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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 cage culture industry in Taiwan (Yeh et al., 1998) and by 1999, four
commercial hatcheries were operating in Taiwan, producing around
Cobia, Rachycentron canadum, is considered one of the most 3 million cobia juveniles annually (Yeh, 2000). Since then, cobia
promising candidates for warm-water marine fish aquaculture in the aquaculture production has been steadily expanding in Asia, primarily
world (Kaiser and Holt, 2004; Liao et al., 2004; Benetti et al., 2007). in Taiwan, Vietnam and China, but also in other Southeast and Indo-
Cobia, also known as lemonfish or ling, is the only member of the Pacific Asian countries including the Philippines, Indonesia, Iran and
family Rachycentridae, and is found in the warm-temperate to tropical Reunion Island. Floating cages continue to be the preferred method
waters of the West and East Atlantic, throughout the Caribbean and in of growout worldwide. Although production is expanding rapidly,
the Indo-Pacific off India, Australia and Japan (Briggs, 1960; Hassler combined production of Asian countries is still rather low, on the
and Rainville, 1975; Shaffer and Nakamura, 1989; Ditty and Shaw, order of tens of thousands of metric tonnes per year (Liao et al., 2004).
1992). In the Eastern Pacific its occurrence has been reported as Most recently, Australia and Marshall Island have begun developing
marginal (Fowler, 1944; Briggs, 1960; Collette, 1999). Cobia are eury- hatchery and cage culture operations for cobia.
thermal and euryhaline, tolerating ranges of temperature and salinity In the Western Hemisphere, cobia aquaculture has also begun to
between 16.8 and 32.2 °C and 5–44.5 ppt, respectively (Shaffer and emerge as a promising industry. Early attempts to culture cobia in the
Nakamura, 1989; Resley et al., 2006). US began in the 1970s using eggs collected from the wild (Hassler
Cobia cage aquaculture was initiated in the early 1990s in Taiwan and Rainville, 1975). Research accelerated in the late 1990s with the
(Yeh, 2000; Liao et al., 2004). By 1997, technology for mass production first successful spawns obtained from captive broodstock fish (Arnold
of juveniles had been developed allowing for the expansion of the 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.,
⁎ Corresponding author. Rosentiel School of Marine and Atmospheric Science,
1997; Franks et al., 2001; Faulk and Holt, 2006; Holt et al., 2007;
University of Miami, FL, USA. Tel.: +1 305 421 4889; fax: +1 305 421 4675. Benetti et al., 2008a,b). During the last 8 years, advanced technology
E-mail address: dbenetti@rsmas.miami.edu (D.D. Benetti). 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 2.1. Growth parameters
AquaSense LLC) using SeaStation (Net Systems LLC) and Aquapod
(Ocean Technologies LLC) submerged cages in exposed sites in Puerto Fish were sampled every 15 to 45 days during the two production
Rico, US (PR) and the Bahamas (BA). Hatchery-reared cobia are also cycles and measured for total length (TL) and wet weight (W). Fish
being cultured in traditional gravity cages in Belize, Panama, Brazil, were caught randomly by divers using nets during feeding events
Dominican Republic, Martinique, and Mexico (Benetti et al., 2008a). and were subsequently sacrificed for sampling. Sample size ranged
Despite the great strides made in cobia aquaculture over the last from 2 to 20 individuals at a time and varied depending on the ability
decade, there remains a paucity of published information regarding of the divers to catch the fish. Sex was determined by visually
cobia growth rates in cages and the available data are highly variable. examining the gonads of each individual sampled. Growth of cobia in
This paper presents data on the growth of cobia reared in submersible weight was analyzed both as absolute and specific growths, and the
cages in Culebra Island, Puerto Rico (Snapperfarm, Inc. U.S.), and in Laird–Gompertz model was fitted to weight data to project estimated
South Eleuthera, The Bahamas (Aquasense LLC and the Cape Eleuthera growth rates over time (Table 2).
Institute). Both operations had technological support from the
University of Miami Aquaculture Program. 2.2. Feeding

2. Materials and methods At both sites, cobia were fed a marine grower pellet (Aquaexcel
5310; 53% crude protein, 10% crude fat) formulated and manufactured
Cobia at both sites were reared in a submersible OceanSpar by Burris Mill, Inc. (Louisiana, USA). Fish were fed twice a day (early
Seastation 3000™ cage. These cages are of a trapezoidal–octagon morning and late afternoon) during the first 60 days post stocking and
shape, with a height of 15 m, a maximum cross-section of 30 m, and a once a day (mid morning) thereafter until harvest. Pellet size was
volume of 2700 m3. Both cages were anchored over sandy bottom at 1.5 mm for the first 21 days and progressively increased to 3.0 mm,
depths of 30 and 27 m in Puerto Rico and the Bahamas, respectively. The 5.0 mm, and 9.0 mm as the fish grew. Fish were initially hand-fed by
top of the cage was approximately 10 m below the surface at the Puerto divers and subsequently by pipes using pumps fitted at a boat from
Rico site and approximately 7 m below the surface at the Bahamas site. the surface. Juveniles held in the nursery cage were fed by hand. Divers
Cobia juveniles, 45 days of age and ranging in weight from 1 to 5 g were used to observe feeding response in order to adjust the ration as
(average 3 g), were shipped from The Aquaculture Center of the required. Throughout the nursery and growout cycle, cobia were fed to
Florida Keys in Florida, U.S. via airfreight. The juveniles stocked at both satiation, corresponding to approximately 5% per day of the total
sites were all F1 from the same parental wild broodstock, spawn batch estimated biomass for the first 3 months and then progressively
and hatchery run. They were initially stocked into nursery nets decreasing to 2% per day during the following 9 months. Food
(volume 600 m3, knotless mesh, opening size 3.25 mm) within the conversion rates (FCR) were estimated by calculating the ratio between
main cages (2700 m3 volume). A total of 8000 juveniles (∼3 fish/m3) the total weight of food given and the total weight of fish harvested.
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 2.3. Water quality and cage maintenance
nursery net into the main growout cages after 1 month (Table 1).
Cobia were grown for 363 days in Puerto Rico and harvested and At the Puerto Rico site, a thorough environmental monitoring study
sold to restaurants in Miami. Cobia were grown in the Bahamas for was conducted by Alston et al. (2005) using two Hydrolabs to collect
346 days before successive entries by sharks into the cage caused an and record water temperature, salinity (conductivity), and dissolved
estimated 90% of the fish to escape within a 3-week period. Growth oxygen data at 15-minute intervals at the cage site and at the control site
rates were accurately determined by regular sampling and measuring located 375 m from the cage. In the Bahamas site, measurements were
individual lengths (cm) and weights (g). Mortalities were determined made daily using a YSI D.O./temperature/conductivity meter at the cage
by collecting and counting individual dead fish from the bottom of and control sites. At both sites, water samples were taken monthly for
the cages, however escapements at both sites required estimation of ammonia-N, nitrite-N, and nitrate-N analyses as well as phosphorous
final stocking densities and feed conversion rates. These estimates concentrations, with no significant differences found in any of these
were made by a combination of visual observations by divers and parameters at both sites throughout the study (Alston et al., 2005;
researchers and by the total amount of feed given daily. Since fish Collins, 2006). Nets were routinely cleaned to reduce drag and maintain
were fed to satiation at know rates ranging from 2 to 5% of their total consistent flow through of the cage. The mesh size on each of the main
biomass per day, escapements were estimated by the proportional cages was 3.8 cm (stretched). The daily water exchange rates through
reduction in the amount of feed given. All growout and harvesting
procedures described here were carried out in accordance with the Table 2
Code of Ethics of the World Medical Association (Declaration of Summary of growth parameters used to express results of growth rates of cobia.

Helsinki) for Animal Experiments. Parameter (unit) Equation

Absolute growth (g) ΔG = W2 − W1


Table 1 Absolute growth rate (g/day) AGR = (W2 − W1) / (t2 − t1)
Summary of cage site and cage characteristics as well as nursery net and main cage Relative growth RG = (W2 − W1) / W1
volumes and corresponding initial stocking densities. Relative growth rate RG = (W2 − W1) / W1(t2 − t1)
Instantaneous growth rate (g/day) IGR = (ln W2 − ln W1) / (t2 − t1)
Parameter Culebra Bahamas
Specific growth rate (%/day) SGR = 100 * (ln W2 − ln W1) / (t2 − t1)
Distance from shore (km) 2 2 Length–weight relationship W = aLb
Water depth (m) 30 25.9 Growth in length (cm) Lt = a + bt
Bottom type Sandy Sandy Laird–Gompertz model in weight (g) Wt = W0 * exp * {(A0 / α) * (1 − exp − αt)}
Top of cage from water surface (m) 10 7
W1 = initial wet weight of fish at stocking.
Cage height (m) 15 15
W2 = final wet weight of fish.
Cage width (m) 20 20
t1 = age at stocking.
Cage volume (m3) 2700 2700
t2 = age at end of grow out period.
Nursery net volume (m3) 600 600
Lt = total length at age t; a and b are constants.
Number of fingerlings stocked 8000 14,000
W0 = initial weight at age = 0.
Initial nursery net stocking density (#/m3) 13.3 23.3
A0 = initial growth rate.
Initial main cage net stocking density (#/m3) 2.4 4.1
α = rate of decline in growth rate with age; (A0/α) is expressed as a constant.
D.D. Benetti et al. / Aquaculture 302 (2010) 195–201 197

both cages was estimated at approximately 3800 m3/day by calculating Table 4


the area of cross-section of the submerged cage and the average current Summary of growth parameter estimates of cobia from the cage culture sites in Puerto
Rico and the Bahamas.
at the location. Both cages were deployed in exposed locations where
current velocities ranged between 0.9 and 2.7 km. These conditions are Parameter Culebra Eleuthera
conducive to excellent water quality for the fish in the cages. Daily Initial age at stocking (dph) 45 45
diving was performed in both locations for observations of the nets and Age at harvest (dph) 408 391
removal of dead fish. Mortalities were accounted for daily. No drugs or Growout duration (days) 363 346
Average initial size (g) 3 3
chemicals were used.
Average final size (g) 6066 3545
Absolute growth (g) 6063 3542
Absolute growth rate (g/day) 16.7 10.2
3. Results and discussion Instantaneous growth rate (g/day) 0.0210 0.0204
Specific growth rate (%/day) 2.1 2.04
A0/α 7.836 7.465
A summary of the water quality parameters at both sites is given in
Rate of decline in growth rate α 0.006194 0.006323
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, Rico would explain 23% of the faster growth rate reported for cobia
5.3 mg/L and 6.1 mg/L, respectively. raised at that site. Average salinity and D.O. were 2.6 ppt and 0.8 mg/L
The fish stocked at the warmer, lower density Puerto Rico site lower at the Culebra site, but these differences are unlikely to account for
grew faster than those at the cooler, higher density Bahamas site. the faster cobia growth rates observed in this site than those of
After being stocked for growout in the submerged cages, cobia grew to temperature and stocking densities. Resley et al. (2006), for example
averages of 6 kg in 363 days and 3.5 kg in 346 days in Puerto Rico and found minimal differences in the growth rate of juvenile cobia cultured
the Bahamas, respectively (Table 4). Results of cobia growth in length between salinities of 5 and 30 ppt.
and weight are expressed in Figs. 1 and 2, respectively. Although the Results of these trials are at the upper end of other published and
Laird–Gompertz model is generally used to express growth rates in anecdotal reports of cobia growth rates, but the available data is
weight of early developmental stages of fish (Kramer and Zweifel, highly variable. Many of the reported growth rates are exceptionally
1970; Cushing, 1995), this model best fitted the data presented to high. Liao et al. (2004) and Benetti et al. (2006, 2007) both reported
show the inflexion point during the onset of maturation, at approx- that cobia may grow to up to 6 kg in one year when stocked at low
imately 10 months of age (300 dph). Indeed, the cobia were observed densities (e.g. 3 kg m− 3) and similar growth rates have been achieved
to engage in courtship behavior and began spawning naturally in the in Mexico with cobia stocked at low densities (≤5 kg m− 3) in
cages starting at approximately 10 months of age, by which time there traditional floating cages off the coast of Campeche in the Gulf of
was a noticeable decrease in growth rates (Fig. 2). Fish in the two Mexico. Additionally, growth rates of cobia raised in floating net cages
cages had similar length–weight relationships (Fig. 3). The final in estuarine waters (15–30 ppt) in São Paulo, Brazil ranged from 2.5 to
stocking densities in the two cages were approximately 5 kg m3 in the 4.5 kg in 1 year and 1 g juveniles stocked at 5–10 kg m− 3 in
Puerto Rico cage and 15 kg m3 in the Bahamas cage. traditional floating cages in Belize have grown to 2.0–4.0 kg in
Both temperature and stocking densities are likely to have had an 1 year (O. Stevens, Marine Farms Belize, personal communication). On
effect on the differences of growth rates of cobia observed in these trials. the other hand, there have also been reports of much lower growth
Sun et al. (2006), for example, compared the growth rate of juvenile rates. Benetti et al. (2008b) reported that cobia stocked in tanks at
cobia (22 g) at temperatures of 23 and 27 °C and reported a 44% densities of 2–3 kg m− 3 with temperature ranges of 20–32 °C reached
improvement in growth over this 4 °C range. Although speculative, only 2 kg in 12 months, and 3–6 kg in 24 months. Cobia have also
assuming that a similar percentage difference in growth rate occurs for been known to grow to as little as 400 g in 1 year under adverse,
larger fish between these temperatures (approximately 11% per crowded conditions in land-based tanks in Northeastern Brazil (M.
degree), then the predicted size difference between fish at the Puerto Carvalho, TWB Brazil, personal communication).
Rico and Bahamas sites over the 2.3 °C water temperature difference Sex related growth differences, usually in favor of females, occur in
would be approximately 25%. Indeed, the Q10 physiological concept many fish species both in the wild and under culture conditions (Felip
(Schmidt-Nielsen, 1983) predicts that the metabolic rate of fish roughly et al., 2001). Cobia, considered gonochoristic, exhibit differential
doubles with every 10 °C. This suggests that, within the specific scope of growth rates between the sexes. Females grow faster and have been
growth of cobia, growth rates would increase proportionally within the shown to be significantly longer and heavier within year classes
range of temperature the fish were exposed to (22–30 °C). Under this (Shaffer and Nakamura, 1989; Franks et al., 1999; Benetti et al., 2007,
assumption, the 2.3 °C higher average temperature observed in Puerto 2008a). This trend in growth performance favoring females was
observed in both growout cycles and corroborates anecdotal reports
Table 3
about differential growth rates.
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). The exponent of the length–weight relationship (b) calculated for
males and females combined in the Bahamas and Puerto Rico are 3.2
Location Culebra Eleuthera
and 3.3, respectively. These values are greater than the range of 2.83
Temperature (°C) to 3.03 reported for wild cobia in the literature (Richards, 1967;
Average 27.8 25.5 Darracot, 1977). This explains the morphological differences observed
Min 26.0 18.7
Max 29.9 29.9
between wild and cultured cobia and prove that cultured cobia has a
higher condition factor than wild cobia. Indeed, wild cobia
Dissolved oxygen (mg/l) are elongated, skinny and streamlined whereas cultured cobia are
Average 5.3 6.1 invariably shorter and fatter, exhibiting an elliptical, oval appearance.
Min 4.5 4.9
The effect of captivity in the fish morphology has been previously
Max 6.9 6.5
demonstrated for flounder and a number of other species by Blaxter
Salinity (g/l) (1988) and for dolphin fish and mutton snapper by Benetti et al.
Average 34.6 37.2 (1995a,b, 2002). The most likely explanations for this pattern are
Min 33.5 37.0 increased food intake and reduced level of activity of captive fish
Max 35.5 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 some semi-pelagic species such as Seriola mazatlana (=S. rivoliana),
such as lipid composition, gonadal and hepatic somatic indices have which reaches 2.0–2.5 kg in 10–12 months after stocking (Benetti et al.,
been reported between wild and cultured gilthead sea bream 1995b). Indeed, growth rates reported for cobia in this paper are several
(Gregorakis et al., 2002) and sea bass (Alasalvar et al., 2002), no times faster (and in some instances an order of magnitude higher) than
such analysis were conducted in this study. Nonetheless, some of those reported for red drum Sciaenops ocellatus (Hopkins, 1990),
these author's findings were corroborated by our observations of mutton snapper Lutjanus analis (Benetti et al., 2002; Watanabe et al.,
excessive intra-peritoneal fat and visible fat deposits in abnormally 2005) European sea bass Dicentrarchus labrax (Kavadias et al., 2003),
enlarged, “fatty” livers in most cobia sampled after 6 months of age. gilthead sea bream Sparus aurata (Barbato et al., 1993; Kissil et al., 2000),
Despite the intra-specific variability in reported growth rates of red sea bream Pagrus major (Ikenoue and Kafuku, 1992), Japanese
cobia, the growth rates exhibited by the fish that were the subject of flounder Paralichthys olivaceus (Matsuoka, 1995), cod Gadus morhua
these trials are generally faster than most other cultured fish, including (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 cause could be an imbalance in the calcium to magnesium ratio of the
considered extraordinary as both bluefin tuna Thunnus thynnus and diet, as magnesium deficiencies have been implicated in nephrocalci-
dolphin fish Coryphaena hippurus grow faster than cobia. Dolphin fish nosis (Stoskopf, 1993). This disease has proven to be quite common in
cultured in tanks in Hawaii, for example, grew to 4.93 kg in 9.5 months cobia both at the laboratory (hatchery) and in growout operations. It
(Benetti et al., 1995a) and juvenile bluefin tuna captured by purse seine occurs indistinctively in cobia of various sizes (including broodstock)
and transferred to a floating cage grew from 5 kg to 45 kg average in being fed pellets of different composition in cages and in tanks at
540 days in the Adriatic Sea (Ticina et al., 2007) (Table 5). hatcheries and those fed fish, squid and shrimp in hatcheries.
Some of the variability in the growth rates of cobia may be due to Subsequently to this trial, the presence of large formations of
disease related stress. Cobia are prone to a number of different diseases calcium carbonate deposits of 2–3 cm in diameter has been commonly
which seem to occur at all stages of their life cycle irrespective of the observed in the intestines of dead juvenile and adult cobia cultured at
method or containment system used for raising them. Before the es- the same site under similar conditions. Occasionally, formations
capements that occurred at both sides in these trials, mortalities were as large as 5–7 cm in diameter were found in the intestines and
estimated at 15 and 10% in Puerto Rico and the Bahamas, respectively. abdominal cavities of 2–4 kg cobia. The causes and consequences
Episodes of mortalities of fish weighing 2–4 kg were reported at both of this condition have not yet been determined or described. It is
sites and subsequently diagnosed as anemia, probably due to a possible that this condition was a factor causing the mortalities of
deficiency of iron in the diet. Upon our request, the feed manufacturer cobia during the growout in the Bahamas site.
increased the level of iron in the diet and mortalities subsequently Survival rates of cobia during growout have varied broadly, between
stopped. At the Puerto Rico site, outbreaks of parasitic (Amyloodinium as low as 10% to as high as 90% throughout the Americas and the
ocellatum) and bacterial (Photobacterium damsela) diseases have been Caribbean. The low survival rates are partly due to the aforementioned
reported during the nursery stage. The ectoparasite Amyloodinium disease problems, but the most serious problem has been escapes due to
caused approximately 50% mortality in a group of juveniles in 2003 shark attacks on the nets of cages. There were losses due to escapements
although it did not affect the fish that were the subject of this trial and at both sites, although the losses in Puerto Rico were considerably lower,
has not occurred since then. Photobacterium could have caused the low- estimated at 20% of the total crop. However, since the end of the trials
rate, continuous mortality (2–4 fish per day) observed in the cage in described, problems with escapements have been brought under control
Puerto Rico during this trial, suggesting that they may have been causing at both sites with the establishment of better management practices, such
chronic stress to the fish. It is worth noting, however, that Photobacter- as efficient collection and removal of mortalities from cages, and cage
ium were present in the Puerto Rico cage despite a relatively low systems with improved anti-predator devices. Indeed, since the instal-
stocking density compared to cage at the Bahamas site. Similar problems lation of anti-predator nets in the Bahamas and in Puerto Rico further to
with pathogenic bacteria were also reported for cobia cultured in these trials, the predator problem has been brought under control.
floating net cages in Taiwan by Liao et al. (2004). As a consequence of disease outbreaks and highly variable survival
Necropsies were not conducted in the Bahamas site and the causes of rates, food conversion ratios (FCR) fluctuated broadly between 1.3
mortalities were undetermined. However, in a subsequent cobia during the juvenile stages and 2.2 at the later stages. The higher
growout trial at the same site, nephrocalcinosis (kidney stones) was estimated FCRs (N2) towards the end of the trials in both Puerto Rico
considered the major cause of mortality during the growout stage. and the Bahamas were mostly driven by the large number of escapees
Necropsies revealed the presence of one to several stones varying in size and, to a lesser degree, to mortalities. The overall estimated FCR of cobia
from 2 mm to 6 mm in diameter in the kidney or blocking the urethra of raised in both sites was 1.8 prior to the escapements at 8–10 months.
sick or dead fish (T. Soligo, Cape Eleuthera Institute, personal Conversely, FCRs as low as 1.01 to 1.45 have been recorded for cobia
communication). Although the mechanisms responsible for the devel- juveniles in tanks with no mortality (Denlinger, 2007). Although FCRs
opment of this condition are not fully understood, Stoskopf (1993) have been steadily decreasing as management strategies improve and
reported that prolonged exposure of fish to free carbon dioxide greater disease related mortalities decrease, combined data and information
than 10–20 mg/L has been implicated as a causative agent. A possible 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 Benetti, D.D., Nakada, M., Shotton, S., Poortenaar, C., Tracy, P.L., Hutchinson, W., 2005.
Aquaculture of three species of yellowtail jacks. In: Kelly, A.M., Silverstein, J. (Eds.),
average compares favorably to the data provided by Nakamura (2007) American Fisheries Society Symposium 46: Aquaculture in the 21st Century,
for cultured cobia fed on pellets in Asia, however the range of 1.3–2.2 is Phoenix, AZ, USA, pp. 491–515.
in accordance with the data provided by that author. Benetti, D.D., Brand, L., Collins, J., Orhun, R., Benetti, A., O'Hanlon, B., Danylchuk, A.,
Alston, D., Rivera, J., Cabarcas, A., 2006. Can offshore aquaculture of carnivorous fish
These trials were the two first demonstration projects of cobia be sustainable? Case Studies from the Caribbean. World Aquac. 37 (1), 44–47.
growout in cages in the Western hemisphere. Due to the nature of Benetti, D.D., Orhun, M.R., Zink, I., Cavalin, F.G., Sardenberg, B., Palmer, K., Denlinger, B.,
the open ocean ventures, logistics and economic constraints (the cost Bacoat, D., O'Hanlon, B., 2007. Aquaculture of cobia (Rachycentron canadum) in the
Americas and the Caribbean. In: Liao, I.C., Leaño, E.M. (Eds.), Cobia Aquaculture:
of purchasing, deploying, stocking and operating each cage was
Research, Development and Commercial Production. Asian Fisheries Society, Manila,
N$500,000/year), only one cage was available for these trials in each Philippines, World Aquaculture Society, Louisiana, USA, The Fisheries Society of Taiwan,
site, not allowing for replications. For this reason, it is not possible to Keelung, Taiwan, and National Taiwan Ocean University, Keelung, Taiwan, pp. 57–78.
Benetti, D.D., Orhun, M.R., Sardenberg, B., O'Hanlon, B., Welch, A., Hoenig, R., Zink, I.,
precisely discern the combined or isolated effects of temperature
Rivera, J.A., Denlinger, B., Bacoat, D., Palmer, K., Cavalin, F., 2008a. Advances in
and stocking densities on the growth rates of cobia in these trials. hatchery and grow-out technology of cobia Rachycentron canadum (Linnaeus).
Nonetheless, results indicate that growout of cobia is technically Aquac. Res. 39, 701–711.
feasible in submerged open ocean cages and suggest that growth rates Benetti, D.D., Sardenberg, B., Welch, A., Hoenig, R., Orhun, M.R., Zink, I., 2008b. Intensive
larval husbandry and fingerling production of cobia Rachycentron canadum.
are negatively influenced by decreasing temperatures and/or increas- Aquaculture 281, 22–27.
ing stocking density. Perhaps more importantly, when compared with Blaxter, J.H.S., 1988. Pattern and variety in development. In: Hoar, W.S., Randall, D.J.
other reported growth rates, these results also reveal the considerable (Eds.), Fish Physiology, vol. 11a. Academy Press, New York, pp. 1–58.
Boisclair, D., Tang, M., 1993. Empirical analysis of the influence of swimming pattern on
amount of uncertainty that plagues the developing cobia industry in the net energetic cost of swimming in fishes. J. Fish Biol. 42 (2), 169–183.
the Americas. Increased knowledge and experience on the various Briggs, J.C., 1960. Fishes of world-wide (circumtropical) distribution. Copeia 3, 171–180.
factors that influence cobia growth rates are required so that uncer- Caylor, R.E., Biesot, P.M., Franks, J.S., 1994. Culture of cobia (Rachycentron canadum):
cryopreservation of sperm and induced spawning. Aquaculture 125, 81–92.
tainties can be eliminated and production planning rationalized. Chambers, M.D., Ostrowski, A.C., 1999. Development of bluefin trevally (Caranx
melampygus) and greater amberjack (Seriola dumerili) for offshore aquaculture. In:
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This research was funded by the National Oceanic and Atmospheric
intake and growth of juvenile Arctic charr, Salvelinus alpinus L., subjected to
Administration under grants No. NA060AR4170238 from the NOAA sustained exercise. Can. J. Zool. 68 (10), 2185–2191.
National Marine Aquaculture Initiative and No. NA08OAR4170826 of the Collette, B.B., 1999. Rachycentridae. The Living Marine Resources of the Western
NOAA Marine Aquaculture Program. The research was also supported by a Central Pacific. : In: Carpenter, K.E., Niem, V.H. (Eds.), Bony Fishes Part 2 (Mugilidae
to Carangidae), vol. 4. FAO, Rome, Italy.
previous grant from the Florida Sea Grant College Program No. RLRA42. Collins, J.F. IV, 2006. Environmental monitoring report, Cape Eleuthera offshore
The authors are indebted to the technical staffs and divers of Snapperfarm, aquaculture demonstration project. M.A. thesis, University of Miami, Rosenstiel
Inc. of Culebra, Puerto Rico and the Island School/Cape Eleuthera Institute School of Marine and Atmospheric Science, Miami, Florida, U.S.A., 56 pp.
Cushing, D.H., 1995. Population Production and Regulation in the Sea: A Fisheries
in South Eleuthera, the Bahamas, for their diligent assistance in feeding, Perspective. University Press, Cambridge, Great Britain.
data collection and daily diving at the respective cage sites. The authors Darracot, A., 1977. Availability, morphometrics, feeding and breeding activity in a multi-
acknowledge and are grateful to Mr. Johann Scheidt of Aquasense LLC for species, demersal fish stock of the western Indian Ocean. J. Fish. Biol. 10 (1), 1–16.
Denlinger, B.I., 2007. Testing aquaculture performance of juvenile cobia Rachycentron
his support and involvement in the demonstration project in South canadum, using diets containing different percentages of protein and fat. M.A. Thesis,
Eleuthera, the Bahamas. We also thank Jorge Alarcón and Owen Stevens of Division of Marine Affairs and Policy. University of Miami, RSMAS, August 2007. 17 pp.
Marine Farms Belize, Marcell Carvalho of TWB Brazil, and Thiago Soligo of Ditty, J.G., Shaw, R.F., 1992. Larval development, distribution, and ecology of cobia
Rachycentron canadum (family: Rachycentridae) in the northern Gulf of Mexico.
the Cape Eleuthera Institute for sharing information about cobia growout. Fish. Bull. 90 (4), 668–677.
Finally, the authors would like to thank Dr. Gavin Partridge from the DuPaul, W.D., Oesterling, M.J., Rose, L.A., 1997. Marine finfish aquaculture in Virginia.
Challenger TAFE Aquaculture Development Unit in Fremantle, Western Species profiles: cobia (Rachycentron canadum). Virginia Marine Research Report
No. 97-4. Virginia Institute of Marine Science, College of William and Mary,
Australia, for his review and contributions to the manuscript while on
Gloucester Point, VA 23062, pp. 23–27.
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held on the culture of Red Drum and Other Warm Water Fishes. Sea Grant Prog., Brian O'Hanlon: Brian O'Hanlon is the founder of
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Ikenoue, H., Kafuku, T., 1992. Modern Methods of Aquaculture in Japan, 2nd. ed. Elsevier fully farm cobia using offshore aquaculture technology. He
Scientific Publishing Company, Amsterdam, The Netherlands. has worked with research institutions such as the
Imsland, A.K., Foss, A., Keodik, R., Folkvord, A., Stefansson, S.O., Jonasses, T.M., 2007. University of Miami since 2003 to develop open ocean
Persistent growth effects of temperature and photoperiod in Atlantic Cod Gadus fish husbandry techniques that are environmentally safe
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Methods of Aquaculture in Japan. Developments in Aquaculture and Fisheries Science that has recently begun operations in the Republic of
vol. 11. Kodansha Ltd., Tokyo and Elsevier, Amsterdam, Holland, pp. 118–129. Panama. His work has been featured on National Geo-
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35, 12–14. general, industry specific, and scientific media.
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culture. In: Stickney, R. (Ed.), Encyclopedia of Aquaculture. John Wyley and Sons, Puerto Rico at Mayagüez. He has worked in different
Inc., New York, pp. 408–416. research positions for the last 36 years as a biologist. His
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problems. World Aquac. 38 (3), 7–13. Dr. Benetti and Mr. O'Hanlon at a GCFI meeting in Key West,
Kramer, D., Zweifel, J.R., 1970. Growth of Anchovy Larvae (Engraulis mordax Girard) in Florida who were in search for a location to start their
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Taiwan: current status and problems. Aquaculture 237 (1–4), 155–165. Rico be considered as a potential site, which was accepted
Matsuoka, S., 1995. A review of the nursery and growout culture techniques for and eventually became the grow-out site from which some
flounder (Paralichthys olivaceus) in Japan. Culture of High-Value Marine Finfish in of the data reported in this paper were gathered.
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Aquaculture. Research, Development and Commercial Production. Asian Fisheries Aaron Welch: Aaron Welch graduated from the Univer-
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Richards, C.E., 1967. Age and growth of cobia, Rachycentron canadum, from Chesapeake Miami's Marine Aquaculture Program where he is currently
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Schmidt-Nielsen, K., 1983. Animal Physiology: Adaptation and Environment, 5th ed. assistant. Aaron has been a part of the University of Miami's
University Press, Cambridge, Great Britain. cobia research team and is currently working to develop
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culture. In: Liao, I.C., Lin, C.K. (Eds.), Cage Aquaculture in Asia: Proceedings of the First
international Symposium on Cage Aquaculture in Asia, Asian Fisheries Society, Manila, Chris Maxey: Chris Maxey graduated from Yale University
and World Aquaculture Society — Southeast Asian Chapter, Bangkok, pp. 97–106. and then served for six years as an officer in the US Navy
Sun, L., Chen, H., Huang, L., Wang, Z., Yan, Y., 2006. Growth and energy budget of SEAL community. Chris founded The Island School in 1999
juvenile cobia Rachycentron canadum relative to ration. Aquaculture 257, 214–220. while serving as a teacher at Lawrenceville School. While
Svasan, T., Ottera, H.M., Tanger, G.I., 2004. Atlantic Cod. The Status and Perspectives for at Lawrenceville he received a Joukowsky Fellowship and
the Species. In: Mokness, E., Kjorsvik, E., Olsen, Y. (Eds.), Culture of Cold-Water he holds an M.A. in Marine Studies from the University of
Marine Fish. Fishing News Books, Blackwell Publishing, Oxford, UK. Miami Rosenstiel School of Marine and Atmospheric
Ticina, V., Katavic, I., Grubisic, L., 2007. Growth indicies of small northern bluefin tuna Sciences. He is the director of the Cape Eleuthera Founda-
(Thunnus thynnus, L.) in growth-out rearing cages. Aquaculture 269 (1–4), 538–543. tion, and in addition to supporting the Island School, he is
Tucker, J.W., 1998. Marine Fish Culture. Kluwer Academic Publishers, Norwell responsible for overseeing the Deep Creek Middle School,
Massachusetts, USA. the Cape Eleuthera Institute, and Cape Systems Limited.
Watanabe, W.O., Benetti, D.D., Feeley, M.W., Davis, A.D., Phelps, R.P., 2005. Status of
artificial propagation of Mutton, Yellowtail, and Red Snapper (family Lutjanidae) in
the Southeastern United States. In: Kelly, A.M., Silverstein, J. (Eds.), American
Fisheries Society Symposium 46: Aquaculture in the 21st Century, Phoenix, AZ
(USA), Aug 22 2001, pp. 517–540. M. Refik Orhun: Dr. Orhun studied biological oceanogra-
Williams, K.C., Barlow, C.G., Rodgers, L., McMeniman, N., Johnston, W., 2000. High phy and zoology at the Universität Kiel, Germany. He
performance grow-out pellet diets for cage culture of Barramundi, (Asian Sea Bass) received his M.S. in Ecology from San Diego State
Lates calcifer. In: Liao, I.C., Lin, C.K. (Eds.), Cage Aquaculture in Asia: Proceedings of the First University in 1989 and completed his Ph.D. in Marine
International Symposium on Cage Aquaculture in Asia. Asian Fisheries Society, Manila, Biology and Fisheries from the University of Miami in
and World Aquaculture Society — Southeast Asian Chapter, Bangkok, pp. 175–191. 2001. Between 2003 and 2007 Dr. Orhun was an associate
Wright, H., 2008. Japan's Kinki first again. Fish Farming International (FFI) May 2008, scientist in the University of Miami Aquaculture Program
pp. 22–23. and Director of the University's Marine Fish Hatchery. He
Yeh, S., 2000. Cobia culture in Taiwan (Rachycentron canadum). The Advocate 3 (2), 67–68. is currently working for NOAA in commercial fishery
Yeh, S.P., Yang, T., Chu, T.W., 1998. Marine fish seed industry in Taiwan. Workshop on statistics and marine aquaculture at the NOAA Southeast
offshore technologies for aquaculture. 13–15 October 1998, Haifa. Fisheries Science Center.

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.

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