Aquaculture 302 (2010) 195–201

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Aquaculture
j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / a q u a - o n l i n e

Growth rates of cobia (Rachycentron canadum) cultured in open ocean submerged

cages in the Caribbean
Daniel D. Benetti a,⁎, Brian O'Hanlon b, José A. Rivera c, Aaron W. Welch a, Christopher Maxey d, M. Refik Orhun e
a
University of Miami, Rosenstiel School of Marine and Atmospheric Science, Aquaculture Program, 4600 Rickenbacker Causeway, Miami, FL 33149, USA
b
Snapperfarm, Inc., P.O. Box 685, Culebra, 00775 Puerto Rico
c
Under contract to NOAA-Fisheries, HC-02, Box 1736, Boqueron, 00622 Puerto Rico
d
The Island School, Cape Eleuthera Institute, P.O. Box 6008, Lawrenceville, NJ 08648, USA
e
SEFSC–NOAA Fisheries Service, 75 Virginia Beach Dr., Miami, FL 33149, USA

a r t i c l e i n f o a b s t r a c t

Article history: Growth rates of hatchery-reared cobia Rachycentron canadum cultured in submersible cages off Puerto Rico and
Received 9 September 2009 the Bahamas were comprehensively studied and are presented, discussed and compared to those of other
Received in revised form 16 February 2010 teleosts. Cobia grew to averages of 6.035 kg (specific growth rate (SGR) = 2.10%/day) in 363 days at the Puerto
Accepted 16 February 2010
Rico site (PR) and 3.545 kg (SGR= 2.04%/day) in 346 days at the Bahamas site (BA). Growth in length is best
expressed by the equations: y = 12+ 0.18x; r2 = 0.59 at PR and y = 12+ 0.16x; r2 = 0.86 at BA. The Laird–
Keywords:
Cobia
Gompertz model was used to represent growth in weight to best express the rate of decline in growth rate with
Growth age (α = 0.006194 PR and α = 0.006323 BA), which occurred at the onset of precocious maturation for this
Condition factor species at 2.0–4.5 kg in 300 days post hatch (dph). The exponents (b) of length–weight relationships calculated
Open ocean aquaculture (3.31 at PR and 3.20 at BA) demonstrate that cultured cobia exhibit greater condition factors than their wild
counterparts (b = 2.8) and explain the morphological differences observed between wild and culture cobia. Final
stocking densities of 5 and 15 kg/m3 were estimated at PR and BA, respectively. Average water temperatures for
the duration of the growout cycle were 27.8 °C at PR and 25.5 °C at BA. Results show that growth rates of cobia
vary widely and suggest a negative effect of lower temperature and increasing stocking density. These trials also
demonstrate, for the first time, that growout of cobia is technically feasible in submerged open ocean cages.
© 2010 Elsevier B.V. All rights reserved.

1. Introduction
Cobia, Rachycentron canadum, is considered one of the most
promising candidates for warm-water marine fish aquaculture in the
world (Kaiser and Holt, 2004; Liao et al., 2004; Benetti et al., 2007).
Cobia, also known as lemonfish or ling, is the only member of the
family Rachycentridae, and is found in the warm-temperate to tropical
waters of the West and East Atlantic, throughout the Caribbean and in
the Indo-Pacific off India, Australia and Japan (Briggs, 1960; Hassler
and Rainville, 1975; Shaffer and Nakamura, 1989; Ditty and Shaw,
1992). In the Eastern Pacific its occurrence has been reported as
marginal (Fowler, 1944; Briggs, 1960; Collette, 1999). Cobia are eury-
thermal and euryhaline, tolerating ranges of temperature and salinity
between 16.8 and 32.2 °C and 5–44.5 ppt, respectively (Shaffer and
Nakamura, 1989; Resley et al., 2006).
Cobia cage aquaculture was initiated in the early 1990s in Taiwan
(Yeh, 2000; Liao et al., 2004). By 1997, technology for mass production
of juveniles had been developed allowing for the expansion of the
⁎ Corresponding author. Rosentiel School of Marine and Atmospheric Science,
University of Miami, FL, USA. Tel.: +1 305 421 4889; fax: +1 305 421 4675.
E-mail address: dbenetti@rsmas.miami.edu (D.D. Benetti).
cage culture industry in Taiwan (Yeh et al., 1998) and by 1999, four
commercial hatcheries were operating in Taiwan, producing around
3 million cobia juveniles annually (Yeh, 2000). Since then, cobia
aquaculture production has been steadily expanding in Asia, primarily
in Taiwan, Vietnam and China, but also in other Southeast and Indo-
Pacific Asian countries including the Philippines, Indonesia, Iran and
Reunion Island. Floating cages continue to be the preferred method
of growout worldwide. Although production is expanding rapidly,
combined production of Asian countries is still rather low, on the
order of tens of thousands of metric tonnes per year (Liao et al., 2004).
Most recently, Australia and Marshall Island have begun developing
hatchery and cage culture operations for cobia.
In the Western Hemisphere, cobia aquaculture has also begun to
emerge as a promising industry. Early attempts to culture cobia in the
US began in the 1970s using eggs collected from the wild (Hassler
and Rainville, 1975). Research accelerated in the late 1990s with the
first successful spawns obtained from captive broodstock fish (Arnold
et al., 2002) and today a large body of knowledge on cobia culture in
the United States continues to accrue (Caylor et al., 1994; DuPaul et al.,
1997; Franks et al., 2001; Faulk and Holt, 2006; Holt et al., 2007;
Benetti et al., 2008a,b). During the last 8 years, advanced technology
has been used to demonstrate the viability of raising hatchery-reared

0044-8486/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.aquaculture.2010.02.021
196 D.D. Benetti et al. / Aquaculture 302 (2010) 195–201
cobia in collaboration with the private sector (Snapperfarm, Inc. and
AquaSense LLC) using SeaStation (Net Systems LLC) and Aquapod
(Ocean Technologies LLC) submerged cages in exposed sites in Puerto
Rico, US (PR) and the Bahamas (BA). Hatchery-reared cobia are also
being cultured in traditional gravity cages in Belize, Panama, Brazil,
Dominican Republic, Martinique, and Mexico (Benetti et al., 2008a).
Despite the great strides made in cobia aquaculture over the last
decade, there remains a paucity of published information regarding
cobia growth rates in cages and the available data are highly variable.
This paper presents data on the growth of cobia reared in submersible
cages in Culebra Island, Puerto Rico (Snapperfarm, Inc. U.S.), and in
South Eleuthera, The Bahamas (Aquasense LLC and the Cape Eleuthera
Institute). Both operations had technological support from the
University of Miami Aquaculture Program.
2. Materials and methods
Cobia at both sites were reared in a submersible OceanSpar
Seastation 3000™ cage. These cages are of a trapezoidal–octagon
shape, with a height of 15 m, a maximum cross-section of 30 m, and a
volume of 2700 m3. Both cages were anchored over sandy bottom at
depths of 30 and 27 m in Puerto Rico and the Bahamas, respectively. The
top of the cage was approximately 10 m below the surface at the Puerto
Rico site and approximately 7 m below the surface at the Bahamas site.
Cobia juveniles, 45 days of age and ranging in weight from 1 to 5 g
(average 3 g), were shipped from The Aquaculture Center of the
Florida Keys in Florida, U.S. via airfreight. The juveniles stocked at both
sites were all F1 from the same parental wild broodstock, spawn batch
and hatchery run. They were initially stocked into nursery nets
(volume 600 m3, knotless mesh, opening size 3.25 mm) within the
main cages (2700 m3 volume). A total of 8000 juveniles (∼3 fish/m3)
were stocked into the cage off Puerto Rico and 14,000 juveniles (∼5.2
fish/m3) into the cage off the Bahamas. Fish were released from the
nursery net into the main growout cages after 1 month (Table 1).
Cobia were grown for 363 days in Puerto Rico and harvested and
sold to restaurants in Miami. Cobia were grown in the Bahamas for
346 days before successive entries by sharks into the cage caused an
estimated 90% of the fish to escape within a 3-week period. Growth
rates were accurately determined by regular sampling and measuring
individual lengths (cm) and weights (g). Mortalities were determined
by collecting and counting individual dead fish from the bottom of
the cages, however escapements at both sites required estimation of
final stocking densities and feed conversion rates. These estimates
were made by a combination of visual observations by divers and
researchers and by the total amount of feed given daily. Since fish
were fed to satiation at know rates ranging from 2 to 5% of their total
biomass per day, escapements were estimated by the proportional
reduction in the amount of feed given. All growout and harvesting
procedures described here were carried out in accordance with the
Code of Ethics of the World Medical Association (Declaration of
Helsinki) for Animal Experiments.
Table 1
Summary of cage site and cage characteristics as well as nursery net and main cage
volumes and corresponding initial stocking densities.
Parameter Culebra Bahamas
Distance from shore (km) 2 2
Water depth (m) 30 25.9
Bottom type Sandy Sandy
Top of cage from water surface (m) 10 7
Cage height (m) 15 15
Cage width (m) 20 20
Cage volume (m3) 2700 2700
Nursery net volume (m3) 600 600
Number of fingerlings stocked 8000 14,000
Initial nursery net stocking density (#/m3) 13.3 23.3
Initial main cage net stocking density (#/m3) 2.4 4.1
2.1. Growth parameters
Fish were sampled every 15 to 45 days during the two production
cycles and measured for total length (TL) and wet weight (W). Fish
were caught randomly by divers using nets during feeding events
and were subsequently sacrificed for sampling. Sample size ranged
from 2 to 20 individuals at a time and varied depending on the ability
of the divers to catch the fish. Sex was determined by visually
examining the gonads of each individual sampled. Growth of cobia in
weight was analyzed both as absolute and specific growths, and the
Laird–Gompertz model was fitted to weight data to project estimated
growth rates over time (Table 2).
2.2. Feeding
At both sites, cobia were fed a marine grower pellet (Aquaexcel
5310; 53% crude protein, 10% crude fat) formulated and manufactured
by Burris Mill, Inc. (Louisiana, USA). Fish were fed twice a day (early
morning and late afternoon) during the first 60 days post stocking and
once a day (mid morning) thereafter until harvest. Pellet size was
1.5 mm for the first 21 days and progressively increased to 3.0 mm,
5.0 mm, and 9.0 mm as the fish grew. Fish were initially hand-fed by
divers and subsequently by pipes using pumps fitted at a boat from
the surface. Juveniles held in the nursery cage were fed by hand. Divers
were used to observe feeding response in order to adjust the ration as
required. Throughout the nursery and growout cycle, cobia were fed to
satiation, corresponding to approximately 5% per day of the total
estimated biomass for the first 3 months and then progressively
decreasing to 2% per day during the following 9 months. Food
conversion rates (FCR) were estimated by calculating the ratio between
the total weight of food given and the total weight of fish harvested.
2.3. Water quality and cage maintenance
At the Puerto Rico site, a thorough environmental monitoring study
was conducted by Alston et al. (2005) using two Hydrolabs to collect
and record water temperature, salinity (conductivity), and dissolved
oxygen data at 15-minute intervals at the cage site and at the control site
located 375 m from the cage. In the Bahamas site, measurements were
made daily using a YSI D.O./temperature/conductivity meter at the cage
and control sites. At both sites, water samples were taken monthly for
ammonia-N, nitrite-N, and nitrate-N analyses as well as phosphorous
concentrations, with no significant differences found in any of these
parameters at both sites throughout the study (Alston et al., 2005;
Collins, 2006). Nets were routinely cleaned to reduce drag and maintain
consistent flow through of the cage. The mesh size on each of the main
cages was 3.8 cm (stretched). The daily water exchange rates through
Table 2
Summary of growth parameters used to express results of growth rates of cobia.
Parameter (unit) Equation
Absolute growth (g) ΔG = W2 − W1
Absolute growth rate (g/day) AGR = (W2 − W1) / (t2 − t1)
Relative growth RG = (W2 − W1) / W1
Relative growth rate RG = (W2 − W1) / W1(t2 − t1)
Instantaneous growth rate (g/day) IGR = (ln W2 − ln W1) / (t2 − t1)
Specific growth rate (%/day) SGR = 100 * (ln W2 − ln W1) / (t2 − t1)
Length–weight relationship W = aLb
Growth in length (cm) Lt = a + bt
Laird–Gompertz model in weight (g) Wt = W0 * exp * {(A0 / α) * (1 − exp − αt)}
W1 = initial wet weight of fish at stocking.
W2 = final wet weight of fish.
t1 = age at stocking.
t2 = age at end of grow out period.
Lt = total length at age t; a and b are constants.
W0 = initial weight at age = 0.
A0 = initial growth rate.
α = rate of decline in growth rate with age; (A0/α) is expressed as a constant.
 D.D. Benetti et al. / Aquaculture 302 (2010) 195–201
both cages was estimated at approximately 3800 m3/day by calculating
the area of cross-section of the submerged cage and the average current
at the location. Both cages were deployed in exposed locations where
current velocities ranged between 0.9 and 2.7 km. These conditions are
conducive to excellent water quality for the fish in the cages. Daily
diving was performed in both locations for observations of the nets and
removal of dead fish. Mortalities were accounted for daily. No drugs or
chemicals were used.
3. Results and discussion
A summary of the water quality parameters at both sites is given in
Table 3. Average water temperatures were 27.8 °C and 25.5 °C at the
Puerto Rico and Bahamas sites, respectively. Salinities at the two sites
averaged 34.6 ppt and 37.2 ppt, respectively and dissolved oxygen,
5.3 mg/L and 6.1 mg/L, respectively.
The fish stocked at the warmer, lower density Puerto Rico site
grew faster than those at the cooler, higher density Bahamas site.
After being stocked for growout in the submerged cages, cobia grew to
averages of 6 kg in 363 days and 3.5 kg in 346 days in Puerto Rico and
the Bahamas, respectively (Table 4). Results of cobia growth in length
and weight are expressed in Figs. 1 and 2, respectively. Although the
Laird–Gompertz model is generally used to express growth rates in
weight of early developmental stages of fish (Kramer and Zweifel,
1970; Cushing, 1995), this model best fitted the data presented to
show the inflexion point during the onset of maturation, at approx-
imately 10 months of age (300 dph). Indeed, the cobia were observed
to engage in courtship behavior and began spawning naturally in the
cages starting at approximately 10 months of age, by which time there
was a noticeable decrease in growth rates (Fig. 2). Fish in the two
cages had similar length–weight relationships (Fig. 3). The final
stocking densities in the two cages were approximately 5 kg m3 in the
Puerto Rico cage and 15 kg m3 in the Bahamas cage.
Both temperature and stocking densities are likely to have had an
effect on the differences of growth rates of cobia observed in these trials.
Sun et al. (2006), for example, compared the growth rate of juvenile
cobia (22 g) at temperatures of 23 and 27 °C and reported a 44%
improvement in growth over this 4 °C range. Although speculative,
assuming that a similar percentage difference in growth rate occurs for
larger fish between these temperatures (approximately 11% per
degree), then the predicted size difference between fish at the Puerto
Rico and Bahamas sites over the 2.3 °C water temperature difference
would be approximately 25%. Indeed, the Q10 physiological concept
(Schmidt-Nielsen, 1983) predicts that the metabolic rate of fish roughly
doubles with every 10 °C. This suggests that, within the specific scope of
growth of cobia, growth rates would increase proportionally within the
range of temperature the fish were exposed to (22–30 °C). Under this
assumption, the 2.3 °C higher average temperature observed in Puerto
Table 3
Summary of annual means and ranges in water quality parameters from the two cobia
cage aquaculture sites (data from Alston et al., 2005; Collins, 2006; Benetti et al., 2006).
Location Culebra
Temperature (°C)
Average 27.8
Min 26.0
Max 29.9
Dissolved oxygen (mg/l)
Average 5.3
Min 4.5
Max 6.9
Salinity (g/l)
Average 34.6
Min 33.5
Max 35.5
197
Table 4
Summary of growth parameter estimates of cobia from the cage culture sites in Puerto
Rico and the Bahamas.
Parameter Culebra Eleuthera
Initial age at stocking (dph) 45 45
Age at harvest (dph) 408 391
Growout duration (days) 363 346
Average initial size (g) 3 3
Average final size (g) 6066 3545
Absolute growth (g) 6063 3542
Absolute growth rate (g/day) 16.7 10.2
Instantaneous growth rate (g/day) 0.0210 0.0204
Specific growth rate (%/day) 2.1 2.04
A0/α 7.836 7.465
Rate of decline in growth rate α 0.006194 0.006323
Rico would explain 23% of the faster growth rate reported for cobia
raised at that site. Average salinity and D.O. were 2.6 ppt and 0.8 mg/L
lower at the Culebra site, but these differences are unlikely to account for
the faster cobia growth rates observed in this site than those of
temperature and stocking densities. Resley et al. (2006), for example
found minimal differences in the growth rate of juvenile cobia cultured
between salinities of 5 and 30 ppt.
Results of these trials are at the upper end of other published and
anecdotal reports of cobia growth rates, but the available data is
highly variable. Many of the reported growth rates are exceptionally
high. Liao et al. (2004) and Benetti et al. (2006, 2007) both reported
that cobia may grow to up to 6 kg in one year when stocked at low
densities (e.g. 3 kg m− 3) and similar growth rates have been achieved
in Mexico with cobia stocked at low densities (≤5 kg m− 3) in
traditional floating cages off the coast of Campeche in the Gulf of
Mexico. Additionally, growth rates of cobia raised in floating net cages
in estuarine waters (15–30 ppt) in São Paulo, Brazil ranged from 2.5 to
4.5 kg in 1 year and 1 g juveniles stocked at 5–10 kg m− 3 in
traditional floating cages in Belize have grown to 2.0–4.0 kg in
1 year (O. Stevens, Marine Farms Belize, personal communication). On
the other hand, there have also been reports of much lower growth
rates. Benetti et al. (2008b) reported that cobia stocked in tanks at
densities of 2–3 kg m− 3 with temperature ranges of 20–32 °C reached
only 2 kg in 12 months, and 3–6 kg in 24 months. Cobia have also
been known to grow to as little as 400 g in 1 year under adverse,
crowded conditions in land-based tanks in Northeastern Brazil (M.
Carvalho, TWB Brazil, personal communication).
Sex related growth differences, usually in favor of females, occur in
many fish species both in the wild and under culture conditions (Felip
et al., 2001). Cobia, considered gonochoristic, exhibit differential
growth rates between the sexes. Females grow faster and have been
shown to be significantly longer and heavier within year classes
(Shaffer and Nakamura, 1989; Franks et al., 1999; Benetti et al., 2007,
2008a). This trend in growth performance favoring females was
observed in both growout cycles and corroborates anecdotal reports
about differential growth rates.
The exponent of the length–weight relationship (b) calculated for
males and females combined in the Bahamas and Puerto Rico are 3.2
Eleuthera
and 3.3, respectively. These values are greater than the range of 2.83
to 3.03 reported for wild cobia in the literature (Richards, 1967;
25.5 Darracot, 1977). This explains the morphological differences observed
18.7
29.9
between wild and cultured cobia and prove that cultured cobia has a
higher condition factor than wild cobia. Indeed, wild cobia
are elongated, skinny and streamlined whereas cultured cobia are
6.1 invariably shorter and fatter, exhibiting an elliptical, oval appearance.
4.9
The effect of captivity in the fish morphology has been previously
6.5
demonstrated for flounder and a number of other species by Blaxter
(1988) and for dolphin fish and mutton snapper by Benetti et al.
37.2 (1995a,b, 2002). The most likely explanations for this pattern are
37.0 increased food intake and reduced level of activity of captive fish
37.6
(Christiansen and Jobling, 1990; Boisclair and Tang, 1993). Although
198 D.D. Benetti et al. / Aquaculture 302 (2010) 195–201

Fig. 1. Growth of cobia in length.

Fig. 2. Growth of cobia fitted to the Laird–Gompertz model.

other important morphological, anatomic and biochemical differences
such as lipid composition, gonadal and hepatic somatic indices have
been reported between wild and cultured gilthead sea bream
(Gregorakis et al., 2002) and sea bass (Alasalvar et al., 2002), no
such analysis were conducted in this study. Nonetheless, some of
these author's findings were corroborated by our observations of
excessive intra-peritoneal fat and visible fat deposits in abnormally
enlarged, “fatty” livers in most cobia sampled after 6 months of age.
Despite the intra-specific variability in reported growth rates of
cobia, the growth rates exhibited by the fish that were the subject of
these trials are generally faster than most other cultured fish, including
some semi-pelagic species such as Seriola mazatlana (=S. rivoliana),
which reaches 2.0–2.5 kg in 10–12 months after stocking (Benetti et al.,
1995b). Indeed, growth rates reported for cobia in this paper are several
times faster (and in some instances an order of magnitude higher) than
those reported for red drum Sciaenops ocellatus (Hopkins, 1990),
mutton snapper Lutjanus analis (Benetti et al., 2002; Watanabe et al.,
2005) European sea bass Dicentrarchus labrax (Kavadias et al., 2003),
gilthead sea bream Sparus aurata (Barbato et al., 1993; Kissil et al., 2000),
red sea bream Pagrus major (Ikenoue and Kafuku, 1992), Japanese
flounder Paralichthys olivaceus (Matsuoka, 1995), cod Gadus morhua
(Svasan et al., 2004; Imsland et al., 2007) and Atlantic Salmon Salmo

Fig. 3. Length–weight relationship of cobia in Puerto Rico and the Bahamas.

 D.D. Benetti et al. / Aquaculture 302 (2010) 195–201 199

salar (Gooley et al., 2000). Cobia's growth rate cannot, however, be
considered extraordinary as both bluefin tuna Thunnus thynnus and
dolphin fish Coryphaena hippurus grow faster than cobia. Dolphin fish
cultured in tanks in Hawaii, for example, grew to 4.93 kg in 9.5 months
(Benetti et al., 1995a) and juvenile bluefin tuna captured by purse seine
and transferred to a floating cage grew from 5 kg to 45 kg average in
540 days in the Adriatic Sea (Ticina et al., 2007) (Table 5).
Some of the variability in the growth rates of cobia may be due to
disease related stress. Cobia are prone to a number of different diseases
which seem to occur at all stages of their life cycle irrespective of the
method or containment system used for raising them. Before the es-
capements that occurred at both sides in these trials, mortalities were
estimated at 15 and 10% in Puerto Rico and the Bahamas, respectively.
Episodes of mortalities of fish weighing 2–4 kg were reported at both
sites and subsequently diagnosed as anemia, probably due to a
deficiency of iron in the diet. Upon our request, the feed manufacturer
increased the level of iron in the diet and mortalities subsequently
stopped. At the Puerto Rico site, outbreaks of parasitic (Amyloodinium
ocellatum) and bacterial (Photobacterium damsela) diseases have been
reported during the nursery stage. The ectoparasite Amyloodinium
caused approximately 50% mortality in a group of juveniles in 2003
although it did not affect the fish that were the subject of this trial and
has not occurred since then. Photobacterium could have caused the low-
rate, continuous mortality (2–4 fish per day) observed in the cage in
Puerto Rico during this trial, suggesting that they may have been causing
chronic stress to the fish. It is worth noting, however, that Photobacter-
ium were present in the Puerto Rico cage despite a relatively low
stocking density compared to cage at the Bahamas site. Similar problems
with pathogenic bacteria were also reported for cobia cultured in
floating net cages in Taiwan by Liao et al. (2004).
Necropsies were not conducted in the Bahamas site and the causes of
mortalities were undetermined. However, in a subsequent cobia
growout trial at the same site, nephrocalcinosis (kidney stones) was
considered the major cause of mortality during the growout stage.
Necropsies revealed the presence of one to several stones varying in size
from 2 mm to 6 mm in diameter in the kidney or blocking the urethra of
sick or dead fish (T. Soligo, Cape Eleuthera Institute, personal
communication). Although the mechanisms responsible for the devel-
opment of this condition are not fully understood, Stoskopf (1993)
reported that prolonged exposure of fish to free carbon dioxide greater
than 10–20 mg/L has been implicated as a causative agent. A possible
cause could be an imbalance in the calcium to magnesium ratio of the
diet, as magnesium deficiencies have been implicated in nephrocalci-
nosis (Stoskopf, 1993). This disease has proven to be quite common in
cobia both at the laboratory (hatchery) and in growout operations. It
occurs indistinctively in cobia of various sizes (including broodstock)
being fed pellets of different composition in cages and in tanks at
hatcheries and those fed fish, squid and shrimp in hatcheries.
Subsequently to this trial, the presence of large formations of
calcium carbonate deposits of 2–3 cm in diameter has been commonly
observed in the intestines of dead juvenile and adult cobia cultured at
the same site under similar conditions. Occasionally, formations
as large as 5–7 cm in diameter were found in the intestines and
abdominal cavities of 2–4 kg cobia. The causes and consequences
of this condition have not yet been determined or described. It is
possible that this condition was a factor causing the mortalities of
cobia during the growout in the Bahamas site.
Survival rates of cobia during growout have varied broadly, between
as low as 10% to as high as 90% throughout the Americas and the
Caribbean. The low survival rates are partly due to the aforementioned
disease problems, but the most serious problem has been escapes due to
shark attacks on the nets of cages. There were losses due to escapements
at both sites, although the losses in Puerto Rico were considerably lower,
estimated at 20% of the total crop. However, since the end of the trials
described, problems with escapements have been brought under control
at both sites with the establishment of better management practices, such
as efficient collection and removal of mortalities from cages, and cage
systems with improved anti-predator devices. Indeed, since the instal-
lation of anti-predator nets in the Bahamas and in Puerto Rico further to
these trials, the predator problem has been brought under control.
As a consequence of disease outbreaks and highly variable survival
rates, food conversion ratios (FCR) fluctuated broadly between 1.3
during the juvenile stages and 2.2 at the later stages. The higher
estimated FCRs (N2) towards the end of the trials in both Puerto Rico
and the Bahamas were mostly driven by the large number of escapees
and, to a lesser degree, to mortalities. The overall estimated FCR of cobia
raised in both sites was 1.8 prior to the escapements at 8–10 months.
Conversely, FCRs as low as 1.01 to 1.45 have been recorded for cobia
juveniles in tanks with no mortality (Denlinger, 2007). Although FCRs
have been steadily decreasing as management strategies improve and
disease related mortalities decrease, combined data and information
from several operations and experimental trials in the Americas and the

Table 5
Comparison of growth in various tropical and temperate marine finfish species from cage aquaculture operations around the world.

Species Temperature Harvest size Age Growth rate Stocking densitya Reference
(°C) (Kg) (Months) (g/month) (kg/m3)

Warm/tropical
Cobia (Rachycentron canadum) 20–30 3–10 12–24 250–500 5–10 Su et al. (2000), current study
Barramundi (Lates calcifer) 24–30 0.7–3 4–24 88–125 5–10 Williams et al. (2000)
Red drum (Sciaenops ocellata) 20–30 1–3 12–20 75–150 5–10 Hopkins (1990)
Greater amberjack (Seriola dumerili) 20–30 0.9–3 7–18 111–167 5–10 Chambers and Ostrowski (1999), Tucker (1998)
Yellowtail/almaco jack (Seriola rivoliana/ 20–30 1–3 9–18 83–250 5–10 Benetti et al. (1995b), Benetti (1997)
mazatlana)
Mutton snapper (Lutjanus analis) 20–30 0.3–1 6–12 44–83 5–10 Benetti et al. (2002), Watanabe et al. (2005)
Dolphin/mahi-mahi (Coryphaena hippurus) 20–30 2–8 6–12 333–667 5–10 Benetti (2001), Benetti et al. (1995a)

Temperate/subtropical
Japanese Hamachi (Seriola quinqueradiata) 15–28 1.5–7 12–24 125–292 5–10 Kafuku and Ikenoue (1992), Benetti et al. (2005)
Kingfish/yellowtail jack (Seriola lalandi/dorsalis) 14–24 1.5–3 8–13 153–230 5–10 Kolkovski and Sakakura (2007), Benetti et al. (2005)
European sea bass (Dicentrarchus labrax) 12–26 0.4–3 12–30 33–125 10–25 Kavadias et al. (2003)
Sea bream (Sparus aurata) 12–26 0.28–2 8–24 35–83 10–25 Barbato et al. (1993)
Bluefin tuna (Thunnus thynnus) 15–25 3.5–45 12–84 291–2380 1–4 Tucker (1998), Wright (2008), Ticina et al. (2007)

Cold/temperate
Cod (Gadus morhua) 10–15 2.2–3.1 24–28 85–129 5–10 Svasan et al. (2004), Imsland et al. (2007)
Atlantic halibut (Hippoglossus hippoglossus) 6–12 3–6 24–36 125–166 100–200b Holm et al. (2004)
Atlantic salmon (Salmo salar) 8–15 2–8 24–26 83–222 5–15 Gooley et al. (2000)
a
Final density at harvest.
b
Stocking density in percent bottom coverage.
200 D.D. Benetti et al. / Aquaculture 302 (2010) 195–201
Caribbean region indicate that the average FCR of cobia is 1.75. This
average compares favorably to the data provided by Nakamura (2007)
for cultured cobia fed on pellets in Asia, however the range of 1.3–2.2 is
in accordance with the data provided by that author.
These trials were the two first demonstration projects of cobia
growout in cages in the Western hemisphere. Due to the nature of
the open ocean ventures, logistics and economic constraints (the cost
of purchasing, deploying, stocking and operating each cage was
N$500,000/year), only one cage was available for these trials in each
site, not allowing for replications. For this reason, it is not possible to
precisely discern the combined or isolated effects of temperature
and stocking densities on the growth rates of cobia in these trials.
Nonetheless, results indicate that growout of cobia is technically
feasible in submerged open ocean cages and suggest that growth rates
are negatively influenced by decreasing temperatures and/or increas-
ing stocking density. Perhaps more importantly, when compared with
other reported growth rates, these results also reveal the considerable
amount of uncertainty that plagues the developing cobia industry in
the Americas. Increased knowledge and experience on the various
factors that influence cobia growth rates are required so that uncer-
tainties can be eliminated and production planning rationalized.
Acknowledgements
This research was funded by the National Oceanic and Atmospheric
Administration under grants No. NA060AR4170238 from the NOAA
National Marine Aquaculture Initiative and No. NA08OAR4170826 of the
NOAA Marine Aquaculture Program. The research was also supported by a
previous grant from the Florida Sea Grant College Program No. RLRA42.
The authors are indebted to the technical staffs and divers of Snapperfarm,
Inc. of Culebra, Puerto Rico and the Island School/Cape Eleuthera Institute
in South Eleuthera, the Bahamas, for their diligent assistance in feeding,
data collection and daily diving at the respective cage sites. The authors
acknowledge and are grateful to Mr. Johann Scheidt of Aquasense LLC for
his support and involvement in the demonstration project in South
Eleuthera, the Bahamas. We also thank Jorge Alarcón and Owen Stevens of
Marine Farms Belize, Marcell Carvalho of TWB Brazil, and Thiago Soligo of
the Cape Eleuthera Institute for sharing information about cobia growout.
Finally, the authors would like to thank Dr. Gavin Partridge from the
Challenger TAFE Aquaculture Development Unit in Fremantle, Western
Australia, for his review and contributions to the manuscript while on
post-doctoral training with the first author.
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Daniel D. Benetti: Dr. Benetti received his M.S. in

Aquaculture and Fishery Management from the University
of Stirling and his Ph.D. in Marine Biology and Fisheries
from the University of Miami. Dr. Benetti has served as the
director of The Mariculture Project in Ecuador, consulted
for aquaculture firms throughout the world, worked as a
research scientist at the Harbor Branch Oceanographic
Institution, and most recently (2003 to 2008) he was the
chair of the Division of Marine Affairs and Policy at the
University of Miami. He currently serves as professor and
director of the aquaculture program at the University of
Miami.