Fishery and Biology of Aristaeomorpha foliacea (Risso, 1827) (Crustacea: Decapoda) in the Northern Tyrrhenian Sea (Western Mediterranean)
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J. Northw. Atl. Fish. Sci., Vol. 31: 195204
Fishery and Biology of Aristaeomorpha foliacea
(Risso, 1827) (Crustacea: Decapoda) in the Northern
Tyrrhenian Sea (Western Mediterranean)
P. Belcari and C. Viva
Dipartimento di Scienze dell'Uomo e dell'Ambiente
Università di Pisa, Via Volta 6, 56126 Pisa, Italy
M. Mori
Dipartimento Territorio e Risorse, Università di Genova, Genova, Italy
and
S. De Ranieri
Centro Interuniversitario di Biologia Marina ed Ecologia Applicata, Livorno, Italy
Abstract
The aim of the study was to provide information about the fishery, population size structure,
breeding period and size at maturity of giant red shrimp, Aristaeomorpha foliacea, of the northern
Tyrrhenian Sea, Italy. Data were collected directly at the local auction of Porto Santo Stefano, the
main landing port of the study area, from 1991 to 1998 and during monthly surveys carried out
during the year 1995. In Porto Santo Stefano the annual landing of red shrimps ranged between 2
and about 17 tons, corresponding to an annual average first sale income of 170200 000 Euros.
The fishery activity depended on season and vessel and the monthly landings per unit effort ranged
from less than 1 to 35 kg/day/boat, although the catch of a single boat could reach values up to 80
kg/day. Size composition of A. foliacea ranged from 14 to 46 mm carapace length (CL) for males
and from 14 to 66 mm CL for females. Recruitment occurred mainly in spring. The overall sex-
ratio observed (1.10:1) was significantly biased towards females. Females begin to be mature in
March, reaching a peak between July and August. From October to March most females had rest-
ing gonads. Mature males were found all year round. Females with spermatophores on the thelycum
were found throughout the year, the highest percentages being recorded from March to Septem-
ber. The average size at sexual maturity was 40.59 mm CL. The average size of maximum repro-
ductive potential was 37.15 mm CL.
Key words: Aristaeomorpha foliacea, Crustacea, Decapoda, Mediterranean, red shrimp fishery,
reproductive biology, Tyrrhenian Sea
Introduction of the dynamics of the reproductive process, which
constitutes the basis of resource renewal, provides the
One goal in studies on reproductive ecology of suitable background for management direction
benthic crustaceans is to make and test generaliza- (Crocos, 1987).
tions about geographical variations in patterns of re-
production and recruitment. Such paradigms are use- Because of its importance on the bathyal fishery
ful for generating hypotheses about the specific envi- grounds of the Mediterranean Sea, a number of pa-
ronmental stimuli (proximate factors) and selective pers have addressed the biology, ecology and fishery
pressures (ultimate factors) that are responsible for of the giant red shrimp Aristaeomopha foliacea
breeding and recruitment patterns observed, as well (Risso, 1827). Most of the information relates to Ital-
as, for making predictions about possible changes in ian waters of the central Mediterranean (Cau and
reproduction and recruitment due to natural fluctua- Deiana, 1982; Cau et al., 1982; Ragonese, 1989; 1995,
tions or anthropogenic alterations in these factors Mura et al., 1992; Bianchini and Ragonese, 1994;
(Bauer and Rivera Vega, 1992). Further, knowledge Ragonese et al., 1994, 1997; Matarrese et al., 1995;196 J. Northw. Atl. Fish. Sci., Vol. 31, 2003
Ragonese and Bianchini, 1995; DOnghia et al., 1998; orbital margin to the mid posterior edge of the cara-
Bianchini et al., 1998; Bianchini, 1999). Only scanty pace. Males were distinguished from females by the
attention is devoted to the western basin (Lumare and presence of the petasma and assigned to two maturity
Utzeri, 1973; Bianchini and Ragonese, 1994). stages, according to the scale proposed by Sardà and
Demestre (1989) for the companion species Aristeus
The northern Tyrrhenian Sea (western Mediter- antennatus. The two stages, immature and mature,
ranean) is an important commercial fishery area for were based on the presence or absence of sperm sacs
bathyal decapod crustaceans and A. foliacea is the on the coxae of the fifth pair of pereiopods. Maturity
most valuable deep-water resource (De Ranieri et al., stages for females were assigned according to the
1994; De Ranieri, 1995; Ardizzone and Corsi, 1997; macroscopic colorimetric scale given by Levi and
Biagi et al., 1998). Despite this, very little informa- Vacchi (1988) and Kao et al. (1999), based on the
tion is available on the life cycle of the species (Mori different colours presented by the ovaries: flesh-
et al., 1994). The aim of present study was to provide colored, immature or spent, Stage I; light gray, ma-
information about the biology and fishery of A. turing, Stage II; dark gray, early mature, Stage III;
foliacea inhabiting the northern Tyrrhenian Sea in pale black, ripe, Stage IV. The presence of spermato-
order to carry out comparative regional studies and phore was checked by visual examination of the
pinpoint differences. thelycum.
Materials and Methods Maturity condition and presence of sperma-
tophores by size were estimated by fitting a logistic
The study was conducted at Porto Santo Stefano, equation; in order to estimate the percentage of ma-
Italy, the main landing point of the northern ture females in each size class, Stage III and Stage IV
Tyrrhenian Sea, western Mediterranean. Landing data were combined. On the basis of two functional crite-
of A. foliacea from the bottom trawl fleet were re- ria (Morizur, 1983), maturity of gonads and presence
corded during three to five days per month at the lo- of spermatophores, the size at the "maximum repro-
cal auction from 1991 to 1998. Monthly total land- ductive potential" was estimated. It was defined as
ings were estimated raising the sampled data to the the size corresponding to the intersection of the go-
total fishing days of the fleet in that month. The Land- nadic maturity curve and the mating complemen-
ing Per Unit of Effort (LPUE) was calculated consid- tary curve (i.e. absence of spermatophore) during
ering the fishing day as a unit of effort. Fishing effort the maximum reproductive activity of the stock
data were calculated monthly by consulting the offi- (Ragonese and Bianchini 1995; D'Onghia et al.,
cial archives. In the years 1995 to 1999 fishing activ- 1998).
ity was monitored, on a seasonal basis, by observa-
tions on board of commercial vessels fishing for Nor- Two moult stages were determined for both sexes,
way lobster and aristeid shrimps. In the sampled pe- intermoult and postmoult, depending on the state of
riod, information from about 500 trawling hours was the exoskeleton: completely calcified in the former
gathered. The collected data were analysed as hourly and at the beginning of calcification with a paper-
yields (kg/hr) and then studied on a seasonal basis. like consistency in the latter. Significant deviations
from the expected 1:1 sex ratio were tested using a χ2
Specimens of the giant red shrimp were collected test (Scossiroli and Palenzona, 1978).
by means of monthly sampling performed between
January and December 1995 off the Tuscan Archi- Results
pelago (northern Tyrrhenian Sea) at depths between
400 and 650 m. The study area is shown in Fig.1. The observations on board of trawlers of Porto
Samples were collected by trawlers using a profes- Santo Stefano allowed the identification of two fish-
sional trawl net with mouth height of about 1 m and ing grounds for red shrimps located on bottoms from
codend width of about 20 m. Mesh size was about about 450 to 650 m. The first is located westward of
40.0 mm stretched. During the months of February Pianosa and Montecristo Isles, the second between the
and December the sampling was not carried out be- Isles of Giglio and Montecristo (Fig. 1). Only a few
cause of bad weather. boats operate in the area westward of Pianosa and
Montecristo Isles. In the second area, the majority of
For each specimen, the carapace length (CL) was trawlers of medium-large size (up to 700 kW engine
measured, to the nearest mm below, from the right power) from Porto Santo Stefano carry out theirBELCARI et al.: Fishery and Biology of Aristaeomorpha foliacea 197
10°00' 10°30' 11°00'
50 m
Elba 100 m Grosseto
Northern Tyrrhenian Sea
Pianosa
200 m
42°30'
42°30'
Porto S. Stefano
Porto Ercole
Giglio
Montecristo
Giannutri
500 m
10°00' 10°30' 11°00'
Fig. 1. Study area in the western Mediterranean.
fishing activity, devoted to red shrimps and Norway The overall sex-ratio observed (1.10:1) was sig-
lobster, mostly during spring and summer. In both nificantly biased towards females (χ2 = 13.5; P198 J. Northw. Atl. Fish. Sci., Vol. 31, 2003
TABLE 1. Seasonal mean yields per hour of red shrimps from 1995 to 1999; SE = standard error.
Winter Spring Summer Autumn
Mean SE Mean SE Mean SE
A. foliacea 3.274 0.765 0.837 0.535 0.973 0.954
A. antennatus 0.505 0.061 0.324 0.301 0.053 0.051
40
1998
1997
kg/day/boat
30 1996
1995
20
1994
10 1993
1992
0
1991
J F M A M J J A S O N D
Fig. 2. Monthly LPUE of red shrimps A. foliacea and A. antennatus recorded at Porto S Stefano, the main landing
point of the northern Tyrrhenian Sea.
The reproductive aspects by size were evaluated March and November, before and after the reproduc-
pooling data from April to September, the period of tive season.
maximum reproductive activity. The size at which
50% of females were mature (Stage III and IV) was Discussion
40.59 mm CL, while 35.54 mm CL corresponded to
the size at which 50% of females had spermatophores In the northern Tyrrhenian Sea A. foliacea is one
(Fig. 7 and 8). Large females without spermatophores of the most valuable resources exploited by deep-sea
were very few; they had probably lost them acciden- fishery. Catches from fishing grounds at bottom depths
tally. The intersection of the logistic function for greater than 450 m showed that this was an impor-
maturity and for absence of spermatophores gave the tant target species. A. foliacea represented up to one
size of the "maximum reproductive potential" of 37.15 third, as biomass, of the total commercial catch, cor-
mm CL (Fig. 9). responding to a high economic income (first sale price
ranging from 20 to 30 Euros/kg). According to the
Moulting national data series recorded by ISTAT (Italian Cen-
tral Institute of Statistics), 43 505 tons of red shrimps
Moulting activity was performed all year long. were landed in the period 198596 (ISTAT, 198899).
Figure 10 shows the percentage of postmoulting males This commercial category includes the two species
and females caught during the monthly samplings. A. foliacea and A. antennatus, making it impossible
In both sexes, catches of postmoulting individuals to assess individual catch levels. Our investigation
were high in spring and autumn, and very low in sum- and experimental trawl surveys conducted over the
mertime. In males, moult took place mainly in May last fifteen years (Biagi et al., 1998) showed that in
and September; in females, moult was greater in the area studied, A. foliacea was much more abundantBELCARI et al.: Fishery and Biology of Aristaeomorpha foliacea 199
Females Males
25 40
20 January 32 January
15 n = 292 24 n = 108
10 16
5 8
0 0
10 15 20 25 30 35 40 45 50 55 60 65 10 15 20 25 30 35 40 45 50 55 60 65
25 40
20 March 32 March
15 n = 129 24 n = 69
10 16
5 8
0 0
10 15 20 25 30 35 40 45 50 55 60 65 10 15 20 25 30 35 40 45 50 55 60 65
25 40
20 April 32 April
15 n = 422 24 n = 257
10 16
5 8
0 0
10 15 20 25 30 35 40 45 50 55 60 65 10 15 20 25 30 35 40 45 50 55 60 65
25 40
20 May 32 May
15 n = 199 24 n = 207
10 16
5 8
0 0
10 15 20 25 30 35 40 45 50 55 60 65 10 15 20 25 30 35 40 45 50 55 60 65
Frequency (%)
25 40
20 July 32 July
15 n = 656 24 n = 666
10 16
5 8
0 0
10 15 20 25 30 35 40 45 50 55 60 65 10 15 20 25 30 35 40 45 50 55 60 65
25 40
20 August 32 August
15 n = 171 24 n = 362
10 16
5 8
0 0
10 15 20 25 30 35 40 45 50 55 60 65 10 15 20 25 30 35 40 45 50 55 60 65
25 40
20 September 32 September
15 n = 164 24 n = 122
10 16
5 8
0 0
10 15 20 25 30 35 40 45 50 55 60 65 10 15 20 25 30 35 40 45 50 55 60 65
25 40
20 October 32 October
15 n = 293 24 n = 372
10 16
5 8
0 0
10 15 20 25 30 35 40 45 50 55 60 65 10 15 20 25 30 35 40 45 50 55 60 65
25 40
20 November 32 November
15 n = 423 24 n = 323
10 16
5 8
0 0
10 15 20 25 30 35 40 45 50 55 60 65 10 15 20 25 30 35 40 45 50 55 60 65
Carapace length
Fig. 3. Size frequency distributions of female and male A. foliacea collected during
monthly samplings in the northern Tyrrhenian Sea. Samples that did not exceed
50 specimens were excluded.200 J. Northw. Atl. Fish. Sci., Vol. 31, 2003
than the companion species A. antennatus, with per- lack of investigation at greater depths in this area.
centages ranging from 72 to 99% of the total red These recent data on the eastern side of the Mediter-
shrimp catch. ranean confirm the general assumption that the abun-
dance of A. foliacea seems to follow longitudinal and
In other areas of the western Mediterranean the latitudinal gradients (Campillo, 1994; D'Onghia et
opposite trend was observed. In the eastern basin, al., 1998). However, a patchy distribution is also ob-
along the Mediterranean coast of Israel (Thessalou- served. Thus the species is scarce or absent in the
Legaki, 1994) and in Greek waters (Kapiris et al., Catalan Sea, in the Gulf of Lions and in the Ligurian
1999, 2001) a significantly higher abundance of Sea (Sardà, 1986, Campillo, 1994; Orsi Relini and
A. foliacea has been reported recently, suggesting that Relini, 1994), abundant in Tunisia (Ben Marien, 1994)
the scanty presence registered previously was due to and weakly present in Algeria (Yahiaoui, 1994), out-
numbers A. antennatus in the northern Tyrrhenian Sea
(Campillo, 1994; Biagi et al. 1998) and in the Sicil-
ian channel (Ragonese and Bianchini, 1995), is
1
present in almost equal proportions in the southeast-
ns
ern Tyrrhenian Sea (Greco et al., 1994) and again
PBELCARI et al.: Fishery and Biology of Aristaeomorpha foliacea 201
36 38 38 37 38 43 43 28 31 37
100
80
Frequency (%)
60
40
20
0
J F M A M J J A S O N D
Month
With spermatophores Without spermatophores
Fig. 6. Monthly percentage of female A. foliacea showing the presence or
absence of spermatophores. For each month, the minimum size at
which females bear spermatophores is indicated above the histograms.
1.0 been attributed to a close correlation between the pres-
R2 = 0.758 ence of the species and variations of hydrological fac-
0.8 tors. Changes in the water salinity rather than in tem-
Proportion
perature could explain its disappearance in certain
0.5
regions of the Mediterranean, as in the Ligurian Sea,
where the "Eastern or intermediate water" disappears
after mixing with the superficial and deep waters
0.3
(Murenu et al., 1994).
0.0 The marked seasonality of the landings observed
12 20 28 36 44 52 60 68 in this study is likely to be related above all to the
Carapace length (mm) fact that adverse weather conditions during winter
Fig. 7. Percentage by size of mature females (Stages III and often prevent fishing vessels from operating at a dis-
IV). tance from the coasts. Size frequency distributions
exhibit a wide size range, with females attaining larger
sizes than males. The distributions reflect those com-
1.0 puted in the same area with specimens gathered by
R2 = 0.923 means of experimental trawl surveys based on strati-
0.8 fied random samplings (Biagi et al., 1998). Percent-
Proportion
age of large individuals is very low. During the spring
0.5
months, an early component can be found, correspond-
0.3
ing to the cohort of trawl net recruits, as already re-
ported in the literature (Ragonese and Bianchini,
0 1995; D'Onghia et al., 1998; Spedicato et al., 1998).
12 20 28 36 44 52 60 68
Carapace length (mm)
Monthly variations in the sex ratio, slightly in
Fig. 8. Percentage by size of females with spermatophores. favour of females or males according to season, indi-202 J. Northw. Atl. Fish. Sci., Vol. 31, 2003
1.0
0.75
Obs. maturity
Exp. maturity
Obs. sperm absence
Proportion
Exp. sperm absence
0.50
0.25
0
10 14 18 22 26 30 34 38 42 46 50 54 58 62 66
Carapace length (mm)
Fig. 9. Size at "maximum reproductive potential" for female A. foliacea caught in the northern
Tyrrhenian Sea.
30 and September. Mating activity is observed all year
25
Females Males long, but the percentage of females bearing
spermatophores is lower during autumn and winter.
20 In the southeastern Mediterranean, the overall repro-
ductive period is similar, but the peak is shifted one
(%)
15
month earlier, in June (Kapiris and Thessalou-Legaki,
10 2001). Our findings on size at first maturity and at
maximum reproductive potential are similar to those
5
estimated for other areas (Ragonese and Bianchini,
0 1995; DOnghia et al.,1998).
J F M A M J J A S O N
Month In the northern Tyrrhenian Sea, the exploitation
Fig.10. Postmoulting males and females of A. foliacea caught of A. foliacea begins in the first juvenile stages or
in the northern Tyrrhenian Sea. before reaching the maximum reproductive potential,
as observed for other areas of the Mediterranean
(Ragonese et al., 1997; D'Onghia et al., 1998;
Spedicato et al., 1998; Bianchini, 1999). Studies pre-
cate very little segregation between sexes, although viously carried out in the northern Tyrrhenian Sea on
the overall sex ratio was biased toward females. The the exploitation state of A. foliacea indicated a con-
increase in number of males in late spring-summer is dition close to the equilibrium (De Ranieri, 1999).
ascribed to their rising out of canyons or from the These results and the recently observed reduction in
deepest areas for mating, as hypothesized by D'Onghia this area of the trawl fleet exploiting this resource,
et al. (1998). indicate that no further modification of the current
exploitation strategy is required.
Data on reproductive activity are consistent with
findings for the central Mediterranean (Cau and
Deiana, 1982; Cau et al., 1982; Mura et al., 1992;
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