Acta Zoologica (Stockholm) 87: 239–245 (October 2006)

Comparative fecundity and oocyte size of Aristaeomorpha

foliacea and Aristeus antennatus in the Greek Ionian Sea
(E. Mediterranean) (Decapoda: Aristeidae)
Kostas Kapiris1 and Maria Thessalou-Legaki2

1
Hellenic Centre for Marine Research,
Institute of Marine Biological Resources,
Aghios Kosmas, Hellinikon, GR-167 77,
Athens, Greece; 2Department of Zoology-
Marine Biology, School of Biology,
University of Athens, Panepistimioupolis,
GR-157 84, Athens, Greece
Keywords:
Aristaeomorpha foliacea, Aristeus antennatus,
fecundity, Ionian Sea
Accepted for publication:
8 June 2006
Abstract
Kapiris, K. & Thessalou-Legaki, M. 2006. Comparative fecundity and oocyte
size of Aristaeomorpha foliacea and Aristeus antennatus in the Greek Ionian Sea
(E. Mediterranean) (Decapoda: Aristeidae). — Acta Zoologica (Stockholm) 87:
239–245
Absolute and relative fecundity of 49 ripe gonads (from the cephalic and the
abdominal sections of the same specimen) of Aristaeomorpha foliacea and 71
ripe gonads of Aristeus antennatus caught in the Ionian Sea (E. Mediterranean)
and their oocyte size were studied. Both average absolute and relative
fecundities estimated for A. antennatus were statistically higher than those
estimated for A. foliacea; the average absolute fecundity of A. foliacea was
151 936 oocytes/female and that of A. antennatus was 200 472 oocytes/female
and the average relative fecundity of A. foliacea was 5477 oocytes/g and that of
A. antennatus was 9386 oocytes/g. Fecundity in both red shrimp species is
linearly related to carapace length, body and gonad weight, although, the most
precise estimation is provided by the relationship of body weight and fecundity.
In the present study, the oocyte size (diameter, volume) of both species could
be considered very large. In conclusion, A. foliacea shows lower fecundity than
A. antennatus, having a lower ovogenetic activity, but it produces larger
oocytes. This is in accordance with the highest value of the gonad weight and
gonad volume of A. foliacea exhibited in relation to A. antennatus.
K. Kapiris, Hellenic Centre for Marine Research, Institute of Marine
Biological Resources, Aghios Kosmas, Hellinikon, GR-167 77, Athens,
Greece. E-mail: kkapir@ath.hcmr.gr

in the Greek Ionian Sea, A. foliacea is much more frequent

Introduction
The deep-water red shrimp populations (Aristaeomorpha
foliacea (Risso 1827) – giant red shrimp – and Aristeus antennatus
(Risso 1816) – red shrimp) are the most economically and
ecologically important deep-water crustacean resource in the
Mediterranean basin, after Nephrops norvegicus. Recently, it
has been shown that these species have the potential to
support a viable fishery in the Greek Ionian Sea (Eastern
Mediterranean) (Papaconstantinou and Kapiris 2001,
2003). Moreover, these species are unpopular to Greek
consumers because (1) fishing in these depths yields a relatively
low economic return; (2) knowledge and gear technology are
limited; and (3) they have a low local value, because they are
unknown to Greek consumers. Regarding their abundances
than A. antennatus. This higher abundance seems to be
related to the particular ecological conditions of this area,
which satisfy better the niche demands of A. foliacea (Papa-
constantinou and Kapiris 2003).
Estimating fecundity in crustaceans can be a very tedious
and time-consuming procedure but is advantageous in
reproductive studies (see Caddy 1990). It is an important
parameter in crustaceans, among others, when determining
the reproductive potential of a species and/or the stock size
of a population (Mantelatto and Fransozo 1997).
Reproduction in Penaeoid shrimps is considered primitive
because fertilized eggs are not brooded but spawned free into
the water. Very scarce data exist on the fecundity and oocyte
size for both red shrimp species in the literature but they are

© 2006 The Royal Swedish Academy of Sciences 

No claim to original US government works
Fecundity of Aristaeomorpha foliacea and Aristeus antennatus • Kapiris et al.
limited to the western and central parts of the Mediterranean
(Relini-Orsi and Pestarino 1980; Orsi Relini 1982; Orsi
Relini and Semeria 1983; Demestre 1990, 1995; Yahiaoui
1994; Kao et al. 1999). This is the first attempt to estimate
the fecundity of both species in the Eastern Mediterranean
area. The purposes of this investigation were (1) to estimate
the fecundity of both species as well as their oocyte size;
(2) to relate their fecundity to carapace length, body and
gonad weight; and (3) to compare their fecundities to provide
basic information concerning their reproductive strategies.
Materials and Methods
The individuals were collected on a monthly basis from
December 1996 to November 1997 in the Greek Ionian Sea
between the Peloponnisos and Zakynthos Island (Fig. 1), in
the frame of the research project ‘Deep waters’, in a depth
range from 446 to 728 m, using a commercial trawl net
(18 mm at the cod end). In the laboratory, the specimens –
preserved in formalin – were sexed, the size of female
shrimps (carapace length, CL, mm) was measured to the
nearest 0.01 mm using calipers, and the body weight (BW, g)
was also measured (with 0.001 g accuracy). Four stages of
gonadal development were established for female shrimp
according to the empirical scale of Levi and Vacchi (1988) for
A. foliacea and the empirical scale of Relini-Orsi and Relini
(1979) for A. antennatus. Then, only the ripe ovary (stage IV)
Fig. 1—Study area and the sampling stations in the Greek Ionian
Sea. —A. Isobath of 800 m —B. Study area.

Acta Zoologica (Stockholm) 87: 239– 245 ( October 2006)
was removed and weighed. The ovary weight (GW, g) was
measured on a digital balance to an accuracy of 0.0001 g.
Gonad volume (GV, mL) was also estimated using the
gravimetric method. After removal of the ovaries, double
sub-samples were taken from the cephalic and abdominal
sections of the same specimen to determine whether the
gonad maturation was simultaneous and homogeneous in all
its regions. These sampling units were enough to estimate the
number of oocytes for each female.
Because quantification of oocytes is only feasible after
separation from connective tissue, many attempts have been
made. In some cases, the use of chemicals has resulted in the
hardening of the ovary tissue which made the separation of
the oocytes much more difficult. In other cases, the oocytes
were destroyed, using several chemical compounds: Gilson’s
solution (Simpson 1951; Staples 1975) Bouin’s solution
(Martosubroto 1974), acetic acid (Barbin and McCleave
1997) and other methods ( Jons and Miranda 1997). After
considering the literature and after personal communication
with several researchers the following technique was used: a
preweighed sub-sample of gonad was transferred to a jar, to
which 5–10% NaClO– solution in seawater was added. After
that, the jars were shaken for 1 h and then the oocytes usually
came cleanly away from the ovarian tissue. This proved to be
an adequate methodology in this red shrimp fecundity study.
The fecundity of the species was estimated from 49 ripe
gonads of A. foliacea and 71 ripe gonads of A. antennatus.
The absolute fecundity (AF) was estimated according to the
formula AF = number of eggs × (GW/weight of sample)
(Bagenal and Braum 1978) and the relative fecundity (RF)
was calculated using the formula: RF = (AF × 100)/BW. The
relative fecundity can be used to compare all individuals
regardless of their size. However, many studies of fecundity,
in fish, have shown that relative fecundity may vary with
the size of individuals. In this situation relative fecundity
is useful only as an indicator of maturation stage of the
individuals. To estimate the volume of the mature oocytes
(stage IV) their diameters were measured with an ocular
micrometer in 32 samples of A. foliacea and 79 samples of
A. antennatus and the oocyte volume were calculated as
spheres in A. antennatus (V = 4/3πR3) (where R is the radius)
and as cylinders in A. foliacea (V = πR2υ), where R is the
radius of the base and υ is the height of the cylinder (Orsi
Relini and Semeria 1983).
The relationship between absolute, relative fecundity/CL,
BW, GW and GW/CL, BW, GV was expressed in terms of
linear regression F = a + bX and the power regression
F = aX b, where F = fecundity or GW, X = CL, BW, GV,
a = a constant and b = an exponent (slope), which is about
3 when fecundity is related to length and about 1 when
fecundity is related to BW and GW (Rhode and Ross 1987).
Both regressions are more generally used to describe fecundity.
It was found that in all cases the linear regression fitted the
data better than the power function in terms of the correlation
coefficient (r), so only the former were used. A logarithmic
© 2006 The Royal Swedish Academy of Sciences
No claim to original US government works
Acta Zoologica (Stockholm) 87: 239–245 (October 2006) Kapiris et al. • Fecundity of Aristaeomorpha foliacea and Aristeus antennatus

Table 1 Comparison between somatic and

reproductive characteristics of Characteristic A. foliacea A. antennatus
Aristaeomorpha foliacea and Aristeus
CL (mm) ± SD (n) 44.61 ± 5.25 (52) 40.63 ± 5.96 (75)
antennatus from the North Greek Ionian Sea
BW (g) ± SD (n) 28.63 ± 9.75 (52) 22.34 ± 7.8 (75)
GW (g) ± SD (n) 2.65 ± 1.21 (52) 1.76 ± 1.18 (75)
GV (mL) ± SD (n) 2.5 ± 11.11 (46) 1.79 ± 1.19 (66)
Absolute fecundity (oocytes/female)
Abdomen 137 963 ± 132 873 (49) 190 131 ± 156 509 (71)
Cephalothorax 165 908 ± 115 140 (49) 207 618 ± 205 977 (71)
Median 152 940 ± 103 276 (49) 199 528 ± 35 282 (71)
Relative fecundity (oocytes/g)
Abdomen 5030 ± 3258 (49) 9321 ± 7801 (71)
Cephalothorax 5923 ± 3855 (49) 9390 ± 5711 (71)
Median 5423 ± 3501 (49) 9390 ± 5711 (71)
Ooocytes
Abdominal lobe, diameter (mm) 0.398 ± 0.040 (32) 0.327 ± 0.036 (79)
Cephalic lobe, diameter (mm) 0.410 ± 0.051 (32) 0.333 ± 0.045 (79)
Abdominal lobe, volume (mm3) 0.034 ± 0.012 (32) 0.019 ± 0.006 (79)
Cephalic lobe, volume (mm3) 0.035 ± 0.015 (32) 0.020 ± 0.006 (79)

CL = carapace length, BW = body weight, GW = gonad weight, GV = gonad volume, SD = standard

deviation, n = number of specimens.

transformation of the data was applied to satisfy the normality Table 2 Relationships between log values of absolute fecundity (AF),
assumptions (Sokal and Rohlf 1981). Differences between relative fecundity ( RF ) , gonad weight (GW) and carapace length (CL)
the slopes of the regressions were examined by analysis of
Variable b SE a n F r
covariance (). To test the slope b of the obtained
regression equations against the isometric slope of 1 or 3
Aristaeomorpha foliacea
Student’s t-test was applied. AF/CL 2.09 0.82 1.66 51 6.47 0.37**
AF/BW 0.82 0.30 3.92 51 7.38 0.39***
AF/GW 0.44 0.21 4.93 51 4.31 0.31**
Results RF/CL −1.36 0.90 5.90 42 2.31 NS
The average ovarian weight (GW) for 52 mature female RF/BW −0.46 0.33 4.31 42 186 NS
RF/GW 0.23 0.18 3.59 42 1.00 NS
A. foliacea (stage IV) was 2.65 g (range 0.8 – 5.8) and for 75
GW/CL 1.83 0.48 −2.64 51 17.27 0.47***
mature female A. antennatus was 1.76 g (range 0.47–6.72) GW/BW 0.69 0.18 −0.61 51 14.43 0.47***
(Table 1). It is worth noting that GW correlated positively GW/GV 0.81 0.04 0.08 45 328.16 0.94***
and significantly with CL, BW and GV in both species Aristeus antennatus
(Table 2). However, the median ripe GW of A. antennatus was AF/CL 1.52 0.51 2.76 65 8.76 0.35***
significantly lower than that of A. foliacea (Mann–Whitney AF/BW 0.67 0.21 4.32 65 9.74 0.37***
test, P < 0.001) and therefore the median GV of A. antennatus AF/GW 0.65 0.16 5.05 65 16.44 0.47***
(range 0.5–6 mL) was also significantly lower than that of RF/CL −0.85 0.52 7.25 65 2.71 NS
A. foliacea (range 0.5 –5 mL) (Mann–Whitney test, P < 0.001) RF/BW −0.36 0.22 6.36 65 2.84 NS
RF/GW 0.16 0.13 5.86 65 1.34 NS
(Table 1). The linear regression (Table 2) shows that the
GW/CL 2.18 0.34 −3.31 74 39.61 0.59***
ovary weight of A. foliacea increases by 0.69 g for each 1 g GW/BW 0.91 0.14 −1.03 74 40.16 0.60***
increase in BW and 1.83 g for each 1 mm increase in CL. GW/GV 0.90 0.04 0.03 65 436.17 0.93***
In A. antennatus these increases were 0.91 g and 2.18 g,
respectively. Body weight (BW), gonad weight (GW) and gonad volume (GV) for both red
Of 17 148 female A. foliacea and 7272 female A. antennatus shrimps according to the F = a + b * X model; b = regression coefficient
(slope); SE = standard error of b; a = intercept; n = number of individual
caught in the Ionian Sea (E. Mediterranean), the fecundity
females examined; F = variance ratio; r = correlation coefficient; **P < 0.05.
of 49 mature females of A. foliacea spanning 36.4 – 61.54 mm
***P < 0.01. NS = non-significant.
CL and the fecundity of 71 females of A. antennatus spanning
27.14–53.26 mm CL was studied (Fig. 2). The CL range of
specimens examined for estimation of fecundity was wide of individuals examined for fecundity encompasses most of
and representative, taking into account that the smallest the mature females in the population. The medians and
mature A. foliacea female was 36 mm and that of A. antennatus the distributions of the absolute as well as of the relative
was 25.59 mm (Kapiris 2004). This suggests that the range fecundity estimated from samples taken from the abdominal

© 2006 The Royal Swedish Academy of Sciences 

No claim to original US government works
Fecundity of Aristaeomorpha foliacea and Aristeus antennatus • Kapiris et al.
Fig. 2—Carapace (CL) size distribution (in mm, size interval:
1 mm) of the specimens of Aristaeomorpha foliacea (white bar) and
Aristeus antennatus (grey bar), which the fecundity was estimated.
lobe did not differ statistically from the median fecundity of
samples taken from the cephalic lobe in both shrimps
(Mann–Whitney test, Kolmogorov–Smirnov test, P > 0.05).
The average absolute fecundity from the abdominal lobe
was 137 963 oocytes (range: 5919 – 681 095), while the same
fecundity estimated from samples of gonad taken from the
cephalic lobe was 165 908 oocytes (range: 19 858 –503 659),
for A. foliacea. In A. antennatus absolute fecundities were
190 131 oocytes (range: 22 219 –967 333) from the abdominal
lobe and 210 812 oocytes (range: 30 149–1269 416). Thus,
data from the two lobes could be pooled. The median of both
absolute and relative fecundities (abdominal, cephalic lobe)
was used. The average absolute fecundity of A. foliacea was
151 936 oocytes/female and that of A. antennatus was 200 472
oocytes/female. On the other hand, the average relative
fecundity in A. foliacea for samples from the abdominal lobe
was 5031 oocytes/g (range: 149–32 939) but from oocytes
taken from the cephalic lobe it was 5923 oocytes/g (range:
437–16 666). In A. antennatus, the estimated relative fecundity
was 9321 oocytes/g (range: 888–32 920) from the abdominal
lobe and 9451 oocytes/g (range: 1537–31 352) from the
cephalic lobe (Table 1). Thus, the average relative fecundity
of A. foliacea was 5477 oocytes/g and that of A. antennatus
was 9386 oocytes/g. The fecundity values estimated for
A. antennatus were statistically higher than those estimated
for A. foliacea (Mann–Whitney test, P < 0.001), median values
included.
Best fits for relationships of absolute and relative fecundity
and CL, BW and GW were given by linear equations
(Table 2). The slope of the equation AF/CL in A. foliacea was
not found to be significantly different from the theoretical
slope of 3.0 (t-test, P > 0.05). Similarly, the slopes of the
equations AF/BW, in both species, were not found to be
significantly different from the theoretical slope of 1.0 (t-test,
P > 0.05) (Table 2). The relationships between AF/CL, BW,
GW were significant in both species (P < 0.05, in most
cases), in spite of the weak correlations (Fig. 3A). Predicted
absolute fecundity ranged from 56 234 and 102 093
oocytes for A. foliacea and A. antennatus female 30 mm CL,
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Acta Zoologica (Stockholm) 87: 239– 245 ( October 2006)
respectively, to 162 181 and 219 785 oocytes for A. foliacea
and A. antennatus female 50 mm CL, respectively. As the
intercept of the regression of AF against weight was positive
(Table 2), the relative fecundity (oocytes/g) declined with
increasing CL and BW, in both species (Fig. 3B), but the
very low correlation coefficients (r) between RF and CL, BW,
GW in both species, indicated the absence of a significant
relationship between size, weight and relative fecundity.
Differences in slopes between the AF, RF, GW/CL, BW,
GV regressions of the two species were tested using ANCOVA.
All differed significantly between the two species (P < 0.05).
Female A. antennatus, for the same length, showed higher
fecundity values than female A. foliacea.
The mean value and the standard deviation of the diameter
and volume of the oocytes of the two species from the two
different gonad regions are given in Table 1. In both species,
no statistically significant difference was found in oocyte size
between the two different gonad regions (Mann–Whitney,
Kolmogorov–Smirnov tests, P < 0.05). In contrast to the
fecundity results, the oocyte size (diameter and, consequently,
volume) was in favour of A. foliacea (Mann–Whitney test,
P < 0.05).
Discussion
A remarkable synchronization and seasonality of reproductive
biology – maturing, mating and release of oocytes of both
studied species and sexes – were shown in the Ionian Sea
(Kapiris 2004). Many similarities, instead of differences,
between the reproductive strategies of both shrimps, were
exhibited. The most crucial reproductive difference between
the two species was that A. antennatus is already sexually
active from its first year of life, instead of the second year of
life as in A. foliacea, having a greater ovogenetic activity than
A. foliacea. In both species, the processes of mating, spawning
and moulting have to be very closely connected, in space and
in time, in such a way that they all depend on each other. The
medium and larger females of both species were the first to
reach sexual maturity, at the start of the reproductive season
(Kapiris 2004) and, according to the relationship between
AF/CL, BW, in the present study, it is clear that these
individuals produce more oocytes that the smaller shrimps.
This seems to agree with the findings of other authors
(e.g. Mori et al. 1994; Ragonese and Bianchini 1995, for
A. foliacea; Relini-Orsi 1980; Relini-Orsi and Pestarino
1980, for A. antennatus) and is commonly accepted for
penaeids (e.g. Menasveta et al. 1994). This more extended
reproductive period could be attributed to the possibility that
the larger specimens carry out one spawning per month (at
least three consecutive spawnings in the same reproductive
season).
Fecundity in red shrimps is linearly related to CL, BW and
GW. These results are in agreement with previous fecundity
studies in other decapods [e.g. in pandalids (Company and
Sardà 1997) in crayfishes (Corey 1987) and in prawns
© 2006 The Royal Swedish Academy of Sciences
No claim to original US government works
Acta Zoologica (Stockholm) 87: 239–245 (October 2006) Kapiris et al. • Fecundity of Aristaeomorpha foliacea and Aristeus antennatus
Fig. 3—Relationship between carapace
length – absolute fecundity (oocytes/
female) (A) and between carapace length –
relative fecundity (oocytes/g) (B) in
Aristaeomorpha foliacea (dotted line)
and Aristeus antennatus (solid line).
(Baelde 1992)]. According to the regression coefficient
value, CL, BW and GW could be used to predict absolute
fecundity of both species but the most precise estimation of
fecundity is provided by the relationship of BW with fecundity,
suggesting that increases of somatic weight could be accom-
panied by proportional increases of the oocyte number. The
ovary weight could also be used, like BW, as an adequate
index of fecundity in both species because it is affected by
oocyte number and size concurrently. However, it is easier
to use BW or CL to estimate fecundity, because these are
simpler to measure than ovary weight.
Bearing in mind that both species present three successive
spawnings in the same reproductive period (Orsi Relini
1982; Orsi Relini and Semeria 1983; Follesa et al. 1998, for
A. foliacea and Orsi Relini 1980; Demestre 1990, for A.
antennatus), the total yearly amount of oocytes in A. foliacea
could be 458 820 oocytes/female and 16 269 oocytes/g.
Similarly, in A. antennatus the absolute fecundity could be
598 584 oocytes/female and the relative fecundity could be
28 170 oocytes/g. The estimated fecundity for both species
was situated among the highest estimated values of penaeid
species, because the maximum reported fecundity ranges
from 250 000 to 950 000 oocytes (e.g. Penn 1980). Since the
© 2006 The Royal Swedish Academy of Sciences
No claim to original US government works
slopes of the regressions between GW/CL, BW were higher
in A. antennatus and its oocyte diameter was lower than that
of A. foliacea oocytes, it is reasonable to expect a higher
number of oocytes for A. antennatus than for A. foliacea.
Calculations of fecundity of both species in this paper
are greater than, lower than or similar to those of other
studies reported in the literature and estimated for other
Mediterranean regions, but, unfortunately, there are no
regression data. In this way, the fecundity of both species
estimated in the Ligurian Sea was significantly higher than
our estimates. Orsi Relini and Semeria (1983) found that in
the Ligurian Sea the fecundity of A. foliacea ranged between
100 000 and 250 000 oocytes and that for A. antennatus
ranged between 140 000–900 000 oocytes in each ovulation.
Moreover, Orsi Relini (1982) pointed out that in the Ligurian
Sea the number of oocytes produced by A. antennatus in a
single ovulation ranged more frequently from 420 000 to
980 000, reaching peaks of 1.5 million. Relini-Orsi and
Pestarino (1980) suggested that an A. antennatus female of
56 mm CL contained 600 000 oocytes in the whole repro-
ductive period. Other studies have overestimated the fecundity
of A. antennatus in comparison to the present study. In the
present study, a female A. antennatus of the same length

Fecundity of Aristaeomorpha foliacea and Aristeus antennatus • Kapiris et al.
(56 mm) was estimated to have 269 153 oocytes. In other
areas the fecundity was underestimated. In Algeria, for
instance, the relative fecundity of A. antennatus was estimated
to be 8000 oocytes/g (Yahiaoui 1994). Our fecundity data
coincide fairly well with those reported by Demestre (1990,
1995) who reported that in the Catalan Sea A. antennatus
released 46 000–606 000 oocytes per ovulation and, in total,
released 138 000–1 800 000 oocytes per reproductive period
(Demestre 1990, 1995). It is worth noting that the same
author counted a significantly higher number of oocytes in
A. antennatus coming from the thoracic region than the
cephalic or the abdominal regions. It is well known that the
fecundity and oocyte size of aquatic invertebrates vary widely
and related species from different habitats or parts of the
world may have oocytes that vary in size (Clarke 1993).
Differences in estimates of fecundity among studies might be
artificial, as the result of differences in technique, or natural,
as the results of differences in fecundity with geographical
location or climatic conditions. Additionally, the different
geomorphological, trophic and fishing efforts in the different
areas could explain these differences (Martosubroto 1974).
However, further investigations including all Mediterranean
regions are needed to deepen our knowledge of their repro-
ductive patterns.
The egg size of aquatic invertebrates varies widely between
even quite closely related species (McEdward 1991). Com-
parative studies of related taxa with different egg or oocyte
size have frequently demonstrated a covariation between egg/
oocyte size and fecundity (Clarke 1993). Since the maximum
mean oocyte diameter of penaeids ranges from 0.25 to
0.35 mm (Tan-Fermin and Pudadera 1989), the size of the
oocyte of both species of the present paper could be considered
very large. The inverse relationship between fecundity and
oocyte size shown by these two deep-water sympatric
shrimps of the South Ionian Sea indicates that they might
follow different reproductive strategies. Thus, A. foliacea
produces fewer, but bigger, oocytes than A. antennatus. This
is in accordance with the highest value of the GW and GV
that the former species exhibited in relation to the latter.
The size of the mature oocytes in the present study does
not differ significantly from those reported by other studies.
Thus, Kao et al. (1999) reported that the mature oocytes of
A. foliacea had a diameter 0.42–0.44 mm and a volume
0.038–0.043 mm3. According to Orsi Relini and Semeria
(1983), the oocyte volume of A. foliacea was 0.024 mm3.
Demestre and Fortuño (1992) reported that the diameter of
the mature oocytes of A. antennatus in the Catalan Sea
ranged from 168 to 330 µm, whereas in Algeria the diameter
ranged between 217 and 301 µm (Yahiaoui 1994). Since
there was no difference in size for oocytes taken from two
different gonadal regions, it could be assumed that the
maturation of the ovary is simultaneous and similar in all
areas. The homogeneous development of ovaries is generally
assumed in penaeid shrimps, but a more detailed analysis is
needed (Ceballos-Vasquez et al. 2003).

Acta Zoologica (Stockholm) 87: 239– 245 ( October 2006)
It is generally accepted that a higher fecundity denotes
a higher mortality rate of young and immature individuals
(Corey 1987). If this is in force, A. antennatus would expect
a high loss of young individuals and A. foliacea would have
lower juvenile mortalities. Moreover, the larger oocytes,
having more yolk, tend to produce larger or more advanced
larvae (Clarke 1993).
This study represents basic data for the two aristeids in the
Ionian Sea; these date were not available in the literature and
can be used in population studies. The data can be used for
elaborating population models that will serve to assess
the effect of the fishery on the reproductive output and to
establish management strategies focused to achieve optimal
exploitation of the red shrimps stocks in this Mediterranean
region.
Acknowledgements
The material was collected during sampling cruises within
the research programme ‘Deep Water Fisheries’ partly
supported by the EU (FAIR 95-0655). Many thanks are due
to Dr C. Papaconstantinou and Mr G. Petrakis (Hellenic
Centre for Marine Research), who are responsible for the
above program in Greece. We also thank all the anonymous
referees whose suggestions improved this manuscript.
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