Synthesis and structure of 1-ethyl-2,4,5-triphenyl-1H-imidazole (Ethyl-Lophine)

Article abstract of DOI:10.5560/ZNB.2013-2333

Ethyl-Lophine, 1-ethyl-2,4,5-triphenyl-1H-imidazole, C₂₃H ₂₀N₂, was synthesized as a precursor for large organic cations in ionic liquids using an improved microwave-assisted method. The title compound and a precursor compound were characterized by NMR, IR, and DSC thermal measurements, as well as elemental analyses. The crystal structure of ethyl-lophine was determined by singlecrystal X-ray structure analysis (triclinic, P1, a = 10:1137(3), b = 12:4935(4), c = 14:6351(4) A, α = 98:182(2), β = 90:694(2), γ = 102:666(2), Z = 4, wR2 = 0:1030 for 611 refined parameters).

Full text of DOI:10.5560/ZNB.2013-2333

A REVIEW OF THE BIOLOGY AND FISHERIES OF THE DEEP WATER
ROSE SHRIMP, PARAPENAEUS LONGIROSTRIS,
IN EUROPEAN ATLANTIC AND MEDITERRANEAN WATERS
(DECAPODA, DENDROBRANCHIATA, PENAEIDAE)
BY
1,5
2
3
4
IGNACIO SOBRINO ), CRISTINA SILVA ), MARIO SBRANA ) and KOSTAS KAPIRIS )
1
) Instituto Español de Oceanografía (Unidad de Cádiz), Muelle Pesquero s/n, Apdo. 2609,
E-11006 Cádiz, Spain
) Instituto de Investigação das Pescas e do Mar (IPIMAR), Av. de Brasília, s/n,
2
P-1449-006 Lisboa, Portugal
) Centro Interuniversitario di Biologia Marina, Viale N. Sauro, 4, I-57128 Livorno, Italy
3
4
) Hellenic Centre for Marine Research, Institute of Marine Biological Resources,
Agios Kosmas Helliniko, GR-166 04, Athens, Greece
ABSTRACT
The deep water rose shrimp, Parapenaeus longirostris, is a very important commercial species for
trawl fleets that fish in the Mediterranean as well as in the Gulf of Cadiz and to the south of Portugal.
Despite its importance, its life cycle has not been thoroughly studied and there are discrepancies as
to certain aspects, depending on the area. Although spawning takes place throughout the year, two
peak periods have been identified between late spring and early autumn. This species is characterized
by a differential growth pattern, with females reaching a larger size, and has a life cycle of two to
three years. Individuals reach sexual maturity at 8 to 10 months at varying sizes, depending on the
area (cephalothorax length 20 to 28 mm in females) and have an average fecundity of 90,000 eggs.
The species feeds on a great variety of prey, its main prey being annelid worms (polychaetes) and
foraminiferans. The fishery involves a large number of boats, about 2,000 over the whole area here
covered. It is basically directed at a single cohort and total catches can reach 10,000 tons a year.
RESUMEN
La gamba blanca, Parapenaeus longirostris, constituye una especie de elevado interés comercial
para las flotas de arrastre que faenan tanto en el Mediterráneo como en el Golfo de Cádiz y sur de
Portugal. A pesar de dicha importancia, su ciclo de vida no es del todo bien conocido existiendo
discrepancia en determinado aspectos en función del área. La puesta tiene lugar a lo largo de todo
el año, si bien presenta dos picos identificados entre final de primavera y principio de otoño. Esta
especie presenta un crecimiento diferencial, alcanzando mayor tamaño las hembras, con un ciclo de
5
) e-mail: ignacio.sobrino@cd.ieo.es
©
Koninklijke Brill NV, Leiden, 2005
Also available online: www.brill.nl
Crustaceana 78 (10): 1153-1184

1154
IGNACIO SOBRINO ET AL.
vida está entre dos y tres años. Alcanzan la madurez sexual entre los 8 y 10 meses variando la talla
en función del área (entre 20 y 28 mm de Longitud de Cefalotórax en las hembras) y un valor medio
de fecundidad de 90000 huevos. Se alimenta de una gran variedad de presas siendo los poliquetos
(polychaetes) y foraminiferos los principales. En esta pesquería están inmersas un elevado número
de embarcaciones, superando los 2000 barcos en el área tratada. Se desarrolla fundamentalmente
sobre en una única cohorte y con valores que pueden alcanzar las 10000 toneladas anuales.
INTRODUCTION
The deep water rose shrimp, Parapenaeus longirostris (Lucas, 1846), family
Penaeidae, is one of the three species in this genus inhabiting the Atlantic
Ocean (Pérez-Farfante & Kensley, 1997). The species shows a wide geographic
distribution, from the eastern Atlantic in the north of Spain (Olaso, 1990) to the
southern waters of Angola (Crosnier et al., 1968), as well as in the Mediterranean
and its adjacent seas (Sea of Marmara) (Karlovac, 1949; Maurin, 1960; Massuti,
1963; Audouin, 1965; Holthuis, 1980).
In the Mediterranean Sea, the bathymetric distribution of the deep water rose
shrimp ranges between 20 and 750 m (Tom et al., 1988), but the species is common
and abundant on sandy-muddy bottoms between 100 and 400 m (Bombace,
1972; Holthuis, 1987). In the Atlantic waters off Portugal and in the Gulf of
Cadiz, the species shows a bathymetric distribution similar to that observed in
the Mediterranean, being found in depths from 80 to 700 m (Ribeiro-Cascalho,
1988) and from 28 to 650 m (Sobrino, 1988), respectively. However, in African
Atlantic waters, this species seems to present a more restricted bathymetric range.
According to Sobrino & Fernandez (1991) it has been found between 20 and 400 m
in the Gulf of Guinea, and from 50 to 500 m off Congo (Crosnier et al., 1970),
as well as in the rest of tropical Africa (Crosnier & Forest, 1973). In Senegalese
waters, it has not been recorded in depths over 500 m (L. Lopez-Abellan, pers.
comm.), while in Mauritanian waters, it ranges from 50 to 400 m (M. Diop, pers.
comm.). Finally, Sobrino & Cardenas (1996) did not find this species beyond a
depth of 400 m in the waters off Angola.
Yet, in all investigated areas the greatest densities in terms of biomass have
been recorded by various authors to range between 100 and 300 m depth. In the
Mediterranean Sea, Froglia (1982) found its greatest abundance between 200 and
350 m, whilst Ardizzone (1990) found the highest densities in the range of 150
to 350 m. Crosnier et al. (1970) recorded the greatest densities in the waters off
Congo between 200 and 300 m, while Sobrino & Cardenas (1996) found them
concentrated in shallower waters (100-200 m).
This species is the main target species of a large fishing fleet working in
the eastern Atlantic. The principal fishing grounds are located to the south of

BIOLOGY AND FISHERIES OF PARAPENAEUS LONGIROSTRIS
1155
Spain and Portugal (Pestana, 1991; Sobrino et al., 1994), as well as in areas off
Morocco, Mauritania, Senegal, Guinea Bissau, Gabon, and Angola (Cervantes &
Goñi, 1986; Cervantes et al., 1991; Sobrino & García, 1991, 1992a, b). In the
Mediterranean Sea, P. longirostris is caught almost exclusively by trawl. Longhurst
(1970) indicated that the species is the most important crustacean resource along
the coasts of Spain, France, and Italy. Also in Algeria, Tunisia, Greece, and Turkey
the species has a commercial value, although on a lesser scale. FAO catches and
landings statistics from 1972 to 1991 indicate that the deep water rose shrimp is
the fifth species in order of importance for biomass among crustaceans landed in
the whole Mediterranean area (Stamatopoulos, 1993).
In spite of its abundance and high economic value, research on the biological
aspects of this species in the Atlantic is scarce. In the Mediterranean Sea, many
studies carried out in the last twenty years allow collecting detailed information
on distribution, abundance, and biology, but there is a lack of knowledge on the
exploitation state of this important resource. The main objective of the present
paper is to describe the actual information available for the Atlantic waters off
Portugal and for the Mediterranean waters off Spain, Italy, and Greece.
Parapenaeus longirostris (Lucas, 1846)
Diagnosis. — Rostrum with 7 dorsal teeth, one epigastric tooth situated con-
spicuously posterior to first rostral tooth. Rostrum very variable in length, longer
in female than in male. Carapace with orbital spine, hepatic spine, and branchioste-
gal spine, and a well-marked carina. Last three abdominal somites ending in spine
and telson armed with a pair of lateral spines. Petasma symmetrical, semiclosed
(fig. 1a). Appendix masculina subovoid (fig. 1b). Thelycum closed, with single
plate of sternite XIV raised in two pairs of lateral prominences (fig. 1c). Semi-
nal receptacles consisting of pairs of longitudinally or obliquely situated sacs. The
morphology of the spermatozoid is typically tack-shaped and includes unistellate
sperm (fig. 1d).
Geographical distribution. — P. longirostris has a wide geographical distribu-
tion in the eastern Atlantic, from the north of Spain to the southern waters of An-
gola, including the Mediterranean Sea. In Atlantic and European Mediterranean
waters, it constitutes a valuable target species for most viable fisheries.
In Spanish waters, this species is mainly caught in the Gulf of Cadiz (south
Spanish Atlantic waters). The bathymetric range of the deep water rose shrimp off
the Gulf of Cadiz is between 40 and 650 m, but the maximum abundance occurs
in depths of 100 to 300 m (Sobrino, 1988).
In Spanish Mediterranean waters, the species is also important in the south
(Alboran Sea) and the abundance decreases towards the north of the Spanish
Mediterranean area.

1156
IGNACIO SOBRINO ET AL.
Fig. 1. Parapenaeus longirostris (Lucas, 1846). a, petasma symmetrical, semiclosed; b, appendix
masculina subovoid; c, thelycum, closed; d, spermatozoid (typical tack-shape).
The deep water rose shrimp is distributed in all Italian waters, with the highest
abundance in the Sicilian Channel and the Ionian Sea, and the lowest density
in the Ligurian Sea and in the central and northern Adriatic. In this latter area,
P. longirostris was recently found to be abundant along the Albanian coast
(D’Onghia et al., 1998). The bathymetric distribution of the species ranges between
depths of 20 and 700 m although the highest densities are observed between 100
and 400 m depth. In the northern and central Tyrrhenian Sea (Mori et al., 1986;
Ardizzone et al., 1990), the vertical distribution of P. longirostris is wide, but the
species is mostly caught over muddy or muddy-sandy bottoms between 70 and
350 m. In the southern Tyrrhenian Sea (Spedicato et al., 1996), P. longirostris is
distributed in depths of 61 to 587 m, with the highest abundance in the 200-450 m
bathymetric stratum. Off western Sicily, Bombace (1972) recognizes a facies of
P. longirostris between 150 and 350 m, and related it to the presence of water
masses of Atlantic origin (temperature of 14◦C).
A size-related bathymetric distribution can be found in this species: after
the dispersion phase in shallow water, the shrimps move toward deeper water

BIOLOGY AND FISHERIES OF PARAPENAEUS LONGIROSTRIS
1157
throughout their life. Juveniles settle at around 100 m depth, although they can be
detected at 300 m, whereas the larger individuals are always found in water deeper
than 350 m (Froglia, 1982; Mori et al., 1986; Ardizzone et al., 1990; D’Onghia
et al., 1998). In the central Tyrrhenian Sea (Ardizzone et al., 1990) most of the
smaller males and females (<20 mm CL) were caught in depths shallower than
250 m. Individuals with a carapace length (CL) of 21-30 mm were collected mostly
between the 250 and 350 m isobaths, while the largest specimens mainly occurred
at greater depths.
P. longirostris is one of the most important species of the crustacean fishery
in Greece. It can be found on muddy or muddy-sandy bottoms, mainly in depths
ranging between 150 and 400 m, being very widespread both in the Aegean and
the Ionian Sea (Koukouras & Kattoulas, 1974; Koukouras et al., 1997; Politou
et al., 1998; Tserpes et al., 1999) In the southern Ionian Sea this species was also
found, in lower abundance, in deep waters (500-700 m) (Kapiris et al., 2002). In
the same area, the larger individuals (CL 26-35 mm) were found mainly in the
deeper stratum (500-700 m) whereas the smaller ones (8-22 mm) occurred in the
shallower zone (300-500 m) (Kapiris et al., 2002).
Trawl surveys performed in the framework of the European Union research
project “MEDITS” (Bertrand et al., 2002), along the European coasts of the
Mediterranean Sea from the Straits of Gibraltar to the Aegean Sea, showed the
highest abundances along the western coast of Sardinia, in the southern Tyrrhenian
Sea, the Sicilian Channel, the northwestern and eastern Ionian Sea, and the
Argosaronikos and Aegean Sea (Abelló et al., 2002). From the samples collected
during day-light hours between 30 and 800 m in spring (May-June) 1994 to 1999,
biomasses were usually higher in the 200-500 m depth stratum than in the 100-
200 m stratum, in accordance with the depth-related size trend found in this
species.
LIFE HISTORY
Growth
The life span of each generation of Parapenaeus longirostris is between two
and three years, and males grow more slowly than females. Length-weight and
total length (TL)-carapace length (CL) relationships are presented by sex (table I).
Von Bertalanffy growth parameters have been estimated for several areas of the
Mediterranean Sea and eastern Atlantic (table II). In Portuguese waters some
estimates of growth parameters were made using Ford-Walford and Gulland plots,
implemented in SLCA and ELEFAN software. Cadima et al. (1995) identified the
estimates presented by Ribeiro-Cascalho (1988) as the most reliable for that area.

1158
IGNACIO SOBRINO ET AL.
TABLE I
Biological characteristics of Parapenaeus longirostris (Lucas, 1846). Total weight — Cara-
pace length (CL) (TW = A*CLˆb) and total length (TL) carapace length (CL) relationships
(TL = A + b*CL)
Sex
A
b
Source
Southern
Portugal
Males [TW (g) — CL (mm)]
0.0075
2.19
Ribeiro-Cascalho &
Arrobas, 1987
Females [TW (g) — CL (mm)]
Males [TL (mm) — CL (mm)]
Females [TL (mm) — CL (mm)]
0.0055
5.454
5.954
2.31
0.90
0.87
Ribeiro-Cascalho &
Arrobas, 1987
Ribeiro-Cascalho &
Arrobas, 1987
Ribeiro-Cascalho &
Arrobas, 1987
Southern
Spain (Gulf
of Cadiz)
Males (weight (gr) — CL (mm))
Females (weight (gr) — CL (mm))
Males (TL (cm) — CL (mm))
Females (TL (cm) — CL (mm))
21.21
16.36
12.91
25.28
1.44
0.76
4.41
3.95
Sobrino, 1998
Sobrino, 1998
Sobrino, 1998
Sobrino, 1998
TABLE II
Von Bertalanffy growth parameters of Parapenaeus longirostris (Lucas, 1846) estimated for eastern
Atlantic Ocean and Mediterranean Sea
Area
Sex
LC (mm)
K
t
Author
∞
0
Southern Portugal
Southern Portugal
Southern Spain (Gulf of Cadiz) Female
Southern Spain (Gulf of Cadiz) Males
Central Tyrrhenian Sea
Central Tyrrhenian Sea
Ionian Sea
Ionian Sea
Greece
Greece
Female
Male
44
36.0
44
0.70 −0.30 Ribeiro-Cascalho, 1988
0.90 −0.30 Ribeiro-Cascalho, 1988
0.74 −0.15 Sobrino, 1998
33
0.95 −0.02 Sobrino, 1998
Female
Males
Female
Male
Female
Male
44.4
33.1
47.7
35.5
37.2
33.7
0.74 −0.13 Ardizzone et al., 1990
0.93 −0.05 Ardizzone et al., 1990
0.74 −0.19 D’Onghia et al., 1998
0.54 −0.29 D’Onghia et al., 1998
0.52 −0.30 Anonymous, 1999
0.62 −0.16 Anonymous, 1999
Sobrino (1998) obtained similar results in the Gulf of Cadiz using Ford-Walford
and Beverton & Holt (1957) methods. In Italian waters, growth parameters were
estimated in the central Tyrrhenian Sea and in the Ionian Sea.
From a comparison of the curves estimated for the various areas, similar
growth rates resulted for females, except for Greek waters, where growth is slower
(table III). As regards males, the specimens analysed in southern Portugal were,
for each age class, larger than those in Mediterranean waters.

BIOLOGY AND FISHERIES OF PARAPENAEUS LONGIROSTRIS
1159
TABLE III
Mean CL (mm) at different ages (in months) estimated for Parapenaeus longirostris (Lucas, 1846)
in southern Portugal and in some Mediterranean areas
Female
Months
Portugal,
Southern
18.9
26.3
31.5
35.2
37.8
39.6
Spain
Gulf of Cadiz
16.8
Italian
Tyrrhenian Sea
16.5
Italian
Ionian Sea
18.8
Greece
6
12
18
24
30
36
12.7
18.3
22.6
26.0
28.5
30.5
25.2
31.0
35.0
37.8
25.2
31.1
35.2
38.1
27.6
33.6
37.8
40.7
39.7
40.0
42.7
Male
Months
Portugal,
Southern
24.8
28.9
31.5
Spain
Gulf of Cadiz
22.4
Italian
Tyrrhenian Sea
20.6
Italian
Ionian Sea
17.8
Greece
12
18
24
30
36
17.3
21.7
24.9
27.2
28.9
26.4
28.9
30.5
31.4
25.3
28.2
30.0
31.2
22.0
25.2
27.6
29.5
33.1
34.2
Reproductive biology
Reproductive season. — In Atlantic water (south of the Iberian Peninsula)
mature females occur in deeper waters all through the year, with a gap in February-
April. Two spawning peaks were recorded, the first in summer and the other
in autumn-winter (Banha, 1982; Ribeiro-Cascalho & Arrobas, 1987; Ribeiro-
Cascalho, 1988; Sobrino, 1998). Caramelo et al. (1996) consider the summer
spawning as the most important and Sobrino (1998) found that the peak of spring
is mainly composed of adult individuals while the second one (summer) comprises
both juveniles and adults. The larger mature females mainly perform spring
spawning, whereas adult females of smaller sizes perform autumn spawning. An
analysis of the composition of maturity stages in each depth stratum sampled
in the surveys, shows that in the shallower strata (less than 100 m), females in
advanced stages are practically absent; thus, the major fraction is composed of
immature specimens. This situation varies with depth, producing an increase of
mature females in the deeper bathymetric strata. In the case of males, immature
specimens are present only in the shallowest stratum (less than 100 m). In the
rest of the deeper strata, practically the whole male population is mature. No
differences in the maturity stages according to depth stratum were observed in the
surveys carried out during March (non-reproductive period) in comparison with

1160
IGNACIO SOBRINO ET AL.
those carried out in October (spawning period). The few larvae found in the area
also suggest two spawning periods (Dos Santos, 1998).
In Italian waters, information on the reproductive season is available for
the Tyrrhenian Sea and Sicilian Channel. In the northern Tyrrhenian Sea, the
reproduction area was found in depths ranging from 150 to 350 m (Mori et al.,
1986, 2000b). In this area, ripe females were found throughout the year with
two maxima of activity, one in spring and another at the beginning of autumn,
confirming the reproductive period previously identified by De Ranieri et al. (1986)
in the same area. The presence of recruits (CL ꢀ 15 mm) throughout the year
confirmed that reproductive activity is almost continuous in this area, although
the larger recruitment seems to occur from July to October (De Ranieri et al.,
1998). By contrast, in the central Tyrrhenian Sea, on the basis of data relative
to the recruitment of young animals to the benthos, Ardizzone et al. (1990)
hypothesized a single winter spawning. Within the area investigated, no sign of
massive recruitment of juveniles was observed. Nevertheless, though it could not
be easily defined, it appeared to be protracted with several peaks, as indicated by
the numerous micro-cohorts among new recruits. This result is in agreement with
the data given by Froglia (1982), showing one generation per year in the southern
Tyrrhenian Sea. In the same area, studies carried out by Spedicato et al. (1996)
identified the highest frequency of mature females in spring-summer, although
mature specimens were found all year round. Along the northwestern Sicilian
coast, Arculeo et al. (1993) have found a long reproductive period, with a major
presence of mature females in winter and spring. In this period the ripe ovaries
contain vitellogenic oocytes with a mean diameter of 231 12.7 µm, while the
gonado-somatic index showed a value of 3.3. In the Sicilian Channel (Levi et al.,
1995) mature females occur in all months of the year, although a wide maturity
peak extends from November to February, with another one in April. The lowest
percentage of mature females appears in June-July, but continuous spawning seems
to occur. In this area, recruits between 11.5 and 12.5 mm CL are present in the
catches all year round, except for August, two months after the lowest percentage
of mature females was found.
In the Greek Ionian Sea, the sex ratio remained close to 1 : 1 in almost all
months in the period December 1996-November 1997. The younger individuals
(CL 15-20 mm) were caught in December, January, March, and July in the
shallower depth zone (300-500 m), reinforcing the view of continuous reproductive
activity (Anonymous, 1999).
Size at maturity. — Along the coasts of southern Portugal and in the whole
Mediterranean Sea, Parapenaeus longirostris reaches sexual maturity in the first
year of life in both sexes (table IV).

BIOLOGY AND FISHERIES OF PARAPENAEUS LONGIROSTRIS
1161
TABLE IV
Size at first maturity of Parapenaeus longirostris (Lucas, 1846) in different areas of the Atlantic and
the Mediterranean Sea
Area
Sex
LC
(mm)
Author
50%
Southern Portugal
Southern Portugal
Southern Spain (Gulf of Cadiz)
Southern Spain (Gulf of Cadiz)
Northern Tyrrhenian Sea
Southern Tyrrhenian Sea
Ionian Sea
Morocco
Gulf of Guinea
Congo
Angola
Female
Male
Female
Male
Female
Female
Female
Female
Female
Female
Female
24
20
22
Ribeiro-Cascalho, 1988
Ribeiro-Cascalho, 1988
Sobrino, 1998
Sobrino, 1998
Mori et al., 2000b
Spedicato et al., 1996
Orsi Relini et al., 1999
Sobrino & García, 1993
Sobrino & Fernandez, 1990
Crosnier et al., 1979
Sobrino & Cardena, 1996
14.8
19.5-24.4
28.4
20-22
24.8
25.5
22
21.6
Ribeiro-Cascalho (1988) estimated the size at first maturity as 20 mm carapace
length for males and 24 mm for females in the south of Portugal, corresponding to
an age of 8-10 months. However, Sobrino (1998) estimated the first maturity for
females to occur between 21.5 and 22 mm CL, whereas in males, the fusion of the
two pieces of the petasma takes place between 12 and 15 mm CL, being always
separated at sizes lower than 11 mm CL. The size at which 50% of the males have
a jointed petasma was estimated at 12.9 mm CL. The spermatophoric mass on
the coxae of the fifth pereiopods was present from a size of 13 mm CL, but the
size at first maturity (50% of males with spermatophoric mass) was estimated at
14.8 mm CL.
In the northern Tyrrhenian Sea (De Ranieri et al., 1998; Mori et al., 2000b),
the smallest females with ripe ovaries were 16 mm CL. The size at onset of
sexual maturity estimated for different years ranged from 24.4 mm CL in 1993
to 19.5 mm CL in 1994. The percentage of post-moulting females increased from
January to August, decreased from September to October, and rose from November
to December. Most post-moulting females were found with immature or post-
spawning ovaries. In the southern Tyrrhenian Sea the smallest females with ripe
ovaries were 18.5 mm CL (Spedicato et al., 1996). The size at onset of sexual
maturity was 28.4 mm and 31.2 mm. In the Ionian Sea the size at first maturity
in females was around 20-22 mm CL and the smallest mature individual was
14-16 mm CL (Relini et al., 1999).
Fecundity. — Fecundity in Parapenaeus longirostris was studied in the Gulf
of Cadiz and in the northern Tyrrhenian Sea. Great differences were observed
comparing the results obtained in the two areas, both regarding the relationship
between CL and egg number and the absolute fecundity.

1162
IGNACIO SOBRINO ET AL.
Fig. 2. Relationship between absolute fecundity and carapace length of Parapenaeus longirostris
(Lucas, 1846).
In the Gulf of Cadiz, absolute fecundity varied from 21,410 eggs for a female of
25.4 mm CL and 186,856 for a female of 34.2 mm CL. The total mean value was
87,766. The average relative fecundity for P. longirostris was 7,486 eggs/g, with
values ranging from 2,048 eggs/g to 13,371 eggs/g. The relationship between size
and weight with absolute fecundity has been fitted to a linear function. Fecundity
increases proportionally with size, increasing by approximately 10,000 eggs for
each mm of CL and increasing approximately by 9,500 eggs/g (fig. 2).
In the northern Tyrrhenian Sea the fecundity of P. longirostris was investigated
by Mori et al. (2000b). The number of oocytes in the ovary was related to the size
of the females and ranged from 23,000 oocytes at 26 mm CL to 204,000 at 43 mm
CL. An exponential relationship was observed between fecundity and carapace
length: Fecundity = 0.0569 CL4.0177 (r = 0.829).
Diet
The feeding habits of Parapenaeus longirostris are characterized by a large
variety of preys. During the hunting phase it feeds on small fish, cephalopods,
and crustaceans, while in a digging phase it searches for preys in the mud, such as
polychaetes, bivalves, echinoderms, and (mostly) foraminiferans (Relini, 1973).
Ribeiro-Cascalho & Arrobas (1983) examined the stomach contents of 40
specimens of P. longirostris, in south Portugal. Fish, crustaceans (Decapoda,
Cirripedia, and Ostracoda), polychaetes and foraminiferans seemed to be the
preferential preys.
Studies by Burukovsky (1969) in samples from the Gulf of Cadiz and off the
northwestern African coast indicate different diets according to age. Thus, at first
foraminiferans and planktonic crustaceans dominate in the food of young speci-

BIOLOGY AND FISHERIES OF PARAPENAEUS LONGIROSTRIS
1163
mens, amphipods and isopods replace them, and later, shrimps, crabs, euphausiids,
mysids, some cephalopods, and fish appear in the stomach content.
In Italian waters, some preliminary results on the natural diet of adult females
inhabiting the continental slope of the northern Tyrrhenian Sea were furnished by
Mori et al. (2000a). The results of that study showed the great variety of organ-
isms on which P. longirostris feeds. The prey items consisted mostly of external
skeletons of bottom organisms, always crushed and often in an advanced state of
deterioration. In addition, some inorganic material was found. Crustaceans domi-
nated the diet both qualitatively and quantitatively; this food group was character-
ized by a high abundance of peracarids, mainly represented by mysids, and above
all Lophogaster typicus (Sars, 1957), and amphipods (Lysianassidae). Molluscs
were represented mostly by juvenile bivalves and gastropods; the small amount
of cephalopods was constituted mainly by the sepiolid, Sepietta oweniana (Or-
bigny, 1840). The small echinoderms were mainly ophiuroids found as fragments
of arms, although sea urchins and holothuroids were also found. The annelids were
mostly represented by polychaetes belonging to Aphroditidae and Nephytidae.
Fishes were also observed among the preys, but quite rarely. Foraminiferans, above
all Globigerinidae, were mainly recorded in foreguts containing small amounts of
food. Organic detritus, mostly as unidentified debris, had a relatively high per-
centage of occurrence, but in low amounts, increasing quantitatively in specimens
caught on bottoms down to 300 m. Also, inorganic and plastic particles were found
mostly in specimens caught in the deepest area.
In Greek waters (Ionian Sea) no clear differentiation in the feeding behaviour in
terms of either diet composition or feeding activity between sexes was observed.
Feeding activity increased in summer for males, and in spring or summer for
females (Kapiris, in press).
Larval ecology
The life cycle of this species is entirely marine. Held (1938) has described the
larval development of Parapenaeus longirostris. We can summarize it at follows:
The development of the egg at a temperature of 16◦C takes 51 to 55 hours, while
at 25-26◦C hatching takes place after 18 hours. The nauplius phase has a length of
0.317 mm in the first stage, up to 0.44-0.51 mm in stage VIII. The next larval stage,
the protozoea phase, has a length of approximately 1 mm. There are three different
protozoeae (I, II, and III). The mysid phase follows the protozoeae. The first mysis
measures 3.9 mm and the larval development ends at mysis XIV with a total length
of 12 mm and a CL of 3.6 mm.
There are few data about larvae in wild conditions, and these only concern larvae
found in Portuguese waters. The occurrence of larvae on the south and southwest

1164
IGNACIO SOBRINO ET AL.
coasts coincides with the area of distribution of the adults. Most larvae occur
around the 100 m isobath that corresponds to the upper limit of the distribution
of the adults. This probably indicates a displacement of the adults during the
spawning period to shallower depths (Dos Santos, 1998).
FISHERY
Parapenaeus longirostris is actually the target species of an important fishery.
The areas in which these fisheries act are very diverse, due to the extensive
geographical range of the species, from the Mediterranean to the eastern central
Atlantic. In all areas in which this fishery is developed, the deep water rose shrimp
is captured by bottom trawl.
The development of this fishery along the Atlantic coast (off the Gulf of Cadiz)
began during the mid-eighteenth century. Martínez de Mora (1977) cites the
presence of 4 trawlers in 1743, the so called “bous”, while Sañez Reguart (1791)
cites the presence of this fishery in the Gulf of Cadiz in 1755, and located its origin
along the Spanish-French Mediterranean coasts.
In the beginning, this fishing activity was mainly centred in the coastal zone.
The first references of shrimp catch by this fleet in the Gulf of Cadiz (Machado,
1847) correspond to the mid-nineteenth century. The expansion of the fleet during
the end of the nineteenth century and the beginning of the twentieth century
led to the development of the fishery off the northern coasts of Morocco, as
indicated by Sancha (1975). All technological advancements, from the viewpoint
of propulsion (sails, steam, and explosion engines) up to fishing gear development
and conservation methods (freezing), have shown their effects on the expansion of
the fishery to its the actual state.
In Portuguese waters, the species is caught in the south-west (south of Cab
Espichel, 38◦25ꢀN) and along the south coasts (Algarve), from 80 to 700 m deep,
with a gradient of sizes according to depth: larger specimens being found in
deeper waters (Ribeiro-Cascalho, 1988). The highest yields are produced during
daytime, when the species forms concentrations, mostly between 100 and 300 m.
The preferential substratum is basically constituted by sandy and muddy sediments
(Viriato & Figueiredo, 1991). The bottom water temperature usually ranges from
12◦ to 14◦C and the salinity in those areas is 35.6-36.1h (Ribeiro-Cascalho, 1988).
The crustacean fishery started along the Portuguese coast in the 1950s, with a
combined fleet of both Spanish and Portuguese vessels. The majority of the catches
were taken by the Spanish component, as the Portuguese fleet component was very
small. In the 1960s, this fleet started operating at greater distances from the coast
and in deeper waters (Ribeiro-Cascalho, 1988; Santana, 1990; Caramelo et al.,
1996). There are very few catch statistics from this first period.

BIOLOGY AND FISHERIES OF PARAPENAEUS LONGIROSTRIS
1165
In 1977, the number of Portuguese vessels entering the fishery started to
increase and, since 1983, after the end of the Portuguese-Spanish bilateral fisheries
agreement, only the Portuguese fleet was allowed to fish for crustaceans in
Portuguese waters (Caramelo et al., 1996). According to Ribeiro-Cascalho et al.
(1986), the increasing interest of the Portuguese fleet in this fishing resource was
due to the decrease in the fish catch, the increase in crustacean abundance after the
end of the agreement, and the improvement of fishing technology that allowed the
landings to increase.
At present, there are important fisheries off the African Atlantic coasts as well.
In Morocco, this species is fished from Larache to Agadir, between the parallels
35◦30ꢀ and 30◦N (Poinsard & Villegas, 1975). The Spanish fleet initially exploited
this fishing ground at the end of the nineteenth century. The Moroccan fishing fleet
started its activity during 1978 (Cervantes et al., 1985, 1991; Anonymus, 1990).
Since then, the Moroccan fleet gradually replaced the Spanish fleet. Nowadays this
resource is wholly exploited by the Moroccan fleet. The mean annual registered
catch for the last 30 years was 8,000 metric tonne.
In the whole region, from the Saharian fishing grounds to the borders of
Mauritania off Cape Blanco (from 29◦ to 21◦N), the presence of P. longirostris
is minimal. As a result, no specific fishery directed towards this species has been
developed. From Cape Blanco onwards, the abundance increases, resulting in a
specific, targeted fishery. This fishery started in the 1960s with the development
of freezing capacity in the fishing vessels (Sancha, 1975), beginning with the
exploitation of the fishing grounds off Mauritania, Senegal, Guinea-Bissau, Gabon,
and Angola. A detailed description of these fisheries can be found in Sobrino &
García (1991, 1992a, b) and Lopéz Abellan et al. (1994).
The mean annual catch on the Mauritanian fishing ground during the last fifteen
years was 1,500 tonnes.
Fleet and gears
At present, the Portuguese fleet comprises two components: trawlers targeting
demersal fish, and trawlers targeting crustaceans. The demersal fish fleet operates
year-round along the entire coast of Portugal. Data on the average fish catch
composition of this fleet show that in recent years horse mackerel and hake were
the most valuable species (ICES, 1999).
There are 35 trawlers licensed for the crustacean fishery in Portugal (table V).
They are smaller than the demersal fish trawlers, part of them being former sardine
purse seines converted into trawlers. On average, these vessels are 25 m in overall
length, 111 t of GRT and 484 HP. Just a few of them make fully use of the freezing
capacity, making trips longer than 3 days (Cadima et al., 1995; Caramelo et al.,
1996).

1166
IGNACIO SOBRINO ET AL.
TABLE V
Profile of fishing fleet for Parapenaeus longirostris (Lucas, 1846) by region
Parameter
Region
Portugal
35
49.3-191.4
111.4
Spain
227
5.4-78.4
25.3
Italy
1600
10-130
35
Greece
344
33-131.5
63.2
Number of vessels
Range in GRT
Average GRT
Modal GRT (ton)
Range in length
Modal length (m)
Range in crew and modal
crew number
150
19-32.3
25
19.9
8.8-22.5
14.2
15-30
20
2-6
15-32.9
23.2
Average: 4
Average: 10
Average: 4
Modal class: 4
Bottom trawl
—
Gear used
Shrimp trawl
—
Shrimp trawl
—
Shrimp trawl
—
Gear units per boat (traps,
length of gill nets)
Multispecies fishery
Targeted or by-catch
Technical fishing aids (GPS,
plotters, etc.)
Y
Both
GPS, radar,
plotter
Y
Both
GPS, radar,
plotter
Y
Y
Both
GPS, radar,
plotter
Targeted
GPS, plotter,
echo sounder
Medium
Dependency on this species
> 50%
< 20%
< 20%
Y = yes.
As stated above, these vessels do not specifically target Parapenaeus lon-
girostris, but are licensed for the crustacean fishery in general. The gear used is
a shrimp trawl with a cod end mesh size of 55 mm. The catches are mostly landed
in Algarve fishing ports in south Portugal, particularly in Vila Real de Santo An-
tónio, where the crustaceans are auctioned.
Until recently, the most important species in this fishery was Nephrops norvegi-
cus (Linnaeus, 1758), but in the last five years, P. longirostris has ranked first in
terms of weight landed. The third important species of this fishery is the red shrimp,
Aristeus antennatus (Risso, 1816).
In the Gulf of Cadiz, as remarked previously, the fishery began in the mid-
nineteenth century (Machado, 1847). Since then, it has suffered multiple modi-
fications, as observed by different researchers (De Miranda, 1921; De Buen, 1922;
Rodriguez-Roda, 1955; Massutí, 1959) till reaching the present period described
by Sobrino et al. (1994). A total of 227 trawlers is operating (table V). These have
a mean tonnage of 25 GRT, a mean HP of 215, and a mean length of 14.2 m. This
fleet exploits a diversity of species, constituting a typical multi-specific fishery in
which the shrimp represents a mean value of 8%. The main landing ports are Isla
Cristina, Huelva, Sanlúcar de Barrameda, and Puerto de Santa María.
In the Spanish Mediterranean, this species only shows a relative importance in
the southern ports, although it can be considered an ancillary species. The total
annual catch shows maximum values of 700 tonnes.

BIOLOGY AND FISHERIES OF PARAPENAEUS LONGIROSTRIS
1167
The exploitation of P. longirostris in the Italian seas is carried out exclusively
by boats using a bottom trawl net, which is the fishing system with industrial
characteristics that is more widespread in Italy. Before the Second World War
this resource was scarcely exploited, because the fishing boats used sails as the
main propulsion system and this allowed only near shore fishing, with beam trawls
and set nets. Two vessels were used for trawling nets with a larger, but not rigid,
mouth. From the 1950s the pair trawl has been left and the otter boards introduced.
The employment of powerful engines and modern technologies allowed to increase
catching capacity and extend the exploitable bottoms, favouring the exploitation of
important resources at greater depth, especially crustaceans such as P. longirostris,
N. norvegicus, A. antennatus, and Aristeomorpha foliacea (Risso, 1827).
According to recent statistics, during 1999 the Italian bottom trawl fleet com-
prised 4094 vessels (Unimar, 2001), representing 25.8% of the total Italian fishing
fleet (table V). On average, these boats showed dimensions (mean GRT = 32 tons)
greater than the national average. It is hard to quantify the number of boats target-
ing P. longirostris, because they usually direct the fishing effort towards different
target species during the year, in accordance with the varying availability of these
resources. For example, in Mazara del Vallo, the most important Italian port for
this resource, about 70 of the total 350 trawlers target P. longirostris during the
whole year on the fishing grounds located in the Sicilian Channel (Levi et al.,
1995). In general, the trawlers with GRT smaller than 10 tons, representing about
the 44% of the Italian trawl fleet by number and 12% of the total tonnage, do not
affect this resource, their activity being restricted to the continental shelf (shal-
lower than 200 m). Among the larger boats, those specialized in crustacean fishery
can divide their activity among P. longirostris, N. norvegicus, A. antennatus, and
A. foliacea. These boats usually utilize some variants of the “Italian” bottom trawl
net, characterized by a low vertical opening of the mouth (about 1 m), very long
wings, and 38-40 mm cod end mesh size. The most important ports are located
along the southern coast of Sicily and in the Ionian Sea. Although to a lesser ex-
tent, P. longirostris is also economically important in the landings of some ports
of the central (e.g., Terracina, Napoli, Salerno) and northern (Porto Santo Stefano)
Tyrrhenian Sea.
The crustacean fishery in Greek waters does not constitute a particular branch
of the Greek fishery, which is mainly multispecies. In Greek waters, the follow-
ing crustaceans can be characterized as target species: N. norvegicus, Melicertus
kerathurus (Forskål, 1775), Palinurus elephas (Fabricius, 1787), Homarus gam-
marus (Linnaeus, 1758), and P. longirostris.
In Greece, the number of trawlers is 344 and their fishing activity is not
directed to P. longirostris but to many other commercial species (table V). The
characteristics of these vessels are: average length 23.2 m, GRT of 63.24, and

1168
IGNACIO SOBRINO ET AL.
390.3 HP. The mean age of these boats in Greece is 22.8 years and that of the
fishermen is 45-50 years. The average number of crew for trawlers is four and the
number of fishing ports in Greece is 42. The common trawl used has a mesh size
at cod end of 32 mm (28-36 mm), with a horizontal opening of about 15 m and
a vertical opening of about 1.5 m. The main fish markets are: Piraeus, Chalkis,
Thessaloniki, Kavala, Chios, Patra, Preveza, and Alexandroupolis. There are no
markets outside the country. In the Ionian Sea, all existent 43 trawlers may catch
this species. Their fishing activity lasts 15-20 h/day. In the Aegean Sea, there are
277 trawlers, working 15-20 h/day, depending on the area.
The fishing gear used for the fishery of this species in Greek waters is the
common trawl. The trawl activity is not directed at P. longirostris but mostly to
other commercial (fish) species. P. longirostris is caught by trawl between 150 and
400 m, on muddy-sandy bottoms from October to May. No clear correlation was
found between depth and size distribution for the deep-water shrimp population in
the Ionian Sea.
Fishing operation regime and fishing area
According to Cascalho (1988), for many years the average fishing regime was
one fishing day per trip, with three hauls a day (of 3 to 4 hours each), and around
17 days per month. This fishing regime has remained constant with only small
variations. The freezer trawlers make longer trips (more than 5 days each) and
the total number of fishing days per month increases to 22. The mean number
of fishing hours per boat also changes among boats and along the year, varying
between approximately 10 and 14 hours per day (Dias, 1992). More recent data
indicate an increase in the average number of fishing hours per day. The trawling
speed varies between 2.5 and 3 knots (approx. 4.5-5.4 km/h). The crustacean fleet
operates along the south-western and southern coasts of Portugal (Alentejo and
Algarve), fishing in the depth range of 100 to 700 meters. Concentrations of the
deep water rose shrimp are found mainly in areas shallower than 400 m. The
southwestern coast is mostly covered by vessels that operate for longer periods
without landing the catches (Ribeiro-Cascalho et al., 1986). The fishing operations
occur on regular bottoms, the substrata being constituted mainly of muddy and
sandy sediments.
In the Gulf of Cadiz, the fleet carries out mostly one-day fishing trips, although
some trips may last two to three days. The species is mainly fished in the area
from Ayamonte (Portuguese border) to the Bay of Cadiz, decreasing its abundance
towards the Strait of Gibraltar. The fishing depth ranges from 90 to 380 m, though
it has been cited in depths beyond 700 m (Sobrino, 1998).
The fishing regime of the trawlers targeting Parapenaeus longirostris is variable
along the Italian coasts, as a consequence of the characteristics of the fleets, the

BIOLOGY AND FISHERIES OF PARAPENAEUS LONGIROSTRIS
1169
geomorphology of the sea bed, and the distance of the fishing grounds. In general,
the trawlers carry out one fishing day per trip, performing three hauls with an
average tow duration of 3 hours. The larger boats, with freezing and refrigeration
equipment on board, can perform trips of 3-5 consecutive days. A particular case
is represented by the fleet of Mazara del Vallo, where large freezing vessels are
able to perform trips of two or three weeks on the fishing grounds of the Sicilian
Channel. The fishing activity per month ranges between 18 and 22 days, depending
on the fishing port. The most important fishing grounds are located in the southern
Tyrrhenian Sea, the Ionian Sea, and the Sicilian Channel, where this species shows
very high abundance indices (Anonymous, 1988, 1993; Tursi et al., 1994). The
bottom is regular and muddy, and the depth range is 200 to 400 m.
P. longirostris, fished all over the Greek seas, constitutes the most important
species (in catch weight, landings, and value) among the commercial crustaceans
in Greece. The higher mean annual landings are found in the ports of Piraeus and
Thessaloniki, followed by the ports of Kavala, Patra, and Alexandroupolis. The
annual catches of P. longirostris in Greece show a great variability in the period
1990-1997, with a periodicity of 4-5 years. The landings of the species clearly
increased during that period. The large amount of catches and the relatively low
value/kg of this species may be the reasons that led fishermen to pass most of
their catches through the official landing ports, since no better price would be
obtained by the direct market, and such volume of catches is difficult to be sold
directly to the small merchants. In fact, the landings of the species represented
23% of the catches in 1990, against the 80% in 1998. The increase in the landings
of P. longirostris was accompanied by a similar increase in their total annual
value, although the value/kg did not indicate a particular trend during that same
period. However, it seems that the value/kg is affected by the volume of landings,
reaching a maximum with minimum landings, and inversely so (Mytilineou et al.,
2001).
Selectivity
Several selectivity experiments have been carried out in Portugal, on board
of commercial vessels and research vessels, using the covered cod end method
(Ribeiro-Cascalho, 1988; Ferreira et al., 1993; Caramelo et al., 1999; Fonseca
et al., 1999, 2000). The nominal cod end mesh sizes tested were 45, 55, 60, 65,
70, and 80 mm. The cod end cover mesh size was 20 mm. Table VI summarizes
the results of the more recent experiments conducted with a commercial crustacean
trawler. The average duration of the tows was 3 hours, at a trawling speed of 3.1-3.2
knots (approx. 5.7 km/h) (Fonseca et al., 1999, 2000).
In the south of Portugal, Campos et al. (2002) experimented with the effect on
the catches of Parapenaeus longirostris by increasing the cod end mesh size from

1170
IGNACIO SOBRINO ET AL.
TABLE VI
Selectivity parameters for Parapenaeus longirostris (Lucas, 1846) in different areas
Area
Mesh size (mm)
L
(mm)
SR (mm)
SF
50
Southern Portugal
38
55
60
70
80
15.5
2.4
13.3
9.8
9.8
10.2
0.41
0.40
0.39
0.40
0.39
23.0
23.5
28.6
30.6
Southern Portugal
Gulf of Cadiz
55
55 square
60
21.8
27.1
24.0
27.1
5.7
9.3
9.3
8.9
0.40
0.49
0.40
0.39
70
35.9
42.8
46
52.7
60.2
66.3
14.3
20.8
19.2
19.3
26.3
29.6
3.1
6.7
6.6
8.4
7.6
13.8
0.40
0.49
0.42
0.37
0.44
0.45
Northern Tyrrhenian
Sea
36
12.4
3.1
0.34
Sicilian Channel
32
16.1
2.3
0.50
55 to 60 and 70 mm, and changing the mesh configuration from diamond to square.
The results indicated a significant increase in L50 when increasing the mesh size or
using a square mesh size cod end (table VI).
In the Gulf of Cadiz selectivity experiments have also been done using the
covered cod end method. Two different selectivity survey designs were applied:
first, using different mesh sizes and a fixed trawl duration (one hour) on board
of an oceanographic vessel (6 surveys) (table VI), and second, carrying out a
selectivity survey on board commercial fishing vessels (4 experiments), using the
standard fishing gear of the fleet at variable trawl duration times. In all cases,
the cod end cover was 20 mm. All results have been published by Sobrino et al.
(2000).
At this moment, there is a lack of information regarding the selectivity of the
commercial trawl net on P. longirostris in Italian waters (table VI). In the northern
Tyrrhenian Sea a trawl survey was carried out in order to estimate the selectivity
of the trawl net used by the fleet (Sbrana et al., in press). Another available value
of L50, estimated through an indirect method, was reported by Levi et al. (1995)
for a commercial trawl net of 32 mm cod-end stretched mesh size in the Sicilian
Channel. Backward projection of the descending arm of the catch curve (after

BIOLOGY AND FISHERIES OF PARAPENAEUS LONGIROSTRIS
1171
logistic transformation), gave for this net L25 = 14.9 CL, L50 = 16.1 mm CL,
and L75 = 17.2 mm CL, with R = 0.9956 for the regression between probabilities
and lengths. Hence, it seems that about two months are needed to reach the CL for
recruitment to the fishery in this area.
MONITORING THE FISHERY
Landings
Fig. 3 presents the development of the landings of Parapenaeus longirostris for
Portugal, Spain, Italy, and Greece, in the period 1990-2000.
In Portugal, the statistics of landings are recorded in the auction places by
vessel, species, and commercial size category. These records also include data
on the product value. Catch-and-effort statistics are available from logbooks, but
the reliability of this information is questionable. The Portuguese landings of
P. longirostris in the last two decades show alternating periods of high and low
abundance, with maxima in 1984, 1989, and 1999. In 1995, due to the decline of
the yields of the Norway lobster, the crustacean fleet redirected its effort to the
deep water rose shrimp grounds. In the period 1998-2000, the high effort on this
species was the result of its highest abundance and produced the highest peak of
landings of this species. The length composition of the landings is estimated by a
sampling program at the main auction place for the crustacean fishery, in Vila Real
de Santo António.
Since 1982, a specific Spanish programme focused on the compilation of
statistical fishery information from the main landing sites of the Gulf of Cadiz
exists. As mentioned before, P. longirostris is one of the target species of a multi-
specific fishery in the region. The development of the catches shows the typical
variation of short-lived species, whose abundance is directly related to recruitment
Fig. 3. Landings of Parapenaeus longirostris (Lucas, 1846) in different countries.

1172
IGNACIO SOBRINO ET AL.
strength. A certain similarity can be observed in the catch trends of the Portuguese
and the Cadiz fisheries (fig. 3), indicating the possibility of a link between these
stocks.
P. longirostris is one of the most important fishery resources of epibathyal
grounds along the Italian coasts. The exploitation of this species takes place all
year round, but peak landings are from March to July (Levi et al., 1995). The
whole catch is marketable, with the biggest specimens being of greater commercial
value. In some areas discard may occur, but at the moment there is no information
about this aspect. A study carried out in the northern Tyrrhenian Sea showed that
discards of P. longirostris were quite negligible, ranging from 0.35 to 1.24% of
the total catch in number of specimens of the species (Sbrana et al., in press).
The size at which 50% of the specimens was discarded was 14.99 mm CL, whilst
almost everything equal to or greater than 20 mm CL was commercialized. Starting
from 1985, ISTAT, Istituto Italiano Nazionale di Statistica, recorded landing data
on Italian seas. From 1997, FAO statistics are available for Italian waters. The
statistics show an evident reduction of biomass landed from 1990 (18,368 t) to
1998 (4,410 t) with a notable recovery in the last year (7,500 t in 2000) (fig. 3).
This trend could be directly influenced by the biomass of this resource at sea,
P. longirostris being a short life cycle species with great abundance fluctuations
from year to year.
In Greek waters the fishing of this species with trawlers takes place between
150 and 400 m, on muddy-sandy bottoms, from October to May. The landing
data mentioned below, come from the ETANAL-Company for the development
of fishery, and are collected from the 10 Greek auction places, independently of
the fishing area. The mean annual crustacean catches in Greek waters increased
from 175 t the period 1928-1934 to 1,260 t in the period 1964-1981 (Stergiou,
1986; NSSG data). In 1999, crustacean catches were about 2,780 t, representing
3.7% of total catches (FAO data). During the same year, the crustacean landings
from Greek waters were about 2,300 t, representing 3.8% of total landings;
their value represented about 8% of the value of the total landings (ETANAL
data).
ASSESSMENT OF THE IMPACT OF FISHING ON THE POPULATION
Assessment and management
Several surveys have been done in Portuguese waters in the period 1981-2000
to estimate abundance indices for crustacean species. Currently, a survey is carried
out once a year, in summer, covering the main area of distribution of these species

BIOLOGY AND FISHERIES OF PARAPENAEUS LONGIROSTRIS
1173
(Alentejo and Algarve). A new series of surveys directed to the deep water rose
shrimp in the recruitment season has started in 2002.
Natural mortality was estimated by several authors using Pauly’s formula, the
Rikhter & Efanov method, and Tanaka’s longevity curve (Banha, 1982; Ribeiro-
Cascalho, 1988; Cadima et al., 1995). The values ranged from 1.20 to 1.87 year−1
for males and from 1.07 to 1.60 year−1 for females.
Based on the geographical distribution of Parapenaeus longirostris on the
Portuguese fishing grounds, Pestana (1991) assumed one stock unit for the whole
area of southern Portugal (Alentejo and Algarve) and presented the results of the
assessment for the period 1984-1989, using age-converted length composition as
input for Virtual Population Analysis. According to this analysis, the stock was
overexploited and a reduction in fishing induced mortality was recommended.
Attention was also called to the high fluctuations in recruitment.
Pestana & Ribeiro-Cascalho (1991) studied the effects of changing trawl mesh
size and fishing effort on the stock, and provided some guidelines for management
purposes, suggesting either a decrease of 10% in the level of fishing effort without
any mesh change, or to keep the same level of effort, increasing trawl mesh size
from 55 to 60 mm.
Cadima et al. (1995) presented an analysis of the trends in total catches and
CPUE for this species for the period 1983-1994. A preliminary stock assessment
was carried out fitting Schaefer & Fox surplus production models to this data set.
The analysis suggested that the deep water rose shrimp stock was overexploited.
Thompson-Bell Yield per Recruit Model, and R. Jones length cohort analysis
were also applied to the rose shrimp data by Cadima et al. (1995). The Y/R model
indicated that the stock was overexploited. The length cohort analysis (applied
separately for males and females) did not indicate a situation of overexploitation
but showed that the fishing level should not be increased.
The shrimp fishery of Cadiz is being monitored since 1993. During this period,
a series of demersal evaluation surveys began with shrimp as one of the main target
species. Since then, abundance indices are estimated, as well as the respective
recruitment indices, which are ultimately used to relate them with the yields of
commercial activity (Sobrino, 1998).
A sampling programme started during the same period has allowed to make
available the necessary information to carry out analytical evaluations (Length
Cohort Analysis), thus obtaining fishing mortality estimates (from 0.3-0.7) by
size range. Fig. 4 shows the results obtained (Sobrino, 1998) during 1994. In
order to assess the effect produced by a modification of the present exploitation
pattern, the model proposed by Thompson & Bell (1934) was used, adapted to
sizes as described by Sparre & Venema (1992). Although the increase in mesh
size could cause immediate losses, these results should be considered cautiously

1174
IGNACIO SOBRINO ET AL.
Fig. 4. Fishing mortality of Parapenaeus longirostris (Lucas, 1846) by sex and length in the Gulf of
Cadiz (year 1994).
with regard to the penaeid fishery, because this is involved with fast-growing and
short-lived species. The immediate losses would be compensated for during the
first year, since the species would take only five months to grow from 17 mm
(L50 with the 40 mm mesh) to 23.7 mm (L50 with the 55 mm mesh). The mean
biomass by size range suffers an important change with mesh increase, especially
in the sizes between 20 and 28 mm. Taking into account that the size at first
maturity of females in this area is at 22.2 mm CL (Sobrino & Gárcia, 1998),
there would be an important biomass stock increase. Depending on the mesh used
(50 or 55 mm), this increase could represent between 200% and 250% (Sobrino
et al., 2000). However, at the present exploitation level, there does not seem to
be a relationship between spawning biomass stock and recruitment. Recruitment
may be influenced more by variable oceanographic conditions than by stock size
(Sobrino, 1998).
National otter trawl surveys were performed in all Italian seas from 1984-
85 until now. All the seas were covered (from 0 to 700 m depth), applying

BIOLOGY AND FISHERIES OF PARAPENAEUS LONGIROSTRIS
1175
a stratified random sampling design. The ten most important demersal species,
including P. longirostris, were chosen and the following data and results were
achieved: length frequency distributions for each survey by stratum, sex ratio,
growth parameters, length/weight relationship, sexual maturity, mortality, and
exploitation state.
From 1994 an international programme (MEDITS — Mediterranean Interna-
tional Trawl Surveys) was financed by the European Commission with the aim
to produce demographic structure and biological data on the demersal resources
along the coasts of the four Mediterranean countries of the European Union (Spain,
France, Italy, and Greece). Each year a campaign was carried out in late spring, be-
ginning of summer, covering all the trawlable areas on the shelves and the upper
slopes (at depths from 10 to 800 m).
In order to compare the national campaigns with MEDITS, the Italian surveys
started to adopt the same sampling protocol as the MEDITS project in 1996.
Utilizing MEDITS data, Abelló et al. (2002) obtained an index of fishing pressure
exerted on each sector covered by the project, from the Strait of Gibraltar to the
Aegean Sea, for the period 1994-1999. The analysis allowed to obtain an estimate
of the instantaneous total mortality rate, Z. High mortality related to small mean
size of the population present at sea was particularly evident in the Sicilian Channel
and the southern Aegean Sea, whereas the inverse relationship was found in the
Ligurian and southern Adriatic seas. The highest values of total mortality were
found in the Sicilian Channel sector, where a high effort targeted at P. longirostris
occurs.
Results of analytical models of evaluation obtained in some areas along the Ital-
ian coasts in the period 1990-1998 showed a generalized, slight overexploitation
status that could be tackled by reducing trawl fishing effort by about 10-20% and/or
increasing the size at first capture (Lc).
In the Sicilian Channel an attempt to assess P. longirostris using one year of
landing data was performed. Total mortality rate (Z), computed by the catch curve
method, ranged between 0.705 and 0.941 according to the various fishing grounds.
A fit of the Beverton & Holt yield per recruit against exploitation rate suggested
that fishing effort should be reduced by about 14% to obtain higher benefits at short
term. On the same data set, the Thompson & Bell model was applied. Compared
with the 14% immediate reduction suggested by the Beverton & Holt model, a
much more serious reduction (some 70%) would seem necessary to reach maximal
Y/R and maximum value of the catch on a long-term basis. The reduction in effort
required to obtain maximum yield in weight in the long run is somewhat less
(about 50%). The possible overall message could thus be that in this area modest
measures taken immediately, would avoid the need of extremely drastic actions in
the future.

1176
IGNACIO SOBRINO ET AL.
TABLE VII
Management measures and structures for Parapenaeus longirostris (Lucas, 1846) by country
MANAGEMENT MEASURES
Greece
Italy
Spain
Portugal
Technical measures
Minimum landing size
No
No
22 mm CL (EU 24 mm CL or
regulation)
No
94 mm TL
No
Sex restrictions
No
No
Mesh regulations in trawls
Trawl devices to exclude by-
catch or undersized fish
28 mm
None
40 mm
None
40 mm
None
55 mm
None
Other technical restrictions Maximum engine
power: 500 HP
None
None
Minimum of
30% of shrimp
species
Effort control
Quota
—
Seasonal or other closure
Closed season 30-45 Consecutive Fishery ban
Closed area
for trawling,
of 4 months
(June-
days of closure
per year. Fishery and Sunday from December
on Saturday
September)
ban on Saturday
and Sunday
to February
Closure zones
Areas prohibited Coastal 3 miles
to trawler or any strip and upper
Trawl ban
< 6 miles
from shore
Trawl ban
< 6 miles
from shore
fishing gear
50 m depth
forbidden
Stop new
licences
Gear limitations
Maintenance of
the total GRT, HP
of the trawl fishery
Stop new
licences
Maximum
number of
licensed
vessels: 35
Centralized,
co-management
MANAGEMENT
STRUCTURES
Centralized
Centralized
Centralized
Table VII summarizes the information available on management measures and
management structures for this fishery in different areas.
DISCUSSION
The deep water rose shrimp, Parapenaeus longirostris is one of the most
important commercial decapod species in bottom trawl fisheries in Atlantic and
Mediterranean waters, where it is caught as target or bycatch. The species plays
an important ecological role in many demersal communities of the continental
shelf and of the upper part of the continental slope. As regards its geographical
pattern, P. longirostris is very abundant along the south western Portuguese coasts
and in the Gulf of Cadiz. In the Mediterranean its quantitative importance is

BIOLOGY AND FISHERIES OF PARAPENAEUS LONGIROSTRIS
1177
rather variable, being scarcely abundant along the Spanish and French coasts. In
Italian seas its importance gradually increases from northern to southern latitudes,
reaching maximum abundance in the Sicilian Channel and the Ionian Sea.
Some aspects of the biology of the deep water rose shrimp are well known
(growth, reproduction), while other information, such as on larval phases, diet, or
abundance variability are, at this moment, poorly investigated. In particular data
on environmental factors related to the biology and abundance of the species are
not available.
Growth shows a similar pattern comparing the investigated Atlantic and Mediter-
ranean areas, with different growth rates between males and females. However,
some differences have been observed: lower growth rates for females in Greek wa-
ters and faster growth rates for males along the Portuguese coasts indicate different
environmental conditions (temperature, food availability, etc.) and/or fishing pres-
sure. In all considered areas, the life span is three years, but the fishery is based
mainly on cohorts of one or two years old. Studies on morphometry and genetics
for the identification of the stocks are totally lacking for this species.
In the Gulf of Cadiz, P. longirostris shows a reproductive pattern with two
spawning peaks, the first one is observed in late spring and corresponds to the
spawning of older individuals. The second peak is observed in autumn, during
which the same older individuals spawn together with those that spawn for the first
time. A similar situation has also been observed by Ribeiro-Cascalho & Arrobas
(1983, 1987) in southern Portugal and by Sobrino & García (1994) in the Atlantic
waters off Morocco, with two spawning peaks (June and October). In the water off
Congo, Crosnier et al. (1970) also defined one main spawning peak between April
and May and a less important one between September and October.
The studies concerning the reproduction of this species carried out in the
Mediterranean show some differences among different areas. Held (1938) de-
scribed a single spawning peak between April and November in Tunisian waters,
while Drobisheva (1970) only found mature females in spring on the Egyptian
coast. On the coasts of Italy, Levi et al. (1995), and D’Onghia et al. (1998), using a
macroscopic maturity scale for the female gonads, described continuous spawning
during the year. Tom et al. (1988), in the waters off Israel, relate oogenesis with the
temperature and distinguish different reproductive periods according to depth. In
shallow waters (between 47 and 73 m), reproductive activity is continuous, while
in deeper waters, spawning ceases from June to August.
The size at first maturity is variable. Ribeiro-Cascalho (1988) estimated the size
at first maturity as 24 mm CL for females in southern Portugal, while Sobrino
(1998) determined this value at around 22 mm CL in the Gulf of Cadiz. In
Moroccan waters, Sobrino & García (1994) obtained values from 24.5 to 27.7 mm
CL. Based on the results of a survey in the Gulf of Guinea, Sobrino & Fernández

1178
IGNACIO SOBRINO ET AL.
(1991) estimated the size at first maturity as 25.5 mm CL, although these authors
pointed out that the non-exploited state of this resource could have an influence on
its reproductive behaviour. The value determined by Crosnier et al. (1970) for the
waters off Congo (22 mm), is very similar to the estimate of Sobrino & Cardenas
(1996) off Angola (21.6 mm) and to the one given by De Ranieri et al. (1996) for
Mediterranean waters (21.5 mm).
The reproductive behaviour of this shrimp in relation to its bathymetric distribu-
tion shows a clear segregation of the various maturity stages. Females in advanced
stages of maturity or imminent spawning are found in depths below 100 m. In
shallower waters, gravid females are practically absent. This behaviour has been
described also in other areas. In the Mediterranean (Tyrrhenian Sea), Mori et al.
(1986) locate the spawning grounds at depths between 150 and 300 m, while in the
Gulf of Guinea they are deeper than 200 m (Sobrino & Fernández, 1991), and in
the waters off Angola they are concentrated between 100 and 400 m depth (Sobrino
& Cardenas, 1996). However, these results differ from those of Tom et al. (1988),
who reported a continuous spawning activity in the waters off Israel in depths from
47 to 73 m, relating this to the water temperature, and suggesting a temperature of
16◦C as the lower limit for a decline of spawning activity. In the Gulf of Cadiz,
within the bathymetric range in which mature females are found (100-400 m),
the water temperature ranges between 16.5◦C in depths of 100-200 m, to 14◦C
at 300 m, and to 13◦C at greater depths. These results are in agreement with Ghi-
dalia & Bourgois (1961) who found a correlation between the abundance of shrimp
and the water temperature, suggesting a lower temperature limit of 13.5◦C and an
optimal temperature between 14 and 15◦C.
The fishery of P. longirostris in the eastern Atlantic (Portugal) and northern
Mediterranean (Spain, Italy, Greece) is well developed and carried out by the
common bottom trawlers involved in a multi-species fishery. Although a historical
data series of about 20-30 years on deep water rose shrimp landings is available
for Portugal and Mediterranean EU countries, frequently these statistics are not
entirely reliable from a quantitative point of view, but they are useful to evaluate
trends in the catches of this species. In addition, data on fishing effort results are
poor or, in some cases, absent. Important data have been recorded in the last ten
years through experimental trawl surveys carried out at national and international
levels. The official statistics on landings show a positive trend in the last decade
both in Portugal and in Spain; an increasing trend, but more fluctuating, is observed
in Greece, whilst an evident decrease is detected in Italy. In the last case this trend
could also be a consequence of the high reduction of the fishing effort in terms of
the number of boats.
An evaluation of the exploitation state of this resource indicates a general over-
or full-exploitation in all areas considered. Global and analytical models applied on

BIOLOGY AND FISHERIES OF PARAPENAEUS LONGIROSTRIS
1179
local populations showed in general the need for a reduction in fishing effort (10-
15%) and an increase of the cod end mesh size in order to reduce fishing mortality
of juveniles. In this sense the future development studies directed to create a fishing
ban on a spatio-temporal basis, and experiments with more selective gears, will
be important points for a better management of this resource. For this reason,
it is also recommended to improve the monitoring of landing and fishing effort
and to increase the knowledge on some poorly known aspects of the biology and
population dynamics of this economically important species.
ACKNOWLEDGEMENT
The authors acknowledge the financial support of the European Commission
through the project “European Decapods Fisheries: Assessment and Management
(EDFAM)” Concert Action QLRT 1999 01272, and the financial support provided
by the IPIMAR project NeoMAv (UE co-financed QCA III-FEDER) for covering
part of the publication expenses. We also thank the referees for their useful
suggestions to improve the paper.
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