Aquaculture 265 (2007) 176 186

www.elsevier.com/locate/aqua-online

The effect of fish stocking density on the growth of California halibut

(Paralichthys californicus) juveniles
German E. Merino a,b,, Raul H. Piedrahita c , Douglas E. Conklin d
a
Departamento de Acuicultura, Facultad de Ciencias del Mar, Universidad Catolica del Norte, Larrondo 1281, Coquimbo, Chile
b
Centro de Estudios Avanzados en Zonas Aridas (CEAZA), Unidad Regional de Desarrollo Cientfico & Tecnolgico (CONICYT),
Colina El Pino s/n, La Serena, Chile
c
Biological and Agricultural Engineering Department, University of California, Davis, California, USA
d
Department of Animal Science, University of California, Davis, California, USA

Received 27 July 2006; received in revised form 12 January 2007; accepted 17 January 2007

Abstract

The ability to raise California halibut (Paralichthys californicus) at a relatively high density, thus maximizing culture area
usage, is of particular importance for future commercial operations. The effect of stocking densities between 100 and 300% of
coverage area (PCA) on growth of juvenile (11.6 g) fish was tested in experiments that lasted 8 to 10 weeks. The experiments were
carried out in a recirculation system in small rectangular tanks and raceways with water depths under 6 cm. It was found that
maximum fish growth was achieved for fish stocked initially at 100% PCA. The results showed that California halibut can be
grown in shallow tanks and raceways at a relatively high stocking density without significantly compromising growth rate and
survivability.
2007 Elsevier B.V. All rights reserved.

Keywords: Stocking density; California halibut; Paralichthys californicus; Surface area; Culture density; Flatfish

1. Introduction rainbow trout (Oncorhynchus mykiss) (Trzebioatowski

The ability to raise fish at a relatively high density, thus
maximizing usage of the fish production infrastructure, is
of importance to commercial aquaculture especially in
recirculation systems. Both positive and negative relation-
ships between stocking density and growth have been
reported and the pattern of this interaction appears to be
species specific. The importance of stocking density on
fish growth has been reported for several species, such us
Corresponding author. Departamento de Acuicultura, Facultad de
Ciencias del Mar, Universidad Catolica del Norte, Larrondo 1281,
Coquimbo, Chile. Tel.: +56 51 209 767; fax: +56 51 209 764.
E-mail address: gmerino@ucn.cl (G.E. Merino).
0044-8486/$ - see front matter 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.aquaculture.2007.01.028
et al., 1981), Nile tilapia (Oreochromis niloticus) (Carro-
Anzalota and McGuinty, 1986), hamoor (Epinephelus
tauvina) (Abdullah et al., 1987), Arctic charr (Salvelinus
alpinus) (Christiansen et al., 1992), African catfish
(Clarias gariepinus) (Kaiser et al., 1995), gilthead sea
bream (Sparus aurata) (Canario et al., 1998), and lake
sturgeon (Acipenser fulvescens) (Fajfer et al., 1999).
There are a few studies on the effect of stocking densities
on the growth of flatfish: turbot (Scophthalmus maximus)
(Martinez-Tapia and Fernandez-Pato, 1991; Irwin et al.,
1999, 2003), olive flounder (Paralichthys olivaceus)
(Jeon et al., 1993), Atlantic halibut (Hyppoglossus
hyppoglossus) (Bjrnsson, 1994), and summer flounder
(Paralichthys dentatus) (King et al., 1998).
 G.E. Merino et al. / Aquaculture 265 (2007) 176186
Flatfish species are bottom dwelling fish and to
enhance growth and survival it has been recommended
to provide an environment with a high bottom area to
water volume ratio (Cripps and Poxton, 1992; Kaiser
et al., 1995; iestad, 1999; Labatut and Olivares, 2004).
Stocking density for flatfish may be expressed either as
biomass per unit of bottom area or as a percentage of
coverage area (PCA = percent ratio of total fish ventral
area to total tank bottom area). Commercial land-based
facilities usually stock their fish at densities as high as
40 kg/m2 for olive flounder (Honda, 1998), 70 kg/m2 for
sole (Liewes, 1984), and 120 kg/m2 for turbot (Person
Le-Ruyet et al., 1991). Bjrnsson (1994) suggested that
the optimal stocking density for 2 kg Atlantic halibut is
between 25 and 50 kg/m2 and between 50 and 100 kg/m2
for 10 kg Atlantic halibut. However, the normal
operational range for flatfish species varies from 15 to
50 kg/m2 depending on rearing conditions (Jones, 1981;
Person Le-Ruyet et al., 1991; Liewes, 1984; Martinez-
Tapia and Fernandez-Pato, 1991; Mallekh et al., 1998;
Silva and Velez, 1998; Kikuchi, 2000). The suggested
safe stocking density, as percentage of coverage area,
for flatfish is between 100 and 200% coverage area (Jeon
et al., 1993; Bjrnsson, 1994; King et al., 1998).
California halibut (Paralichthys californicus) is a
gregarious species in the wild and when stocked in
tanks, therefore high stocking densities might be
possible for a commercial aquaculture system. To date,
no studies have been published that examine stocking
density for California halibut. The ability to raise
California halibut at a relatively high density, thus
maximizing culture area usage, is of particular impor-
Fig. 1. Schematic diagram of an experimental tank used for the density experiment. All measurements are in cm. Each slot to introduce nets or weirs is
0.6 cm wide (measurement not shown). Figure is not to scale. Large arrow indicates the direction of the water flow. Water depth was 6.0 cm (not
shown). Tanks were built from dark grey acrylic plate, 0.6 cm thick.
177
tance for future commercial operations. Consequently,
this research was planned to estimate the effect that
stocking density may have on the growth rate of
California halibut juveniles.
2. Materials and methods
A study was carried out to test the impact of three
stocking densities nominally 100, 200, and 300%
PCA on fish performance. Stocking densities were
tested in triplicate in small tanks also referred to as
experimental tanks (Fig. 1), attached to the California
Halibut Recirculating Hatchery (Conklin et al., 2003;
Merino, 2004). Fish morphometrics (total length, TL;
total surface area with fins, TSA; body surface area
without fins, BSA) were determined at the beginning of
the experiment and at Week 8 by image analysis
(Merino, 2004). Fish average mass (W, g) was measured
at the beginning of the experiment and every other week
thereafter. The stocking density experiment lasted
10 weeks.
Three of the raceways (Fig. 2) within the California
Halibut Recirculating Hatchery (Conklin et al., 2003;
Merino, 2004) were stocked at nominally 100, 200,
300% PCA as were the experimental tanks. The purpose
was to validate the use of the experimental tanks by
comparing W for the fish population from raceways and
experimental tanks. Stocking densities were not repli-
cated for the raceways. After Week 6, the raceways
stocked at 200 and 300% PCA started to have water
quality problems due to limitations in flow capacity and
the comparison was terminated.
178 G.E. Merino et al. / Aquaculture 265 (2007) 176186

Fig. 2. Schematic diagram of a raceway. Water flow was from left to right (arrow). Water depth was between 3.5 and 4.5 cm. Figure is not to scale.

2.1. Fish stock maintained at 6 cm by a weir located at the end of the

For this study 171 days post hatch (dph) weaned
California halibut grown at the UC Davis California
Halibut Recirculating Hatchery were used. The juve-
niles (11.6 g wet mass) were reared in raceways on a
recirculating seawater system with a constant tempera-
ture of 22 1 C and salinity of 30 g/L (Gadomski and
Caddell, 1991; Madon, 2002). The fish were fed daily at
1.2 % body mass with commercial dry pelleted feeds
(EWOS Alpha #2; EWOS Pacific #3; http://www.
ewos.com). Light was provided by overhead fluorescent
tubes on a 16 L: 8 D (L = light; D = dark) photoperiod
(Boeuf and Le Bail, 1999; Klokseth and iestad, 1999).
2.2. Experimental tanks
Rectangular tanks with a total internal length of
60 cm, and an effective experimental length of 41 cm,
were used for the stocking density experiment (Fig. 1).
Three tank widths, 9.3 cm, 12.3 cm, and 15.3 cm,
having a surface area of 381. 3 cm2, 504.3 cm2, and
627.3 cm2, respectively, were used to achieve the
desired stocking densities. Screens (5 mm aperture)
were placed at both ends of a tank to hold the fish within
the experimental area. The water depth in each tank was
tank (second weir in Fig. 1). The first weir was placed
with the effluent screen (Fig. 1). A quiescent zone was
created between the weirs, allowing for observation of
the amount of feed uneaten by the fish. Tanks were
flushed every day in the morning and before feeding by
removing the second weir for a few seconds. The
function of the first weir was to keep a constant water
level of 2 cm during the flushing process.
Water velocity in the experimental tanks stocked at
100% PCA was adjusted to 0.08 0.01 body lengths
per second (bl/s) to match the corresponding velocity
(0.09 bl/s) calculated for the 100% PCA raceway
(Table 1). A velocity of 0.09 bl/s corresponded to a flow
rate of about 5 L/min for an experimental tank stocked at
100% PCA, resulting in a loading rate of 21.6 L/min kg.
This loading rate for the 100% PCA was used to
determine the flow rates for the 200% (8 L/min) and
300% (9 L/min) PCA experimental tanks, resulting in
velocities of 0.16 0.01 bl/s and 0.25 0.01 bl/s for the
200 and 300% PCA, respectively. Thereafter and as the
fish grew, the experimental tank flow rates were adjusted
every other week to keep the initial relative swimming
velocities constant over time. Water velocities within
experimental tanks induced relative swimming velocities
below 1 bl/s as recommended by Merino (2004).

Table 1
Culture conditions at time of stocking for raceways stocked at nominal densities of 100%, 200%, and 300% PCA
Nominal Biomass Fish W TSA TL Initial Flow Loading rate Velocity
PCA (%) (g) number (g) (cm2) (cm) PCA (%) (L/min) (L/min kg)
cm/s bl/s a
100 2404 214 11.23 30.6 9.9 107 9 3.7 1.1 0.09
200 4579 392 11.68 31.4 10.1 201 12 2.6 1.4 0.13
300 6795 616 11.03 30.0 9.9 306 15 2.2 1.8 0.16
Raceway surface area 6105 cm2.
a
Body length per second.
 G.E. Merino et al. / Aquaculture 265 (2007) 176186 179

2.3. Raceways (Lifegard model AF2 from Aquatic Eco-Systems Inc;

http://www.aquaticeco.com/) during the light hours
The raceways used were 2.41 m long, 0.28 m wide (0900 to 2200 h) 7 days/week for 10 weeks. Total fish
with a water depth between 3.5 and 4.5 cm (Fig. 2). A mass was measured at stocking time and then every
19 cm quiescent zone was created at the effluent section. other week to adjust the amount of feed, maintaining a
The tank surface area available for fish rearing, ration of 1.2% body mass per day. Fish mortalities were
excluding the quiescent zone was 6105 cm2. Cylin- replaced at sampling time with fish that matched the
drical nets (5 mm aperture) were placed at the influent average population morphological data for the respec-
and effluent to prevent fish from escaping and to provide tive replicate.
a frame support for a dissolved oxygen sensor.
Water flow rates in the raceways were set at stocking 2.5. Fish growth in raceways
time to have 5 mg/L of dissolved oxygen at the effluent,
resulting in 9, 12, and 15 L/min for the 100%, 200%, and This experiment was set up to compare growth
300% PCA treatment, respectively (Table 1). A maximum tendencies between fish stocked within the experimental
flow of 15 L/min per raceway could be achieved in the
system. The flow rates were maintained throughout the Table 2
six week duration of the test. Water velocities within the Summary of data from California halibut reared at three stocking
densities in experimental tanks for Week 0 (Initial), Week 8 (Final),
raceways induced relative swimming velocities below
and for selected parameters on Week 10 (Final at Week 10)
1 bl/s as recommended by Merino (2004).
Nominal stocking density as PCA (%)
2.4. Morphometrics and growth in experimental tanks 100 200 300
Average SD Average SD Average SD
California halibut juveniles to be stocked in the tanks W (g)
were obtained from a population (n = 1840) with an Initial 11.8 0.4 11.1 0.2 11.2 0.4
average W of 11.6 g, determined by counting and Final 23.9a 1.0 20.9b 0.4 19.0c 0.7
weighing the fish as a group. For simplicity and to Final at Week 10 28.5a 1.6 24.4b 0.6 21.2c 0.6
TL (cm)
minimize fish handling, the fish to be stocked in the
Initial 10.4 0.03 10.3 0.1 10.3 0.1
experimental tanks were selected from the population on Final 12.9a 0.1 12.5b 0.1 12.1c 0.2
the basis of total length (TL). The target TL for Wof 11.6 g TSA (cm2/fish)
was 10.0 cm and was determined as (Merino, 2004): Initial 33.3 0.3 33.0 0.6 32.9 0.4
Final 51.2a 1.2 48.1b 0.6 44.6c 1.2
TLcm 4:79W g0:302 ; r2 0:99: 1 BSA (cm2/fish)
Initial 21.7 0.5 21.5 0.4 21.4 0.4
The TSA corresponding to an average fish of 11.6 g Final 33.3a 0.2 31.9b 0.4 29.5c 0.8
CV (%)
was 31.2 cm2 and was determined as (Merino, 2004):
Initial 12.7a 2.3 12.9a 3.3 11.9a 1.8
Final 26.4a 1.8 23.0b 1.1 26.7a 1.1
TSAcm 7:08W g0:6056 ;
2
r 0:99:
2
2 Final at Week 10 29.4a 2.6 25.8a 3.6 30.1a 2.3
PCA (%)
The number of fish (N) to be stocked in each of the Initial 106 1 209 3 311 4
experimental tanks was estimated based on a TSA of Final 163 4 302 7 421 12
31.2 cm2/fish, the culture area available (A, cm2), and Final at Week 10 165 6 311 5 425 8
the target stocking density. The three nominal stocking Increment (%) 56 49 37
BFA (kg/m2)
densities, 100, 200 and 300% PCA, were established by Initial 3.8 0.1 7.1 0.1 10.6 0.4
stocking 20, 32, and 36 individuals, respectively, into Final 7.6 0.3 13.3 0.2 17.9 0.6
each of three replicate experimental tanks per treatment Final at Week 10 9.1 0.5 15.5 0.4 20.0 0.6
(Table 2). The percentage of covered area (PCA) was Increment (%) 140 118 89
determined as: BFV (kg/m3)
Initial 62.5 1.9 117.5 1.9 176.0 5.9
TSAcm2 N 100
Final 127.2 5.5 221.4 3.9 298.2 10.7
PCA% : 3 Final at Week 10 151.6 8.4 274.1 6.2 333.5 9.8
Acm2 Increment (%) 143 133 90
A different letter in a row indicates significant differences ( p b 0.05).
The fish were fed commercial dry pelleted feeds SD = Standard deviation; Increment = [((final at week 10) (initial)) /
(EWOS 1 and 2 mm) by automatic carrousel feeders (initial)] 100.
180 G.E. Merino et al. / Aquaculture 265 (2007) 176186
tanks and fish stocked within the raceways for 6 weeks.
California halibut juveniles were taken randomly from a
population of 1580 fish with a mean W of 11.6 g and an
estimated TSA of 31.2 cm2/fish (Eq. (2)). The number
of fish to be stocked in each raceway was determined
based on the calculated average TSA of 31.2 cm2/fish,
the tank culture area available (6105 cm2), and the
nominal stocking density (100, 200, or 300% PCA).
Each group of fish was weighed and the average W was
used to calculate the corresponding TSA (Eq. (2)) and
the actual PCA (Eq. (3)) (Table 1).
Raceways were cleaned daily in the morning before
feeding. Mortalities were removed and counted daily
and were not replaced. Fish biomass was recorded every
other week to adjust the amount of feed and maintain a
ration of 1.2% body mass per day. The fish were fed
commercial dry pelleted feeds (EWOS 1 and 2 mm)
by automatic belt feeders (model BFS12A from Aquatic
Eco-Systems, Inc.) during the light hours (0900 to
2200 h) 7 days/week for 6 weeks.
2.6. Water quality measurements
Dissolved oxygen (DO) concentrations in the
effluents from the tanks were measured three times per
day. In the case of the raceways, effluent oxygen
concentrations were measured every 10 min with
dissolved oxygen sensors (Sensorex model DO6000;
http://www.sensorex.com). Temperature was monitored
with a copper-constantan thermocouple (Omega;
http://www.omega.com) placed in the system reservoir.
Data from oxygen sensors and thermocouples were
recorded with a micrologger (Model 21X, Campbell
Scientific, Inc.; http://www.campbellsci.com). TAN,
nitrite and nitrate were measured using a Hach
Odyssey spectrophotometer (model DR/2500; http://
www.hach.com). Readings of pH were taken with a
Fisher Scientific Accumet pHmeter (model 50; https://
www1.fishersci.com). Alkalinity was measured by
titration (La Motte Chemical test kit, model WAT-DR;
http://www.lamotte.com). Salinity was measured with a
YSI SCT meter (model 33; http://www.ysi.com). TAN,
nitrite, nitrate, pH, salinity, and alkalinity were moni-
tored daily at 9:00 h before fish feeding. Sodium
bicarbonate was added daily at 9:00 h and 17:00 h to
maintain alkalinity.
2.7. Fish sampling
All fish were weighed as groups every other week to
determine biomass so that feeding rations could be
adjusted. Individual morphological data (TL; TSA; and
BSA) were gathered through image analysis (Merino,
2004) only for fish within the experimental tanks at
initial stocking and at Week 8. Fish were not fed 24 h
before sampling. Feeding was resumed the day after
sampling.
Fish growth as percent gain with respect to the values
at experiment initiation was calculated based on mass
(PWG, %), total surface area (TSAG, cm2), and body
surface area (BSAG, cm2). Other values determined
were specific growth rate (SGR, %/day, Fonds et al.,
1995), feed efficiency (relative to feed offered, FE, g/g,
Forster and Ogata, 1996), coefficient of variation for the
fish within the culture tanks (mass based, CV, %),
culture density as biomass of fish per area (BFA, kg/m2)
or per water volume (BFV, kg/m3 ), and Fulton's
condition factor (K) (Innis, 1990) (Eq. (4)).
W g
K 100 4
TLcm3
2.8. Statistical analysis
The response variables analyzed were W, TL, TSA,
BSA, SGR, PWG, TSAG, BSAG, FE and CV. Response
variables were analyzed for significant differences
among treatments using a one-way complete model
analysis of variance (Dean and Voss, 1999) with SAS
statistical software, release 8.02 Level 02M0 (SAS
Institute Inc., Cary, NC; http://www.sas.com). The
stated null hypothesis (Ho) was that there is no
difference in fish response between stocking density
levels, and the corresponding alternative hypothesis (Hi)
was that at least one level is different. Significance was
analyzed using Duncan's multiple range test and
Tukey's studentized range test at a significance of
0.05 (Dean and Voss, 1999).
The three raceways stocked at densities similar to
those of the experimental tanks were used to validate the
use of the small tanks by comparing the tendencies for
growth data from raceways and experimental tanks
stocked at similar densities.
3. Results
Salinity during the experiments was 30.1 1.3 g/L,
and temperature was 22.4 0.7 C. Temperature in-
creased during Week 6 and remained at 24.1 1.0 C
until Week 9 due to a chiller failure. During Weeks 9 and
10 the temperature was lowered to 19.6 1.2 C. During
the higher temperature period fish behaved normally but
TAN concentration increased from less than 1.0 mg/L to
5.5 mg/L. Nitrite nitrogen concentration varied between
 G.E. Merino et al. / Aquaculture 265 (2007) 176186 181

Table 2 lists the growth performance of the fish reared

Fig. 3. Mean mass (W ) of California halibut juveniles reared at three
densities during 10 weeks. Vertical lines represent standard deviations.
0.2 and 0.9 mg/L. Water pH remained unchanged over
time, having an average of 7.3 0.2. Alkalinity varied
between 2 and 3 meq/L and had an average value of
2.46 0.3 meq/L.
The lowest dissolved oxygen in the influent water to
both the experimental tanks and raceways at stocking time
was 6.4 0.1 mg/L. Effluent dissolved oxygen concentra-
tions from the raceways reached 4.2 0.3, 4.4 0.4, and
4.9 0.4 mg O2/L for 300% PCA, 200% PCA, and 100%
PCA, respectively, during Week 6, when the raceway
experiment was terminated. The lowest dissolved oxygen
concentration in the effluents from the experimental tanks
was 5.8 0.2 mg/L and was observed at the end of the
experiment (Week 10) in the tanks stocked at 300% PCA.
at the three stocking densities. At stocking time the fish
were chosen to provide a uniform TSA. There were no
significant differences ( p N 0.05) between treatment
means for W, TL, TSA, BSA, and CV at the start of the
experiment. Mean W was significantly different for the
three treatments for Weeks 8 ( p b 0.05) and 10 ( p b 0.05),
with the mean W for the 100% PCA treatment being
greater than that for the 200% PCA treatment, which was
in turn greater than that for the 300% PCA (Table 2).
Significant differences ( p b 0.05) between treatments
were seen on Week 8 for TL, TSA, BSA, and CV
(Table 2). The coefficient of variation (CV) increased
significantly ( p b 0.05) from the start of the experiment
to Week 10 for the three stocking densities. However,
there was no clear trend in terms of the evolution over
time of CV differences between the three stocking
densities. In fact, although CVs were significantly
different ( p b 0.05) among stocking densities at Week
8, no differences were found at Week 10 (Table 2).
At the end of the experiment the largest fish were
those stocked originally at 100% PCA, and the smallest
those stocked at 300% PCA. The increase in fish
biomass per area (BFA) at Week 10 was 140%, 118%,
and 89% for the 100% PCA, 200% PCA, and 300%
PCA, respectively (Table 2). The increase in fish
biomass per volume (BFV) at Week 10 was 143%,
133%, and 90% for the 100% PCA, 200% PCA, and
300% PCA, respectively (Table 2). Growth in W for the
duration of the experiment was represented well by
exponential curves (Fig. 3).
Average PWG, SGR, TSAG, and BSAG were
affected significantly by stocking density (Table 3).
Significant decreases ( p b 0.05) in PWG, SGR, TSAG,
and BSAG were observed as density was increased.

3.1. Morphometrics and growth in experimental tanks

Table 3
Fish growth and feed efficiency for California halibut reared at three
A total of five fish were replaced in replicates of the stocking densities for 8 weeks in experimental tanks (mean valuestandard
300% PCA treatment, two of which died in the tanks deviation)
and the other three jumped out of the tanks. Five fish Nominal stocking density (%)
were replaced in replicates of the 200% PCA treatment, 100 200 300
one of which died in the tank and the others jumped out
Average SD Average SD Average SD
of the culture tanks. No fish had to be replaced in
replicates of the 100% PCA treatment. PWG (%) 103.5a 3.8 88.4b 3.4 69.5c 6.4
SGR (%/d) 1.18a 0.03 1.06b 0.03 0.88c 0.06
Some fish in the 300% PCA treatment tended to
TSAG (%) 53.9a 3.9 45.9b 1.4 35.5c 2.7
attach themselves to the side walls of the tanks. A total BSAG (%) 53.4a 2.3 48.2a 1.6 37.7b 2.8
of three fish with damaged tails were found in the 300% FE (g biomass/g feed) 1.22a 0.03 1.10b 0.02 0.89c 0.06
PCA treatment tanks from Weeks 2 to 10. No injured Values followed by a different letter on the same row are significantly
fish were seen in any of the 100% and 200% PCA different ( p b 0.05).
treatment tanks. SD = Standard deviation.
182 G.E. Merino et al. / Aquaculture 265 (2007) 176186

The FE was affected significantly by stocking similar and larger than for the fish stocked at 300%
density (Table 3). A significant decrease ( p b 0.05) in PCA.
FE occurred as stocking density was increased, from a
maximum of 1.22 g/g in fish initially stocked at 100% 4. Discussion
PCA to 0.89 g/g for fish stocked at 300% PCA (Table 3).
The condition factor, K, at stocking time for all PCAs Water velocities within the tanks and raceways
was 1.0. At Week 8, K was 1.11, 1.06 and 1.08 for 100, induced swimming velocities which were less than
200, and 300% PCA, respectively. 1.0 bl/s. Merino (2004) found that California halibut
juveniles grew faster and made more efficient use of the
3.2. Fish growth comparison between raceways and feed provided at relative swimming velocities of 0.5 and
experimental tanks 1.0 bl/s than at 1.5 bl/s. Bengston and Alves (2002)
found that summer flounder (124 g, 257 g, and 387 g),
This experiment was terminated on Week 6 due to grew best when reared at 0.5 bl/s in comparison to
low oxygen concentrations in the raceways stocked at those grown at relative swimming velocities less than
200 and 300% PCA. Growth data for fish reared at three 0.3 bl/s or greater than 1.3 bl/s. Ogata and Oku (2000)
stocking densities in the raceways and experimental found that water flow velocities corresponding to a
tanks are shown in Table 4. relative swimming velocity beyond 2.1 bl/s (19.5 to
Fish were stocked at about the same BFA in both 23.5 C) led to growth retardation and lower feed
vessel types but water depths were lower in the efficiency for all olive flounder sizes tested and reported
raceways resulting in higher volumetric densities in that the optimal water flow velocity for growth occurred
comparison to the experimental tanks (Table 4). The at about 1.0 bl/s for a 12.5 cm/body length (5.7 g/fish).
overall tendency for W at Week 6 was to decrease with Therefore, there is enough evidence to support that the
stocking density with the exception of fish stocked in swimming velocities used in these experiments do not
raceways at 100 and 200% PCA for which the W were affect the growth performance of California halibut
juveniles.
Table 4 All water quality parameters were within values
Growth summary and observed volumetric density for California recommended in the literature for other flatfish species.
halibut juveniles reared during six weeks at three stocking densities
In the case of TAN, the worst condition measured was
within two types of culture vessels
5.5 mg TAN/L (salinity 30.1 g/L; 24.1 C; pH 7.3)
Nominal stocking density (%) which is equivalent to about 0.05 mg/L un-ionized
100 200 300 ammonia as nitrogen (NH3N), Juvenile turbot ( 20 g)
Average SD Average SD Average SD reduced their food intake when un-ionized ammonia
Raceways was over 0.117 mg NH3N/L, and a reduction in body
W (g) mass gain per day occurred when the level was over
Initial 11.2 11.7 11.0 0.108 mg NH3N/L (pH 8, 16 C, 28 g/L salinity)
Final 20.6 20.5 18.2
(Rasmussen and Korsgaard, 1996). Person-Le Ruyet
BFA (kg/m2)
et al. (1997) reported that for 13, 23, and 104 g turbot
Initial 3.9 7.5 11.1
Final 7.1 13.1 17.5 reared in water having in the effluent over 80% oxygen
BFV (kg/m3) saturation (6 mg/L), the growth was not affected at
Initial 101 165 228 concentrations of 0.21, 0.18, and 0.09 mg NH3N/L,
Final 181 286 359
while growth stopped immediately for all groups above
0.8 mg NH3N/L ( pH 8, 17 C, 34.5 g/L salinity).
Experimental tanks
W (g) Turbot survival was not affected at levels below 0.4 mg
Initial 11.8 0.4 11.1 0.2 11.2 0.4 NH3N/L (13 mg TAN/L) (Person-Le Ruyet et al.,
Final 20.7 0.9 18.3 0.4 17.0 0.7 1997). Therefore the worst concentration of un-ionized
BFA (kg/m2) ammonia as nitrogen recorded in this experiment is
Initial 3.7 0.1 7.1 0.1 10.6 0.4
within a safe range. In the case of dissolved oxygen, the
Final 5.6 0.2 9.8 0.4 14.2 0.4
BFV (kg/m3) stocking density trial carried out in the raceways was
Initial 62.5 1.9 117.5 1.9 176.0 5.9 terminated at the end of Week 6 when dissolved oxygen
Final 109.8 4.7 193.9 4.5 267.6 10.3 in the raceway effluent dropped below 60% of oxygen
Tank depth was 6 cm and raceway depth was 3.64.5 cm. saturation, the value recommended as an allowable limit
There was no SD for raceways as they were not replicated. for turbot (Brown et al., 1984). Effluent dissolved
 G.E. Merino et al. / Aquaculture 265 (2007) 176186
oxygen in the experimental tanks remained above 75%
of saturation throughout the experiment.
Fish stocked in the experimental tanks at 200% and
300% PCA did not eat all the feed offered between
Weeks 8 and 10. The increase in pellets left in the
quiescent zones of the tanks loaded at 200% and 300%
PCA started when the stocking density reached 283%
PCA and 397% PCA, respectively. California halibut
reared in the experimental tanks had a drop in SGR
during the last 2 weeks of experimentation, regardless of
the tank stocking density. A reduction of feeding activity
and a drop in SGR can indicate a possible stressful
condition.
The culture density of flatfish is frequently reported
as the PCA (Bjrnsson, 1994; Klokseth and iestad,
1999). A common practice to estimate surface area of
flatfish in aquaculture is by placing the fish over a paper
grid, drawing the fish edges, and counting the area
enclosed by the drawing (Bjrnsson, 1994; King et al.,
1998; Fairchild and Howell, 2001). The limits of this
current methodology are numerous. Foremost is that
since the process is labor intensive and time consuming
only a few animals can be sampled. Sampled fish are
subjected to excessive manipulation, with a high
potential for injury. Merino (2004) used photographic
and computer image analysis techniques to perform an
accurate and fast method to estimate surface area of
flatfish which he called total surface area (TSA) (area of
fish edges) as well as body surface area (BSA)
(excluding the fins) of California halibut.
Fulton's condition factor (K ) is an indicator of
general fish well-being. California halibut in the present
study had a K value between 1.06 and 1.11, which
according to Fulton's condition definition will be
considered as a healthy fish.
4.1. Morphometrics and growth in experimental tanks
Morphometric analysis was planned originally for a
period of 10 weeks, but a mechanical problem with the
chiller resulted in an increase in the water temperature
from 22 to 24 C during Week 6. Therefore, it was
decided to do a morphometric analysis at the next
scheduled sampling (beginning of Week 8). Madon
(2002) who experimented with California halibut
juveniles of similar sizes to the ones used in this
research did not detect differences in growth for
temperatures ranging from 20 to 28 C.
In the experiments described here, fish stocked at
100% PCA (final 163% PCA; 127.2 kg/m3) grew larger
than those stocked at 200% PCA (final 302% PCA;
221.4 kg/m3 ) and 300% PCA (final 421% PCA;
183
298.2 kg/m3) (Table 2). Fish held at the lowest densities
had the largest SGR (Table 3). Most studies on flatfish
stocking densities have found that higher densities result
in lower growth. Irwin et al. (1999) reported that turbot,
with initial biomass of 8.62 g/fish, were stocked at four
different densities (10%, 14%, 17%, and 21% PCA),
and after 45 days fish reared at the higher stocking
densities (final density 149.3% PCA) showed signifi-
cantly slower growth rates than those held at lower
densities (final density 67.4% PCA). Jeon et al. (1993)
reported that olive flounder with initial biomass of 42 g/
fish were stocked at five different densities (33%, 50%,
100%, 200%, and 300% PCA), and after 35 days of
experimentation it was found that fish at the highest
stocking density (final density 460% PCA) showed a
significantly slower growth rate than those held at the
lowest density (final density 45% PCA). King et al.
(1998) summer flounder with initial biomass of 7.8 g/
fish, were stocked at three different densities (100%,
150%, and 200% PCA), and after 58 days the fish reared
at the highest stocking density (final density 493%
PCA) showed significantly slower growth rates than
those held at the lowest density (final density 355%
PCA).
Feed efficiency was affected when California halibut
were stocked at higher densities as has been described
for olive flounder (Jeon et al., 1993). In the current
study, a considerable amount of feed was found in the
quiescent zone of tanks stocked at 200% and 300% PCA
starting on Week 8. There are reports that with rising
stocking density, an increase in aggressive behavior,
growth, and feed efficiency was observed for the
dominant fish while stress, fin damage, and metabolic
rates increased in subordinate fish (Refstie and Kittel-
sen, 1976; Purser and Hart, 1991; Canario et al., 1998).
A study done with turbot did not find differences on
growth nor aggressive behavior between equally sized
or mixed animals (Sunde et al., 1998). Fish stocked in
the present study were pre-selected by size and only one
case of damaged tail was observed in each of the 300%
PCA replicates. Bjrnsson (1994) found that for
Atlantic halibut, more of the feed consumed was used
for metabolism than for growth with increasing stocking
densities. Thus it is possible that an increase in
metabolic rate, which was not measured, at higher
stocking densities might have been the cause for the feed
efficiency and growth reduction found here with
California halibut.
A direct effect of stocking density on growth
variability has been reported by a number of authors
(Andrews et al., 1971; Irwin et al., 1999; Lambert and
Dutil, 2001). The relative size difference between
184 G.E. Merino et al. / Aquaculture 265 (2007) 176186
members of a population usually increases with
increasing stocking density due to social interactions,
development of hierarchies, and establishment of
territorial borders (Lambert and Dutil, 2001). Further-
more, an increase in the CV for fish within a population
is considered to be indicative of the establishment of
hierarchies (Jobling, 1994; Irwin et al., 1999; Lambert
and Dutil, 2001). The CV for turbot was found to
increase more rapidly in groups held at higher stocking
densities (Irwin et al., 1999). It was also found that the
CV for turbot increased with increasing size of fish both
for graded and ungraded groups (Sunde et al., 1998). In
this study, the CV increased significantly for all groups
of California halibut during the 10 weeks of experi-
mentation, indicating that growth rates were affected at
all stocking densities analyzed. However, there was no
clear trend in terms of the evolution over time of CV
differences among stocking densities (Table 2).
4.2. Fish growth comparison between raceways and
experimental tanks
Experimental tanks were used in triplicate to set up a
density experiment and three raceways were stocked at
the same densities (100%, 200%, and 300% PCA) and
used as controls against the experimental tanks. Results
for the experimental tanks agreed well with those from
the raceways (Table 4). Raceways, which were 5.6 times
larger, 23 times wider, and stocked with 9 to 16 times
more biomass than experimental tanks had growth
results similar to those in the tanks.
Because of the shallowness of the water in the tanks
and raceways used, BFV in this research (Table 4) are
larger than the 75 kg/m3 maximum reported for turbot
(Irwin et al., 1999; Martinez-Tapia and Fernandez-Pato,
1991). In addition BFV and BFA in the current research
were also larger than the 200.8 kg/m3 and 8.03 kg/m2,
respectively, reported recently by Labatut and Olivares
(2004) for 7.63 g turbot reared at 262 PCA% in shallow
raceways with a rearing area of 36,000 cm2 and a
velocity of 0.64 bl/s. Areal densities reported for conven-
tional aquaculture vessels are up to 29.5 kg/m2 for
juvenile summer flounder (King et al., 1998), 36.3 kg/m2
for olive flounder (Chang et al., 1995), 68 kg/m2 for turbot
(Martinez-Tapia and Fernandez-Pato, 1991), and 95 kg/
m2 for halibut (Bjrnsson, 1994) which suggests that
flatfish can be raised at high densities.
4.3. Practical implications
The practical implications of the present results are
that California halibut can be grown in shallow rect-
angular tanks and raceways at a relatively high stocking
density without significantly compromising survivabil-
ity. In this study, the number of fish was held constant
for the duration of the experiment, and biomass density
(BFA and BFV) was allowed to increase with time due
to the growth of the fish. Initial stocking densities of
100% PCA produced a higher individual fish growth
over time than the other stocking densities tested, which
indicates that stocking density does affect the growth of
California halibut juveniles. The results indicate that, of
the densities tested, the optimal one is somewhere
between 100% and 200% PCA. Therefore, tanks with
California halibut juveniles should be split when the
stocking density reaches a maximum target PCA to
reduce the BFA and prevent a significant reduction in
SGR.
Volumetric biomass densities for California halibut
juveniles can be relatively high. In this study, BFV
reached values between 151.6 and 333.5 kg/m3, which are
larger than those reported for other flatfish (Irwin et al.,
1999; Iglesias et al., 1987; Jones et al., 1981; Labatut and
Olivares, 2004). The high BFV reached in this study were
the result of rearing the fish in shallow tanks.
In this study relatively low BFA of 9.1 and 20.0 kg/m2
were tested due to the small size of the fish used, but
work with olive flounder (Honda et al., 1993; Honda,
1998), turbot (Person Le-Ruyet et al., 1991), and
Atlantic halibut (Bjrnsson, 1994) suggest that values
of 48 (for a 120 g fish), 25 to 50 kg/m2 (for fish N 300 g),
and 25 to 50 (for a 2 kg fish), respectively, are possible.
The BFA can be estimated by knowing W, the
corresponding TSA, and the PCA. The TSA can be
measured for flatfish by using image analysis (Merino,
2004). For California halibut, TSA can be estimated
using Eq. (2) (Merino, 2004). Having W and TSA, an
Fig. 4. Projected stocking density of California halibut corresponding
to 100%, 200%, and 300% PCA related to biomass of fish.
 G.E. Merino et al. / Aquaculture 265 (2007) 176186
equation can be written that relates fish biomass (W )
with stocking density:
 
W g
BFA a k 5
TSAcm2
where is the stocking factor and is equal 1 for 100%
PCA, 2 for 200% PCA, and so on; and k is a conversion
factor (10) to express g/cm2 as kg/m2. Increases in W will
result in the corresponding rises in BFA for a given PCA
as shown in Fig. 4. The graph does not represent growth,
merely the calculated biomass of fish stocked per surface
area (BFA) for a given fish size (W ) and desired PCA. For
example it would take 10 kg of 200 g California halibut
(50 fish) for a 100% PCA or 20 kg of 1000 g California
halibut (20 fish) for a 100% PCA. Thus one layer of small
fish will weigh less than one layer of large fish.
5. Conclusions
Growth of California halibut decreased as stocking
density was increased from 100% to 300% PCA.
Maximum SGRs were obtained when California halibut
juveniles were initially stocked at 100% PCA.
The results of this study could be used to plan yield-
maximizing farming strategies for juvenile California
halibut. Results from this study are appropriate for the
size of fish tested. Requirements and growth of
California halibut may change for fish of different
sizes. Further observations and research will refine the
understanding of optimum stocking density for other
California halibut sizes.
Ultimately, the optimal stocking density from the
point of view of a fish farmer will depend on capital cost
per unit of rearing area, availability and cost of
juveniles, fish growth, market price and size of fish,
among other factors, to maximize the biomass and size
of fish produced for a target market.
Acknowledgements
The fish used in this work were supplied as eggs or
larvae by the California Halibut Hatchery Program in
Redondo Beach, California. The assistance of Jean Benoit
Muguet, Doug Bush, Dr. Margarita Cervantes-Trujano,
and Dr. Enric Gisbert is gratefully acknowledged.
This work was primarily funded by the National Sea
Grant College Program of the U.S. Department of
Commerce's National Oceanic and Atmospheric Ad-
ministration under NOAA Grant no. NA06RG0142,
project number R/A-116, through the California Sea
Grant College Program, and in part by the California
185
State Resources Agency. The views expressed herein are
those of the authors and do not necessarily reflect the
views of NOAA or any of its sub-agencies. The U.S.
Government is authorized to reproduce and distribute
for governmental purposes.
Other sponsors were the Chilean Government
through the Beca Presidente de la Republica, the
Organization of American States (OAS) through a
fellowship under the Regular Training Program (PRA),
and the University of California at Davis through a
Jastro-Shields Research Scholarship.
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