A global review of the cosmopolitan flathead mullet Mugil cephalus Linnaeus 1758 (Teleostei: Mugilidae), with emphasis on the biology, genetics, ecology and fisheries aspects of this apparent species complex

Abstract

This study reviews published information on Mugil cephalus from around the world, with recent genetic studies indicating that the flathead mullet may indeed be a species complex. Disciplines that are covered range from the taxonomy, genetics and systematics, through a variety of biological and ecological attributes, to biomarker and fisheries studies. The eurytopic nature of M. cephalus is emphasized, with the migratory life history covering a succession of very different aquatic environments (e.g. rivers, estuaries, coastal lakes/lagoons, marine littoral, open ocean), each of which is occupied for varying lengths of time, depending on the population characteristics within a region and the life-history stage of the species. Interpretation of these movements over time has been greatly enhanced by the use of otolith micro-chemistry which has enabled scientists to map out the different habitats occupied by individual fish at the different life stages. The range of physico-chemical attributes within these environments necessitates a wide tolerance to differing conditions, especially with regard to salinity, turbidity, dissolved oxygen and temperature, all of which are discussed in this review. The importance of M. cephalus to the ecological functioning of coastal systems is emphasized, as well as the pivotal role that this species fulfills in fisheries in some parts of the world. The parasites range from internal trematode and cestode infestations, to external branchyuran and copepod parasites, which use M. cephalus as either an intermediate or final host. The value of the flathead mullet as a biomarker for the monitoring of the health of coastal habitats is discussed, as well as its potential as an indicator or sentinel species for certain ecosystems.

Full text

1 23
Reviews in Fish Biology and Fisheries
ISSN 0960-3166
Volume 22
Number 3
Rev Fish Biol Fisheries (2012) 22:641-681
DOI 10.1007/s11160-012-9263-9
A global review of the cosmopolitan
flathead mullet Mugil cephalus Linnaeus
1758 (Teleostei: Mugilidae), with emphasis
on the biology, genetics, ecology and
fisheries aspects of this apparent species
complex
A.K.Whitfield, J.Panfili & J.-D.Durand

1 23
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REVIEWS
A global review of the cosmopolitan flathead mullet Mugil
cephalus Linnaeus 1758 (Teleostei: Mugilidae),
with emphasis on the biology, genetics, ecology and fisheries
aspects of this apparent species complex
A. K. Whitfield •J. Panfili •J.-D. Durand
Received: 2 February 2012 / Accepted: 6 March 2012 / Published online: 27 March 2012
ÓSpringer Science+Business Media B.V. 2012
Abstract This study reviews published information
on Mugil cephalus from around the world, with recent
genetic studies indicating that the flathead mullet may
indeed be a species complex. Disciplines that are
covered range from the taxonomy, genetics and
systematics, through a variety of biological and
ecological attributes, to biomarker and fisheries stud-
ies. The eurytopic nature of M. cephalus is empha-
sized, with the migratory life history covering a
succession of very different aquatic environments (e.g.
rivers, estuaries, coastal lakes/lagoons, marine littoral,
open ocean), each of which is occupied for varying
lengths of time, depending on the population charac-
teristics within a region and the life-history stage of the
species. Interpretation of these movements over time
has been greatly enhanced by the use of otolith micro-
chemistry which has enabled scientists to map out the
different habitats occupied by individual fish at the
different life stages. The range of physico-chemical
attributes within these environments necessitates a
wide tolerance to differing conditions, especially with
regard to salinity, turbidity, dissolved oxygen and
temperature, all of which are discussed in this review.
The importance of M. cephalus to the ecological
functioning of coastal systems is emphasized, as well
as the pivotal role that this species fulfills in fisheries in
some parts of the world. The parasites range from
internal trematode and cestode infestations, to
external branchyuran and copepod parasites, which
use M. cephalus as either an intermediate or final host.
The value of the flathead mullet as a biomarker for
the monitoring of the health of coastal habitats is
discussed, as well as its potential as an indicator or
sentinel species for certain ecosystems.
Keywords M. cephalus Genetics Biology 
Ecology Life-history Fisheries
Introduction
The flathead mullet Mugil cephalus L. (Fig. 1)is
cosmopolitan, occurring in tropical, subtropical and
temperate coastal waters in all the world’s major
oceans (Briggs 1960; Thomson 1966), mainly
between latitudes 42°N and 42°S (Fig. 2). This species
occupies a wide variety of marine, estuarine and
A. K. Whitfield (&)
South African Institute for Aquatic Biodiversity (SAIAB),
Private Bag 1015, Grahamstown 6140, South Africa
e-mail: a.whitfield@saiab.ac.za
J. Panfili
Institut de Recherche pour le De
´veloppement (IRD),
UMR 5119 ECOSYM, LABEP-AO, BP 1386,
18524 Dakar, Senegal
J.-D. Durand
Institut de Recherche pour le De
´veloppement (IRD),
UMR 5119 ECOSYM, Bat.24 Cc.093, Universite
´
Montpellier 2, Place E. Bataillon, 34095 Montpellier
Cedex 5, France
123
Rev Fish Biol Fisheries (2012) 22:641–681
DOI 10.1007/s11160-012-9263-9
Author's personal copy

freshwater environments but spawning occurs in the
sea (Thomson 1955; Iba
´n
˜ez and Gutie
´rrez-Benı
´tez
2004). As expected with the above distribution
pattern, M. cephalus is a strongly euryhaline species
capable of living in waters ranging from fresh to
hyperhaline (Wallace 1975a; Cardona 2006; Young
and Potter 2002). The flathead mullet is also found in
both clear and turbid areas, sandy and muddy habitats,
and can survive in waters with a wide range of
dissolved oxygen levels (Thomson 1963; Hoese 1985).
The above eurytopic characteristics, together with its
foraging at the base of the food web, enable this
species to be abundant and attain a high biomass in
many parts of its range. These same characteristics
have meant that it is a popular fishery and aquaculture
species, especially in countries such as Greece and
Taiwan where it is of considerable commercial value
(Maitland and Herdson 2009). Indeed the demand for
mullet roe in many parts of the world has grown
considerably in recent decades and elevated the status
of grey mullet to be being called ‘‘grey gold’’ by
fishermen (Hung and Shaw 2006).
Mugil cephalus is the most widespread species
among the family Mugilidae, which comprises a total
of 20 genera and 70 valid species (11 of which belong
the genus Mugil) (Eschmeyer and Fricke 2011). Despite
its global spread in both hemispheres, M. cephalus
has a discontinuous distribution and has been success-
fully introduced to areas where it did not occur
naturally, e.g. the Caspian Sea (Whitehead et al. 1986).
Fig. 1 The flathead mullet
M. cephalus Linnaeus 1758
(drawing of a 17 cm SL
specimen from South Africa
by Dave Voorvelt)
Fig. 2 Global distribution records of M. cephalus (modified from http://www.fishbase.org)
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Given its wide distribution, and similar morphometric
characteristics to closely related mugilids, it is perhaps
not surprising that it has been confused with species
such as Mugil liza (Menezes et al. 2010) which is now
regarded as a synonym of Mugil platanus (Fraga et al.
2007). Indeed, there is increasing evidence that
M. cephalus is part of a species complex (Shen et al. 2011)
and there is a distinct possibility that at least 14 Mugil
species are part of that complex (Durand et al. 2012).
Perhaps another indication of the confusion regarding
M. cephalus is the choice of the common name ‘‘sea
mullet’’ to describe this fish in some parts of the world.
In most areas diadromous migrations are a common
feature for this species (e.g. Wallace 1975a; Bok 1979)
and the name sea mullet is therefore inappropriate.
Considerable research has been conducted on
M. cephalus in both hemispheres, much of which is
quoted in this review. These studies, which have
expanded rapidly since the 1950s (e.g. Idyll and Sutton
1952; Cadenat 1954; Thomson 1955; Tung 1959),
covering everything from the basic biology to fisheries
aspects of the species. However, they have generally
been poorly co-ordinated and often do not take full
cognisance of published work on the same topic from
other parts of the world. A broad review of information
on grey mullet, including M. cephalus, was produced
by Thomson (1966) and a brief review of grey mullet,
including M. cephalus, was published more than three
decades ago (de Silva 1980).
The purpose of the current review is to draw on
more than half a century of literature on M. cephalus
and to give a global perspective that will provide a
benchmark of our current knowledge and facilitate a
more directed focus on future research needs associ-
ated with this ubiquitous and ecologically important
fish species. We also wish to attract attention to the
fact that mullet fisheries are just as susceptible to
overfishing as other marine fish species, and that the
recent collapse of M. cephalus stocks in Taiwanese
coastal waters is an early warning sign in this regard
(Hung and Shaw 2006).
Taxonomy, systematics and genetics
Taxonomy and systematics
Mugil cephalus belongs to the family Mugilidae
(Order: Mugiliformes) which are Actinopterygian
teleosts. It is a robust grey mullet with a broad head
and a thick, soft, transparent adipose eyelid that is well
developed and covers most of the eye, except for a
vertical elliptical shaped opening (Fig. 1) that allows
most of the pupil direct visual contact with the
surrounding medium. Indeed, the adipose eye tissue
in M. cephalus appears to be more developed than any
of the other mugilid species.
The upper lip is thin and there is an absence of
papilli that are found on the upper lips in some grey
mullet, e.g. Crenimugil crenilabis. The labial setiform
teeth in the upper jaw are small, straight and dense,
and can be used to identify the juveniles of this species
at an early stage of development (van der Elst and
Wallace 1976; Trape 2009). Other scientists have used
melanophore patterns on the ventral side of the head to
identify mugilid fry, with M. cephalus having only a
lightly pigmented ventro-opercular and gular region
(Minos et al. 2002). The same authors have high-
lighted the acute angle of the dentary symphysis as a
distinguishing feature in this species. The cleft mouth
of M. cephalus ends below the posterior nostril
(Whitehead et al. 1986) and the preorbital bone is
shorter than the upper jaw (Maitland and Herdson
2009). The underside of the lower jaw has a broad
midline space at the front.
Dorsal and anal fin spine and ray counts cannot be
used to distinguish M. cephalus from other mugilids
because of the close overlap in counts within the family
(Smith and Heemstra 1986). However, a large axillary
scale is located at the base of each of the short pectoral
fins and is approximately one third of the fin length
(Whitehead et al. 1986). The scales are a typical
percomorph type (Thomson 1966) and are cycloid in
the early juveniles becoming ctenoid by the end of the
first year. The lateral series of scales can range between
36 and 45 for M. cephalus (Whitehead et al. 1986)
which overlaps with the counts for some of the other
mugilid species (Smith and Heemstra 1986). Although
geometricand morphometric analysesof mugilid scales,
or the ctenii of the scales, can be used to distinguish
species (Iba
´n
˜ez and Gallardo-Cabello 2005;Iba
´n
˜ez et al.
2007), these methods are seldom used in practise.
The scales on the flanks of M. cephalus are usually
streaked and can give an overall impression of stripes
(Figs. 1,3), hence the common name of striped mullet
in some regions of the world. Once again, this is a poor
choice of common name because there are other
mugilid species with much stronger stripe patterns on
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the flanks, e.g. Liza tricuspidens. The back of M. cephalus
is usually an olive green colour and the belly is white.
A blue colouration is sometimes visible at the base of the
pectoral fins (Fig. 3a).
An indication of the taxonomic confusion around
M. cephalus is the occurrence of subspecies within this
apparent species complex (Table 1). One example is
provided by Mugil ashanteensis Bleeker, 1863, which
was originally described as a separate species but
subsequently regarded by Cadenat (1954) and later
Trewavas and Ingham (1972) as a subspecies of
M. cephalus. This subspecies has a mainly West African
distribution with yellow in the anal fin and sometimes
in the caudal fin as well (Fig. 3b). Further examples of
variation within M. cephalus are the morphometric
differences that exist within its distributional range
(Corti and Crosetti 1996) and led to separate species
being described for Mugil japonicus Temminck &
Schlegel 1845 from Japanese waters, Mugil peruanus
Hildebrand 1946 from Peru, Mugil galapagensis
Ebeling 1961 from the Galapagos, and numerous
others examples, all of which have been synonymised
with M. cephalus (Table 1). Despite the above appar-
ent external variation within the species, M. cephalus
otoliths collected from all over the world appear to
belong to the same species and this is reflected in both
the similar otolith shape and the relationship between
individual fish fork length and otolith diameter, as well
as other otolith dimensions (Panfili et al. 2007).
Genetics
The unusually wide distributional range for a coastal
species such as M. cephalus, with limited dispersion
Fig. 3 Pictures of aM. cephalus (40 cm TL) from Okinawa
Island, Japan (photographer: J. Randall), showing the ‘normal’
anal fin coloration and bM. cephalus (31 cm TL) from Dakar,
Senegal (photographer: J.-D. Durand), showing the bright
yellow anal fin that is characteristic of certain M. cephalus
populations in the central and west African regions
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abilities across large open oceans, has given rise to
questions regarding its systematic status. Initial stud-
ies tried to estimate broad genetic differences between
remote populations and thus provide information
about the level of gene flow and dispersal history of
the species. While the cytogenetic approach showed
that scattered populations of M. cephalus had the same
chromosome number and morphology, with no
changes detected in heterochromatin and NORs (Rossi
et al. 1996), genetic studies have identified popula-
tions with ‘‘private’’ haplotypes and specific allozy-
mic/enzymatic allele frequencies (Fig. 4a, b) (Crosetti
et al. 1993,1994; Rossi et al. 1998a,b; Rocha-Olivares
et al. 2000; Fraga et al. 2007; Heras et al. 2009;
Livi et al. 2011). Mugil cephalus appears to consist of
highly isolated populations characterised by specific
mitochondrial lineages, with each of these lineages
(Fig. 5) being highly divergent and sometimes
exceeding the level of intraspecific diversity expected
within a species (Rocha Olivares et al. 2000,2005).
Some studies have demonstrated that M. cephalus is
monophyletic, which is in agreement with the pres-
ence of a single species worldwide (Rossi et al. 1998a;
Livi et al. 2011), but others stress the paraphyly of
M. cephalus with M. liza (Fraga et al. 2007; Heras et al.
2009). Interestingly, Fraga et al. (2007) concluded that
Table 1 A list of mugilid species that were synonymised with M. cephalus Linnaeus, 1758 by both Thomson (1997) and Eschmeyer
and Fricke (2011)
Species Author Location
Mugil albula Linnaeus, 1766 Charleston, USA
Mugil ashanteensis Bleeker, 1863 Ashantee, Ghana
Mugil borbonicus Valenciennes, 1836 Bourbon River, Re
´union, France
Mugil catalarum Whitley, 1951 Bogny River, New Caledonia
Mugil cephalotus Valenciennes, 1836 Malabar and Pondicherry, India
Mugil ciliilabis Valenciennes, 1836 Lima, Peru
Mugil constantiae Valenciennes, 1836 Constance, Europe
Mugil dobula Gu
¨nther, 1861 Perth, Australia
Mugil galapagensis Ebeling, 1961 Galapagos Islands, Ecuador
Mugil gelatinosus Klunzinger, 1872 Murray River, Australia
Mugil grandis Castelnau, 1875 New South Wales, Australia
Mugil hypselosoma Ogilby, 1897 New South Wales, Australia
Mugil japonicus Temminck & Schlegel, 1845 Nagasaki, Japan
Mugil lineatus Valenciennes, 1836 New York, USA
Mugil marginalis De Vis, 1885 Brisbane, Australia
Mugil mexicanus Steindachner, 1876 Acapulco, Mexico
Mugil mu
¨elleri Klunzinger, 1879 King George Sound, Australia
Mugil ou
¨rForsska
˚l, 1775 Red Sea
Mugil peruanus Hildebrand, 1946 Independencia Bay, Peru
Mugil plumieri Bloch, 1794 St Vincent
Mugil provensalis Risso, 1810 Var River, Nice, France
Mugil rammelsbergii Tschudi, 1845 San Lorenzo Island, Peru
Mugil tongae Gu
¨nther, 1880 Tongatabu, Kingdom of Tonga
Myxus barnardi Gilchrist & Thompson, 1914 Durban, South Africa
Myxus caecutiens Gu
¨nther, 1876 Rodrigues Island, Mauritius
Myxus flavus Mohr, 1927 Mazatla
´n, Mexico
Myxus lepidopterus Mohr, 1927 Peru
Myxus niger Mohr, 1927 Lima, Peru
Myxus superficialis Klunzinger, 1870 Al-Qusair, Egypt
Myxus tincoides Mohr, 1927 Cape Hatteras, USA
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M. liza may be a population of the globally distributed
M. cephalus due to the limited level of genetic
divergence (0–1 % for 16S and 2–5 % for cytochrome
b) between the two species. In contrast, Heras et al.
(2009) considered that the absence of shared haplo-
types among M. cephalus lineages and M. liza support
the species complex assumption. However, the bio-
logical species concept cannot be applied due to the
allopatric distributional range of M. liza and
M. cephalus preventing the testing of potential inter-
breeding between these populations (Gilbert 1993;
Menezes et al. 2010).
North West and South West Pacific M. cephalus
populations have two to three different lineages
(Huang et al. 2001; Jamandre et al. 2009; Ke et al.
2009; Liu et al. 2009,2010; Livi et al. 2011; Shen et al.
2011; Sun et al. 2012, Durand et al. 2012) whereas
elsewhere only one lineage has been recorded. This
constitutes an interesting case study to determine the
evolutionary significance of these lineages, i.e. intra-
specific diversity versus the cryptic species concept.
Shen et al. (2011), using two classes of molecular
markers, demonstrated that no gene flow occurs among
individuals harboring different mitochondrial lineages
in the North West Pacific which clearly demonstrate
the existence of cryptic species within this region. The
above study sheds new light on the taxonomy of
M. cephalus and points to a species complex rather than
a single cosmopolitan species. However, the evolu-
tionary history of this species complex is still essen-
tially unresolved as no phylogenetic trees have been
constructed that highlight a dispersal history. Never-
theless, according Livi et al. (2011), the centre of origin
of the M. cephalus species complex may indeed be in
the Indo-West Pacific but this conclusion is based
solely on 300 bp from the cytochrome bgene.
Galapagos
Galapagos
Galapagos
Galapagos
Australia (West)
Australia (West)
Australia (West)
Hawaii
Hawaii
Hawaii
Hawaii
USA (East)
USA (East)
USA (East)
USA (East)
South Africa
South Africa, Mozambique
Mauritania
South Africa
South Africa
Australia (East)
Australia (East)
Australia (East)
Mauritania
Mauritania
Mauritania
Mauritania
Egypt
Italy (East)
Italy (East)
Italy
Turkey
Italy (West)
Italy (West), Asov Sea
Italy (West)
Taiwan
Taiwan
Japan, Taiwan
Japan, Taiwan, China
0.16 0.12 0.08 0.04 0.0
6.0 4.5 3.0 1.5 0.0 3.0 2.0 1.0 0.0
outgroups
0.1 substitutions
(a) (b)
(c) (d)
Fig. 4 a UPGMA phenogram for M. cephalus haplotypes based
on Mahalanobis distances (Corti and Crosetti 1996); bUPGMA
phenogram based on restriction polymorphisms using 13
enzymes from the mitochondrial genome (Crosetti et al. 1994);
cPhenogram generated by UPGMA cluster analysis using
unbiased genetic distances estimated from 27 allozyme loci
(Rossi et al. 1998a,b); dMaximum likelihood phylogram of
M. cephalus haplotypes using 300 bp from the cytochrome
bgene (Livi et al. 2011)
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Few studies have investigated the genetic structure
of M. cephalus on a regional scale. With exception of
the North West Pacific, there are only two areas where
comparable information is available, namely the Gulf
of Mexico (Campton and Mahmoudi 1991; Rocha-
Olivares et al. 2000) and the Mediterranean Sea
(Crosetti et al. 1994; Rossi et al. 1998b; Blel et al.
2010; Livi et al. 2011). The above studies highlight the
(a)
(b)
Fig. 5 a Phylogenetic tree
depicting the genetic
diversity within M. cephalus
and M. liza species from
Durand et al. (2012).
Numbers on the branches
are ML bootstrap values for
those above 50 %. Asterisks
indicate nodes with an a
posterior probability from
partitioned Bayesian
analysis of [0.95.
bDistributional range of 14
species within the M.
cephalus complex
considering molecular
phylogenetics results from
South America (Fraga et al.
2007), North West Pacific
(Shen et al. 2011), Durand
et al. (2012), and Genbank
sequences DQ185446 and
FJ384686 from India
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presence of only one mtDNA lineage and absence of
genetic heterogeneity in these two regions. They all
conclude that apparent panmixy in these areas is
consistent with the suspected dispersion abilities of
M. cephalus, especially during the pelagic larval
stages. Another ‘tool’ which may provide an estima-
tion of the dispersive ability of M. cephalus are
microsatellite markers for population genetic studies
at regional and fine scales of distribution (Miggiano
et al. 2005; Andree et al. 2010).
Biology and life history traits
Life-history information
Detailed information on the life-history traits of
M. cephalus on a global scale is generally lacking
(Whitfield 1990). However, continental information is
available from certain regions, e.g. America (Iba
´n
˜ez
and Gallardo-Cabello 1996a; Iba
´n
˜ez et al. 1999;
Nordlie 2000; McDonough et al. 2003; McDonough
and Wenner 2003), Asia (Tung 1981; Chang et al.
2000), Oceania (Smith and Deguara 2003), Europe
(Cardona 2000) and Africa (Bok 1979; Blaber 1987;
Hamza 1999). This information points towards a life-
history cycle that encompasses mainly marine and
estuarine environments but can also extend into
freshwaters (Fig. 6, Wang et al. 2010). Indeed, it has
been recorded penetrating high up into rivers of
Australia, South Africa, USA, India and Israel
(Thomson 1963,1966; Bok 1983) with some speci-
mens located more than 100 km from the coast
(Bok 1979). The versatility of M. cephalus in occupying
varied aquatic environments at different stages of its
life cycle have made it an attractive species to use in
both freshwater aquaculture and mariculture projects
(Tamaru et al. 2005).
In most parts of the world M. cephalus spawns in
the nearshore marine environment (Wallace 1975b),
the egg and early larval stages are spent drifting in
ocean currents, and there is an onshore migration at the
postflexion larval stage which is followed by tempo-
rary occupation of the surf zone as early juveniles
(Strydom and d’Hotman 2005). Schools of these fry
then enter estuaries after about a month at sea
(Hsu et al. 2009) and colonise the entire length of these
systems, sometimes extending into adjoining river
catchments. In some river systems, colonisation of
freshwater areas can be extensive enough for the term
catadromous species to be used in connection with
certain M. cephalus populations (Bok 1979).
The juvenile and sub-adult life stages are spent
mainly in estuarine waters (Lawson and Jimoh 2010)
and adults then emigrate to the sea to spawn (Wallace
1975b). Some adults return to estuaries following
spawning (Whitfield and Blaber 1978a) and others
may remain within the marine environment (Fig. 6).
However, not all M. cephalus populations follow this
pattern and in some parts of the world, especially
where river flow and estuaries are intermittent features
of the coast (e.g. Shark Bay in Australia), this species
conducts its entire life cycle within the marine
environment.
In an attempt to study nursery area associations by
juvenile M. cephalus in Taiwan, otolith trace elemen-
tal composition was used as an indicator of site fidelity
to different estuaries (Wang et al. 2010). Out of twelve
elemental concentrations (Li, Na, Mg, K, Mn, Fe, Ni,
Cu, Zn, Sr, Ba and Pb), five elements (Mn, Ni, Zn, Sr
and Ba) were found to be significantly different for the
fish among estuaries. The canonical discriminant
function using Ca also indicated that 10 of the 12
elemental ratios (i.e. excluding Li:Ca and Cu:Ca)
played a significant role in discriminating the juvenile
mullet among the different estuaries. Among them,
Mn:Ca, Ni:Ca and Zn:Ca contributed approximately
50 % to the first discriminant function, while Ba:Ca
and Sr:Ca contributed 33 % to the second function.
As a result, 84 % of the M. cephalus could be assigned
to their recruited estuaries using their otolith chemical
signatures, thus indicating that the otolith elemental
composition can be used as a natural tag to trace
individuals to their specific nursery areas (Wang et al.
2010).
Global evidence points to a range in life history
traits and gives rise to a number of questions linked to
the adaptive capability of this species in contrasting
situations, e.g. is M. cephalus a generalist capable of
adapting its life history according to local conditions
or is it integrated into a species complex comprising
a number of specialized species with different life
history traits? Evidence from Taiwan strongly sug-
gests that M. cephalus in these waters can be divided
into those that have a freshwater component to their
life cycle and those that do not (Chang et al. 2004a).
However, the full answer to the above question will
require detailed genetic and physiological studies on
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what we perceive to be M. cephalus with different life
histories from around the world.
Physiological tolerances
Tolerance to a wide range of salinities is key to the
global success of M. cephalus as a species. Walsh et al.
(1991) investigated the combined effects of temper-
ature and salinity on embryonic development on
fertilised M. cephalus eggs transferred directly from
the spawning tank and found that normal embryonic
development occurred at water temperatures from 20
to 30 °C and a salinity range of 15–36. Hatching time
was affected by temperature but not by salinity. The
optimum yield of normal larvae from fertilised
M. cephalus eggs occurred at a water temperature of
25 °C and salinity of 36. In contrast, Sylvester et al.
(1975) tested the hatching success of flathead mullet
eggs and larval survival under different salinity
conditions and determined that the optimum salinity
at a water temperature of 20 °C was 30–32 for the eggs
and 26–28 for preflexion larvae.
Fig. 6 Typical life cycle
of the flathead mullet
M. cephalus (see text for
details)
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Early M. cephalus juveniles have a wide salinity
tolerance range and under experimental condi-
tions 20–39 mm SL specimens survived immediate
transfer from seawater to all lower salinities except
freshwater (Nordlie et al. 1982; Walsh et al. 1991).
However, a gradual transfer of fry approximately
28 mm SL into freshwater over a 48 h acclimation
period increased survival rates considerably (Ciccotti
et al. 1994). In addition, M. cephalus fry survived
direct transfer from a salinity of 20 to higher salinities
but mortalities were recorded when placed in salinities
above 45 (Hotos and Vlahos 1998). If gradually
acclimated, these fry were able to tolerate experimen-
tal salinities as high as 126 for up to 4 days but
probably comes as the cost of great physiological
stress to the fish (Khe
´riji et al. 2003). Nevertheless, the
fact that specimens \20 mm SL have been located in
West African waters with a salinity more than double
seawater (Trape et al. 2009) suggests that this species
is highly euryhaline at a very small size. Similarly the
adults of M. cephalus can tolerate salinities from
freshwater to hyperhaline (Simmons 1957, Whitfield
et al. 1981) and were recorded surviving Lake St Lucia
salinities as high as 80 for an extended period
(Whitfield et al. 2006). Despite the above wide ranges,
the preferred salinity range of M. cephalus for
maximum growth performance appears to be in
oligohaline or mesohaline waters (Bok 1983; Cardona
2006). This view is supported by research conducted
by Nordlie and Leffler (1975) who determined that the
energetic cost of the osmoregulatory process in
M. cephalus is high when the external environment
is distinctly hyperosmotic with respect to the blood of
the fish, and negligible when the environmental
concentration is less than that of the blood.
There is strong evidence which indicates that the
chloride cells in the gills of this species adapt rapidly
to any salinity change, e.g. Hossler (1980) showed that
the gill filament epithelium of M. cephalus underwent
ultrastructural changes as early as 6 h after the fish had
been transferred into oligohaline waters and that this
process was completed within 24 h of the salinity
change. Johnson (1972) noted wide variability in the
plasma electrolytes of M. cephalus exposed to a range
of salinities, with individuals from freshwater showing
higher inter-renal activity than those in seawater. The
pituitary/inter-renal system is probably important in
the control of salt and water balances in this species
(Abraham 1971), with evidence also being presented
that prolactin cells are stimulated in freshwater
adapted M. cephalus (Blanc-Livini and Abraham
1970).
The eggs and larvae of M. cephalus are tolerant of
relatively low dissolved oxygen levels, with egg
survival only declining when these levels fall below
5.0 g l
-1
and for larvae when levels decline below
5.4 g l
-1
(Sylvester et al. 1975). Larval oxygen
consumption rates did not vary among salinities
between 10 and 35 but acute water temperature
increases did elicit significant increases in oxygen
consumption by the larvae (Walsh et al. 1989). Some
habitats occupied by M. cephalus can have very low
dissolved oxygen concentrations at times and this
species is then able to jump or gulp air into the upper
pharyngeal chamber as a supplementary oxygen source
(Hoese 1985). Support for this mechanism is provided
by the fact that jumping frequencies are inversely
correlated with dissolved oxygen concentration and
that the pharyngobranchial organ is capable of holding
gas (Hoese 1985). Under hypoxic conditions, the
oxygen content of the swimbladder decreases to a level
proportional to the dissolved oxygen content of the
surrounding water; however it is unlikely that this gas
plays an important role in respiration because of the
very small volume involved (Moore 1970).
The flathead mullet appears tolerant of very low
dissolved oxygen levels and may even switch to
anaerobic metabolism in order to swim through
hypoxic waters (Vagner et al. 2008). However, under
hypoxic conditions, M. cephalus usually move to the
water surface to ventilate water in contact with the
air, an adaptive process known as aquatic surface
respiration (ASR). ASR, hypoxic bradycardia and
branchial hyperventilation are stimulated by fish
chemoreceptors that are sensitive to both low systemic
and low dissolved oxygen levels (Shingles et al. 2005).
Sight of a model avian predator inhibited reflex
hyperventilation under hypoxic conditions but not
ASR. However, the presence of the model predator did
modify M. cephalus behaviour by causing them to
surface under surface objects in order to avoid the
predator. Turbid water abolished the fear response and
effects of the predator on both gill ventilation rates and
timing of ASR (Shingles et al. 2005).
The flathead mullet appears to tolerate a wide range
of water turbidities but generally prefers waters of
10–80 Nephelometric Turbidity Units (Cyrus and
Blaber 1987). The physiology of M. cephalus vision
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was studied by Munz (1958) who found that the
photosensitivity of the retinal pigments of this species
was higher than that of most fishes that had been
tested. He concluded that this sensitivity, which was
near the maximum transmission range of turbid water,
indicated that the vision of M. cephalus is adapted to
murky conditions.
Although detailed temperature tolerance tests do
not appear to have been conducted, Marais (1978) did
establish that M. cephalus can tolerate temperatures
between 13 and 33 °C. However, Bok (1983) recorded
M. cephalus in farm dams where the littoral water
temperature sometimes declined to 6 °C which
matches minimum winter in the Black Sea where this
species also occurs (Apekin and Vilenskaya 1978).
The general distribution indicates a predominantly
warm water species and this is supported by Radovich
(1961) who recorded an absence of M. cephalus from
San Diego Bay when average summer temperatures
were \18 °C. Bok (1983) recorded a significant
movement of M. cephalus from freshwater reaches
of two Eastern Cape rivers into the adjacent estuaries
during winter, and attributed this downstream migra-
tion to an avoidance of lower water temperatures in
the river during winter. Flathead mullet appear to be
susceptible to sudden declines in water temperatures,
especially under freshwater or oligohaline conditions.
This is reflected in the mortalities of M. cephalus in
subtropical Lake St Lucia, South Africa, when water
temperatures suddenly declined to 12 °C and the
salinity was 1 (Blaber and Whitfield 1976).
In northern hemisphere temperate waters M. ceph-
alus will often leave the estuaries and bays in autumn
(Thomson 1966), presumably to move to warmer
waters in the south where they can avoid cold winter
temperatures. At high temperatures M. cephalus
shows increased haemoglobin concentration, increased
haematocrit and mean erythrocyte volume (Cameron
1970), all which suggests an altered blood oxygen
capacity to cope with reduced dissolved oxygen
levels at elevated water temperatures. Another study
(Sylvester 1975) showed that the flathead mullet
exhibited increasing critical thermal maxima towards
midday, declining thereafter, an attribute that has
distinct benefits for this species in tropical waters.
Although this species does occupy certain tropical
coastal areas in most oceans (Fig. 2), it appears to be
most abundant in subtropical and warm temperate
waters (Thomson 1963; Wallace 1975a).
Routine oxygen consumption tests on M. cephalus
at different temperatures and salinities have shown
that both small (10 g) and larger (100 g) specimens
consume more oxygen when temperatures increase
(Marais 1978). In addition, this same study revealed
that M. cephalus oxygen consumption at a salinity of 1
was 16–18 % higher than at a salinity of 35, suggest-
ing that metabolic energy requirements are higher
under stressful salinity conditions that require active
osmoregulation.
Movements and migrations
Mullet are essentially schooling fish and M. cephalus
schools vary in size seasonally, being smaller in the
feeding and post-spawning phases than in the period of
pre-spawning accumulation and migration (Thomson
1955). Flathead mullet tend to spread over tidal flats
and shallow banks of estuaries during the flood tide
and may feed in scattered groups or singly, often
aggregating into schools if alarmed. Light appears to
be necessary for schooling (Breder 1959). According
to Thomson (1955) the shape of a mullet school may
change rapidly from a compact oval mass to a long
narrow band. A swerve by the leading fish is usually
replicated by the followers but a change in direction
may also be initiated from the mid-flanks of the
school. Such a movement may temporarily split the
school which usually reunites rapidly (Thomson
1966).
As has been mentioned, M. cephalus can migrate
along coasts and between continental and open
seawater environments during its life cycle. Juvenile
and sub-adult stages occur mainly in freshwater and/or
brackish waters (estuaries, coastal lakes and lagoons).
The main migration by adult M. cephalus usually
occurs during the breeding season in different parts of
the world (Bacheler et al. 2005), with spawning
migrations corresponding to movements of mature
adults from their coastal feeding areas (often estuaries)
to the open sea (Wallace 1975b; Iba
´n
˜ez and Gutie
´rrez-
Benı
´tez 2004). The downstream migration of
M. cephalus to their spawning grounds can be disrupted
by physical barriers such as weirs, even if these
structures are provided with sluice gates (Russell
1991). Often the shoals of ripe mullet congregate in
the mouth region of estuaries before moving out to sea
(Wallace 1975b), with the seaward spawning migra-
tion being triggered by strong offshore winds in
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Australia (Thomson 1955). Whilst congregating in the
St Lucia Estuary mouth region, M. cephalus exhibit
intensive bouts of leaping into the air as they are
pursued by a variety of predators, including pelicans,
crocodiles and sharks (Whitfield and Blaber 1979a,b).
However, in some parts of the world (e.g. sections of
Mauritania and Australia) flathead mullet adults are
found mainly in sheltered coastal waters and not
estuaries, thus providing support to the proposal that
M. cephalus is a complex of species.
Detailed studies of the marine movements of large
schools of reproductively active M. cephalus is limited
but in some parts of the world (e.g. east and west coasts
of Australia) they have been recorded swimming
against the prevailing ocean currents (Smith and
Deguara 2003). The extent of this migration varies
between geographical areas: migrations of over
700 km have been reported in Australian and Chinese
waters, whereas on the coast of Florida (USA) 90 % of
tagged fish were recaptured within 32 km from the site
of release. It should be noted that Lester et al. (2009)
contest the 740 km distance purported to be covered
by migrating adult M. cephalus in Australia. Along
West African coasts, adult flathead mullet can under-
take reproductive migrations of over 400 km between
Mauritania and the Senegal Estuary (Bernardon and
Vall 2004). The longest confirmed migration by an
adult M. cephalus was 240 km (Idyll and Sutton 1952)
and similar results have been reported for M. cephalus
in North Carolina (Bacheler et al. 2005).
Not all fish follow the same spawning migratory
route; while northward migrations are dominant along
the east and west coasts of Australia (Thomson 1951;
Kesteven 1953) and southward migrations are dom-
inant in North Carolina and West Africa (Thomson
1966; Vall 2004), a small portion of the above
populations were recaptured whilst moving in the
opposite direction to the mainstream migration.
Thomson (1966) went so far as to suggest that the
up-current migration has evolved so that the down-
stream drift will bring the eggs and larvae back to the
estuaries where the adults developed. There does not
appear to be a return migration following spawning on
the Australian east coast, with some spent individuals
entering estuaries up to 725 km to the north of their
starting point and others returning to the estuaries they
left (Thomson 1966). This latter finding is similar to
the situation in Lake St Lucia, South Africa, where a
small portion of the M. cephalus leaving the estuary to
spawn return to the lake as spent individuals (Wallace
1975b).
The M. cephalus spawning migration down the
Lake St Lucia system under hyperhaline and mesoh-
aline conditions has been documented by Wallace
(1975b) and Whitfield and Blaber (1978a) respec-
tively. The above studies eliminated salinity as the
primary directional driver in terms of locating the
marine spawning environment and suggested that the
co-ordination of these migrations is a relatively
complex phenomenon. In addition, the regular mass
jumping behaviour at all hours by adult M. cephalus
during the spawning migration could not be deter-
mined and it was speculated that, in addition to
predator avoidance, it served a role in co-ordinating
the build-up of the shoals of mature adults that were
ready to breed (Whitfield and Blaber 1978a). Other
possible explanations are that the jumping habit
dislodges parasites or that is the result of sheer joie
de vivre ! (Thomson 1966). Of course the reason why
some mugilid species tend to jump and others do not is
unknown.
Following hatching in the natural environment the
M. cephalus larvae exhibit positive phototaxis by
actively swimming towards the water surface.
Although larvae are mainly found near the sea surface
between the coast and the continental slope over the
shelf (Belyanina 1995; Ditty and Shaw 1996), the
presence of larval M. cephalus has been reported from
depths between 10 and 900 m off the south-eastern
United States (Collins and Stender 1989). Larvae do
not seem to undertake diel vertical migrations, yet
some may have changes in activity rates and are most
often recorded in surface waters at night (Collins and
Stender 1989). It is distinctly possible that surface
ocean currents may be an important factor in promot-
ing long distance larval dispersal and thus gene flow
over a wide geographic scale (Campton and Mahmo-
udi 1991; Jamandre et al. 2009; Shen et al. 2011).
Flathead mullet exhibit strong schooling tendencies
from late larval to adult stages (Wallace 1975a), with
mean speeds and mean turning frequencies declining
significantly with increased shoal size (Fitzsimmons
and Warburton 1992). Movements of the early larval
stages (Fig. 6) tend to be dictated by the dominant
oceanographic currents in a region (Chang et al. 2000)
but postflexion larvae can form schools and then
migrate towards coastal waters, as indicated by a
pattern of inshore movement with growth (Collins and
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Stender 1989). In South Africa these late larvae tend to
arrive in the coastal surf zone at about 8–12 mm SL
(Whitfield 1989, Strydom and d’Hotman 2005).
Similar sized specimens have been found in bays
associated with rocky shores (Strydom 2008) but these
postflexion larvae may have been en route to estuarine
nursery areas.
Flathead mullet first enter estuaries when individ-
uals are between 10 and 30 mm SL (Kilby 1955; Grant
and Spain 1975;Tang1975; Fujita et al. 2002;
Strydom and Wooldridge 2005) with most recruitment
occurring at a size of about 15–25 mm SL (Anderson
1958; Wallace 1975a; Bok 1979; De Silva and Silva
1979; Chubb et al. 1981). Pe
´rez-Rufaza et al. (2004)
found that not all M. cephalus recruit into estuaries as
early juveniles; indeed some were recorded as pre-
flexion and flexion larvae in plankton samples from
the Mar Menor lagoon (Eastern Spain) and must have
entered the system at a very early stage of develop-
ment. Johnson and McClendon (1970) claimed that the
presence of 30 M. cephalus fry (28–40 mm SL),
captured 192 km from the mouth of the Colorado
River, was evidence of a freshwater spawning by this
species. However, Whitfield and Blaber (1978a)
contested this view and suggested that these fry were
probably a product of a marine spawning and could
have swam up the river after entering the estuary at
about 10–15 mm SL.
Recruitment into estuaries is usually protracted and
takes place over several months and is linked to the
serial spawning of M. cephalus which also takes place
over an extended period (Wallace 1975b). However,
there is also evidence that successful recruitment by
M. cephalus early juveniles into estuaries that are only
open for short periods each year is possible (Young
et al. 1997). An indication that estuaries are the
preferred nursery habitat for M. cephalus is provided
by the work of Potter et al. (1997) who showed that
this species is found mainly in the highly saline
Leschenault Estuary, despite the close proximity of the
more marine Koombana Bay as an alternative habitat.
Within many estuaries it would appear that the upper
freshwater dominated reaches harbour the highest
densities of 1?,2?and 3?M. cephalus (Bok 1983;
Marais 1983a), with Marais (1981) and Chubb et al.
(1981) suggesting that this distribution pattern may be
linked to riverine inputs of detritus into the upper
reaches. However, in other systems large M. cephalus
may be more abundant in the mouth region, e.g. Blaber
(1977) found that individuals [200 mm SL were
more abundant in the euhaline lower reaches of the
Kosi Estuary where preferred food items such as
Foraminifera were readily available.
In some parts of the world the timing of M. cephalus
recruitment into estuaries and lagoons appears to be
geared towards matching their arrival to the onset of
favourable conditions within the nursery area (Silva
and De Silva 1981). This often coincides with the end
of the rainy season (Wallace 1975a; Payne 1976;De
Silva and Silva 1979) which ensures productive
marine larval habitats and also means that fry are less
likely to be washed out of estuarine nursery areas.
However, in other parts of the world recruitment can
coincide with the onset of winter (Bartulovic
´et al.
2011) or the dry season when an estuary can be
hyperhaline (Trape et al. 2009). Survival of artificially
raised 0?M. cephalus juveniles was highest when
they were released in spring to coincide with maxi-
mum juvenile recruitment of wild stock M. cephalus
into a Hawaiian estuary (Leber et al. 1996,1997). This
suggests that selection pressures are at work to
optimize recruitment periodicity for this species or
that the different recruitment timing described above
is further evidence for a species complex rather than a
single species.
Within an estuary, juvenile M. cephalus are usually
concentrated within the littoral zone, extending into
deeper offshore waters with increase in size (Whitfield
and Blaber 1978a). In Australia, juveniles \50 mm
SL are primarily found in shallow estuarine areas
(Chubb et al. 1981), a pattern repeated in other parts of
the world. Young recruits initially colonise the lower
part of estuaries and often progressively extend their
distribution up these systems towards the head of the
estuary (Bok 1979). The movement of M. cephalus fry
up an estuarine system can be very rapid and may
exceed 0.7 km per day (Whitfield and Blaber 1978a).
Larger juveniles can move even faster, with a swim-
ming speed of 47 cm s
-1
(Nanami 2007) that will
enable the M. cephalus to escape even rapidly moving
piscivorous fishes. It is perhaps significant that
‘‘escape latency’’ is independent of body size in
M. cephalus (Turesson and Domenici 2007), which
means that small fish can respond just as quickly as
large fish to an approaching threat.
The reasons why some early juvenile M. cephalus
are attracted to certain South African river systems and
not others is unknown, e.g. in certain Eastern Cape
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systems large numbers of 0?juveniles colonise the
river systems (Bok 1979) whereas in many Western
Cape and KwaZulu-Natal systems M. cephalus are
confined to estuaries. This colonisation of watersheds
is facilitated by the osmotic regulation capabilities of
M. cephalus postflexion larvae and juveniles which
can tolerate conditions ranging from fresh to hyper-
haline (Trape et al. 2009). The urge to colonise river
catchments may drive juvenile fish to actively over-
come minor physical barriers during their upstream
migration (Kowarsky and Ross 1981; Russell 1991)
but weirs and dams are usually too much of an obstacle
to overcome (Bok 1984a). It has been shown that
juvenile M. cephalus actively avoid polyhaline and
euhaline waters, preferring freshwater and oligohaline
waters in a stratified study carried out in several inland
estuaries of the Island of Menorca (Western Mediter-
ranean) (Cardona 2000). However, occupation of
freshwater is not obligatory for newly recruited
M. cephalus and abundant evidence is available from
around the world of this species remaining in estuaries
until mature and then returning to the sea (Thomson
1959; Wallace and van der Elst 1975).
The cruising speed of subadult M. cephalus
approximately 15 cm TL was 7 body lengths per
second whilst they were migrating along the mid-
Atlantic coast of North America, which is consider-
ably higher than typical cruising speeds for other
fishes at 2–3 body lengths per second (Peterson 1976).
Monitoring of M. cephalus movements in freshwater
areas have shown that individual fish tagged in
Australia had mean monthly travelled distances of
approximately 5 km, with a maximum distance of
20 km (Gehrke et al. 2001). Downstream movements
were generally followed by upstream returns to the site
of tagging, as evidenced by the narrow home range of
individual mullet (Gehrke et al. 2001).
When not foraging on the bottom for food,
M. cephalus usually swims near the water surface.
Although this behaviour is suited to the downward
angled eyes which would detect piscivorous fishes
approaching from the side or below, it is a disadvan-
tage in terms of bird predation from above (Whitfield
and Blaber 1978b). Despite its ability to jump,
M. cephalus is unable to surmount river weirs (Bok
1984a) but does have the ability to migrate between
different estuaries or catchments as shown by tagging
studies (Thomson 1955). Occasionally, juveniles and
adults are flushed out of estuaries in what seems to be a
forced export of estuarine residents caused by river
flooding. Some of these individuals may return to the
same or different estuaries at a later stage. However, it
is sometimes difficult to distinguish whether these
events are actually coinciding with a natural move-
ment of M. cephalus between the estuarine and marine
environment, or whether they are being forced out of
the system by the floodwaters.
One of the main tools for studying flathead mullet
migrations between waters with different salinities is
the use of otolith Strontium:Calcium (Sr:Ca) ratios,
leading to a better understanding of this part of life
history (Chang et al. 2004a,b). According to the
temporal change of otolith Sr:Ca ratios in M. cephalus
from Taiwanese waters, the migratory history of adult
flathead mullet beyond the juvenile stage can be
classified into two types (Fig. 7). Type 1 M. cephalus,
with otolith Sr:Ca ratio ranges of 4.0–13.9 910
-3
,
indicates that they migrate between estuarine and
marine waters but rarely enter freshwater habitats
(Fig. 7a). Type 2 otolith Sr:Ca ratios decreased to a
minimum value of 0.4 910
-3
which indicates that
these M. cephalus migrated to and occupied freshwa-
ter habitats for an extended period. Flathead mullet
beyond the juvenile stage usually left the estuary for
0
5
10
15 Type 1
0
5
10
15
0 1000 2000 3000 4000 5000 6000
0 1000 2000 3000 4000 5000 6000
Type 2
Sr:Ca ratio (×10-3)
Distance from primordium to otolith edge (µm)
Fig. 7 Temporal changes in the Sr:Ca ratios in M. cephalus
otoliths of Type 1 and Type 2 from Taiwanese waters (solid
triangles =onset of estuarine life-history phase in year 1; open
triangles =annuli). The grey bands between the Sr:Ca ratios
3–7 910
-3
indicate occupation of estuarine waters, above
7910
-3
occupation of marine waters and below 3 910
-3
occupation of fresh waters (modified from Chang et al. 2004a)
654 Rev Fish Biol Fisheries (2012) 22:641–681
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the sea, but a few of the juveniles \2 years old do not
appear to have left the freshwater habitat during this
phase in their life cycle (Fig. 7b). Mugil cephalus
collected from nearshore and offshore marine areas
comprised mainly Type 1 specimens, while those
collected from Taiwanese estuaries were a mixture of
Type 1 and 2. Therefore, the migratory patterns of
M. cephalus in Taiwan are more complicated than
previously thought (Chang et al. 2004a,b), with the
presence of cryptic species in North West Pacific
M. cephalus populations (Shen et al. 2011) shedding
new light on these results and suggesting the possi-
bility of species specific migratory behaviour.
The otolith microstructure, incremental width and
Sr:Ca ratios indicate that otoliths from juvenile M.
cephalus in Taiwan can be used to sort specimens into
marine larval, estuarine pre-juvenile, intermediate-
juvenile and post-juvenile stages (Chang et al. 2004a).
Relative abundance, total length, otolith radius and
daily age of the estuarine pre-juvenile stage display a
geographic cline that increases from the central-
western towards the northern and southern estuaries
of Taiwan. This suggests that ocean currents and tidal
movements in the Taiwan Strait play an important role
in estuarine recruitment dynamics of flathead
mullet along the western coast of Taiwan (Chang
et al. 2000).
Parasites can sometimes provide evidence of the
movements and migrations of M. cephalus, e.g. the
supposed northward movements of flathead mullet
from New South Wales to Queensland waters was
disproved by Lester et al. (2009) using the trematode
Plethorchis acanthus which infect juvenile M. ceph-
alus in Queensland waters but not those in New South
Wales.
Food and feeding
Larval M. cephalus are planktonic feeders in the
offshore marine environment (Zisman et al. 1975;
Brownell 1979), surf zone (Inoue et al. 2005) and
when they first enter estuaries (Gisbert et al. 1996).
Although the larvae are planktivorous (Nash et al.
1974), between 10 and 20 mm SL the early
M. cephalus juveniles undergo a change in diet, initially
feeding on small invertebrates that undergo vertical
migrations within the water column and later feeding
mainly on benthic organisms (Suzuki 1965; Blaber
and Whitfield 1977). This pattern was confirmed by
De Silva and Wijeyaratne (1977) who recorded no
sand or detritus in specimens smaller than 25 mm in
length, and the percentage occurrence of detritus and
sand increased with increase in body length above this
size. Also reflecting the above switch, the diet of
juveniles between 20 mm and 100 mm SL shows a
progressive increase in similarity to the food items
available within the sediment, thus indicating that the
smaller individuals may be browsing more selectively
than the larger fish (Eggold and Motta 1992).
Calanoid and cyclopoid copepods feature strongly
in the diet of late larvae and early juvenile M. cephalus
and this leads to strong competitive interactions with
other fish species at the same stage of development
(Gisbert et al. 1995; Whitfield 1985). Apart from the
similarity in early diet between mullet fry (Pisarevs-
kaya and Aksenova 1991), food competition is also
increased because several mugilid species often
migrate into estuaries at the same time (Bartulovic
´
et al. 2011). The feeding periodicity of the larvae is
unknown but M. cephalus fry in the mouth of the Arno
River have been shown to feed mostly at dusk
(Torricelli et al. 1981). On the basis of existing studies
at the time of their review De Silva (1980) concluded
that young mugilids tend to feed throughout the day,
although at different intensities. Although juvenile
M. cephalus are predominantly benthic foragers, they
are also able to feed on items within the water column,
especially at the water/air interface (Odum 1968).
The larger juvenile and adult M. cephalus feed
mainly on detritus (including particulate organic
matter) and benthic microalgae (especially diatoms),
together with foraminiferans, filamentous algae, pro-
tists, meiofauna and small invertebrates (Thomson
1963; Blaber 1976; Payne 1976; Marais 1980; Lawson
and Jimoh 2010). Although diatoms made up about
20–30 % of the organic carbon in M. cephalus
stomachs from seagrass flats in Australia, bacteria
were also important and comprised 15–30 % of the
organic carbon (Moriarty 1976). The fact that total
organic carbon was between 2 and 3 % of ash weight,
an increase of 10–20 fold over that in the in situ
sediments, suggests that M. cephalus is very effective
at actively selecting these very small food items.
Laboratory experiments have also shown that
M. cephalus predation results in a significant decrease
in abundances of total meiofauna but not macrofa-
una (Service et al. 1992), thus indicating that this
species selects small as opposed to large prey items.
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In addition, the evidence suggests that the diet of
larger juveniles and adults does not change with
increasing body size or between shallow and deeper
waters (Platell et al. 2006).
The food value of microalgae is generally higher
than that of detritus and this leads to elevated growth
rates when it is readily available (Jana et al. 2004).
There is also evidence to suggest that M. cephalus
actively seeks out benthic diatoms as a food source
(De Silva and Wijeyaratne 1977; Michaelis 1993;
Rueda 2002) but can switch to less nutritious detritus
when necessary. Evidence to support this view is
provided by M. cephalus feeding on a dinoflagellate
bloom near Sapelo Island, Georgia (Odum 1968).
Although the organic content of the mullet diet was
higher during the bloom (due primarily to the absence
of sediment particles in the stomach contents) than
during periods of normal diet consisting of detritus
particles and benthic diatoms, the assimilation effi-
ciency for the flagellates was low. Another example of
an alternative food source, or deviation from a natural
mullet diet, has been described by Porter et al. (1996),
Katz et al. (2002) and Lupatsch et al. (2003) who
demonstrated that M. cephalus can be used success-
fully to process organically enriched sediments below
marine fish farm cages in the Gulf of Aqaba, primarily
due to their ability to consume and digest the
particulate organic matter that settles on the sea floor.
When foraging M. cephalus angle their heads
downwards and create typical conical depressions in
the substratum as they take a mouthful of sediment and
associated food material. After working the material
between the pharyngeal bones they reject the most
indigestible portion (mostly sand) through their
operculae and mouth in a characteristic ‘coughing
and spitting’ action (Thomson 1954). If the fish
densities are high enough, the ejected sand and mud
produced in this way can even cloud the water, thereby
providing fishermen with an indication of where to
‘shoot’ their nets (Thomson 1954). An interesting
abnormal behavioural trait of mugilids, including
M. cephalus captured in gill nets, is the regurgitation
of stomach food contents and the ingestion of gill raker
filaments that break loose during the struggle to be free
of the net (Luther 1964; Blaber 1975).
Some studies have shown that the distribution
patterns of juvenile M. cephalus in an estuary are often
correlated with shallow muddy substrates that have a
rich microphytobenthos (Mwandya et al. 2010).
Pennate and centric diatoms are an important part of
the diet of M. cephalus and the species composition of
the diatom assemblage gives a good indication of
where the fish have been feeding, e.g. freshwater and
euryhaline diatoms dominated the diet of M. cephalus
from the upper reaches of the Swartkops Estuary
whereas marine diatoms were dominant in the diet of
specimens from the lower reaches (Masson and Marais
1975). Large amounts of sediment, mostly comprising
sand granules ranging in size from 125 to 500 lm
(Blaber 1977; Mariani et al. 1987), are usually
consumed together with the food items and this is
used to break down the food in the muscular pyloric
stomach. In addition, Anderson et al. (1958) have
shown that clay minerals passed through the gut of
M. cephalus are altered in their nature.
The inclusion of blue-green algae in the diet of
mugilids has raised the question as to whether they can
digest such material (De Silva 1980). Payne (1976) has
presented indirect evidence to suggest that M. ceph-
alus can digest blue-green algae and Stickney and
Shumway (1974) experimentally demonstrated cellu-
lose activity in this species. Odum (1968) has
suggested that bacteria and protozoa attached to
ingested detritus may also assist in the breakdown of
plant material.
It would appear that M. cephalus feeds during both
the day and night. Whilst Kuthalingham (1966) found
little variation in feeding intensity in relation to time of
day, Blaber (1976) determined that M. cephalus
forages at reduced intensities during nocturnal hours.
Similar results were obtained by Collins (1981) who
found that maximum feeding intensity was at
1100 hours, with little or no foraging at night. De
Silva and Wijeyaratne (1977) also recorded a peak in
feeding activity around midday but also recorded a
secondary peak at dawn, both of which were unrelated
to tidal phase. However, other studies have indicated
that tidal phase may influence M. cephalus foraging
activity and both Thomson (1966) and Odum (1970)
found a marked increase in ingestion rates during
incoming and high tides. Consumed food took
between 2 and 6 h to pass through the alimentary
system, with a mean of between 4 and 5 h (Odum
1970). The work of Perere and De Silva (1978) has
shown similar evacuation rates but highlighted the
effect of body size and salinity, with the rate of gut
evacuation by juvenile M. cephalus increasing with
increased body mass and salinity.
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The chemical composition of the alimentary canal
contents of M. cephalus from the Swartkops Estuary in
South Africa indicated a mean protein content of
4.9 % and carbohydrate content of 3.3 % (Marais and
Erasmus 1977a). The above study also determined that
smaller M. cephalus tended to consume more nutri-
tious food material than larger fish. The body compo-
sition of M. cephalus from the same estuary was
determined by Marais and Erasmus (1977b) who
found that overall this species comprised 17.5 %
protein, 2.7 % fat, 4.9 % ash and 74.5 % moisture.
These figures are very similar to those quoted by
Thomson (1966), except for ash content which was
1.2–1.8 % and fat 2.7–6.8 % in his review. When
compared to other fish species, M. cephalus can be
regarded as having a relatively high fat composition
(De Silva 1980; Marais and Erasmus 1977b; Perera
and De Silva 1978).
Obviously the actual condition of individual fish is
related to feeding intensity, which in some parts of the
world is linked to rainfall events (De Silva and Silva
1979) or available food resources. Bok (1983)
recorded heavier M. cephalus for equivalent lengths
in the nutrient and food rich Swartkops and Great Fish
estuaries when compared to the Kowie Estuary, and
also recorded faster growth rates in the former two
systems. The condition factor in adult fish is also likely
to be influenced by the spawning cycle, with pre-
spawning individuals having a higher condition
(length-weight relationship) than spent fish due to
the large size of the gonads in this species. This view is
supported by Bok (1983) who recorded a significant
decline in the relative condition of mature M. cephalus
in the Kowie Estuary between May and July, which
coincided with the spawning season of this species.
Blaber (1976) attempted to determine if there was
any interspecific competition between the mugilid
species, including M. cephalus, in Lake St Lucia. He
was able to ascertain considerable dietary overlap but
that little spatial segregation occurred, all of which
suggests that these species are consuming a non-
limiting and abundant resource (mainly detritus and
particulate organic matter). One of the potential ways
of partitioning the other food resources between the
mugilid species, including M. cephalus, may be linked
to differential selection according to inorganic particle
size (Blaber 1977). Co-existence and potential com-
petitive interactions of five mugilid species (including
M. cephalus) in the Western Mediterranean have been
studied by Cardona (2001) who found that niche
theory did not apply to these species, with results
indicating that the mullet species were below carrying
capacity of the food resources and hence competitive
exclusion did not operate in the study area. Similarly,
Iba
´n
˜ez (1993) examined coexistence of M. cephalus
and Mugil curema in a coastal lagoon of the Gulf of
Mexico and found no major difference in diet. Luther
(1962) came to the same conclusion following an
investigation of the food habits of M. cephalus and
Liza macrolepis (now Chelon macrolepis).
Potential dietary competition between M. cephalus
and two other non-mugilid detritivorous fish species
(Oreochromis mossambicus and Chanos chanos)in
Lake St Lucia, determined that the diet of these three
species did not overlap, even to the extent of having
different diatom species in their stomach contents
(Whitfield and Blaber 1978c). Darnell (1961)
attempted to determine the potential competition
between M. cephalus and other iliophagous feeders,
including invertebrates. Although 77 % of the volume
of food consumed by M. cephalus was detritus,
compared to 8–99 % for 24 other species of fish and
two species of crustaceans, the study was not able to
answer the question as to whether the pressure on the
detrital resource was great enough to create significant
competition between the species.
Although dietary overlap between mugilids has
been firmly established by the above studies, there are
large differences in the ratios of intestinal length:
standard length for mugilids, e.g. M. cephalus has a
ratio of 4.5–5.2:1 whereas three other mullet species in
the Swartkops Estuary had ratios of 2.0–2.6:1 (Marais
1980). Whilst M. cephalus has an average of two
pyloric caeca, which are the source of diatase and
trypsin-like enzymes, the other three species from the
Swartkops had an average of six which must signif-
icantly increase the surface area of the intestinal
epithelium and may compensate for the relatively
shorter intestine in these species. Thomson (1966) has
summarized the number of pyloric caeca in mugilids
and they range from 2 to 22 per species, with most taxa
having between 2 and 8 caeca. Clearly the digestive
system of M. cephalus is very different to that of other
mugilid species but the significance of these differ-
ences has not been fully explored (Thomson 1954).
The relatively long intestine of M. cephalus would
deal very effectively with diatoms in its diet (Odum
1970) but is also adapted to extracting nourishment
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from less digestible plant detritus and blue green algae
(Hickling 1970).
Due to the nature of the food consumed by
mugilids, documentation of actual assimilation,
potential dietary overlap between sympatric mullet
species and trophic level information all require
confirmation using modern analytical techniques.
The stable carbon isotope signature (d
13
C) of
M. cephalus from the Mngazana and Mngazi estuaries
in South Africa (Mbande et al. 2004) averages about
-16%which, in terms of the known food sources for
this species in those estuaries, most closely reflects a
mixture of epiphytic microalgae (-11%) and a more
enriched d
13
C source (e.g. microphytobenthic algae
-20%). An almost identical result was obtained by
Hadwen et al. (2007) for M. cephalus (-16%)in
Belongil Creek, an intermittently open estuary in
Australia, and the inferred primary food source was
given as epilithic algae (-15%). In contrast,
M. cephalus from the nearby freshwater dominated
Tallows Creek had a much more depleted d
13
C value
(-23%), with the diet in this estuary being dominated
by a much higher proportion of seston (-23%) from
the water column (Hadwen et al. 2007).
Studies which support planktonic food consumption
by M. cephalus include Cardona et al. (1996) who
showed that this species can obtain much of their food
from the zooplankton in freshwater fish ponds. Since
M. cephalus uses a pump-filter action when foraging,
and can select particles as small as 20 lm (Cardona
1996), filtering of small invertebrates from open waters
should not be too difficult. The filtering of very small
particles from the water column by M. cephalus has
been shown to occur under both laboratory and natural
environmental conditions, with ‘marine snow’ being
actively consumed but zooplankton preferred when
both food sources were available (Larson and Shanks
1996). Despite the foraging flexibility exhibited by
M. cephalus there are conditions where growth can be
inhibited due to sub-optimal habitats and food
resources (Brusle and Cambrony 1992).
More recently, fatty acids have been used to trace
the transfer of organic matter in aquatic food webs.
A study by Alikunhi et al. (2010), which included
M. cephalus from an estuarine mangrove system,
showed that the fish were attracted to mangrove leaf
litter that was approximately 40 days old and that this
coincided with a peak in branched fatty acids associ-
ated with high microbial biomass on the leaf material.
This link was not only attributable to enrichment
created by the microbial biomass but also the
enhancement of nutrients associated with the leaves,
an important dietary requirement for detritivorous fish
(Rajendran and Kathiresan 2007).
Growth
Numerous published studies have been undertaken on
flathead mullet growth in aquaculture (e.g. Silva and
Perera 1976;Oren1981; Carr and Aldrich 1982; Kraul
1983; Jana et al. 2004). Unfortunately, the widespread
distribution of M. cephalus, together with the lack of
co-ordinated studies on this species, has meant that
research methods have not been uniformly applied for
estimating the age and growth, and therefore make
global comparisons difficult. Nevertheless, useful
information can be gleaned from aquaculture studies,
e.g. maximum growth efficiency of juvenile
M. cephalus fed to excess was found to occur at a
salinity of 20 and percentage conversion efficiency
was highest at a salinity of 10 (De Silva and Perera
1976). Similarly Barman et al. (2005) recorded
maximum M. cephalus growth, feed conversion
efficiency and intestinal enzyme activity at a salinity
of 10, thus reinforcing the role of estuarine salinities in
optimizing the growth potential of this species.
A universal and validated method for age estima-
tion of M. cephalus does not exist in the literature. Fish
scales were originally used for mullet age estimation
in the 1950s (Tung 1959), but since the 1970s both
scales (Tung 1981; Hamza 1999) and otoliths (Smith
and Deguara 2003) have been used to estimate age and
growth based on seasonal macro-increments. Iba
´n
˜ez
and Gallardo-Cabello (1996b) compared scales and
otoliths for age determination purposes and reported
that fish scales could be used for juveniles but that
otoliths provided better resolution for the older size
classes. Van der Kooy and Guindon-Tisdel (2003)
tried to standardized otolith preparation methods for
age determination of several mullet species from the
Gulf of Mexico, including M. cephalus, and recom-
mended using a transverse section of the sagittal
otolith. This approach is supported by Panfili et al.
(2007) who recorded otolith macro-increments from
flathead mullet individuals from different parts of the
world using transverse sections (300 lm thick) under
transmitted light in order to reveal thin opaque bands
highlighted in the otolith sulcus area (Fig. 8).
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Whole otoliths have been used by Chang et al.
(2000) for the detection of daily micro-increments in
larvae and early 0?juveniles from Taiwanese waters
and further work in this field is required from other
parts of the world. According to Kuo et al. (1973) the
larvae hatch at approximately 2.6 mm in length and
attain 17.7 mm by 42 days. However, both water
temperature and available food resources are likely to
affect M. cephalus larval and postlarval growth rates
and this will be reflected in the otoliths of these small
individuals.
Growth rate data on juvenile and adult M. cephalus
are relatively scarce in the primary literature but seem to
be more readily available in the less accessible grey
literature which is astonishing for a cosmopolitan
species such as the flathead mullet. Even in the only
study focused on age validation (Smith and Deguara
2003), growth rate data and the variability thereof is not
given. Nevertheless, the few studies involving
M. cephalus growth have highlighted the variable recorded
rates that appear to be dependent upon the local climate
and available food resources (Wallace and van der Elst
1975; Bok 1983; Iba
´n
˜ez and Gallardo-Cabello 1996b).
For example, Anderson (1958) recorded juvenile
growth rates of about 5 mm per month during
November to January (winter) along the Atlantic coast
of the USA, but this increased to about 17 mm per
month from March to October (spring and summer)
which was similar to the mean growth rates of juvenile
M. cephalus in more tropical Florida waters (Kilby
1949). Overall growth is rapid in the first year, with fish
usually attaining 140–180 mm SL in tropical and
subtropical waters (Thomson 1963; Wallace and van
der Elst 1975), and 130–160 mm SL in more temperate
regions (Whitfield and Kok 1992). These growth rate
variations have also been confirmed in aquaculture
experiments (e.g. Oren 1981) which highlights the
need for considerably more research on M. cephalus
growth at all stages of its life history (Bok 1983).
Hendricks (1961) recorded a maximum size for
M. cephalus of 62 cm FL and Wallace (1975a)
recorded similar sized specimens of 68–72 cm TL in
the subtropical waters of Lake St Lucia. Thomson
(1966) has been able to summarize the annual lengths
of M. cephalus as being approximately 15, 25, 32 and
38 cm FL in the first 4 years. In addition, Thomson
(1951) gives estimates of 46 and 51 cm FL for
M. cephalus in temperate Western Australian waters at
the end of years five and six. Thomson (1966)
indicates the age of 54 cm FL individuals as being
7 years but points out that other authors give older age
estimates for similar sized specimens. These differ-
ences in length at age information may be partly
attributable to variations in growth within the
M. cephalus species complex but also the methods
used in ageing the fish. In addition, differences in
growth rates between the sexes have been identified by
several authors, with females generally growing faster
than males (Oren 1981; Bok 1983). Distinct morpho-
metric differences between male and female
M. cephalus have also been documented in some
regions, for example in the Gulf of Mexico (Iba
´n
˜ez
and Lleonart 1996), but the consistency of these
differences on a global scale has yet to be investigated.
Reproduction and larval development
The detailed structure and development of M. ceph-
alus gonads have been studied by Stenger (1959). The
size at sexual maturity recorded for this species ranges
very widely, with males usually maturing between 25
and 30 cm SL and females slightly larger at 27–35 cm
SL (Ameur et al. 2003). These size classes are
generally regarded as being approximately 3 years
old but some studies have given higher and lower ages
at 50 % sexual maturity (Bok 1983; Ameur et al.
2003). The wide differences in the above estimates can
probably be partially ascribed to individual research-
ers having slightly different perspectives and methods
of determining both the reproductive state and age for
this species. Even within particular studies results can
be ambiguous depending on an author’s interpretation
of the term sexual maturity, e.g. McDonough et al.
(2005) determined that female M. cephalus first
matured at about 29 cm TL (age 2 years) but 100 %
maturity was only recorded at 40 cm TL (age 4 years).
The possibility that ranges in sexual maturity esti-
mates were based on different taxa within the
M. cephalus species complex also cannot be ruled out.
Although it has been suggested that M. cephalus
may exhibit protandric hermaphrodism (Stenger
1959), more recent studies (e.g. McDonough et al.
2005) have shown that this species is gonochoristic but
capable of exhibiting non-functional hermaphroditic
characteristics in differentiated mature gonads. Once
again it may be that only certain taxa within the
M. cephalus species complex exhibit hermaphroditic
tendencies.
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COUNTRY Whole otolith Transverse section
(transmitted light)
Transverse section
(reflected light)
Benin
1 opaque increment
Greece
1 opaque increment
Mauritania
2 opaque increments
Senegal
1 opaque increment
South Africa
1 opaque increment
Spain
4 opaque increments
Taiwan
6 opaque increments
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The sex ratios of adult M. cephalus shoals are
usually well balanced, e.g. Bartulovic
´et al. (2011) and
Iba
´n
˜ez and Gallardo-Cabello (2004) recorded a male
to female ratio of 1.1:1.0 for fish in Croatian and
Mexican waters, Lawson and Jimoh (2010) had a ratio
of 1.0 male to 1.09 females in Nigeria, Katselis et al.
(2005) an overall ratio of 1:1 for mature individuals
undergoing a seaward spawning migration in Greece,
and Silva and De Silva (1981) had a 1:0.95 ratio of
males to females in Sri Lanka. Bok (1983) also
documented a 1:1 sex ratio for M. cephalus in the
Kowie and Great Fish estuaries of South Africa but
recorded a 1:0.2 male to female ratio in the Kowie
River and 1:0.3 male to female ratio in the Great Fish
River. When examining the composition of mature
M. cephalus in an isolated freshwater farm dam (grown
from previously introduced fry), Bok (1983) found an
almost 1:1 sex ratio. He therefore suggested that fry
recruitment into the Kowie and Great Fish rivers takes
place equally by males and females but that most
females depart the river before sexual development
occurs.
Despite the similar sex ratios outlined above,
M. cephalus males and females usually have very differ-
ent maximum gonadosomatic indices (GSI) when
fully mature, e.g. in Taiwanese waters males have a
recorded GSI of 19 and females 21 at the peak of the
spawning season (Hsu et al. 2007). However, even
higher fecundity (GSI [24) has been recorded by
female M. cephalus in Queensland (Grant and Spain
1975), South Carolina estuaries (maximum GSI =28)
(McDonough et al. 2003) and in the Black Sea and
Gulf of Mexico where the GSI reached 40 (Apekin and
Vilenskaya 1978; Iba
´n
˜ez and Gallardo-Cabello 2004).
Actual fecundity is related to fish size, with 1,082,200
eggs being recorded for a 32 cm (TL) female from
Lagos Lagoon (Lawson and Jimoh 2010) and
2,632,000 eggs recorded for a 54 cm (SL) specimen
participating in the spawning migration at Lake St
Lucia (Whitfield and Blaber 1978a). A review of
M. cephalus fecundity by Thomson (1963)givesanormal
range of 1.2–2.8 910
6
ova for this species but some
authors have recorded considerably more (Grant and
Spain 1975; Apekin and Vilenskaya 1978).
Bok (1983) has documented that M. cephalus in
Eastern Cape (South Africa) have limited gonadal
development in the freshwater areas of rivers, with a
maximum GSI of 1.3 being recorded. A similar lack of
reproductive development in freshwater has been
recorded under aquaculture conditions (Yashouv 1969).
However, gonadal development up until an advanced
stage occurs in estuarine waters but the ‘ripe running’
state is generally only attained in the marine environ-
ment (Bok 1979; Wallace 1975b). If mature fish are
denied access to the sea during the spawning season,
these individuals tend to resorb their gonads (Wallace
1975b). Although seawater salinities are not required
for the full maturation of the gonads (Tamaru et al.
1994), the success of fertilisation and larval survival of
M. cephalus does depend on environmental salinity
(Lee et al. 1992). Laboratory research has shown that
fertilised M. cephalus eggs can develop to the
embryonic stage within a salinity range of 5–60 and
that hatching occurs in salinities from 10 to 55.
However, no larvae survived at 10 or 55 and optimal
survival occurred between 30 and 40, with a peak at 35
(Lee and Menu 1981).
Water temperature appears to be an important
determinant in vitellogenesis (Kuo et al. 1974) which
means that there are different spawning periods across
the distributional range of this species. Most recorded
M. cephalus spawnings appear to occur in water close
to 20 °C (Brownell 1979; Shyu and Lee 1986). This
means that the species needs to be flexible in which
season to spawn, depending on geographical location.
For example, M. cephalus in the cooler temperate
regions of Europe tend to have a summer spawning
period whereas spawning reaches a peak during winter
in the subtropical regions (Table 2). Indeed, it would
appear that flathead mullet avoid spawning in seasons
when water temperatures are either too hot or cold
(Table 2) for the survival of the eggs and larvae, e.g. in
the Black Sea where winter temperatures can decline
as low as 3–6 °C and spawning therefore occurs
during summer when water temperatures are much
higher (Apekin and Vilenskaya 1978). A good exam-
ple of the influence of temperature is in South Africa
where M. cephalus in sub-tropical waters spawn
mainly between May and August (winter) (Wallace
1975b), whereas in the cool-temperate region
Fig. 8 Examples of M. cephalus otoliths from different parts of
the world viewed whole under reflected light on a dark
background and in transverse section (300 lm thickness) under
transmitted or reflected light. Scale bar 1 mm (whole otolith)
and 500 lm (otolith sections). Modified from Panfili et al.
(2007)
b
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spawning occurs predominantly during January and
February (summer) (Brownell 1979).
Photoperiod is also important in initiating gonadal
development in M. cephalus (Kuo et al. 1974). Given,
the different spawning seasons outlined above, one
must assume that the populations in the different
biogeographic areas would have different photoperiod
triggers. If so, this could be further evidence in favour
of a species complex for M. cephalus. In some parts of
the world the spawning of M. cephalus often occurs
after wet seasons and this is reflected by the seasonal
catches of fishermen in the Gulf of Mexico (Iba
´n
˜ez and
Gutie
´rrez-Benı
´tez 2004). Similar observations have
been made on the southeast coast of Africa where
M. cephalus shoals start spawning in early winter, i.e. at
the end of the summer rainy season (Wallace 1975b).
Such a spawning strategy would ensure that postlarval
recruitment into estuaries would occur during rela-
tively stable conditions, i.e. in the absence of river
flooding.
As has been mentioned, the spawning grounds of
M. cephalus are usually located in the sea (Bartulovic
´
Table 2 Spawning seasons (months with black borderline and
bold temperatures) of M. cephalus from various parts of the
world (numbers represent the mean monthly coastal seawater
temperatures in °C for each region; SST data source
http://fr.surf-forecast.com/)
Areas Spawning periods References
JJASONDJFMAM
Black Sea 20 23 24 20 16 12 8 6 5 6 9 15 Apekin and Vilenkaya (1978)
Turkey
(Mediterranean) 24 26 28 26 24 20 18 17 16 16 18 20 Erman (1959)
Egypt
(Mediterranean) 24 26 27 26 25 22 20 18 17 17 18 20 Faouzi (1938)
Morocco
(Atlantic Ocean) 21 23 23 23 21 20 18 17 17 17 17 19 Ameur et al. (2003)
Caspian Sea 20 24 24 21 16 12 9 6 6 6 9 14 Avanesov (1972)
Adriatic Sea 22 25 26 22 19 16 14 12 11 11 13 18 Morovic (1963)
Greece
(Agean Sea) 22 24 25 23 20 17 15 13 13 13 14 18 Koutrakis (2004)
Greece
(Mediterranean) 22 25 26 24 22 19 17 15 15 15 16 19 Katselis et al. (2005)
Tunisia
(Mediterranean) 21 24 26 25 22 19 17 15 14 15 16 18 Brusle and Brusle (1977)
USA
(Atlantic Ocean) 26 28 28 26 24 23 22 20 19 20 21 24 Bacheler et al. (2005)
Mexico
(Gulf of Mexico) 28 28 29 28 26 23 21 20 19 19 22 25 Ibáñez and Gutierrez-Benitez (2004)
India
(Indian Ocean) 29 28 28 29 29 28 27 27 27 29 30 30 Mohanraj et al. (1994)
Mauritania
(Atlantic Ocean) 23 26 27 28 27 25 22 20 19 19 19 20 Brulhet (1975)
Sri Lanka
(Indian Ocean) 29 28 28 28 28 28 28 27 28 29 30 30 De Silva and Silva (1979)
USA
(Atlantic Ocean) 26 28 28 27 25 23 22 20 20 20 20 24 McDonough et al. (2005)
USA
(Atlantic Ocean) 26 28 29 27 25 23 21 23 21 20 23 26 Kilby (1955)
Australia
(Pacific Ocean) 22 21 21 22 22 24 25 26 26 26 25 24 Kesteven (1942)
South Africa
(Indian Ocean) 22 22 21 21 22 23 24 26 26 26 25 24 Wallace (1975b)
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et al. 2011), even though adults can spend almost their
entire life in estuaries and/or lagoons (Cardona 2000).
Spawning has been documented in inshore and
offshore waters of regions such as Gulf of Mexico,
Israel and South Africa where salinities of between 30
and 40 have been recorded at the time of spawning
(Ditty and Shaw 1996; van der Horst and Erasmus
1981). In Taiwan, the mullet spawning stock consists
mainly of individuals migrating between the estuary
and marine zone (Chang et al. 2004a) but some
populations have never entered an estuary and have a
purely marine life cycle. Once again this is an
indication that different M. cephalus species exhibit
different life-history patterns in Taiwanese waters.
There are only a few published studies on the actual
reproductive behaviour of M. cephalus from different
parts of the world, with some information from the
Gulf of Mexico (Iba
´n
˜ez and Gutie
´rrez-Benı
´tez 2004;
Iba
´n
˜ez and Gallardo-Cabello 2004) and the Mediter-
ranean area (Cardona 2000). Actual spawning has
seldom been observed but Arnold and Thompson
(1958) give a detailed account of apparent nocturnal
spawning in surface waters by small groups of
M. cephalus in the Gulf of Mexico. Males appeared to
congregate around single females and then break away
from the school, swimming close to the surface in an
erratic manner. Periodically one or more of the males
would move beside or below the female and nudge her
abdomen with their head and body, quivering and
ceasing to swim momentarily. Although mullet ova
were collected from the sea during the above event,
they did not appear to have been fertilized. Thomson
(1957) suggested that M. cephalus produce only one
set of ova per year and Stenger (1959) concluded,
based on specimens from Florida, that this species is
capable of spawning more than once in a season.
Mugilid eggs are externally fertilised and both eggs
and larvae (Fig. 9) are pelagic within the marine
environment (Brownell 1979). The effective temper-
ature range for the development of M. cephalus eggs is
11–24 °C, with an optimum of 24 °C (Nash and
Koningsberger 1981). Although eggs are positively
buoyant in seawater, they become negatively buoyant
after fertilisation and this may be a good reason to
spawn over deeper offshore waters (Arnold and
Thompson 1958). Indeed, Sylvester and Nash (1975)
have speculated that the considerable tolerance of
M. cephalus eggs to fluctuating temperatures may be
an adaptation to changing water temperatures as they
sink. However, slow surface currents and eddies in the
ocean are more than sufficient to keep the fertilised
eggs in suspension (Nash et al. 1974). Ripe, unferti-
lised eggs are rounded and colourless and the surface
of the fertilised egg is smooth and approximately
870 ±30 lm in diameter with an oil globule of
350 lm (El-Gharabawy and Assem 2006). However,
other authors give a slightly larger mean diameter for
fertilised M. cephalus eggs, ranging from 930 to
950 lm in diameter with an oil globule diameter
ranging from 330 to 380 lm (Tang 1964; Kuo et al.
1973; Finucane et al. 1978).
Hatching occurs about 26 ±4 h after fertilisation
at 25 ±1°C and a salinity of 34, with the newly
hatched larvae being approximately 1.97 ±0.23 mm
in length (El-Gharabawy and Assem 2006). The larvae
of M. cephalus undergo two periods of sinking if water
currents are absent, between day 2 and 3 and between
day 6 and 7 (Kuo et al. 1973). Larvae began feeding
3–5 days after hatching in the laboratory, which
coincided with the completion of eye and mouth
development and straightening of the body axis (Nash
Fig. 9 Illustrations showing early larval development of M.
cephalus in seawater at a temperature of 15 °C (after Brownell
1979); aM. cephalus egg (0.98 mm in diameter), bM. cephalus
soon after hatching (2.7 mm NL alive), cM. cephalus at 2 days
(3.3 mm NL alive), dM. cephalus at 4 days (3.5 mm NL alive)
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et al. 1974; El-Gharabawy and Assem 2006). Rates of
larval development appear to be influenced by water
temperature (Thomson 1966) and laboratory experi-
ments have also indicated that larval growth is
adversely affected if they have not fed by 84 h
posthatch (Eda et al. 1990). Postlarval development
has been summarised by Thomson (1966) who
indicated that scales develop at 11 mm in length but
M. cephalus have to reach 20 mm to be at the
querimana stage before all the main features of the
species are present, except that it has two anal spines
instead of three and no adipose eyelid.
Parasites
The flathead mullet acts as a host for a wide variety of
parasites (Valles-Rı
´os et al. 2000). These include
Monogenea in the gills of M. cephalus in the Atlantic,
Mediterranean and Japan Sea (Baker et al. 2005;
Rubtsova et al. 2006,2007); Digenea in specimens
from Egypt and Australia (Elsheikha and Elshazly
2008; Lester et al. 2009); myxosporidean parasites
infecting M. cephalus gills in India, Turkey and Spain
(Eiras and D’Souza 2004; Umur et al. 2010; Maı
´llo-
Bello
´n et al. 2011), scales and mesenteric vessels
in Tunisia (Bahri and Marques 1996), the heart
in Senegal (Faye et al. 1997), intestinal musculature in
Japan (Maeno et al. 1993), brain and gall bladder in
India (Narasimhamurti and Kalavati 1979, Kalavati
and Anuradha 1995); trichodinid parasites in the gills
of M. cephalus from Turkey (O
¨zer and O
¨ztu
¨rk 2004);
acanthocephalan parasites in the intestines of adult fish
from India (Jithendran and Kannappan 2010); para-
sitic copepods on the gills of M. cephalus from South
Africa and the USA (Oldewage and van As 1988;
Baker et al. 2005); trypanorhynch plerocercoid infes-
tation of the body muscles in South Africa (Schramm
1991); branchiuran parasites from the USA (Rawson
1977); haemoprotozoan infections in the blood of
M. cephalus from estuaries in South Africa (Smit et al.
2002); and parasitic nematodes from the Mediter-
ranean (Merella and Garippa 2001) and Australia
(Shamsi et al. 2011).
The above parasites infect a variety of organs and
tissues of M. cephalus, ranging from the brain, blood
vessels, gills, skin, alimentary canal, liver, muscles to
mesentery tissue. Whilst the dynamics and distribution
of ectoparasites on the body of the fish are strongly
influenced by external conditions such as salinity
(Baker et al. 2008), the environmental physico-chem-
ical drivers of endoparasites are less clear (Merella
and Garippa 2001). Nevertheless, Moravec and Libos-
va
´rsky
´(1975) found that the incidence and intensity of
intestinal helminth infection of individual M. cephalus
in brackish Lake Borullus, Egypt, gradually declined
after the mullet entered the lake from the sea. The
trend of altered parasitic infections in grey mullet with
changing foraging behaviour has been highlighted by
Paperna (1975) who found that specimens \30 mm
arriving in estuaries from coastal marine waters are
heavily infected with larval trematodes, cestodes and
nematodes, but that these parasites are then replaced
by other species (mainly trematodes and acanthoceph-
alans) following the mugilid transition from a plank-
tonic to benthic feeding habit. Similar sentiments were
expressed in the mugilid review by Thomson (1966)
who highlighted that early pelagic life stages are
infested by endoparasites requiring an intermediate
host, whereas the assumption of an iliophagous
benthic diet results in replacement of most of these
species by endoparasites and ectoparasites with direct
life cycles.
There are a number of parasites, especially trem-
atodes, which use M. cephalus as an intermediate host
(Thomson 1966). The definitive hosts of many of these
parasites appear to be piscivorous birds, e.g. the final
host of the trematodes Mesostephanus appendiculo-
ides and Phagicola longa include the brown pelican
Pelicanus occidentalis, the night heron Nycticorax
nycticorax and American egret Casmerodius albus
(Hutton and Sogandares-Bernal 1959). The intensity
of infection of individual mullet has not been recorded
but in the above study the proportion of mullet infected
by M. appendiculoides was 89 % in 1956 and 60 % in
1957, with P. longa infecting 76 % of the mullet
examined in 1957.
An important side effect of parasite infections is
that they can set up conditions in the host fish that
facilitate secondary infection by bacteria, e.g. red spot
disease in M. cephalus from New South Wales
estuaries (Callinan and Keep 1989). Red spot disease
in this species has also been shown to be linked to low
dissolved oxygen concentrations (Callinan et al. 1989)
and high river flows which lead to dermal ulcers on the
fish (Virgona 1992). However, Rodgers and Burke
(1988) have suggested that ectoparasites, together
with a variety of primary and secondary bacterial
pathogens, may interact with environmental and
664 Rev Fish Biol Fisheries (2012) 22:641–681
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behavioural factors to determine the prevalence of
external lesions on M. cephalus in southern Queens-
land estuaries.
In general, there is an increase in the number of
parasite species with increased age of the host
M. cephalus, with the initial infection being influenced
by a variety of factors, e.g. closeness of association of
the different mullet age classes (Rawson 1976). The
issue of competition between different parasite species
on the same fish has been investigated by some
authors. For example, El Hafidi et al. (1998) found that
two monogenean species that are parasitic on the gills
of M. cephalus tend to occupy different portions of the
gills on the same fish, and also that one species has a
marked preference for the left side and the other for the
right side. Rawson (1977) also studied co-existence of
different gill parasites on M. cephalus and found a
high degree of habitat subdivision between the various
crustacean parasites, but no competitive exclusion.
Given that there are at least 40 different species of
parasitic copepods known to be hosted by M. cephalus
(Ho and The Do 1982), it is likely that competition
between parasites on this fish species is likely to be
intense.
Thomson (1963) lists 27 parasite species on
M. cephalus that had been detected by various investi-
gators in the early part of the last century, 18 of which
were trematodes. Whether this indicates the relative
importance of trematodes as parasites on this species
or merely the abundance of scientists studying trem-
atodes is uncertain. Some species of trematode appear
to parasitize M. cephalus and other species of grey
mullet (Thomson 1966) and a similar situation is
found with some species of copepod parasites on the
gills of mugilids (e.g. Bere 1936). Removal of
ectoparasites by ‘cleaner’ fishes, as is prevalent in
many coral reef environments, is seldom possible in
the more turbid estuarine environments where flathead
mullet are abundant. However, Breder (1962)
recorded small Lagodon apparently picking off para-
sites from large M. cephalus.
Ecological importance and biomarker studies
Ecological importance
Most M. cephalus populations undergo regular migra-
tions between coastal and marine areas in many parts
of the world and are therefore very important in energy
and nutrient transfer between these adjacent aquatic
ecosystems (Ray 2005; Lebreton et al. 2011).
Although a quantification of these transfers has yet
to be undertaken M. cephalus, by virtue of its
abundance and biomass, is likely to be one of the
more important fish species providing connectivity
within the coastal zone. An indication of this impor-
tance is provided by Thomson (1959) who estimated
that 1?and older M. cephalus in the 44 km
2
Lake
Macquarie, New South Wales, had a population of
2.693 910
6
(806 910
3
kg) or 61 individuals ha
-1
(18 g m
-2
).
The flathead mullet is situated at the base of the
food pyramid and, by virtue of its consumption of
particulate organic matter, detritus and benthic mic-
roalgae, this species is able to ‘telescope’ the food
chain and make high quality fish protein available to
top predators. For example, in Lake St Lucia
M. cephalus is a dominant fish species that is exten-
sively preyed upon by the bull shark (Carcharhinus
leucas), fish eagle (Haliaeetus vocifer), osprey
(Pandion haliaetus), white pelican (Pelecanus ono-
crotalus) and Nile crocodile (Crocodylus niloticus)
(Tooth 1946; Whitfield and Blaber 1978b,1979a,b;
Clancy 2005). In more temperate regions piscivorous
birds are major predators of the flathead mullet
(Martuccia et al. 1993; Liordos and Goutner 2009)
and in some estuaries bottlenose dolphins (Tursiops
aduncus) enter these systems to feed on M. cephalus
which is a favoured food of these predators (Fury and
Harrison 2011). According to Thomson (1963) at least
13 species of fish and six species of bird are known to
include grey mullet in their diet, and this figure is
likely to be much higher if a complete global inventory
was undertaken.
Mugil cephalus are responsible for bioturbation of
sediments whilst foraging and may also affect micro-
phytobenthos composition and benthic invertebrate
abundance within these habitats. In addition, there is
indirect evidence that they may alter phytoplankton
stocks within an ecosystem by selective predation on
small vertically migrating planktonic invertebrates. In
the latter context, a laboratory investigation by Torras
et al. (2000) showed that predation by M. cephalus on
cladocerans in freshwater tanks favoured the devel-
opment of phytoplankton. Since phytoplankton
appears to be an essential ecosystem component that
ensures larval survival and growth (Harel et al. 1998),
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the ecological role of M. cephalus for larval fish
survival may be important.
Loss of flathead mullet populations from coastal
ecosystems as a result of mass mortalities or the
breakdown of migration routes can have severe
consequences for the associated ecosystems. A major
fish kill involving M. cephalus in the Tanshui Estuary,
Taiwan, affecting approximately 50 tonnes of this
species, would undoubtedly have affected ecological
pathways within this system (Wang et al. 2011).
Similarly, a major fish kill in the Indian River (Florida)
affected M. cephalus in this system and was caused by
a dinoflagellate bloom (Steidinger et al. 1998). The
loss of marine connectivity in the St Lucia system,
South Africa, due to redirecting Mfolozi River flow
(Whitfield and Taylor 2009) has resulted in hyperh-
aline conditions in the lake and a cessation of the large
M. cephalus spawning migrations on which a large
number of predators depended for food. Long-term
freshwater deprivation in an estuary can also result in
poor M. cephalus stocks (Marais 1983b) when com-
pared to nearby estuaries with moderate river inflow
(Marais 1983a). The opposite effect, namely hurricane
induced river flooding of an estuarine system, can
also result in a temporary decline in the abundance
M. cephalus (Switzer et al. 2006).
The introduction of exotic fish species that compete
with M. cephalus for food can also have a detrimental
ecological effect. For example, Cardona et al. (2008)
have shown that the introduction of the exotic
Cyprinus carpio played a pivotal role in creating a
‘‘juvenile bottleneck’’ for mugilid fry, including
M. cephalus, recruiting into 42 Western Mediterranean
estuary sites. This effect then persists through the
juvenile and adult stages by altering overall fish
assemblage composition within the affected estuaries.
Biomarker studies
The use of biomarkers can offer a more integrated
evaluation of the effects of pollutants on an ecosystem
(Richardson et al. 2011). As already indicated,
M. cephalus is a widespread species that can occupy
a range of aquatic habitats, thereby possessing several
characteristics that are required in an indicator species,
e.g. broad salinity tolerance and a distribution that
covers a wide variety of habitats. Unfortunately few
biomarker studies have been performed using grey
mullet but Ferreira et al. (2005) showed that the
presence of pollutants induced oxidative stress
responses in M. cephalus. Ferreira et al. (2004) also
assessed the impact of organochlorine contaminants
on M. cephalus in a Portuguese estuary and Maruya
et al. (2005) focused on toxophene contaminants on
the same species in a United States estuary, with both
studies pointing to the suitability of this fish as a
‘sentinel’ species for environmental monitoring.
Similarly Frodello et al. (2001) suggested that the
accumulation of macrophage aggregates in the liver of
M. cephalus can also be an indication of environmen-
tal stress.
Tsangaris et al. (2011) assessed oxidative stress and
genotoxicity biomarker responses in M. cephalus from
Saronikos Gulf, Greece, and concluded that this
species could be useful in assessing pollution impacts
in coastal environments. Oxidative stress in the liver
mitochondria of M. cephalus can also indicate expo-
sure of fish to contaminated waters. For example,
oxidative stress induced responses were detected in
M. cephalus from the polluted Ennore Estuary when
compared to the same species from the unpolluted
Kovalam Estuary (Padmini and Vijaya Geetha 2009).
The decrease in antioxidant enzyme activities
observed in mugilids following transfer to clean water
after exposure to oxidative stress due to environmental
pollutants suggests that cell response can change when
transferred to an unpolluted environment (Ferreira
et al. 2007). The effect of long-term depuration in
M. cephalus that had been exposed to pollutants in
Douro Estuary confirmed that this species can recover
if the source of environmental contamination is
removed.
A wide range of other pollutants have been detected
in M. cephalus from a variety of localities. Hassani
et al. (2006) have assessed butyltin and phenyltin
pollution of M. cephalus in the western Mediterranean.
Similarly, the bioaccumulation of heavy metals by
M. cephalus has been documented for Camlik Lagoon
(Dural et al. 2006) and Iskenderun Bay (Yilmaz 2003;
Tu
¨rkmen et al. 2006) in the eastern Mediterranean,
Pulicat Lake in India (Laxmi Priya et al. 2011) and
Lake Macquarie in Australia (Kirby et al. 2001a,b).
Fortunately for humans, heavy metals appear to
concentrate in the liver, kidney or gill tissue of this
species and not to the same extent in the body muscles
(Sultana and Rao 1998; Bahnasawy et al. 2009).
There also appears to be differential absorption of
pollutants by M. cephalus, e.g. although total mercury
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levels were low in fish from northwest Florida (North
America), elevated PCB levels were recorded in the
same area (Karouna-Renier et al. 2011). PCBs and
other pesticides have also been recorded from
M. cephalus samples in a South African estuary
(Grobler et al. 1996) and Ebro Delta in the western
Mediterranean (Pastor et al. 1996). In the latter study
DDT and HCB (hexachlorobenzene) levels were
higher in flathead mullet than red mullet and sea bass,
probably because M. cephalus were captured in
coastal lagoons where concentrations of these pollu-
tants were highest (Pastor et al. 1996).
Perhaps one of the more novel approaches towards
using M. cephalus as an indicator of environmental
change comes from the study of Boglione et al. (2006).
These authors examined skeletal malformations in
mugilids, including M. cephalus, on the basis that a
stressed environment should induce alterations in
normal skeletal development within these species.
Their results showed significant differences among
the type and rate of skeletal abnormalities in the fish
samples, as well as differences in abnormalities
between sites. Another study describes tumorlike
formations in the olfactory organ of M. cephalus in
Sovetskaya Gavan’ Bay and attributed them to
carcinogenic industrial wastes and radioactive con-
tamination of the bay (Byankin 2001).
Parasites on mugilids may also provide an indica-
tion of ecosystem pollution. A study by Dzikowski
et al. (2003) showed that grey mullet in a coastal
polluted site had a significantly lower degree of
parasite infestation when compared to a nearby
unpolluted site. This was attributed to the fact that
some parasites have complex life cycles that usually
involve more than one host and therefore renders them
sensitive to perturbed environments. From the above
discussion it is apparent that there are a wide variety of
options in using mullet species, such as M. cephalus,
as a biomarker tool to help monitor the health of
coastal aquatic environments, with the final choice
being made mainly according to available skills,
technology and financial resources.
Fisheries and aquaculture
Grey mullet are widely exploited (Bacheler et al.
2005) using many different fishing gears such as beach
seines, gill nets, purse seines, trammel nets, staked
nets, shoreline traps, barrage traps, dip nets and cast
nets (Thomson 1963). The harvested mugilids will
vary in species composition, depending primarily on
the availability of different taxa within each region,
but M. cephalus is often an important species in the
catch (Katselis et al. 2003; Chaoui et al. 2006). Adults
are targeted mainly by small-scale fisheries, while
larvae are captured for aquaculture in certain areas.
However, the ability to artificially spawn M. cephalus
for aquaculture purposes was developed more than
40 years ago (Shehadeh and Ellis 1970; Shehadeh
et al. 1973) and the fry have been used to stock
aquaculture ponds in some parts of the world such as
Taiwan (Kuo et al. 1973; Liao 1981; Chang et al.
2000). High mortalities of captured wild fry are a
serious problem and can reduce the viability of
aquaculture operations and local M. cephalus popula-
tions (Tang 1975; Ben-Yami 1981; Bok 1983). The
commercial importance of mugilids depends on the
country, ranging from highly esteemed in Tunisia,
Egypt and Taiwan, to being of little value in parts of
Spain, France and Australia (Gray et al. 2004). In some
parts of the world they are highly prized as bait and in
many countries (e.g. South Africa) they are exten-
sively used as live bait for large piscivorous fishes.
Total world harvesting of mullet species (fisheries
and aquaculture) shows an interesting trend, with
M. cephalus landings increasing significantly between
the mid-1990s and early 2000s but there has been a
decrease in yield since 2004 (Fig. 10). Indeed, the
M. cephalus stocks in eastern Australia by the mid-1900s
were considered by Thomson (1953) to already be
fished to capacity whilst Kesteven (1941) considered
them to be depleted. As so often happens when coastal
fishery yields of a particular species declines, aqua-
culture production of that species then shows an
increasing contribution to the total harvest (Fig. 10).
Flathead mullet represent almost 50 % of all mugilids
harvested globally but these data are only an estimate
due to the inability of fishermen and mariculture
operations in many areas to separate out the different
mullet species. Most mugilid fisheries operate in
coastal lagoons and estuaries, although some fisheries
target marine populations (Panfili et al. 2006).
In certain Asian countries the M. cephalus post larvae
and early juveniles are captured for use in aquaculture,
e.g. in Taiwan the 0?juveniles are cultured to sexual
maturity in order to harvest the roe which has a high
economic value in the region. In other parts of the
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world mullet are sometimes exported to countries
where these species have a higher economic value.
In Greece there are 72 lagoons covering approxi-
mately 55,000 ha, all of which are public domain but
are rented out by the State to individuals or cooper-
atives for fishing. One of the most important fishery
species in these lagoons is M. cephalus, primarily due
to its relatively high commercial value as an edible fish
and because the mature ovaries are used for the
production of dried salted roe, a well-known delicacy
in the Mediterranean region (Katselis et al. 2005).
There are two peaks in M. cephalus landings from fish
barrier traps in the Messolongi-Etoliko lagoon com-
plex, the first from August to October which corre-
sponds to the spawning migration of this species to the
sea, and the second peak during November and
December when a post-spawning inshore migration
occurs (Katselis et al. 2003). Although annual catches
of M. cephalus in the Messolonghi-Etoliko complex
do not show any particular trend during recent years, it
is reported that total annual catches have decreased
from 1,500 to 2,000 tonnes in the 1960s to
1,300–1,500 tonnes in the 1990s (Katselis et al.
2005). Moreover, Koutrakis (2004) has indicated that
fish catches in the lagoons of northern Greece have
decreased further, mainly due to pollution. These
trends, as well as the fact that lagoon fisheries are
based on a practice that negatively impacts the life
cycle of migratory species by removing potential
spawner stock, underline the necessity for integrated
management of lagoonal fishing activities.
Mugil cephalus is a very important species for both
fisheries and aquaculture in Taiwan. During winter,
adult mullet migrate southward from the coastal
waters of mainland China to the offshore waters of
southwestern Taiwan to spawn. Here they are targeted
by fishermen who supply the important mullet roe
industry, which add salt to the female ovaries to make
Botarga caviar, a favourite delicacy in Taiwan and
Japan (Tung 1981). The eggs and larvae that survive
this fishery disperse with the coastal current to the
estuarine nursery areas where fishermen collect the
early juveniles for stocking of aquaculture facilities
(Chang and Tzeng 2000).
Catches of grey mullet in Taiwanese waters fluc-
tuate widely from one year to the next but a general
trend of increasing catches from 1967 until about 1985
was recorded, followed by a virtual collapse of the
fishery, with less than 100,000 fish harvested in 2005
(Panfili et al. 2006). The sharp decline in Taiwanese
catches has been attributed to an expansion in the sizes
of the fishing fleets targeting grey mullet, especially
the recent establishment of a grey mullet fishery by
China (which targets stocks before they migrate into
Taiwanese waters), as well as an increase in sea
surface temperatures associated with global warming
(Hung and Shaw 2006). In contrast to declining wild
stocks of grey mullet, the Taiwanese production of
M. cephalus from aquaculture has shown a consider-
able increase, and is now approximately four times
greater than the yield from capture fisheries. Harvest-
ing of juvenile M. cephalus for restocking of the
aquaculture ponds fluctuates widely, with most being
collected from the northwest and southwest coasts of
Taiwan (Panfili et al. 2006). The clearly defined
emigration period and visibility of adult flathead
mullet shoals as they move through estuaries en route
to their spawning grounds, and the concentrated shoals
of fry immigrating into estuaries, makes this fish
species very vulnerable to over-exploitation.
Fig. 10 Global production of M. cephalus in the world between
1950 and 2009, in terms of fishing activities and aquaculture,
as recorded by the Food and Agriculture Organization
(http://www.fao.org)
668 Rev Fish Biol Fisheries (2012) 22:641–681
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Apart from Taiwan, attempts have also been made
at developing an indigenous fish farming culture using
M. cephalus as one of the species in Eastern Cape
(South Africa) farm dams (Bok 1984b). The flathead
mullet is also one of the fish species produced by farms
in Israel where approximately 1,000 tonnes were
marketed in 1995 (Sarig 1996). In addition, Lake
Kinneret in Israel has been used to stock M. cephalus
for commercial purposes (Bar-Ilan 1975). Similar
plans have been proposed for the Ivory Coast (Albaret
and Legendre 1985) and there is also an extensive
aquaculture in Tunisia based industry that relies on the
transfer of mullet fry (including M. cephalus) from
marine coastal waters to stock continental freshwater
lakes (Khe
´riji et al. 2003). Greece also has an
important mullet aquaculture industry, much of which
is focused on avgotaracho roe production (Katselis
et al. 2005). In contrast to the above extractive
approaches to mullet farming, M. cephalus in Hawaii
are cultured and the fry released into coastal waters in
an attempt to boost future commercial catches of this
species in the wild (Leber and Arce 1996).
Mugilids are the most important group of fishes in
the Mauritanian fishery, with landings estimated
above 14,000 tonnes annually (Bernardon and Vall
2004). This represents 18 % of the total catch, 50 % of
which comprise Mugil capurii and 20 % M. cephalus.
The landings in the Banc d’Arguin National Park are
almost exclusively M. cephalus (98 %) and mullet
catches by pelagic trawlers are estimated at approx-
imately 15,000 tons per year. The mullet fishery in
Mauritania is extremely important economically, with
20 % of the total number of fishermen focused on this
fishery which is the second most consumed fish in
Mauritania, with 95 % of the landings being con-
sumed within the subregion (Bernardon and Vall
2004). In recent years, increasing pressure has been
placed on mullet populations in Mauritania due to the
collection of fish roe for export to Europe.
In Mexico, mugilids are an important gill net
fisheries resource for fishermen along both the Pacific
and Gulf of Mexico coasts. Catches from about 800
fishing boats in estuarine and coastal waters along the
Pacific coast declined from 4,000 tons in 1986 to 3,500
tons in 2002 (SARGARPA 2006). Mugil cephalus is
considered to be fully exploited in this region and
fishing effort regulations are mostly centred around
minimum fish size (31 cm for M. cephalus) and closed
fishing seasons (which differ according to locality).
Mugilid fisheries in the Gulf of Mexico are dominated
on the north-western side by catches of M. cephalus
and by M. curema in Veracruz State waters.
The largest fishery for M. cephalus in South African
waters is found at Lake St Lucia where both legal and
illicit gill net fisheries operate (Mann 1995). Although
M. cephalus is not the primary targeted species, large
numbers of these fish were captured by these opera-
tions since they are concentrated in the northern
compartments of Lake St Lucia where adult
M. cephalus used to be abundant (prior to the closure
of the St Lucia Estuary mouth in 2000). Current South
African government policy aims to create more
equitable access to marine resources and there is
pressure to increase the inshore gill-net fishing effort.
At present, the gill-net fishery in the Western Cape is
confined to the cool temperate west coast where the
southern mullet Liza richardsonii is targeted (Hutch-
ings and Lamberth 2003). Although L. richardsonii
dominates catches in the region, at least 33 by-catch
species (including M. cephalus) have been recorded.
Less formal small-scale fisheries operate in west,
central and east Africa and M. cephalus is often one of
many targeted species. Fisheries in many estuaries
and lagoons on the continent are unregulated and
can be characterised by a multiplicity of gears and a
‘‘free-for-all’’ approach (Koranteng et al. 1998). The
flathead mullet is also harvested by recreational and
subsistence fishermen in some countries using rod and
line. For example, M. cephalus was the most com-
monly captured fish species by anglers in both the
Durban Harbour and Mgeni Estuary shore-fisheries in
South Africa (Pradervand et al. 2003).
There is little doubt that both overfishing and
habitat alteration are resulting in major declines in
M. cephalus stocks around the world. For example,
high fishing pressure in the estuaries and lagoons of
the eastern Adriatic coastal zone, together with land
reclamation and water pollution, have resulted in this
species becoming rare and endangered within the
region (Bartulovic
´et al. 2011). Similarly, increased
nutrient loading and declining salinities in Lake
Manzala (Egypt) have been proposed as possible
reasons for the decline in grey mullet, including
M. cephalus, from 65 % of the total catch during the
1920s to only 2 % of the total catch in the early 1980s
(Khalil 1997). These declines in natural populations
have led to increasing interest in the culture of
M. cephalus (Tamaru et al. 2005).
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Some conclusions and future research directions
The available evidence indicates that M. cephalus
consists of a cluster of ecologically specialized
mugilids, perhaps even a species complex, each of
which has a slightly different ecological niche that is
yet to be defined. In the North West Pacific it would
appear that the three different M. cephalus species
have slightly different temperature preferences, as
well as displaying very different migratory behaviour
between freshwater, estuarine and marine systems.
Future research needs to clarify the ecological niche of
the above species in order to understand the origin and
maintenance of this biodiversity. On a global scale,
this would help to correctly interpret these close
species relationships. All genetic studies to date have
failed to reveal any clear phylogeographic pattern and
new approaches, such as restricted site associated
DNA (RAD) tags (Emerson et al. 2010), are required
to further investigate this issue. Overall, three different
approaches are needed to understand the origin and
biology of M. cephalus:
1. Intra-family phylogenetic relationships may be
investigated using a set of representative species
and genera of the Mugilidae. Specific attention to
the Mugil genus may be guided by results of
Caldara et al. (1996) which suggested a specific
evolution rate within this particular lineage. The
above phylogenetic study would focus on
M. cephalus sampled in the different areas from
within its distributional range.
2. Inferences about the evolutionary processes that
have shaped the genetic diversity and structure
within M. cephalus require a phylogeographic
approach which needs to operate on both a global
and regional scale. In order to provide an evolu-
tionary perspective regarding mechanisms influ-
encing M. cephalus genetics it is important to
identify the detailed phylogeographic structure
(location of genetic discontinuities as well as the
areas of genetic homogeneity) to infer geological
events or physical barriers that impacted past gene
flow within this species. With the above data it
would then be possible to estimate a divergence
time among different M. cephalus populations
and/or species around the world.
3. More population genetic studies are needed at
regional and fine scales using microsatellite
markers to provide an estimation of the dispersive
ability of the species. This approach would also
allow for the delineation of contemporary barriers
to M. cephalus gene flow which is important to
understanding life history traits and the inter-
pretation of variation in terms of plasticity or
adaptive features.
Apart from the potential genetic studies outlined
above, the technology now exists to monitor the
detailed movements of individual fish using acoustic
telemetry. This technology has been implemented
very successfully for a number of larger estuary-
associated marine fish species (Childs et al. 2008) and
it would be appropriate to monitor the movements
(particularly the spawning migrations) of M. cephalus
individuals from around the world. Such studies would
provide us with detailed information on habitat
preferences, home ranges and migrations that are
currently unavailable.
Ecological niche delineation, or migration patterns
by species belonging to the M. cephalus species
complex, can be studied using analyses of specific
isotopes within the otolith or acoustic telemetry. In
addition, stable isotope and fatty acid analyses of body
tissue can assist in identifying food pathways used by
M. cephalus in marine, estuarine and freshwater
environments. All these methods are necessary to
better understand life history variation observed
within the M. cephalus species complex. Indeed, it
would be premature to use M. cephalus species as an
‘indicator’ of littoral environmental changes until the
details surrounding the species complex are unravelled.
Already the analysis of M. cephalus otolith Sr:Ca
ratios have been successfully used as tracers of water
salinity in Taiwan, thus contributing to an understand-
ing of salinity preferences by individuals and popu-
lations at different life-history stages. This technique
now needs to be used more widely on M. cephalus in
order to obtain a better understanding of the environ-
mental requirements and preferences of individual
species within the complex. Studies have revealed that
otolith oxygen isotopes can also be used to trace fish
migrations across different salinities at different stages
of their life cycle (e.g. Meyer-Rochow et al. 1992),
thus opening up a new area for mugilid research.
Integrated studies on the reproductive cycles of the
different M. cephalus species throughout the world are
now required in order to highlight the relationship
670 Rev Fish Biol Fisheries (2012) 22:641–681
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between environmental parameters and the spawning
season of each species. Similarly, studies need to be
undertaken on the possible effects of the environment
on reproductive traits such as fecundity, size at first
maturation, oocyte size, etc. All the above work needs
to be conducted using standardized techniques that
allow for species comparisons at a later stage.
The mechanisms of colonisation and habitat selec-
tion by M. cephalus species within continental waters
are poorly understood. For example, what is the role of
olfactory cues in guiding larval and early juvenile
M. cephalus from the marine environment into
estuarine and riverine nursery habitats? Arising from
the above question, why are the early juveniles of
some flathead mullet species attracted to freshwater
areas, whereas others remain within the estuarine
environment of the same system? Indeed, there may
also be M. cephalus species that do not enter estuaries
or freshwater systems at all and this marine trait may
be a distinguishing feature for these particular species.
Linked to the above distributional patterns is the
differential availability of suitable food resources for
M. cephalus in marine, estuarine and freshwater
environments, and how this affects the trophic
dynamics of the species.
This review has identified the fact that M. cephalus
populations in different parts of the world do not have
uniform life-history traits and that this may be partially
explained if the flathead mullet is a species complex
and not a single species. This is particularly evidenced
by the widely differing spawning seasons, nursery
areas, occupied habitats and migratory behaviour,
even between what appear to be geographically
overlapping populations. In addition, the degree of
connectivity between populations, as evidenced by
gene flow measurements, still requires resolution.
Similarly, the degree to which variations in life history
traits are localised adaptations by the same population
to different environments, or whether they are based
on fundamental genetic differences, is largely
unknown. The reference list at the end of this review
indicates that much work has been conducted on
M. cephalus life-history traits around the world but it
would appear that a proper understanding of these
traits will only be possible once the species complex
issues are resolved.
It is very apparent that further phylogenetic
and population genetic investigations are needed to
understand the origin, distribution and biology of
M. cephalus on a global basis. Collating the different
adaptive responses in term of growth and reproduction
(e.g. fecundity, size at first maturation, oocyte size) for
different populations could constitute a basis for the
characterization of the environmental drivers and
response by the species to these drivers. However, it
is important to note that the different methods used for
life-history trait assessments (e.g. age and reproductive
state) need be standardized in order to facilitate
comparisons at the global level (Panfili et al. 2006,
2007.
There is preliminary evidence to suggest that
M. cephalus represents a good candidate as an ‘indicator’
or ‘sentinel’ species in order to follow littoral envi-
ronmental changes around the world, e.g. representa-
tives of this species show much promise as a possible
indicator of climate change in Queensland, Australia
(Meyncke et al. 2006; Meynecke and Lee 2011) and
Oksyuzyan and Sokolovsky (2003) suggested that
M. cephalus would be a good indicator species for the
state of the environment in the Tumen Estuary (Sea of
Japan). A global observation network coordinating
the use of this species complex as an indicator of the
state of coastal areas could be particularly relevant.
However, as indicated above, it is important that a
resolution of the species complex issue is addressed
before the indicator species can be defined and used as
planned.
Acknowledgments This review was greatly facilitated by
access to published information on M. cephalus collated during
the European Commission 6th Framework Programme, INCO-
CT-2006-026180, MUGIL Project. The funding provided by the
European Commission (EC) and National Research Foundation
(NRF) of South Africa is gratefully acknowledged. We also
thank Susan Abraham for her assistance with the drawing of
Figs. 2,5,6.
References
Abraham M (1971) The ultrastructure of the cell types of the
neurosecretory innervation in the pituitary of Mugil ceph-
alus L. from fresh water, the sea and hypersaline lagoon. 1.
The rostral pars distalis. Gen Comp Endocrinol 17:334–350
Albaret JJ, Legendre M (1985) Biologie et e
´cologie des Mugil-
idae en lagune Ebrie
´(Co
ˆte d’Ivoire). Inte
´re
ˆt potentiel pour
l’aquaculture lagunaire. Rev Hydrobiol Trop 18:281–303
Alikunhi NM, Narayanasamy R, Kandasamy K (2010) Fatty
acids in an estuarine mangrove system. Rev Biol Trop
58(2):577–587
Rev Fish Biol Fisheries (2012) 22:641–681 671
123
Author's personal copy

Ameur B, Bayed A, Bennazou T (2003) Ro
ˆle de la communi-
cation de la lagune de Merja Zerga (Gharb, Maroc) avec
l’Oce
´an Atlantique dans la reproduction d’une population
de Mugil cephalus L. (Poisson Mugilidae). Bull Inst Sci,
Rabat, Sect Sci Vie 25:77–82
Anderson WW (1958) Larval development, growth and
spawning of striped mullet (Mugil cephalus) along the
South Atlantic coast of the United States. Fish Bull Fish
Wildl Serv (US) Res Rep 58:501–519
Anderson AE, Jones EC, Odum HT (1958) Alteration of clay
minerals by digestive process of marine organisms. Sci-
ence 127:190–191
Andree K, Axtner J, Bagley MJ, Barlow EJ, Beebee TJC,
Bennetzen JL, Bermingham E et al (2010) Permanent
genetic resources added to molecular ecology resources
database 1 April 2010 to 31 May 2010. Mol Ecol Res
10:1098–1105
Apekin VS, Vilenskaya NI (1978) A description of the sexual
cycle and state of the gonads during the spawning migra-
tion of the striped mullet, Mugil cephalus. J Ichthyol
18(3):446–456
Arnold EL, Thompson JR (1958) Offshore spawning of the
striped mullet, Mugil cephalus, in the Gulf of Mexico.
Copeia 2:130–132
Avanesov EM (1972) Present spawning conditions of mullets
(genus Mugil) in the Caspian Sea. J Ichthyol 12:419–425
Bacheler NM, Wong RA, Buckel JA (2005) Movements and
mortality rates of striped mullet in North Carolina. N Am J
Fish Manag 25:361–373
Bahnasawy M, Khidr AA, Dheina N (2009) Seasonal variations
of heavy metals concentrations in mullet Mugil cephalus
and Liza ramada (Mugilidae) from Lake Manzala. Egypt.
J Appl Sci Res 5(7):845–852
Bahri S, Marques A (1996) Myxosporean parasites of the genus
Myxobolus from Mugil cephalus in Ichkeul Lagoon,
Tunisia: Description of two new species. Dis Aquat Org
27(2):115–122
Baker TG, Pante E, De Buron I (2005) Co-occurrence of Nao-
branchia lizae (Copepoda) and Metamicrocotyla macra-
cantha (Monogenea), gill parasites of the striped mullet
Mugil cephalus. Parasitol Res 97(6):515–520
Baker TG, Pante E, Levesque EM, Roumillat WA, De Buron I
(2008) Metamicrocotyla macracantha, a polyopisthocoty-
lid gill parasite of the striped mullet, Mugil cephalus:
Population dynamics in South Carolina estuaries. Parasitol
Res 102(5):1085–1088
Bar-Ilan M (1975) Stocking of Mugil capito and Mugil cephalus
and their commercial catch in Lake Kinneret. Aquaculture
5(1):85–89
Barman UK, Jana SN, Garg SK, Bhatnagar A, Arasa ART
(2005) Effect of inland water salinity on growth, feed
conversion efficiency and intestinal enzyme activity in
growing grey mullet, Mugil cephalus (Linn.): Field and
laboratory studies. Aquacul Int 13(3):241–256
Bartulovic
´V, Dulc
ˇic
´J, Matic
´-Skoko S, Glamuzina B (2011)
Reproductive cycles of Mugil cephalus,Liza ramada and
Liza aurata (Teleostei: Mugilidae). J Fish Biol 78(7):
2067–2073
Belyanina TN (1995) Ichthyoplankton of Syrian surface waters
(from a fall survey 1993). J Ichthyol 35(3):78–85
Ben-Yami M (1981) Handling, transportation and stocking of
fry. In: Oren OH (ed) Aquaculture of grey mullets. Inter-
national Biological Programme 26, Cambridge University
Press, Cambridge
Bere R (1936) Parasitic copepods from Gulf of Mexico fish. Am
Midl Nat 17:577–625
Bernardon M, Vall M (2004) Le mulet en Mauritanie: biologie,
e
´cologie, pe
ˆche et ame
´nagement. FIBA IUCN PRCM,
Arles
Blaber SJM (1975) The shedding of gill raker filaments in grey
mullet (Mugilidae). S Afr J Sci 71:377
Blaber SJM (1976) The food and feeding ecology of Mugilidae
in the St Lucia lake system. Biol J Linn Soc 8:267–277
Blaber SJM (1977) The feeding ecology and relative abundance
of mullet (Mugilidae) in Natal and Pondoland estuaries.
Biol J Linn Soc 9:259–275
Blaber SJM (1987) Factors affecting recruitment and survival of
mugilids in estuaries and coastal waters of southern Africa.
Amer Fish Soc Symp 1:507–518
Blaber SJM, Whitfield AK (1976) Large scale mortality of fish
at St Lucia. S Afr J Sci 72:218
Blaber SJM, Whitfield AK (1977) The feeding ecology of
juvenile mullet (Mugilidae) in south east African estuaries.
Biol J Linn Soc 9:277–284
Blanc-Livini M, Abraham M (1970) The influence of environ-
mental salinity on the prolactin and gonadotrophin
secreting regions in the pituitary of Mugil (Teleostei). Gen
Comp Endocrinol 14:184–197
Blel H, Panfili J, Guinand B, Berrebi P, Said K, Durand J-D
(2010) Selection footprint at the first intron of the Prl gene
in natural populations of the flathead mullet (Mugil ceph-
alus, L. 1758). J Exp Mar Biol Ecol 387:60–67
Boglione C, Costa C, Giganti M, Cecchetti M, Dato PD, Scardi
M, Cataudella S (2006) Biological monitoring of wild
thicklip grey mullet (Chelon labrosus), golden grey mullet
(Liza aurata), thinlip mullet (Liza ramada) and flathead
mullet (Mugil cephalus) (Pisces: Mugilidae) from different
Adriatic sites: meristic counts and skeletal anomalies. Ecol
Ind 6(4):712–732
Bok AH (1979) The distribution and ecology of two mullet
species in some freshwater rivers in the eastern Cape, South
Africa. J Limnol Soc S Afr 5:97–102
Bok AH (1983) The demography, breeding biology and man-
agement of two mullet species (Pisces: Mugilidae) in the
Eastern Cape, South Africa. PhD dissertation, Rhodes
University, Grahamstown
Bok AH (1984a) Freshwater mullet in the Eastern Cape.
A strong case for fish ladders. Nat 28:31–35
Bok AH (1984b) Extensive culture of two mullet species in
freshwater impoundments in the eastern Cape. S Afr J Zool
9:31–36
Breder CM (1959) Studies on social groupings in fishes. Bull
Am Mus Nat Hist 117:399–481
Breder CM (1962) Interactions between the fishes Mugil and
Lagodon. Copeia 1962(3):662–663
Briggs JC (1960) Fishes of worldwide (circumtropical) distri-
bution. Copeia 1960:171–180
Brownell CL (1979) Stages in the early development of 40
marine fish species with pelagic eggs from the Cape of
Good Hope. Ichthy Bull JLB Smith Inst Ichthyol 40:1–84
672 Rev Fish Biol Fisheries (2012) 22:641–681
123
Author's personal copy

Brulhet J (1975) Observations on the biology of Mugil cephalus
ashenteensis and the possibility of its aquaculture on the
Mauritian coast. Aquaculture 5:271–281
Brusle J, Brusle S (1977) Les muges de Tunisie: pe
ˆche lagunaire
et biologie de la reproduction de trois espe
`ces (Mugil
capito,Mugil cephalus et Mugil chelo) des lacs d’Ichkeul
et de Tunis. Rapp Comm Int Expl Mer Me
´dit 24:101–130
Brusle J, Cambrony M (1992) Mediterranean lagoons: favour-
able nurseries and/or dangerous traps for juvenile euryha-
line fish? Critical analysis of the growth of grey mullet
populations from several brackish water lagoons in the
Languedoc-Roussillon during the first year of life. Vie
Milieu 42(2):193–205
Byankin AG (2001) Tumorlike formations in the olfactory organ
of the striped mullet Mugil cephalus from Sovetskaya Ga-
van’ Bay, Sea of Japan. Russ J Mar Biol 27(5):333–335
Cadenat J (1954) Notes d’ichthyologie ouest-africaine. VII. sur
les mulets de la co
ˆte occidentale d’Afrique. Bull Inst Fr Afr
2:584–591
Caldara F, Bargelloni L, Ostellari L, Penzo E, Colombo L,
Patarnello T (1996) Molecular phylogeny of grey mullets
based on mitochondrial DNA sequence analysis: evidence
of a differential rate of evolution at the intrafamily level.
Mol Phylogen Evol 6:416–424
Callinan RB, Keep JA (1989) Bacteriology and parasitology of
red spot disease in sea mullet, Mugil cephalus L., from
eastern Australia. J Fish Dis 12(4):349–356
Callinan RB, Fraser GC, Virgona JL (1989) Pathology of red
spot disease in sea mullet, Mugil cephalus L. from eastern
Australia. J Fish Dis 12(5):467–479
Cameron JN (1970) The influence of environmental variables on
the hematology of pinfish (Lagodon rhomboides) and
striped mullet (Mugil cephalus). Comp Biochem Physiol
32(2):175–192
Campton DE, Mahmoudi B (1991) Allozyme variation and
population structure of striped mullet (Mugil cephalus)in
Florida. Copeia 1991:485–492
Cardona L (1996) Microalgae selection by mullets (Mugil
cephalus and Liza ramada) in Israeli semi-intensive fish
ponds. Isr J Aquacult-Bamid 48(3):165–173
Cardona L (2000) Effects of salinity on the habitat selection and
growth performance of Mediterranean flathead grey mullet
Mugil cephalus (Osteichthyes, Mugilidae). Est Coast Shelf
Sci 50:727–737
Cardona L (2001) Non-competitive coexistence between Med-
iterranean grey mullet: evidence from seasonal changes in
food availability, niche breadth and trophic overlap. J Fish
Biol 59:729–744
Cardona L (2006) Habitat selection by grey mullets (Oste-
ichthyes: Mugilidae) in Mediterranean estuaries: the role of
salinity. Sci Mar 70:443–455
Cardona L, Torras X, Gisbert E, Castello
´F (1996) The effect of
striped grey mullet (Mugil cephalus L.) on freshwater
ecosystems. Isr J Aquacult-Bamid 484(4):179–185
Cardona L, Hereu B, Torras X (2008) Juvenile bottlenecks and
salinity shape grey mullet assemblages in Mediterranean
estuaries. Est Coast Shelf Sci 77(4):623–632
Carr BA, Aldrich DV (1982) Population density effects on the
behaviour and feeding of young striped mullet (Mugil
cephalus) in 37.8-liter aquaria. J World Mar Soc
13:254–260
Chang CW, Tzeng WN (2000) Species composition and sea-
sonal occurrence of mullets (Pisces, Mugilidae) in the
Tanshui estuary northwest Taiwan. J Fish Soc Taiwan
27:253–262
Chang CW, Tzeng WN, Lee YC (2000) Recruitment and
hatching dates of grey mullet (Mugil cephalus L.) juveniles
in the Tanshui estuary of northwest Taiwan. Zool Stud
39:99–106
Chang CW, Iizuka Y, Tzeng WN (2004a) Migratory environ-
mental history of the grey mullet Mugil cephalus as
revealed by otolith Sr:Ca ratios. Mar Ecol Progr Ser
269:277–288
Chang CW, Lin SH, Iizuka Y, Tzeng WN (2004b) Relationship
between Sr:Ca ratios in otoliths of grey mullet Mugil
cephalus and ambient salinity: validation, mechanisms,
and applications. Zool Stud 43:74–85
Chaoui L, Kara MH, Faure E
´, Quignard J-P (2006) The fish
fauna of Mellah Lagoon (north-east Algeria): Diversity,
production and commercial catches analysis. Cybium
30(2):123–132
Childs A-R, Cowley PD, Næsje TF, Booth AJ, Potts WM,
Thorstad EB, Økland F (2008) Do environmental factors
influence the movement of an estuarine fish? A case study
using acoustic telemetry. Est Coast Shelf Sci 78:227–236
Chubb CF, Potter IC, Lenanton RC, Wallace J (1981) Age
structure, growth rates and movements of sea mullet, Mugil
cephalus L., and yellow-eye mullet, Aldrichetta forsteri
(Valenciennes), in the Swan-Avon River system, Western
Australia. Aust J Mar Freshw Res 32:605–628
Ciccotti E, Marino G, Pucci P, Cataldi E, Cataudella S (1994)
Acclimation trial of Mugil cephalus juveniles to freshwa-
ter: morphological and biochemical aspects. Environ Biol
Fish 43:163–170
Clancy GP (2005) The diet of the osprey (Pandion haliaetus)on
the north coast of New South Wales. Emu 105(1):87–91
Collins MR (1981) The feeding periodicity of striped mullet,
Mugil cephalus L., in two Florida habitats. J Fish Biol
19(3):307–315
Collins MR, Stender BW (1989) Larval stripped mullet (Mugil
cephalus) and white mullet (Mugil curema) off the south-
eastern United States. Bull Mar Sci 45:580–589
Corti M, Crosetti D (1996) Geographic variation in the grey
mullet: a geometric morphometric analysis using partial
warp scores. J Fish Biol 48:255–269
Crosetti D, Avise JC, Placidi F, Rossi AR, Sola L (1993) Geo-
graphic variability in the grey mullet Mugil cephalus:
preliminary results of mtDNA and chromosome analyses.
Aquaculture 111:95–101
Crosetti D, Nelson WS, Avise J (1994) Pronounced genetic
structure of mitochondrial DNA among populations of the
circumglobally distributed grey mullet (Mugil cephalus
Linnaeus). J Fish Biol 44:47–58
Cyrus DP, Blaber SJM (1987) The influence of turbidity on
juvenile marine fishes in estuaries. Part 1. Field studies at
Lake St Lucia on the southeastern coast of Africa. J Exper
Mar Biol Ecol 109:53–70
Darnell RM (1961) Trophic spectrum of an estuarine commu-
nity, based on studies of Lake Pontchartrain, Louisiana.
Ecology 42:553–568
De Silva SS (1980) Biology of juvenile grey mullet: a short
review. Aquaculture 19:21–36
Rev Fish Biol Fisheries (2012) 22:641–681 673
123
Author's personal copy

De Silva SS, Perera PAB (1976) Studies of the young grey mullet,
Mugil cephalusL. I. Effects of salinityon food intake, growth
and food conversion. Aquaculture 7(4):327–338
De Silva SS, Silva EIL (1979) Biology of young grey mullet,
Mugil cephalus L., populations of a coastal lagoon in Sri
Lanka. J Fish Biol 15:9–20
De Silva SS, Wijeyaratne MJS (1977) Studies on the biology of
young grey mullet, Mugil cephalus L. II. Food intake and
feeding. Aquaculture 12(2):157–167
Ditty JG, Shaw RF (1996) Spatial and temporal distribution of
larval striped mullet (Mugil cephalus) and white mullet (M.
curema, Family: Mugilidae) in the northern Gulf of Mex-
ico, with notes on mountain mullet Agonostomus monti-
cola. Bull Mar Sci 59(2):271–288
Dural M, Goksu MZL, Ozak AA, Derici B (2006) Bioaccu-
mulation of some heavy metals in different tissues of
Dicentrarchus labrax L, 1758, Sparus aurata L, 1758 and
Mugil cephalus L, 1758 from the Camlik lagoon of the
eastern coast of Mediterranean (Turkey). Environ Monit
Assess 118:65–74
Durand J-D, Shen K-N, Chen W-J, Jamandre B-W, Blel H, Diop
K, Nirchio M, Garcı
´a de Leo
´n FJ, Whitfield AK, Chang
C-W, Borsa P (2012) Systematics of the grey mullet (Te-
leostei: Mugiliformes: Mugilidae): molecular phylogenetic
evidence challenges two centuries of morphology-based
taxonomic studies. Mol Phylogen Evol. doi:10.1016/
j.ympev.2012.03.0006
Dzikowski R, Paperna I, Diamant A (2003) Use of fish parasite
species richness indices in analysing anthropogenically
influenced coastal marine ecosystems. Helgoland Mar Res
57:220–227
Eda H, Murashige R, Oozeki Y, Hagiwara A, Eastham B, Bass
P, Tamaru CS, Lee C-S (1990) Factors affecting intensive
larval rearing of striped mullet, Mugil cephalus. Aquacul-
ture 91:281–294
Eggold BT, Motta PJ (1992) Ontogenetic dietary shifts and
morphological correlates in striped mullet, Mugil cephalus.
Environ Biol Fish 34(2):139–158
Eiras JC, D’Souza J (2004) Myxobolus goensis n. sp. (Myxozoa,
Myxosporea, Myxobolidae), a parasite of the gills of Mugil
cephalus (Osteichthys, Mugilidae) from Goa, India. Para-
site 11(3):243–248
El Hafidi F, Berrada-Rkhami O, Benazzou T, Gabrion C (1998)
Microhabitat distribution and coexistence of Microcotyli-
dae (Monogenea) on the gills of the striped mullet Mugil
cephalus: Chance or competition. Parasitol Res 84(4):
315–320
El-Gharabawy MM, Assem SS (2006) Spawning induction in
the Mediterranean grey mullet Mugil cephalus and larval
development stages. Afr J Biotech 5(19):1836–1845
Elsheikha HM, Elshazly AM (2008) Preliminary observations
on infection of brackish and fresh water fish by heterophyid
encysted metacercariae in Egypt. Parasitol Res 103(4):
971–977
Emerson KJ, Merz CR, Catchen JM, Hohenlohe PA, Cresko
WA, Bradshaw WE, Holzapfel C (2010) Resolving post-
glacial phylogeography using high-throughput sequencing.
Proc Nat Acad Sci 107:16196–16200
Erman F (1959) Observations on the biology of the common
grey mullet Mugil cephalus L. Proc Gen Fish Coun Medit
5:157–169
Eschmeyer WN, Fricke R (eds) (2011) Catalog of fishes.
http://research.calacademy.org/research/ichthyology/
catalog/fishcatmain.asp
Faouzi H (1938) Quelques aspects de la biologie des Muges en
Egypte. Rapp Comm Int Expl Sci Mer Me
´diter 11:63–68
Faye N, Kpatcha TK, Fall M, Toguebaye BS (1997) Heart
infections due to myxosporean (myxozoa) parasites in
marine and estuarine fishes from Senegal. Bull Eur Assoc
Fish Pathol 17:115–117
Ferreira M, Antunes P, Gil O, Vale C, Reis-Henriques MA
(2004) Organochlorine contaminants in flounder (Platich-
thys flesus) and mullet (Mugil cephalus) from Douro
Estuary, and their use as sentinel species for environmental
monitoring. Aquat Toxicol 69:347–357
Ferreira M, Moradas-Ferreira P, Reis-Henriques MA (2005)
Oxidative stress biomarkers in two resident species, mullet
(Mugil cephalus) and flounder (Platichthys flesus), from a
polluted site in River Douro Estuary, Portugal. Aquat
Toxicol 71:39–48
Ferreira M, Moradas-Ferreira P, Reis-Henriques MA (2007)
The effect of long-term depuration on levels of oxidative
stress biomarkers in mullets (Mugil cephalus) chronically
exposed to contaminants. Mar Environ Res 64(2):181–190
Finucane JH, Collins LA, Barger LE (1978) Spawning of the
striped mullet Mugil cephalus in the northwestern Gulf of
Mexico. Northeast Gulf Sci 2:148–150
Fitzsimmons SD, Warburton K (1992) Fish movement behav-
iour: variability within and between groups. Behav Process
26:211–216
Fraga E, Schneider H, Nirchio M, Santa-Brigida E, Rodrigues-
Filho LF, Sampaio I (2007) Molecular phylogenetic anal-
yses of mullets (Mugilidae, Mugiliformes) based on two
mitochondrial genes. J Appl Ichthyol 23:598–604
Frodello JP, Raqbi A, Mattei X, Viale D, Marchand B (2001)
Quantification of macrophage aggregates in the liver of
Mugil cephalus. J Submicr Cytol Path 33(4):473–476
Fujita S, Kinoshita I, Takahashi I, Azuma K (2002) Species
composition and seasonal occurrence of fish larvae and
juveniles in the Shimanto Estuary, Japan. Fish Sci
68(2):364–370
Fury CA, Harrison PL (2011) Seasonal variation and tidal
influences on estuarine use by bottlenose dolphins (Tursi-
ops aduncus). Est Coast Shelf Sci 93(4):389–395
Gehrke PC, Gilligan DM, Barwick M (2001) Fish communities
and migration in the Shoalhaven River—before construc-
tion of a fishway. NSW Fisheries Final Report Series 26,
Cronulla Fisheries Centre, Australia
Gilbert CR (1993) Geographic distribution of the striped mullet
(Mugil cephalus Linnaeus) in the Atlantic and eastern
Pacific oceans. Fla Sci 56:204–210
Gisbert E, Cardona L, Castello
´F (1995) Competition between
mullet fry. J Fish Biol 47:414–420
Gisbert E, Cardona L, Castello
´F (1996) Resource partitioning
among planktivorous fish larvae and fry in a Mediterranean
coastal lagoon. Est Coast Shelf Sci 43(6):723–735
Grant CJ, Spain AV (1975) Reproduction growth and size
allometry of Mugil cephalus Linnaeus (Pisces Mugilidae)
from north Queensland inshore waters. Aust J Zool
23:181–201
Gray CA, Johnson DD, Young DJ, Broadhurst MK (2004)
Discards from the commercial net fishery for dusky
674 Rev Fish Biol Fisheries (2012) 22:641–681
123
Author's personal copy

flathead, Platycephalus fuscus, in New South Wales,
Australia: Spatial variability and initial effects of change in
minimum legal length of target species. Fish Manag Ecol
11(5):323–333
Grobler DF, Badenhorst JE, Kempster PL (1996) PCBs, chlo-
rinated hydrocarbon pesticides and chlorophenols in the
Isipingo Estuary, Natal, Republic of South Africa. Mar
Pollut Bull 32(7):572–575
Hadwen WL, Russell GL, Arthington AH (2007) Gut content-
and stable isotope-derived diets of four commercially and
recreationally important fish species in two intermittently
open estuaries. Mar Freshw Res 58(4):363–375
Hamza AK (1999) A study on some biological characteristics of
Mugil cephalus (L.) in Bardawil Lake, Egypt. J Appl Ich-
thyol 15:135–137
Harel M, Atia SB, Zlotkin V, Tandler A (1998) Mass production
of grey mullet, Mugil cephalus: Effects of environmental
and nutritional factors on larval performance. Isr J Aqua-
cult-Bamid 50(3):91–98
Hassani LH, Frenich AG, Benajiba MH, Rodriguez MJG, Vidal
JLM, Debdoubi A (2006) Assessment of butyltin and
phenyltin pollution in the sea mullet, Mugil cephalus, along
the Moroccan and Spanish coasts (Mediterranean Sea).
Arch Environ Con Tox 51:608–614
Hendricks LJ (1961) The striped mullet, Mugil cephalus Lin-
naeus. In: Walker BW (ed) The ecology of the Salton Sea,
California, in relation to the Sportfishery. Fish Bull Calif
113:95–104
Heras S, Rolda
´n MI, Castro MG (2009) Molecular phylogeny
of Mugilidae fishes revised. Rev Fish Biol Fish 19(2):
217–231
Hickling CF (1970) A contribution to the natural history of the
English grey mullets (Pisces, Mugilidae). J Mar Biol Assoc
UK 50:609–633
Ho J-S, The Do T (1982) Two species of Ergasilidae (Copepoda:
Poecilostomatoida) parasitic on the gills of Mugil cephalus
Linnaeus (Pisces: Teleostei), with the proposition of a new
genus Dermoergasilus. Hydrobiologia 89:247–252
Hoese HD (1985) Jumping mullet—the internal diving bell
hypothesis. Environ Biol Fish 13(4):309–314
Hossler FE (1980) Gill arch of the mullet, Mugil cephalus. III.
Rate of response to salinity change. Am J Physiol-Reg I
7(2):160–164
Hotos GV, Vlahos N (1998) Salinity tolerance of Mugil ceph-
alus and Chelon labrosus (Pisces: Mugilidae) fry in
experimental conditions. Aquaculture 167:329–338
Hsu CC, Han YS, Tzeng W-N (2007) Evidence of flathead
mullet Mugil cephalus L. spawning in waters northeast of
Taiwan. Zool Stud 46:717–725
Hsu CC, ChangCW, Iisuka Y, Tzeng W-N (2009) A growth check
deposited at estuarine arrival in otoliths of juvenile flathead
mullet (Mugil cephalus L.). Zool Stud 48(3):315–324
Huang C-S, Weng C-F, Lee S-C (2001) Distinguishing two
types of grey mullet, Mugil cephalus L. (Mugiliformes:
Mugilidae), by using glucose-6-phosphate isomerase (GPI)
allozymes with special reference to enzyme activities.
J Comp Physiol B 171:387–394
Hung C-M, Shaw D (2006) The impact of upstream catch and
global warming on the grey mullet fishery in Taiwan: A
non-cooperative game analysis. Mar Res Econ 21:285–300
Hutchings K, Lamberth SJ (2003) Likely impacts of an eastward
expansion of the inshore gill-net fishery in the Western
Cape, South Africa: implications for management. Mar
Freshw Res 54:39–56
Hutton RF, Sogandares-Bernal F (1959) Further notes on
Trematoda encysted in Florida mullets. Q Jl Fla Acad Sci
21:329–334
Iba
´n
˜ez AL (1993) Coexistence of Mugil cephalus and Mugil
curema in a coastal lagoon of the Gulf of Mexico. J Fish
Biol 42(6):959–961
Iba
´n
˜ez AL, Gallardo-Cabello M (1996a) Total and natural
mortality of Mugil cephalus and M. curema V. (Pisces:
Mugilidae) in Tamiahua lagoon, Veracruz. I. Selectivity.
Hydrobiol 6:9–16
Iba
´n
˜ez AL, Gallardo-Cabello M (1996b) Age determination
of the grey mullet Mugil cephalus and the white mullet
M. curema V. (Pisces: Mugilidae) in Tamiahua lagoon,
Veracruz. Cienc Mar 22:329–345
Iba
´n
˜ez AL, Gallardo-Cabello M (2004) Reproduction of Mugil
cephalus and M. curema (Pisces: Mugilidae) from a coastal
lagoon in the Gulf of Mexico. Bull Mar Sci 75:37–49
Iba
´n
˜ez AL, Gallardo-Cabello M (2005) Identification of two
Mugilidae species, Mugil cephalus and M. curema (Pisces:
Mugilidae), using the ctenii of their scales. Bull Mar Sci
77(2):305–307
Iba
´n
˜ez AL, Gutierrez-Benitez O (2004) Climate variables and
spawning migrations of the striped mullet and white mullet
in the north-western area of the Gulf of Mexico. J Fish Biol
65:622–631
Iba
´n
˜ez AL, Gutie
´rrez-Benı
´tez O (2004) Climate variables and
spawning migrations of the striped mullet and white mullet
in the north-western area of the Gulf of Mexico. J Fish Biol
65:822–831
Iba
´n
˜ez AL, Lleonart J (1996) Relative growth and comparative
morphometrics of Mugil cephalus L. and M. curema V. in
the Gulf of Mexico. Sci Mar 60:361–368
Iba
´n
˜ez AL, Gallardo-Cabello M, Carrara XC (1999) Growth
analysis of striped mullet, Mugil cephalus, and white
mullet, M. curema (Pisces: Mugilidae), in the Gulf of
Mexico. Fish B-NOAA 97:861–872
Iba
´n
˜ez AL, Cowx IG, O’Higgins P (2007) Geometric morpho-
metric analysis of fish scales for identifying genera, spe-
cies, and local populations within the Mugilidae. Can J Fish
Aquat Sci 64(8):1091–1100
Idyll CP, Sutton JW (1952) Results of the first year’s tagging of
mullet, Mugil cephalus L., on the west coast of Florida.
Trans Am Fish Soc 81:69–77
Inoue T, Suda Y, Sano M (2005) Food habits of fishes in the surf
zone of a sandy beach at Sanrimatsubara, Fukuoka Pre-
fecture, Japan. Ichthyol Res 52(1):9–14
Jamandre BW, Durand J-D, Tzeng WN (2009) Phylogeography
of the flathead mullet Mugil cephalus in the north-west
Pacific as inferred from the mtDNA control region. J Fish
Biol 75(2):393–407
Jana SN, Garg SK, Patra BC (2004) Effect of periphyton on
growth performance of grey mullet, Mugil cephalus
(Linn.), in inland saline groundwater ponds. J Appl Ich-
thyol 20:110–117
Jithendran KP, Kannappan S (2010) A short note on heavy
infection of acanthocephalan worm (Neoechinorhynchus
Rev Fish Biol Fisheries (2012) 22:641–681 675
123
Author's personal copy

agilis) in grey mullet, Mugil cephalus. J Paras Dis 34(2):
99–101
Johnson DW (1972) Variations in the interrenal and corpuscle of
Stannius of Mugil cephalus from the Colorado River and
estuary. Gen Comp Endocrinol 19:7–25
Johnson DW, McClendon EL (1970) Differential distribution of
striped mullet, Mugil cephalus Linnaeus. Calif Fish Game
56(2):138–139
Kalavati C, Anuradha I (1995) A new myxosporean, Bipteria
indica sp. n. (Myxospora: Sinuolineidae) from the gall
bladder of the striped mullet, Mugil cephalus Linnaeus.
Acta Protozool 34(4):307–309
Karouna-Renier NK, Snyder RA, Lange T, Gibson S, Allison
JG, Wagner ME, Ranga Rao K (2011) Largemouth bass
(Micropterus salmoides) and striped mullet (Mugil ceph-
alus) as vectors of contaminants to human consumers in
northwest Florida. Mar Environ Res 72(3):96–104
Katselis G, Koutsikopoulos C, Dimitriou E, Rogdakis Y (2003)
Spatial patterns and temporal trends in the fishery landing
of the Messolonghi-Etoliko lagoon system (western Greek
coast). Sci Mar 67:501–511
Katselis G, Koutsikopoulos C, Rogdakis Y, Lachanas T, Dim-
itriou E, Vidalis K (2005) A model to estimate the annual
production of roes (avgotaracho) of flathead mullet (Mugil
cephalus) based on the spawning migration of species. Fish
Res 75:138–148
Katz T, Herut B, Genin A, Angel DL (2002) Gray mullets
ameliorate organically enriched sediments below a fish
farm in the oligotrophic Gulf of Aqaba (Red Sea). Mar Ecol
Progr Ser 234:205–214
Ke HM, Lin WW, Kao HW (2009) Genetic diversity and dif-
ferentiation of grey mullet (Mugil cephalus) in the coastal
waters of Taiwan. Zool Sci 26:421–428
Kesteven GL (1941) Conserving the mullet catch. Fish Bull
West Aust 1:1–14
Kesteven GL (1942) Studies on the biology of the Australian
mullet. 1. Account of the fishery and preliminary statement
of the biology of Mugil dobula Gunther. Bull Coun Scient
Ind Res Melb 157:1–147
Kesteven GL (1953) Further results of tagging sea mullet. Aust J
Mar Freshw Res 4:251–306
Khalil MT (1997) Changes in the mullet fishery of Lake
Manzala, Egypt. Int J Salt Lake Res 5(3):241–251
Khe
´riji S, El Cafsi M, Masmoudi W, Castell JD, Romdhane MS
(2003) Salinity and temperature effects on the lipid com-
position of mullet sea fry (Mugil cephalus, Linn, 1758).
Aquacult Int 11(6):571–582
Kilby JD (1949) A preliminary report on the young striped
mullet (Mugil cephalus Linnaeus) in two gulf areas of
Florida. Quart J Fla Acad 11:7–23
Kilby JD (1955) The fishes of two gulf coastal marsh areas of
Florida. Tulane Stud Zool 2:175–247
Kirby J, Maher W, Krikowa F (2001a) Selenium, cadmium,
copper, and zinc concentrations in sediment and mullet
(Mugil cephalus) from the southern basin of Lake Macqua-
rie, NSW, Australia. Arch Environ Con Tox 40(2):246–256
Kirby J, Maher W, Harasti D (2001b) Changes in selenium,
copper, cadmium, and zinc concentrations in mullet (Mugil
cephalus) from the southern basin of Lake Macquarie,
Australia, in response to alteration of a coal-fired power
station fly ash handling procedures. Arch Environ ConTox
41(2):171–181
Koranteng KA, Ofori-Danson PK, Entsua-Mensah M (1998)
Comparative study of the fish and fisheries of three coastal
lagoons in West Africa. Int J Ecol Environ Sci 24(4):
371–382
Koutrakis E (2004) Temporal occurrence and size distribution
of grey mullet juveniles (Pisces, Mugilidae) in the estua-
rine systems of the Strymonikos Gulf (Greece). J Appl
Ichthyol 20:76–78
Kowarsky J, Ross AH (1981) Fish movement upstream through
a central Queensland (Fizroy River) coastal fishway. Aust J
Mar Freshw Res 32:93–109
Kraul S (1983) Results and hypotheses for the propagation of the
grey mullet, Mugil cephalus L. Aquaculture 30:273–284
Kuo C-M, Shedadeh ZH, Milesen KK (1973) A preliminary
report on the development, growth and survival of labo-
ratory reared larvae of grey mullet Mugil cephalus L. J Fish
Biol 5:459–470
Kuo C-M, Nash CE, Shehadeh ZH (1974) The effects of tem-
perature and photoperiod on ovarian development in cap-
tive grey mullet (Mugil cephalus L). Aquaculture 3:25–43
Kuthalingham MDK (1966) A contribution to the life history
and feeding habits of Mugil cephalus L. Treubia 27:11–32
Larson ET, Shanks AL (1996) Consumption of marine snow by
two species of juvenile mullet and its contribution to their
growth. Mar Ecol Progr Ser 130:19–28
Lawson EO, Jimoh AA-A (2010) Aspects of the biology of grey
mullet, Mugil cephalus, in Lagos Lagoon, Nigeria. AACL
Bioflux 3(3):181–193
Laxmi Priya S, Senthilkumar B, Hariharan G, Paneer Selvam A,
Purvaja R, Ramesh R (2011) Bioaccumulation of heavy
metal in mullet (Mugil cephalus) and oyster (Crassostrea
madrasensis) from Pulicat lake, south east coast of India.
Toxicol Ind Health 27(2):117–126
Leber KM, Arce SM (1996) Stock enhancement in a commercial
mullet, Mugil cephalus L., fishery in Hawaii. Fish Manag
Ecol 3(3):261–278
Leber KM, Arce SM, Sterritt DA, Brennan NP (1996) Marine
stock enhancement potential in nursery habitats of striped
mullet, Mugil cephalus, Hawaii. Fish B-NOAA 94(3):
452–471
Leber KM, Blankenship HL, Arce SM, Brennan NP (1997)
Influence of release season on the size-dependent survival
of cultured striped mullet, Mugil cephalus, in a Hawaiian
estuary. Fish B-NOAA 95(2):267–279
Lebreton B, Richard P, Parlier EP, Guillou G, Blanchard GF
(2011) Trophic ecology of mullets during their spring
migration in a European saltmarsh: A stable isotope study.
Est Coast Shelf Sci 91:502–510
Lee C-S, Menu B (1981) Effects of salinity on egg development
and hatching in the grey mullet, Mugil cephalus L. J Fish
Biol 19(2):179–188
Lee C-S, Tamaru CS, Kelley CD, Moriwake A, Miyamoto GT
(1992) The effect of salinity on the induction of spawning
and fertilization in the striped mullet, Mugil cephalus.
Aquaculture 102:289–296
Lester RJG, Rawlinson SE, Weaver LC (2009) Movement of sea
mullet Mugil cephalus as indicated by a parasite. Fish Res
96:129–132
676 Rev Fish Biol Fisheries (2012) 22:641–681
123
Author's personal copy

Liao IC (1981) Cultivation methods. In: Oren OH (ed) Aqua-
culture of grey mullets. Cambridge University Press,
Cambridge
Liordos V, Goutner V (2009) Sexual differences in the diet of
great cormorants Phalacrocorax carbo sinensis wintering
in Greece. Eur J Wildl Res 55(3):301–308
Liu J-Y, Brown CL, Yang T-B (2009) Population genetic
structure and historical demography of grey mullet, Mugil
cephalus, along the coast of China, inferred by analysis of
the mitochondrial control region. Biochem Syst Ecol
37(5):556–566
Liu J-Y, Brown CL, Yang T-B (2010) Phylogenetic relation-
ships of mullets (Mugilidae) in China seas based on partial
sequences of two mitochondrial genes. Biochem Syst Ecol
38(4):647–655
Livi S, Sola L, Crosetti D (2011) Phylogeographic relationships
among worldwide populations of the cosmopolitan marine
species, the striped gray mullet (Mugil cephalus), investi-
gated by partial cytochrome b gene sequences. Biochem
Syst Ecol 39(2):121–131
Lupatsch I, Katz T, Angel DL (2003) Assessment of the removal
efficiency of fish farm effluents by grey mullets: A nutri-
tional approach. Aquac Res 34(15):1367–1377
Luther G (1962) The food habits of Liza macrolepis (Smith) and
Mugil cephalus L. (Mugilidae). Indian J Fish 9:604–626
Luther G (1964) On the shedding of gill-raker processes in grey
mullets. J Mar Biol Assoc India 6:251–256
Maeno Y, Nagasawa K, Sorimachi M (1993) Kudoa intestinalis
n. sp. (Myxosporea: Multivalvulida) from the intestinal
musculature of the striped mullet, Mugil cephalus, from
Japan. J Parasitol 79(2):190–192
Maı
´llo-Bello
´n P-A, Marques A, Gracia-Royo MP (2011)
Myxosporean infection of grey mullet in the Ebro Delta:
Identification and ultrastructure of Myxobolus ichkeulensis
Bahri and Marques, 1996 infecting the gills of Mugil
cephalus L. Acta Protozool 50(1):65–69
Maitland PS, Herdson D (2009) Key to the marine and fresh-
water fishes of Britain and Ireland. Environment Agency,
Bristol
Mann BQ (1995) Quantification of illicit fish harvesting in the
Lake St Lucia Game Reserve, South Africa. Biol Conserv
74:107–113
Marais JFK (1978) Routine oxygen consumption of Mugil
cephalus,Liza dumerili and L. richardsoni at different
temperatures and salinities. Mar Biol 50:9–16
Marais JFK (1980) Aspects of food intake, food selection, and
alimentary canal morphology of Mugil cephalus (Lin-
naeus, 1958), Liza tricuspidens (Smith, 1935), L. richard-
soni (Smith, 1846), and L. dumerili (Steindachner, 1869).
J Exp Mar Biol Ecol 44:193–209
Marais JFK (1981) Seasonal abundance, distribution and catch
per unit effort using gillnets, of fishes in the Sundays
estuary. S Afr J Zool 16:144–150
Marais JFK (1983a) Fish abundance and distribution in the
Gamtoos estuary with notes on the effects of floods. S Afr J
Zool 18:103–109
Marais JFK (1983b) Seasonal abundance, distribution and catch
per unit effort of fishes in the Krom estuary, South Africa.
S Afr J Zool 18:96–102
Marais JFK, Erasmus T (1977a) Chemical composition of ali-
mentary canal contents of mullet (Teleostei: Mugilidae)
caught in the Swartkops estuary near Port Elizabeth, South
Africa. Aquaculture 10:263–273
Marais JFK, Erasmus T (1977b) Body composition of Mugil
cephalus,Liza dumerili,Liza richardsoni and Liza tricu-
spidens (Teleostei: Mugilidae) caught in the Swartkops
estuary. Aquaculture 10:75–86
Mariani A, Panella S, Monaco G, Cataudella S (1987) Size
analysis of inorganic particles in the alimentary tracts of
Mediterranean mullet species suitable for aquaculture.
Aquaculture 62(2):123–129
Martuccia O, Pietrellib L, Consiglioc C (1993) Fish otoliths as
indicators of the cormorant Phalacrocorax carbo diet
(Aves, Pelecaniformes). B Zool 60(4):393–396
Maruya KA, Francendese L, Manning RO (2005) Residues of
toxaphene decrease in estuarine fish after removal of con-
taminated sediments. Estuaries 28(5):786–793
Masson H, MaraisJFK (1975) Stomach content analyses of mullet
from the Swartkops Estuary. Zool Afr 10(2):193–207
Mbande S, Froneman W, Whitfield A (2004) The primary car-
bon sources utilised by fishes in the Mngazi and Mngazana
estuaries, South Africa: a preliminary assessment. Afr J
Aquat Sci 29(2):195–204
McDonough CJ, Wenner CA (2003) Growth, recruitment, and
abundance of juvenile striped mullet (Mugil cephalus)in
South Carolina estuaries. Fish B-NOAA 101:343–357
McDonough CJ, Roumillat WA, Wenner CA (2003) Fecundity
and spawning season of striped mullet (Mugil cephalus L.)
in South Carolina estuaries. Fish B-NOAA 101:822–834
McDonough CJ, Roumillat WA, Wenner CA (2005) Sexual
differentiation and gonad development in striped mullet
(Mugil cephalus L.) from South Carolina estuaries. Fish
B-NOAA 103:601–619
Menezes NA, De Oliveira C, Nirchio M (2010) An old taxo-
nomic dilemma: The identity of the western south Atlantic
lebranche mullet (Teleostei: Perciformes: Mugilidae).
Zootaxa 2519:59–68
Merella P, Garippa G (2001) Metazoan parasites of grey mullets
(Teleostei: Mugilidae) from the Mistras Lagoon (Sardinia,
Western Mediterranean). Sci Mar 65(3):201–206
Meyer-Rochow VB, Cook I, Hendy CH (1992) How to obtain
clues from the otoliths of an adult fish about the aquatic
environment it has been in as a larva. Comp Biochem
Physiol 103A:333–335
Meynecke J-O, Lee SY (2011) Climate coastal fisheries rela-
tionships and their spatial variation in Queensland, Aus-
tralia. Fish Res 110(2):365–376
Meynecke J-O, Lee SY, Duke NC, Warnken J (2006) Effect of
rainfall as a component of climate change on estuarine fish
production in Queensland, Australia. Est Coast Shelf Sci
69(3–4):491–504
Michaelis H (1993) Food items of the grey mullet Mugil ceph-
alus in the Banc d’Arguin area (Mauritania). Hydrobiolo-
gia 258:175–183
Miggiano E, Lyons RE, Li Y, Dierens LM, Crosetti D, Sola L
(2005) Isolation and characterization of microsatellite loci
in the striped mullet, Mugil cephalus. Mol Ecol Notes
5:323–326
Minos G, Katselis G, Ondrias I, Harrison IJ (2002) Use of
melanophore patterns on the ventral side of the head to
identify fry of grey mullets (Teleostei: Mugilidae). Isr J
Aquacult-Bamid 54(1):12–26
Rev Fish Biol Fisheries (2012) 22:641–681 677
123
Author's personal copy

Mohanraj G, Nammalwar P, Kandaswamy S, Sekar AC (1994)
Availability of grey mullet spawners in Adyar Estuary and
Koovalum Backwater around Madras, India. J Mar Biol
Ass India 36:167–180
Moore RH (1970) Changes in the composition of the swim-
bladder gas of the striped mullet, Mugil cephalus, during
hypoxia. Comp Biochem Physiol 34(4):895–899
Moravec F, Libosva
´rsky
´J (1975) Intestinal helminths from grey
mullet, Mugil capito Cuv. and M. cephalus L., of Lake
Borullus, A.R.E. Folia Parasit 22(3):279–281
Moriarty DJW (1976) Quantitative studies on bacteria and algae
in the food of the mullet Mugil cephalus L. and the prawn
Metapenaeus bennettae (Racek and Dall). J Exp Mar Biol
Ecol 22(2):131–143
Morovic D (1963) Contribution a
`la connaissance du de
´but de la
premie
`re maturite
´sexuelle et de la pe
´riode de ponte chez
Mugil cephalus L. et Mugil chelo Cuv. en Adriatique
(Dalmatie). Rapp Comm Int Expl Mer Me
´dit 17:779–786
Munz FW (1958) The photosensitive retinal pigments of fishes
from relatively turbid coastal waters. J Gen Physiol
42:4455–4459
Mwandya AW, Mgaya YD, O
¨hman MC, Bryceson I, Gullstro
¨m
M (2010) Distribution patterns of striped mullet Mugil
cephalus in mangrove creeks, Zanzibar, Tanzania. Afr J
Mar Sci 32(1):85–93
Nanami A (2007) Juvenile swimming performance of three fish
species on an exposed sandy beach in Japan. J Exp Mar
Biol Ecol 348:1–10
Narasimhamurti CC, Kalavati C (1979) Kudoa tetraspora n.sp.
(Myxosporidae: Protozoa) parasitic in the brain tissue of
Mugil cephalus. Proc. Anim Sci 88(1):85–89
Nash CE, Koningsberger RS (1981) Artificial propagation. In:
Oren OH (ed) Aquaculture of grey mullets. International
Biological Programme 26, Cambridge University Press,
Cambridge
Nash CE, Kuo C-M, McConnel SC (1974) Operational proce-
dures for rearing larvae of the grey mullet (Mugil cephalus
L.). Aquaculture 3(1):15–24
Nordlie FG (2000) Patterns of reproduction and development of
selected resident teleosts of Florida salt marshes. Hydro-
biologia 434:165–182
Nordlie FG, Leffler CW (1975) Ionic regulation and the ener-
getics of osmoregulation in Mugil cephalus Lin. Comp
Biochem Physiol 51(1A):125–131
Nordlie FG, Szelistowski WA, Nordlie WC (1982) Ontogenesis
of osmotic regulation in the striped mullet, Mugil cephalus
L. J Fish Biol 20:79–86
Odum WE (1968) Mullet grazing on a dinoflagellate bloom.
Chesapeake Sci 9(3):202–204
Odum WE (1970) Utilization of the direct grazing and plant
detritus food chains by the striped mullet, Mugil cephalus.
In: Steele JH (ed) Marine food chains. Oliver and Boyd,
Edinburgh, pp 222–240
Oksyuzyan EB, Sokolovsky AS (2003) The marine component
of ichthyofauna in the mouth area and lower course of the
Tumen River. Russ J Mar Biol 29(1):18–22
Oldewage WH, van As JG (1988) Two new species of Erga-
silidae (Copepoda: Poecilostomatoida) parasitic of Mugil
cephalus L. in southern Africa. Hydrobiologia 162(2):
135–139
Oren OH (1981) Aquaculture of grey mullets. Cambridge Uni-
versity Press, New York
O
¨zer A, O
¨ztu
¨rk T (2004) Trichodina puytoraci Lom, 1962 and
Trichodina lepsii Lom, 1962 (Peritrichida: Ciliophora)
infestations on mugilids caught at the Black Sea coast of
Sinop in Turkey. Turk J Zool 28(2):179–182
Padmini E, Vijaya Geetha B (2009) Impact of season on liver
mitochondrial oxidative stress and the expression of
HSP70 in grey mullets from a contaminated estuary.
Ecotoxicology 18(3):304–311
Panfili J, Aliaume C, Berrebi P, Casellas C, Chang CW, Diouf
PS, Durand J-D, Flores Herna
´ndez D, Garcı
´a de Leo
´nF,
Lale
`ye
`P, Morales-Nin B, Toma
´s J, Tzeng, WN, Vassi-
lopoulou V, Wang CW, Whitfield AK (2006) State of the
art of Mugil research. European Commission 6th Frame-
work Programme, INCO-CT-2006-026180, MUGIL
Deliverable 1, 21 pp
Panfili J, Aliaume C, Flores Herna
´ndez D, Hsu CC, Jacquart A,
Lale
`ye
`P, Morales-Nin B, Ramos Miranda J, Toma
´sJ,
Tzeng WN, Vassilopoulou V, Whitfield AK (2007) Mugil
cephalus life-history trait studies. European Commission
6th Framework Programme, INCO-CT-2006-026180,
MUGIL Deliverable 3, 33 pp
Paperna I (1975) Parasites and diseases of the grey mullet
(Mugilidae) with special reference to the seas of the near
East. Aquaculture 5:3–8
Pastor D, Boix J, Ferna
´ndez V, Albaige
´s J (1996) Bioaccumu-
lation of organochlorinated contaminants in three estuarine
fish species (Mullus barbatus,Mugil cephalus and Dicen-
trarcus labrax). Mar Pollut Bull 32(3):257–262
Payne AI (1976) The relative abundance and feeding habits of
the grey mullet species occurring in an estuary in Sierra
Leone, West Africa. Mar Biol 35(3):277–286
Perera PAB, De Silva SS (1978) Studies on the biology of young
grey mullet (Mugil cephalus) digestion. Mar Biol 44(4):
383–387
Pe
´rez-Rufaza A, Quispe-Becerra JI, Garcı
´a-Charton JA, Marcos
C (2004) Composition, structure and distribution of the
ichthyoplankton in a Mediterranean coastal lagoon. J Fish
Biol 64:202–218
Peterson CH (1976) Cruising speed during migration of the
striped mullet (Mugil cephalus L.): an evolutionary
response to predation. Evolution 32(2):393–396
Pisarevskaya II, Aksenova EO (1991) Feeding of Black Sea
mullets during early ontogeny. J Ichthyol 31(3):22–30
Platell ME, Orr PA, Potter IC (2006) Inter- and intra-specific por-
tioning of food resources by six large and abundant fish spe-
cies in a seasonally open estuary. J Fish Biol 69(1):243–262
Porter CB, Krost P, Gordin H, Angel DL (1996) Preliminary
assessment of grey mullet (Mugil cephalus) as a forager of
organically enriched sediments below marine fish farms.
Isr J Aquacult-Bamid 48(1):47–55
Potter IC, Tiivel D, Valesini FJ, Hyndes GA (1997) Compari-
sons between the ichthyofaunas of a temperate lagoon-like
estuary and the embayment into which that estuary dis-
charges. Int J Salt Lake Res 5(4):337–358
Pradervand P, Beckley LE, Mann BQ, Radebe PV (2003)
Assessment of the line fishery in two urban estuarine sys-
tems in KwaZulu-Natal, South Africa. Afr J Mar Sci
25:111–130
678 Rev Fish Biol Fisheries (2012) 22:641–681
123
Author's personal copy

Radovich J (1961) Relationships of some marine organisms of
the northeast Pacific to water temperatures, particularly
during 1957 through 1959. Fish Bull Calif 112:1–62
Rajendran N, Kathiresan K (2007) Microbes associated with
submerged leaf litter of mangroves. Rev Biol Trop 55:
393–400
Rawson MV (1976) Population biology of parasites of striped
mullet Mugil cephalus L. I. Monogenea. J Fish Biol 9(2):
185–194
Rawson MV (1977) Population biology of parasites of striped
mullet Mugil cephalus L, Crustacea. J Fish Biol 10(5):
441–451
Ray GC (2005) Connectivities of estuarine fishes to the coastal
realm. Est Coast Shelf Sci 64:18–32
Richardson N, Gordon AK, Muller WJ, Whitfield AK (2011) A
weight-of-evidence approach to determine estuarine fish
health using indicators from multiple levels of biological
organization. Aquat Conserv 21:423–432
Rocha-Olivares A, Garber NM, Stuck KC (2000) High genetic
diversity, large inter-oceanic divergence and historical
demography of the striped mullet. J Fish Biol 57:
1134–1149
Rocha-Olivares A, Garber NM, Garber AF, Stuck KC (2005)
Structure of the mitochondrial control region and flanking
tRNA genes of Mugil cephalus. Hydrobiolo
´gica 15:
139–149
Rodgers LJ, Burke JB (1988) Aetiology of the ‘red spot’ disease
(vibriosis) with special reference to the ectoparasitic
digenean Prototransversotrema steeri (Angel) and the sea
mullet Mugil cephalus (Linnaeus). J Fish Biol 32(5):
655–663
Rossi AR, Crosetti D, Gornung E, Sola L (1996) Cytogenetic
analyses of global populations of Mugil cephalus (striped
mullet) by different staining techniques and fluorescent in
situ hybridization. Heredity 76(1):77–82
Rossi AR, Capula M, Crosetti D, Sola L, Campton DE (1998a)
Allozyme variation in global populations of striped mullet,
Mugil cephalus (Pisces: Mugilidae). Mar Biol 131:203–212
Rossi AR, Capula M, Crosetti D, Campton DE, Sola L (1998b)
Genetic divergence and phylogenetic inferences in five
species of Mugilidae (Pisces: Perciformes). Mar Biol
131:213–218
Rubtsova NY, Balbuena JA, Sarabeev VL, Blasco-Costa I,
Euzet L (2006) Description and morphometrical variability
of a new species of Ligophorus and of Ligophorus chab-
audi (Monogenea: Dactylogyridae) on Mugil cephalus
(Teleostei) from the Mediterranean basin. J Parasitol
92:486–495
Rubtsova NY, Balbuena JA, Sarabeev VL (2007) Three new
species of Ligophorus (Monogenea: Dactylogyridae) on
the gills of Mugil cephalus (Teleostei: Mugilidae) from the
Japan Sea. J Parasitol 93:772–780
Rueda PS (2002) Stomach content analysis of Mugil cephalus
and Mugil curema (Mugiliformes: Mugilidae) with
emphasis on diatoms in the Tamiahua Lagoon, Mexico.
Rev Biol Trop 50(1):245–252
Russell DJ (1991) Fish movements through a fishway on a tidal
barrage in subtropical Queensland. Proc Roy Soc Qld
101:109–118
SARGARPA (2006) Carta Nacional Pesquera. Report 25, Dia-
rio Oficial de la Federacio
´n Pesquera, Me
´xico
Sarig S (1996) The fish culture industry in Israel in 1996. Isr J
Aquacul-Bamid 48(3):158–164
Schramm M (1991) Grillotia perelica (Cestoda: Trypanorhyn-
cha) plerocercoids in mullet (Pisces: Mugilidae) from
estuaries in Transkei, southern Africa. S Afr J Mar Sci
11:169–178
Service SK, Feller RJ, Coull BC, Woods R (1992) Predation
effects of three fish species and a shrimp on the macro-
benthos and meiobenthos in microcosms. Est Coast Shelf
Sci 34(3):277–293
Shamsi S, Gasser RB, Beveridge I (2011) Mutation scanning-
coupled sequencing of nuclear ribosomal DNA spacers as a
tool for the specific identification of different Contracae-
cum (Nematoda: Aniskidae) larval types. Mol Cell Probe
25(1):13–18
Shehadeh ZH, Ellis JN (1970) Induced spawning of the striped
mullet, Mugil cephalus L. J Fish Biol 2:355–360
Shehadeh ZH, Kio CM, Millisen KK (1973) Induced spawning
of the striped mullet Mugil cephalus with fractionated
salmon pituitary extract. J Fish Biol 5:471–478
Shen K-N, Jamandre BW, Hsu C–C, Tzeng W-N, Durand J-D
(2011) Plio-Pleistocene sea level and temperature fluctua-
tions in the northwestern Pacific promoted speciation in the
globally-distributed flathead mullet Mugil cephalus. BMC
Evol Biol 11(1):Article No 83
Shingles A, McKenzie DJ, Claireaux G, Domenici P (2005)
Reflex cardiovascular responses to hypoxia in the flathead
gray mullet (Mugil cephalus) and their behavioural mod-
ulation by perceived threat of predation and water turbid-
ity. Physiol Biochem Zool 78(5):744–755
Shyu CZ, Lee TJ (1986) Hydrographic studies on the area of the
Taiwan Strait in winter using satellite IR imagery and its
relationship to the formation of grey mullet fishing
grounds. J Fish Soc Taiwan 12(2):98–122
Silva EIL, De Silva SS (1981) Aspects of the biology of grey
mullet, Mugil cephalus L., adult populations of a coastal
lagoon in Sri Lanka. J Fish Biol 19:1–10
Silva SSD, Perera PAB (1976) Studies on the young grey mullet,
Mugil cephalus L. 1. Effects of salinity on food intake,
growth and food conversion. Aquaculture 7:327–338
Simmons EG (1957) An ecological survey of the Upper Laguna
Madre of Texas. Publ Inst Mar Sci Univ Tex 4:156–200
Smit NJ, Eiras JC, Ranzani-Paiva MJT, Davies AJ (2002) A
Desseria sp. from flathead mullet in South Africa. J Mar
Biol Assoc UK 82(4):675–676
Smith KA, Deguara K (2003) Formation and annual periodicity
of opaque zones in sagittal otoliths of Mugil cephalus
(Pisces: Mugilidae). Mar Freshw Res 54:57–67
Smith MM, Heemstra PC (1986) Smiths’ sea fishes. Macmillan
South Africa, Johannesburg
Steidinger KA, Landberg JH, Truby EW, Roberts BS (1998)
First report of Gymnodium pulchellum (Dinophyceae) in
North America and associated fish kills in the Indian River,
Florida. J Phycol 34(3):431–437
Stenger AH (1959) A study of the structure and development of
certain reproductive tissues of Mugil cephalus Linnaeus.
Zoologica 44(2):53–70
Stickney PR, Shumway SE (1974) Occurrence of cellulose
activity in the stomachs of fishes. J Fish Biol 6:779–790
Strydom NA (2008) Utilization of shallow subtidal bays asso-
ciated with warm temperate rocky shores by the late stage
Rev Fish Biol Fisheries (2012) 22:641–681 679
123
Author's personal copy

larvae of some inshore fish species, South Africa. Afr Zool
43(20):256–269
Strydom NA, d’Hotman BD (2005) Estuary-dependence of
larval fishes in a non-estuary associated South African surf
zone: evidence for continuity of surf assemblages. Est
Coast Shelf Sci 63:101–108
Strydom NA, Wooldridge TH (2005) Diel and tidal variations in
larval fish exchange in the mouth region of the Gamtoos
Estuary, South Africa. Afr J Aquat Sci 30(2):131–140
Sultana R, Rao DP (1998) Bioaccumulation patterns of zinc,
copper, lead, and cadmium in grey mullet, Mugil cephalus
(L.), from harbour waters of Visakhapatnam, India. Bull
Environ Contam Toxicol 60(6):949–955
Sun P, Shi Z-H., Yin F, Peng S-M (2012) Genetic variation
analysis of Mugil cephalus in China Sea based on mito-
chondrial COI gene sequences. Biochem Genet 50:
181–190
Suzuki K (1965) Biology of the striped mullet, Mugil cephalus
L. I. Food contents of young. Prefectural University of Mie,
Faculty of Fisheries Report vol 5, pp 296–305
Switzer TS, Winner BL, Dunham NM, Whittington JA, Thomas
M (2006) Influence of sequential hurricanes on nekton
communities in a southeast Florida estuary: Short-term
effects in the context of historical variations in freshwater
flow. Est Coast 29(6A):1011–1018
Sylvester JR (1975) Critical thermal maxima of three species of
Hawaiian estuarine fish: a comparative study. J Fish Biol
7(2):257–262
Sylvester JR, Nash CE (1975) Thermal tolerance of eggs and
larvae of Hawaiian striped mullet, Mugil cephalus L. Trans
Am Fish Soc 104:104–147
Sylvester JR, Nash CE, Emberson CR (1975) Salinity and
oxygen tolerances of eggs and larvae of Hawaiian striped
mullet, Mugil cephalus L. J Fish Biol 7:621–629
Tamaru CS, Lee CS, Kelley CD, Miyamoto G, Moriwake A
(1994) Oocyte growth in the striped mullet Mugil cephalus
L. maturing at different salinities. J World Aquacult Soc
25:109–114
Tamaru CS, Tamaru CC-T, FitzGerald WJ, Ako H, Sato VT
(2005) Advances in the culture of striped mullet. Am Fish
Soc Symp 46:439–456
Tang YA (1964) Induced spawning of striped mullet by hor-
mone injection. Jpn J Ichthyol 12:23–28
Tang YA (1975) Collection, handling distribution of grey mullet
fingerlings in Taiwan. Aquaculture 5:81–84
Thomson JM (1951) Growth and habits of the sea mullet Mugil
dobula (Gunther), in West Australia. Aust J Mar Freshw
Res 2:193–225
Thomson JM (1953) Status of the fishery for sea mullet (Mugil
cephalus Linnaeus) in eastern Australia. Aust J Mar
Freshw Res 4:41–81
Thomson JM (1954) The organs of feeding and the food of some
Australian mullet. Aust J Mar Freshw Res 5:469–485
Thomson JM (1955) The movements and migrations of mullet
(Mugil cephalus L.). Aust J Mar Freshw Res 6:328–347
Thomson JM (1957) The size at maturity and spawning times of
some Western Australian estuarine fishes. Fish Bull West
Aust 8:1–8
Thomson JM (1959) Some aspects of the ecology of Lake
Macquarie, N.S.W., with regard to an alleged depletion of
fish. X. The movements of fish. Aust J Mar Freshwat Res
10:375–387
Thomson JM (1963) Synopsis of biological data on the grey
mullet, Mugil cephalus Linnaeus, 1758. CSIRO Aust Fish
Oceanogr Fish Synop 1:1–66
Thomson JM (1966) The grey mullets. Oceanogr Mar Biol Ann
Rev 4:301–355
Thomson JM (1997) The Mugilidae of the world. Mem Qld Mus
41(3):457–562
Tooth T (1946) When the pelicans come to St Lucia. S Afr
Angler 1(1):9
Torras X, Cardona L, Gisbert E (2000) Cascading effects of the
flathead grey mullet Mugil cephalus in freshwater eutro-
phic microcosmos. Hydrobiologia 429:49–57
Torricelli P, Tongiorgi P, Almansi P (1981) Migration of grey
mullet fry into the Arno River: Seasonal appearance, daily
activity, and feeding rhythms. Fish Res 1:219–234
Trape S (2009) Les Mugilidae des co
ˆtes ouest-africaines:
Identification mole
´culaire, phyloge
´nie et bio-e
´cologie du
recrutement des juve
´niles dans un estuaire hypersale
´. PhD
dissertation, University of Montpellier, France
Trape S, Durand J-D, Guilhaumon F, Vigliola L, Panfili J (2009)
Recruitment patterns of young-of-the-year mugilid fishes
in a West African estuary impacted by climate change. Est
Coast Shelf Sci 85(3):357–367
Trewavas E, Ingham SE (1972) A key to the species of Mugil-
idae (Pisces) in north-eastern Atlantic and Mediterranean,
with explanatory notes. J Zool Lond 167:15–29
Tsangaris C, Vergolyas M, Fountoulaki E, Nizheradze K (2011)
Oxidative stress and genotoxicity biomarker responses in
grey mullet (Mugil cephalus) from a polluted environment
in Saronikos Gulf, Greece. Arch Environ Con Tox
61(3):482–490
Tung IH (1959) Age determination of the grey mullet. China
Fish Mon 80:2–10
Tung IH (1981) On the fishery biology of grey mullet, Mugil
cephalus L., in Taiwan. Institute of Fisheries Biology
Report, Ministry of Economic Affairs, National Taiwan
University vol 3, pp 38–102
Turesson H, Domenici P (2007) Escape latency is size inde-
pendent in grey mullet. J Fish Biol 71(1):253–259
Tu
¨rkmen A, Tu
¨rkmen M, Tepe Y, Mazlum Y, Oymael S (2006)
Metal concentrations in blue crab (Callinectes sapidus) and
mullet (Mugil cephalus) in Iskenderun Bay, northern east
Mediterranean, Turkey. Bull Environ Con Tox 77:186–193
Umur S¸ , Pekmezci GZ, Beyhan YE, Gu
¨rler AT, Ac¸ici M (2010)
First record of Myxobolus muelleri (Myxosporea: Myxo-
bolidae) in the flathead grey mullet Mugil cephalus (Te-
leostei: Mugilidae) from Turkey. Ankara Universitesi
Veteriner Fakultesi Dergisi 57(3):205–207
Vagner M, Lefranc¸ois C, Ferrari RS, Satta A, Domenici P
(2008) The effect of acute hypoxia on swimming stamina at
optimal swimming speed in flathead grey mullet Mugil
cephalus. Mar Biol 155:183–190
Vall M (2004) Etude de la dynamique des systemes d’exploi-
tations et de l’eco biologie de la reproduction, de trois
Mugilide
´s: Mugil cephalus (Linnaeus, 1758), Liza aurata
(Perugia, 1892) et Mugil capurrii (Risso, 1810), analyse de
leurs strategies d’occupations des secteurs littoraux Mau-
ritaniens et de leurs possibilites d’amenagement. PhD
680 Rev Fish Biol Fisheries (2012) 22:641–681
123
Author's personal copy

dissertation, Nice Sophia Antipolis University, France,
133 pp
Valles-Rı
´os ME, Ruiz-Campos G, Galavı
´z-Silva L (2000)
Prevalencia e intensidad parasitaria en Mugil cephalus
(Pises: Mugilidae), del Rı
´o Colorado, Baja California,
Me
´xico. Rev Biol Trop 48:495–501
van der Elst RP, Wallace JH (1976) Identification of the juvenile
mullet of the east coast of South Africa. J Fish Biol
9:371–374
van der Horst G, Erasmus T (1981) Spawning time and
spawning grounds of mullet with special reference to Liza
dumerili (Steindachner, 1869). S Afr J Sci 77:73–78
Van der Kooy SJ, Guindon-Tisdel K (2003) A practical hand-
book for determining the ages of Gulf of Mexico fishes.
Gulf States Marine Fisheries Commission, Ocean Springs,
USA
Virgona JL (1992) Environmental factors influencing the
prevalence of a cutaneous ulcerative disease (red spot) in
the sea mullet, Mugil cephalus L., in the Clarence River,
New South Wales, Australia. J Fish Dis 15(5):363–378
Wallace JH (1975a) The estuarine fishes of the east coast of
South Africa. I. Species composition and length distribu-
tion in the estuarine and marine environments. II. Seasonal
abundance and migrations. Invest Rep Oceanogr Res Inst
40:1–72
Wallace JH (1975b) The estuarine fishes of the east coast of
South Africa. III. Reproduction. Invest Rep Oceanogr Res
Inst 41:1–51
Wallace JH, van der Elst RP (1975) The estuarine fishes of the
east coast of South Africa. IV. Occurrence of juveniles in
estuaries. V. Ecology, estuarine dependence and status.
Invest Rep Oceanogr Res Inst 42:1–63
Walsh WA, Swanson C, Lee CS, Banno JE, Eda H (1989)
Oxygen consumption by eggs and larvae of striped mullet,
Mugil cephalus, in relation to development, salinity and
temperature. J Fish Biol 35(3):347–358
Walsh WA, Swanson C, Lee CS (1991) Combined effects of
temperature and salinity on embryonic development and
hatching of striped mullet, Mugil cephalus. Aquaculture
97:281–289
Wang C-H, Hsu C–C, Chang C-W, You C-F, Tzeng W-N (2010)
The migratory environmental history of freshwater resident
flathead mullet Mugil cephalus L. in the Tanshui River,
northern Taiwan. Zool Stud 49(4):504–514
Wang C-H, Hsu C–C, Tzeng W-N, You C-F, Chang C-W (2011)
Origin of the mass mortality of the flathead grey mullet
(Mugil cephalus) in the Tanshui River, northern Taiwan, as
indicated by otolith elemental signatures. Mar Pollut Bull
62(8):1809–1813
Whitehead PJP, Bauchot M-L, Hureau J-C, Nielson J, Tortonese
E (1986) Fishes of the north-eastern Atlantic and Medi-
terranean, vol III. UNESCO, Paris
Whitfield AK (1985) The role of zooplankton in the feeding
ecology of fish fry from some southern African estuaries.
S Afr J Zool 20:166–171
Whitfield AK (1989) Ichthyoplankton in a southern African surf
zone: nursery area for the postlarvae of estuarine associated
fish species? Est Coast Shelf Sci 29:533–547
Whitfield AK (1990) Life history styles of fishes in South
African estuaries. Environ Biol Fish 28:295–308
Whitfield AK, Blaber SJM (1978a) Distribution, movements
and fecundity of Mugilidae at Lake St Lucia. Lammergeyer
26:53–63
Whitfield AK, Blaber SJM (1978b) Feeding ecology of pisciv-
orous birds at Lake St Lucia. Part 1: Diving birds. Ostrich
49:185–197
Whitfield AK, Blaber SJM (1978c) Resource segregation
among iliophagous fish in Lake St Lucia, Zululand. Envi-
ron Biol Fish 3:293–296
Whitfield AK, Blaber SJM (1979a) Feeding ecology of pisciv-
orous birds at Lake St Lucia. Part 3: Swimming birds.
Ostrich 50:10–20
Whitfield AK, Blaber SJM (1979b) Predation on striped mullet
(Mugil cephalus)byCrocodylus niloticus at St. Lucia,
South Africa. Copeia 2:266–269
Whitfield AK, Kok HM (1992) Recruitment of juvenile marine
fishes into permanently open and seasonally open estuarine
systems on the southern coast of South Africa. Ichthy Bull
JLB Smith Inst Ichthyol 57:1–39
Whitfield AK, Taylor R (2009) A review of the importance of
freshwater inflow to the future conservation of Lake St
Lucia. Aquat Conserv 19:838–848
Whitfield AK, Blaber SJM, Cyrus DP (1981) Salinity ranges of
some southern African fish species occurring in estuaries.
S Afr J Zool 16:151–155
Whitfield AK, Taylor RH, Fox C, Cyrus DP (2006) Fishes and
salinities in the St Lucia system—a review. Rev Fish Biol
Fish 16:1–20
Yashouv A (1969) Preliminary report on induced spawning of
Mugil cephalus reared in captivity in freshwater ponds. Isr
J Aquacult-Bamid 21:19–24
Yilmaz AB (2003) Levels of heavy metals (Fe, Cu, Ni, Cr, Pb,
and Zn) in tissue of Mugil cephalus and Trachurus medi-
terraneus from Iskenderun Bay, Turkey. Environ Res
92:277–281
Young GC, Potter IC (2002) Influence of exceptionally high
salinities, marked variations in freshwater discharge and
opening of estuary mouth on the characteristics of the
ichthyofauna of a normally-closed estuary. Est Coast Shelf
Sci 55(2):223–246
Young GC, Potter IC, Hyndes GA, De Lestang S (1997) The
ichthyofauna of an intermittently open estuary: Implica-
tions of bar breaching and low salinities on faunal com-
position. Est Coast Shelf Sci 45(1):53–68
Zismann L, Berdugo V, Kimor B (1975) The food and feeding
habits of early stages of grey mullet in the Haifa Bay
region. Aquaculture 6:59–75
Rev Fish Biol Fisheries (2012) 22:641–681 681
123
Author's personal copy