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Crowding female red claw

crayfish, Cherax
quadricarinatus, under smalltanks hatchery conditions:
What is the limit?
ARTICLE in AQUACULTURE JANUARY 2000
Impact Factor: 1.88 DOI: 10.1016/S0044-8486(99)00235-5

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Aquaculture 181 2000. 235240

www.elsevier.nlrlocateraqua-online

Crowding female red claw crayfish, Cherax

quadricarinatus, under small-tanks hatchery
conditions: what is the limit?
Assaf Barki ) , Ilan Karplus
Department of Aquaculture, Institute of Animal Science, Agricultural Research Organization, The Volcani
Center, P.O. Box 6 Bet Dagan 50250, Israel
Accepted 1 June 1999

Abstract
Previous studies on reproduction in the red claw crayfish, Cherax quadricarinatus, have
successfully employed broodstock densities of up to 20 crayfish my2 . This study attempted to
increase the density of broodstock in order to intensify production of young in indoor hatcheries.
Adult crayfish were stocked in 0.3 m2 tanks at a 1:5 M:F sex ratio at three densities: 6, 12 and 18
crayfish per tank corresponding to 20, 40 and 60 my2 .. The number of berried females was noted
at weekly intervals during 14 weeks. There was no difference among densities in survival rate and
percentage of spawning and molting, nor in clutch size pleopodal egg counts. of females. The
results indicated that breeding red claw females are insensitive to crowding over a wide range of
densities. They suggest that a broodstock density higher than 60 crayfish my2 is feasible under
indoor hatchery conditions. q 2000 Elsevier Science B.V. All rights reserved.
Keywords: Reproduction; Broodstock density; Crayfish; Cherax quadricarinatus

1. Introduction
Culture of the red claw crayfish Cherax quadricarinatus. in warm temperate
climates is constrained by a growth season of 57 months. Although this period may be
sufficient to attain marketable size crayfish Rouse, 1995.. To this end, however,
juveniles must be stocked into growout ponds as soon as water temperature is high
enough for growth. Development of hatchery and nursery systems is necessary to control
)

Corresponding author. Fax: q972-3-9605-667; E-mail: vlaqua@agri.gov.il

0044-8486r00r$ - see front matter q 2000 Elsevier Science B.V. All rights reserved.
PII: S 0 0 4 4 - 8 4 8 6 9 9 . 0 0 2 3 5 - 5

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A. Barki, I. Karplusr Aquaculture 181 (2000) 235240

the timing of stocking with young of appropriate size and quality, as well as to conduct
genetic programs.
Owing to the relatively low reproductive output of female crayfish, a large number of
brooders must be maintained in order to produce a significant amount of young. Since
space and water volume in indoor systems is limited, brooder density is an important
factor to consider in maximizing production of young. Low density of females is not
space efficient; however, reproductions at high densities may be adversely affected e.g.,
in Astacus leptodactylus, Sevilla, 1988.. The effect of crowding may be indirect through
its effect on environmental parameters such as oxygen, water quality and food availability, or crowding may act through increased competition and social interactions among
crayfish Lowery, 1988.. Crowding may increase competition for food and thus result in
poor nutrition for less successful females. This, in turn, reduces their egg production
Huner and Lindqvist, 1991.. Aggressive interactions may cause loss of eggs in
ovigerous red claw females Levi, 1997.. Furthermore, injuries and cannibalism mainly
of newly-molted females. may increase under crowding. While we can control many
environmental factors in hatchery systems, it is very difficult to eliminate social effects
of crowding.
Production of young red claw in Australia is usually practiced in earthen ponds at low
densities of 5001000 females hay1 Curtis and Jones, 1995.. However, red claw
crayfish also readily breed indoors. Previous studies addressing reproduction in red claw
under indoor conditions have employed broodstock densities ranging between 6 and 20
my2 , with sex ratios of 1:11:5 male:female Yeh and Rouse, 1994, 1995; Jones, 1995;
Barki et al., 1997; Sagi et al., 1997; Austin, 1998.. There was no evidence for a
reduction in spawning at a density as high as 20 my2 Yeh and Rouse, 1995.. These
studies imply that a higher density of broodstock is possible in indoor hatcheries.
The aim of this study was to extend previous results Yeh and Rouse, 1995. in an
attempt to increase density of broodstock and thereby to intensify production of young
in hatcheries.

2. Materials and methods

The experiment was conducted in nine 120-l plastic tanks, each with a bottom area of
0.30 m2 . Each tank contained two shelters, consisting of three plastic egg trays, one
above the other, with two pairs of PVC tubes between the trays. These shelters occupied
46% of the water volume. We have successfully used such shelters for red claw
reproduction in previous studies Barki et al., 1997; Sagi et al., 1997.. To avoid possible
effects between densities due to water-borne substances, the crayfish were held in three
separate recirculating water systems, one for each treatment. The water of each
treatment recirculated through a tank containing a biofilter, a thermostat-controlled
heater and a submerged pump. Minimum and maximum temperatures were similar in all
densities, averaging 26.2 " 0.7 and 27.6 " 0.98C. We tested reproduction using groups
of crayfish at three densities, 6, 12 and 18 crayfish per tank, in three replicates. A
male-to-female ratio M:F. of 1:5 was used in all densities, yielding groups of 1 and 5, 2

A. Barki, I. Karplusr Aquaculture 181 (2000) 235240

237

and 10, and 3 and 15, males and females, respectively. These densities correspond to 20,
40, and 60 crayfish my2 . The experimental tanks were stocked early in the breeding
season April 7, 1998. using sexually mature crayfish that have over-wintered in outdoor
earthen ponds. Mean "s.d.. female wet weight at stocking was 43.4 " 8.2 g and all
females were intact, unberried, and hard-shelled. The males were on average larger
51.1 " 6.3 g.. Each female was individually tagged using a colored tag glued to the
carapace. To maintain constant densities throughout the experiment, dead females and
males were replaced. The crayfish were fed 2% of biomass three times a week using
commercial trout pellets 40% protein, 4 mm in diameter. and once weekly with ground
carrots.
The experiment lasted 14 weeks. All tanks were inspected daily for the presence of
exuvia, and newly molted females were re-tagged. Once a week the shelters were
removed and all berried ovigerous. females were noted. Three weeks after a female was
first observed to be berried, her pleopodal eggs were carefully stripped, counted, and a
sample of five eggs from each clutch was examined under a stereo microscope for
viability and developmental stage. The embryonic stage of the eggs ranged between
stages 4 and 6 Jones, 1995.. In four females the eggs had hatched stage 7. before
stripping.
Only females that survived throughout the experiment were included in the analysis.
Differences in survival and percentage of berried females and molting of the original
number of females. were analyzed using one-way ANOVA on data transformed to
arcsine of the square root. The number of females per tank was used as a weighing
variable in these analyses. Difference in clutch size i.e., number of pleopodal eggs. was
analyzed using Analysis of Covariance ANCOVA. with the weight of female as a
covariate. Statistical analysis was carried out using JMP statistical package version
3.1.5.

3. Results
Survival of females declined mainly during the first 6 weeks and remained stable
after nine weeks. Mean survival rates ranged from 75 to 80% and were similar for all
densities P ) 0.05. Fig. 1..
First spawnings occurred between weeks 3 and 12 with peak spawning between
weeks 4 and 6 in all densities Fig. 2.. There was no difference among densities in the
cumulative percentage of first spawnings during the experiment, with mean values
ranging between 67 and 75.6% after 14 weeks P ) 0.05. Fig. 2.. Second spawnings
occurred from week 11. The cumulative percentage of second spawnings was similar
among all densities P ) 0.05. 23.326.7% of the females. Fig. 2..
The percentage of females molting ranged from 20 to 30% after 14 weeks excluding
one exceptional tank where 3 of 5 females molted., with no significant difference among
densities P ) 0.05..
ANCOVA revealed high dependence of the number of pleopodal eggs on female
weight P - 0.001. and no significant density-weight interaction effect P ) 0.05.,
indicating a similar slope of regression line for all densities. There was no significant

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A. Barki, I. Karplusr Aquaculture 181 (2000) 235240

Fig. 1. Mean percentage survival of C. quadricarinatus females of the original number of females. held in 0.3
m2 tanks at a 1:5 M:F sex ratio during 14 weeks, for each of three densities total MqF per m2 ..

difference among densities in clutch size after eliminating the effect of weight P ) 0.05..
The mean size-standardized clutch size across densities was 7.5 " 2.7 eggs gy1 of
female, which is consistent with other reports on red claw see Austin, 1998 and
references therein..

Fig. 2. Mean cumulative spawning percentage of the original number of females. for first spawnings upper
lines. and second spawnings of C. quadricarinatus females during 14 weeks, held in 0.3 m2 tanks at a 1:5
M:F sex ratio, for each of three densities total MqF per m2 ..

A. Barki, I. Karplusr Aquaculture 181 (2000) 235240

239

4. Discussion
The results clearly indicated that increasing density in the 2060 crayfish my2 range
did not influence spawning rate and fecundity, nor did it affect survival of females. This
finding holds for a density that is three-fold higher than densities reported earlier for
reproduction in red claw Yeh and Rouse, 1995; Sagi et al., 1997.. Thus, red claw
brooders seem to be insensitive to crowding over a wide range of densities. Although we
focused on one breeding cycle, the fact that a similar proportion of females spawned for
the second time during the last 3 weeks of the experiment in all densities suggests that
the lack of density effect would have extended for the whole breeding season.
In order to determine the optimal density for breeding, the tested range should
include a threshold density above which a reduction in egg production per female is
evident. The highest tested density 60 crayfish my2 . apparently was within the range
where the net return from increasing density did not decline. Thus, this density was less
than, or equal to, optimum. Furthermore, in indoor breeding systems where water
volume is a limiting factor, density of females may exceed optimum at a cost of
reduction in mean production per female. as long as the total production per tank
increases. Thus, it seems feasible to crowd red claw females to a density higher than 60
my2 . Studies on reproduction in other crayfish species have usually reported stocking
densities not exceeding 50 my2 e.g., Dube and Portelance, 1992; Sevilla and Tubert,
1993; Carral et al., 1994; Mitchell and Collins, 1995.. At higher densities reproduction
decreased substantially Sevilla, 1988., and aggression mainly of males. increased,
leading to mortality Woodlock and Reynolds, 1988.. The above studies involved
crayfish that were of smaller size than Cherax quadricarinatus.
The mean number of pleopodal eggs per tank obtained under the high-density
condition 60 my2 . was 3710 for one breeding cycle during 12 weeks, which is 12,367
eggs my2 . The number of the eventual independent young would have been smaller than
pleopodal egg counts Huner and Lindqvist, 1991; Yeh and Rouse, 1994; Austin, 1998..
On the other hand, the number of eggs that may be produced in our system is potentially
higher than 12,367 my2 , since we did not include in the calculation the substitute
females. Furthermore, larger females bearing larger clutch sizes may have been used.
Since the results refer to the specific environmental conditions that prevailed in our
system e.g., size of tank and type of shelters., extrapolation to larger tanks of females
of a different size should be made with caution. The use of small tanks in indoor
breeding systems has an advantage over large ones since they facilitate individual
monitoring of the crayfish provided they are tagged.. This enables efficient management of brooders by tracking the reproductive history of individuals and replacing
non-fecund and berried females, thus continually maintaining high production. On the
other hand, very small tanks housing individual mating pairs may yield lower spawning
rates than large tanks with grouped crayfish, as has been demonstrated with a related
species, C. albidus Mitchell and Collins, 1995..
Finally, it is possible that the sex ratio used in this study and previous ones 1:5 M:F.
may be reduced to increase the number of females at a given total density without
damaging the reproduction, thus increasing overall production of young. For example, a
single male noble crayfish A. astacus . was able to fertilize 23 females on average

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A. Barki, I. Karplusr Aquaculture 181 (2000) 235240

during 3040 days Svensson and Gydemo, 1997.. Although the potential gain from
changing the sex ratio is lower than from increasing the density, it may further
contribute to maximization of production of young in the hatchery.
Acknowledgements
We wish to thank the reviewers for their suggestions on the earlier version of the
manuscript. This study was supported by the Chief Scientist, Ministry of Agriculture and
Rural Development, grant number 358.0285.
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