Aquaculture Potential of the Common Octopus

www.elsevier.com/locate/aqua-online

Aquaculture 238 (2004) 221–238

Aquaculture potential of the common octopus

(Octopus vulgaris Cuvier, 1797): a review

Paulo Vaz-Pires*, Pedro Seixas, Alexandra Barbosa

ICBAS-Institute of Biomedical Sciences Abel Salazar, University of Porto, Largo Prof. Abel Salazar, 2,

4099-003 Oporto, Portugal

CIIMAR-Interdisciplinary Centre for Marine and Environmental Research, Rua dos Bragas, 289,

4050-123 Oporto, Portugal

Received 17 February 2004; received in revised form 4 May 2004; accepted 7 May 2004

Abstract

The potential for aquaculture of the cephalopod species Octopus vulgaris is evaluated, taking into

consideration biological and physiological characteristics, as well as some economic and marketing

aspects, which may be relevant for the future development of octopus farming. O. vulgaris, a

widespread, strictly marine species meets many of the requirements to be considered as a candidate

for industrial culture: easy adaptation to captivity conditions, high growth rate, acceptance of low-

value natural foods, high reproductive rate and high market price. The life cycle from eclosion of

eggs to settlement or beginning of the benthonic adult phase is not commercially viable, but the

published results from laboratory and pilot scales are promising. Comments are also made on general

research lines needed to improve the use of octopus as farmed species in the future.

D 2004 Elsevier B.V. All rights reserved.

Keywords: Octopus vulgaris; Reproduction; Paralarvae; Ongrowing

1. Introduction: the cephalopods

Cephalopods are considered as the most active and specialised class of molluscs.

They may have a chambered shell (e.g., Nautilus), an internal shell, as in squid (e.g.,

0044-8486/$ - see front matter D 2004 Elsevier B.V. All rights reserved.

doi:10.1016/j.aquaculture.2004.05.018

* Corresponding author. Instituto de Ciencias Biomedicas de Abel Salazar, Universidade do Porto, Largo

Prof. Abel Salazar. 2, 4099-003 PORTO, Portugal. Tel.: +351-222062200; fax: +351-222062232.

E-mail address: [email protected] (P. Vaz-Pires).

P. Vaz-Pires et al. / Aquaculture 238 (2004) 221–238222

Loligo) and cuttlefish (e.g., Sepia) or no shell, as in octopods (e.g., Octopus, Eledone).

They are almost all fast-swimming carnivores and live pelagically. Cephalopods share

certain characteristics with more highly developed vertebrates, such as eyes with lens,

pupil and eyelid, a well-developed nervous system and the capability to learn (de

Groot, 1995).

Only a few of the cephalopod species are commercially fished on a large scale

(Kreuzer, 1984). Squid is by far the main cephalopod species, representing 73% of

cephalopod world catches. Cuttlefish is the second and octopus the third, with 15% and

8.8%, respectively. Cephalopod total landing reached a peak of 3.6 million tonnes in 2000.

As regards Octopus catches, Morocco is the world leader with 35% of the total production,

followed by Japan, Thailand, Spain and Mexico (FAO, 2003a).

The main cephalopod consuming countries are Japan, Korea, Argentina, Taiwan and

China, followed by a group that includes Spain, Portugal, Morocco, Mauritania, Greece

and Italy (Baldrati, 1989). Such geographical preference is associated with, but does not

exactly match, the proximity of cephalopod fishing areas, due to imports and consumption

or processing traditions.

During the second half of the last century, Octopus vulgaris and other cephalopods

were considered as less conventional resources, and consequently, the capture of these

species was recommended as a way of diversifying the fishing effort (Pedrosa-Menabrito

and Regenstein, 1988).

Cephalopod fisheries are among the few which still show some local potential for

expansion. As ground fish landings have declined globally, cephalopod landings have

increased (Caddy and Rodhouse, 1998). These authors also postulated that the heavy

fishing pressure on finfish stocks could induce a reaction from the ecosystem that could

include increases in cephalopod abundance, apart from the increased market demand for

these species. Conclusions were, however, based mainly on squid fisheries.

In a review published at the end of the 1980s, Boucaud-Camou (1989) indicated four

possible directions for the marketing of farmed cephalopods: direct consumption in

countries where the value is high (Japan, Spain, Italy, France and Portugal, which are

consequently among the first countries where the aquaculture of some species is being

attempted); the production of juveniles for natural stock reconstitution; neuro-physiolog-

ical, for work on giant neurological cells (mainly squid); and finally as ornamental species.

These remain the main possibilities, since the culture of these species is only moderately

developed.

Nowadays, there is a renewed interest in the farming of new species, stimulated by a

need to diversify the marine farming industry, which is suffering from relative market

saturation for some species like sea bass (Dicentrarchus labrax) and sea bream (Sparus

aurata). A high proportion of the first farming trials with new species, for example

Senegalese sole (Solea senegalensis) took place in southern Europe, notably Spain and

Portugal (Dinis et al., 1999; Aragao et al., 2004); other trials include the black spot

seabream (Pagellus bogaraveo) (Peleteiro et al., 2000). This was also the case of octopus

farming.

Although the Octopus genus includes approximately 200 species, this review will focus

mainly on O. vulgaris Cuvier, 1797 (order Octopoda, suborder Incirrata), which is one of

the most important species in terms of landings and commercial value.

P. Vaz-Pires et al. / Aquaculture 238 (2004) 221–238 223

2. Octopus characteristics

O. vulgaris is a benthic, neritic species occurring from the coast line to the outer edge of

the continental shelf, in depths from 0 to 200 m, where it is found in a variety of habitats,

such as rocks, coral reefs and grass.

In what concerns the behaviour of octopuses as predators in nature, a complete

overview was published by Mather (1993). It was concluded that young O. vulgaris

do not normally modify their movement in the presence of others of the same species;

they do not appear to attract them, they maintain an individual distance and have

specific colours and postures for communication; they do not defend any area, and

they do not stay in a location very long. They occupy a home range for several days

and then move. They can be described as exploratory and opportunistic, but inactive

(Mather and O’Dor, 1991).

Most of the bibliographical data on octopus behaviour and biology is based on

laboratory observations (Nixon, 1966; Wells et al., 1983; Andreu-Moliner and Cachaza,

1984), but some data have been taken directly from octopus caught in nature (Mangold

and Boletzky, 1973). The role of the home in the behaviour of octopus in tanks, including

observations on when and how they occupy brick pots and plastic buckets, was described

by Boyle (1980). This kind of data is important for future aquaculture engineering

dedicated to this species, namely for appropriate tank and home design. O. vulgaris, like

many other species of octopus, ejects shells and other prey remains from the den (home);

this can be considered an advantage in culture, as the remains do not foul the den

(Anderson et al., 1999).

General biometry data, including the relationship between live body weight and total

and dorsal mantle length were published by Nixon (1970), both for specimens kept in

captivity and for animals directly collected from nature. The biometry of several octopus

species is also the subject of articles by Guerra and Manrıquez (1980) working on

Mediterranean octopus caught near Barcelona (Spain), Cunha and Pereira (1995) on O.

vulgaris from Azores Islands (Portugal) and Mangold (1998) for Eastern Atlantic Ocean

and Mediterranean individuals.

3. Octopus aquaculture

3.1. General characteristics

The short life cycle of 12–18 months, rapid growth of up to 13% body weight per day

and food conversion rates of 15–43% are considered the most relevant basic character-

istics which have influenced O. vulgaris culture (Mangold and Boletzky, 1973; Mangold,

1983; Navarro and Villanueva, 2003). Octopus shows a rapid and easy adaptation to life in

captivity (Iglesias et al., 2000a) in aquaria, cylindrical–conical containers, raceways and

floating cages. This includes a high resistance to transport and handling stresses, easy and

rapid feeding in tanks, and the rapid onset of reproduction behaviour (Villanueva, 1995).

Handling of these species, however, can be more complex due to their ability to attach

to any surface, for example in weighing operations. Escape from tanks can be avoided by


the use of a porous surface layer surrounding the tank walls above water level, like foam,

which prevents suckers from attaching to the walls.

Iglesias et al. (2000a) published a very complete review on experiments performed at

the Spanish Institute of Oceanography, based in Vigo (Galicia, Spain). The experiments

included reproduction, paralarvae rearing and ongrowing of subadults at different densities

and separated by sexes.

3.2. Water requirements

The most important water quality parameters are temperature, salinity, pH, O2,

ammonia (NH3), nitrite (NO2�) and nitrate (NO3

�). In open systems, only temperature

and salinity are likely to fluctuate rapidly, whereas in closed systems, the other parameters

are more likely to vary (Boletzky and Hanlon, 1983).

Octopus is a strictly marine species, showing very low tolerance to low concentrations

of salts. O. vulgaris live in nature at salt concentrations of around 35 g l� 1; their minimum

salt concentration is around 27 g l� 1 (Boletzky and Hanlon, 1983). This means slight

fluctuations due to freshwater (e.g., proximity of rivers, strong rain or freshwater from

natural subterranean layers) can be fatal for them.

Preliminary results about post-prandial ammonia production have been obtained in

individual octopus (O. vulgaris) and correlated with the protein intake (Cerezo et al.,

2003). It seems that ammonia excretion is very important in this species compared with

others, like sea bass and gilthead sea bream. Ammonia production per body weight was

found to be much higher in octopuses in some cases. Post-prandial oxygen consumption

after a single meal, with crabs and until satiation, was also recently studied by Cerezo and

Garcıa Garcıa (2004) in common octopus with body weights between 0.22 and 3.26 kg

and at temperatures of 13.8 and 22.2 jC during a period of 3 days. These authors observed

an approximate twofold increase in oxygen consumption, with the maximum value being

attained 6–16 h after the meal ingestion. Ammonia and oxygen are thus important

parameters to be taken into account when planning octopus water systems.

Ongrowing temperature should be kept ideally between 10 and 20 jC, but growth is

higher at higher temperatures in this range.

This species shows a preference for live food, but it also accepts dead whole marine

organisms. Thus, the water systems should be designed in order to facilitate self-cleaning,

due to the high amount of residues produced, like crustacean shells and fish bones. This

will help to keep the quality of the water at an acceptable level.

3.3. Reproduction

O. vulgaris produces an estimated number of 100000–500000 eggs per female

(Mangold, 1983). Iglesias et al. (1997) obtained a maximum number of 605000 eggs in

their reproduction experiments with octopus. The reproduction stocking comprised a 1:1

ratio of males and females, with water temperature and salinity conditions established in

the range of 13–20 jC and 32–35 g l� 1, respectively.

Reproductive behaviour is shown by the copulatory activity of the males, which insert

the hectocotylus into the internal mantle cavity of the females. When the latter are ready to


deposit the spawn, they hide in dens, placing the clusters on the walls and roofs of the

tubes or boxes (Iglesias et al., 2000a).

Usually, females take care of the eggs alone, and then die when the eggs finally

eclode. The spawning season depends on the region. Two spawning peaks per year can

be observed throughout its distributional range: In the Mediterranean and the Inland Sea

of Japan, the first occurs in April/May, corresponding to the group migrating in shore

in spring (most important in the Mediterranean) and the second in October,

corresponding to the group migrating in autumn (most important in Japan) (FAO,

2003b).

Temperature is described as one of the primary factors mediating embryonic develop-

ment in cephalopods (Boletzky, 1989). In the Mediterranean, Villanueva (1995) observed

a period of 34 days after the onset of spawning until the first hatched paralarvae, when the

water temperature was raised to 20F 1 jC. Iglesias et al. (2000a) under laboratory

conditions, observed that in Galicia, Spain the spawning period occurs between February

and November and the embryonic development lasts between 80 and 135 days. Recent

data pointed an incubation period of 47 days at 17–19 jC (Iglesias et al., in press).

3.4. Paralarvae and subadult phases

The family Octopodidae contains the largest number of known octopus species. In

some species, hatchlings are large and immediately benthic like the adults and thus are

referred to as juveniles (Villanueva, 1995). However, this is not the case in O. vulgaris.

This species has a planktonic posthatching stage termed paralarvae by Young and Harman

(1988). At hatching, this species has very small hatchlings (2 mm mantle length)

(Boletzky, 1987).

The biological characteristics of the early life stages of O. vulgaris were reviewed by

Nixon and Mangold (1998). Several experiments on the complete control of octopus

paralarvae have appeared in the literature since the nineteen sixties, when the classic article

by Japanese researchers (Itami et al., 1963) was published. Working on the northwestern

Pacific O. vulgaris, the authors succeeded in the rearing of hatchlings until settlement,

with a survival rate of 8% at day 45 and 5% at day 60 at mean water temperature of 24.7

jC. Several years after, Imamura (1990) reported new advances and high survival rates to

settlement, on O. vulgaris of the same geographical area.

Villanueva (1995) successfully reared Mediterranean O. vulgaris from hatchling to

settlement, feeding the planktonic paralarvae with zoeae of two crustacean species,

Liocarcinus depurator and Pagurus prideaux. The survival rate observed until day 40

was 32.1% at mean water temperature 21.2 jC. A supply of Carcinus maenas ovaries was

given from day 42, when some presettlement reflexes were already noted in paralarvae

behaviour. The author described octopuses in the presettlement stage as those individuals

that predominantly were planktonic but that intermittently rested on the bottom with arms

adhering to the wall or bottom of the tank and (or) that crawled by their arms for short

distances along the wall or bottom of the tank. Survival rate at day 60 was 0.8%.

Villanueva et al. (2002) studied growth and proteolytic activity of paralarvae fed with

Artemia nauplii (supplemented with vitamin complexes) and millicapsules. Starting with a

rearing density of 32 paralarvae l� 1 and a food ration of 4 nauplii ml� 1 day� 1, a survival


of 38% at day 20 and a doubling time of 14 days were obtained. After this early period, a

larger prey or suitable microdiet is required.

Carrasco et al. (2003) cited a survival rate of the paralarvae between 89.6% and 93.5%

at day 20 in two experiments conducted in 2002, at mean water temperature 21.2 jC.Maia

squinado zoeae and Artemia were used as living prey for the planktonic individuals. The

settlement of paralarvae was noted on day 52 and at day 60 all individuals were benthic,

with a survival rate of 3.4%.

Paralarvae is the limiting step in the culture of this species, which means that, currently,

aquaculture is commercially confined to the growth of subadults obtained from fisheries.

Growth from egg to subadult is only possible at laboratory and pilot scales, according to

the available scientific publications.

The main factors to be tested in future experiments, to increase survival of paralarvae

are prey availability (Villanueva, 1994, 1995) and temperature. Temperature is believed to

have a strong influence on settlement which occurs when paralarvae reach a critical size

(>7.5 mm of mantle length, irrespective of age) (Forsythe, 1993).

In order to clarify the nutritional requirements more precisely, the fatty acid compo-

sition of the paralarvae (Navarro and Villanueva, 2000) and ovaries, late eggs and wild

subadults (Navarro and Villanueva, 2003) were analysed. These authors found a close

relationship between the fatty acid profile of the dietary components and the resulting fatty

acid profile of the reared individuals. Poor growth and high mortalities seem to be

associated with a nutritional imbalance in the fatty acid profile, namely the docoxahex-

aenoic acid/eicosapentaenoic acid (DHA/EPA) ratio in artificial feeding. These authors

also concluded that co-feeding techniques based on the use of polar lipid and PUFA

enriched Artemia, together with palatable pellets, seemed to be a possible way to improve

paralarvae and subadult cephalopod culture beyond the experimental scale.

It is important to emphasise that a high proportion of the published studies on octopus

growth is only available from Mediterranean congresses and seminars in Spain and Italy

(Table 1), which makes their use difficult. This fact was noted by cephalopod workers and

originated several complete compilations: Santos (1999a), focused on ‘‘grey literature’’

between 1996 and 1999, and Santos (1999b).

Lee (1994) stated that dissolved gases and nutrients might contribute significantly to

meeting the metabolic and nutritional requirements of cephalopods, especially hatchlings.

These dissolved nutrients could be absorbed actively across the epidermis and then either

be used immediately for metabolism in the mantle tissue or enter the semi-closed

circulatory system for distribution.

3.5. Ongrowing

In nature, octopuses attack prey when they perceive movement. Perception is generally

monocular and accidental (Boucaud-Camou and Boucher-Rodoni, 1983). Octopuses

prefer to be fed slowly; this characteristic must be respected when planning feeding in

tanks. While some authors reported O. vulgaris to be more active during the night, being

considered dim-light feeders, others found this species more active when darkness

approaches (Boucaud-Camou and Boucher-Rodoni, 1983). These authors also point out

that this species seems to be opportunistic, prepared to feed at any time.


Octopus species often switch their food preference from small crustaceans to larger

ones during growth (Mangold, 1983). The quantity of food eaten is regulated in all

cephalopods that have been studied: they all reject any excess food. It seems to be

impossible, by offering food, to overfeed a cephalopod experimentally. Although they

prefer live food, they can be adapted to accept dead food like pieces of crabs, fish or

molluscs (Boucaud-Camou and Boucher-Rodoni, 1983).

An advantage of this species is its easy adaptation to captivity after the benthic stage,

which includes high acceptance of natural foods. This is important not only because there

is still no satisfactory artificial diet for cephalopods, but also because the potential for the

production of a more natural food exists. This could help to distinguish the farmed octopus

from other farmed species, which could increase the image of this new product for

consumers (for example, leading to the creation of ‘‘biologically produced octopus’’ or the

like). However, it should be noted, from the nutritional point of view, that the aquacultural

potential of this species will involve a change from natural to commercial dehydrated

foods. Lee et al. (1991) performed growth trials with pelleted diets developed for

cephalopods and analysed their palatability and acceptance on octopus and cuttlefish.

Authors observed that the texture of the dried pellets (10% moisture content) might be a

major factor affecting ingestion on Octopus bimaculoides. For raw, live and pureed diets

(40% moisture content), texture did not appear to be as important as for dried pellets,

although the mean-latency-to-grab was lower in live and raw diets (both composed by

shrimp and chicken). Thus, moisture content may be the most important property affecting

ingestion.

The nutrition of cephalopods was reviewed by Lee (1994). Aspects like the biochemical

composition of cephalopods, their feeding behaviour, digestibility and assimilation of

nutrients, as well as the importance of proteins on their growth and as a source of energy

were focused. This author stated that the feeding behaviour (pursuit and capture) in

cephalopods in initiated primarily by visual stimuli, but ingestion is affected by both

chemical and textural qualities of the food. Continued ingestion depends on the properties

of the food (pre-ingestinal factors) as well as the nutritional quality of the diet (post-

ingestinal factors).

An important group of publications with results on O. vulgaris ongrowing in captivity

appeared in Spain (Iglesias et al., 1997, 1999, 2000a), Portugal (Sendao et al., 1998) and

Italy (Cagnetta, 1999; Cagnetta and Sublimi, 2000). Several types of food were tested,

including crabs (C. maenas, Polybius henslowi) (Iglesias et al., 1997, 2000a), sardines

(Sardina pilchardus) and bogues (Boops boops) (Garcıa Garcıa and Aguado, 2002), of

which crabs, especially when live, seem to be the most desirable in terms of growth

(Cagnetta and Sublimi, 2000). When crustaceans are given as food in tanks, a high volume

of discarded material is produced (external shells); this problem should be minimized by

appropriate tank design, automatic separation of rejected materials and regular cleaning

procedures.

Some other important conclusions taken from recent ongrowing studies are that initial

octopus sizes must be similar, initial density should not exceed 10 kg/m3 (Otero et al.,

1999), males and females must be cultured separately and artificial structures for hiding

must be present in the tanks, in numbers similar to the number of octopus in each tank.

There are no important problems of cannibalism or competition for food.

Table 1

Summary of the more important results from seminars, technical magazines and internal technical reports on octopus culture

Factor, phase Major observations or summary Language Reference Kind

General culture procedures System design, water and food needs for laboratory

maintenance of several species including O. vulgaris

Spanish Forsythe, 1987 Seminar proceedings

Adult behaviour in nature Feeding behaviour, mechanisms and diets of cephalopods English Nixon, 1988 Seminar proceedings

General biology and culture

potential

Biology, fishing and farming Portuguese Gonc�alves, 1993 Equivalent to MSc thesis

Culture potential Overall view of O. vulgaris as candidate for aquaculture English Iglesias et al., 1996 Report

Biology, biometry Biometry parameters used to distinguish two different

populations

English Cunha and Pereira, 1995 Seminar proceedings

General culture procedures Evaluation of parameters that make O. vulgaris a

promising candidate for aquaculture

English Cagnetta et al., 1998 Seminar proceedings

Reproduction and hatching Reproduction behaviour, hatching and paralarvae

development

Spanish Moxica et al., 1999 Seminar proceedings

(abstract of oral presentation)

Hatching 100% mortality from eggs to 40 days (before subadult

phase)

Spanish Carrasco and

Rodrıguez, 1999

Seminar proceedings


Hatching Effect of Artemia enrichment with lipid and

protein sources and density on paralarvae survival

Spanish Martın et al., 1999 Seminar proceedings


Paralarvae and ongrowing 0.5–1.0 growth rates and low mortality for subadult;

high mortalities for paralarval growth

Spanish Iglesias et al., 1997 Seminar proceedings

Paralarvae chemical

composition

Paralarvae amino acid profile, relationship with nutrients

and absorption of amino acids through skin

Spanish Villanueva et al., 2003 Seminar proceedings

Reproduction, paralarvae and

ongrowing (laboratory and

cages)

Parameters for the control of the reproduction phase and

paralarvae growth; ongrowing from 750 g until 2.5–3 kg

in 3–4 months, mortality 10–15%

English Iglesias et al., 1999 Seminar proceedings

Subadult Low-cost closed circuit maintenance in captivity for

laboratory studies

Spanish Andreu-Moliner and

Cachaza, 1984

Internal report

Transportation and ongrowing Effect of temperature in handling and farming; sensitivity

to high temperatures

Spanish Aguado et al., 1999 Seminar proceedings


Ongrowing Separated sex cultures give best results; males grow faster

than females; 3 kg (males) and 2.5 kg (females) are

recommended as maximum ongrowing weight in

separated sex cultures

English Sanchez et al., 1998 Internal report

Ongrowing Crab diet resulted in faster growth than sardine or mullet

diets

English Sendao et al., 1998 Seminar proceedings

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Ongrowing Octopus grow better if sufficient space and freedom for

movement is provided

English Cagnetta, 1999 Seminar proceedings

Ongrowing From several monodiets tested, crab showed best results English Cagnetta and Sublimi, 1999 Seminar proceedings

Ongrowing Influence of dissolved oxygen in oxygen consumption

and ventilatory frequency; 13% saturation (0.8 mg/l) of

oxygen is the lower limit

Spanish Garcıa Garcıa et al., 1999a Seminar proceedings


Ongrowing Influence of octopus weight and water temperature in

oxygen consumption

Spanish Garcıa Garcıa et al., 1999b Seminar proceedings


Ongrowing Density of 10 kg/m3 is recommended as maximum Spanish Otero et al., 1999 Seminar proceedings


Ongrowing High weight gain and low accumulated mortalities in

rectangular-shaped tanks; specific growth rate 1.3%

Spanish Rodrıguez and

Carrasco, 1999

Seminar proceedings


Ongrowing Post-prandial oxygen consumption Spanish Cerezo and

Garcıa Garcıa, 2003

Seminar proceedings

Ongrowing Post-prandial ammonia production Spanish Cerezo et al., 2003 Seminar proceedings

Ongrowing in cages Raft-suspended cages can be used, 1 m3 are recommended

as maximum; PVC tubes are better shelters than

pneumatics or plastic baskets

Spanish Rama-Villar et al., 1997 Seminar proceedings

Ongrowing in cages Analysis of a period of 2 years of several ongrowing

parameters in 35 cages

Spanish Luaces-Canosa and

Rey-Mendez, 1999

Seminar proceedings


Subadult and adult pathology Skin ulcers of several Cephalopod species, subsequent

pathologies and bacterial agents; minimizing of wall

contact is advised

English Hanlon et al., 1988 Seminar proceedings

Processing Octopus marinating process English Baldrati, 1989 Technical magazine

Preservation, quality evaluation Oscillatory pressurization at 400 MPa at 7 and 40 jC to

octopus muscle resulted in reduced microbial load,

TMA-N, TVB-N, proteolytic activity, softening and WHC

English Hurtado et al., 1998 Seminar proceedings

Preservation, quality evaluation Musky octopus (Eledone moschata); sensory, chemical

and microbiological analysis during storage

Italian Civera et al., 1999 Technical magazine

Quality evaluation Octopine is considered as better quality indicator than

TVB-N, K value and polyamines

Spanish Respaldiza et al., 1997 Technical magazine

Reference list Cephalopod internal reports, seminars, congresses and

other ‘‘grey’’ literature (1996–1999)

English Santos, 1999a Seminar working document

Reference list Cephalopod scientific references (1996–1999) English Santos, 1999b Seminar working document

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The description of the feeding behaviour and techniques regarding live crustaceans

(Grisley and Boyle, 1988), bivalve prey (McQuaid, 1994) and the special case of broody

females (Wodinsky, 1978) is interesting as general biology and behaviour understanding of

this species.

Benthic octopuses exhibit negative phototaxis and reclusive behaviour (Villanueva,

1995). Consequently, higher growth rates were obtained when these characteristics

were respected by installing artificial shelters in the growth tanks, also known as

‘‘homes’’, ‘‘sheltering homes’’ or ‘‘dens’’ (Mather, 1994). They also prefer dark and

opaque dens, with no light inside at all (Anderson et al., 1999). Mather (1994) also

stated that octopuses in nature do not just seek suitable homes, but also chose

unsuitable ones and modify them, especially by moving stones to reduce the aperture

to around 12 cm in diameter.

From the industrial point of view, this is the phase of the life cycle showing the highest

potential: The species shows no important signs of cannibalism or competition for food,

and it is possible to attain a commercial size of 2.5–3 kg (from 750-g specimens) in 3 or 4

months, with mortality not exceeding 10–15% (Iglesias et al., 2000a).

Industrial ongrowing of small octopus in floating cages was predicted in the late

nineties (Rey-Mendez, 1998; Iglesias et al., 2000a) and is now a reality in Galicia (Spain),

where one company is rearing O. vulgaris. Some experiments, performed by the

University of Santiago de Compostela group with this company, resulted in growth rates

of 0.3–0.8 kg/month and low mortality (5.7%) using low-value frozen feeds including

sardine (S. pilchardus), scad (Trachurus trachurus), blue whiting (Micromesistius pou-

tassou), bogue (B. boops), mackerel (Scomber scombrus) and mussels (Mytilus sp.)

(Rama-Villar et al., 1997).

Octopus culture is now a strong area of study in Spain; first published results on

ongrowing involved cylindrical or square shaped cages, with individual dens (on the walls

or in the centre) for 150 individuals (Luaces-Canosa and Rey-Mendez, 1999). The

ongrowing process lasts 4 months, which means three fattening cycles theoretically can

be conducted per year. General calculations indicate a company with 25 cages would be

able to produce around 11000 octopuses per year (Iglesias et al., 2000b).

Separation of sexes at the ongrowing phase is recommended, as non-fecundated

females continue to grow until commercial size; in separate-sex culture, males grow

faster than females. Recommended attainable weight in separate-sex cultures is 3 kg for

males and 2.5 kg for females, as beyond this point, an increasing rate of mortality reduces

the yield of the ongrowing process (Sanchez et al., 1998).

As it is easy to feed octopuses in captivity with low-value natural food like live, fresh or

frozen crustaceans and fish, it seems that the development of a pelleted feed was not of

main concern or line of research until now. Cephalopods can be adapted to pellet foods,

but the costs and labour should be evaluated with care. Specific artificial foods for O.

vulgaris are not yet commercially available, but they will probably follow the complete

control of the life cycle of this species. On the other hand, as pellet foods can be used for

oral administration of antibiotics and food supplements, there will be a need for the

production of commercial feeds for these species (Lee et al., 1991).

O. vulgaris has a very rapid digestive rate (12 h at 18–19 jC) compared with other

truly benthonic octopuses like Eledone cirrhosa, depending on temperature, sex and


sexual maturation (Boucaud-Camou and Boucher-Rodoni, 1976; Boucher-Rodoni and

Mangold, 1977). Explanations for this rapid digestive rate and for several nutritional

characteristics of these species were published by Lee (1994). High growth rates are

explained by a very efficient amino-acid metabolism; lipids are not, as in vertebrate

carnivores, the predominant long-term energy store. Details on digestion and factors that

could interfere with it can be found in Boucher-Rodoni and Mangold (1977).

In some growth studies, sex did not appear to have any influence on the growth rate, but

there was some influence on the feeding rate, which was higher in females (Garcıa Garcıa

and Aguado, 2002). Some authors observed that males reach higher body weights than

females (Mangold, 1983; Iglesias et al., 2000a) because females experience stronger

metabolic needs during sexual maturation. However, prior to maturation, females grow as

rapidly as males (Garcıa Garcıa and Aguado, 2002).

Protein synthesis and growth were studied by Houlihan et al. (1990), who concluded

that rapid growth rates in O. vulgaris are brought about by high rates of protein synthesis

and high efficiencies of retention of synthesized protein and, therefore, little protein

degradation.

Major conclusions from another study (Aguado and Garcıa Garcıa, 2002) were that

growth or food intake were not affected by sex, optimum temperature for growth was 17.5

jC, food intake was higher with crab diet, but food efficiency was better for animals fed on

fish, which was reached at 16.5 jC for both diets tested. When temperature was above 23

jC, weight losses and mortality occurred. Taking into account all data obtained, optimum

performance of O. vulgaris growth is between 16 and 21 jC; recirculation in closed

systems with temperature control is probably a choice to consider.

4. Final overview

The complete life cycle of O. vulgaris under culture conditions was attained for the first

time in the year 2001 by Iglesias et al. (2002). Using Artemia and spider crab (Maja

squinado) zoeas, the survival during the paralarvae rearing was 31.5% per day after

hatching. These authors give weights of 0.5–0.6 kg at the age of 6 months and 2 months

later average weights of 1.6 kg (Iglesias et al., in press).

Iglesias et al. (1999, 2000a) presented a review on common octopus culture. The

main conclusion was that in order to reduce paralarval mortality rates and thus, to

close the culture cycle for this species, it will be necessary to focus future research on

finding prey food with a suitable nutritional profile and size. Using Artemia nauplii in

the first week of life followed by Artemia metanauplii, the survival rate was 10% until

140 Ag dry weight, but as high as 100% at the end of the experiments (maximum

duration of 32 days). Since paralarvae are the main difficulty in the octopus cycle,

focus will probably continue on components like fatty and amino acids, as some

results until now are promising.

O. vulgaris, like almost all species from fisheries, is a carrier in nature of several types

of parasite (Pascual et al., 1996), but these are not frequently cited as a problem in the

farming of this species. References to other pathologies in captivity are rare (Forsythe et

al., 1987, 1990). External pathologies (Hanlon et al., 1988) and fatal penetrating skin


ulcers were described by Hanlon et al. (1984) for cephalopods reared in captivity for

laboratory use.

5. Octopus processing

5.1. General considerations

While octopus production systems have been tested extensively, processing and

marketing strategies have received little attention. This is also the case with other recently

farmed species like tilapia in saline waters (Suresh and Lin, 1992).

After death, cephalopods enter a state of high protein degradation by both endogenous

and bacterial enzymes. Such rapid protein degradation results in the release of high levels of

nitrogen from the muscle, promoting bacterial growth and leading to rapid decomposition.

Consequently, the shelf life of an octopus is extremely limited, typically 6–7 days after catch

even at a low storage temperature of 2.5 jC (Hurtado et al., 1999) or 8 days at 0 jC (Barbosa

and Vaz-Pires, 2003). Farmers and processors must consider this difference from other

species.

One biological peculiarity of cephalopodmeat is the high solubility of its fibrilar proteins,

causing loss in nutritive value by the leaching out of a considerable amount of protein when

in contact with water. Washing, bleaching, brining, thawing in water, chilling, etc., need

careful attention in the processing plants if nutritive quality and flavour are to be retained.

Cephalopod muscle, in general, gains in weight when in contact with cold water but loses

nutrients quickly, much more readily than finfish muscle.

Spanish researchers performed experiments on the extension of octopus shelf life in ice

and texture improvement using exposure to high pressure as pre-treatment (Hurtado et al.,

1998) and combinations of heat and high pressure (Hurtado et al., 2001a,b), but although

some quality-related chemical and microbiological parameters were positively affected by

this method, no softening effects on the muscle texture were observed.

The presence of chromatophores in the skin (pigment organs) also creates a series of

processing problems, mainly in handling, freezing, cold storage, thawing and drying

(Kreuzer, 1984). After death, the muscles attached to the chromatophores are no longer

controlled, the chromatophores remain expanded and the muscles relax slowly, causing

skin colour changes from dark to light within a few hours of death. This process seems to

be concluded with the onset of rigor mortis.

Within the official sensory schemes of the European Union (Council Regulation, 1996),

the table for cephalopods only applies to cuttlefish (Sepia officinalis and Rossia macro-

soma). However, recent efforts have tested sensory tables (Barbosa and Vaz-Pires, 2003),

microbial counts and physical instruments (Vaz-Pires and Barbosa, 2003), chemical

evaluations like agmatine (Yamanaka et al., 1987; Ohashi et al., 1991) and octopine

(Respaldiza et al., 1997), and also microbial counts of psychrophilic bacteria like

Photobacterium phosphoreum and Pseudoalteromonas (Paarup et al., 2002). All these

are methods recently recommended for quality evaluation. Some work has also been

published in Italy on the chemical and microbiological characterisation of cephalopods

during storage (Civera et al., 1999).

5.2. Processing yield and edible parts

Due to lack of bones, the average edible portion of the cephalopods is 80–85% of the

total body, very high when compared with crustaceans (40–45%), teleosts (40–75%) and

cartilaginous fish (25%) (Kreuzer, 1984). This emphasises the potential of the species of

this group, as the rejected tissues are very low in percentage.


6. Octopus marketing

It is not too risky to predict that octopus will be included in the list of farmed species in

a relatively short time. The authors believe that the next 5–10 years will represent a very

good opportunity to create an appropriate market position for this species, as farmed

octopus will co-exist with octopus caught in nature for a long time. Two different

approaches are possible: to introduce farmed octopus in the same competition level as

the octopus caught in nature (as was the case for most other farmed species), or to create a

different product, and consequently, a different market for this new product. The authors

are convinced this second option is much more likely to be successful, as a good set of

advantages can be used to educate consumers and increase their respect for farmed

octopus. These include the already cited dietary, yield, convenience and environmental

advantages, but also the natural food farmers now use to grow octopus in captivity, far

different from the artificial foods used to grow many other animals for human consump-

tion. The creation of special guarantees and labels to emphasise this characteristic would

be of great interest for all involved in octopus aquaculture.

Farmed octopus marketing will also depend on the development of new products and

processing methods, and recovering of traditional products that were abandoned or have

only local importance. These include octopus canning, common in countries like Portugal

and Spain and marinating, as described by Baldrati (1989).

7. Future

Scientific results obtained with octopus are not as common as for other cephalopods

such as squid and cuttlefish. A great part of the bibliography is presented in a form of

reports produced by the research organizations, mainly for internal use, or as posters or

short communications at scientific meetings, which are always more difficult to find and

use; part of the information is not available in English: Spanish, Italian, Japanese and

Portuguese are quite common languages in the cephalopod field. It is advisable for authors

to increase their range of target readers by publishing in English, in accepted international

scientific journals.

From the available bibliography in English and in Spanish, it is reasonable to suppose

that the life cycle of O. vulgaris is now understood, but paralarvae rearing is only possible

under laboratory conditions and mortality is still too high. Main questions for future

research are paralarvae nutrition and the correct combination of physical parameters like

temperature, salinity and other water quality factors.


Ongrowing of subadult wild individuals is the only industrialized phase of the life

cycle, both in tanks and floating cages, with promising technical and financial results.

Production in NW Spain was estimated at around 32 tonnes year� 1 in 1998 and 1999

(FAO, 2001, 2002). Easy adaptation to captivity and feeding based on low-value foods, as

well as a rapid growth and high commercial value, are the main reasons for being

optimistic about the future aquaculture of this species.

As a conclusion, it can be said that the future research required to move forward in the

topic of O. vulgaris aquaculture will be focused on the need for stardardisation of

paralarvae rearing methods, especially on live prey versus inert diets. Experiments on the

survival during the weaning process and studies directed to the development of dry diets

for subadult growing will also be of major importance.

Acknowledgements

The authors gratefully acknowledge the support from the EU program ‘‘Iniciativa

Comunitaria-Pequenas e Medias Empresas’’ and Agencia de Inovac�ao (Eng. Joao Santos

Silva), Lisbon, Portugal, who financed the author Alexandra Barbosa (project ‘‘The Use of

the Crab P. henslowi as Food for Aquaculture’’). The authors also thank the invaluable

advices and detailed revision work kindly offered by Professor Graham A.E. Gall,

University of California, Davis, USA.

References

Aguado, F., Garcıa Garcıa, B., 2002. Growth and food intake models in Octopus vulgaris Cuvier (1797):

influence of body weight, temperature, sex and diet. Aquac. Int. 10 (5), 361–377.

Aguado, F., Gonzalez, R.F.M., Nicolas, E.M.A., Llorente, H.M.D., Munuera, F.F., Garcıa Garcıa, B., 1999.

Efecto de la temperatura sobre la supervivencia en el transporte, estabulacion y engorde de Octopus vulgaris

Cuvier (1797) en el Mediterraneo Occidental. Libro de resumenes, 7j Congreso Nacional de Acuicultura. Las

Palmas de Gran Canaria, 19–21 Mayo.

Anderson, R.C., Hughes, P.D., Mather, J.A., Steele, C.W., 1999. Determination of the diet of Octopus rubescens

Berry, 1953 (Cephalopoda: Octopodidae), through examination of its beer bottle dens in puget sound.

Malacologia 41 (2), 455–460.

Andreu-Moliner, E.S., Cachaza, A.N., 1984. Tecnica para el mantenimiento de Octopus vulgaris en acuarios

marinos con circulacion cerrada de agua. Inf. Tec. Inst. Esp. Oceanogr. 28 (11 pp.).

Aragao, C., Conceic�ao, L., Martins, D., Rønnestad, I., Gomes, E., Dinis, M.T., 2004. A balanced dietary amino

acid profile improves amino acid retention in post-larval Senegalese sole (Solea senegalensis). Aquaculture

233, 293–304.

Baldrati, G., 1989. Handling, marketing and processing of cephalopods in Italy. Ind. Conserve 64, 353–355.

Barbosa, A., Vaz-Pires, P., 2003. Quality index method (QIM): development of a sensorial scheme for common

octopus (Octopus vulgaris). Food Control 15 (3), 161–168.

Boletzky, S.v., 1987. Embryonic phase. In: Boyle, P.R. (Ed.), Cephalopod Life Cycles. Comparative Reviews,

vol. II. Academic Press, London, pp. 23–25.

Boletzky, S.v., 1989. Recent studies on spawning, embryonic development and hatching in the Cephalopoda.

Adv. Mar. Biol. 25, 85–115.

Boletzky, S.v., Hanlon, R.T., 1983. A review of the laboratory maintenance, rearing and culture of cephalopod

molluscs. Mem. Natl. Mus. Vic. 44, 147–187.


Boucaud-Camou, E., 1989. L’aquaculture des cephalopodes: evaluation et perspectives. Haliotis 19, 201–214.

Boucaud-Camou, E., Boucher-Rodoni, R., 1976. Digestive absorption in Octopus vulgaris (Cephalopoda:

Mollusca). J. Zool. (Lond.) 179, 261–271.

Boucaud-Camou, E., Boucher-Rodoni, R., 1983. Feeding and digestion in cephalopods. In: Wilbur, A.S.M. (Ed.),

The Mollusca. Physiology, Part 2, vol. 5. Academic Press, London. 189 pp.

Boucher-Rodoni, R., Mangold, K., 1977. Experimental study of digestion in Octopus vulgaris (Cephalopoda:

Octopoda). J. Zool. (Lond.) 183, 505–515.

Boyle, P.R., 1980. Home occupancy by male Octopus vulgaris in a large seawater tank. Anim. Behav. 28,

1123–1126.

Caddy, J.F., Rodhouse, P.G., 1998. Cephalopod and groundfish landings: evidence for ecological change in

global fisheries? Rev. Fish Biol. Fish. 8, 431–444.

Cagnetta, P., 1999. The effect of 3 different rearing strategies on the productive responses of the common octopus

(Octopus vulgaris C). Seminar on the Mediterranean Marine Aquaculture Finfish Species Diversification.

Zaragoza, Spain, 24–27 May.

Cagnetta, P., Sublimi, A., 1999. Productive performance of the common octopus (Octopus vulgaris C.) when fed

on a monodiet. Seminar on the Mediterranean Marine Aquaculture Finfish Species Diversification. Zaragoza,

Spain, 24–27 May.

Cagnetta, P., Sublimi, A., 2000. Productive performance of the common octopus (Octopus vulgaris C.) when fed

on a monodiet. Recent Advances in Mediterranean Aquaculture Finfish Species Diversification. Cahiers

Options Mediterraneennes, vol. 47, pp. 331–336.

Cagnetta, P., Zezza, L., Perniola, R., 1998. Preliminary trials on octopus (Octopus vulgaris C.) rearing under

controlled conditions. Atti del 33j International Symposium on new species for Mediterranean aquaculture.

Alghero, Italy, 23–24 Aprile.

Carrasco, J.F., Rodrıguez, C., 1999. Cultivo larvario del pulpo (Octopus vulgaris, Cuvier). Evolucion de la

supervivencia. Libro de resumenes, 7j Congreso Nacional de Acuicultura. Las Palmas de Gran Canaria,

19–21 Mayo 1999.

Carrasco, J.F., Rodrıguez, C., Rodrıguez, M., 2003. Cultivo intensivo de paralarvas de pulpo (Octopus vulgaris,

Cuvier 1797) utilizando como base de la alimentacion zoeas vivas de crustaceos. IX Congreso Nacional de

Acuicultura, Cadiz, 12–16 Mayo 2003.

Cerezo, J., Garcıa Garcıa, B., 2003. Descripcion del patron de consumo de oxıgeno postprandial en el pulpo de

roca (Octopus vulgaris Cuvier, 1797). IX Congreso Nacional de Acuicultura, Cadiz, 12–16 Mayo 2003.

Cerezo, J., Garcıa Garcıa, B., 2004. Influence of body weight and temperature on post-prandial oxygen con-

sumption of common octopus (Octopus vulgaris). Aquaculture 233, 599–613.

Cerezo, J., Gomez, E., Garcıa Garcıa, B., 2003. Resultados preliminares sobre la produccion de amoniaco en el

pulpo de roca (Octopus vulgaris Cuvier, 1797) obtenidos mediante un electrodo de ion selectivo. IX Congreso

Nacional de Acuicultura, Cadiz, 12–16 Mayo 2003.

Civera, T., Grassi, M.A., Pattono, D., 1999. Caractteristiche chimiche e microbiologiche di molluschi cefalopodi

nel corso della conservazione. Ind. Aliment. 38, 933–937.

Council Regulation (EC), 1996. No 2406/96 of 26 November 1996 laying down common marketing standards for

certain fishery products. Off. J. L. 334, 1–15 (23/12/1996).

Cunha, M.M., Pereira, J.M.F., 1995. Octopus vulgaris Cuvier, 1797 from Sao Miguel (Azores)—a biometrical

approach. Proceedings of the Second International Workshop of Malacology and Marine Ecology.

Acoreana (Supplement 1995). Sociedade de Estudos Acoreanos ‘‘Afonso Chaves’’, Azores, Portugal,

pp. 297–311.

de Groot, S.J., 1995. Edible species. In: Ruiter, A. (Ed.), Fish and Fishery Products: Composition, Nutritive

Properties and Stability. CAB International, UK, pp. 31–76.

Dinis, M.T., Ribeiro, L., Soares, F., Sarasquete, C., 1999. Cultivation potential of sole Solea senegalensis in

Portugal and Spain. Aquaculture 176, 27–38.

FAO, 2001. Fish and Fisheries Products. Food Outlook 1, February, vol. 11. FAO/GIEWS, Rome, Italy.

FAO, 2002. The State of the World Fisheries and Aquaculture 2002. FAO, Rome.

FAO, 2003a. Cephalopods commodity update. http://www.globefish.org/publications/commodityupdate/200311/

200311.htm.

FAO, 2003b. Species identification sheets. http://www.fao.org/figis/servlet/FiRefServlet?ds=species&fid=3571.

http:\\www.globefish.org\publications\commodityupdate\200311\200311.htm


Forsythe, J.W., 1987. Principales consideraciones en el cultivo de pulpos en laboratorio. In: Solis-Ramirez, M.J.

(Ed.), Memoria del Simposio sobre Investigaciones de Pulpos y Calamares, Yacalpeten, Yacatan, Mexico, 10

al 11 de Nov. Instituto Nacional de la Pesca, Yucalpeten, Yucatan, Mexico, pp. 25–46.

Forsythe, J.W., 1993. A working hypothesis of how seasonal temperature change may impact the field growth of

young cephalopods. In: Okutani, T., O’Dor, R.K., Kubodera, T. (Eds.), Recent Advances in Fisheries Biology.

Tokay Univ. Press, Tokyo, pp. 133–143.

Forsythe, J.W., Hanlon, R.T., Lee, P.G., 1987. A synopsis of cephalopod pathology in captivity. Proceedings of

the 18th Annual IAAAM Conference 1987 (Monterey, California) 1 (4), 130–135.

Forsythe, J.W., Hanlon, R.T., Lee, P.G., 1990. A formulary for treating cephalopod mollusc diseases. In: Perkins

F., Cheng, T. (Eds.), Pathology in Marine Science. Academic Press, San Diego, pp. 51–63.

Garcıa Garcıa, B.G., Aguado, F., 2002. Influence of diet on ongrowing and nutrient utilization in the common

octopus (Octopus vulgaris). Aquaculture 211, 171–182.

Garcıa Garcıa, B., Aguado, F., Llorente, H.M.D., Gonzalez, R.F.M., Nicolas, E.M.A., 1999a. Efecto de la

concentracion de oxıgeno disuelto sobre el consumo de oxıgeno y la frecuencia ventilatoria, y nivel letal

en el pulpo comun (Octopus vulgaris Cuvier, 1797). Libro de resumenes, 7j Congreso Nacional de Acui-

cultura. Las Palmas de Gran Canaria, 19–21 Mayo.

Garcıa Garcıa, B., Aguado, F., Nicolas, E.M.A., Llorente, H.M.D., Gonzalez, R.F.M., 1999b. Influencia del

peso y la temperatura sobre el consumo de oxıgeno de rutina en el pulpo comun (Octopus vulgaris

Cuvier, 1797). Libro de resumenes, 7j Congreso Nacional de Acuicultura. Las Palmas de Gran Canaria,

19–21 Mayo.

Gonc�alves, J.M.A., 1993. Octopus vulgaris Cuvier 1797 (polvo-comum): sinopse da biologia e explorac�ao.Provas de Aptidao Pedagogica e Capacidade Cientıfica. University of Ac�ores, Horta, Portugal.

Grisley, M.S., Boyle, P.R., 1988. Recognition of food in Octopus digestive tract. J. Exp. Mar. Biol. Ecol. 118,

7–32.

Guerra, A., Manrıquez, M., 1980. Parametros biometricos de Octopus vulgaris. Investig. Pesq. 44 (1), 177–198.

Hanlon, R.T., Forsythe, J.W., Cooper, K.M., Folse, D.S., Kelly, M.T., 1984. Fatal penetrating skin ulcers in

laboratory-reared octopuses. J. Invertebr. Pathol. 44, 67–83.

Hanlon, R., Forsythe, J., Lee, P., 1988. External pathologies of cephalopods in captivity. Proceedings 3rd

International Colloq. Pathology in Marine Aquaculture. Gloucester Point, Virginia, pp. 2–8.

Houlihan, D.F., McMillan, D.N., Agnisola, C., Genoino, I.T., Foti, L., 1990. Protein synthesis and growth in

Octopus vulgaris. Mar. Biol. 106, 251–259.

Hurtado, J.L., Montero, P.E., Borderıas, A.J., 1998. Chilled storage of pressurized octopus (Octopus vulgaris).

28th Western European Fish Technologists Association Conference, October 4–7, Tromso, Norway.

Hurtado, J.L., Borderıas, J., Montero, P., An, H., 1999. Characterization of proteolytic activity in octopus

(Octopus vulgaris) arm muscle. J. Food Biochem. 23, 469–483.

Hurtado, J.L., Montero, P., Borderıas, J., 2001a. Behavior of octopus muscle (Octopus vulgaris) under a process

of pressure– time– temperature combinations. Food Sci. Technol. Int. 7, 259–267.

Hurtado, J.L., Montero, P., Borderıas, J., Solas, M.T., 2001b. High-pressure/temperature treatment effect on the

characteristic of octopus (Octopus vulgaris) arm muscle. Eur. Food Res. Technol. 213, 22–29 (doi: 10.1007/

s002170100321).

Iglesias, J., Sanchez, F.J., Otero, J.J., 1996. The octopus (Octopus vulgaris Cuvier): a candidate for aquaculture?

International Council for the Exploration of the Sea (ICES), C.M. 1996/F, vol. 10. ICES, Copenhagen,

Denmark (5 pp).

Iglesias, J., Sanchez, F.J., Otero, J.J., 1997. Primeras experiencias sobre el cultivo integral del pulpo (Octopus

vulgaris Cuvier) en el Instituto Espanol de Oceanografıa. Actas del VI Congreso Nacional de Acuicultura.

Cartagena, Spain, 9, 10 and 11 July.

Iglesias, J., Sanchez, F.J., Otero, J.J., Moxica, C., 1999. Culture of octopus (Octopus vulgaris, Cuvier): present

knowledge, problems and perspectives. Seminar on the Mediterranean Marine Aquaculture Finfish Species

Diversification. Zaragoza, Spain, 24–27 May.

Iglesias, J., Sanchez, F.J., Otero, J.J., Moxica, C., 2000a. Culture of octopus (Octopus vulgaris, Cuvier): present

knowledge, problems and perspectives. Recent advances in Mediterranean Aquaculture Finfish Species

Diversification. Cahiers Options Mediterraneennes, vol. 47, pp. 313–322.

Iglesias, J., Sanchez, F.J., Otero, J.J., Moxica, C., 2000b. Ongrowing, reproduction and larvae rearing of octopus


(Octopus vulgaris Cuvier), a new candidate for aquaculture in Galicia (NW Spain). Workshop on New

Species for Aquaculture, Faro, Portugal, November.

Iglesias, J., Otero, J.J., Moxica, C., Fuentes, L., Sanches, F.J., 2002. Paralarvae culture of octopus (Octopus

vulgaris) using artemia and zoeas, and first data on juvenile growth up to eight months of age. Seafarming

Today and Tomorrow-Extended Abstracts and Short Communications. Presented at the Aquaculture

Europe 2002, Trieste, Italy, October 16–19. EAS Special Publication, vol. 32. European Aquaculture

Society, EAS, Oostende, Belgium, pp. 268–269.

Iglesias, J., Otero, J.J., Moxica, C., Fuentes, L., Sanches, F.J., in press. The complete life cycle of octopus

(Octopus vulgaris Cuvier) under culture conditions: paralarvae rearing using Artemia and zoeae, and first data

on juvenile growth up to eight months of age. Aquac. Int.

Imamura, S., 1990. Larval rearing of octopus (Octopus vulgaris Cuvier). The progress of technological devel-

opment and some problems remained. Collect. Breed. 52, 339–343 (Japanese text, English abstract).

Itami, K., Izawa, Y., Maeda, S., Nakai, K., 1963. Notes on the laboratory culture of the octopus larvae. Bull. Jpn.

Soc. Sci. Fish., vol. 29, pp. 514–520. Japanese text, English abstract.

Kreuzer, R., 1984. Cephalopods: handling, processing and products. FAO Fish. Tech. Pap. 254 (108 pp.).

Lee, P.G., 1994. Nutrition of cephalopods: fuelling the system. Mar. Freshw. Behav. Physiol. 25, 35–51.

Lee, P.G., Forsythe, J.W., DiMarco, F.P., DeRusha, R.H., Hanlon, R.T., 1991. Initial palatability and growth trials

on pelleted diets for cephalopods. Bull. Mar. Sci. 49 (1–2), 362–372.

Luaces-Canosa, M., Rey-Mendez, M., 1999. El engorde industrial de pulpo (Octopus vulgaris) en jaulas: analisis

de dos anos de cultivo en la Rıa Camarinas (Galicia). Libro de resumenes, 7j Congreso Nacional de

Acuicultura. Las Palmas de Gran Canaria, 19–21 May.

Mangold, K., 1983. Food, feeding and growth in cephalopods. Mem. Natl. Mus. Vic. 44, 81–93.

Mangold, K., 1998. The Octopodinae from the Eastern Atlantic Ocean and the Mediterranean Sea. In: Voss, N.A.,

Veccione, M., Toll, R.B., Sweeney, M.J. (Eds.), Systematics and Biogeography of Cephalopods. Smithsonian

Contributions to Zoology, vol. II. 586 pp.

Mangold, K., Boletzky, S.v., 1973. New data on reproductive biology and growth of Octopus vulgaris. Mar. Biol.

19, 7–12.

Martın, A.Y., Izquierdo, M., Fernandez-Palacios, H., Hernandez-Cruz, C.M., Castro, J., Hernandez-Garcıa, V.,

1999. Variacion del enriquecimiento de Artemia y la densidade de estabulacion en el cultivo larvario deOctopus

vulgaris. Libro de resumenes, 7j Congreso Nacional de Acuicultura. Las Palmas de Gran Canaria, 19–21May.

Mather, J.A., 1993. Octopuses as predators: implications for management. In: Okutani, T., O’Dor, R.K., Kubo-

dera, T. (Eds.), Recent Advances in Fisheries Biology. Tokai Univ. Press, Tokyo, pp. 275–282.

Mather, J.A., 1994. ‘Home’ choice and modification by juvenile Octopus vulgaris, (Mollusca: Cephalopoda):

specialized intelligence and tool use? J. Zool. (Lond.) 233, 359–368.

Mather, J.A., O’Dor, R.K., 1991. Foraging strategies and predation risk shape the natural history of juvenile

Octopus vulgaris. Bull. Mar. Sci. 49 (1/2), 256–269.

McQuaid, C.D., 1994. Feeding behaviour and selection of bivalve prey by Octopus vulgaris Cuvier. J. Exp. Mar.

Biol. Ecol. 177, 187–202.

Moxica, C., Otero, J.J., Iglesias, J., Sanchez, F.J., 1999. Comportamiento reproductor, puestas e desarrollo

embrionario del pulpo (Octopus vulgaris, Cuvier) en cautividad. Libro de resumenes, 7j Congreso Nacional

de Acuicultura. Las Palmas de Gran Canaria, 19–21 May.

Navarro, J.C., 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., Villanueva, R., 2003. The fatty acid composition of Octopus vulgaris paralarvae reared with live

and inert food: deviation from their natural fatty acid profile. Aquaculture 219, 613–631.

Nixon, M., 1966. Changes in body weight and intake of food by Octopus vulgaris. J. Zool. (Lond.) 150, 1–9.

Nixon, M., 1970. Some parameters of growth in Octopus vulgaris. J. Zool. (Lond.) 163, 277–284.

Nixon, M., 1988. The feeding mechanisms and diets of Cephalopods—living and fossil. In: Wiedmann, J.,

Kulmann, J. (Eds.), Cephalopods. Present and Past (Second International Cephalopod Symposim. E. Schwei-

zerbartsche Verlagsbuchhandlung (Nagelle u. Obermiller). Schweizerbart’sche Verlagsbuchhandlung, Stutt-

gart, Germany, pp. 641–652.

Nixon, M., Mangold, K., 1998. The early life of Sepia officinalis, and the contrast with that of Octopus vulgaris

(Cephalopoda). J. Zool. (Lond.) 245, 407–421.


Ohashi, E., Okamoto, M., Ozawa, A., Fujita, T., 1991. Characterization of common squid using several freshness

indicators. J. Food Sci. 56 (161–163), 174.

Otero, J.J., Moxica, C., Sanchez, F.J., Iglesias, J., 1999. Engorde de pulpo (Octopus vulgaris Cuvier) a diferentes

densidades de estabulacion. Libro de resumenes, 7j Congreso Nacional de Acuicultura. Las Palmas de Gran

Canaria, 19–21 May.

Paarup, T., Sanchez, J.A., Moral, A., Christensen, H., Bisgaard, M., Gram, L., 2002. Sensory, chemical and

bacteriological changes during storage of iced squid (Todaropsis eblanae). J. Appl. Microbiol. 92, 941–950.

Pascual, S., Gestal, C., Estevez, J.M., Rodrıguez, H., Soto, M., Abollo, E., Arias, C., 1996. Parasites in commer-

cially-exploited cephalopods (Mollusca Cephalopoda) in Spain: an update perspective. Aquaculture 142, 1–10.

Pedrosa-Menabrito, A., Regenstein, J.M., 1988. Shelf-life extension of fresh fish—a review: part I. Spoilage of

fish. J. Food Qual. 11, 117–127.

Peleteiro, J.B., Olmedo, M., Alvarez-Blasquez, B., 2000. Culture of Pagellus bogaraveo: present knowledge,

problems and perspectives. Recent advances in Mediterranean aquaculture finfish species diversification.

Cahiers Options Mediterraneennes, vol. 47, pp. 141–152.

Rama-Villar, A., Faya-Angueira, V., Moxica, C., Rey-Mendez, M., 1997. Engorde de pulpo (Octopus vulgaris)

en batea. Actas del VI Congreso Nacional de Acuicultura, Cartagena, Espana, 9–11 July.

Respaldiza, E.E., Delgado, M.C., Moral, A., 1997. Determinacion del nivel de octopina en cefalopodos comes-

tibles y su uso potencial como ındice de frescura. Aliment. Equipos Tecnol., 85–88 October.

Rey-Mendez, M., 1998. Cultivo de polbo en Galicia: dos resultados experimentais o cultivo industrial. In:

Fernandez Casal, J., Rey-Mendez, M., Cervino, A. (Eds.), Foro dos Recursos Marinos e da Acuicultura

das Rıas Galegas. O Grove, Spain, pp. 67–81.

Rodrıguez, C., Carrasco, J.F., 1999. Engorde del pulpo (Octopus vulgaris, Cuvier). Resultados de crecimiento,

supervivencia y reproduccion. Libro de resumenes, 7j Congreso Nacional de Acuicultura. Las Palmas de

Gran Canaria, 19–21 May.

Sanchez, F.J., Iglesias, J., Moxica, C., Otero, J.J., 1998. Growth of octopus (Octopus vulgaris) males and females

under culture conditions. International Council for the Exploration of the Sea (ICES), CM 1998/M: 47. ICES,

Copenhagen, Denmark (3 pp).

Santos, M.B., 1999a. Cephalopod grey literature (1996–1999). Working document. ICES Working Group of

Cephalopod Fisheries and Life History, Heraklion, 25–27 March.

Santos, M.B., 1999b. Cephalopod reference list (1996–1999). Working document. ICES Working Group of

Cephalopod Fisheries and Life History, Heraklion, 25–27 March.

Sendao, J.C., Carvalho, V., Borges, T.C., 1998. Rearing octopus (Octopus vulgaris, Cuvier) with three different

diets. Livro de resumos, VI Congresso Nacional de Aquacultura. Viana do Castelo, Portugal, 15–16 October.

Suresh, A.V., Lin, C.K., 1992. Tilapia culture in saline waters: a review. Aquaculture 106, 201–226.

Vaz-Pires, P., Barbosa, A., 2003. Sensory, microbiological, physical and nutritional properties of iced common

octopus (Octopus vulgaris). Lebensm.-Wiss. Technol. 37 (1), 105–114.

Villanueva, R., 1994. Decapod crab zoeae as food for rearing cephalopod paralarvae. Aquaculture 128, 143–152.

Villanueva, R., 1995. Experimental rearing and growth of planktonic Octopus vulgaris from hatching to settle-

ment. Can. J. Fish. Aquat. Sci. 52, 2639–2650.

Villanueva, R., Koueta, N., Riba, J., Boucaud-Camou, E., 2002. Growth and proteolytic activity of Octopus

vulgaris paralarvae with different food rations during first feeding, using Artemia nauplii and compound diets.

Aquaculture 205, 269–286.

Villanueva, R., Riba, J., Ruız Capillas, C., Gonzalez, A.V., Baeta, M., 2003. Composicion de aminoacidos en

Octopus vulgaris durante su desarrollo inicial. IX Congreso Nacional de Acuicultura, Cadiz, Espana, 12–16

Mayo 2003.

Wells, M.J., ODor, R.K., Mangold, K., Wells, J., 1983. Diurnal changes in activity and metabolic rate in Octopus

vulgaris. Mar. Behav. Physiol. 9, 275–287.

Wodinsky, J., 1978. Feeding behaviour of broody female Octopus vulgaris. Anim. Behav. 26, 803–813.

Yamanaka, H., Shiomi, K., Kikuchi, T., 1987. Agmatine as a potential index of freshness of common squid

(Todarodes pacificus). J. Food Sci. 52, 936–938.

Young, R.E., Harman, R.F., 1988. ‘‘Larva’’, ‘‘paralarva’’ and ‘‘subadult’’ in cephalopod terminology. Malaco-

logia 29 (1), 201–207.

Aquaculture Potential of the Common Octopus

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