, 1915 [(unpaginated), Brasil]; Ringuelet & Aramburu, 1960: 57 (Uruguay and Argentina);
1961: 57 (Argentina, listed); Menezes, 1983: 5 (Rio de Janeiro, Brazil to Argentina); Menezes & Figueiredo, 1985:
24 (Rio de Janeiro, Brazil to Argentina); 2003: 65 (Rio de Janeiro, Brazil to Argentina); Eiras-Stofella, Chavert-
Almeida, Fanta &Vianna, 2001: 122 (southern Brazil; ultrastructure of gills): Lopez, Miquelarena & Menni 2003: 78
(Argentina); Menni, 2004; 85 (Argentina); Cousseau, Castro, Figueroa & Gosztoni, 2005: 138–139 (Argentina; Rio
Grande do Sul, Brazil); Castro, Heras, Cousseau & Roldán, 2007: 322–324 (southern Brazil and Argentina).
Mugil cephalus (not of Linnaeus, 1758); Ribeiro, 1915 [(unpaginated), Brazil, listed]; Thomson, 1978 [(unpaginated),
only specimens originating from the West Indies to Brazil]; 1997, 483 (only specimens from Brazil); Cervigón,
1992: 361 (northern South America); Keith, Le Ball & Planquette, 2000: 22 (French Guiana); Camargo & Isaac,
2001: 141 (northern Brazil); Harrison, 2003: 1079 (only specimens originating from the West Indies to Argentina).
Material examined: MZUSP 76404, 1, SL 54.3 mm, Venezuela, Falcón: mangrove near road San Juan de los
Cayos-Capadare, 11º10’N, 63º26’W; LBP 6064, 6, SL 188–207 mm, Isla Margarita, Nueva Esparta: Boca de
Rio, approximately 11º00’S, 64º10’W; MZUSP 67422, 1, SL 280 mm, Brazil, Maranhão: coast of Maranhão;
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MZUSP 67415, 67419, 8, SL134–395 mm, Alagoas: Lagoa Mundaú, 09º37’S, 35º36’W; MZUSP 67421, 1,
SL 365 mm, Sergipe: rio Sergipe, 10º51’S, 37º00’W; MZUSP 67418,1, SL 23.3 mm, Rio de Janeiro: Atafona,
21º37’S, 41º01’W; MZUSP 49087, 55474, 67431, 8, SL 68–137 mm: Cabo Frio, 22º52’S, 42º01’W; MZUSP
2319, 1, SL 146 mm: Ilha Grande, 23º09’S, 44º13’W; MZUSP 28291-292, 17, SL 80–292 mm: mouth of rio
da Guarda, Itaguaí, 22º57’S, 43º11’W; MZUSP 67416, 1, SL 193 mm: Copacabana, 22º58’S, 43º11’W;
MZUSP 67433, 1, SL 319 mm, São Paulo: Ubatuba, 23º26’S, 45º04’W; MZUSP 67423, 1, SL 203 mm: São
Sebastião, 23º45’S, 45º24’W; MZUSP 67429, 1, SL 407 mm: Ilha dos Búzios, 1, SL 407 mm: 23º48’S,
45º08’W; MZUSP 67445, 1, SL 48 mm: Santos, 23º57’, 46º19’W; MZUSP 67424, 1, SL 290 mm: São
Vicente, 23º57’S, 46º23’W; MZUSP 67426, 1, SL 218 mm: Guarujá, 23º59’S, 46º59’W; MZUSP 44703-704,
2, SL 263–284 mm: Itanhaém, 24º11’S, 46º41’W; MZUSP 67417, 1, SL 260 mm: Valo Grande, 24º41’S,
47º34’W; MZUSP 67430, 74843, 3, SL 54.5–152 mm: Cananéia, 25º01’S, 47º55’W; MZUSP 67427, 1, SL
287 mm, Paraná: Guaraguaçu, Baía de Paranaguá, 25º27’S, 48º25’W; MZUSP 6531, 3, SL 241–332 mm:
market in Paranaguá, 25º31’S, 48º27’W; MZUSP 14328-329, 14331, 14334-336, 14338-343, 67425, 67434,
67436, 64, SL 31–317 mm, Rio Grande do Sul; Tramandaí, 29º58’S, 50º10’W; MZUSP 67446, 4, SL 228–
366 mm: Saco do Justino, Lagoa dos Patos, 31º09’S, 51º25’W; MZUSP 67447, 4, SL 243–328 mm: Praia do
Cassino, 8 kilometers south of Molhes, 32º10’S, 52º10’W; MZUSP 67440, 9, SL 24.5–37.5 mm , Rio Grande:
Saco do Justino, south of Lagoa Mangueira, 32º03’S, 52º05’W; MZUSP 67441, 6, SL 24–40 mm: stream in
praia of Sarita, about 12 kilometers from light, 32º38’S, 52º27’W; MZUSP 2318, 4, SL 172–314 mm,
Uruguay: market in Montevideo, 34º53’S, 56º04’W; LBP 6279, 1, SL 343 mm, Argentina: 36º10’S, 56º43’W.
FIGURE 1. Mugil liza, LBP 6064, 188 mm SL, left lateral view.
Diagnostic characters. Morphometric data presented in table 1. First dorsal fin with 4 slender spines.
Second dorsal fin with with 1 unbranched and 8 branched rays. Pectoral fin with 1 unbranched and 14–17
branched rays. Pelvic fin with 1 slender spine and 5 branched rays. Anal fin with 3 spines and 8 branched rays
in adult specimens and 2 spines and 9 branched rays in specimens smaller than 30 mm SL. Dorsal and anal
fins mostly naked with small scales only at their basal portions. Transverse scale rows 29–39 (32–39 in the
studied material). Horizontal scale rows around caudal peduncle 17–20. Gill rakers close-set, 22–80 on
ceratobranchial portion of first gill arch, increasing in number with increase in standard length (Fig. 2).
Vertebrae 24–25 (n = 24, mean = 24.6, SD = 0.5).
Body dark dorsally, silvery on sides and whitish on abdominal region. Variable number of dark stripes along sides of
body aligned with horizontal scale rows along sides of body. All fins pale. Dorsal, pectoral anal and caudal fins with
scattered dark chromatophores.
Discussion
No meristic differences were detected among the samples representing the species in the analysis of all
characters with exception of number of transverse scale rows along the body (Fig. 3) This feature has been
consistently used as the main character to distinguish two species in the western south Atlantic. For the
purpose of comparison, the samples were grouped as indicated in the section “Material and methods”.
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THE IDENTITY OF THE LEBRANCHE MULLET
Comparing only the sample from Venezuela with that of Uruguay and Argentina, the respective ranges and
medians are significantly different, with little overlap (p = 0.008). However, the same values for the
intermediate samples from northeastern Brazil, southeastern Brazil and southern Brazil bridge the gap. In
other words there are no significant differences between the samples from Venezuela and northeastern Brazil
(p = 0.118), northeastern Brazil and southeastern Brazil (p = 0.215), southeastern Brazil and south Brazil (p =
0.205) and south Brazil and Uruguay and Argentina (p = 0.546). A pattern of latitudinal variation probably
associated with a North to South latitudinal decrease in coastal water temperature in the western south
Atlantic is indicated instead of sharp differences that would justify the recognition of more than one species.
TABLE 1. Morphometrics of Mugil liza. Standard length expressed in mm; measurements through head length are
percentages of standard length; last seven entries are percentages of head length.
The question of identifying the species of lebranche mullet in the Atlantic coast of South America was
never addressed properly either because less than adequate number of specimens were available (Menezes,
1983; Menezes and Figueiredo, 1985; Thomson, 1997; Harrison, 2003) or only a section of the distribution
area of occurrence was considered (Cousseau et al., 2005; Castro et al., 2008: Heras et al., 2009; Vazzoler et
al. 1990). The presence of Mugil liza, ranging from the Caribbean sea region to the eastern coast of South
America was never questioned, but its co-occurrence with a second species raised some questions. If M.
cephalus is distributed from the West Indies to Argentina (Harrison, 2003) it would be sympatric with M. liza,
for example in coastal waters of northeastern Brazil and those species should have been collected together
during any survey. However, samples from this area revealed specimens with a unimodal distribution for
number of transverse scale rows ranging from 32 to 37, including thus values within the total range of the
samples from Venezuela (Fig. 3). Although the sample from Venezuela is represented by few specimens, data
from the literature (Aguilera, 1998, Bencomo et al., 1997, and Marin, 2000) confirm the lower values
obtained herein. Examining 1,072 specimens from estuarine waters in the region of Cananéia on the coast of
the São Paulo state, Brazil, Scorvo Filho et al. (1988) found that the number of transverse scale rows ranged
from 32 to 41 and a single distribution mode was obtained. Vazzoler et al. (1990) obtained the same values for
1,152 specimens from the same region and additionally did not detect any variation among specimens
analyzed via electrophoretic patterns of eye-lens general proteins.
Characters nrange mean SD
Standard length 155 24.0–412.0 179.3
Body depth 132 22.0–032.5 026.0 2.8
Snout to dorsal-fin origin 156 70.0–079.0 074.7 1.6
Snout to pectoral-fin origin 156 25.3–032.7 028.2 1.9
Snout to pelvic-fin origin 156 35.2–045.3 040.6 2.1
Snout to anal-fin origin 156 69.0–074.7 072.5 1.4
Caudal peduncle depth 156 09.0–011.8 010.6 0.5
Pectoral-fin length 155 16.4–021.7 018.5 1.1
Pelvic-fin length 155 13.6–019.1 016.4 1.0
Head length 155 24.0–030.8 027.3 1.9
Head width 155 60.0–071.7 066.6 2.5
Head depth 155 60.0–070.0 064.8 2.8
Horizontal orbital diameter 153 16.5–026.6 022.0 1.9
Snout length 155 20.0–025.5 022.3 1.3
Least interorbital width 155 32.5–038.8 035.7 1.6
Upper jaw length 155 27.0–032.0 029.6 1.1
Mouth width 155 20.0–025.0 023.4 1.0
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FIGURE 2. Regression of number of gill rakers versus standard length for specimens of Mugil liza within the range of
the species.
FIGURE 3. Comparative plots of number of transverse scale rows for population samples of Mugil liza within the range
of the species.
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THE IDENTITY OF THE LEBRANCHE MULLET
FIGURE 4. Comparative plots of number of transverse scale rows between Mugil liza and M. cephalus.
FIGURE 5. Comparative plots of number of horizontal scale rows between Mugil liza and M. cephalus.
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FIGURE 6. Comparative plots of number of circumpeduncular scale rows between Mugil liza and M. cephalus.
In light of the sum of evidence discussed herein we are convinced that a single species of lebranche mullet
should be recognized in the western south Atlantic and the oldest available name that should be used is Mugil
liza Valenciennes, 1836. Nonetheless, two studies based on the analysis of DNA sequences of mitochondrial
genes reached different conclusions. Fraga et al. (2007) mentioned that their results were “elucidative for the
taxonomy of Brazilian mugilids” and concluded that “Mugil liza and Mugil platanus should be treated as a
single species or even populations of M. cephalus”, but did not include specimens from across the entire
distribution of the group. Furthermore, for comparative purposes they used specimens from Chile identified as
Mugil cephalus. These might represent a species distinct from M. cephalus from the Mediterranean, where
type specimens of the species apparently originated. Heras et al. (2009) on the other hand, suggested that
Mugil platanus and M. liza should be considered a single species, and in spite of examining specimens only
from a limited area of distribution, they concluded that they form a continuum and are closely related but
different from M. cephalus “which itself appears to comprise a grouping of multiple and closely related
species.” Because they did not use taxonomic procedures nor apply the rules of nomenclature the decision
about the species name that should be used was not followed.
Morphometric data failed to provide any significant differences between Mugil liza and M. cephalus when
compared through regression analysis, but specimens of the latter are too small (MNCN 065674-65700, SL
33.5–57.0 mm) for a suitable comparison. Nonetheless, comparison of meristic data pertaining to scale
numbers (all samples of M. liza combined) revealed very significant statistical differences (p < 0.001) in
number of transverse scale rows, horizontal scale rows, and circumpeduncular scale rows (Figures 4, 5, and 6)
clearly indicating they are different species. These and other differences correlate with molecular data
showing differences between the two species as noted by Castro et al. (2008) and Heras et al. (2009). Thus the
name M. cephalus cannot be used for the species occurring in the western south Atlantic. Records of M.
cephalus in the West Indies are extremely rare and its occurrence in the area was even questioned by Harrison
(2007: 1079). The presence of M. liza north of the West Indies (Menezes 1983, 2003; Menezes & Figueiredo
1985) and together with M. cephalus (Rivas, 1980; McEachran & Fechhelm, 1998; Greenfield & Thomerson,
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THE IDENTITY OF THE LEBRANCHE MULLET
1997; Castro Aguirre et al., 1999; Smith Vaniz et al., 1999) needs to be investigated. Specimens from this
region might simply represent a population of M. liza in waters colder than in the West Indies.
Acknowledgments
We are grateful to Marc Puigcerver and Jesús Matallanas, Universidad Autonoma de Barcelona, who provided
assistance in acquiring specimens of Mugil cephalus from museums in Spain and especially to Gema Solís
collection manager, Museo Nacional de Ciencias Naturales, Madrid, Spain, who kindly loaned us specimens
of the species for study. Thanks are also due to José Lima de Figueiredo (MZUSP) for reading and criticizing
the manuscript and Richard P. Vari National Museum of Natural History, Smithsonian Institution,
Washington, D.C., U.S.A., who also read and offered suggestions to improve the manuscript and the English
text. José Birindelli, Leandro Sousa, André Netto-Ferreira, Eduardo G. Baena, and Rogério R. Silva (MZUSP)
provided assistance in the use of computer programs. Leandro Sousa also prepared figure 1.
Literature cited
Aguilera, O. (1998) Los peces marinos del occidente de Venezuela. Acta Biológica Venezuelica, (18), 43–47.
Ayres, M., Ayres Jr., M, Ayres, D.L. & dos Santos, A. de A.S. (2007) BioEstat. Aplicações estatísticas nas áreas das
ciências biomédicas. Belém, Pará: xvii + 359 pp.
Bencomo, E.J.G., Borges, J.A. & Castillo, E.C. (1997) Ictiofauna del sector San Carlos del Lago de Maracaibo,
Venezuela. Boletin del Centro de Investigaciones Biológicas, Universidad del Zulia, 31(20), 151–179.
Camargo, M. & Isaac, V. (2001) Os peixes estuarinos da região norte do Brasil: lista de espécies e considerações sobre
sua distribuição geográfica. Boletim do Museu Paraense Emílio Goeldi, Nova Série, Zoologia, 17(2), 133–157.
Castro, G.M., Heras, S., Cousseau, M.B. & Roldán, M.I. (2007) Assessing species validity of Mugil platanus Günther,
1880 in relation to Mugil cephalus Linnaeus, 1758 (Actinopterygii). Italian Journal of Zoology, 75(3), 319–325.
Cervigón, F. (1992) Tiburones, peces batoideos y peces óseos. Pp. 163–465. In: Cervigón, F., R. Ciprioni, W. Fischer, L.
Garibaldi, M. Hendrickx, A.J. Lemus, R. Marques, J.H. Poutiers, G. Robaina & B. Rodriguez. Fichas FAO de
identificación de especies para los fines de pesca. Guia de campo de las espécies comerciales, marinas y de águas
salobres de la costa septentrional de Sur América, FAO, Rome, 513 p.
Cervigón, F. (1993) Los peces marinos de Venezuela. Fundación Científica Los Roques, 2, 1–499.
Cousseau, M.B., Castro, M.G., Figueroa, D.E. & Gosztonyi, A.E. (2005) Does Mugil liza Valenciennes 1836 (Teleostei:
Mugiliformes) occur in Argentinean waters? Revista de Biologia Marina y Oceanografia, 40(2), 133–140.
Eiras-Stofella, D.R., Chavert-Almeida Fanta, P.E. & Vianna, A.C.C. (2001) Surface ultrastructure of the gills of the
mullets Mugil curema, M. lisa and M. platanus (Mugilidae, Pisces). Journal of Morphology, 247, 122–133.
Eschmeyer, W.N. & Fricke, R. (eds). (2009) Catalog of Fishes electronic version (9 September 2009)http://
research.calacademy.org/ichthyology/catalog/fishcatmain.asp
Evermann, B.W. & Marsh, M.C. (1902) The fishes of Puerto Rico. Buletin of the United States Fish Commission, 20, 49–
350.
Fowler, H.W. (1903) New and little known Mugilidae and Sphyraenidae. Proceedings of the Academy of Natural
Sciences of Philadelphia, 55, 743–752.
Fowler, H.W. (1936) The marine fishes of West Africa. Bulletin of the American Museum of Natural History, 70(1), 1–
605.
Fraga, E.H., Schneider Nirchio, M., Santa Brígida, E., Rodrigues Filho, L.F. & Sampaio, I. (2007) Molecular
phylogenetic analysis of mullets (Mugilidae) based on two mitochodrial genes. Journal of Applied Ichthyology,
23(5), 598–604.
Greenfield, D.W. & Thomerson, J.E. (1997) Fishes of the continental waters of Belize. 1997: iii–xxiii+1–311.
Günther, A. (1880) A contribution to the knowledge of the fish fauna of the Río de la Plata. Annals and Magazine of
Natural History, (5)7, 7–13.
Harrison, I.J. (2003) Mugilidae, pp 1071–1085. In: Carpenter, K.E. (ed.). The living marine resources of the Western
Central Atlantic. FAO species identification guide for fishery purpose, 2: vii+602–1373.
Heras, S., Roldán, M.I. & Castro, M.G. (2009) Molecular phylogeny of Mugilidae fishes revised. Reviews in Fish
Biology and Fisheries, 19, 217–231.
Jordan, D.S. & Evermann, B.W. (1896) A descriptive catalogue of the fishes of North and Central America, part I.
Bulletin of the United States National Museum, 47, 1–1240.
Jordan, D.S. & Swan, J. (1884) A review of the American species of marine Mugilidae. Proceedings of the United States
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National Museum, 7, 261–275.
Keith, P., Le Ball, P.Y. & Planquette, P. (2000) Atlas des poisons d’eau douce de Guyane. Tome 2 – fascicule I.
Batrachoidiformes, Synbranchiformes, Perciformes, Pleuronectiformes, Tetraodontiformes. 1–286.
López, H.L., Morgan, C.C. & Montenegro, M.J. (2002) Ichthyological ecoregions of Argentina. ProBiota, FCNyM,
UNLP. Series Technica y Didactica, (1), 1–68.
López, H.L., Miquelarena, A.M. & Menni, R.C. (2003) Lista comentada de los peces continentales de la Argentina.
ProBiota. Serie Técnica y Didactica, (5), 1–85.
McEachran, J.D. & Fechhelm, J.D. (1998) Fishes of the Gulf of Mexico, Volume 1: Myxiniformes to Gasterosteiformes.
University of Texas Press, Austin, 1–112.
Robins, C.R. & Ray, G.C. (1986) A field guide to Atlantic coast fishes of North America. The Peterson Field Guide
Series. Boston: iii–xi, 1–354.
Marin, G. (2000) Ichthyofauna and fisheries of the Unare Lagoon, Estado Anzoategui, Venezuela. Acta Biologica
Venezuelica, 20(30), 61–92.
Meek, S.E. & Hildebrand, S.F. (1923) The marine fishes of Panama. Pt. 1. Publications of the Field Museum of Natural
History, 10, 333–338.
Menezes, N.A. (1983) Guia prático para o conhecimento e identificação das tainhas e paratis (Pisces: Mugilidae) do
litoral brasileiro. Revista Brasileira de Zoologia, 2(1), 1–12.
Menezes, N.A. (2003) Mugilidae, p. 65. In: Menezes, N.A., Buckup, J.L. Figueiredo, J.L. & Moura, R.L. de (eds.),
Catálogo das espécies de peixes marinhos do Brasil. São Paulo, Museu de Zoologia da Universidade de São Paulo,
160 p.
Menezes, N.A. & Figueiredo, J.L. (1985) Manual de peixes marinhos do sudeste do Brasil. V. Teleostei (4). São Paulo,
Museu de Zoologia da Universidade de São Paulo, 105 p.
Menni, R.C. (2004) Peces y ambientes en la Argentina. Monografias del Museo Argentino de Ciências Naturales,
Buenos Aires, (5), 1–316.
Poey, F. (1860) Poissons de Cuba, espèces nouvelles. Pp. 97–336. In: Memorias sobre la Historia natural de la isla de
Cuba, 2 Barcina: Havana.
Ribeiro, A. de M. (1915) Fauna Brasiliense. Peixes. V (Eleutherobranchios Aspirophoros) Physoclisti. Archivos do
Museu nacional, Rio de Janeiro, 17, 679 p.
Ringuelet, R.A. & Aramburu, R.A. (1960) Peces marinos de la Republica Argentina. Agro-Publicacion Técnica, (5), 1–
141.
Ringuelet, R.A. & Aramburu, R.A (1961) Peces argentinos de agua dulce. Agro-Publicacion Técnica, (7), 1–98.
Rivas, L.R. (1980) Synopsis of knowledge on the taxonomy, biology, distribution, and fishery of the Gulf of México
mullets (Pisces: Mugilidae), in: Flandorfer, M and L. Skupien (eds.) “Proceedings” of a workshop for potential
fishery resources of the northern Gulf of Mexico, pp. 34–53, New Orleans, LA: Mississippi-Alabama Sea Grant
Consortium. Publ. MASGP-80-012.
Robins, C.R. &. Ray, G.C. (1986) A field guide to Atlantic coast fishes of North America. The Peterson Field Guide
Series. Boston, 1986: iii–xi, 1–354
Schultz, L.P. (1949) A further contribution to the ichthyology of Venezuela. Proceedings of the United States National
Museum, 99, 1–211.
Scorvo Filho, J.D., Serralheiro, P.C.S., Paiva, P. de., Godinho, H.M., Almeidsa Dias, E.R., Oliveira, I.R. & Ito, K. (1988)
Variação do número de escamas laterais em tainha Mugil sp na região lagunar de Cananéia, S.P. In: Resumos do
Congresso Brasileiro de Zoologia, SBZ/UFPR, Curitiba, PR, p. 404.
Smith, C.L., Tyler, J.C., Davis, W.P., Jones, R.S., Smith, D.G. & Baldwin, C.C. (2003) Fishes of the Pelican Cays, Belize.
Atol Research Bulletin No. 497: ii+1–88.
Smith-Vaniz, W.F., Collette, B.B. & Luckhurst, B.E. (1999) Fishes of Bermuda: history, zoogeography, annotated
checklist, and identification keys. American Society of Ichthyologists and Herpetologists, Special Publication No. 4:
i–x+1–424, 1–12.
Spix, J.B. von & Agassiz, L. (1829–31) Selecta genera et species piscium quos in itinere per Brasiliam annos
MDCCCXVII-MDCCCXX jussu et auspiciis Maximiliani Josephi I … colleget et pingendso curavit Dr. J.B. Spix
… :Monachi. Part 1: i–xvi+i–ii+1–82, Pts. 1–48; Part 2: 83–138, pls. 49–101.
Steindachner, F. (1876) Ichthyologische Beitrage. 3. Sitzungsberichte der Akademie der Wissenschaften, Wien, 72, 29–
96.
Steindachner, F. (1879) Zur fisch-fauna des Magdalenen-Stromes. Denkschrift der Akademie der Wissenschaften, Wien,
60, 19–78.
Thomson, J.M. (1978) Mugilidae. In: W. Fischer (ed.). FAO species identification sheets for fishery purposes. Western
Central Atlantic (fishing area 31). Vols. 1–7, Rome, FAO, unpaginated.
Thomson, J.M. (1997) The Mugilidae of the world. Memoirs of the Queensland Museum, 41(3), 457–562.
Vazzoler, A.E.A., Lizama, M.A.P. & Cohen, M.R.G (1990) Caracterização bioquímica das tainhas (Mugil sp) da região
estuarino-lagunar de Cananéia, São Paulo, Brasil. Boletim do Instituto de Pesca, 17, 37–52.
