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Gonad maturation in Octopus vulgaris during ongrowing, under different conditions of sex ratio
Juan Estefanell, Juan Socorro, Francisco J. Roo, Rafael Guirao, lito Ferna ndez-Palacios, and Marisol Izquierdo Hipo
ndez-Palacios, H., and Izquierdo, M. 2010. Gonad maturation in Octopus vulgaris during ongrowing, Estefanell, J., Socorro, J., Roo, F. J., Ferna under different conditions of sex ratio. ICES Journal of Marine Science, 67: 1487 1493.

Octopus vulgaris is a suitable candidate for aquaculture, but there are problems with breeding in captivity, such as aggressive behaviour among males and the frequent death of females after the eggs hatch. To avoid these problems and further understand the sexual maturation of common octopus in captivity, males and females were reared together and separately under similar culture conditions. In all trials, the initial rearing density was 10 kg m23. Females (n 15, sex ratio 0:1) and males (n 11, sex ratio 1:0) were kept in circular tanks, and a mixed group (n 209, sex ratio 4:1) in oating cages. Trials started in November 2008 and octopuses from each treatment were examined macroscopically and histologically in December and January to assess sexual maturation. All the males matured, regardless of the sex ratio during rearing, as did all females in the mixed group. In contrast, a large proportion of the females kept isolated from males was still immature in December and January. Although maturation was successful in oating cages, there was 76% mortality there, in contrast to the zero mortality in tanks. Moreover, most of the dead octopuses from the cages were in post-reproductive condition, with a low digestive gland index, suggesting that this was natural post-reproductive mortality. Therefore, sex segregation is deemed advantageous to avoiding early mortality.
Keywords: digestive gland, gonad maturation, mortality, Octopus vulgaris. Received 19 October 2009; accepted 8 June 2010. ndez-Palacios, and M. Izquierdo: Grupo de Investigacio n en Acuicultura, Instituto Canario de Ciencias J. Estefanell, J. Socorro, F. J. Roo, H. Ferna Marinas i Instituto Universitario de Sanidad Animal y Seguridad Alimentaria, PO Box 56, E-35200 Telde Las Palmas de Gran Canaria, Spain. timo Pesquero de Las Palmas, Simo n Bol var 15, E-35007 Las Palmas de Gran Canaria, Spain. R. Guirao: CANEXMAR SL, J. Socorro: IES Mar de Tirajana, Las Palmas de Gran Canaria, Spain. Correspondence to J. Estefanell: Palangre s/n nave 1, Castillo del Romeral, San Bartolome tel: + 34 928 132900; fax: + 34 928 132908; e-mail: juanestefanell@iccm.rcanaria.es.

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Introduction
The common octopus (Octopus vulgaris) is a merobenthic species that lives from the coast to the outer edge of the continental shelf, up to 200 m deep (Guerra, 1992), all along the western and eastern Atlantic coasts, in the Mediterranean Sea, and in the Northwest ller et al., 2000; Warnke Pacic, around Taiwan and Japan (So et al., 2004). It has been proposed as a candidate for diversication of European aquaculture, the further development of which is constrained by market saturation because of the small number of species produced commercially (Vaz-Pires et al., 2004). The common octopus commands high market prices and there is increasing demand for it in Europe, South America, and Asia (Vaz-Pires et al., 2004). It grows rapidly (Mangold, 1983) and easily adapts to culture conditions (Iglesias et al., 2000), so would seemingly have an excellent potential for culture (Socorro guez et al., 2006; Garc a et al., 2005; Chapela et al., 2006; Rodr a et al., 2009). Despite unsolved problems in rearing the Garc larvae (Navarro and Villanueva, 2000, 2003; Villanueva et al., 2002, 2004, 2009; Iglesias et al., 2004, 2006, 2007; Villanueva and Bustamante, 2006), a few companies based in Spain (Galicia and the Canary Islands) already ongrow wild subadults in oating cages, using low-price species as feed.
# 2010

One factor inuencing the industrial development of octopus culture is sexual maturation under rearing conditions. Male octopuses die after mating, and females die immediately after the eggs a et al., 2002) and can lose ndez-Garc hatch (Guerra, 1992; Herna as much as 30 60% of their initial body weight during egg-laying (Iglesias et al., 2000). Although there have been several successful ongrowing experiments with males and females reared together guez et al., 2006; Garc a Garc a et al., (Socorro et al., 2005; Rodr 2009), the effect of sex segregation during broodstock maturation has not been studied well, and the results are contradictory. Whereas Chapela et al. (2006) stated that both males and females grow at similar rates even though 60% of females spawn ndez et al. (2003) documented slower during summer, Rey Me growth and greater survival in females than in males. Sexual maturation of the common octopus in wild populations ndez Nun ez et al., 1996; has been extensively studied (Ferna a et al., 2002; Silva et al., ndez Garc Quetglas et al., 1998; Herna guez de la Ru a et al., 2005; Otero et al., 2007), but 2002; Rodr little attention has been paid thus far to evaluating sexual maturation under rearing conditions (Mangold and Boletzky, 1973; Cerezo et al., 2007). The aim of the present work was to study gonad maturation in males and females reared together and

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separately under similar culture conditions, compared with animals taken from local sheries. The effect of sexual maturation on the digestive gland index (DGI, an indicator of condition; Cerezo Valverde et al., 2008) and under two dietary treatments is also discussed.

J. Estefanell et al. Valverde, 2006). The other oating cage (M:Fsh) was fed a monospecic diet of bogue (Boops boops). The daily feeding rate was 6% of initial biomass for bogue, and 10% for crab. The bogue used in the trials were provided by local farms as discard species from offshore sea-bream cages. The blue crab was purchased from a local trader. The trials lasted 11 weeks in the oating cages (1 November 200822 January 2009) and 8 weeks in the tanks (26 November 200824 January 2009). Mean water temperature and oxygen levels, measured once a day, were 18.4 + 0.48C and 7.1 + 0.2 ppm in tanks and 19.0 + 1.08C and 7.0 + 0.3 ppm in oating cages, respectively. Light intensity, measured at the water surface with a portable luxometer (Model HT170N; Italy), was 30 50 lux in the tanks and 250 300 lux in the oating cages. Octopuses were observed by scuba three times per week; dead animals were removed daily.
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Material and methods


Capture and acclimation of the stock
n (Gran Canaria, Canary Islands). Octopuses were caught in Moga Local shers used cylindrical trawls (1.5 m diameter, 0.4 m high, with a metallic net of 31.6 mm mesh) placed 20 30 m deep. Octopuses were kept on board in open ow-through seawater reservoirs and transported to the laboratory by truck in 500-l square tanks. This operation took 60 80 min and oxygen was provided so that its concentration would not be limiting ( . 12 ppm). Acclimatization was in rectangular 1.5 m3 tanks, and (previously frozen) blue crab (Portunus pelagicus) was provided ad libitum once daily. After 1 week, the octopuses were PIT-tagged subcutaneously on left arm III, using immersion in seawater with 1.5% ethanol (96%) as anaesthetic, following the procedures described by Estefanell et al. (2007). One week later, tagging was veried as successful by reading the PIT digits in all octopuses, prior to the start of the experiment.

Sampling procedure
One tank per treatment (n 5) was randomly sampled in December, and all the tanks were sampled in January. From each cage, six octopuses were sampled randomly in December (by scuba), and all surviving animals at the end of the experimental period. We also dissected any dead octopuses collected each month from the cages that showed no signs of cannibalism. Animals were sacriced by immersion in ice-cold seawater. Finally, during the experiment, 11 males and 1 female from the local shery (wild group) were sacriced after catching to assess the biological parameters in wild populations.

Experimental design
For males and females reared separately, octopuses were kept in circular tanks of 1 m3 and the water level was kept up to 500 l to prevent the octopuses from escaping. The initial density in these tanks was 9.9 + 1.2 kg m23 of water. In all, 11 males were split between two tanks (M1 and M2) and 16 females among three tanks (F1, F2, and F3; Table 1). Where males and females were being reared together (M:Fcontrol and M:Fsh), two galvanized stainless-steel oating cages (3 m long, 1.5 m wide, and 3 m high) with 20 mm mesh size and a total internal volume of 10 m3 were used. More than 100 octopuses, male:female sex ratio 4:1, were placed in each cage, and the initial density was 10.9 + 0.2 kg m23 of water (Table 1). Different rearing systems were used for practical reasons. The relative scarcity of females provided by the local shers (natural sex ratio 4:1), along with the difculty gathering the octopuses, led us to place the single-sex groups in tanks.

Biological parameters
Mortality was recorded daily. After sacrice, all animals were weighed and the sex was determined. The following indices were calculated, and data were expressed in relation to a monthly mean value of each parameter. (i) Reproductive status in males [where WN is the Needham complex + spermatophoric sac weight (in g), WT the testis weight (in g), and W the octopus weight (in g)]: (a) the Hayashi index, H, as modied by Guerra (1975), i.e. HM WN/(WN + WT): (b) the gonadosomatic index, GSI (Otero et al., 2007): GSIM (WN/W 2 WN) 100. (ii) Indices for the reproductive status of females [where WOG is the oviducal gland weight (in g) and WO the ovary weight (in g)]: (a) the Hayashi index, H, as modied by Guerra (1975), i.e. HF WOG/(WOG + WO): (b) the gonadosomatic index, GSI (Otero et al., 2007): GSIF (WO/W 2 WO) 100.

Rearing conditions
Regardless of rearing system, shelters (PVC T-shaped tubes, 160 mm diameter) and shadowing nets were provided (Villanueva, 1995; Hanlon and Messenger, 1996). In tanks, an open ow-through seawater system was adjusted to 500 l h21. All treatments were fed ad libitum 6 d per week under the natural photoperiod. Tank treatments and one oating cage (M:Fcontrol) were fed a control diet based on a 40 60% a Garc a and Cerezo bogue crab mix on alternate days (Garc

Table 1. The rearing system used for the experiment, showing the initial body weight of males (Wm) and females (Wf ) + s.d.
Rearing system Tank Floating cage Treatment Mn Fn M:Fcontrol M:Fsh Number of tanks or cages 2 3 1 1 Number per treatment 11 15 109 105 Number of males 11 0 88 84 Number of females 0 15 21 21 Wm (g) 1 122 + 267 1 041 + 249 1 060 + 326 Wf (g) 865 + 226 916 + 322 888 + 267

Gonad maturation in Octopus vulgaris during ongrowing in culture (iii) Digestive gland index: DGI (WDG/W ) 100, where WDG is the digestive gland weight (in g). In addition, a macroscopic maturation stage (Dia and Goutschine, 1990) was assigned to every octopus collected (I, immature; II, maturing; III, mature; IV, post-reproductive), for both males and females.

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found regardless of treatment. The DGI decreased after mating and the GSI remained static. No mortality was recorded in any of the male-only or female-only treatments, but in cages, mortality reached 76 and 77% for M:Fcontrol and M:Fsh, respectively. The numbers of individuals dissected per treatment, either sacriced or collected dead, nal weight, H index, GSI values, and DGI values are listed in Table 3 for males and in Table 4 for females. Macroscopic maturity stages in males and females from all treatments are shown in Figures 1 and 2, respectively. Males were consistently mature throughout the experiment, and octopuses collected dead, regardless of sex and diet, were mainly in the post-reproductive stage (IV). In contrast, a smaller proportion of the males sacriced was at stage IV. In terms of females, data were absent from random sampling in oating cages in December, and wild specimens (those dissected immediately after capture) were scarce throughout the experimental period. The females sampled from the cages in January were mainly at stage IV, in contrast to females kept separately from males in tanks. In fact, in the tank treatment, immature females represented 40 and 80% of those sampled in December and January, respectively. The DGI (%) of octopuses fed on the control diet was signicantly lower in those at the post-reproductive stage (Figure 3), both males and females. The H index was correlated with DGI in males and females fed the control diet (Figure 4). In females, the DGI increased with sexual maturation, then collapsed after spawning, and in males, it decreased from mature to postreproductive animals. In terms of histology, Figure 5 shows a mature and a postreproductive testis and digestive gland. The lumen was full of spermatozoa in the mature testis (Figure 5a), whereas few spermatozoa and abundant empty spaces in the lumen were indicative of the spermatozoa having been expelled in the post-mating state

Histological and statistical analyses


Histological studies were performed on gonad and digestive gland from sacriced octopuses obtained from the oating cages. The organs were xed in 10% neutral-buffered formalin, embedded in parafn, then stained with haematoxylin and eosin (H&E) for a del Moral, 1993). Micrographs of optical examination (Garc gonads were taken from the parafn sections using a Nikon Microphot-FXA microscope and an Olympus DP50 camera. In terms of the statistics, means and standard deviations were calculated for initial body weight, initial rearing density, and temperature. Data from each treatment were submitted to the Levene test, and when that showed no difference in variance among groups ( p 0.05), individual means were compared using the one-way ANOVA along with the Tukey test for multiple comparisons (Sokal and Rolf, 1995).

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Results
No differences in initial body weight density (Table 1) and rearing temperature were found among treatments. The mean values for each index calculated for males and females are listed in Table 2. Daily observations revealed mating from the start of the experiment. In females, HF decreased with maturity, but then increased sharply after spawning. In males, HM increased with maturity. In females, both GSI and DGI increased with maturity, but collapsed after egg-laying. In males, no immature or maturing stages were

Table 2. Mean value ( + s.d.) for each index calculated for female and male O. vulgaris, the Hayashi index modied by Guerra (H ), the GSI, and the DGI.
Sex Female Maturity stage I II III IV III IV Description Immature Maturing Mature Post-reproductive Mature Post-reproductive H 0.21 + 0.06 0.11 + 0.02 0.05 + 0.02 0.47 + 0.06 0.54 + 0.08 0.78 + 0.05 GSI (%) 0.20 + 0.09 1.06 + 1.15 3.45 + 2.05 0.46 + 0.14 0.39 + 0.10 0.58 + 0.17 DGI (%) 4.07 + 0.48 4.21 + 1.70 4.52 + 1.35 0.98 + 0.32 2.71 + 1.25 1.14 + 0.58

Male

Table 3. The number of male O. vulgaris dissected (sacriced or collected dead, n), weight, Hayashi index modied by Guerra (H ), GSI, and DGI in December and January (mean values + s.d.).
Month December How collected Sacriced Treatment M1 M:Fcontrol M:Fsh Wild M:Fcontrol M:Fsh M2 M:Fcontrol M:Fsh Wild M:Fcontrol M:Fsh n 5 6 6 4 23 18 6 14 12 7 14 12 Weight (g) 1 648 + 529 2 943 + 1 369 2 117 + 596 1 187 + 78 1 400 + 453 1 624 + 609 2 168 + 777 3 021 + 1 373 2 957 + 930 1 883 + 344 2 358 + 1 250 1 697 + 514 H 0.52 + 0.12 0.66 + 0.16 0.65 + 0.16 0.43 + 0.09 0.76 + 0.07 0.77 + 0.10 0.50 + 0.09 0.65 + 0.12 0.59 + 0.11 0.41 + 0.04 0.73 + 0.10 0.80 + 0.06 GSI (%) 0.43 + 0.08 0.37 + 0.10 0.44 + 0.06 0.38 + 0.17 0.61 + 0.13 0.65 + 0.26 0.45 + 016 0.36 + 0.07 0.41 + 0.10 0.33 + 0.09 0.52 + 0.14 0.57 + 0.09 DGI (%) 2.93 + 1.61 1.89 + 1.64 1.74 + 1.16 4.39 + 1.42 1.52 + 0.86 1.20 + 0.40 3.77 + 1.15 1.70 + 0.83 2.52 + 1.62 3.46 + 0.68 1.28 + 0.70 1.06 + 0.37

Collected dead January Sacriced

Collected dead

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Table 4. The number of female O. vulgaris dissected (sacriced or collected dead, n), weight, Hayashi index modied by Guerra (H ), GSI, and DGI in December and January (mean values + s.d.).
Month December How collected Sacriced Treatment F1 M:Fcontrol M:Fsh Wild M:Fcontrol M:Fsh F2 3 M:Fcontrol M:Fsh Wild M:Fcontrol M:Fsh n 5 0 0 0 4 4 10 1 1 1 5 3 Weight (g) 1 348 + 349 831 + 163 861 + 285 2 009 + 188 800 1 120 1 192 923 + 123 745 + 176 H 0.12 + 0.11 0.24 + 0.18 0.37 + 0.21 0.10 + 0.03 0.42 0.45 0.08 0.44 + 0.04 0.55 + 0.03 GSI (%) 2.99 + 2.89 1.53 + 1.57 1.42 + 2.07 0.87 + 0.73 0.75 0.49 0.65 0.35 + 0.07 0.56 + 0.10 DGI (%) 4.73 + 0.44 1.23 + 0.62 0.97 + 0.40 4.92 + 0.64 1.28 0.61 4.71 0.99 + 0.23 1.24 + 0.56

Collected dead January Sacriced

Collected dead

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Figure 3. DGI (%) in octopuses fed the control diet at the stages mature (III) and post-reproductive (IV). Figure 1. Macroscopic maturation of male O. vulgaris in December and January.

Figure 4. Relationship between the DGI and the Hayashi index modied by Guerra (H ) in sacriced males and female O. vulgaris, from all treatments.

Discussion
Under rearing conditions, O. vulgaris breed frequently regardless of the time of year (Mangold and Boletzky, 1973; Iglesias et al., 2000; Estefanell, 2006). In the Canary Islands, O. vulgaris in the wild breed all year-round, with two periods of main activity, one from January to July, with a peak in April, and a second in a et al., 2002). ndez-Garc autumn (OctoberNovember; Herna In the current experiment, males were constantly mature throughout the period of sampling, irrespective of treatment (Figure 1), a nding that supports data obtained under rearing conditions (Mangold and Boletzky, 1973; Cerezo et al., 2007) and several previous reports from wild populations in the Atlantic Ocean ` re et al., 2002; Silva et al., 2002; (Guerra, 1979; Caverivie Carvalho and Sousa Reis, 2003; Oosthuizen and Smale, 2003; guez de la Ru a et al., 2005; Otero et al., 2007) and Rodr

Figure 2. Macroscopic maturation of female O. vulgaris in December and January.

(Figure 5b). Histological examination revealed evidence of spermatogonia, spermatocytes, spermatids, and spermatozoa in mature testes. In the digestive gland, octopuses in the postreproductive state showed degenerative vacuolization in the parenchyma (Figure 5d) compared with that of mature animals (Figure 5c).

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Figure 5. Transverse sections ( 10) of testis and digestive gland at different stages of maturity. (a) Mature testis; (b) post-reproductive testis; (c) mature digestive gland; (d) post-reproductive digestive gland. Mediterranean (Quetglas et al., 1998). In contrast, a large proportion of the females kept isolated from males remained immature or maturing (Figure 2), suggesting that the presence of males may promote female sexual maturation. Indeed, sex steroid hormones have been reported in O. vulgaris of both sexes (DAniello et al., 1996; Di Cosmo et al., 2001), and they could act as pheromones if released to the water, coordinating maturation in both sexes. Therefore, in sex-segregated females, the absence of hormones from the opposite sex could postpone maturation. On the other hand, females matured in cages where the light intensity at the surface was nine times higher than in tanks inside the experimental facilities, contradicting the statement of Mangold (1987) that high light intensity can retard sexual maturation and that low light intensity can stimulate gonad development in aquarium-held octopuses. The few females caught in random sampling in cages during December was probably the result of breeding activity then, because females remain in their dens cleaning, ventilating, and caring for the egg strings (Guerra, 1992). It may also explain the few females caught in the wild before and during the experiment. Progressively lower values of HF were recorded from the start of winter to spring in females living in the wild around Gran a et al., 2002), and deviations from a ndez Garc Canaria (Herna 1:1 sex ratio were attributed to reproductive behaviour or the sampling strategy (Silva et al., 2002). In the cages, large proportions of octopuses in postreproductive state were found (Figures 1 and 2) contrary to the ndings of Cerezo et al. (2007). Daily observations showed mating behaviour throughout the experiment, which agrees with the ndings of Iglesias et al. (2000). Occasionally, two males were seen introducing a hectocotylus into the same female at the same time. Most octopuses collected dead from either treatment were at the post-reproductive stage, which demonstrates a clear relationship between mating processes under rearing conditions and mortality, recorded here for the rst time in oating cages, a ndez Garc supporting earlier observations in tanks (Herna et al., 2002). Histological examination conrmed the maturity stages obtained from macroscopic observation and the Hayashi index. Therefore, the presence of spermatogonia/spermatocytes in the tubular wall and spermatids/spermatozoa in the central lumen are evidence of maturing and mature stages in male O. vulgaris, and empty spaces in the lumen indicate that spermatozoa have guez de la Ru a et al., 2005). been expelled after mating (Rodr On the other hand, the presence of cells at all stages of maturation suggests different phases of gonad development at the various maturity stages of the testes, as found in the testis of maturing or mature Octopus mimus (Avila-Poveda et al., 2009). In the present experiment, female GSI and DGI increased with sexual maturation, but this was not the case for males (Table 2; guez de la Ru a et al., 2005; Otero et al., Silva et al., 2002; Rodr 2007). The signicantly higher DGI in mature than in post-mating O. vulgaris (Figure 3) supports other authors ndings (Otero et al., 2007). A decrease in DGI in post-mating O. mimus (Cortez et al., 1995) and Sepia ofcinalis (Castro et al., 1992) has been documented. In contrast to the ndings of Otero et al. (2007), the correlation between H and GSI was very poor, probably related to the fact that few females in an immature or maturing state were analysed here and that few post-reproductive animals were available to Otero et al. On the other hand and also according to Otero et al. (2007), DGI and H are inversely correlated in both sexes (Figure 4). For egg production, O. vulgaris uses energy directly from food, rather than from stored resources (Rosa et al., 2004; Otero et al.,

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2007). In our work, sexual maturation of both males and females did not seem to be inuenced by diet, because most, irrespective of diet, were in a mature or post-reproductive stage. To conclude, under the experimental conditions described, there was a clear increase in mortality attributable to the reproductive process. Deterioration of the gonads was conrmed by histological examination (Figure 5) and by a massive decrease in digestive gland weight in post-reproductive animals. Consequently, further work needs to evaluate the benets of segregation by sex under rearing conditions if mortality reduction and an increase in protability of octopus culture are desired.

J. Estefanell et al.
de la calidad de la puesta. Minor thesis, University of Las Palmas de Gran Canaria. 137 pp. rez, M., Ferna ndezEstefanell, J., Roo, J., Socorro, J., Afonso, J. M., Sua n de dos aneste Palacios, H., and Izquierdo, M. S. 2007. Evaluacio a de marcaje individual con microchip subcuta sicos y metodolog neo (PIT) en Octopus vulgaris (Cuvier, 1797). In Proceedings of the 11th Spanish Congress of Aquaculture, 1, pp. 655 658. Ed. by A. . Guerra, and C. Perez. ISBN TOMO I: o, A Cervin 978-84-611-9086-7. ndez-Nu n ndez-Gonza lez, C. L., Raya, C. P., and ez, M. M., Herna Ferna as, E. 1996. Reproductive biology of Octopus vulgaris Balguer Cuvier, 1797, from North Western African Coast (218N- 268N). ICES Document CM 1996/K: 15. 19 pp. a del Moral, R. 1993. Te a cnicas de Laboratorio de Anatom Garc gica. Interamericana/McGraw-Hill. 654 pp. Patolo a Garc a, B., and Cerezo Valverde, J. 2006. Optimal proportions Garc of crabs and sh in diet for common octopus (Octopus vulgaris) ongrowing. Aquaculture, 253: 502 511. a Garc a, B., Cerezo Valverde, J., Aguado-Gime a nez, F., Garc Garc a, J., and Herna ndez, M. 2009. Growth and mortality of Garc common octopus Octopus vulgaris reared at different stocking densities in Mediterranean offshore cages. Aquaculture Research, 40: 1202 1212. . 1975. Determinacio n de las diferentes fases del desarrollo Guerra, A sexual de Octopus vulgaris Lamarck, mediante un ndice de n Pesquera, 39: 397 416. madurez. Investigacio . 1979. Fitting a von Bertalanffy expression to Octopus vulGuerra, A n Pesquera, 43: 319 326. garis growth. Investigacio . 1992. Mollusca, Cephalopoda. In Fauna Ibe rica, 1. Ed. by Guerra, A . s, J. G. Gonsa lbez, A M. A. Ramos, J. A. Tercedor, X. Belle n, et al. Museo Nacional de Guerra, E. Macpherson, F. Mart Ciencias Naturales, CSIC, Madrid. 327 pp. Hanlon, R. T., and Messenger, J. B. 1996. Cephalopod Behaviour. Cambridge University Press, Cambridge, UK. 232 pp. a, V., Herna ndez-Garc ndez-Lo pez, J. L., and Castro Herna ndez, J. J. 2002. On the reproduction of Octopus vulgaris Herna off the coast of the Canary Islands. Fisheries Research, 57: 197 203. nchez, J., Otero, J. J., Moxica, C., and Lago, Iglesias, J., Fuentes, L., Sa M. J. 2006. First feeding of Octopus vulgaris Cuvier, 1797 paralarvae using Artemia: effect of prey size, prey density and feeding frequency. Aquaculture, 261: 817 822. nchez, F. J. 2004. Iglesias, J., Otero, J. J., Moxica, C., Fuentes, L., and Sa The completed life cycle of the octopus (Octopus vulgaris, Cuvier) under culture conditions: paralarval rearing using Artemia and zoeae, and rst data on juvenile growth up to 8 months of age. Aquaculture International, 12: 481 487. nchez, F. J., Bersano, J. G. F., Carrasco, J. F., Dhont, J., Iglesias, J., Sa Fuentes, L., Linares, F., et al. 2007. Rearing of Octopus vulgaris paralarvae: present status, bottlenecks and trends. Aquaculture, 266: 1 15. nchez, F. J., Otero, J. J., and Moxica, C. 2000. Culture of Iglesias, J., Sa octopus (Octopus vulgaris, Cuvier): present, knowledge, problems and perspectives. Recent Advances in Mediterranean Aquaculture Finsh Species Diversication. Cahiers Options diterrane ennes, 47: 313 322. Me Mangold, K. 1983. Food, feeding and growth in cephalopods. Memoirs of the National Museum of Victoria, 44: 81 93. Mangold, K. 1987. Reproduction. In Cephalopod Life Cycles. 2. Comparative Reviews, pp. 157 200. Ed. by P. R. Boyle, and G. J. Pierce. Academic Press, London. 441 pp. Mangold, K., and von Boletzky, S. 1973. New data on reproductive biology and growth of Octopus vulgaris. Marine Biology, 19: 7 12. Navarro, J. C., and Villanueva, R. 2000. Lipid and fatty acid composition of early stages of cephalopods: an approach to their lipid requirements. Aquaculture, 183: 161 177. Navarro, J. C., and Villanueva, R. 2003. The fatty acid composition of Octopus vulgaris paralarvae reared with live and inert food:

Acknowledgements
The work was nanced by JACUMAR Spanish National Plans for n del engorde de pulpo Octopus vulAquaculture (Optimizacio garis, 2007-09). We thank the CANEXMAR staff for kindly providing the bogue feed for the work, and the reviewers and editors for their valued input.

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