Aquaculture 254 (2006) 293 300 www.elsevier.

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Common octopus (Octopus vulgaris Cuvier, 1797) juvenile ongrowing in floating cages
Carmen Rodrguez a,, Jos F. Carrasco a , Juan C. Arronte b , Montse Rodrguez b
b

Consejera de Medio Rural y Pesca, Direccin General de Pesca, Centro de Experimentacin Pesquera, El Muelle S/n, 33760, Castropol, Asturias, Spain Universidad de Oviedo, Facultad de Medicina, Departamento de Gentica, C/ Julin Clavera S/n, 33071, Oviedo, Asturias, Spain Received 28 October 2004; received in revised form 26 October 2005; accepted 28 October 2005

Abstract The study of Octopus vulgaris culture has recently undergone a significant increase in Spain. The need to diversify the available number of farmable marine species, taking into consideration biological and economical aspects, makes the common octopus a potential candidate for industrial culture. In this article, results for four growth trials using O. vulgaris, which were held in captivity in Asturias (Spain) during 2002 and 2003, are presented. A rectangular floating cage of 4 m3 containing PVC tubes as refuges was used in each trial. Each trial lasted 3 months. A mixed diet of fish, crabs and mussels was used to feed the animals. Different initial densities (24.5, 10.7, 12.6 and 11.9 kg m 3) were tested. These produced 41.0%, 17.0%, 36.0% and 18.7% mortality, 0.51%, 1.46%, 0.64% and 1.54% specific growth and 7.02, 29.39, 8.69 and 33.18 g daily weight increase, respectively. There were no serious problems of cannibalism. Best results concerning growth and survival were obtained in summerearly autumn, with a mean water temperature of 15.9 1.6 and 16.7 1.7 C. Time of the year and diet seem to be the main factors that have influence on O. vulgaris commercial culture success. 2005 Elsevier B.V. All rights reserved.
Keywords: Aquaculture; Culture; Farming; Floating cage; Octopus vulgaris; Ongrowing

1. Introduction The common octopus (Octopus vulgaris) has a worldwide distribution in tropical, subtropical and warm waters. It is a benthic, neritic species occurring from the coastal line to the outer edge of the continental shelf, at depths of up to 200 m and in very diverse habitats (Guerra, 1992). In some Octopodidae species, hatchlings are immediately benthic like the adults and

Corresponding author. Tel./fax: +34 985 365020. E-mail address: menchurr@princast.es (C. Rodrguez). 0044-8486/$ - see front matter 2005 Elsevier B.V. All rights reserved. doi:10.1016/j.aquaculture.2005.10.053

thus are referred to as juveniles (Villanueva, 1995). But in the case of O. vulgaris this species has a planktonic post-hatching stage termed paralarvae (Young and Harman, 1988). It is considered to be an opportunistic predator (Mangold, 1983) that feeds on a wide variety of species (Mather, 1991). The shelter, known as home or den (Mather, 1994), plays an important role in octopus behaviour and biology (Hanlon and Messenger, 1996) and this data must be taken into account for the culture of this species. The need to diversify marine aquaculture products, which have reached market saturation in the case of some species, has stimulated the farming of new

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species, such as O. vulgaris (Vaz-Pires et al., 2004). This species meets many of the characteristics for industrial culture, such as rapid and easy adaptation to captivity conditions (Iglesias et al., 2000), short life cycle (up to 24 months) (Mangold, 1997), rapid growth and high fecundity (Mangold, 1983). Added to this are the acceptance of non-living, non-motile foods (Boucaud-Camou and Boucher-Rodoni, 1983), high food conversion rate (Mangold and Boletzky, 1973), high market prices and an increasing demand in European, South American and Asian countries. Growth from egg to sub-adult has been attained successfully only at laboratory and experimental scale (Itami et al., 1963; Villanueva, 1995; Carrasco et al., 2003; Iglesias et al., 2004), which means that octopus aquaculture is currently restricted to the ongrowing of sub-adults captured by fisheries. Several ongrowing trials have been carried out in tanks (Iglesias et al., 1997; Otero et al., 1999; Rodrguez and Carrasco, 1999; Garca and Aguado, 2002) and in floating cages (Rama-Villar et al., 1997; Tun et al., 2001; Oltra et al., 2004) with different results regarding growth and food intake rates. O. vulgaris must be reared with natural food due to the fact that specific formulated foods are unavailable, however cephalopods have shown their adaptability to feed on inert food (Lee et al., 1991). Octopuses usually have been fed on crabs, fish and molluscs in an attempt to replicate its natural diet (Guerra, 1978; Mather, 1991). Natural food for O. vulgaris growth is not standardized and there are seasonal and geographical differences in the diet (Hanlon and Messenger, 1996). Water temperature also has great influence on feeding (Mangold and Boletzky, 1973). The differences observed in growth and food intake between the above mentioned trials may be a result of these facts. The main objectives of this work were to ongrow O. vulgaris in floating cages and provide information about the production responses of the common octopus under confinement conditions at different times of the year. For this, the octopuses were fed with discarded and low market value species, which make up the basis of its natural diet. Success would enable the marketing of octopus with high weights at high prices when demand is greatest. At the same time, the ongrowing in floating cages would entail a reduction in terms of costs and maintenance with regard to the culture in inland facilities, since individuals are reared directly in the sea. However, the cage farming has a series of disadvantages such as fuel and transportation costs, boats, losses of individuals due to bad weather conditions and subadult supply shortage.

2. Material and methods For the trials, a rectangular floating cage (1.70 1.20 1.95 m length, height and width, respectively) with a capacity of approximately 4 m3 was designed (Fig. 1). The cage is made entirely of galvanized steel with the exception of two fibreglass floats and octopus refuges, which are made of polyvinyl chloride (PVC). It is made up of two independent pieces, one external and one internal. The external has a base where the mobile internal fits. In the internal piece, 98 PVC tubes measuring 36 cm long by 16 cm in diameter were placed as shelters. A galvanized steel net was welded around the cage. The mesh of the metallic net is circular and it is 20 mm in diameter. The cage was anchored at Tapia de Casariego (4334N; 657W), a well-protected port in Asturias (Spain). Four trials were carried out from April to July and from July to October in two consecutive years (2002 and 2003). O. vulgaris specimens were captured from their natural environment by professional fishing boats using lobster pots. Until arrival at the port, they were kept in tanks provided with flowing seawater. In all the trials prior to stabling, the sexes were separated. Only males were used thus avoiding the fertilization and natural death of females, which occurs when the eggs finally hatch. The photoperiod was natural and salinity varied from 33 to 35. All trials lasted 86, 96, 90 and 91 days, respectively. The experimental conditions are shown in Table 1. The octopuses were fed from Monday to Friday with a mixture of fish and live crabs and molluscs since these

Fig. 1. Floating cage used during the trials; (a) external piece with two fibreglass floats, (b) internal piece with PVC tubes as hiding places.

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are the main foods of O. vulgaris in the sea (Guerra, 1978). On Friday, the amount of food provided was double the daily ration and it was a base of live crabs and mussels to reduce the amount of uneaten food since octopuses present external digestion when feeding on crabs, leaving a whole and empty exoskeleton (Nixon, 1984). Food supply was based on low-value feeds, including blue whiting (Micromesistius poutassou), horse mackerel (Trachurus trachurus), bogue (Boops boops), grey mullet (Mugil sp.), mackerel (Scomber scombrus), pouting (Trisopterus luscus), sardine (Sardina pilchardus), common crab (Carcinus maenas) and mussel (Mytilus sp.). Food was supplied during daylight hours only, and was distributed homogeneously, as best as possible. The amount of food supplied ranged between 9% and 3% of the total octopus biomass, varying according to amounts of uneaten food. After each trial, the octopuses were examined to determine the occurrence of skin lesions at the edge of the mantle, as well as the presence of specimens with eaten arms, indicative of cannibalism. Weight was used to assess the change in size of O. vulgaris, since it is a more sensitive measure of growth in this species than length (Nixon, 1966). For each experiment, the following indices were calculated: Specific growth rate (SGR): (lnWaf lnWai)100/t Feed efficiency (FE): IW 100/IF Daily weight increase (DWI): (Waf Wai)/t Food conversion (FC): IF/IW where Wai = average initial weight (g); Waf = average final weight (g); t = time in days; IF = ingested food (g)
Table 1 Initial experimental conditions Year 2002 Exp. 1 Period of fattening Number of individuals Initial biomass (kg) Initial density (kg m 3) Initial mean weight (g) Range (g) Mean temperature (C) Range (C) Food supplied Initial (%) Final (%) Fish (%) Crab (%) Mussel (%) 17/0412/07 90 98.1 24.5 1090 220.8 9001400 14.3 1.1 12.017.0 8 3 44 33 23

(difference between food supplied and food not consumed); IW = increase in weight (g) (difference between final and initial biomass). In the calculation of FC, IF was estimated by taking into account the bivalve shells, fish heads and bones, empty crab carcasses and the IW including the weight of the octopuses that died during each experiment. Anderson-Darling normality tests were carried out to test whether final weight data came from a normal distribution. Since the assumption of normality was not met, data were compared using the Mann-Whitney U-test. Differences in mortality between trials were examined by 2 test. Statistical analyses were done with Minitab version 13.0 software (Minitab Inc.). 3. Results The final experimental conditions are shown in Table 2. In trial 1, mortality rate was constant during the whole ongrowing process, with a peak of instant mortality on day 80 in individuals of 23 kg, resulting in a mortality rate of 41.1%. In trial 2, the casualties occurred during the first month, after which there were practically none, resulting in a mortality of 17.0%. In trial 3 a constant loss of octopuses in the ongrowing period was observed, reaching the higher values in the last 10 days and resulting in 36.0% mortality. Finally, in trial 4 a peak in mortality was observed at day 60, with no more losses afterwards resulting in a final mortality of 18.7% (Fig. 2). In all trials the weekend period did not produce detectable changes in mortality rate. Two groups can be considered: the first, made up of the springearly summer trials (1 and 3) and the second,

Year 2003 Exp. 2 25/0729/10 47 43.1 10.7 917 231.6 6001600 15.5 1.6 13.018.0 9 5 50 35 15 Exp. 3 10/0409/07 50 50.4 12.6 1009 242.1 8001400 15.3 1.2 14.017.0 8.7 6.6 30 37 33 Exp. 4 23/0722/10 48 47.8 11.9 996 237.9 7001300 16.7 1.7 13.020.0 8.3 3.3 54 26 20

Values S.D.; Initial and final food supplied = % total octopus biomass.

296 Table 2 Final experimental conditions

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Year 2002 Exp. 1 Days of trial Number of individuals Final biomass (kg) Final density (kg m 3) Final mean weight (g) Range (g) DWI (g day 1) SGR (%) FC FE (%) Mortality rate (%) Values S.D. 86 53 89.8 22.5 1694 461.3 7002800 7.02 0.51 10.5 9.5 41.1 Exp. 2 96 39 145.8 36.5 3739 889.5 15005500 29.39 1.46 4.03 24.7 17.0 Exp. 3 90 32 53.7 13.4 1790 604.5 7003400 8.69 0.64 9.2 10.8 36.0

Year 2003 Exp. 4 91 39 155.3 38.8 3982 818.7 16005000 33.18 1.54 3.8 25.8 18.7

composed of the summerearly autumn trials (2 and 4). Statistically significant differences in mortality were observed between these time periods (2, p b 0.01). However, no significant differences were recorded between both springearly summer trials and both summerearly autumn trials (2, p N 0.05). However, there were significant differences in the final weight between the two groups (U, p b 0.01), but not between both summerearly autumn and both springearly trials (U, p N 0.05). In trial 1, during the ongrowing process, some individuals lost weight, resulting at the end of the experiment in a total biomass lower than at the outset. That is, the loss of biomass was not offset because of both the low growth rate and high mortality. Marked differences between experiments were observed, the summerearly autumn trials showing both

better results regarding SGR and FC than the spring early summer experiments. An improvement in the incidence of skin lesions was observed with decreasing animal densities, falling from 77% in the first trial to 23%, 20% and 12% in the other three, respectively. Mortality due to skin lesions was not observed. In all cases, the surface of the skin lesion was smaller than 2 cm2. 4. Discussion Despite the fact that O. vulgaris continues to grow in captivity, and in relatively high densities, this species is inherently territorial and requires a minimum space for its development and growth. Likewise, since octopuses need a home for taking shelter proper hiding places should be used in ongrowing experiences. Rama-Villar

45 40 35

Mortality (%)

30 25 20 15 10 5 0 0 10 20 30 40 50 60 70 80 90 95

Days
Exp.1 Exp.2 Exp.3 Exp.4

Fig. 2. Mortality of Octopus vulgaris recorded in the four ongrowing trials.

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et al. (1997) pointed out that the presence of shelters seems to improve the growth rate. O. vulgaris shows low tolerance to concentrations of salts lower than 29 30 and requires fairly large quantities of clean, oxygenated seawater (Boletzky and Hanlon, 1983). However, in our trials it was not considered an important fact since the cage was placed in oceanic waters. Salinity must be taken into account when floating cages are placed in areas with salinity fluctuations due to freshwater. During the trials, to check whether there was sufficient oxygen within the cage to maintain the metabolic rate of the octopus, irregular samplings of dissolved oxygen levels inside and outside the cage were conducted, displaying no differences in levels of oxygen (unpublished data). Studies on water circulation within the cages could be very useful to see if they might improve the conditions for continued growth. In all trials, cannibalism was not a significant problem, although it has been found to be problematic when food is not sufficiently supplied or when octopuses display a marked difference in size (Hanlon and Messenger, 1996). These situations can be partially avoided by supplying enough food, separating individuals according to sex and stabling with octopuses of similar sizes. High densities might provoke competition for space, stressing the animals and preventing the smaller individuals from eating. This was observed in trial 1, where the initial density was high and the amount of food consumed from the outset was very low. Consumption did not change throughout the trial despite the fact that the number of octopuses in the cage decreased markedly. The high percentage of skin lesions recorded in this experiment seems to be due to the high initial density. Skin lesions were most likely produced by friction with home walls and/or by the interactions among octopuses in a high-density culture. Damaged skin is invaded by opportunistic bacteria, which kills untreated octopuses (Hanlon et al., 1984). Although the authors do not consider it a serious problem in the farming of this species, it must be taken into account since damaged octopuses present lower market prices. On the other hand, a negative influence on octopus growth cannot be ruled out. The low growth and high mortality observed in trial 1 cannot be solely attributed to the initial density. In fact, despite that initial density was practically reduced to half after the first trial, low survival and growth were attained in the trial carried out the following year during the springearly summer. Considering all the trials together, it becomes evident that initial culture density does not itself explain the lower growth rates. Time of the year (e.g. photoperiod and water temperature) and/

or the quality and composition of the food provided also seem to affect octopus growth. When comparing results it is important to distinguish between trials carried out in tanks and those carried out in floating cages. In tanks, Aguado and Garca (2002) recorded mortalities of 43% and 50% at the end of their assays, while Iglesias et al. (1997, 2000) obtained lower mortality (5.6% and 10.8%) in the laboratory than that recorded in these four trials. However, these authors, following Nixon (1969), did not consider the mortality due to the spawning of females and so it is difficult to compare the results. In floating cages, the rate and mortality pattern recorded by Tun et al. (2001) were similar to the present study. In their springearly summer trials they obtained mortalities from 34.5% to 51.5%, while during the summer trials, mortality ranged from 16.0% to 25.0%. Rama-Villar et al. (1997) reported mortality rates of 5.7% after 4 months. Oltra et al. (2004) recorded a mortality of 40.6% and 43.3% in two trials carried out in the Mediterranean Sea. These authors pointed out that the high sea temperatures recorded during certain times of the ongrowing, along with the location of the floating cage, seem to have affected the survival of the individuals. Above 23 C there is an increase in weight loss and mortality rate (Aguado and Garca, 2002). Fasting does not seem to have an effect on mortality neither in tanks (Garca et al., 2003) nor in floating cages (this study). In all the trials feed efficiency was lower than that observed by Garca and Aguado (2002) in a period of 5 months of ongrowing and with octopuses distributed individually in tanks and fed to satiation. In the present study, FE was higher in summerearly autumn trials, a situation which in principle could be due to a higher temperature, although the authors mentioned above did not find significant correlation between FE and temperature or body weight. In the same way, food conversion is also independent of temperature and of size of the individuals, but appears to be dependent upon diet quality and shows high individual variability (Mangold and Boletzky, 1973). If we start with the assumption that diet quality was similar in all trials and that temperature does not affect FE or FC, the differences found in these indices between both times of the year may be due to physical factors related to the time of the year, such as luminosity or photoperiod, or with the physiological state of the individuals. The best growth rates were obtained in the summer early autumn trials. These results agree with results obtained from studying O. vulgaris in tanks and under laboratory conditions (Nixon, 1966; Mangold and

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Boletzky, 1973), where higher temperatures cause greater food intake and growth (Mangold, 1983). Our values of SGR in the summerearly autumn trials were higher than those obtained by Iglesias et al. (2000) (0.77% and 0.95%) with specimens displaying an average weight of 873 and 883 g, respectively, at temperatures ranging from 13 to 16 C. This was slightly higher than that recorded by Rodrguez and Carrasco (1999) (1.3%) after 5 months and with water temperature varying from 11 to 18 C. In both trials octopuses were fed on crustaceans, fish and mussels. Smale and Buchan (1981) obtained higher values (from 0.64% to 2.08%) and they attributed faster growth of the specimens to both high mean temperature and food provided. These differences might be due to the fact that all the trials mentioned above were carried out in tanks and that initial weights were lower than that used in this study. However, the SGR results were better than those recorded in the laboratory by Cagnetta and Sublimi (1999) (1.07%) and Garca and Aguado (2002) (0.73%) using a monodiet of fish and slightly lower than Cagnetta and Sublimi (1999) feeding with a monodiet of crab (Carcinus mediterraneus) (1.72%) and tattler (Illex coindetii) (1.77%). Single foods should only be used during short periods during ongrowing, since mixed diets meet the octopus's nutritional requirements better than a monodiet (Smale and Buchan, 1981; Cagnetta and Sublimi, 1999). In floating cages, Oltra et al. (2004) recorded values of SGR of 1.38% and 1.69% using specimens with an average body weight of 532 and 783 g, respectively. Growth is dependent upon body weight, diet and temperature (Aguado and Garca, 2002) and seems to decrease when body weight increases. There is no influence of fasting either on the SGR or on the growth but it does seem to affect FC (Garca et al., 2003). On the other hand, the growth rates would probably have been better in all trials if octopuses were fed mainly on crabs instead of fish, given the different intake and conversion rate that this species presents for this kind of food (Smale and Buchan, 1981; Mangold, 1983; Aguado and Garca, 2002). In all the trials, it was observed that live crabs were the first to be taken and were devoured immediately. The preference of O. vulgaris for crabs compared to other types of food could be explained not only by visual stimuli but also by both the chemical and the textural qualities of the food (Lee et al., 1991). Proteins play a decisive role in the growth and as a source of energy in O. vulgaris since this species has a special capacity to digest and assimilate protein rapidly (Lee, 1994) along with a low and inefficient lipid digestion (O'Dor et al., 1983). Although

frozen crab can be purchased and stored at a lower cost than live crabs, if there is enough supply, the use of live food is advisable since it produces higher growth rates (DeRusha et al., 1989). Cephalopod growth rates are high regardless of diet composition, providing that it takes part of its natural diet (Forsythe and Van Heukelem, 1987). For optimum growth results, Otero et al. (1999) have recommended that initial density should not exceed 10 kg m 3. However, such data must be treated carefully since they were obtained in two tanks with initial densities of 20 and 10 kg m 3 and working with a small sample size (37 and 18 individuals, respectively). As we mentioned above, initial density does not seem to be the main factor for obtaining satisfactory results. For industrial culture, and from an economic point of view, the growth rate is not the only factor that determines the profitability of the cages (Garca et al., 2001). According to Aguado and Garca (2002), the optimum temperature range for commercially viable growth is 1621 C. However, even at a given temperature, growth rate is very variable between the different specimens. This variability in growth may be attributed to both genetic and behavioural facts. This variation is in agreement with the results of this study, where, within the same trial, the final weights varied from 1500 to 5500 g, in agreement with trial 2. The large variation in individual growth rates is due to extreme differences in the way individuals respond to the environmental conditions (Smale and Buchan, 1981). In view of the results obtained in the present study, temperature seems to have a positive effect on growth. In accordance with the results obtained from SGR, the best results for DWI were obtained in the summerearly autumn (29.39 and 33.18 g day 1) than in springearly summer trials (7.02 and 8.69 g day 1). Similar results were obtained by Tun et al. (2001), reporting weight increases that varied between 9.22 and 13.48 g day 1 in the late winterearly summer trial and from 15.65 to 21.58 g day 1 in the summer trials. In the summer early trials, the daily increase obtained was slightly lower than that recorded in the laboratory by Iglesias et al. (1997) (36.5 g day 1) after 10 months and beginning with individuals of initial mean weight of 1340 g. However, these results were better than those obtained by Rodrguez and Carrasco (1999) (27.8 g day 1), although in this case with considerably lower densities (1.0 kg m 3). From the four experiments carried out in 2002 and 2003, it can be concluded that industrial ongrowing of sub-adult wild individuals in floating cages shows promising results. Beginning with octopuses weighing

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around 1 kg, at an initial density of 1012 kg m 3, temperature ranging between 15 and 21 C and a mixed diet with a minimum of 25% crustaceans, it was possible to attain individuals of 34 kg in ongrowing cycles of 3 months, with a growth rate close to 1.5% and with less than 20% mortality. However, from an economic point of view, the farming of O. vulgaris is limited by the capture of wild individuals from the sea. Several facts can affect the specimen supply such as bad weather conditions or changes in the target species by the inshore fleet. Therefore, the future of octopus ongrowing involves both the improvement and standardisation of paralarvae culture to reduce the production costs. Acknowledgements This work was funded by JACUMAR Spanish National Plans for Aquaculture. The authors would like to thank R.M. Babbitt and J. Bell for their assistance in editing and to J. Iglesias for his technical assistance provided throughout this study. References
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