||Mugil cephalus Linnaeus, 1758
Other Taxonomic Groupings
Division Teleostei in the Class Actinopterygii
replaces the former class Osteichthyes as the newer taxonomic grouping for the
ray-finned fishes (Compagno 1991, Nelson 1994, Bond 1996). The teleost fishes
are further divided into subdivisions, with the mullets ordered as follows:
Subdivison Euteleostei, Superorder Acanthopterygii, Series Mugilomorpha, Order
The striped mullet, Mugil cephalus, can attain 18" in length and
reach approximately 3 pounds. Body shape is cylindrical anteriorally, becoming
somewhat laterally compressed toward the posterior. Adult coloration is bluish-gray or
greenish above, becoming silver along the sides of the body, and white on the
ventral surface. There are 6-7 black horizontal bars along the sides of the
body, and no obvious lateral line. The pectoral fins are placed high on the
shoulders, and the pelvic fins are abdominal. M. cephalus has a blunt
snout, and a small, somewhat upturned mouth.
Nine species of mullet occur in the west
central Atlantic ocean (Ditty and Shaw 1996). In Florida, M. cephalus is
the most common of the mullet species, but also occurs with M. curema,
the white mullet, and M. gyrans, the fantail mullet. Differences in fin
rays and fin morphology help separate species. There is an apparent seasonality
of larvae, which also assists in separating larvae of M. cephalus from
other species, especially M. curema. Larvae of M. cephalus are
most abundant in the northern Gulf of Mexico from November through December,
while those of M. curema are most abundant from April through May.
HABITAT AND DISTRIBUTION
Mugil cephalus occurs worldwide from approximately 42°N to
42°S Latitude (Bok 1979, Render et al. 1995), where it inhabits estuarine
intertidal, freshwater and coastal marine habitats. In the western Atlantic
Ocean, M. cephalus ranges from Cape Cod to Brazil, including the Gulf of
Mexico, Caribbean, and West Indies (Amos and Amos 1997).
Mugil cephalus occurs lagoon-wide, with juvenile
fishes most common in impounded areas, around mangroves, in seagrass beds, and
offshore throughout the late fall and winter.
LIFE HISTORY AND POPULATION BIOLOGY
Age, Size, Lifespan
Mugil cephalus attains an adult size of 46 cm (18 inches). In the
first year, it grows to 17.8 - 22.2 cm (7 - 8.5 inches), and weighs 64 - 119 g
(2.3 - 4.2 oz.).
Prejuvenile mullet from 0.17 - 0.35 cm (0.07 -
0.14 inches) standard length (SL) are a distinct silvery color, with evident
countershading (i.e., they are generally darker on the dorsal surface than on
the ventral surface). Prejuveniles and small juveniles form loose schools of tens to
hundreds of individuals that occupy shallow, warm, near-shore water where they feed intensely and continuously. Prejuveniles undergo metamorphosis
to the juvenile stage before they reach 0.50 cm (0.2 inches). The most striking
change seen after metamorphosis is the loss of silver body color , especially
along the dorsal side. Countershading is still evident, however. Other
metamorphic changes in juvenile mullet include elongation and convolution of the intestine,
development of adipose eyelids, transformation of some soft anal fin rays into
spines, and changes in the morphology of the teeth and lips (Major
1978). The ontogenetic shift in the diet of M. cephalus from feeding
primarily on copepods and other small zooplankton, to feeding on detritus and
algae is coincident with metamorphic changes in the intestine, teeth and lips of
the fish as it becomes a juvenile (Major 1978).
M. cephalus is the most abundant of the mullet species
throughout much of its range, especially in fresh water and estuarine areas.
Rulifson (1977), in tests to assess maximum burst swimming
speed of mullet, found that most juveniles between 2.5-6.5 cm (1 - 2.5
inches) SL could sustain maximum swimming speeds of at least 12.7 body lengths
(L) per second, for 30 seconds. These findings led Rulifson to suggest that M.
cephalus juveniles could reach a maximum burst speed of over 20 L/s for
approximately 2 seconds.
Female mullet reach sexual maturity in their fourth year, when they are
between 40 - 42 cm (15.8 - 16.5 inches). Males mature in their third year, once
they reach a size of 33 - 38 cm (13 - 15 inches). The minimum spawning size of
females is between 31 - 34 cm (12.2 - 13.4 inches) (Apekin and Vilenskaya
1979). The general reproductive pattern of Mugil cephalus involves
migration from either fresh or estuarine waters to offshore waters where they
spawn in large schools. Larvae and prejuveniles then migrate to inshore
estuaries where they inhabit shallow, warm water in the intertidal zone.
Beginning in the early fall, large schools of
mullet aggregate in the lower reaches of estuaries and at river mouths in
preparation for offshore migration to spawning grounds. Environmental cues
such as falling water temperatures, passage of cold fronts and falling
barometric pressure are thought to trigger aggregation and subsequent migration
(Mahmoudi 2000). Spawning occurs in deep, offshore waters from mid-October
through late January, with peak spawning occurring in November and December
(Ditty and Shaw 1996).
are isochronal spawners, with all oocytes reaching maturity at the same time.
However, based on the size of the female body cavity, it is unlikely that a
female's entire store of eggs is hydrated at the same time in preparation for
spawning. Rather, females are likely to hydrate eggs in batches (Thompson 1958,
Render et al. 1995) and spawn on successive evenings until their supply of
yolked eggs is depleted. Female fecundity ranges from 270,000 - 1.6
million eggs per individual per season (Render et al. 1995); absolute fecundity
is between 2.9 - 16 million eggs (Apekin and Vilenskaya 1979).
M. cephalus has a generally well defined and short recruitment period throughout its range.
In South Africa, Bok (1979) found that recruitment of 0.15 - 0.40 cm (0.06 -
0.16 inches) fork length (FL) fry takes place from July to October, with fewer
numbers in May, June and November. Chubb et al. (1981), studying mullet in
Australia, found that, based on the first appearance of small juveniles between
0.20 - 0.30 cm (0.08 - 0.12 inches), spawning in Australia occurs between March
and September. In Hawaii, the reproductive season of the striped mullet is
between September and March (Kelly 1990). In Florida, M. cephalus spawns
offshore from October through mid-January, with spawning completed by late
February (Render et al. 1995, Ditty and Shaw 1996).
Larvae become abundant in the waters of the
northern Gulf of Mexico between November and December (Ditty and Shaw 1996) in
water temperatures between 23-25°C. Tag returns along the U.S.
Gulf of Mexico coast indicate that M. cephalus do not make extensive migrations
in this region, but instead remain in a relatively small area and return to
their original bay system after spawning (Funicelli 1989, Mahmoudi et al. 1989,
Ditty and Shaw 1996).
The eggs of Mugil cephalus contain a single oil globule. Egg size
varies according to location, and with water temperature. Apekin and Vilenskaya
(1979) measured oocyte diameter in Black Sea M. cephalus as between 425 -
632 um. In Hawaii, Shehadeh et al. (1973) measured oocytes between 650-700 um.,
with mature eggs reaching up to 930 um. Kou et al (1974), in a later study,
determined the egg size of Hawaiian mullet to range from 0.621 mm - 1.09 mm.
Eggs are shed and fertilized in the water column, and
hatch within 48 hours (Render et al. 1995). Newly hatched larvae of M. cephalus
measure approximately 2.2 - 2.6 mm (0.87 - 1.0 inch) (Bensam 1987; Eda et al.
1990). Larval pigmentation consists of thick, stellate chromatophores covering
the body, except in the posterior region. Additionally, larvae and early
postlarvae of M. cephalus possess a midlateral row of stellate
melanophores. This pigmentation pattern helps distinguish M. cephalus
larvae from those of other mullet genera (Bensam 1987).
The mouths of larval mullet are open by the
second day of post hatch, with the yolksac fully absorbed by the fifth day.
Active feeding begins prior to full absorption of the yolksac, as early as 70
hours post hatching, with young larvae beginning to take rotifers and microalgae
as food. Adverse effects from withholding food become evident as early as 3.5
days after hatching. Larval mullet die within 192 hours (8 days) if not fed (Eda
et al. 1990).
Embryos of M. cephalus develop optimally at temperatures of
approximately 21°C. Greatest hatching success was achieved at 22 - 25°C
(Sylvester et al. 1974, 1975). Growth of embryos was retarded at temperatures above 26°C (Kou et al.1974). Critical thermal maxima (CTM) for juvenile mullet ranges
from 30° C to 42.5° C depending on acclimation temperature.
Acclimation to higher temperatures appears to
be a selective advantage to prejuvenile and juvenile mullet, which routinely
choose shallow, warm waters in the intertidal zone. In studies of M. cephalus
in the Pacific Ocean, Major (1978) and Chubb et al. (1981) found that
prejuvenile mullet approximately 20 mm SL leave the open ocean to enter
estuaries where they select shallow waters with extensive diel fluctuations in
both temperature and salinity. Prejuvenile fish < 50 mm (1.97 in.) SL
were found in shallow pools with high, often near lethal temperatures of between
34 - 42.5° C, and salinities ranging from 2 - 30 ppt. (Major 1978). This
choice of marginal habitats, in conjunction with the schooling behaviors observed
in prejuveniles and juveniles, is thought to be beneficial as a predation refuge
for small fish. As mullet grow larger and their predation risk is lessened, they
are then able to move away from the high intertidal zone into open waters where
temperature and salinity are more stable. When mullet complete their metamorphosis to the juvenile
stage (at approximately 50 mm SL), they
begin moving into somewhat deeper areas of the intertidal zone. In a laboratory
experiment, Major (1978) showed that as body size increased, the temperature
range within the chosen habitat tends to decrease. Older juveniles (> 50 mm
SL) maintained themselves seaward of the tide line in waters of lower
temperature and more uniform salinity (Major 1978). These observations have lead
some authors to speculate that the biochemical and hormonal changes that result
from metamorphosis result in the preference of older juveniles and adults for
Kulikova, Shekk and Rudenko (1986) studied M.
cephalus juveniles under conditions of cold stress. Findings from this study
showed that juvenile mullet are active at temperatures above 10 - 12°C. With
decreasing temperature, they begin to sink toward the bottom where they form
dense schools. At temperatures below 5°C, juveniles respond sluggishly to
food, and will eventually cease feeding. At 2 - 3°C, they remain nearly
motionless at the bottom. During the first week under these conditions, red
hemorrhages appear on the snout and fins. In the next week, hemorrhages appear
under the skin, and on the internal organs, followed by tissue necrosis, and eventually, death.
The Kulikova, Shekk and Rudenko (1986) study
also found that short, cold periods have little long-term effect on mullet.
Older mullet were found to overwinter in deep bays where temperatures sometimes
reach as low as 7°C. In the laboratory, young mullet were able to endure
brief cooling to temperatures of 1 - 1.5°C for up to 1 day. There were no
adverse effects on mullet when they were gradually warmed to their original
acclimation temperature. Cold stress did occur, however, if fish were repeatedly
subjected to alternate cold/warm cycles.
Changes in red blood cells also occur in
thermally stressed mullet. Low temperatures decrease the functionality of blood
hemoglobin and lead to asphyxia. Additionally, lipids in the body are not
catabolized (Kulikova, Shekk and Rudenko 1986) to maintain bodily function.
Rather, mullet that are cold stressed increase the catabolism of bodily
proteins. This results in the alteration of blood chemistry with respect to increased
ammonia nitrogen excretion.
Adult M. cephalus are highly euryhaline, and survive in a range of
salinities from 0 ppt. in fresh water, to hypersaline waters with salinity as
high as 90 ppt. (Lee and Menu 1981). However, Hu and Liao (1981) showed that
female M. cephalus are generally unable to ovulate in fresh water.
Before being spawned, the eggs of M.
cephalus are isosmotic to the blood of the parent (Lee and Menu 1981). In
the laboratory, fertilization rates were optimal at 22.5 ppt. and eggs
incubated at temperatures between 22 - 24°C hatched optimally at salinities
between 22 - 23 ppt. (Hu and Liao 1981). Hatching was successful at salinities
from 15 - 42 ppt., but increased to over 50% at salinities above 19 ppt. At
11 ppt., the majority of fertilized eggs died, and those that did develop
further, ultimately died in the gastrula stage. However, Lee and Menu (1981)
showed that naturally spawned eggs developed to the embryonic stage at
salinities between 5 - 60 ppt. Hatching in this study occurred at salinities
between 10 - 55 ppt., but no larvae survived at either 10 or 55 ppt. In the Lee
and Menu (1981) study, the optimal salinity range at an incubation temperature
of 22 to 25°C, was between 30 - 40 ppt., peaking at 35 ppt.
In aquaculture trials, young M. cephalus
subjected to various salinites (Murashige et al. 1991) showed short-term
differences in growth. Fish raised in tanks with a salinity of 22-23 ppt. grew
faster than those held at higher salinity. However, in the long term, there was
no significant difference in growth rates among the groups. This finding led the
authors to suggest that there is no particular advantage or disadvantage to
maintaining uniform salinity levels for tank-reared mullet.
Other Physical Tolerances
Mullet are particularly susceptible to red tide
organisms. It has been estimated that as many as 16 different pathologies
can be involved in cases of red tide-induced death in mullet, and that
concentrations of 250,00 cells/L of Gymnodidium breve are sufficient to
cause mortaility in mullet (Mahmoudi 2000).
Mugil cephalus is a heterotroph that, as an adult, is primarily a
detritus feeder. Mullet are highly flexible in their food habits, with possible
ontogenetic shifts in the diet as they age and make the transition from
post-larva to adult. The mullet has a gizzard-like pyloric stomach and intestine
that is 3-5 times the length of the body, indicating a principally herbivorous
diet (Service et al. 1992). Many studies of feeding habits in mullet (Suzuki
1965, Odum 1968 and 1970, Zismann et al. 1975, Bishop and Miglarese 1978) found
that juvenile M. cephalus (< 30 mm) are primarily carnivorous.
However, De Silva and Wijeyarante (1977) found that young mullet (20-55 mm)
feed primarily on diatoms (55.5%), followed by green algae (22.3%), Xanthophycea
(15.5%), Cyanobacteria (6.1%) and animal matter (principally foraminiferans and
copepods, 0.6 %).
In one study, detritus and sand first began to appear in
mullet over 25 mm in length (0.9 in.), and increased in percent occurrence as
body length increased (De Silva and Wijeyarante 1977). This finding led the
authors to conclude that 25 mm may represent a transitional size in mullet where
they gradually begin to alter their trophic mode from being primarily planktonic
or carnivorous feeders to being primarily benthic feeders. The findings of De
Silva and Wijeyarante (1977) contradicted other studies in that these authors
found a diurnal pattern of feeding in mullet, with peaks of activity occurring
around dawn and midday, regardless of the state of the tide. Others (Odum 1970)
found that M. cephalus feeds almost continuously throughout the day, and
their feeding intensity varies with tidal state.
Bishop and Miglarese (1978) found that the
principal food sources of adult mullet are detritus and epiphytic algae.
However, these authors also observed Mugil cephalus feeding
opportunistically on swarming polychaetes of the Nereis genus. This observation
lead the authors to suggest that since M. cephalus lack the mouthparts
for tearing and cutting, predation must be limited to bite-sized prey, or to
prey items which break apart easily. Odum (1970) made a similar observation
about the opportunistic nature of feeding in mullet, stating that mullet will
select food with higher caloric value whenever presented with the opportunity.
Mullet also actively ingest "marine
snow," a composite material which consists of detritus, mineral grains,
phytoplankton, microorganisms, and small nematode worms all bound together in a
mucous matrix. Particles range in size from 0.5 mm to several cm in size. Marine
snow is nearly always present in coastal and estuarine environments (Larson and
Shanks 1996), and is a valuable food source to detritivores such as M.
Juvenile mullet > 50 mm, as well as adults, both appear to compete with
prejuveniles in estuarine regions, with some evidence suggesting that limited habitat partitioning occurs. Major (1978) observed larger mullet moving into an
intertidal estuarine region during high tide to feed on the same food resources
used by smaller mullet during low tide. As the larger mullet moved in on the
high tide, the smaller fishes moved closer to the shoreline in the high
Lizardfish, needlefish, crabs, etc. prey on juvenile M. cephalus. Larger
mullet are subject to larger predators such as snook, snappers, barracuda,
dolphins, etc. In the presence of large predators, prejuvenile, juvenile and
adult mullet organize into tightly formed schools and tend to cease feeding
activity. In the absence of predators, schools become more loosely organized,
and individuals will feed constantly during low tides (Major 1978).
Juveniles occupy the high intertidal zone of
estuaries where water temperatures and salinity fluctuate greatly. Older mullet
inhabit deeper, more stable waters.
Benefit in the IRL
M. cephalus is an important commercial and recreational
fishery species in the Indian River Lagoon. Adult mullet are line caught as food fish
and for roe, while juveniles are commonly used as bait for larger sportfish.
In the Western Atlantic, M. cephalus is
commercially valuable on the east coast of Florida and in the Gulf of Mexico,
especially in western Florida and Louisiana, where historically a large portion of the
commercial catch consists of mullet (Render et al. 1995). Florida's fishery for
mullet is primarily centered around fresh and smoked mullet, as well as roe
harvesting for shipment to Asian markets (Render et al. 1995, Ditty and Shaw
Mugil cephalus is one of the most important animal protein sources
for people in the Pacific Basin, Southeast Asia, India, the Mediterranean,
Eastern Europe, Central America and South America (Nash 1978). It has gained
popularity as a widely cultured food fish throughout Europe and Asia (Lee and
The striped mullet is a high value fishery species within Florida. The statewide commercial catch of
Mugil cephalus between the years 1987 - 2001 was 232.9 million pounds, with a
dollar value of over $115.2 million. Over the same time period within the 5 county area
encompassing the IRL (Volusia, Brevard, Indian River, St. Lucie and Martin
Counties) the commercial catch of M. cephalus
accounts for approximately 10% of the statewide total, with a harvest of 23.9
million pounds, and a value in excess of $11 million. This
ranks the striped mullet eleventh in commercial value within the IRL, and sixth in
Figure 1 and Table 1 below show the dollar value of the
fishery to IRL counties by year. As shown, commercial catch ranged from a
low of $456,700 in 1991 to a high of over $1.1 million in 1994.
Volusia County annually accounted for the largest percentage of the catch with
approximately 39% of the total (Figure 2, Table 2), followed by Brevard County, which
accounts for 23% of the total. Indian River, St. Lucie and Martin Counties
account for approximately 14%, 15%, and 10% of the harvest respectively.
Of interest is the change in the mullet harvest
following implementation of the gill-net ban in 1995. With the exception
of 1996, landings of mullet within IRL counties decreased, but have held at a
relatively stable level, with a commercial value of approximately $600,00
annually. This trend is reflective of the mullet harvest throughout
Florida. Mahmoudi (2000) reported that fishing effort was reduced 54% from
1995 - 1999 following the ban on gill-netting.
Figure 1. Annual dollar value of the commercial catch of striped mullet to the 5-county area of the Indian River Lagoon.
Figure 2. Total striped mullet dollar value and percentage by county for the years 1987 - 2001.
Table 1. Total dollar value of the IRL harvest of striped mullet, Mugil cephalus, between 1987 -2001.
||Value to IRL ($)
Table 2. By-county percentage of the striped mullet harvest for the years 1987-2001.
Table 3. By county cumulative dollar value and percentage of total for the IRL striped mullet harvest from 1987 - 2001.
Recreational fishery data for mullet have been collected from survey data within
the 5-county area encompassing the Indian River Lagoon, and were provided by the
National Marine Fisheries Service. Due to the high number of records for
unidentified mullet, data for all members of the genus Mugil have been
The recreational fishery for mullet has grown in Indian
River Lagoon counties since 1997, likely in response to the 1995 ban on
commercial gill netting practices. Reduced fishing pressures on mullet
have allowed stocks to rebound significantly (Mahmoudi 2000). As shown in Figures
3 and 4, the bulk of the recreational harvest of mullet was taken from the
Indian River Lagoon (34.8%) and other inland waters (35.7%). Lesser
numbers of mullet are harvested in nearshore waters less than 3 miles from the
coast (25.8%), and from offshore waters to 200 miles (3.6%).
Based on survey information, an average of 3.4 million
mullet per year are harvested recreationally by anglers in the 5 county area
that encompasses the Indian River Lagoon. Within the Lagoon itself, over
1.18 million mullet are harvested annually (Table 4). The harvest from
inland waters other than the Indian River Lagoon rivals the catch from lagoon
waters at 1.21 million fish per year, while the figures for the nearshore
fishery and the offshore fishery drop to approximately 876,000 and 123,000
Figure 3. Survey data for the recreational fishery of Mugil species, primarily striped and white mullet, showing the number of fishes harvested in East Florida waters from 1997 - 2004.
Figure 4. Summary of the recreational harvest of striped and white mullet and percentage of total by area from 1997 - 2004.
Table 4. Summary data for recreational fishery in Eastern Florida waters for Mugil species, including the striped mullet (M. cephalus) and the white mullet (M. curema) from 1997 - 2004. Data provided by National Marine Fisheries Service, Fisheries Statistics Division, NOAA.
||To 3 Miles
||To 200 Miles
Table 5. By-county annual and cumulative percentages of mullet harvest, including striped and white mullet, for the years 1997 - 2001. Data provided by National Marine Fisheries Service, Fisheries Statistics Division, NOAA.
||To 3 Miles
||To 200 Miles
Table 6. Summary of the recreational fishery for mullet, including striped and white mullet, harvested recreationally and percentage of total fish captured in each area from 1997 - 2004. Data provided by National Marine Fisheries Service, Fisheries Statistics Division, NOAA.
||To 3 Miles
||To 200 Miles
Cost in the IRL
Because much of the fishery for striped mullet targets
gravid females prior to spawning, mullet may be susceptible to overfishing (Ditty and Shaw 1996) as they make the migration from the Indian
River Lagoon to offshore waters.
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Report by: K. Hill,
Smithsonian Marine Station
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