Aquaculture 219 (2003) 471 – 483

www.elsevier.com/locate/aqua-online

Culture experiments with Oithona oculata Farran,

1913 (Copepoda: Cyclopoida), and its advantages as
food for marine fish larvae
O.G. Hernández Molejón a, L. Alvarez-Lajonchère b,*
a
Empresa Pesquera Industrial PESCASILDA, Sancti Spiritus, Cuba
b
Delegación Provincial de Ciudad de La Habana, Ministerio de Ciencia, Tecnologı́a y Medio Ambiente,
Avenida 17 No. 5008, Playa, Ciudad de La Habana C.P. 11300, Cuba
Received 27 February 2002; received in revised form 25 September 2002; accepted 19 November 2002

Abstract

Culture experiments were made with Oithona oculata Farran, 1913, a coastal water copepod, at
Tunas de Zaza and Santa Cruz del Sur, Cuba. A 1 1/2-year semicontinuous experimental-scale
polyculture with the rotifer Brachionus rotundiformis (type ‘‘S’’) in outdoor tarpaulin 2.5-m3 tanks
was established, with Nannochloropsis oculata (20  106 cells ml 1) and baker’s yeast (1 – 2 g/106
rotifers day 1) as food, reaching a mean density of 5 copepods ml 1 and a maximum density of 8
copepods ml 1, with a daily harvest of 25% of the total volume. A 1-year batch-culture pilot-scale
system was also established in indoor concrete 20-m3 tanks using N. oculata (8 – 20  106 cells
ml 1) as food, reaching a mean density of 7 copepods ml 1 and a maximum density of 10 copepods
ml 1. Copepod monocultures maintained in polycarbonate 1000-l outdoor tanks were fed a mixture
of five species of microalgae (Chaetoceros ceratosphorum, Tetraselmis tetrathele, Chlorella spp.,
Dunaliella tertiolecta and N. oculata) at an average density of 2  106 cells ml 1. In these
nonaerated cultures, final concentrations of 13 copepods ml 1 were obtained in 15-day cultures.
Recommendations for improving pelagic copepod culture are presented and their advantages as live
food for rearing marine fish larvae are analyzed.
D 2003 Elsevier Science B.V. All rights reserved.

Keywords: Copepod culture; Live food; Oithona oculata

* Corresponding author. Present address: Calle 41 No. 886 entre 24 y Avenida 26, Nuevo Vedado, Plaza,
Ciudad de La Habana C.P. 10600, Cuba. Tel.: +53-7814985.
E-mail address: lajonchere@yahoo.com (L. Alvarez-Lajonchère).

0044-8486/03/$ - see front matter D 2003 Elsevier Science B.V. All rights reserved.
doi:10.1016/S0044-8486(02)00644-0
472 O.G. Hernández Molejón, L. Alvarez-Lajonchère / Aquaculture 219 (2003) 471–483

1. Introduction

Copepods are the main food source of marine fish larvae in nature (May, 1970;
Zismann et al., 1975; McMichel and Peten, 1989). In aquaculture, copepods have been
proven to be the preferred and most adequate food for many marine fish larvae (Houde,
1973; May et al., 1974; Kraul, 1983a, 1989, 1993), and are also used for shrimp larvae and
postlarvae (Shamsudin and Saad, 1993). Good results in terms of fish larval growth and
survival using natural plankton are due to the presence of copepods and their role as the
main food component (May, 1970; Bent, 1993).
The larvae of many marine fish require prey about 50 – 100 Am wide at first feeding
(Detwyler and Houde, 1970; Yúfera and Pascual, 1984). Even the rotifer Brachionus
rotundiformis, type ‘‘S’’, is too large in many cases (Houde, 1973; May et al., 1974; Doi
and Singhagraiwan, 1993). Copepods have also been satisfactorily used after a period of
using rotifers and before introducing Artemia nauplii (Kraul, 1993; Alvarez-Lajonchère et
al., 1996), and also offered simultaneously (Leu and Chou, 1996).
Copepods offer a great variety of sizes, species and qualities (Kinne, 1977; Yúfera and
Pascual, 1984; Delbare et al., 1996), and have high levels of protein, highly unsaturated
fatty acids (HUFA), carotenoids and other essential compounds (Kraul et al., 1992).
Watanabe et al. (1983) reported that culture media did not influence the copepod chemical
composition, although Delbare et al. (1996) reported that copepod n 3 HUFA reflected
the culture diet. A positive correlation of the n 3 HUFA levels and highest nauplii
production was reported by Norsker and Støttrup (1994). Other good characteristics of
copepods are their swimming movements as a larval visual stimulus, the tank-cleaning
performance primarily by benthic harpacticoids, which are grazers (Støttrup et al., 1995),
their high digestive-enzyme content (Delbare et al., 1996) and a possible enhancement of
feeding rates with improved growth and survival (Støttrup and Norsker, 1995, 1997).
HUFA are essential for marine fish larvae (Watanabe et al., 1983; Witt et al., 1984;
Sorgeloos et al., 1988; Watanabe and Kiron, 1994; Kanazawa, 1995). Docosahexaenoic
acid (DHA; 22:6 (n 3)) has a significant influence on larval stress resistance (Kraul et al.,
1991, 1993). DHA content is higher in copepods than in recently hatched Artemia nauplii
and gives better results in terms of survival, growth and stress resistance (Fujita, 1979).
Superior larval stress resistance can be achieved with copepods, even when DHA content
is less than enriched Artemia nauplii (Kraul et al., 1992). Although Artemia enrichment
has greatly improved commercial aquaculture, the most advanced rotifer and Artemia
bioencapsulation techniques have not matched the results with copepods, and their
composition is now taken as the standard for improving enrichment techniques for rotifers
and Artemia (Kraul et al., 1988, 1992; Sorgeloos and Léger, 1992).
Most of the large-scale copepod culture systems are based on outdoor semicontrolled
polyculture techniques, although several attempts have been made to culture some species
in intensive systems (Støttrup et al., 1986; Støttrup and Norsker, 1997).
In Japan, copepods are cultured together with rotifers, and Tigriopus japonicus is the
main species (Kuronuma and Fukusho, 1984; Fukusho, 1980, 1991). Copepods of the
genus Oithona are abundant and permanent in coastal lagoons (Zacarı́as and Zoppi de
Roa, 1981). This highly prolific genus has short life cycles and wide geographic
distribution, making it a good candidate for culture and an excellent substitute for Artemia
 O.G. Hernández Molejón, L. Alvarez-Lajonchère / Aquaculture 219 (2003) 471–483 473

nauplii by their better nutritional value, especially for larvae during metamorphosis (Bryan
and Madraisau, 1977). Omori (1973) considered Oithona as an ideal candidate for culture
in Japan.

2. Materials and methods

Oithona oculata Farran, 1913, a pelagic coastal water copepod with a mean length of
65 Am for nauplius I, 100 Am for copepodite I and 625 Am for adults, was isolated from
Tunas de Zaza, Sancti Spiritus Province, Cuba, and monoculture experiments, lasting 14 –
16 days each, were done in three outdoor polycarbonate 1000-l tanks. Inocula of 2
copepods ml 1 were fed a mixture of five microalgae: Nannochloropsis oculata (75 –
80%), and similar densities of Chaetoceros ceratosphorum, Tetraselmis tetrathele,
Chlorella spp. and Dunaliella tertiolecta, to an overall algal density of 2  106 cells
ml 1. No aeration was applied. The starting volume was 150 l and was increased by 150 l
every 2 days up to 900 l.
A semicontinuous experimental-scale polyculture of the rotifer B. rotundiformis and O.
oculata, with a daily harvest of 25% of the total volume, was done at Tunas de Zaza in four
outdoor 2.5-m3 tarpaulin tanks with PVC pipe supports, 0.70-m depth, and two 0.25-m3
inocula tanks for a total culture volume of 10.5 m3. Water was pumped from near shore and
filtered through 10- and 5-Am cartridges. Tap water was added daily to restore the
evaporation of 1– 2 cm from the culture tanks. An airflow of 20 l min 1 at 0.10 bar was
supplied. The polyculture was fed N. oculata at 15 – 20  106 cells ml 1 initial density and
1 –2 g of baker’s yeast per 106 rotifers day 1. The culture cycle with partial harvest usually
lasted 20 days in each tank.
A batch-culture pilot-scale system was also established in four concrete 20-m3 (0.5 m
deep) production tanks with total harvest of one tank each day after a 4-day growth cycle.
The production tanks were inside a building with a translucent fiberglass roof, and were
inoculated with 1.3-m3 or 2.5-m3 outdoor fiberglass tanks, at a shrimp pilot-scale hatchery
at Santa Cruz del Sur, Camagüey Province. Water was directly pumped from near shore
and filtered through 5-Am cartridges, and tap water also added daily to restore the
evaporated water. Constant aeration was provided with 120 l min 1 airflow at 0.10 bar for
each production tank and 20 l min 1 m 3 airflow at 0.17 bar for each of the inoculant
tanks. These cultures were fed N. oculata at an initial density between 8 and 20  106 cells
ml 1.
Copepods were harvested with 40 Am mesh-size screens, and supplied to striped
patao Eugerres brasilianus larvae in two consecutive trials in experimental (5 m3;
Tunas de Zaza) and pilot-scale (10 m3; Santa Cruz del Sur) units. The larval-rearing
techniques at these sites have been described by Alvarez-Lajonchère et al. (1994) and
Alvarez-Lajonchère et al. (1996). In all the trials, the rotifer B. rotundiformis was
supplied from day 2 posthatch (d2ph) to d32ph at densities of 3– 10 rotifers ml 1,
and Artemia nauplii from d13ph to d54ph at densities of 2 – 8 nauplii ml 1. Rotifers
and Artemia nauplii were enriched with a cod liver oil emulsion for the experimental
trials only (Watanabe et al., 1982). In the experimental trials, copepods were supplied
daily to two 5-m3 larval-rearing tanks at a density of 0.5 copepod ml 1 starting at
474 O.G. Hernández Molejón, L. Alvarez-Lajonchère / Aquaculture 219 (2003) 471–483

d8ph until d32ph. In two pilot-scale 10-m3 larval-rearing tanks, the density of
copepods was increased to 1– 3 copepods ml 1 starting on d2ph and the supply
of Artemia nauplii was delayed until day 18 posthatch. A third pilot-scale 10-m3
larval-rearing tank was used as a control with rotifers as the sole diet during two
rearing cycles. Stomach contents of larvae were examined daily during the first trial
only.
In each of the experiments temperature, salinity and oxygen were monitored
twice daily (0800 and 1600 h), and in the pilot-scale trials, pH was measured once
every 2 days. Correlation coefficient significance tests between the environmental
parameters and rotifer and copepod harvest densities were carried out (Sokal and
Rohlf, 1981).

3. Results

The semicontinuous experimental system worked in 15 –20-day cycles carried out

during 1 1/2 years at Tunas de Zaza with mean temperatures of 23.3 F 2 jC in the
winter dry season and 31.2 F 3 jC in the wet summer season, with a minimum of
20.0 jC and a maximum of 39.5 jC in the whole period. The salinity ranged
between 37 and 41 ppt (38 F 1 ppt) in the outdoor culture tanks, and dissolved
oxygen was always between 5.5 and 8.9 mg l 1. Rotifer harvest densities were
between 50 and 120 rotifers ml 1, averaging 92 F 8 rotifers ml 1 in summer and
65 F 6 rotifers ml 1 in winter. Rotifer harvest densities were significantly and
positively correlated ( P < 0.01) with temperature. Copepods reached between 1 and
8 copepod ml 1 with a mean of 5 copepods ml 1, averaging 5.8 F 1.5 copepods
ml 1 in summer and 3.1 F 1 copepods ml 1 in winter, but not significantly
correlated ( P>0.05) with temperature. On several occasions, a sharp decrease in
harvest density of both culture organisms occurred together with a decrease in water
temperature to 20 jC. Microalgae depletion correlated with lowered rotifer densities
and increased copepod densities. Baker’s yeast induced ciliate proliferation and
coincided with rotifer density decreases.
The pilot-scale system worked with 4-day batch culture cycles during 1 year at
Santa Cruz del Sur. Water quality problems arose due to operational difficulties with
the cartridge filters, as well as failure with the tap water used to reduce the high
salinity levels of 39 F 2 ppt (35 – 42 ppt). Dissolved oxygen was between 4.6 and 8.7
mg l 1, pH, between 6.5 and 8.9, and temperature range was 23 –32 jC with mean
temperatures of 24.5 F 1.5 jC in winter and 32.3 F 2 jC in summer. On one occasion,
temperature reached 17 jC, coinciding with mass mortality and culture collapse.
Average harvest densities were 7 F 2.5 copepods ml 1 (1 –10 copepods ml 1), with
8.6 F 1 copepods ml 1 in summer and 4.8 F 2 copepods ml 1 in winter, and
100 F 14 rotifers ml 1 (50 – 200 rotifers ml 1), with 166 F 9 rotifers ml 1 in summer
and 97 F 6 rotifers ml 1 in winter. Two-day average rotifer and copepod productions
during a typical winter 20-day culture cycle are presented in Fig. 1, together with the
water temperature to show the positive and significant correlation ( P < 0.01) between
temperature and rotifer production.
 O.G. Hernández Molejón, L. Alvarez-Lajonchère / Aquaculture 219 (2003) 471–483 475

Fig. 1. Daily production in terms of total numbers of rotifers (squares) and copepods (triangles) harvested from
20-m3 tanks during a culture cycle, together with average temperature (crosses). Average of 2 days.

Small-scale monoculture experiments were carried out during April to July 1997 with
an average temperature of 32 F 3 jC, salinity of 39.8 F 1 ppt and dissolved oxygen
between 5.3 and 9 mg l 1. Final densities after 15 days of culture were 10 –13 copepods
ml 1 (12.2 F 0.6 copepods ml 1).
Survival of E. brasilianus larvae in the experimental-scale tanks were 6% in rotifer-fed
larvae and 12.5% in copepod-fed larvae, with harvest densities of 5.1 and 10.2 juveniles
l 1 and the same growth of 0.41 mm day 1 at temperatures of 29 –30 jC. Survival in the
copepod-fed pilot-scale tanks was 30% and 50%, with a mean harvest density of 6.1
juveniles l 1 at d50ph and a mean growth of 0.3 mm day 1. Survival in the pilot-scale
control tank (no copepods) was 12% and 18% with a mean harvest density of 5.1 juvenile
l 1, with the same growth rate.
476 O.G. Hernández Molejón, L. Alvarez-Lajonchère / Aquaculture 219 (2003) 471–483

4. Discussion

Copepod monoculture results were satisfactory and in agreement with those of other
authors that used several species of microalgae (Zillioux, 1969; Nassogne, 1970; Girin and
Devauchelle, 1974; Iwasaki and Kamiya, 1977; Iwasaki et al., 1977; Zurlini et al., 1978).
The semicontinuous system used in the experimental-scale polyculture is the most
popular method in Japanese hatcheries (Hino, 1993). In our experiments, smaller tanks
were used with much smaller capacity than are typical of batch culture, although volumes
of 0.5– 2 m3 have been used in that system with good results (Coves et al., 1990). Small
volume and the external location caused great temperature fluctuation, which could have
negatively influenced our production densities.
The mean daily harvest densities in our semicontinuous system were similar to those
obtained in Japan since the 1980s (Kuronuma and Fukusho, 1984; Fukusho, 1991). Daily
partial harvests with culture media exchange had a positive influence, in agreement with
Nassogne (1970) and Zurlini et al. (1978). A poor culture turnover results in a low water
quality and low number of eggs per female, and organic matter sticks to appendages,
causing breakage and mortality. Clean water reduces faecal particle ingestion and provides
high oxygen levels, good pH values, constant food concentrations (Kahan et al., 1982) and
low ammonia – nitrogen levels (Fengqi, 1996). Zurlini et al. (1978) recommended culture
systems with large water exchanges.
There are few reports of copepod culture with N. oculata as food. It has been supplied
to T. japonicus (Kitajima, 1973; Koga, personal communication to Kinne, 1977), Acartia
clausi (Kitajima, 1973), and Acartia tsuensis (Ohno et al., 1990). Euterpina acutifrons
seems unable to digest N. oculata cell wall (Kraul, 1998, personal communication).
The microalgal densities were high in our experiments, although N. oculata is very
small, so high number of cells per milliliter were expected, as in Japan (Kuronuma
and Fukusho, 1984; Morizane, 1991). Most experimental work with copepods is
carried out with lower microalgal densities, although at lower temperatures than those
of the present study. Ohno et al. (1990) used 4– 5  104 cells ml 1 for A. tsuensis
culture, while Nassogne (1969) found the highest egg production and best feeding
efficiencies for E. acutifrons at 5  104 cells ml 1. Kraul (1989) recommended densities
between 5  104 and 2  105 cells ml 1, while Zurlini et al. (1978) had their best
results at 6  105 cells ml 1 with the same species. Kahan and Azoury (1981) used 1–
3  105 cells ml 1 as the best density for Nitocra spinipes, and Iwasaki et al. (1977)
estimated optimum concentrations for A. clausi were a 1:1 combination of Isochrysis
galbana and Monochrysis lutheri at 1  106 cells ml 1 (15 jC) or 1.5  106 cells ml 1
(20 jC).
Use of baker’s yeast promoted ciliate, bacteria and detritus proliferation, but in the
present study, there were no negative effects for copepods because these products are also
food sources for copepods (Zillioux, 1969). Copepod culture based only on bacteria has
been tried successfully (Lee and Taga, 1988). Baker’s yeast has been used with good
results in Japan for copepod culture (Omori, 1973; Fukusho, 1980; Fukusho et al., 1980).
Artificial feeds also work well (Omori, 1973; Kitajima, 1973; Kahan and Azoury, 1981).
The best survival and growth of some fish larvae were obtained with copepods reared with
N yeast (Fukusho et al., 1980), possibly by their higher nutritional value due to their high
 O.G. Hernández Molejón, L. Alvarez-Lajonchère / Aquaculture 219 (2003) 471–483 477

eicosapentaenoic acid (EPA; 20:5 (n 3)), DHA and total n 3 HUFA content (Watanabe
et al., 1983; Fukusho et al., 1980).
In our experiments, the quantity of baker’s yeast was the same as that used by other
authors who combined it with N. oculata or other microalgae (Fulks and Main, 1991). To
improve culture conditions, the yeast should be divided into two or more partial rations
during the day to lessen water quality deterioration, as recommended by Cittolin and
Guidastri (Servizi Tecnici in Maricoltura, 1992, personal communication).
Difficulties in the pilot-scale culture included fluctuations of water quality and
environmental parameters, and filtration system problems. However, results were better
with the total harvest method, which was more laborious but also more effective,
productive, and reliable than the semicontinuous method used in the experimental-scale
trials. This is in agreement with other reports on rotifers (Morizane, 1991). The batch
culture system is more effective for T. japonicus combined with B. plicatilis in Japan
(Fukusho, 1991). Greater tank volume also improved feeding and temperature stability,
especially in indoor locations (Fulks and Main, 1991). Microalgae concentrations at the
end of the third day of the batch culture system were notably lower and the copepod
reproduction was favored in accordance with Zurlini et al. (1978) and Kraul (1989).
Fengqi (1996) pointed out that copepod abundance was favored by conditions improper
for rotifers. Our results confirm other positive results of polyculture using rotifers with
other copepod species (Girin and Devauchelle, 1974) and these methods are common in
Japan (Kuronuma and Fukusho, 1984), although Kraul (1989) recommended not mixing
rotifers with E. acutifrons because rotifers will affect culture procedures for best efficiency.
Harpacticoid copepod culture reports are generally characterized by very high densities
in volumes of 15 l or less, among which those of Kahan and Azoury (1981), Kahan et al.
(1982), and the tray system of Støttrup and Norsker (1997) are the best, though their
application in commercial mass production of marine fish juveniles is impractical.
Densities around 10 copepods ml 1 reached in our monoculture experiments are
among the highest of the several hundred liter volume category, except the 20– 50 or more
copepods ml 1 obtained with E. acutifrons by Kraul (1983a, 1989). Our pilot-scale results
are similar to the highest among those of several cubic meter culture volumes with
densities between 5 and 10 copepods ml 1. These last results and those of Kraul (1983a,
1989) can be applied to feed marine fish larvae on a commercial scale.
Survival in E. brasilianus was more than double when fed with copepods than without
in experimental- and pilot-scale experiments. There was no growth difference between
rotifer- and copepod-fed larvae, although growth was slightly lower in the pilot-scale trials
compared with the experimental-scale trials. We observed improved larval fish survival in
bigger tanks, and with the presence of copepods, especially when copepods were
introduced early in the larval phase and in higher quantities. The copepod densities
supplied were within the reported range for several marine species, from 0.5 to 1.0
copepod ml 1 (Saifulina, 1991) up to 3 copepods ml 1 (Houde, 1973) and in agreement
with those recommended by Kraul (personal communication) of 1 – 2 copepods ml 1 for
densities between 1 and 10 larvae l 1. Our results are in agreement with other reports on
marine fish larvae fed with copepods (Kraul, 1993).
The sizes of O. oculata are satisfactory for larval rearing of marine fish because their
nauplii are small, and are adequate as a first food for many small-mouth species, among
478 O.G. Hernández Molejón, L. Alvarez-Lajonchère / Aquaculture 219 (2003) 471–483

which the ornamentals are abundant. The sizes of the first copepodite stages are similar to
those of rotifer type ‘‘S’’, and the size of cultivated adults are within the range reported by
Campos Hernández and Suarez Morales (1994) for the copepod species in the Gulf of
Mexico and the Caribbean Sea and may partially replace rotifer type ‘‘L’’ and Artemia
nauplii.
The cultivation environments for pelagic copepods are homogeneous and relatively
easy to operate and to scale up, but most of them use calanoids, and up to now can only
support very low densities (Støttrup and Norsker, 1997). On the other hand, benthic
copepods (mostly harpacticoids) are euriplastic species that are easier to cultivate at high
densities, have less specific nutritional requirements, and readily grow and reproduce with
short life cycles under a large variety of culture conditions (Kinne, 1977; Delbare et al.,
1996). However, their proliferation depends on the area of a solid substratum and such an
environment is not homogenous and, hence, is considerably more difficult to manage and
scale up (Støttrup and Norsker, 1997).
Adult benthic copepods tend to stay close to the bottom and walls of the culture
reservoir and, together with their rapid movements; this makes them difficult for the fish
larvae to feed on them. Pelagic nauplii and copepodite stages are more readily available to
the fish larvae (Kinne, 1977).
Kitajima (1973) stated that it is necessary to search for some suitable pelagic species
belonging to Calanoida and Cyclopoida. From the results of our cultivation trials, it can be
inferred that suitable species are those euriplastic pelagics with short life cycles, high
reproductive capacity that could feed on different food sources, especially microalgae and
some artificial feed. Using these criteria, copepods of the genus Oithona seem suitable for
large-scale hatchery culture of fish.
Based on our experience with O. oculata and those of Kraul (Hawaii, USA) with E.
acutifrons (Kraul, 1983b, 1989), we recommend several improvements for pelagic
copepod culture: (1) assessment of the effect of temperature, salinity, pH, and levels of
ammonia; (2) use of several cubic meter culture reservoirs with no more than 1-m depth
without additional substrates, best located indoors or outdoor in shade, and with sufficient
aeration to suspend microalgae but not in excess to affect sexual activities (approximately
2– 4 l min 1 m 3); (3) use baker’s yeast as a food source and a microalgae mixture of
different classes that does not settle to avoid slimy surfaces which can trap nauplii; we
recommend N. oculata, T. tetrathele, Isochrysis spp. (clone T. ISO), and a diatom such as
Chaetoceros muelleri. The proportion and density of the microalgae should be estimated
because the size and other characteristics can influence the daily feeding rate (Iwasaki and
Kamiya, 1977) (a maximum of 1  106 cells ml 1 is recommended); and (4) the batch and
semicontinuous systems should be evaluated, with partial harvest proportional to the
culture growth rate (approximately 10 – 30% total volume per day) to maintain the culture
in the best growth stage (a minimum of one turnover every week with more than 10
copepods ml 1, or every 2 weeks at concentrations of less than 1 copepod ml 1), and a
change to a clean tank every several weeks.
Judging by the results obtained in the present study, and those in the literature
(especially Japan), we recommend the establishment of a semicontinuous mass polyculture
of rotifers in great volumes together with the copepod O. oculata as a secondary species.
Although Delbare et al. (1996) considered T. japonicus to be the only species that can be
 O.G. Hernández Molejón, L. Alvarez-Lajonchère / Aquaculture 219 (2003) 471–483 479

cultured with rotifers, densities of 5– 10 copepods ml 1 can be obtained of O. oculata

with these techniques. This can sustain an adequate supply to larval-rearing tanks if their
volume keeps the 1:1 proportion of rotifer to larval rearing tank volumes recommended by
Tsujigado and Lee (1993).
Annual sales of Artemia cysts marketed for crustacean and larval fish food has
increased to more than 2000 t (Sorgeloos et al., 1994). This figure may be underestimated
considering the report of Stael et al. (1995) that 600 t was used in Thailand just for shrimp
culture in 1993. These high consumption levels can lead to a situation where the demand
exceeds the supplies (Stael et al., 1995), with consequent costs increases.
According to Lavens and Sorgeloos (1998), the current situation of the last few years is
characterized by poor yields from the main natural harvest site, the Great Salt Lake in Utah
(USA) that resulted in cyst shortage and increased cyst prices. It is expected that this
situation will not improve much during the next few seasons due to rather unfavorable
conditions for Artemia production in the Lake as well as increasing hatchery demand
(Lavens and Sorgeloos, 1998). This is known as the Artemia crisis (Sorgeloos, 2000).
There are reports of partial and total Artemia replacement with copepods (Liao, 1975;
Houde et al., 1976; Fujita, 1979; Witt et al., 1984; Yúfera and Pascual, 1984; James and
Al-Khars, 1985). Copepod culture has been considered commercially impractical or
technically difficult due to high cost by many authors (Girin and Person-Le Ruyet,
1977; Kraul, 1989; Kraul et al., 1992; Delbare et al., 1996), especially considering that
doubling times for most copepods are 5 –10 times longer than for rotifers (Tucker, 1992).
However, the densities needed in fish-rearing tanks are 5 – 10 times lower than those
usually used with rotifers. Copepods are one of the basic organisms in larval food
schedules in Asia (Fukusho, 1980, 1991; Lee et al., 1993; Kraul, 1993) and there are
reports of juvenile commercial production in Europe based on copepods as food (Bent,
1993), as well as recent research projects (European Commission, 1988) highlighting their
importance (Støttrup and Norsker, 1997; Sargent et al., 1997, 1999).
As a general conclusion, we propose that small pelagic copepod species with short life
cycles and fast growth could be cultured with high-yield semiintensive technologies. In
tropical and perhaps subtropical countries with low costs of technically qualified labor and
land, copepods could give good economic results in terms of highly stress-resistant larvae,
good survival, growth and biomass productions when compared with enriched Artemia.

Acknowledgements

This research was done at the Experimental Mariculture Station at Tunas de Zaza,
Sancti Spiritus Province, and at the Pilot-Scale Research Station at Santa Cruz del Sur,
Camagüey Province, Center for Fishery Research. The authors wish to express their
gratitude to Estela Beltran Estepe and Felipe Barbeira Docurro for their technical support,
and to Blanca Anderes and Amparo Campos for their interest and assistance on copepod
collection and identification. The authors are indebted to Dr. Sydney A. Kraul (HI, USA)
and Dr. Josianne G. Støttrup (Danish Institute for Fisheries Research) for their valuable
comments and recommendations and to Dr. Ellis Glazier (CIBNOR, La Paz, Mexico) for
editing of the English-language text of an earlier version of the manuscript.
480 O.G. Hernández Molejón, L. Alvarez-Lajonchère / Aquaculture 219 (2003) 471–483

References

Alvarez-Lajonchère, L., Pérez Sánchez, L., Torres Gómez, E., Hernández Molejón, O.G., 1994. Producción de
juveniles del patao Eugerres brasilianus (Pisces, Gerreidae) en Santa Cruz del Sur, Camagüey. Rev. Cuba.
Investig. Pesq. 18 (4), 25 – 27.
Alvarez-Lajonchère, L., Pérez Sánchez, L., Hernández Molejón, O.G., Torres Gómez, E., 1996. Mass production
of striped patao Eugerres brasilianus juveniles in Cuba. J. World Aquac. Soc. 27 (3), 347 – 352.
Bent, U., 1993. Methods for production of turbot fry based on copepods as food organisms. In: Carrillo, M.,
Dahle, L., Morales, J., Sorgeloos, P., Svennevig, N., Wyban, J. (Eds.), From Discovery to Commercializa-
tion, Oosteden. Eur. Aquac. Soc. Spec. Publ., vol. 19. European Aquaculture Society, Oostende, Belgium,
p. 609.
Bryan, P., Madraisau, B.B., 1977. Larval rearing and development of Siganus lineatus (Pisces: Siganidae) from
hatching through metamorphosis. Aquaculture 10, 243 – 252.
Campos Hernández, A., Suárez Morales, E., 1994. Copépodos pelágicos del Golfo de México y Mar Caribe: I.
Biologı́a y sistemática Centro de Investigaciones de Quintana Roo, Chetumal. 353 pp.
Coves, D., Audineau, P., Nicolas, J.L., 1990. Rotifers-rearing technology. In: Barnabe, G. (Ed.), Aquaculture,
vol. 1. Ellis Horwood, New York, pp. 232 – 245.
Delbare, D., Dhert, P., Lavens, P., 1996. Zooplankton. In: Lavens, P., Sorgeloos, P. (Eds.), Manual on the
Production and Use of Live Food for Aquaculture. FAO Fish. Tech. Pap., vol. 361. FAO, Rome, Italy,
pp. 252 – 282.
Detwyler, P., Houde, E.D., 1970. Food selection by laboratory-reared larvae of the scaled sardine Harengula
pensacolae (Pisces, Clupeidae) and the bay anchovy Anchoa mitchilli (Pisces, Engraulidae). Mar. Biol. 7,
214 – 222.
Doi, M., Singhagraiwan, T., 1993. Biology and culture of the red snapper, Lutjanus argentimaculatus. The
Research Project of Fishery Resource Development in the Kingdom of Thailand, Thailand. 51 pp.
European Commission, 1988. Progress in aquaculture R&D projects sponsored by the European Commission.
Aquac. Eur. 22 (3), 46 – 47.
Farran, G.P., 1913. Plankton from Christmas Island, Indian Ocean: 2. On Copepoda of the genera Oithona and
Paroithona. Proc. Zool. Soc. Lond., 181 – 193.
Fengqi, L., 1996. The use of rotifers in aquaculture. Aquac. Mag. 22 (3), 16 – 22.
Fujita, S., 1979. Culture of red sea bream, Pagrus major and its food. Spec. Publ. - EMS (14), 183 – 197.
Fukusho, K., 1980. Mass production of a copepod, Tigriopus japonicus, in combination culture with a rotifer
Brachionus plicatilis, fed-yeast as food source. Bull. Jpn. Soc. Sci. Fish. 46 (5), 625 – 629.
Fukusho, K., 1991. Review of the research status of zooplankton production in Japan. In: Fulks, W., Main, K.L.
(Eds.), Rotifer and Microalgae Culture Systems. Proceedings of U.S. – Asia Workshop, Honolulu, Hawaii,
January 28 – 31, 1991. Argent Laboratories, Redmond, pp. 55 – 60.
Fukusho, K., Arakawa, T., Watanabe, T., 1980. Food value of a copepod, Tigriopus japonicus, cultured with N-
yeast for larvae and juveniles of mud dab Limanda yokohamae. Bull. Jpn. Soc. Sci. Fish. 46 (4), 499 – 503.
Fulks, W., Main, K.L. (Eds.), 1991. Rotifer and Microalgae Culture Systems. Proceedings of U.S. – Asia Work-
shop, Honolulu, Hawaii, January 28 – 31, 1991. Argent Laboratories, Redmond. 364 pp.
Girin, M., Devauchelle, B., 1974. Production du rotifer Brachionus plicatilis O.F. Muller en elevage mixte avec
le copepod Tisbe furcata (Baird). Actes Colloq. CNEXO (1), 87 – 99.
Girin, M., Person-Le Ruyet, J., 1977. L’élevage larvaire des poissons marins: chaines alimentaires et aliments
composés. Bull. Fr. Piscic. 264, 88 – 101.
Hino, A., 1993. Present culture systems of the rotifer (Brachionus plicatilis) and function of microorganisms. In:
Lee, C.-S., Su, M.S., Liao, I.-C. (Eds.), Finfish Hatchery in Asia. Proceeding of Finfish Hatchery in Asia ’91.
Tungkang Marine Laboratory Conf. Proc., vol. 3. The Oceanic Institute, Hawaii, USA, pp. 51 – 59.
Houde, E.D., 1973. Some recent advances and unsolved problems in the culture of marine fish larvae. Proc.
World Maric. Soc. 3, 83 – 112.
Houde, E.D., Berkeley, S.A., Klinovsky, J.J., Schekter, R.C., 1976. Culture of larvae of the white mullet, Mugil
curema Valenciennes. Aquaculture 8, 365 – 370.
Iwasaki, H., Kamiya, S., 1977. Cultivation of marine copepod, Pseudodiaptomus marinus Sato. Bull. Plankton
Soc. Jpn. 24 (1), 44 – 54.
 O.G. Hernández Molejón, L. Alvarez-Lajonchère / Aquaculture 219 (2003) 471–483 481

Iwasaki, H., Katoh, H., Fujiyama, T., 1977. Cultivation of marine copepod, Acartia clausi Ciesbrecht: I. Bull.
Plankton Soc. Jpn. 24 (1), 55.
James, C.M., Al-Khars, A.M., 1985. Studies on the production of planktonic copepods for aquaculture. Annu.
Res. Rep. - Kuwait Inst. Sci. Res. 1984, 45 – 48.
Kahan, D., Azoury, M., 1981. Laboratory cultivation of Nitocra spinipes Boeck from the solar pond (Gulf of
Aoaba—Red Sea). Spec. Publ. - EMS 6, 117 – 131.
Kahan, D., Uhlig, G., Schwenzer, D., Horowitz, L., 1982. A simple method for cultivating harpacticoid copepods
and offering them to fish larvae. Aquaculture 26, 303 – 310.
Kanazawa, A., 1995. Effects of docosahexaenoic acid and phospholipids on stress tolerance of fish larvae.
In: Lavens, P., Jaspers, E., Roelants, I. (Eds.), Larvi’95 Fish and Shellfish Larviculture Symposium. Spec.
Publ. - Eur. Aquac. Soc., vol. 24. European Aquaculture Society, Ghent, Belgium, p. 105.
Kinne, O., 1977. Marine Ecology. Vol. III, Cultivation: Part 2. Wiley, New York. 1923 pp.
Kitajima, C., 1973. Experimental trials on mass culture of copepods. Bull. Plankton Soc. Jpn. 20 (1), 54 – 60.
Kraul, S., 1983a. Results and hypotheses for the propagation of the grey mullet, Mugil cephalus L. Aquaculture
30, 273 – 284.
Kraul, S., 1983b. Instructions for Culturing Euterpina acutifrons. Waikiki Aquarium, Hawaii. 1 p.
Kraul, S., 1989. Production of live prey for marine fish larvae. Actes Colloq. IFREMER (9), 595 – 607.
Kraul, S., 1993. Larviculture of the mahimahi Coryphaena hippurus in Hawaii, USA. J. World Aquac. Soc. 24
(3), 410 – 421.
Kraul, S., Nelson, A., Brittain, K., Wenzel, D., 1988. Feeding and handling requirements for hatchery culture of
the mahimahi Coryphaena hippurus. Quart. Univ. Hawaii Sea Grant Program 10 (3), 1 – 6.
Kraul, S., Ako, H., Brittain, K., Ogasawara, A., Cantrell, R., Nagao, T., 1991. Comparison of copepods and
enriched Artemia as feeds for larval mahimahi, Coryphaena hippurus. In: Lavens, P., Sorgeloos, P., Jaspers,
E., Ollevier, F. (Eds.), Larvi’91 Fish and Crustacean Larviculture Symposium. Eur. Aquac. Soc. Spec. Publ.,
vol. 15. European Aquaculture Society, Ghent, Belgium, pp. 45 – 47.
Kraul, S., Ako, H., Nelson, A., Brittain, K., Ogasawara, A., 1992. Evaluation of live feeds for larval and
postlarval mahimahi, Coryphaena hippurus. J. World Aquac. Soc. 23 (1), 299 – 306.
Kraul, S., Brittain, K., Cantrell, R., Nagao, T., Ako, H., Ogasawara, A., Kitagawa, H., 1993. Nutritional
factors affecting stress resistance in the larval mahimahi Coryphaena hippurus. J. World Aquac. Soc. 24 (2),
186 – 193.
Kuronuma, K., Fukusho, K., 1984. Rearing of marine fish larvae in Japan. International Development Research
Centre, Ottawa, IDRC-TS 47e. International Development Research Centre, Ottawa, Canada. 109 pp.
Lavens, P., Sorgeloos, P., 1998. Present status and prospects of the use of Artemia cysts and biomass in shrimp
farming. Anais do Aquicultura Brasil’98, vol. 1, Recife, 2 a 6 de Novembro de 1998. Latin American Chapter
of the World Aquaculture Society, Pernambuco, Brazil, pp. 147 – 162.
Lee, W.-J., Taga, N., 1988. Investigation of marine bacteria for the food of Tigriopus japonicus Mori (Harpacti-
coida). Bull. Korean Fish. Soc. 21 (1), 50 – 56.
Lee, C.-S., Su, M.S., Liao, I.-C. (Eds.), 1993. Finfish Hatchery in Asia: Proceedings of Finfish Hatchery in Asia
’91. Tungkang Marine Laboratory, Taiwan Fisheries Research Institute, Tungkang, Pingtung, Taiwan. 252 pp.
Leu, M.-M., Chou, Y.-H., 1996. Induced spawning and larval rearing of captive yellowfin porgy, Acanthopagrus
latus (Houttuyn). Aquaculture 143, 155 – 166.
Liao, I.C., 1975. Experiments on induced breeding of the grey mullet in Taiwan from 1963 to 1973. Aquaculture
6, 31 – 58.
May, R.C., 1970. Feeding larval marine fishes in the laboratory: a review. Calif. Mar. Res. Commun. Cal. COFI
Rep. 14, 76 – 83.
May, R.C., Popper, D., McVey, J.P., 1974. Rearing and larval development of Siganus canaliculatus (Park)
(Pisces: Siganidae). Micronesica 10 (2), 285 – 298.
McMichel Jr., R.H., Peten, K.M., 1989. Early life history of spotted seatrout, Cynoscion nebulosus (Pisces:
Sciaenidae), in Tampa Bay, Florida. Estuaries 12 (2), 98 – 110.
Morizane, T., 1991. A review of automation and mechanization used in the production of rotifers in Japan. In:
Fulks, W., Main, K.L. (Eds.), Rotifer and Microalgae Culture Systems. Proceedings of U.S. – Asia Workshop,
Honolulu, Hawaii, January 28 – 31, 1991. Argent Laboratories, Redmond, pp. 79 – 88.
Nassogne, A., 1969. La coltura dei copepodi in laboratorio. Pubbl. Stn. Zool. Napoli 37, 203 – 218.
482 O.G. Hernández Molejón, L. Alvarez-Lajonchère / Aquaculture 219 (2003) 471–483

Nassogne, A., 1970. Influence of food organisms on the development and culture of pelagic copepods. Helgol.
Wiss. Meeresunters. 20, 333 – 345.
Norsker, N.-H., Støttrup, J.G., 1994. The importance of dietary HUFAs for fecundity and HUFA content in the
harpacticoid, Tisbe holothuriae Humes. Aquaculture 125, 155 – 166.
Ohno, A., Takahashi, T., Taki, Y., 1990. Dynamic of exploited populations of the calanoid copepod Acartia
tsuensis. Aquaculture 84, 27 – 39.
Omori, M., 1973. Cultivation of marine copepods. Bull. Plankton Soc. Jpn. 20 (1), 3 – 11.
Saifulina, E., 1991. Larval rearing of the mullet Mugil so-iyu. In: Lavens, P., Sorgeloos, P., Jaspers, E., Ollevier,
F. (Eds.), Larvi’91 Fish and Crustacean Larviculture Symposium. Eur. Aquac. Soc. Spec. Publ., vol. 15.
European Aquaculture Society, Ghent, Belgium, pp. 317 – 318.
Sargent, J.R., McEvoy, L.A., Bell, J.G., 1997. Requirements, presentation and sources of polyunsaturated fatty
acids in marine fish larval feeds. Aquaculture 155, 117 – 127.
Sargent, J., McEvoy, L., Estevez, A., Bell, G., Bell, M., Henderson, J., Tocher, D., 1999. Lipid nutrition of marine
fish during early development: current status and future directions. Aquaculture 179, 217 – 229.
Shamsudin, L., Saad, C.R., 1993. Live food organisms used in Malaysia for mass propagation of marine shrimp
larvae, Penaeus monodon. In: Carrillo, M., Dahle, L., Morales, J., Sorgeloos, P., Svennecvcig, N., Wyban, J.
(Eds.), From Discovery to Commercialization, Oosteden. Eur. Aquac. Soc. Spec. Publ., vol. 19. European
Aquaculture Society, Oostende, Belgium, p. 170.
Sokal, R.R., Rohlf, F.J., 1981. Biometry. Freeman, San Francisco, CA.
Sorgeloos, P., 2000. President’s column. World Aquac. 31 (1) 3, 71.
Sorgeloos, P., Léger, P., 1992. Improved larviculture outputs of marine fish, shrimp and prawn. J. World Aquac.
Soc. 23 (4), 251 – 264.
Sorgeloos, P., Léger, P., Lavens, P., 1988. Improved larval rearing of European and Asian seabass, seabream,
mahi-mahi, siganid and milkfish using enrichment diets for Brachionus and Artemia. World Aquac. 19 (4),
78 – 79.
Sorgeloos, P., Dehasque, K.M., Dhert, P., Lavens, P., 1994. Larviculture of marine finfish: the current status.
Infofish Int. 4/94, 49 – 54.
Stael, M., Sanygontanagtit, T., van Ballaer, E., Puwapanieh, N., Tungsutapanich, A., Lavens, P., 1995. Decap-
sulated cysts and Artemia flakes as alternative food sources for the culture of Penaeus monodon postlarvae.
In: Lavens, P., Jaspers, E., Roelants, I. (Eds.), Larvi’95 Fish and Shellfish Larviculture Symposium. Eur.
Aquac. Soc. Spec. Publ., vol. 24. European Aquaculture Society, Ghent, Belgium, pp. 342 – 345.
Støttrup, J.G., Norsker, N.H., 1995. Production and use of copepods in marine fish larviculture. In: Lavens, P.,
Jaspers, E., Roelants, I. (Eds.), Larvi’95 Fish and Shellfish Larviculture Symposium. Eur. Aquac. Soc. Spec.
Publ., vol. 24. European Aquaculture Society, Ghent, Belgium, p. 320.
Støttrup, J.G., Norsker, N.H., 1997. Production and use of copepods in marine fish larviculture. Aquaculture 155,
231 – 247.
Støttrup, J.G., Richardson, K., Kirkegaard, E., Pihl, N.J., 1986. The cultivation of Acartia dana for use as a live
food source for marine fish larvae. Aquaculture 52, 87 – 96.
Støttrup, J.G., Gravningen, K., Norsker, N.H., 1995. The role of different algae in the growth and survival of
turbot larvae (Scophthalmus maximus L.) in intensive rearing systems. ICES Mar. Sci. Symp. 201, 173 – 186.
Tsujigado, A., Lee, C.-S., 1993. The design of a typical marine finfish hatchery in Japan using Mie prefectural
fish farming center as an example. In: Lee, C.-S., Su, M.S., Liao, I.-C. (Eds.), Finfish Hatchery in Asia.
Proceeding of Finfish Hatchery in Asia ’91. Tungkang Marine Laboratory Conf. Proc., vol. 3. The Oceanic
Institute, Hawaii, USA, pp. 27 – 39.
Tucker Jr., J.W., 1992. Feeding intensively-cultured marine fish larvae. Proceedings of the Aquaculture Nutrition
Workshop, Salamander Bay, 15 – 17 April 1991. NSW Fisheries, Brackish Water Fish Culture Research
Station, Salamander Bay, Australia, pp. 129 – 146.
Watanabe, T., Kiron, V., 1994. Prospects in larval fish dietetics. Aquaculture 124, 223 – 251.
Watanabe, T., Ohta, M., Kitajima, M.C., Fujita, S., 1982. Improvement of dietary value of brine shrimp Artemia
salina for fish larvae by feeding them on omega-3 highly unsaturated fatty acids. Bull. Jpn. Soc. Sci. Fish. 48,
1775 – 1782.
Watanabe, T., Kitajima, C., Fujita, S., 1983. Nutritional value of live organisms used in Japan for mass prop-
agation of fish: a review. Aquaculture 34, 115 – 143.
 O.G. Hernández Molejón, L. Alvarez-Lajonchère / Aquaculture 219 (2003) 471–483 483

Witt, U., Quantz, G., Kuhlmann, D., Kattner, G., 1984. Survival and growth of turbot larvae Scophthalmus
maximus L. reared on different food organisms with special regard to long-chain polyunsaturated fatty acids.
Aquac. Eng. 3, 190 – 1771.
Yúfera, M., Pascual, E., 1984. La producción de organismos zooplanctónicos para la alimentación larvaria en
acuicultura marina. Inf. Téc. Inst. Investig. Pesq. 119, 3 – 27.
Zacarı́as, D., Zoppi de Roa, E., 1981. Desarrollo larval y postlarval de Oithona hebes Giesbrecht (Copepoda:
Ciclopoida). Acta Biol. Venez. 11 (2), 109 – 123.
Zillioux, E.J., 1969. A continuous recirculating culture system for planktonic copepods. Mar. Biol. 4, 215 – 218.
Zismann, L., Berdugo, V., Kimor, B., 1975. The food and feeding habits of early stages of grey mullets in the
Haifa bay region. Aquaculture 6 (1), 59 – 76.
Zurlini, G., Ferrari, I., Nassogne, A., 1978. Reproduction and growth of Euterpina acutifrons (Copepoda:
Harpacticoida) under experimental conditions. Mar. Biol. (46), 59 – 64.