a rearing system for the culture of ornamental decapod crustacean larvae

A rearing system for the culture of ornamental

decapod crustacean larvae

R. Caladoa,*, L. Narcisoa, S. Moraisa, A.L. Rhyneb, J. Linb

aLaboratorio Marıtimo da Guia—IMAR, Estrada do Guincho, 2750-642 Cascais, PortugalbDepartment of Biological Sciences, Florida Institute of Technology, 150 University Boulevard,

Melbourne, FL 32901, USA

Received 9 May 2002; received in revised form 15 August 2002; accepted 16 October 2002

Abstract

The design and operation of a small research scale and a mass commercial scale rearing system

for the culture of marine ornamental decapod crustacean larvae are described in the present paper.

Preliminary data on the culture of the Mediterranean cleaner shrimp (Lysmata seticaudata),

peppermint shrimp (Lysmata wurdemanni), blue-white partner shrimp (Periclimenes sagittifer),

sponge crab (Cryptodromiopsis antillensis) and green emerald crab (Mithraculus sculptus) are also

presented. The use of these ‘‘plantonkreisel’’ based systems allowed the complete larval

development of the above-mentioned species, inducing minimal mechanical stress while keeping

an excellent water quality. Higher survival rates (up to 70% and 60% for L. seticaudata and L.

wurdemanni, respectively) to the post-larval stage and a shorter larval stage duration (27 and 22 days

for L. seticaudata and L. wurdemanni, respectively) were achieved, in comparison to conventional

rearing systems. This culture technology may play a key role in the realisation of a commercial

culture of these highly priced crustacean species and therefore the reduction of wild specimen

collection.

D 2003 Elsevier Science B.V. All rights reserved.

Keywords: Marine ornamentals; Crustacean culture; Larvae; Aquaculture systems

1. Introduction

Marine ornamental species trade is a multi-million-dollar industry. Along with corals,

marine tropical decapod crustaceans are among the most popular invertebrate species in

0044-8486/03/$ - see front matter D 2003 Elsevier Science B.V. All rights reserved.

PII: S0044 -8486 (02 )00583 -5

* Corresponding author. Tel.: +351-21-4869211; fax: +351-21-4869720.

E-mail address: [email protected] (R. Calado).

www.elsevier.com/locate/aqua-online

Aquaculture 218 (2003) 329–339

the aquarium trade industry. In recent years, researchers, traders, collectors and

hobbyists have began a worldwide effort to minimise the growing pressure on natural

populations of marine ornamental species and to promote the sustainable use of these

highly valued resources (Corbin, 2001). Nevertheless, this goal will only be achieved if

wild specimen collection is significantly replaced by the artificial rearing of these

species.

The development of artificial culture methodologies for marine ornamental decapod

crustaceans has been focused on a limited number of shrimp species. Among them, the

tropical species of the genus Lysmata, Periclimenes, Hymenocera and Stenopus have

received special attention, mainly due to their growing demand in the aquarium trade

industry (Fiedler, 1994; Fletcher et al., 1995; Palmtag and Holt, 2001; Lin et al., 2001).

Recently, Calado et al. (2001a) have started to evaluate the rearing potential of certain

species of those genera present in temperate and subtropical Eastern Atlantic and

Mediterranean Sea. This approach may be of particular commercial interest for Western

European countries, which are only second to the USA in marine ornamentals imports

(Lem, 2001).

In order to reduce the gap between supply and increasing demand through the

commercialisation of captive raised organisms, one special constraint must be over-

come—larval mass rearing. Lysmata, Hymenocera, Periclimenes, and Stenopus are

known, or at least believed, to have a large number of larval stages (Caroli, 1918; Kurata,

1970; Goy, 1991; Fiedler, 1994; Fletcher et al., 1995; Wunsch, 1996; Zhang et al., 1998).

Under unsuitable rearing conditions, larval stages may last longer and larval substages and

mark-time moulting (a series of moults where small or no morphological changes take

place) may also occur (Gore, 1985; Calado et al., 2001b). Long larval periods with high

mortality rates impair the production of several other commercially important species (e.g.

scyllarid and palinurid lobsters). In addition, the presence of frail long paddle-like

appendages (e.g. in Lysmata larvae) and carapace spines (e.g. in Cryptodromiopsis and

Mithraculus larvae) must be taken into account when designing a suitable larval rearing

system.

Rearing systems based on Greve’s (1968) ‘‘planktokreisel’’ proved to be very

appropriate for the culture of the frail spiny lobster larvae (Illingworth et al., 1997;

Kittaka, 1997). The main characteristic of these systems is the maintenance of larvae and

food in suspension only through water upwelling motion. Water aeration may induce

damage to the larvae and sometimes does not provide an adequate water circulation, which

will cause late stage larvae to sink in the rearing tanks. The larval aggregation in the

bottom of the tanks will certainly cause ‘‘tangling’’, damage larval appendages or even the

death of the larvae. The ‘‘planktokreisel’’-based system allows larvae to develop with

minimal mechanical stress, while providing adequate water renewal and circulation.

The present paper describes the design and operation procedures of a small research

scale and a mass commercial scale larval rearing systems for ornamental decapods.

Preliminary results on the captive rearing of the subtropical Mediterranean cleaner shrimp

(Lysmata seticaudata) and blue-white partner shrimp (Periclimenes sagittifer) and the

tropical peppermint shrimp (Lysmata wurdemanni), sponge crab (Cryptodromiopsis

antillensis), and green emerald crab (Mithraculus sculptus) are presented and compared

to previous studies using conventional rearing techniques.

R. Calado et al. / Aquaculture 218 (2003) 329–339330

2. Materials and methods

2.1. Small research scale tanks

White cylindrico-conical fiberglass tanks were designed and manufactured with a

diameter of 0.25 m, 0.4 m of total height, and a conical slope at 0.23 m from the top. The

approximate volume was 12 l and a 12-mm diameter drain valve was fitted to the base.

Round Nitexk mesh screen (80-mm diameter) was mounted 0.1 m from the water surface

and attached to a 13-mm diameter outlet, placed 45 mm from the top of the tank. Mesh

size was dependent on the used prey (50 and 150 Am when using Brachionus and Artemia,

respectively).

The tanks were connected in parallel to a recirculation system composed of a strongly

aerated 30 l sump, filled with biological media and a protein skimmer. A 19-mm drain

valve was fitted at the base of the sump to allow for the complete drainage of the system.

Water was pumped to an 80-l head tank with a pump rated at 6000 l h� 1 and passed

through a 40-W ultra-violet filter and a 10-Am cartridge filter. The head tank’s water was

returned to the sump and distributed to the culture tanks through 19-mm diameter

polyvinyl chloride (PVC) pipes. Each culture tank was gravity-fed water from the head

tank through a 19-mm valve manifold. A 5-mm-diameter inlet connected to the valve

manifold was positioned in the deepest point of the rearing tank. Rearing tank water

passed through the tank screen and was returned to the sump through a 19-mm-diameter

PVC pipe (Fig. 1).

Larvae of L. wurdemanni and M. sculptus were hatched in the laboratory from captive

ovigerous females. C. antillensis larvae were obtained from a wild caught ovigerous

female. The most active larvae (the ones displaying pronounced positive phototactic

responses—approximately 90%) were selected and transferred to the rearing tanks. Three

rearing tanks per species were stocked with 100 larvae (about 8 larvae l� 1). Five

haphazardly selected larvae per tank were sampled and staged every other day to determine

stage duration. L. wurdemanni larvae were staged according to Kurata (1970), while Rice

and Provenzano (1966) and Wilson et al. (1979) were followed for C. antillensis and M.

sculptus larvae, respectively. After staging, the larvae were returned to the rearing tanks.

Seawater was 1 Am-filtered, salinity was maintained at 35F 1x and temperature was

kept at 26F 1 jC through a heating/cooling system. The tanks were illuminated from

above with fluorescent light, with an intensity of 1000 lux at the water surface and a

photoperiod of 14 h light/10 h dark.

During the first larval stage, the larvae were fed on Brachionus sp. grown with Culture

Selcok at a density of 15000 nauplii l� 1. Water flow in this period was maintained at 0.5

l min� 1. The following larval stages were fed daily on Artemia franciscana (Kellogg,

1906) (Brine Shrimp Direct) metanauplii, enriched with DHA Selcok at a density of 5000

metanauplii l� 1 and the water flow was increased to 1 l min� 1. Beyond the fifth larval

stage, the water flow was increased to 2 l min� 1, in order to keep the larvae suspended in

the water column.

When the last zoeal stage was reached, two strips of indoor/outdoor carpeting (0.05 wide

by 0.25m long) were submerged in each rearing tank. Besides providing an extra area for lar-

val settlement, these strips also provided shelter for negatively phototatic post-larval stages.

R. Calado et al. / Aquaculture 218 (2003) 329–339 331

To flush the culture tanks of uneaten prey, an 800-ml beaker was put inside the rearing

tank, in order to displace a certain amount of water through the drain outlet. The 50-

or150-Am mesh screens were then replaced by a 500-Am mesh screen (Fig. 2). Rearing

tanks were allowed to flush for 2 h daily and a 50-Am mesh bag filter was placed in the

sump to collect any uneaten prey.

2.2. Commercial mass scale tanks

White cylindrico-conical fiberglass tanks were also designed and manufactured, with a

maximum diameter of 0.8 m, 1 m of total height, and a conical slope at .0.7 m from the

Fig. 1. Small research scale rearing system: (1) water pump; (2) sump filled with biological media, with a protein

skimmer; (3) ultra-violet light filter; (4) cartridge filter; (5) head tank; (6) inflow pipes; (7) outflow pipes; (8)

rearing tanks. Grey and black arrows represent in and outflow, respectively.


top. The tanks had an approximate volume of 200 l and a 15-mm-diameter drain valve was

fitted to the base. Round Nitexk mesh screen (0.2-m diameter) was fitted 0.3 m from the

water surface and connected to a 25-mm-diameter outlet, placed 0.1 m from the top of the

tank (Fig. 3). Mesh size was as previously described for the small research scale screens.

Fig. 3. Commercial mass scale rearing system. A: (1) water pump; (2) sump filled with biological media, with a

protein skimmer; (3) rearing tanks; (4) inflow pipe; (5) outflow pipe. Rearing tank detail. B: (6) fluorescent light;

(7) inflow pipe with inlet at the bottom of the rearing tank; (8) mesh screen; (9) outflow pipe. Grey and black

arrows represent in and outflow, respectively.

Fig. 2. Mesh screen changing. (A) The rearing tank has a regular water level and a fine mesh screen. (B) A 800-ml

beaker is put inside the rearing tank raising the water level. (C) The beaker causes displaced water to be drained,

lowering the water level bellow the water outlet. (D) The fine mesh screen may be detached. (E) The fine mesh

screen may be replaced by a larger mesh screen. (F) Water raises to regular level fluxing uneaten prey.


The commercial mass scale rearing tanks were also connected in parallel to a recirculation

system made up of a strongly aerated tank (approximate volume of 220 l) filled with

biological media, to a protein skimmer and to a water pump providing a flux of 5700 l

h� 1. A 30-mm drain valve was fitted at the base of the recirculation tank, in order to allow

the complete drainage of the rearing system.

Ultra-violet and 10-Am filtered water was supplied. Water was fed from 25-mm supply

lines positioned in the deepest point of the rearing tank. Water was returned to the sump as

previously described for the small research scale system.

Larvae of L. seticaudata and P. sagittifer were obtained from wild caught ovigerous

females. As previously described, only the most active larvae were chosen and transferred

to the rearing tanks. Six rearing tanks (three per species) were stocked with approximately

3500 larvae each (about 18 larvae l� 1), while other three tanks were stocked with 5000 L.

seticaudata larvae each (about 25 larvae l� 1). Thirty haphazardly selected larvae from

each tank were sampled daily and developmental stage assessed (modified from Caroli,

1918 for L. seticaudata and modified from Bourdillon-Casanova, 1960 for P. sagittifer).

Dead larvae were removed daily with a siphon, after stopping the water flow in the rearing

tanks, and recorded.

All other rearing conditions were the same as for the small research scale tanks, with

the exceptions of water temperature (which was kept at 22F 1 jC since these two species

are subtropical) and flow (2.5 l min� 1 on rotifers and 5 l min� 1 on Artemia metanauplii),

Artemia enrichment product (Algamac 2000k) and settlement strips (0.1 m wide by 0.5 m

long).

To compare the effect of different larval densities on the mass culture of L. seticaudata,

a t-test analysis was used. All results were considered statistically significant at the 0.05

probability level (Sokal and Rohlf, 1995).

In the present study, all caridean shrimp larval stages will be named zoea and the term

mysis will not be used to designate late stage larvae displaying pleopods (Williamson,

1969).

3. Results

3.1. Small research scale tanks

All the species were cultured to the post-larval stage. Each larval stage of C. antillensis

andM. sculptus lasted 2 and 3 days, respectively. The megalopa stage for both crab species

presented an average duration of 6 days (Table 1). All larval stages of L. wurdemanni

lasted 2 days (Table 1). Due to the fast decomposition of dead larvae, data on mean

survival rate by larval stage could not be accurately obtained. Therefore, only the mean

percentage of larvae reaching the post-larval stage was recorded. The survival rates to the

post-larval stage for C. antillensis, M. sculptus and L. wurdemanni were 20.6%, 22.1% and

60.2%, respectively.

The sponge crab (C. antillensis) megalopa was the only larval stage displaying strong

cannibalistic behaviour, mainly preying on zoea VI larvae and other smaller megalopa and

showing almost no interest in the much smaller Artemia metanauplii.


3.2. Commercial mass scale tanks

With the exception of the last zoeal stage, all larval stages of L. seticaudata lasted an

average of 3 days (Table 2). In order to determine the survival rate and the percentage of

larvae displaying ‘‘mark-time’’ moulting, all the larvae were removed from the rearing

tanks 20 days after the first post-larva were recorded. This procedure did not allow us to

estimate the average stage duration for the 9th larval and only the minimum duration (3

days) was recorded. Nevertheless, our observation indicated that most of the larvae

metamorphosed to the post-larval stage within 10 days after the first larva on the 9th zoeal

stage was recorded. Larval stage duration was not significantly different between the two

tested stocking densities ( p= 0.534).

Similar to the small research scale larval cultures, the rapid decomposition of dead

larvae impaired the accurate estimation of mean survival rate by larval stage. Therefore,

only the mean percentage of larvae reaching the post-larval stage and the percentage of

larvae displaying ‘‘mark-time’’ moulting (never metamorphosed into post-larval stage)

were recorded (Table 2). There was an indication that most mortality on the larviculture of

L. seticaudata occurred after Zoea 4. The rearing tanks stocked with 18 larvae l� 1

displayed a significantly lower ( p = 0.016) number of larvae ‘‘marking-time’’ (1.1% vs.

12.3%) and consequently a significantly higher ( p = 0.0007) number of larvae reaching the

post-larval stage than the ones stocked with 25 larvae l� 1 (61.1% vs. 52.4%).

Table 2

Preliminary results on the larviculture of L. seticaudata (at different densities) and of P. sagittifer: average stage

duration (days) and percentage of larvae that displayed mark-time moulting (meanF S.D.) and that survived to

post-larvae (meanF S.D.)

Species Number of

larval stages

Average stage duration Marking time Survival rate (%)

to post-larvae

Total

L. seticaudata

(at 18 larvae l� 1)

9 Zoeas 3 days per zoeal stage,

except the last stagea1.1F 0.6 61.1F 5.2 62.3F 8.33

L. seticaudata

(at 25 larvae l� 1)

9 Zoeas 3 days per zoeal stage,

except the last stagea12.3F 3.0 52.4F 4.6 64.8F 5.2

P. sagittifer 8 Zoeas 2 days per zoeal stage,

except the last stagea73.2F 5.4 0a 73.2F 5.4

a See text for explanation.

Table 1

Preliminary results on the larviculture of L. wurdemanni, C. antillensis andM. antillensis: number of larval stages,

average stage duration (days) and average survival rate to the post-larval stage (%) (meanF S.D.)

Species Number of

larval stages

Average stage duration Survival rate to

post-larval stage (%)

L. wurdemanni 11 Zoeas 2 days per zoeal stage 60.2F 3.1

C. antillensis 6 Zoeas and

1 Megalopa

2 days per zoeal stage;

6 days in megalopa

20.6F 6.5

M. sculptus 2 Zoeas and

1 Megalopa

3 days per zoeal stage;

6 days in megalopa

22.1F 6.9


With the exception of the last stage, each larval stage of P. sagittifer lasted only 2 days

(Table 2). However, all the larvae displayed ‘‘mark-time’’ moulting in the last stage and

none of them reached the post-larval stage 20 days after moulting to Zoea 8. After this

period, 10 larvae were haphazardly chosen and placed in a small mesh basket in a 70-l

tank stocked with adults and their host anemone Anemonia sulcata. A single zoea survived

and metamorphosed to the post-larval stage in 7 days.

4. Discussion

Kurata (1970) and Goy (1991) cultured L. wurdemanni larvae using glass bowls and

obtained an average larval duration of 43 and 110 days, respectively, with survival

rates to the post-larval stage lower than 15%. Zhang et al. (1998) reported to L.

wurdemanni larvae raised in 4-l bottles 29 to 36 days of larval duration, with survival

rates of approximately 67%. However, Lin (2000) recognized that the cultured species

was actually the closed related species Lysmata rathbunae. Recently, Rhyne (2002)

questioned the validity of L. rathbunae and hypothesized the existence of a puzzling

and complex group of closely related forms of ‘‘peppermint shrimp’’ in the Caribbean

and the Gulf of Mexico, which have certainly been misidentified in the past as L.

wurdemanni. In this way, until this systematic issue is solved (Rhyne et al., work in

progress), every comparison between the present study and previous ones should be

regarded with caution. Rice and Provenzano (1966) reared C. antillensis to the post-

larval stage in plastic trays, in 30 to 40 days with a survival rate of only 11%.

Although no other data are presently available for M. sculptus larviculture, Wilson et

al. (1979) reared the closely related species M. forceps to the post-larval stage in

plastic compartment trays in 14 days, and with survival rates lower than 5%. When

compared to the studies mentioned above, using conventional rearing systems, all these

species displayed much shorter larval stage duration and generally higher survival rates

when cultured on our small research scale system.

Couturier-Bhaud (1974) reared L. seticaudata in small beakers, reaching the last larval

stage (Zoea 9) in 43 days. However, all the larvae died before metamorphosing to the post-

larval stage. Bourdillon-Casanova (1960) raised an unknown Mediterranean species of

Periclimenes in glass beakers and a single larva was cultured to the last zoeal stage (Zoea

8) but failed to metamorphose to the post-larval stage. In the present work, several

thousand L. seticaudata post-larvae were mass raised in a short period of time, with high

survival rates using this alternative rearing system. The poor results on the larval rearing of

P. sagittifer can probably be explained by the absence of settlement cues in the rearing

system. Goy (1991) obtained higher survival rates to the post-larval stage when raising

Periclimenes pedersoni and P. yucatanicus larvae (21% for both species) exposed to their

host anemone exudates. Since the larvae of P. sagittifer, only metamorphosed after being

exposed to adults and their host anemone (A. sulcata) exudates, future studies should

evaluate the importance of each of these settlement cues. Although the larvae should not

be in direct contact with the host anemone, as this would result in the death of the larvae

(Goy, 1991). This can be easily avoided by placing the host anemone, the adults, or both in

a mesh basket in the systems head tank or sump.


It is well known that the nutritional value of enriched brine shrimp metanauplii decreases

rapidly 24 h after being supplied to the larvae (Narciso et al., 1999; Navarro et al., 1999).

Therefore, the daily cleaning and maintenance of the rearing tanks, particularly the daily

washing and exchanging of the 200- and 500-Am mesh screens, not only allowed the

maintenance of good water quality but also the daily replacement of 24-h-old enriched

metanauplii by newly enriched ones. This procedure played a vital role during the entire

rearing process since it enabled the larvae to feed on highly nutritional preys. The

traditional water change of other rearing systems often requires larval manipulation and

induces stress to the larvae. Water change in our system is not only far less stressful to the

larvae but also less time consuming. One of the potential solutions for the cannibalistic

behaviour presented by the sponge crab megalopa, which may also occur in other

ornamental decapod species, is to present bigger prey (such as older enriched brine shrimp

or live mysids) for late larval stages.

The control of water velocity of the upwelling flux in each rearing tank was also

very important for the successful larval culture of these species. The upwelling water

flux kept in suspension the larvae of all stages, including the large last larval stages

of L. seticaudata and L. wurdemanni and the bulky sponge crab megalopa. This

upwelling flow prevented larval accumulation on the bottom of the rearing tank and

the consequent larval ‘‘tangling’’. An unequivocal evidence of this was the presence

of both frail paddle shape 5th pereiopods in about 95% of late stages Lysmata

larvae.

The artificial grass strips were very useful in the commercial mass scale rearing tanks,

not only for increasing the tank settlement area, but also for easy removal of newly settled

shrimp post-larvae that firmly clanged onto this structure. This process avoided draining

the culture tank through the bottom valve, which could obviously damage the zoeal stages.

The present work provides some clues on the mechanisms responsible for mark-time

moulting on caridean larvae. In this study, the initial stocking density of larvae seemed to

play the major role on the occurrence of this type of moulting. The sole reduction of the

rearing density from 25 to 18 larvae l� 1 was enough to drastically reduce the percentage

of larvae marking-time. The importance of settlement cues for shortening larval duration,

particularly the exudates of the hosts of species displaying close symbiotic relations, was

already stressed.

Despite the fact that only caridean shrimp and true crab larvae were tested in the present

study, morphological similarities have been shown between dromioid crab and anomuran

hermit crab larvae, as well as caridean shrimp and other Pleocyemata larvae (e.g.

stenopodids and nephropids) (Williamson and Rice, 1996). Since the ‘‘planktokreisel’’-

based systems confirmed to be very appropriate for the culture of Palinura phyllosoma

larvae (Kittaka, 1997), the described rearing systems may be used to culture other

ornamental decapods: shrimp, hermit crabs, true crabs and lobsters. For some species,

e.g. hermit crabs and some true crabs, production may be optimised if the larvae are raised

to the megalopa stage in our systems and then moved to other rearing systems where

settlement induction can be better achieved (Davis and Stoner, 1994; Forward et al., 2002).

This would allow the use of important settlement cues, e.g. substrate for true crabs

(O’Connor, 1991) and gastropod shells for hermit crabs (Harvey, 1996), shortening the last

larval stage duration and improving survival rates.


Future studies using this methodology should focus on the species elected by traders as

the ‘‘most desirable to be cultured’’: the fire shrimp Lysmata debelius, the skunk shrimp

Lysmata amboinensis, the coral banded shrimp Stenopus hispidus, the harlequin shrimp

Hymenocera picta and reef lobsters Enoplometopus spp. (Moe, 2001).

Although the main efforts for the development of rearing techniques for these highly

priced ornamental species are taking place in the importing countries (namely the USA,

Japan and EU countries), exporting countries in Southeast Asia, Caribbean, Eastern Africa

and Red Sea regions have strong potential for ornamental species production, with

excellent climate conditions and low production costs. The transfer of culture technology

to these countries will certainly allow local authorities to establish more effective

conservation programs while still generating important financial incomes (Fletcher et

al., 1999).

Acknowledgements

The authors would like to thank the Luso-American Foundation for Development

and the Fundac�ao para a Ciencia e a Tecnologia (scholarship SFRH/BD/983/2000 and

research project POCTI/BSE/43340/2001) from the Portuguese government for their

financial support. We would also like to acknowledge American Eagle Canoesk and

Fernando Ribeirok for their enthusiastic support in the design and manufacture of the

smaller rearing tanks and system. We also thank Jennifer Stefaniak, Rhian Resnick and

Marcus Zokan for the technical support and Catia Bartilotti for the rearing system

drawings.

References

Bourdillon-Casanova, L., 1960. Le meroplancton du Golfe de Marseille: les larves des crustaces decapodes. Recl.

Trav. Stn. Mar. Endoume 30, 1–286.

Calado, R., Morais, S., Narciso, L., 2001a. Temperate shrimp: perspective use as ornamental species. Book of

Abstracts, Marine Ornamentals 2001. University of Florida, Sea Grant Florida and National, Florida Depart-

ment of Agriculture, Orlando, USA, pp. 118–119.

Calado, R., Martins, C., Santos, O., Narciso, L., 2001b. Larval development of the Mediterranean cleaner shrimp

Lysmata seticaudata (Risso, 1816) (Caridea: Hippolytidae) fed on different diets-costs and benefits of mark—

time molting. In: Hendry, C.I., Van Stappen, G., Wille, P., Sorgeloos, P. (Eds.), Larvi ’01 Fish and Crustacean

Larviculture Symposium. European Aquaculture Society, Special Publication, vol. 30, pp. 96–99. Belgium.

Caroli, E., 1918. Miersia clavigera Chun, stadio misidiforme di Lysmata seticaudata Risso. Pubbl. Stn. Zool.

Napoli 2, 177–189.

Corbin, J.S., 2001. Marine Ornamentals ’99, conference highlights and priority recommendations. In: Burgess, P.

(Ed.), Marine Ornamental Conference, Hawaii. Aquarium Sciences and Conservation, vol. 3, pp. 3–11.

Couturier-Bhaud, Y., 1974. Cycle biologique de Lysmata seticaudata Risso (Crustace, Decapode): III. Etude du

developpement larvaire. Vie Milieu, Ser. A Biolog. Mar. 24, 431–442.

Davis, M., Stoner, A.W., 1994. Trophic cues induce metamorphosis of queen conch larvae (Strombus gigas

Linnaeus). J. Exp. Mar. Biol. Ecol. 180, 83–102.

Fiedler, G.C., 1994. The larval stages of the Harlequin shrimp, Hymenocera picta (Dana). MS thesis, University

of Hawaii at Manoa, US. 101 pp.

Fletcher, D., Kotter, I., Wunsch, M., Yasir, I., 1995. Preliminary observations on the reproductive biology of

ornamental cleaner prawns. Int. Zoo Yearb. 34, 73–77.


Fletcher, D.J., Wood, E., Jones, D.A., 1999. Marine ornamental culture and reef conservation. Aquaculture 99

Book of Abstracts. World Aquaculture Society, Sydney, Australia, p. 265.

Forward Jr., R.B., Tankersley, R.A., Welch, J.W., 2002. Selective tidal-stream transport of the blue crab Calli-

nectes sapidus: an overview. Bull. Mar. Sci. (in press).

Gore, R., 1985. Molting and growth in decapod larvae. In: Wenner, A.M. (Ed.), Crustacean Issues, vol. 2—Larval

Growth. AA Balkema Publishers, Rotterdam, Netherlands, pp. 1–65.

Goy, J., 1991. Components of reproductive effort and delay of larval metamorphosis in tropical marine shrimp

(Crustacea:Decapoda: Caridea and Stenopodidea). PhD Thesis, Texas A&M University, College Station,

USA. 193 pp.

Greve, W., 1968. The ‘‘planktonkreisel’’, a new device for culturing zooplankton. Mar. Biol. (Berl.) 1, 201–203.

Harvey, A.W., 1996. Delayed metamorphosis in Florida hermit crabs: multiple cues and constraints (Crustacea:

Decapoda: Paguridae and Diogenidae). Mar. Ecol., Prog. Ser. 141, 27–36.

Illingworth, J., Tong, L., Moss, G., Pickering, T., 1997. Upwelling tank for culturing rock lobster (Jasus

edwardsii) phyllosomas. Mar. Freshw. Res. 48, 911–914.

Kittaka, J., 1997. Application of ecosystem method for complete development of phyllosomas of spiny lobster.

Aquaculture 155, 319–331.

Kurata, H., 1970. Studies on the life histories of decapod crustacea of Georgia. PhD Thesis, University of

Georgia, Sapelo Island, GA, USA. 274 pp.

Lem, A., 2001. International trade in ornamental fish. Book of Abstracts, Marine Ornamentals 2001. University

of Florida, Sea Grant Florida and National, Florida Department of Agriculture, Orlando, USA, p. 26.

Lin, J., 2000. Errata. J. World Aquac. Soc. 31, 476.

Lin, J., Zhang, D., Creswell, R., 2001. Aquaculture of marine ornamental shrimps: an overview. Aquaculture

2001 Book of Abstracts. World Aquaculture Society, Orlando, FL, USA, p. 378.

Moe Jr., M.A., 2001. Culture of marine ornamentals: for love, for money and for science. Book of Abstracts,

Marine Ornamentals 2001. University of Florida, Sea Grant Florida and National, Florida Department of

Agriculture, Orlando, USA, pp. 27–28.

Narciso, L., Pousao-Ferreira, P., Passos, A., Luıs, O., 1999. HUFA content and DHA/EPA improvements of

Artemia sp. with commercial oils during different enrichment periods. Aquac. Res. 30, 21–24.

Navarro, J.C., Henderson, R.J., McEvoy, L.A., Bell, M.V., Amat, F., 1999. Lipid conversions during enrichment

of Artemia. Aquaculture 174, 155–166.

O’Connor, N.J., 1991. Flexibility in timing of the metamorphic molt by fiddler crab megalopae Uca pugilator.

Mar. Ecol., Prog. Ser. 68, 243–247.

Palmtag, M.R., Holt, G.J., 2001. Captive rearing of fire shrimp (Lysmata debelius). Texas Sea Grant College

Program Research Report, pp. 1–4.

Rhyne, A.L., 2002. Improvements in marine ornamental culture. MS thesis, Florida Institute of Technology,

Melbourne, FL, USA. 76 pp.

Rice, L.A., Provenzano Jr., A.J., 1966. The larval development of the west Indian sponge crab Dromidia

antillensis (Decapoda: Dromiidae). J. Zool. (Lond.) 149, 297–319.

Sokal, R., Rohlf, F., 1995. Biometry—The Principles and Practice of Statistics in Biological Research, 3rd ed.

Freeman, New York, USA. 887 pp.

Williamson, D.I., 1969. Names of larvae in the Decapoda and Euphasiacea. Crustaceana 16, 210–213.

Williamson, D.I., Rice, A., 1996. Larval evolution in the crustacea. Crustaceana 69, 267–287.

Wilson, K.A., Scotto, L.E., Gore, R.H., 1979. Studies on decapod Crustacea from the Indian River region of

Florida: XIII. Larval development under laboratory conditions of the spider crab Mithrax forceps (A. Milne-

Edwards, 1875) (Brachyura: Majidae). Proc. Biol. Soc. Wash. 92, 307–327.

Wunsch, M., 1996. Larval development of Lysmata amboinensis (De Man, 1888) (Decapoda: Hippolytidae)

reared in the laboratory with a note on L. debelius (Bruce, 1983). Ms Thesis, University of Wales, Bangor,

United Kingdom. 110 pp.

Zhang, D., Lin, J., Creswell, R., 1998. Effects of food and temperature on survival and development in the

peppermint shrimp Lysmata wurdemanni. J. World Aquac. Soc. 29, 471–476.


a rearing system for the culture of ornamental decapod crustacean larvae

Documents