Living Resources Committee ICES CM 2002/G:04 Ref. ACFM, ACE

REPORT OF THE

Working Group on Cephalopod Fisheries and Life History

Rome, Italy 10–12 April 2002

This report is not to be quoted without prior consultation with the General Secretary. The document is a report of an expert group under the auspices of the International Council for the Exploration of the Sea and does not necessarily represent the views of the Council.

International Council for the Exploration of the Sea Conseil International pour l’Exploration de la Mer

Palægade 2–4 DK–1261 Copenhagen K Denmark TABLE OF CONTENTS Section Page

1 INTRODUCTION...... 1 1.1 Terms of Reference...... 1 1.2 Attendance ...... 1 1.3 Opening of the Meeting and Arrangements for the Preparation of the Report ...... 2 2 LANDINGS AND EFFORT STATISTICS AND SURVEY DATA (TOR A)...... 3 2.1 Compilation of Landing Statistics...... 3 2.2 General Trends...... 3 2.3 National Trends...... 4 2.4 Comparison of ICES and FAO data...... 8 2.5 Conclusions...... 9 3 STOCK IDENTIFICATION AND POPULATION SIZE ESTIMATION (TOR B)...... 9 3.1 Introduction...... 9 3.2 Estimates of population trends based on indices...... 10 3.3 Total stock size assessments ...... 12 3.4 General discussion ...... 13 4 GEAR SELECTIVITY (TOR C) ...... 13 5 POSSIBLE PRECAUTIONARY APPROACHES TO MANAGEMENT (TOR D)...... 14 5.1 General considerations...... 14 5.2 Management in European cephalopod fisheries...... 14 6 ENVIRONMENTAL FACTORS AFFECTING RECRUITMENT AND DISTRIBUTION PATTERNS (TOR E)15 7 CEPHALOPOD LITERATURE RELEVANT TO FISHERIES (TOR F) ...... 16 8 RESEARCH PRIORITIES ...... 16 8.1 Funding for data collection ...... 16 8.2 Research priorities ...... 17 9 THE FUTURE PROGRAMME OF WGCEPH AND RECOMMENDATIONS ...... 18 9.1 Terms of reference ...... 18 9.2 Next WGCEPH meeting ...... 18 10 OTHER BUSINESS AND CLOSING OF THE MEETING ...... 18 10.1 Exchanges with scientists studying cephalopods in the Mediterranean...... 18 10.2 Information about scientific meetings...... 19 11 ACKNOWLEDGEMENTS ...... 19 12 REFERENCES...... 19 ANNEX 1 ...... 30 ANNEX 2 ...... 32 ANNEX 3 ...... 42 ANNEX 4(I) ...... 52 ANNEX 4(II)...... 61 WORKING DOCUMENT N° 1...... 64 WORKING DOCUMENT N° 2...... 69 WORKING DOCUMENT N° 3...... 73 WORKING DOCUMENT N° 4...... 83 WORKING DOCUMENT N° 5...... 91

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1 INTRODUCTION

1.1 Terms of Reference

As indicated in Part 2 of the ICES Annual Report for 2000, ICES Council Resolution C.Res. 2001/2G02 stated that the Working Group on Cephalopod Fisheries and Life History [WGCEPH] (Chair: Dr .J.P. Robin, France) would meet in Rome, Italy from 10-12 April 2002 to: a) update currently available landing statistics and information on fishing effort and discards; explore existing resource survey databases for information about sampled cephalopods in the ICES area; b) compile methods and results available for stock identification and estimation of population size of fished cephalopods; c) compile available data on gear selectivity for cephalopods; d) identify possible precautionary approaches to the management of cephalopod resources; e) compile available data on relationships between abundance and environmental conditions, factors affecting recruitment, migration and distribution patterns of juveniles and adults, and trophic interactions; f) update the bibliographic database of cephalopod literature relevant to fisheries, including grey literature;

WGCEPH will report by 30 April 2002 for the attention of the Living Resources Committee, and ACFM and ACE.

1.2 Attendance

The WGCEPH meeting at the ICRAM headquarters, Rome, 10-12 April 2002, was attended by 8 of the currently appointed WGCEPH members. These participants represented 5 ICES member states (France, Germany, Portugal, Spain, UK).

Participants: 1) WGCEPH members

Name Address Telephone and Fax E-mail

Dr Angel F. ECOBIOMAR, Instituto de Investigacions Tel +34-986-231930 [email protected] González Mariñas, Eduardo Cabello 6, E-36208 Vigo, (Ext. 181) Spain Fax: 34 986 292762 Dr Noussithé Université de Caen, 14032 Caen Cedex, Tel +33 231 538016 [email protected] Koueta France Fax +33 231 538009 3Dr. João IPIMAR, Avenida de Brasília, S/N, 1449-006 Tel +351-1-3027000 [email protected] Pereira Lisbon, Portugal Fax +351-1-3015948 Mr. João IPIMAR, Avenida de Brasília, S/N, 1449-006 Tel +351-1-3027000 [email protected] Pereira Lisbon, Portugal Fax +351-1-3015948 Dr Uwe Institut für Meereskunde, Universität Kiel, Tel +49-431-5973908 [email protected] Piatkowski Düsternbrooker Weg 20, D-24105 Kiel, Fax +49-431-565876 kiel.de Germany Dr. Inaki AZTI, Txatxarramendi Ugartea z/g, 48369 Tel +34 946 870700 [email protected] Quincoces Sukarrieta (Bizkaia), Spain Fax +34 946 870006 Dr Jean-Paul Université de Caen, I.B.B.A., Biologie et Tel +33 231 538017 [email protected] Robin (Chair) Biotechnologies Marines, F-14032 Caen Fax +33 231 538009 Cedex, France Dr Ignacio Instituto Español de Oceanografía Tel +34 956261333 [email protected] Sobrino Unidad de Cádiz, Apdo. 2609, 11006 CADIZ, Fax +34 956263556 Spain Dr Robert Renewable Resources Assessment Group, Tel +44 20 7594 7310 [email protected] Wakeford Imperial College, Royal School of Mines, Fax +44 20 7589 5319 Prince Consort Road, LONDON, SW7 2BP, UK

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The WGCEPH meeting was held in Rome in order to provide an opportunity to contact FAO Fisheries Scientists and to improve contacts with Mediterranean Cephalopod research. Four observers (including local host) contributed to the meeting.

Participants: 2) observers

Name Address Telephone and Fax E-mail

Dr Patrizia Jereb ICRAM, Via di Casalotti 300 Tel +39 06 61570491 [email protected] (meeting host) 00166 - Roma, Italy fax +39 06 61561906

Pr. John F. Caddy Senior Research Fellow, [email protected] Dept Environmental Science and Technology, Imperial College, University of London And Professor, CINVESTAV, Merida, Mexico Dr. Jorge Csirke Chief, Marine Resources Service Tel.: +39 0657056506 [email protected] Fishery Resources Division Fax: +39 0657053020 Food and Agriculture Organization of the United Nations (FAO) Viale delle Terme di Caracalla 00100 Rome, Italy Dr. Luca Fishery Statistician (Capture Fisheries) Fax: +39-0657052476 [email protected] Garibaldi Fishery Information, Data and Statistics Unit g (FIDI) Food and Agriculture Organization of the United Nations (FAO)

A full list of members of WGCEPH (provided by ICES in February 25th 2002) is given in Annex 1. It was indicated to new members (and also to persons who were not wishing to continue to participate) that for any change on that list they had to contact their ICES national delegate (and not WGCEPH chairman).

1.3 Opening of the Meeting and Arrangements for the Preparation of the Report

The meeting was hosted by the Istituto Centrale per la Ricerca Scientifica e Tecnologica Applicata al Mare (ICRAM). The agenda of the meeting is given at the end of this report.

Prior to the meeting, responsibility for preparation, collation and presentation of material for the T.o.R. was delegated to the following members, João Pereira (Portugal), Uwe Piatkowski (Germany), Robert Wakeford (UK), and Begoña Santos (UK).

It was agreed that amended text, updated during and following the meeting, would be submitted electronically to the Chairman, who undertook to write and circulate a final draft to members and attendees.

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2 LANDINGS AND EFFORT STATISTICS AND SURVEY DATA (TOR A)

2.1 Compilation of Landing Statistics

The present report updates landing statistics from 1995 to 2000 and provides preliminary catch data of 2001 for cephalopod groups caught in the ICES area (Tables 2.1 to 2.6). The data largely originate from the ICES STATLANT database and from additional national and more precise information supplied by Working Group members. It should be noted that several ICES member countries/regions could yet not supply information for 2001 (i.e., Faroe Islands, Netherlands and partly Spain). In these cases the 2001 catch information was marked as “not available” (n.a.) in the tables. It is anticipated that these data will be available for the 2003 report. In general, we feel that all 2001 data should be considered as preliminary, and they are marked as such in the tables (“P”).

The data compiled in this report represent the most precise information on cephalopod landings within the ICES area that can be obtained to date. For all major fishery nations (i.e., France, Portugal, Spain, UK) we relied on the statistical information provided by the Working Group members. This information is – as in previous years – not necessarily identical to the data officially reported to the ICES ATATLANT database and stresses the inaccuracy with which cephalopod statistics are still handled.

Tables 2.1 to 2.4 give information on annual catch statistics (1995-2001) per cephalopod group in each ICES division or subarea, separately for each nation. The cephalopod groups listed in the tables comprise the following species:

• Table 2.1. Cuttlefish (Sepiidae). The majority of landings summarised in this table are catches of Sepia officinalis, the common cuttlefish, plus small amounts of S. elegans and S. orbignyana. WGCEPH considers that no bobtail squids (Sepiolidae) occur in the reported catches.

• Table 2.2. Common squid (including the long-finned squids Loligo forbesi, L. vulgaris, Alloteuthis subulata and A. media). The majority of common squid landings are L. forbesi and L.vulgaris.

• Table 2.3. Short-finned squid (Illex coindetii and Todaropsis eblanae), European Flying squid (Todarodes sagittatus), and Neon Flying squid (Ommastrephes bartrami).

• Table 2.4. Octopods (including Eledone cirrhosa, E. moschata and Octopus vulgaris).

A compilation separated into single species is still not possible as all countries report landings for cephalopod groups, mostly in the format as given in the tables.

Table 2.5 summarises total annual cephalopod landings in the whole ICES area for major cephalopod groups. Table 2.6 provides information of total annual cephalopod landings in the whole ICES area for major cephalopod groups separated for each fishing nation.

2.2 General Trends

Total reported annual cephalopod landings within the ICES region varied between 47531 t in 1995 and 56867 t in 2000 (see Table 2.5). Data for 2001 (39383 t) are very provisional and definitely too low, particularly because data of several fishing nations for that year are yet not available. It is striking, however, that the overall catch has increased constantly from year to year in the period of 1995 to 2000. This clearly emphasises the greater importance that cephalopods have gained as a fishery resource within the ICES area.

In terms of yields, cuttlefish are currently the most important cephalopod group taken in the ICES area. Their landings increased remarkably from 1995 to 2000 (19601 t and 24008 t, respectively), with the exception of 1997 when the total catch of this group dropped to about 16652 t (Table 2.5 and 2.6). As previously reported by WGCEPH this is mostly due to an increase of catches in the English Channel taken by France and the UK. France was by far the most important fishery nation with annual catches of always more than 60% of the overall annual cuttlefish catch (Table 2.6).

Total landings of common squid (Loliginidae) were more stable in recent years and in the range of ca. 10000 t from 1995 to 2000 (Tables 2.2; 2.5; and 2.6). The yield of short-finned squid (comprised of the species Illex coindetii, Todaropsis eblanae und Todarodes sagittatus) showed the most obvious fluctuations ranging from 1703 t in 1995 to 7719 t in 1999 (see Tables 2.3; 2.5; and 2.6). During the period reported here Spain was by far the major fishery nation of this group taking more than 80% of the total catch in most years.

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After a considerable decrease in 1998, octopod catches increased again since 1999 and will probably also be seen to have further increased in 2001, once more comprehensive catch data can be obtained (Tables 2.4; 2.5; 2.6). They were composed nearly exclusively of the two species Eledone cirrhosa, and Octopus vulgaris. It should be noted here that Portugal and Spain together took more than 97% of the catch in each reported year. This reflects the high economic value of this resource in the coastal fisheries of both nations.

In terms of total cephalopod landings, the most active nations in 2000 were France (23161 t), followed by Spain (14693 t) and Portugal (11650 t) which all conduct a target fishery on cephalopods and which together took the major share within the ICES region (90%). Next to them the UK followed with a total of 5152 t (9%). The remaining 1% was mainly taken by Ireland, the Netherlands and Belgium. The latter two catch cuttlefish and common squid as a valuable by-catch. In the following paragraphs some information and trends for the major fishing nations are provided.

2.3 National Trends

France

Updated landings data were obtained for 1999-2001 from the official data-base (Centre Administratif des Affaires Maritimes, St. Malo) although 2001 is still a provisional underestimate.

Cuttlefish landings increased from 1997 to 2000, when they reached a level similar to that observed in 1990. It is worth noting that twelve years ago fish markets seemed to have difficulties with such an abundant cohort whereas in 2000 cuttlefish remained throughout the fishing season a desirable resource for the French Fishery.

Loliginid squid landings were higher in 1999 than in 2000 (5825 t. and 5180 t respectively). This is an average level when compared to 1989 and 1993 peaks (above 6000 t) and 1996 sag (below 4000 t). Inter-annual trends in French common squid landings look rather stable over the last decade, however variations from one cohort to the next can be buffered when this group of species (Loligo forbesi and Loligo vulgaris) is observed at the annual scale. Monthly landings (Figure 2.1) suggest that yields in the 1999 fishing season (June 1999 – May 2000) were as high as in the 1993 fishing season whereas the 2000 fishing season was almost as low as 1998 but was nevertheless twice higher than 1996.

Short finned squid and octopod landings by France are much smaller though both groups increased in 1999 and 2000.

Geographic origin of landings underlines that English Channel cuttlefish and Loliginid squid represent the major stocks exploited by France (and largest exploited populations for these species in Europe). The proportion of landings for this area (English Channel) in all French data seems rather constant in time (about 2/3 of France cephalopod landings).

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Cuttlef ish

3000 2500 2000 1500 1000 500 0

Loliginid squid

1 400 1 200 1 000 800 600 400 200 0

Short Finned Squid 250 200 150 100 50 0

Octopod 50 40 30

20 10 0

Figure 2.1 Monthly cephalopod landings (in tonnes) by France in the period 1989-2001.

Ireland

Cephalopod landings by the Irish fleet indicate that highest yields were observed in 1997 in common squid and 1998 in short finned squid (Table 2.6). Geographic differences (Tables 2.2 and 2.3) suggest that decrease in recent years is mainly observed in the Celtic Sea and SW of Ireland whereas catches in Porcupine Bank and the West of Ireland are stable or increasing.

Portugal

Landing statistics in Portugal are generally made available by March each year up to the end of the preceding year. These data are preliminary until approximately mid-year but are usually very close to the final figure. Currently, the provisional 2001 data are under analysis.

As a group, cephalopods have yielded average landings in the order of 11000 tonnes per year throughout the last decade (1992-2001), with fluctuations between c. 9000 (1994) and 13500 tonnes (1996). These are among the highest values within the historical series available from 1927 (Figure 2.2), and are part of the stable sub-series started in 1987,

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following the introduction of technical modification to the pattern of the fisheries, which greatly enhanced the catchability of the fleets targeting these organisms. Therefore a rough analysis of these data show an apparent sustainable yield within each of three sub-series corresponding to technical modifications affecting the catchability. Apparently cephalopod populations in Portugal have been able to provide increasing yields when subjected to greater fishing pressure, contrary to what has been observed with finfish stocks. Under closer scrutiny, however, each of the two latest sub-series shows a slight decreasing tendency from former to latter years. In the present sub-series, the four last years have all been in the lower 50% group of landings. Since this tendency is statistically non-significant, it may not warrant any specific measure, but we feel it is worth mentioning.

All Cephalopods

16000

14000

12000

10000 Tonnes) 8000

6000

Landings ( 4000

2000

0 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010 Year

Figure 2.2 Cephalopod landings (in tonnes) by Portugal in the period 1927-2001.

Dissecting the series into its four major statistical components, which by family correspond to the Ommastrephid squid, the Loliginid squid, Octopods and Cuttlefish, landings in Portugal are, as always have been, strongly dominated by the octopods, of which the most part is Octopus vulgaris.

The group of the octopods in general and Octopus vulgaris in particular, is in fact responsible for each and every alteration in the tendency apparent in the general statistics shown in Figure 2.2. Since 1992, the species accounted for 78% of cephalopod landings in the country with a minimum of 67% in 1998 and a maximum of 84% in 1996. It is, in addition highly priced among the most abundant species landed in the country (finfish included), driving a strong interest in the fisheries sector. The fishing pressure has been high and there are reasons to suspect that the official statistics are a significant underestimation of the fishing yield. There is a popular belief among fishermen in the Algarve that the species has produced lower CPUE’s in present years but overall and country-wide, we do not have indications of this. There have however been alterations in the availability pattern which may be responsible for this belief, since by geographic region, the relative importance of the landings has changed from a more southern concentration to a gradual shift northwards presently resulting in a roughly equal distribution of landings around the coast (Figure 2.3). Since most octopus catches are made by the fleet operating locally, there seems to have been a real change in the distribution of the species.

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7000

6000

Algarve .) 5000

Tons 4000 (

3000 ndings

La 2000 Lisbon, Tagus Basin & Alentejo 1000

0 North & Center 1970 1975 1980 1985 1990 1995 2000 Year

Figure 2.3 Octopod landings (in tonnes) by statistical region in Portugal in the period 1973-2000.

The remaining three groups of species display stable trends in the past years, with the possible exception of the long- finned squid, with somewhat decreasing landings in the beginning of the 90s (Figure 2.4). The decrease of that group at that time, coincides with a documented disappearence of the species Loligo forbesi from Portuguese waters. It is perhaps worth examining a longer series for that group (Figure 2.5).

2000 Long-Finned Squid 1800 Short-Finned Squid Cuttlefish 1600

1400

1200

1000

800 Landings (Tonnes) 600

400

200

0 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 Year

Figure 2.4 Landings (in tonnes) in Portugal of Long- and Short-finned squid and of cuttlefish in the period 1990-2001.

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2500

2000 s) e n

n 1500 o gs (T n i

d 1000 Lan

500

0 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010 Year

Figure 2.5 Landings (in tonnes) in Portugal of Long-finned squid in the period 1927-2001.

Generally, Long-finned squid were landed in relatively low weights up to the end of the 1960’s, having had an increment since then, but most importantly two peaks, one in 1965 and another in 1991. After the first EU funded cephalopod research project in the early 1990’s, results suggest that there may be periodic inputs of Loligo forbesi from its usual distribution further north, possibly following periods of high abundance in its main distribution grounds. This may affect catches in Portugal, which later return to lower levels.

Spain

During the last seven years the cuttlefish Spanish landings belong to Subdivisions IX (71.3%) and VIII (23.9%). The landings were at their maximum level in years 98-99, and the lowest catches were recorded in 2001 (provisional data). For Common Squid, as described in cuttlefish, the bulk of the fisheries are placed in Subdivision IX (66.9%) and VIII (22.1%). Their landings had a peak in 97-98, with a decreasing trend in the last three years.

The Ommastrephid fishery takes mainly place in Subdivision IX and VIII as described in the previous species (47% and 35.8% respectively) and also in the Subdivision VII (15%). Highest landings were recorded in 1999 and the lowest ones in the last two years.

Octopods are the most important species (mainly Octopus vulgaris) from the total landings of cephalopods for the Spanish fleet. Subdivision IX is the main fishery area for the species (62.7%), with Subdivision VIII like a secondary area, accounting only a 32.1% of total catches. The trend in the landing series, show a rather stable pattern with an important decrease in the last year (provisional data)

United Kingdom

Data collected for 2001 do not mention the proportion of each nation's landings in UK data (Scotland, England and Wales and Northern Ireland, Isle of Man, Channel Islands). Although this can be known for some fishing grounds (off Scottish waters for instance) provisional 2001 figures were put somewhat arbitrarily in the continuation of previous data (Tables 2.1 to 2.4). Temporally, it seems that landings in recent years (2000 – 2001) were slightly lower than in 1995- 1999. In some areas this may be a result of fleet interactions in international fishing grounds: for instance English Channel cuttlefish landings by the UK were still high (2500 – 3000 T) although not as high as in 1995 – 1996 when interest for this resource started to increase in England. From a geographical point of view it is worth noting that UK landings are no more dominated by Northern squid stocks fished by Scotland. Landings from the English Channel and Celtic Sea represent almost an equivalent amount.

2.4 Comparison of ICES and FAO data

The participation of FAO scientists provided the opportunity to compare of cephalopod data integrated in ICES and FAO databases. A series of tables were prepared which are annexed to this document (annex 2).

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FAO statistics included in these tables are those available in the March 2002 version of FISHSTAT+ (downloadable at http://www.fao.org/fi/statist/FISOFT/FISHPLUS.asp). ICES data have been extracted from the December 2001 version of the "Integrated Atlantic captures", a data file assembled by EUROSTAT under the aegis of the Coordinating Working Party on Fishery Statistics (CWP). As a consequence of this exercise, some of the resulting discrepancies will be corrected in the next release of the FAO capture database.

Information was also provided on the procedure applied by FAO in case of inconsistent record declaration by one country (and the commitments of FAO to stick to official figures).

The working group noticed that discrepancies between the two data bases were generally low in the period 1987 – 1998 (a part from 1000 T of 1995 Irish landings not included in the FAO data set). In recent years highest differences were related to the fact that France statistics for 1999, were missing in the ICES data base (which did not yet include year 2000 landings). Minor cephalopod fishing countries like Lithuania also seem to be better taken into account in the FAO data base.

2.5 Conclusions

The fact that the overall catch has increased constantly from year to year in the period of 1995 to 2000 clearly emphasises the greater importance that cephalopods have gained as a fishery resource within the ICES area.

The compilation of landings statistics is a recurrent task for the working group which leads logically to update the series of annual tables prepared in the same format as in previous reports. At this stage, a series of remarks are proposed with the aim to make this work more useful:

+WGCEPH members have developed GIS applications which enable to display trends by species and ICES divisions (or groups of divisions) on a series of maps. An example of such outputs is given in the annex 3 which contains recent temporal trends and year 2000 yields per country.

+Although TOR a mentions "landings and effort" statistics, effort data is not compiled by WGCEPH. At the national scale information on effort trends can be used by members to analyse inter-annual trends (2.3). However, in spite of the fact that WGCEPH can contribute to identify fishing effort targeted at cephalopods it can hardly handle any European fishing effort likely to land cephalopods as a by-catch. Comprehensive records of fishing effort are not available in every country for every fleet but WGCEPH can underline any new situation where simultaneous catch and effort data enable to derive abundance indices (see TOR b in this report).

+Annual landings statistics represent a first step in the analysis of cephalopod fishery trends. However, no short living species like cephalopods can be assessed with annual data only and assessments of European cephalopod stocks (Pierce et al, 1996; Royer et al 2002) all require data on (at least) a monthly basis. Thus collection of fishery statistics on a finer time scale should be encouraged among WGCEPH members. An interesting view of the progress in this direction is provided with map n°6 (annex 3): year 2000 landings by country can be completed by the information that monthly statistics on a "monthly and per ICES rectangle" basis can be available now for the main countries (i.e. UK, France, Ireland) weekly statistics are collected by Portugal and some Spanish landings by selected fleets are also studied on a monthly basis.

3 STOCK IDENTIFICATION AND POPULATION SIZE ESTIMATION (TOR B).

3.1 Introduction

This section presents recent work on cephalopod stock assessments (either new assessments or new outputs). The update of assessments based on historical data depends both on the continuation of data collection and on the availability of fishery statistics.

The term of reference is as follows: Compile methods and results available for stock identification and estimation of population size of fished cephalopods.

The situation at the end of the EC-funded Study Project 99/063 (Data collection for Assessment of Cephalopod Fisheries) which involved participants from the UK, France, Spain Portugal and Greece was summarized by the project co-ordinator.

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Although the knowledge of stock exploitation was improved in every country two different stages were achieved as far as population trends are concerned. In some rather well known fisheries (Scottish waters, English Channel) estimates of the entire stock size have been obtained for several cohorts in which recruitment could be compared. In other areas the lack of comprehensive catch data (or problems related with geographic origin of landings) did not allow to derive estimates of total population size, nevertheless abundance indices based on catch rate were computed (these indices are assumed to be proportional to population size).

3.2 Estimates of population trends based on indices.

Abundance indices for Octopus vulgaris

Portugal

An estimate of weekly landing per unit effort (LPUE) has been calculated as an index of abundance for O. vulgaris between 1997 and 2000 (Figure 3.1). This is an octopus-directed artisanal fishery that operates throughout the year with a multitude of gear types but mostly dependent on two, octopus-specific creels and clay pots, the first represented in the vast majority of operations.

To obtain estimates of LPUE from the daily landings record, several procedures are followed in steps. Firstly the following assumptions are made: 1 - each vessel has a similar working pattern in a year; 2 - gears will remain the same within a day for each vessel; 3 - individual daily effort patterns will remain constant for the period of one civil year. Therefore several standardisations are made: time = 1 day; vessel = each; gear = same set per vessel per year.

From a database of daily landings per vessel, the second landing of any two consecutive landings per vessel is selected. Of those, the average daily landing of each vessel throughout one civil year is used as the representation of its fishing capacity. Variations in the series of each boat’s landings in relation to the average landing are recorded as positive and negative contributions of a magnitude equivalent to the difference from the respective mean, divided by the standard deviation of the mean of that vessel’s landings. All the contributions of each vessel (positive or negative) are added up for each period of a week and averaged taking into consideration the total number of contributions for the week, with a result that can either be positive or negative. This last figure is the “standardisation” index (SI). A new figure is arrived at which is the weekly sum of the total landings made from the second daily landings, selected as above (unit = kg), divided by the number of contributions made to that sum (unit = day), i.e. a number of kg/day (dL) is derived. Finally, the standardised LPUE is obtained by adding the product between the weekly standardisation index and the weekly addition of the selected daily landings (WdL) to that last number (LPUE=WdL+SI.WdL).

The results show a seasonal trend in the weekly abundance of octopus, which reaches a peak value during the winter period. Interestingly, the relative abundance of octopus has increased towards the end of 1999 and 2000. The latter may reflect a period of good recruitment to the population.

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1997 1998

80 10,000 80 50,000 Catch Catch 9,000 45,000 70 LPUE 70 LPUE 8,000 40,000 60 60 7,000 35,000 50 50 6,000 SV) 30,000

40 5,000 day/ 40 25,000 (kg/

4,000 E 20,000 30 30 LPUE (kg/day/SV) 3,000 LPU 15,000 Second daily landings (kg) 20 20 Second daily landings (kg) 2,000 10,000 10 10 1,000 5,000

0 0 0 0 0 5 10 15 20 25 30 35 40 45 50 0 5 10 15 20 25 30 35 40 45 50 Week Week

1999 2000

180 100,000 180 250,000 Catch Catch 160 90,000 160 LPUE LPUE

140 80,000 140 200,000 g) g)

70,000 k k ( ( ) 120 ) 120 V V ngs ngs /S 60,000 /S 150,000 100 100 andi andi l l

g/day 50,000 g/day y y l l k k

( 80 ( 80 40,000 100,000 UE UE LP LP 60 ond dai 60 ond dai 30,000 c c e e S S 40 20,000 40 50,000

20 10,000 20

0 0 0 0 0 5 10 15 20 25 30 35 40 45 50 0 5 10 15 20 25 30 35 40 45 50 Week Week

Figure 3.1 Weekly abundance indices (kg/day/SV) and total second daily landings (kg) and for the Portuguese artisanal fishery targeting Octopus vulgaris between 1997 and 2000. Note different scales on vertical axes. SV relates to a standardised vessel unit.

Spain

Sobrino et al (2002) analysed landings, effort and landing per unit effort (LPUE) patterns of common octopus (Octopus vulgaris) from a time-series of eighteen years. They show that both LPUE and landing may be used as suitable indices of abundance by analysing correlations between landings, LPUE and fishing effort.

They used the approach suggested by Pierce et al. (1994) for the UK Loligo forbesi fishery. According to these authors, the problem of identification of cephalopod species as target or by-catch in a fishery, as well as that of the use of its overall landing per unit effort (LPUE) as a reliable estimator of the species abundance, may be solved by examining the correlation between landings, overall effort (fishing days) and the resulting LPUE in a fishery.

In annex 9 of Report of the Working Group on cephalopod Fisheries and Life History (Anon, 2001) information was presented about the surveys in the Gulf of Cadiz. We now present two abundance indexes in the Gulf of Cadiz for Octopus vulgaris: landing and survey yields. Figure 3.2 shows total landings from 1993/1994 to 2000/2001 and an index of abundance from both the March (1993 to 2002) and November (1997 to 2001) surveys.

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Landing Survey-Mar. Survey-Nov 6000 12000

9000 4000

6000 hour Tn Gr/ 2000 3000

0 0 92/93 93/94 94/95 95/96 96/97 97/98 98/99 99/00 00/01 01/02 Years

Figure 3.2 Total annual landings (tonnes) from 1993 to 2002, and an average index of abundance (grammes/hour) from both the March (1993 to 2002) and November (1997 to 2001) research surveys.

The results indicate a good relationship exists between the total annual landings and the average index of abundance calculated from the March surveys (0.88 correlation coefficient). Furthermore, there is some evidence to suggest a relationship exists between the total annual landings and the index of abundance obtained during the November surveys. This latter relationship may prove particularly important, since it could help to predict the level of recruitment at the start of the fishery during October – November. More data is required in to test this hypothesis.

3.3 Total stock size assessments

Loliginid squid

Scotland

A synthesis of population assessments of the Loligo forbesi stock off the West of Scotland has been prepared by Graham Pierce (University of Aberdeen) and has been submitted to Fisheries Research.

The manuscript title is: "How many squid (Loligo forbesi) are there on the West Coast of Scotland" (authors: Young I.A.G., G.J. Pierce, H.I. Daly, M.B. Santos, L.N. Key, N. Bailey, J.-P. Robin, A.J. Bishop, G. Stowasser, M. Nyegaard, S.K. Cho, M. Rasero and J.M.F. Pereira).

In this paper abundance of cohorts 1995-2000 in ICES division VIa is estimated with depletion methods.

English Channel

The last population assessments for English Channel Loligo forbesi and Loligo vulgaris have been presented to the ICES 2001 Annual Science Conference in Oslo. The paper by Royer et al is now in press in the ICES J. Mar Sci. (to be published in 2002, issue 59). In this analysis, based on fishery statistics for 1993-1998, two methods (depletion methods and cohort analysis) provided similar estimates for the number of recruits. Fishing mortality vectors derived from cohort analysis were used in simulations in order to obtain a diagnostic of the exploitation of cohorts 1993 to 1996. Loligo forbesi resulted close to optimal exploitation and a slight growth over-fishing was detected in Loligo vulgaris. No cohort could give much higher yields with a reduced fishing effort.

The update of these assessments was limited by the availability of capture statistics. It is worth noting that a longer series of population estimates will also be useful in the analysis of stock-recruitment relationships.

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Cuttlefish

English Channel

Fish market sampling carried out by the University of Caen enabled to apply cohort analysis to English Channel Sepia officinalis. This study is presented in the working document n° 5 annexed to this report (Royer and Robin). Diagnostics for cohorts 1995 and 1996 should be looked with caution since exploitation rate can increase in periods of low recruitment like 1998.

3.4 General discussion

In an increasing number of cephalopod stocks the collection of quantitative data enables to monitor population trends in abundance. It is important to note that even when total population size estimates are not available yet trends in relative abundance indices are useful both to understand yield variation and to study stock-recruitment relationships.

Almost all assessments are derived from fishery statistics and biological sampling of commercial catches. When available surveys are used in comparisons of inter-annual trends but never seem to be carried out at an appropriate time scale for the analysis of cephalopod cohorts. The timing of surveys often does not coincide with the main fishery and the frequency of cruises is generally not adapted to annual species. In this context, the need to continue data collection programmes should be stressed.

At the present time, shared stocks (exploited by several European countries) are only assessed via biological sampling carried out by one laboratory (Scottish waters by Aberdeen, English Channel by Caen, etc.). National fisheries laboratories should be encouraged to extend landing sampling schemes to cephalopod resources. The analysis of landings structure is also necessary to better understand fleets interactions. In Southern areas exploited by Spain or Portugal there seems to be little interaction between countries (and besides Portugal national landings sampling scheme does cover cephalopods). Nevertheless fleet interaction studies are desirable within each country (in particular between offshore fishing boats and inshore, artisanal, fisheries).

The analysis of historical data can be developed with two different objectives:

(i) To understand how cohorts have been exploited, and (ii) To obtain predicting tools.

Parameters which are only available late in the fishing season have of course little predictive value though they can be useful to understand what happened. Depletion methods are more likely to provide predictions of recruitment strength at the beginning of a fishing season (Agnew et al, 2001) whereas age-based methods will remain more time consuming.

It is worth noting that in a number of situations cephalopod stock structure is rather simple and a small number of age- groups are being fished at the same time. It has already been pointed out by Pierce and Guerra (1994) that in this context the two methods were rather close (differing mainly by the set of assumptions used).

4 GEAR SELECTIVITY (TOR C)

Although there was no new data on gear selectivity for cephalopod provided to the 2002 working group meeting, there is still a need to take into account gear selectivity. Two consequences of variations in gear selectivity can justify specific studies of this problem:

1) most abundance indices are applied with the hypothesis of constant catchability

2) fleet interactions ought to be analysed in the light of selectivity of the different fishing gears.

In Portugal, a study on species selectivity of the traps used in the octopus fishery which was conducted within the CephAssess Study Project. Results of this analysis have been presented in the report submitted to the EU. Data were also gathered within the same project on size selectivity of the same gear but they are considered insufficient to publish at this stage.

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5 POSSIBLE PRECAUTIONARY APPROACHES TO MANAGEMENT (TOR D)

5.1 General considerations

Without a reliable way of forecasting abundance, cephalopod fisheries in European waters are unlikely to be controlled setting annual TAC. Nevertheless, the increasing importance of cephalopod resources for fishermen and the widespread distribution of cephalopod stocks are in favour of an integration of these resources within European Community's Common Fisheries Policy. In some areas, the depletion of traditional quota species is so dramatic that fishermen's income depend mainly on national resources and/or international non quota species like cephalopods.

Also it should be noted that in several fishing grounds fishermen are asking for quotas which they see as a mean to keep the proportions of cephalopod resources shared by each country in a statu quo.

A review of management tools applied in other cephalopod stocks in the world is going to be prepared within the CEPHSTOCK concerted action (beginning at the end of this year). In the meantime, WGCEPH can underline aspects of cephalopod life cycle and biology which will influence the choice of management options.

Stocks of semelparous species depend on the escapement of adults in sufficient numbers in spite of the fact that juveniles are fished. Escapement is generally achieved via fishing effort limitations which may concern only a part of the fishing season or a limited area.

Migrating populations use a range of habitats and at some stage exploitation can be reduced when the area is less suitable for fishing. Such situations are likely to change in time for instance with the development of gears able to fish on rocky bottom. More generally changes in mechanical equipment in artisanal fisheries should be taken into account.

The analysis of survival curves through the whole life span (including early life stages) should be developed in search of critical stages. Any measure increasing ecosystem carrying capacity in the critical stage is likely to have a higher impact on subsequent survivors than less targeted options.

When fluctuating populations can be described with a rather constant core (representing a source) and a more variable extended part (acting as a sink) management measures must concentrate on the protection of the core. An example of this situation in European cephalopods could be the extension of English Channel stocks into the Southern North Sea.

5.2 Management in European cephalopod fisheries

In the absence of management measures defined on a scientific basis for European cephalopod stocks WGCEPH discussed tools suggested by fishers.

Licences:

In several countries (France, Portugal) licences are more and more used by fishermen's organisations to identify fleets involved in the exploitation of cephalopods. At this stage it does not seem that these licences are likely to represent a tool in fishing effort limitation (as it is in the Falkland Islands fishery). Nevertheless, it can represent a useful source of information about boat activities and fishing gear. In the case of multi-species trawlers it can also help the identification of "métiers" (periods in the fishing season when trawlers are targeting cephalopods).

The availability of this information for scientific analysis remains problematic since the official system generally does not take into account licences.

Stock enhancement and protected areas:

An experiment is being carried out in the Western coast of Normandy to prevent cuttlefish to lay their eggs on the fishing traps (which are removed out of the water before eggs-hatching) by adding in the spawning ground egg laying devices. Very simple equipment (ropes partially buried in the sand) is used by cuttlefish females who attach their eggs to it. This experiment was partly initiated by fishermen using traps. However, consequences of these efforts on recruitment are not known and should be studied scientifically because density-dependent survival or growth in early life stages are likely to reduce their effect. Besides, traps catch only adult cuttlefish but the inshore trap fishery is dominated by offshore trawlers.

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Since other cephalopod species attach their eggs to the substrate, egg laying grounds could be protected (or the occurrence of egg laying ground taken into account in the definition of Marine Protected Areas which are more and more discussed). It should be noted that Fishermen in Normandy did not define a protected area rather than tried to continue to practice trap fishing in the spawning period.

A better knowledge of egg laying areas is desirable in which fishermen can co-operate although their indications can be biased. Again the protection of the core of a spawning ground can be an objective whereas egg masses observed outside as observed in Loligo forbesi by Lordan and Casey (1999) cannot be taken into account.

6 ENVIRONMENTAL FACTORS AFFECTING RECRUITMENT AND DISTRIBUTION PATTERNS (TOR E)

The term of reference is as follows: Compile available data on relationships between abundance and environmental conditions, factors affecting recruitment, migration and distribution patterns of juveniles and adults, and trophic interactions

Presentations made during the meeting correspond to two kinds of papers. New results of field work and experiments (Gonzalez et al working document n°2 and Koueta working document n° 4) and reflections and suggestions by J. Caddy (working document n° 1).

Although there is no ongoing international project on cephalopod environmental variation this year, attention was also given to local analyses like trawl survey data off Sicily (Jereb et al, 2001), environmental influences on fishing efficiency (Hill and Wakeford, 2001) and relationships between abundance and with temperature in the Gulf of Cadiz (Sobrino et al, 2002).

Relationship between environmental variable and abundance of Octopus and Sepia in the Gulf of Cadiz

Sobrino et al (2002) has studied the correlations between environmental variables and abundance of two species of benthic cephalopods in the Gulf of Cadiz. Primarily, they aim to reveal putative links between rainfall rates, river discharges, sea surface temperature and landings as indices of abundance. They analysed landings, effort and landing per unit effort (LPUE) patterns of common octopus (Octopus vulgaris) and cuttlefish (Sepia officinalis) from a time- series of eighteen years. They showed that both LPUE and landing may be used as suitable indices of abundance by analysing correlations between landings, LPUE and fishing effort. Pearson coefficients of correlation showed that octopus abundance is highly correlated with the rainfall previous to the fishing season, the river discharges of December, and sea surface temperature of may and June. Cuttlefish did not show correlations with any variable. While Octopus seems to be very much affected by the environment, particularly in early-life stages, cuttlefish survival is less susceptible to environmental changes or fluctuations. A multivariate regression linear model including Rain-1, SST5 (May) and SST6 (June) as predictors were built up.

Results from the multiplicative linear regression models

-1 2 (Ln (landing octopus) = Intercept + B1*Rain + B2*SST5 + B3*SST6 ) are significantly correlated (R = 0.76). The average of standardised residual is 0.001 and variance is 0.88. Figure 1 shows observed and predicted landing values. They must highlight the similarity of both data series.

Landing Estimation 4000

3000 T o 2000 n s 1000

0 Year 83-84 85-86 87-88 89-90 91-92 93-94 95-96

Figure 1 Observed and predicted values of the multivariate regression linear model for O. vulgaris.

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Octopus vulgaris paralarvae are sampled in Galician waters with zooplankton bongo nets, their occurrence is analysed in the context of a wind-driven upwelling system and preliminary results seem to indicate that more paralarvae are observed in enriched cold waters. Overall estimates of the number of paralarvae are computed and the figures should be compared with adult abundance (which shows correlation with the upwelling index with a 1-2 years time lag.

Sepia officinalis juvenile have been reared with live and frozen shrimps and also using enrichment with fatty acids or proteins. Results indicate that the quality of the diet enrichment during juvenile cuttlefish rearing depends on the age of the animal. Poly Unsaturated Fatty Acids appears very important until 20 days old after what protein enrichment is more important.

Suggestions for future analyses

Reflections presented by J. Caddy to WGCEPH are related to the idea that natural and fishing mortality rates in juvenile concentration areas can lead to the selection of faster growing individuals. Cephalopods are known as fast growing species and individual growth rates can show high variations. The hypothesis presented could be tested in species in which individual age is determined with statoliths.

7 CEPHALOPOD LITERATURE RELEVANT TO FISHERIES (TOR F)

Information on literature relevant to cephalopod fisheries published during the last calendar year (2001-2002) was downloaded from bibliographic databases and supplied by WGCEPH members. This information is presented in Annex 4and divided in two sections (journal papers and grey literature).

8 RESEARCH PRIORITIES

8.1 Funding for data collection

It is important to repeat here that WGCEPH relies on a wide range of scientists working outside the traditional government fisheries laboratory. Considerable increase in the knowledge of Northeast Atlantic cephalopod fisheries has been acquired. Assessments and exploitation diagnostics are now being developed in a context of low interest from still many national fisheries research laboratories. A direct consequence of this is that most biological data collection and some of the basic fishery data collection is project-based rather than part of national fishery data collection scheme.

The new management of EU funding for fishery data collection (Council Resolution 1543/2000) is a quite significant change in cephalopod fisheries monitoring. Its negative effects must be underlined and all the consequences of downgraded cephalopod data collection must be known.

WGCEPH believes that the low priority assigned to collection of data on cephalopods is not consistent with their current importance as fishery resources and will seriously impede progress towards assessment and management of these stocks in the future. Fishery statistics compiled in this report as well as comparisons of catch statistics from Northeast Atlantic and from European Mediterranean confirm that data collection priorities are not consistent with the importance of Atlantic cephalopod stocks for the fishing industry.

The list of species and areas to be sampled (Appendix XIV of the DG Fisheries Report on Council Resolution 1543/2000, see Table 7.1) was prepared in a commission where only those WGCEPH members from Spain and Portugal (working in national fishery institutes) had any involvement in the process of setting priorities.

According to the Council Resolution the list of species should be updated every year.

WGCEPH believes that this is an opportunity for DG Fisheries to adopt a more realistic approach of cephalopod species for which data collection is needed. For instance Eledone cirrhosa and Eledone moschata were included in priority data collection in the Mediterranean (1000 T landed by Italy in 1999) whereas in the Bay of Biscay, English Channel, Celtic Sea, and west coasts of Scotland and Ireland, Octopus vulgaris, Loligo vulgaris, and Sepia officinalis (20 000T of cuttlefish caught in 1999) appear only in the list of optional species. Loligo forbesi is not even on the list although it represents about 80% of English Channel squid landings (4 200 T in 1999) and is the only squid exploited off Scotland (1250 T in 1999). Besides Loligo forbesi is the first species for which different Northeast Atlantic stocks have been assessed (Pierce et al 1996, Royer et al 2002) within the framework of EU funded study projects.

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Another general concern is that the nature of the sampling programme does not take into account the biological differences between cephalopods and most quota finfish species. Namely, cephalopods are short-lived (generally annual) species, with extended periods of recruitment and spawning and complex migratory movements. They show wide interannual fluctuations in distribution and abundance, which will not easily be detected or predicted by low- resolution sampling. Landings and market sample data are needed on (minimally) a monthly basis to allow assessment by depletion methods. Collection of landings data by ICES rectangle is desirable since understanding of the complex migratory patterns and relationships with environmental factors is not possible without these baseline data.

A third point is the fact that Northeast Atlantic cephalopod stocks are indeed European shared resources. They are not managed yet by the EU (and TAC may not be the best tool for such populations) but assessments are being developed for these stocks which can be more important for fishermen than actual quota species. This "international" context is clear off Scotland and in the English Channel. It is also the case in the Bay of Biscay where the lack of biological sampling is the main impediment towards stock assessments.

A series of more specific recommendations were listed in 2001 WGCEPH report. There is no need to repeat them here although they are still valid. In particular, WGCEPH considers that scientists who have demonstrated expertise in cephalopod fisheries should be consulted by decision makers so that 10 years of EC funded research on exploited cephalopod stocks do not be lost.

8.2 Research priorities

A rather comprehensive list of priorities was presented in the 2001report. WGCEPH considers that all these points are still very worth studying and would all contribute usefully to the definition of sustainable exploitation.

This year's WGCEPH meeting (in the light of fishery trends as well as research recent outputs) suggests however to focus the attention on two points:

The assessment of most important stocks (the major stocks like English Channel cuttlefish, Octopuses off Portugal and Spain and English Channel common squid are significant resources shared by several countries which is also the case in common squid off Scotland). In spite of the lack of information about some population parameters (like natural mortality) the comparison of results obtained with methods based on different sets of assumptions should be encouraged. Stock assessments are not only useful for fisheries management but are also desirable for an approach of ecosystem changes.

The ecological study of early life stages. Almost all studies of environmental factors affecting cephalopod abundance have underlined that recruitment variations were the most important. Pre-recruit survival and growth are assumed to be key parameters. Difficulties in collecting juvenile have limited the available material however improved methodology and better knowledge of paralarvae distribution and behaviour can help to solve this. Also, back-calculated age with statolith readings, or rearing experiments can provide information about early life history.

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9 THE FUTURE PROGRAMME OF WGCEPH AND RECOMMENDATIONS

9.1 Terms of reference

WGCEPH considers that, broadly speaking, the present terms of reference continue to be relevant, although some minor points could be grouped under different headings (see below "gear selectivity").

The working group wishes to continue to gather expertise on European cephalopod fisheries and to make available to ICES Advisory Commission for Fisheries Management any progress in stock assessments. Also, the working group considers that progress in the understanding of cephalopod life history and of the role of cephalopod populations in changing ecosystems should be of interest to the Advisory Commission for Environmental Management.

The following terms of reference are recommended: a) update currently available landing statistics and information on fishing effort and discards and gear selectivity ; explore existing resource survey databases for information about sampled cephalopods in the ICES area; This activity remains fundamental to the work of the group. The broadening of the remit to include effort, discard and survey data was useful but improved data, and improved access to data, are needed. b) compile methods and results available for stock identification and estimation of population size of fished cephalopods; Arguably there is no need to repeat the compilation of methods, but it remains useful to continue to compile results. Updated assessments require both that fishery statistics become more quickly available and that biological sampling be continued or extended. c) identify possible precautionary approaches to the management of these cephalopod resources; and provide management options. a This task remains useful, since the outcome will be update as new results become available under tor b d) compile available data and identify relationships between abundance and environmental conditions, factors affecting recruitment, migration and distribution patterns of juveniles and adults, and trophic interactions; This tor essentially concerns review of ecological data needed to underpin progress on assessment and management. Recent analyses of European cephalopod populations have underlined the influence of environment during the pre- recruit stage on cohort strength, further progress require that biological and environmental data be collected simultaneously. e) review cephalopod culture techniques and results and their interest in the understanding of biological phenomena. This tor is suggested in order to analyse key life history stages in population assessments. The first results are expected in juvenile growth and survival. f) update the bibliographic database of cephalopod literature relevant to fisheries, including grey literature; This provides a useful service to the research community as well as to WGCEPH.

9.2 Next WGCEPH meeting

WGCEPH, more than most ICES Working Groups, relies on participation from a wide range of scientists working outside the traditional government fisheries laboratories in ICES countries and has, indeed, benefited enormously over the last 10 years from the input of other scientists working often in universities where no funding is available for participation in ICES activities. This should be taken into account in the organisation of the next WGCEPH meeting. In particular, the opportunity to use project funding should be seized. The EU funded Concerted Action CEPHSTOCK is starting in the last quarter of 2002 and so WGCEPH suggest to hold its next meeting in combination with the first CEPHSTOCK meeting in Lisbon in January 2003. Should this appear unrealistic, then WGCEPH would work by correspondence in 2003 and report to ICES by the end of May 2003.

10 OTHER BUSINESS AND CLOSING OF THE MEETING

10.1 Exchanges with scientists studying cephalopods in the Mediterranean

The WGCEPH meeting in Rome provided the opportunity to discuss with Mediterranean scientists about cephalopod fisheries in both areas (Northeast Atlantic and the Mediterranean). Contributions to this discussion appear in a

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comparison of capture statistics in FAO areas 27 and 37 presented by Luca Garibaldi (see Annex 2 FAO tables). A more detailed analysis of the Mediterranean case was prepared by Patrizia Jereb (Working Document n°3). This document mentions a series of figures which have not been included for space constrains but which can be obtained from the author ([email protected]). However, the fishery statistics table from which these figures were prepared is included in the document.

Comparisons between Northeast Atlantic and Mediterranean cephalopod productions indicate that landings from the ICES area have increased in the last decade while Mediterranean landings were decreasing. European Atlantic catches seem now to exceed Mediterranean (see graphs in Annex 2 page 10). This is mainly due to the increase in cuttlefish and squid Atlantic landings, whereas octopus landings are still more important in the Mediterranean.

Detailed analysis of the Mediterranean case (Patrizia Jereb, Working Document n°3) underlines the decreasing trend in Mediterranean catches in the last fourteen years and shows that in most groups of species (but Sepia officinalis) 1999 records reach their lowest values since the 70's. Although cephalopods have been heavily fished in the Mediterranean, one cannot tell now whether observed fluctuations result from changes in fishing pressure or correspond to environmental changes. Since Mediterranean countries play a major role in cephalopod consumption and trade changes in this area should be of interest to all European cephalopod fisheries.

10.2 Information about scientific meetings

(1) The 2nd International Workshop and Symposium of Pacific Squids will be held from November 25 - 29, 2002 at La Paz, northwest Mexico.

See http://www.cicimar.ipn.mx/eventos/Avicorrg.htm for details.

(2) "Coleoid cephalopods through time: neontological approaches to their palaeobiology in the light of the fossil record" from September 16 - 19, 2002 at Berlin, Germany.

See http://userpage.fu-berlin.de/~palaeont/Coleoid-Symposium.html. Switzerland.

The chairman closed the meeting at 1.00 pm on April 12th.

11 ACKNOWLEDGEMENTS

WGCEPH wishes to thank Patrizia Jereb and her colleagues at ICRAM for hosting the meeting, João Pereira (Portugal), Uwe Piatkowski (Germany), Robert Wakeford (UK), and Begoña Santos (UK) for assistance with drafting the report, and staff at ICES, notably Mette Bertelsen, for assistance with provision of data and for general assistance during 2001- 02.

12 REFERENCES

Agnew, D.J., Hill, S.L. Beddington, J.R. & R.C. Wakeford, 2001. Using predictions of recruitment strength to manage single-cohort stocks of an annual squid species: Loligo gahi in the Falkland Islands. ICES C.M. 2001/K:32 (poster).

Anonymous, 2001. Report of the working group on cephalopod fisheries and life history. ICES CM 2001/G:5. 73 p.

Hill, S.L. & R.C. Wakeford, 2001. Environmental influences on vessel efficiency in the Falkland Islands Loligo gahi fishery. ICES C.M. 2001/K:14 (poster).

Jereb P., Massi D., Norrito G. and Fiorentino F., 2001. Preliminary observations of environmental effects on spatial distribution and abundance of Eledone cirrhosa and Illex coindetii in the Strait of Sicily (Central Mediterranean Sea). ICES CM 2001/K:34 (Poster)

Lordan C., and Casey J., 1999. The first evidence of offshore spawning in the squid species Loligo forbesi. J.Mar. Biol. Ass. U.K., 79, 379-381.

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Pierce G. J. and Guerra A., 1994. Stock assessment methods used for cephalopod fisheries. Fisheries Research, 21, 255-286.

Pierce, G.J., Boyle, P.R., Hastie, L.C. and Shanks, A., 1994. Distribution and abundance of the fished population of Loligo forbesi in UK waters: analysis of fishery data. Fisheries Research, 21: 193-216.

Pierce, G.J., Bailey, N., and Robin, J.P. 1996. Stock assessment for Loligo spp. in the Northeast Atlantic. ICES CM 1996/K:23.

Royer J., Peries P., and Robin J.P. 2002. Stock assessments of English Channel Loliginid squid: updated depletion methods and new analytical methods. ICES J. Mar. Sci., 59, (in press).

Sobrino I., Silva L., Bellido J.M. and Ramos F. 2002. Rainfall, river discharges and sea temperature as factors affecting abundance of two coastal benthic cephalopod species in the Gulf of Cadiz. Bull. Mar. Sci. (In Press)

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Table 2.1 Landings (in tonnes) of Cuttlefish (Sepiidae). Country 1995 1996 1997 1998 1999 2000 2001P

ICES Division IVb (Central North Sea) Belgium 11233 7 4 Netherlands +++++ 2 n.a.

ICES Division IVc (Southern North Sea) Belgium 155445 12 22 England, Wales & Northern Ireland 163 90 22 28 22 14 5 France 234 174 135 140 231 420 184 Netherlands +++++ 97 n.a.

ICES Division VIa,b (NW coast of Scotland and North Ireland, Rockall) England, Wales & Northern Ireland + + 0 + 0 0 5 France 131017 1 + Spain +111416 0 0 0

ICES Division VIIa (Irish Sea) Belgium 21111 1 2 England, Wales & Northern Ireland 198111 1 + France 1100+ 1 1

ICES Divisions VIIb, c (West of Ireland and Porcupine Bank) England, Wales & Northern Ireland 00043 0 0 Spain +101314 0 0 0

ICES Divisions VIId, e (English Channel) Belgium 19 11 6 15 9 35 25 Channel Islands 1 11 8 20 22 26 4 England, Wales & Northern Ireland 3925 4038 1634 2449 2014 2910 2600 France 8869 8012 5742 7530 8266 10894 6932 Netherlands +++++ 2 n.a.

ICES Division VIIf (Bristol Channel) Belgium 4 1 1 + 1 1 1 England, Wales & Northern Ireland 42 64 44 39 9 12 7 France 14 33 29 36 23 22 10

ICES Divisions VIIg-k (Celtic Sea and SW of Ireland) Belgium 52334 2 3 England, Wales & Northern Ireland 188 367 464 220 206 139 79 France 18 34 21 946 872 965 768 Spain + 46 57 181 122 110 18

ICES Subarea VIII (Bay of Biscay) Belgium + + 0 0 1 1 + England, Wales & Northern Ireland 2 40 37 19 4 0 + France 3878 4058 5118 4363 5031 5118 4903 Portugal 0 11 8 11 5 8 10 Spain 194 260 368 593 829 511 342

ICES Subarea IX Portugal 981 1625 1415 1723 1156 1357 1338 Spain 1025 819 1504 1916 1868 1339 765

Grand Total 19601 19736 16652 20275 20725 24008 18028

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Table 2.2 Landings (in tonnes) of Common Squid (includes Loligo forbesi, L. vulgaris, Alloteuthis subulata and A. media). Country 1995 1996 1997 1998 1999 2000 2001P

ICES Division IIIa (Skagerrak and Kattegat) Denmark 11686 7 2 Sweden 2+111 + +

ICES Division IVa (Northern North Sea) Denmark 11253 3 2 England, Wales & Northern Ireland +0032 3 0 France 0010+ + + Germany +++++ + + Scotland 268 279 453 844 712 547 350

ICES Division IVb (Central North Sea) Belgium 14 9 7 11 16 24 11 Denmark + + 9 3 18 10 1 England, Wales & Northern Ireland 22 21 39 144 65 29 146 Germany 31355 3 2 Netherlands +++++ 4 n.a. Scotland 25 14 66 214 144 87 ?

ICES Division IVc (Southern North Sea) Belgium 153 87 39 36 72 121 109 England, Wales & Northern Ireland 103322 4 12 France 188 85 123 93 151 165 244 Germany 62161 2 2 Netherlands +++++ 758 n.a.

ICES Division Vb (Faroe Grounds) England, Wales & Northern Ireland + 0 0 + + + + Faroe Islands + + 5 32 23 + n.a. Scotland +1112 2 8

ICES Division VIa (NW coast of Scotland and North Ireland) England, Wales & Northern Ireland 16 49 40 7 3 2 ? France 98 132 82 136 88 56 8 Ireland 85 114 140 99 106 38 38 Scotland 267 287 301 285 334 210 195 Spain +++78 + 1

ICES Division VIb (Rockall) England, Wales & Northern Ireland 2 8 5 14 1 + ? Ireland 106122 3 0 Scotland 6 19 5 27 13 5 35 Spain 2617649 2 + +

ICES Division VIIa (Irish Sea) Belgium 28253 3 2 England, Wales & Northern Ireland 156 218 125 173 40 31 107 France 14 9 5 17 11 12 22 Ireland 7 9 6 22 13 5 8 Isle of Man 73222 + + Scotland 22322 2 ?

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Table 2.2. continued.

Country 1995 1996 1997 1998 1999 2000 2001P

ICES Divisions VIIb, c (West of Ireland and Porcupine Bank) England, Wales & Northern Ireland 96 307 228 162 59 40 49 France 22 84 80 60 34 76 7 Ireland 50 48 42 34 40 26 49 Scotland 176457134 27 ? Spain + 55 69 51 0 2 18

ICES Divisions VIId, e (English Channel) Belgium 220 163 77 133 113 254 223 Channel Islands 216511 9 n.a. England, Wales & Northern Ireland 672 392 496 419 641 449 437 France 2636 2033 2518 2689 3417 3227 2643 Netherlands +++++ 11 n.a.

ICES Division VIIf (Bristol Channel) Belgium 1312666 8 12 England, Wales & Northern Ireland 132 39 77 29 68 16 55 France 275 164 193 126 147 88 245

ICES Divisions VIIg-k (Celtic Sea and SW of Ireland) Belgium 26 63 10 13 9 5 3 England, Wales & Northern Ireland 1002 1381 924 505 377 202 162 France 118 50 69 325 402 259 224 Ireland 80 143 168 158 123 67 16 Scotland 1 121 127 128 109 100 ? Spain 29 241 302 225 352 68 1

ICES Subarea VIII (Bay of Biscay) Belgium 40 46 14 49 3 48 8 England, Wales & Northern Ireland 55 46 68 8 3 + + France 1565 1419 1489 829 1571 1256 1076 Portugal 02221 1 1 Spain 196 418 505 811 826 217 274

ICES Subarea IX France ++++4 42 4 Portugal 908 463 848 1011 329 619 862 Spain 245 236 1301 1043 540 875 753

ICES Subarea X (Azores Grounds) Portugal* 250 200 303 98 45 58 48

Grand Total 10001 9632 11519 11245 11115 10186 8471

*Landings consist exclusively of Loligo forbesi.

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Table 2.3 Landings (in tonnes) of Short-finned Squid (Illex coindetii and Todaropsis eblanae), European Flying Squid (Todarodes sagittatus), and Neon Flying Squid (Ommastrephes bartrami). Country 1995 1996 1997 1998 1999 2000 2001P

ICES Subarea I + II (Barents Sea and Norwegian Sea) Norway* 352 + 190 2 + + n.a.

ICES Division Va (Iceland Grounds) Iceland* 113543 1 n.a.

ICES Division VIa, b (NW coast of Scotland and North Ireland, Rockall) England, Wales & Northern Ireland + + + 3 5 + 0 Ireland ++++0 + 38 Spain 0 43 112 177 3 + 0

ICES Division VIIa (Irish Sea) England, Wales & Northern Ireland 00000 + 7 Ireland 17 23 + + 0 0 8

ICES Divisions VIIb, c (West of Ireland and Porcupine Bank) England, Wales & Northern Ireland 0 0 8 39 18 35 0 France 00001 29 5 Ireland 21 36 + 52 + 29 93 Spain + 38 97 150 69 84 ?

ICES Divisions VIId, e (English Channel) England, Wales & Northern Ireland +0100 0 1 France 11112 3 4

ICES Divisions VIIg-k (Celtic Sea and SW of Ireland) England, Wales & Northern Ireland 29 13 14 251 181 151 0 France 0 0 2 49 72 122 45 Ireland 167 312 + 295 9 83 60 Spain + 38 97 150 69 84 0

ICES Subarea VIII (Bay of Biscay) England, Wales & Northern Ireland 60300 0 0 France 136 139 372 166 228 185 66 Portugal 011151 2 1 Spain 360 1830 2013 1806 1453 652 560

ICES Subarea IX Portugal 101 121 353 383 325 325 232 Spain 149 1495 2536 1800 4476 699 283

Grand Total 1703 4221 6145 5841 7719 6425 1709

*Landings consist exclusively of Todarodes sagittatus.

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Table 2.4 Landings (in tonnes) of Octopods (Eledone spp. and Octopus vulgaris). Country 1995 1996 1997 1998 1999 2000 2001P

ICES Division IVa (Northern North Sea) Scotland 2 2 6 13 17 15 6

ICES Division IVb (Central North Sea) Belgium 0 + + 2 5 5 5 England, Wales & Northern Ireland 00011 1 + Scotland 00011 + +

ICES Division IVc (Southern North Sea) Belgium 2 0 2 + 2 1 1 England, Wales & Northern Ireland 8 4 1 + + + +

ICES Division VIa, b (NW coast of Scotland and North Ireland, Rockall) Belgium 0011+ + 0 England, Wales & Northern Ireland 00020 + 0 Ireland 1 1 + 0 1 1 + Scotland 4110+ 0 + Spain 0273542 0 + 0

ICES Division VIIa (Irish Sea) Belgium 14 3 18 26 4 5 11 England, Wales & Northern Ireland 2 0 1 + + + 0 Ireland 1 + 0 1 0 + 0

ICES Divisions VIIb, c (West of Ireland and Porcupine Bank) England, Wales & Northern Ireland +4353 4 2 France 0000+ 8 1 Ireland 22402 4 4 Spain + 27 33 41 34 9 0

ICES Divisions VIId, e (English Channel) Belgium 6 1 1 + + + + Channel Islands 0000+ + + England, Wales & Northern Ireland 77 75 37 17 9 22 + France 4523738 13 3

ICES Division VIIf (Bristol Channel) Belgium 96633 13 1 England, Wales & Northern Ireland 86934 10 0 France 2210+ + +

ICES Divisions VIIg-k (Celtic Sea and SW of Ireland) Belgium 27 17 13 11 10 16 6 England, Wales & Northern Ireland 144 127 66 58 16 78 9 France 20198 32 19 Ireland 425327 7 7 Scotland 05191 5 0 Spain 452 116 145 179 348 286 326

ICES Subarea VIII (Bay of Biscay) Belgium 314417 4 5 England, Wales & Northern Ireland + 5 23 1 + 0 0 France 68 49 84 78 225 167 72 Portugal 107 113 75 57 156 250 70 Spain 1779 2486 2448 2787 1261 1057 1272

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Table 2.4. continued

Country 1995 1996 1997 1998 1999 2000 2001P

ICES Subarea IX Portugal 9708 11523 9078 6350 9098 9019 7177 Spain 3741 2991 3630 3298 4490 5205 2178

ICES Subarea X (Azores Grounds) Portugal* 8 16 64 39 12 11 n.a.

Grand Total 16226 17658 15801 13043 15743 16248 11175

*Landings consist exclusively of Octopus vulgaris.

Table 2.5 Total annual cephalopod landings (in tonnes) in whole ICES area separated into major cephalopod species groups.

Cephalopod Group 1995 1996 1997 1998 1999 2000 2001P

Cuttlefish 19601 19736 16652 20275 20725 24008 18028 Common Squid 10001 9632 11519 11245 11115 10186 8471 Short-finned Squid 1703 4221 6145 5841 7719 6425 1709 Octopods 16226 17658 15801 13043 15743 16248 11175

Total 47531 51247 50117 50404 55302 56867 39383

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Table 2.6 Total annual cephalopod landings (in tonnes) in whole ICES area by country and separated into major cephalopod species groups. Country 1995 1996 1997 1998 1999 2000 2001P

(a) Cuttlefish (Sepiidae)

Belgium 46 21 17 26 24 59 57 Channel Islands 1 11 8 20 22 26 4 England, Wales & N. Ireland 4339 4607 2202 2760 2259 3076 2696 France 13015 12315 11046 13015 14440 17421 12798 Netherlands + + + + + 101 n.a. Portugal 881 1636 1423 1734 1161 1365 1348 Spain 1219 1146 1956 2720 2819 1960 1125 Total 19601 19736 16652 20275 20725 24008 18028

(b) Common Squid (Loliginidae)

Belgium 468 388 155 253 222 463 368 Channel Islands 2 1 6 5 11 9 ?? Denmark 2 2 17 16 27 20 5 England, Wales & N. Ireland 2163 2464 2005 1466 1261 776 968 Faroe Islands + + 5 32 23 + n.a. France 4916 3976 4560 4275 5825 5181 4469 Germany 9 3 4 11 6 5 4 Ireland 232 320 357 315 284 139 111 Isle of Man 7 3 2 2 2 + n.a. Netherlands + + + + + 773 n.a. Portugal 1158 665 1153 1111 375 678 911 Scotland 570 799 1001 1572 1350 980 588 Spain 472 1011 2253 2186 1728 1162 1047 Sweden 2 + 1 1 1 + + Total 10001 9632 11519 11245 11115 10186 8471

(c) Short-finned Squid (Ommastrephidae)

England, Wales & N. Ireland 35 13 26 293 204 186 8 France 137 142 375 216 303 339 120 Iceland 11 3 5 4 3 1 n.a. Ireland 205 371 + 347 9 112 199 Norway 352 + 190 2 + + n.a. Portugal 101 122 364 388 326 327 233 Spain 862 3570 5185 4591 6874 5014 1149 Total 1703 4221 6145 5841 7719 6425 1709

(d) Octopods (Octopodidae)

Belgium 61 28 45 47 41 44 29 Channel Islands 0 0 0 0 + + n.a. England, Wales & N. Ireland 239 221 140 87 33 115 11 France 117 74 93 90 241 220 95 Ireland 8 28 7 3 10 12 11 Portugal 9823 11652 9217 6446 9266 9280 7247 Scotland 6 8 8 23 19 20 6 Spain 5972 5647 6291 6347 6133 6557 3776 Total 16226 17658 15801 13043 15743 16248 11175

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AGENDA

ICES Working Group on Cephalopod Fisheries and Life History [WGCEPH]

(Chair: J-P. Robin, France) Meeting in Rome, Italy from 10-12 April 2002 hosted by Patrizia Jereb (ICRAM)

Agenda

Day 1: Wednesday 10th (afternoon)

Opening of meeting 2 pm at ICRAM

(Participants are invited to meet at the Battistini Metro station at 12:30 am, Patrizia Jereb organising a special shuttle service from here to the institute).

Introduction and practical arrangements (J.P. Robin, P. Jereb)

Presentation of ICRAM (P. Jereb)

Presentation of recent results in cephalopod research

Preparation of links/contacts with other scientific organisations interested in cephalopods

(FAO, ICRAM)

Preparation of tor f (Literature compilation)

End of this afternoon session by 6 pm.

Day 2: Thursday 11th

Morning

(9:30 at ICRAM shuttle service proposed by Patrizia at 8:45 from the Battistini Metro station)

Review of WGCEPH Terms of Reference tor a) update currently available landing statistics and information on fishing effort and discards; explore existing resource survey databases for information about sampled cephalopods in the ICES area. comparison of ICES and FAO data sets (discussion with FAO scientist) tor b) compile methods and results available for stock identification and estimation of population size of fished cephalopods tor c) compile available data on gear selectivity for cephalopods

Afternoon tor d) identify possible precautionary approaches to the management of cephalopod resources; tor e) compile available data on relationships between abondance and environmental conditions, factors affecting recruitment, migration and distribution patterns of juveniles and adults, and trophic interactions;

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Day 3: Friday 12th (9:30)

Review of WGCEPH Terms of Reference (continued)

Morning:

Discussion about future projects (for example links between WGCEPH and CEPHSTOCK) Round table about differences/links and peculiarities of Mediterranean cephalopod studies (discussion with FAO and ICRAM scientists)

End of the meeting at ICRAM by 1 pm.

List of Annexed Documents

Annex 1: list of WGCEPH members provided by ICES secretariat (February 25th, 2002). 3 p.

Annex 2: FAO tables (1970 - 2000 capture statistics for cephalopods in areas 27 and 37 and comparison with ICES data). 10 p.

Annex 3: Sample visualisation (maps) of WGCEPH fishery statistics tables, inter-annual trends per area and year 2000 landings per area and per country. 10 p.

Annex 4: Compilation of cephalopod literature references and grey publications for the period 2001-2002. 10p.

Working Document n° 1: Caddy J. Suggestions for an experiment on size-selective mortality, especially for species subject to small mesh selection during juvenile aggregations. 6 p.

Working Document n° 2: Gonzalez A., Guerra A., Otero J., and Rocha F.J. Influence of oceanographic factors and abundance of Octopus vulgaris in Galician waters (NE Atlantic). 6 p.

Working Document n° 3: Jereb P. Recent trends in Mediterranean cephalopods capture production. 9 p.

Working Document n° 4: Koueta N. Experimental study of the effect of enriched frozen diet on survival and growth of juvenile cuttlefish Sepia officinalis L. 9 p.

Working Document n° 5: Royer J. and Robin J.P. Assessment of the English Channel cuttlefish stock based on analytical methods. 6 p.

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ANNEX 1

02-25-2002 List for WGCEPH/ Mr E.G. Dawe Dr Lawrence Jackobson Dept. of Fisheries & Oceans Northeast Fisheries Science Center Dr J.P. Andrade Northwest Atlantic Fisheries NMFS/NOAA Universidade do Algarve, UCTRA Centre Woods Hole, MA 02543-1026 Campus de Gambelas P.O. Box 5667 USA 8000-810 Faro St John's, Nfld A1C 5X1 [email protected] Portugal Canada TEL: +15084952317 [email protected] [email protected] FAX: +15084952393 no telephone number available no telephone number available no fax number available no fax number available Dr Noussithé Koueta Lab. de Biologie et Dr Alexander Arkhipkin Dr S. Desclers Biotechnolgies Marines Fisheries Department Jackson Environment Institute Esplanade de la Paix Falkland Islands Government University College London B.P. 5186 P.O. Box 598 Stanley 5 Gower Street 14032 Caen Cedex Falkland Islands London WC1E 6HA France [email protected] [email protected] [email protected] no telephone number available no telephone number available TEL: 33 (0)2 31 53 80 16 no fax number available no fax number available FAX: 33 (0)2 31 53 80 09

Dr N. Bailey Mr E. Gaard Dr Han-Lin Lai Fisheries Research Services Faroese Fisheries Laboratory Northeast Fisheries Science Center Marine Laboratory Nóatún NMFS/NOAA P.O. Box 101 P.O. Box 3051 Woods Hole, MA 02543-1026 Victoria Road FO-110 Tórshavn USA Aberdeen AB11 9DB Faroe Islands [email protected] United Kingdom Denmark TEL: +1 508 4952238 [email protected] [email protected] FAX: +1 508 4952393 TEL: +44 1224 876544 no telephone number available FAX: +44 1224 295511 no fax number available Dr C. Lordan The Marine Institute Dr T.C. Borges Dr Simeon Hill Fisheries Res. Centre Universidade do Algarve, UCTRA Renewable Resources Abbotstown Campus de Gambelas Assessment Group Dublin 15 8000-810 Faro T.H. Huxley School of Ireland Portugal Environment [email protected] [email protected] Imperial College no telephone number available TEL: 351.289 800 924 8 Prince's Gardens no fax number available FAX: 351.289 818 353 London SW7 1NA United Kingdom Mr P. Lucio Mr Steve Cadrin [email protected] AZTI Northeast Fisheries Science Center no telephone number available Txatxarramendi Irla NMFS/NOAA no fax number available 48935 Sukarrieta/Pedernales Woods Hole, MA 02543-1026 Spain USA Mr T. Howell [email protected] [email protected] Fisheries Research Services no telephone number available TEL: +1 508 495 2335 Marine Laboratory no fax number available FAX: +1 508 495 2393 P.O. Box 101 Victoria Road Dr W.K. Macy Dr M. Collins Aberdeen AB11 9DB Graduate School of Oceanography University of Aberdeen United Kingdom University of Rhode Island Zoology Department [email protected] South Ferry Road Tillydrone Avenue TEL: +44 1224 876544 Narragansett, RI 02882 Aberdeen AB9 2TN FAX: +44 1224 295511 USA United Kingdom [email protected] [email protected] TEL: +1 401 8746174 FAX: +1 401 8746853

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Mr J. Martinez-Portela Dr G. Pierce Ms Marina Santurtun Inst. Español de Oceanografía Dept of Zoology AZTI Centro Oceanográfico de Vigo University of Aberdeen Txatxarramendi Irla Apdo 1552 Tillidrone Avenue 48935 Sukarrieta/Pedernales E-36280 Vigo Aberdeen AB24 2TZ SPAIN Spain United Kingdom [email protected] [email protected] [email protected] no telephone number available no telephone number available TEL: 44 (0)1224 272459 no fax number available no fax number available FAX: 44 (0)1224 272396 Mr Ignacio Sobrino Ms A. Moreno Dr Jean-Paul Robin Unidad Oceanográfica de Cádiz IPIMAR Biologie & Biotechnologies Muelle de Levante, s/n Avenida de Brasilia Marines 11071 Cádiz P-1449-006 Lisbon I.B.B.A., Universite de Caen Spain Portugal 14032 Caen Cedex [email protected] [email protected] France no telephone number available no telephone number available [email protected] no fax number available no fax number available TEL : 33 2 31 53 80 17 FAX : 33 2 31 53 80 09 Dr M. Vecchione Mr J.M.F. Pereira NMFS/NOAA IPIMAR Dr P. Rodhouse Systematics Laboratory Avenida de Brasilia BAS National Museum of Natural P-1449-006 Lisbon High Cross, Madingley Road History Portugal Cambridge CB3 OET Washington D.C. 20560 [email protected] United Kingdom USA no telephone number available [email protected] [email protected] no fax number available no telephone number available no telephone number available no fax number available no fax number available Dr U. Piatkowski Institut für Meereskunde Dr B. Santos K Zumholz an der Universität Kiel University of Aberdeen Institut für Meereskunde Düsternbrooker Weg 20 Dept. of Zoology an der Universität Kiel D-24105 Kiel Tillydrone Avenue Düsternbrooker Weg 20 Germany Aberdeen AB9 2TZ D-24105 Kiel [email protected] United Kingdom Germany no telephone number available [email protected] [email protected] no fax number available no telephone number available no telephone number available no fax number available no fax number available

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32

ANNEX 2

FAO Table 1: 1970-2000 FAO capture statistics for cephalopods in area 27 (in metric tons)

Country Scientific name English name Area 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 Belgium Sepia officinalis Common cuttlefish 27 ...... Belgium Loligo spp Common squids nei 27 200 100 100 100 90 206 97 278 86 37 22 58 116 308 238 241 Belgium Octopodidae Octopuses, etc. nei 27 ...... Channel Islands Sepia officinalis Common cuttlefish 27 ------...... 1 3 Channel Islands Loligo spp Common squids nei 27 0 0 0 0 0 1 1 1 1 1 1 1 0 3 2 1 Denmark Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 27 0 0 0 0 0 - - 0 11 9 15 20 23 9 6 5 Denmark Illex illecebrosus Northern shortfin squid 27 ...... - - Faeroe Islands Illex illecebrosus Northern shortfin squid 27 ------France Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 27 ...... 3 945 6 591 3 715 6 737 4 223 1 933 2 378 3 805 4 555 9 191 7 648 7 583 France Illex coindetii Broadtail shortfin squid 27 ------33 137 - - 130 413 136 71 France Loliginidae, Ommastrephidae Various squids nei 27 7 900 11 500 7 400 11 500 3 374 4 094 4 432 3 902 3 182 432 2 563 2 036 3 991 5 361 2 418 2 988 France Octopodidae Octopuses, etc. nei 27 ... - - - - 8 15 - 9 - 1 - 9 22 9 26 Germany Illex illecebrosus Northern shortfin squid 27 ------4 5 2 0 1 Germany Loliginidae, Ommastrephidae Various squids nei 27 ------Iceland Todarodes sagittatus European flying squid 27 ------436 16 7 13 4 1 634 2 Ireland Loligo spp Common squids nei 27 ------7 166 70 51 100 253 392 198 264 274 Ireland Octopodidae Octopuses, etc. nei 27 ------4 13 - - Isle of Man Loligo spp Common squids nei 27 ...... 1 3 8 16 4 6 5 Isle of Man Octopodidae Octopuses, etc. nei 27 ------Italy Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 27 ------10 - - - - - Italy Loligo spp Common squids nei 27 ... ------208 - 485 1 264 - 100 - - - Italy Loliginidae, Ommastrephidae Various squids nei 27 ------588 - 475 - - - Italy Octopodidae Octopuses, etc. nei 27 ------8 - - - - - Japan Loliginidae, Ommastrephidae Various squids nei 27 - 0 0 100 66 - 1 ------Korea, Republic of Loliginidae, Ommastrephidae Various squids nei 27 ------Lithuania Loliginidae, Ommastrephidae Various squids nei 27 ...... Netherlands Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 27 100 0 0 0 49 36 75 84 7 5 12 25 39 366 - - Netherlands Loliginidae, Ommastrephidae Various squids nei 27 ------Norway Todarodes sagittatus European flying squid 27 - 400 0 - 0 0 - 260 346 1 668 2 977 9 780 18 385 18 025 7 803 13 819 Portugal Sepia officinalis Common cuttlefish 27 400 1 300 1 300 1 200 1 176 896 824 1 243 1 405 1 071 937 875 849 979 1 045 1 140 Portugal Illex illecebrosus Northern shortfin squid 27 ------Portugal Loligo spp Common squids nei 27 700 600 1 500 1 300 1 122 669 591 1 041 716 835 1 440 1 240 784 1 441 932 770 Portugal Octopodidae Octopuses, etc. nei 27 1 800 2 300 3 000 6 400 2 459 4 461 4 471 3 598 6 551 2 756 5 356 6 916 4 145 3 215 3 382 3 916 Spain Sepia officinalis Common cuttlefish 27 5 600 5,600 F 4 200 4 000 3 585 1 974 1 869 1 681 30 284 458 133 222 431 570 196 Spain Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 27 ------Spain Illex illecebrosus Northern shortfin squid 27 ... - - - - 1 706 - - 2 469 13 1 178 878 8 4 - - Spain Todarodes sagittatus European flying squid 27 2 400 2,400 F 2 300 ...... - 1 658 ... 182 387 - - - 372 1 785 965 Spain Loligo spp Common squids nei 27 3 200 3,200 F 2 000 2 400 4 200 2 786 2 423 3 083 3 048 158 1 202 963 91 111 849 702 Spain Loliginidae, Ommastrephidae Various squids nei 27 - - - - - ...... 98 831 1 188 30 25 Spain Octopodidae Octopuses, etc. nei 27 17 800 17,800 F 5 200 9 300 4 970 6 530 3 120 3 752 4 394 4 528 7 725 5 319 4 902 5 075 5 138 4 052 Sweden Loliginidae, Ommastrephidae Various squids nei 27 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 Un. Sov. Soc. Rep. Loliginidae, Ommastrephidae Various squids nei 27 - - - - - 40 - - - - - 2 100 342 53 - - United Kingdom Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 27 ...... 761 1 114 639 United Kingdom Loligo spp Common squids nei 27 1 100 1 400 600 800 238 421 581 393 360 98 127 330 554 487 654 675 United Kingdom Loliginidae, Ommastrephidae Various squids nei 27 300 300 100 200 230 650 908 1 041 316 256 273 391 637 215 353 5 United Kingdom Octopodidae Octopuses, etc. nei 27 ------1 26 70

TOTAL 41 500 46 900 27 700 37 300 25 504 31 069 24 788 27 468 27 439 15 581 28 654 35 240 41 618 48 254 36 043 38 174

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Annex 2 (Cont’d)

FAO table 1: 1970-2000 FAO capture statistics for cephalopods in area 27 (in metric tons) –continued–

Country Scientific name English name Area 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 Belgium Sepia officinalis Common cuttlefish 27 ...... 33 64 84 57 45 21 16 25 23 59 Belgium Loligo spp Common squids nei 27 271 187 265 391 544 89 171 215 290 467 386 154 252 222 463 Belgium Octopodidae Octopuses, etc. nei 27 ...... 46 44 37 40 62 28 46 49 41 44 Channel Islands Sepia officinalis Common cuttlefish 27 1 1 3 F 4 4 F 4 F 4 4 F 2 2 F 12 10 15 22 26 Channel Islands Loligo spp Common squids nei 27 1 2 2 F 2 2 F 2 F 1 1 F 2 2 F 1 6 5 11 9 Denmark Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 27 0121832225352320 - 2620 Denmark Illex illecebrosus Northern shortfin squid 27 ------1716 -- Faeroe Islands Illex illecebrosus Northern shortfin squid 27 ------1 - - 2 32 23 ... France Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 27 6 643 4 992 10 684 12 156 21 576 12 943 10 634 13 832 11 932 14 642 14 532 11 982 13 015 14 465 18 550 France Illex coindetii Broadtail shortfin squid 27 209 298 200 1 188 474 551 833 762 702 571 395 411 216 304 364 France Loliginidae, Ommastrephidae Various squids nei 27 3 244 3 785 5 402 6 781 4 805 4 135 5 394 6 881 6 140 6 017 3 852 4 413 4 289 5 946 5 422 France Octopodidae Octopuses, etc. nei 27 44 101 119 103 54 38 121 226 106 126 67 75 90 246 480 Germany Illex illecebrosus Northern shortfin squid 27 000000 -0311 - - - - - Germany Loliginidae, Ommastrephidae Various squids nei 27 ------3--2411 65 Iceland Todarodes sagittatus European flying squid 27 00------11354 31 Ireland Loligo spp Common squids nei 27 730 167 198 431 183 149 260 364 277 298 481 442 610 282 322 Ireland Octopodidae Octopuses, etc. nei 27 - - - 1 1 0 1 4 6 25 13 7 3 10 ... Isle of Man Loligo spp Common squids nei 27 31621127151567322 22 F Isle of Man Octopodidae Octopuses, etc. nei 27 00------Italy Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 27 ------Italy Loligo spp Common squids nei 27 ------Italy Loliginidae, Ommastrephidae Various squids nei 27 ------Italy Octopodidae Octopuses, etc. nei 27 ------Japan Loliginidae, Ommastrephidae Various squids nei 27 ------Korea, Republic of Loliginidae, Ommastrephidae Various squids nei 27 ------1------Lithuania Loliginidae, Ommastrephidae Various squids nei 27 ...... - - - - 241 ------Netherlands Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 27 ------101 Netherlands Loliginidae, Ommastrephidae Various squids nei 27 ------773 Norway Todarodes sagittatus European flying squid 27 89 3 936 1 183 5 - - - - - 352 0 190 2 0 0 Portugal Sepia officinalis Common cuttlefish 27 1 336 1 460 1 905 1 580 1 621 1 208 1 234 1 209 1 125 987 1 636 1 422 1 734 1 162 1 357 Portugal Illex illecebrosus Northern shortfin squid 27 84 926 420 353 322 512 777 ------Portugal Loligo spp Common squids nei 27 918 1 604 1 087 1 634 1 675 1 870 1 646 879 616 1 267 787 1 514 1 499 701 1 003 Portugal Octopodidae Octopuses, etc. nei 27 4 045 9 075 8 261 10 395 7 028 7 522 9 631 7 219 7 479 9 836 11 652 9 181 6 445 9 265 9 696 Spain Sepia officinalis Common cuttlefish 27 360 544 ... 377 510 561 - - - - 176 336 1 078 1 411 1 602 Spain Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 27 - - - 171 72 105 1 637 1 267 1 196 1 588 971 1 621 1 446 1 322 254 Spain Illex illecebrosus Northern shortfin squid 27 - 5 237 2 621 3 234 3 049 3 306 3 172 2 089 2 272 2 359 2 600 2 426 762 1 932 1 503 Spain Todarodes sagittatus European flying squid 27 1 915 ------973 2 508 2 137 2 481 2 417 Spain Loligo spp Common squids nei 27 743 849 560 853 690 709 1 647 1 251 1 280 1 135 1 015 2 520 2 190 1 376 1 038 Spain Loliginidae, Ommastrephidae Various squids nei 27 63 81 414 ------1 459 2 267 942 Spain Octopodidae Octopuses, etc. nei 27 2 625 8 524 6 547 5 331 6 757 4 829 8 670 8 135 5 822 7 318 5 650 6 292 6 350 4 374 5 731 Sweden Loliginidae, Ommastrephidae Various squids nei 27 0000113400011 1 - Un. Sov. Soc. Rep. Loliginidae, Ommastrephidae Various squids nei 27 ------United Kingdom Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 27 370 257 646 1 398 3 566 773 1 157 2 164 2 089 3 631 4 607 2 202 2 760 2 260 3 076 United Kingdom Loligo spp Common squids nei 27 1 648 1 294 1 702 1 211 934 697 1 227 1 655 1 947 2 272 3 264 3 004 3 039 2 611 1 757 United Kingdom Loliginidae, Ommastrephidae Various squids nei 27 0 1 17 1 946 1 445 892 1 085 543 273 666 13 26 301 208 189 United Kingdom Octopodidae Octopuses, etc. nei 27 68 110 122 156 145 151 137 182 313 333 229 148 111 63 135

TOTAL 25 410 43 433 42 366 49 740 55 502 41 155 49 860 49 030 43 978 54 033 53 371 50 983 49 948 53 066 57 341

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Annex 2 (Cont’d)

FAO table 2: 1970-2000 FAO capture statistics for cephalopods in area 37 (in metric tons)

Country Scientific name English name Area 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 Albania Sepia officinalis Common cuttlefish 37 ...... Albania Loligo spp Common squids nei 37 ...... Albania Octopus vulgaris Common octopus 37 ...... Algeria Sepia officinalis Common cuttlefish 37 ------...... Algeria Loligo spp Common squids nei 37 ------...... Algeria Octopodidae Octopuses, etc. nei 37 ...... Algeria Cephalopoda Cephalopods nei 37 ...... - - Croatia Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 ...... Croatia Loligo spp Common squids nei 37 ...... Croatia Octopus vulgaris Common octopus 37 ...... Croatia Cephalopoda Cephalopods nei 37 ...... Cyprus Sepia officinalis Common cuttlefish 37 66 55 62 52 45 44 45 53 64 57 54 74 70 91 95 80 Cyprus Octopodidae Octopuses, etc. nei 37 60 50 62 52 45 44 51 60 64 66 58 59 68 188 157 101 Egypt Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 346 384 442 338 304 322 255 337 560 743 356 644 691 990 780 640 France Sepia officinalis Common cuttlefish 37 ...... 274 296 298 383 312 199 258 322 347 397 398 296 239 France Illex coindetii Broadtail shortfin squid 37 ...... France Todarodes sagittatus European flying squid 37 ...... 18 21 10 France Loligo spp Common squids nei 37 217 961 1 400 1 073 216 288 390 320 423 529 369 381 301 372 203 325 France Loliginidae, Ommastrephidae Various squids nei 37 ------9 2 ... France Octopus vulgaris Common octopus 37 - - - - 957 1 605 1 329 1 163 1 001 925 1 044 1 061 1 064 823 699 647 France Cephalopoda Cephalopods nei 37 ------Gaza Strip(Palestine) Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 ...... Gaza Strip(Palestine) Loligo spp Common squids nei 37 ...... Greece Sepia officinalis Common cuttlefish 37 428 399 353 452 493 687 765 743 780 627 709 562 544 564 714 847 Greece Loligo spp Common squids nei 37 ...... 221 245 359 398 475 448 600 512 443 489 498 580 455 477 Greece Loliginidae, Ommastrephidae Various squids nei 37 549 562 282 321 293 219 181 175 183 185 203 255 275 301 248 236 Greece Octopus vulgaris Common octopus 37 ...... 427 368 294 459 Greece Octopodidae Octopuses, etc. nei 37 763 833 754 650 776 932 836 787 918 996 1 069 958 684 769 697 697 Israel Cephalopoda Cephalopods nei 37 ------Italy Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 12 081 11 620 14 279 12 017 13 212 11 820 10 666 10 449 8 272 9 025 13 570 10 955 12 564 12 795 11 653 14 354 Italy Todarodes sagittatus European flying squid 37 2 343 2 241 2 276 2 464 2 679 2 693 3 025 4 047 2 946 2 779 2 871 3 623 3 115 3 920 5 730 7 002 Italy Loligo spp Common squids nei 37 5 212 4 850 4 795 3 934 4 834 4 792 4 426 4 497 4 082 4 280 4 558 4 331 3 128 3 530 4 640 5 230 Italy Eledone spp Horned and musky octopuses 37 3 080 2 740 3 459 2 650 2 681 2 612 2 497 2 369 3 134 2 375 2 070 2 032 2 586 4 310 2 724 3 671 Italy Octopus vulgaris Common octopus 37 8 122 7 864 9 439 8 148 8 289 9 202 9 390 8 718 8 974 8 313 8 868 9 033 8 925 9 329 9 140 9 419

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Annex 2 (Cont’d)

FAO table 2: 1970-2000 FAO capture statistics for cephalopods in area 37 (in metric tons) –continued–

Country Scientific name Area 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 Lebanon Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 ------...... Lebanon Octopus vulgaris Common octopus 37 ...... Malta Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 0 0 0 0 2 3 18 16 5 9 10 13 8 5 10 57 Malta Todarodes sagittatus European flying squid 37 ...... - - Malta Loligo spp Common squids nei 37 00006 535533112 1 18 Malta Octopodidae Octopuses, etc. nei 37 0 0 0 100 39 34 33 29 28 17 13 8 7 17 22 38 Morocco Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 23 53 0 100 64 62 65 47 48 97 174 52 124 98 142 157 Morocco Loliginidae, Ommastrephidae Various squids nei 37 ...... 14 12 20 Morocco Octopodidae Octopuses, etc. nei 37 ...... 226 529 717 Slovenia Sepia officinalis Common cuttlefish 37 ...... Slovenia Loligo spp Common squids nei 37 ...... Slovenia Octopus vulgaris Common octopus 37 ...... Spain Sepia officinalis Common cuttlefish 37 ------Spain Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 2 356 2 897 2 500 1 700 1 500 1 282 1 485 1 571 2 038 1 526 1 884 1 220 1 273 1 419 1 350 1 409 Spain Todarodes sagittatus European flying squid 37 ------Spain Loligo spp Common squids nei 37 2 502 2 619 1 200 1 000 1 100 1 236 972 1 024 1 958 1 719 1 835 980 1 125 1 192 1 200 928 Spain Loliginidae, Ommastrephidae Various squids nei 37 ...... 100 ... 124 143 150 26 ------Spain Octopodidae Octopuses, etc. nei 37 4 066 4 233 5 000 4 300 4 200 4 590 5 441 4 871 3 947 3 003 4 717 4 650 6 082 6 628 5 918 7 111 Spain Cephalopoda Cephalopods nei 37 ------...... Tunisia Sepia officinalis Common cuttlefish 37 336 281 340 715 541 767 2 173 964 1 133 1 318 1 115 1 388 2 374 1 829 2 559 4 005 Tunisia Loligo spp Common squids nei 37 87 78 100 100 104 104 167 323 305 400 384 152 171 138 160 145 Tunisia Eledone spp Horned and musky octopuses 37 ...... Tunisia Octopus vulgaris Common octopus 37 105 203 252 900 700 2 144 2 201 3 312 3 675 3 254 4 022 2 819 2 222 3 328 4 851 5 192 Tunisia Cephalopoda Cephalopods nei 37 1 069 1 110 1 839 1 718 1 455 ...... Turkey Sepia officinalis Common cuttlefish 37 0 0 300 400 300 200 213 229 141 175 245 268 71 174 368 256 Turkey Loligo spp Common squids nei 37 51 2 0 100 100 50 64 30 10 38 56 127 45 169 290 322 Turkey Octopus vulgaris Common octopus 37 87 9 0 0 10 10 27 27 17 25 54 123 60 137 229 252 Yugoslavia SFR Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 151 157 186 171 206 233 261 208 182 150 213 170 156 188 164 159 Yugoslavia SFR Loligo spp Common squids nei 37 234 239 210 191 271 263 207 263 298 300 264 221 228 260 253 353 Yugoslavia SFR Octopus vulgaris Common octopus 37 253 218 132 99 98 85 107 133 132 122 107 106 114 113 100 150 Yugoslavia, Fed. Rep. Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 ...... of Yugoslavia, Fed. Rep. Loligo spp Common squids nei 37 ...... of Yugoslavia, Fed. Rep. Octopus vulgaris Common octopus 37 ...... of

TOTAL 44 587 44 658 49 883 44 264 46 275 47 024 48 275 47 673 46 272 43 852 51 660 47 102 49 398 55 292 56 706 65 723

35

O:\scicom\LRC\WGCEPH\REPORTS\2002\WGCEPH02.doc 35 36

Annex 2 (Cont’d)

FAO table 2: 1970-2000 FAO capture statistics for cephalopods in area 37 (in metric tons) –continued–

Country Scientific name English name Area 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 Albania Sepia officinalis Common cuttlefish 37 ...... 39 33 33 51 51 50 Albania Loligo spp Common squids nei 37 ...... 7 47 34 93 93 90 Albania Octopus vulgaris Common octopus 37 ...... 66 75 59 93 90 85 Algeria Sepia officinalis Common cuttlefish 37 ...... 424 400 F 350 F 400 F 500 F 600 F 351 567 619 492 312 300 F Algeria Loligo spp Common squids nei 37 ...... 331 250 F 200 F 250 F 250 F 300 F 101 53 138 202 240 230 F Algeria Octopodidae Octopuses, etc. nei 37 ...... 382 190 185 543 305 300 F Algeria Cephalopoda Cephalopods nei 37 ------70 29 30 F Croatia Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 Croatia Loligo spp Common squids nei 37 ...... 167 203 118 118 102 151 215 145 138 Croatia Octopus vulgaris Common octopus 37 ...... 263 330 268 273 233 287 290 170 127 Croatia Cephalopoda Cephalopods nei 37 ...... 155 231 187 193 180 293 435 ...... Cyprus Sepia officinalis Common cuttlefish 37 ...... 552 799 455 431 483 443 265 646 873 Cyprus Octopodidae Octopuses, etc. nei 37 90 92 95 79 108 106 103 174 218 153 111 89 146 140 122 Egypt Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 67 108 119 201 208 170 258 288 474 300 190 199 228 179 166 France Sepia officinalis Common cuttlefish 37 1 030 724 941 631 614 659 787 1 067 1 031 1 097 1 365 1 370 1 152 1 449 1 503 France Illex coindetii Broadtail shortfin squid 37 275 245 215 192 260 194 254 152 130 100 85 85 85 88 88 France Todarodes sagittatus European flying squid 37 ...... 34 42 France Loligo spp Common squids nei 37 6 23 ... 38 74 75 49 11 21 19 87 87 87 - - France Loliginidae, Ommastrephidae Various squids nei 37 311 277 278 244 262 262 261 220 136 150 135 465 465 241 223 France Octopus vulgaris Common octopus 37 ... 28 ... 112 66 107 86 86 67 ... 43 43 43 - - France Cephalopoda Cephalopods nei 37 806 659 643 1 163 1 400 1 250 1 196 1 161 873 1 146 706 439 439 1 493 1 354 Gaza Strip(Palestine) Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 ------55 46 Gaza Strip(Palestine) Loligo spp Common squids nei 37 ...... 41 62 98 144 145 F 145 F Greece Sepia officinalis Common cuttlefish 37 ...... 15 23 30 61 60 F 60 F Greece Loligo spp Common squids nei 37 1 202 1 026 1 514 1 483 1 996 1 736 1 780 1 941 2 853 2 516 1 819 2 381 1 807 2 732 1 609 Greece Loliginidae, Ommastrephidae Various squids nei 37 482 588 678 779 668 1 064 1 135 1 369 1 101 945 856 623 426 397 560 Greece Octopus vulgaris Common octopus 37 236 285 285 390 480 446 703 567 678 906 812 513 566 675 870 Greece Octopodidae Octopuses, etc. nei 37 624 631 830 834 1 356 2 296 3 563 2 456 2 886 2 856 2 919 2 952 1 673 1 910 2 193 Israel Cephalopoda Cephalopods nei 37 641 517 445 298 430 489 708 780 1 166 1 451 809 740 732 794 878 Italy Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 - - 95 87 F 91 F 76 F 68 64 50 50 50 98 76 120 117 Italy Todarodes sagittatus European flying squid 37 14 591 11 438 13 417 8 734 8 291 10 106 7 984 7 682 13 616 11 150 7 974 7 382 6 839 5 720 5 534 Italy Loligo spp Common squids nei 37 7 739 7 412 7 237 8 243 8 215 7 418 7 800 6 451 5 737 4 789 4 672 2 614 3 995 2 056 2 516 Italy Eledone spp Horned and musky octopuses 37 5 450 5 352 4 758 4 224 4 743 5 132 4 728 4 372 5 282 5 483 5 366 4 141 2 237 1 909 1 890 Italy Octopus vulgaris Common octopus 37 2 748 2 870 3 562 2 470 2 972 3 366 3 632 3 542 3 928 2 500 1 939 1 590 1 506 1 041 1 621

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Annex 2 (Cont’d)

FAO table 2: 1970-2000 FAO capture statistics for cephalopods in area 37 (in metric tons) –continued–

Country Scientific name English name Area 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 Lebanon Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 10 996 10 184 10 354 9 145 10 163 10 508 10 300 11 565 9 661 9 757 9 057 8 907 9 478 6 999 7 800 Lebanon Octopus vulgaris Common octopus 37 ... 25 25 25 20 F 25 25 F 25 25 25 25 50 25 50 25 Malta Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 ...... 25 25 25 Malta Todarodes sagittatus European flying squid 37 4 4 4 6 2 2 0 1 10533 5 4 Malta Loligo spp Common squids nei 37 ------2 2 2 Malta Octopodidae Octopuses, etc. nei 37 5 1 0 1 1 1 0 0 00222 2 3 Morocco Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 7 9 6 10 4 2 3 3 4 6 11 11 9 11 9 Morocco Loliginidae, Ommastrephidae Various squids nei 37 141 273 81 97 129 153 205 215 139 265 178 162 132 157 179 Morocco Octopodidae Octopuses, etc. nei 37 13 11 28 63 17 37 40 25 122 32 10 2 2 3 6 Slovenia Sepia officinalis Common cuttlefish 37 492 205 80 98 8 72 78 71 25 69 36 98 68 115 112 Slovenia Loligo spp Common squids nei 37 ...... 12 21 4 10 6 5 18 18 11 Slovenia Octopus vulgaris Common octopus 37 ...... 4 4 42243 2 3 Spain Sepia officinalis Common cuttlefish 37 ...... 11 25 6 34 2 7 - - - Spain Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 ------499 440 483 985 - Spain Todarodes sagittatus European flying squid 37 1 239 1 033 919 1 354 1,300 F 1,200 F 1,100 F 1,000 F 900 F 800 F 746 780 834 446 1 311 Spain Loligo spp Common squids nei 37 51 ------140 149 119 144 338 Spain Loliginidae, Ommastrephidae Various squids nei 37 1 099 1 245 1 035 1 006 900 F 900 F 800 F 800 F 700 F 700 F 505 462 405 1 002 691 Spain Octopodidae Octopuses, etc. nei 37 ------592 547 747 673 791 Spain Cephalopoda Cephalopods nei 37 6 770 5 627 6 125 5 573 5,600 F 5,600 F 5,700 F 5,700 F 5,700 F 5,700 F 5 594 4 898 5 134 5 565 7 889 Tunisia Sepia officinalis Common cuttlefish 37 ...... 490 362 ------20 90 Tunisia Loligo spp Common squids nei 37 4,187 F 6,422 F 6 191 6 593 5 909 5 879 7 053 6 315 5 121 3 517 4 340 6 479 4 935 6 622 6 002 Tunisia Eledone spp Horned and musky octopuses 37 152 F 233 F 252 185 159 157 201 214 277 240 230 253 288 348 310 Tunisia Octopus vulgaris Common octopus 37 ...... 668 252 392 369 Tunisia Cephalopoda Cephalopods nei 37 5,427 F 8,325 F 12 295 9 135 6 417 6 337 7 544 3 350 1 841 1 868 3 460 5 681 3 129 2 349 2 103 Turkey Sepia officinalis Common cuttlefish 37 ...... Turkey Loligo spp Common squids nei 37 304 430 4 600 5 416 4 363 349 611 526 717 933 644 900 750 537 550 Turkey Octopus vulgaris Common octopus 37 408 405 3 398 4 123 3 232 267 557 397 579 331 364 420 500 360 400 Yugoslavia SFR Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 274 460 1 732 2 133 482 336 579 472 659 602 802 1 000 1 450 510 680 Yugoslavia SFR Loligo spp Common squids nei 37 154 190 235 214 296 169 F ------Yugoslavia SFR Octopus vulgaris Common octopus 37 228 197 205 269 374 214 F ------Yugoslavia, Fed. Rep. Sepiidae, Sepiolidae Cuttlefish,bobtail squids nei 37 154 159 179 180 211 121 F ------of Yugoslavia, Fed. Rep. Loligo spp Common squids nei 37 ...... 2 6 5 9 10 9 10 10 10 of Yugoslavia, Fed. Rep. Octopus vulgaris Common octopus 37 ...... 4 10 7 13 12 13 13 12 13 of

TOTAL 68 403 67 713 83 346 76 945 72 471 67 831 71 713 65 447 68 676 62 545 59 266 60 139 54 281 50 692 53 496

37 O:\scicom\LRC\WGCEPH\REPORTS\2002\WGCEPH02.doc 37 38

Annex 2 (Cont’d) FAO table 3: 1970-2000 ICES capture statistics for all cephalopods in area 27 Country Source 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 Belgium ICES 0 0 0 155 90 206 97 278 86 37 22 58 116 308 238 Channel Is ICES 0 0 0 0 0 1 1 1 2 9 2 2 0 3 5 Denmark ICES 0 0 0 0 0 0 0 0 11 9 17 21 23 9 5 Faeroe Islands ICES 0 0 0 0 0 0 0 0 11 49 504 2 132 249 200 1 147 France ICES 0 0 0 11 500 7 319 10 693 8 162 10 638 7 448 2 502 4 942 5 841 8 685 14 988 10 211 Germany ICES 0 0000000 0004520 Iceland ICES 0 0 0 0 0 0 0 0 0 436 16 7 13 4 1 634 Ireland ICES 0 0 0 0 0 0 7 170 70 51 100 253 396 211 264 Isle of Man ICES 0 0 0 0 0 0 0 0 0 1 3 9 16 4 6 Italy 0 0000000 0000000 Japan 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Korea, Republic of 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Lithuania 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Netherlands ICES 0 0 0 31 49 36 75 84 7 5 12 25 39 366 0 Norway ICES 0 0 0 0 0 0 0 260 345 1 667 2 978 9 780 18 385 18 024 7 802 Portugal ICES 0 0 0 0 4 757 6 026 5 886 5 882 8 672 4 662 7 733 9 031 5 778 5 635 5 359 Spain ICES 0 0 0 15 731 12 783 12 997 9 070 8 516 10 124 5 370 10 563 7 391 6 054 7 181 8 372 Sweden ICES 0 0000000 0000020 Un. Sov. Soc. Rep. ICES 0 0 0 0 0 40 0 0 0 0 0 2 100 342 53 0 UK ICES 0 0 0 959 635 1070 1489 1434 675 353 399 720 1191 1465 2155 TOTAL 0 0 0 28 376 25 633 31 069 24 787 27 263 27 451 15 151 27 291 37 374 41 292 48 455 37 198

Country Source 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 Belgium ICES 242 270 186 265 392 546 170 280 336 387 578 437 216 326 287 Channel Is ICES 4 23061515 22313162434 Denmark ICES 5 02218322254 5122171627 Faeroe Islands ICES 32 01100000 010023223 France ICES 10 668 10 140 9 176 16 405 20 222 26 897 17 659 16 982 21 688 18 868 21 356 18 825 16 881 17 610 0 Germany ICES 0 0000000 231134116 Iceland ICES 2 0000000 00113543 Ireland ICES 274 730 167 196 432 174 149 261 367 283 1 363 494 448 613 292 Isle of Man ICES 5 3062112715 15673222 Italy 0 0000000 0000000 Japan 0 0000000 0000000 Korea, Republic of 0 0000000 0000000 Lithuania 0 0000000 0000000 Netherlands ICES 0 0000000 0000000 Norway ICES 13 819 88 3 937 1 1825000 00352019020 Portugal ICES 5 826 6 383 13 065 11 673 13 962 10 646 11 112 13 277 9 307 9 220 12 090 14 075 12 118 9 678 11 126 Spain ICES 5 940 5 705 15 233 10 140 9 966 11 078 9 510 15 126 12 742 10 570 12 400 11 385 15 703 15 422 15 163 Sweden ICES 0 0000113 0000111 Un. Sov. Soc. Rep. ICES 0 0000000 0000000 UK ICES 1388 2087 1663 2520 4 705 6 083 2 513 3 606 4 493 4 619 6 867 8 099 5 374 6 203 5 138 TOTAL 38 205 25 408 43 443 42 389 49 729 55 484 41 144 49 609 48 957 43 960 55 040 53 339 50 977 49 944 32 102

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Annex 2 (Cont’d) FAO table 4: 1970-2000 FAO capture statistics for all cephalopods in area 27 Country Source 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 Belgium FAO 200 100 100 100 90 206 97 278 86 37 22 58 116 308 238 241 Channel Islands FAO 0 0000111 1111033 4 Denmark FAO 0 0000000 11915202396 5 Faeroe Islands FAO 0 0000000 0000000 0 France FAO 7 900 11 500 7 400 11 500 7 319 10 693 8 162 10 639 7 447 2 502 4 942 5 841 8 685 14 987 10 211 10 668 Germany FAO 0 0000000 0004520 1 Iceland FAO 0 0000000 04361671341 634 2 Ireland FAO 0 000007166 70 51 100 253 396 211 264 274 Isle of Man FAO 0 0000000 01381646 5 Italy FAO 0 000000208 0 485 1 870 0 575 0 0 0 Japan FAO 0 0010066010 0000000 0 Korea, Republic of FAO 0 0000000 0000000 0 Lithuania FAO 0 0000000 0000000 0 Netherlands FAO 100 0 0 0 49 36 75 84 7 5 12 25 39 366 0 0 Norway FAO 0 40000000260 346 1 668 2 977 9 780 18 385 18 025 7 803 13 819 Portugal FAO 2 900 4 200 5 800 8 900 4 757 6 026 5 886 5 882 8 672 4 662 7 733 9 031 5 778 5 635 5 359 5 826 Spain FAO 29 000 29 000 13 700 15 700 12 755 12 996 9 070 8 516 10 123 5 370 10 563 7 391 6 054 7 181 8 372 5 940 Sweden FAO 0 0000000 0000020 0 Un. Sov. Soc. Rep. FAO 0 00004000 0002 100342530 0 United Kingdom FAO 1 400 1 700 700 1 000 468 1 071 1 489 1 434 676 354 400 721 1 191 1 464 2 147 1 389 TOTAL 41500 46900 27700 37300 25504 31069 24788 27468 27439 15581 28654 35240 41618 48254 36043 38174

Country Area 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 Belgium FAO 271 187 265 391 544 168 279 336 387 574 435 216 326 286 566 Channel Islands FAO 2 3 5 6 6 6 5 5 4 4 13 16 20 33 35 Denmark FAO 0 1 2 18 32 22 53 5 2 3 2 17 16 26 20 Faeroe Islands FAO 0 0 0 0 0 0 0 0 1 0 0 2 32 23 0 France FAO 10 140 9 176 16 405 20 228 26 909 17 667 16 982 21 701 18 880 21 356 18 846 16 881 17 610 20 961 24 816 Germany FAO 0 0 0 0 0 0 0 3 3 11 2 4 11 6 5 Iceland FAO 0 0000000 01135431 Ireland FAO 730 167 198 432 184 149 261 368 283 323 494 449 613 292 322 Isle of Man FAO 3 1 6 21 12 7 15 15 6 7 3 2 2 2 2 Italy FAO 0 0000000 0000000 Japan FAO 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Korea, Republic of FAO 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 Lithuania FAO 0 0 0 0 0 0 241 0 0 0 0 0 0 0 0 Netherlands FAO 0 0 0 0 0 0 0 0 0 0 0 0 0 0 874 Norway FAO 89 3 936 1 183 5 0 0 0 0 0 352 0 190 2 0 0 Portugal FAO 6 383 13 065 11 673 13 962 10 646 11 112 13 288 9 307 9 220 12 090 14 075 12 117 9 678 11 128 12 056 Spain FAO 5 706 15 235 10 142 9 966 11 078 9 510 15 126 12 742 10 570 12 400 11 385 15 703 15 422 15 163 13 487 Sweden FAO 0 0001134 0001110 Un. Sov. Soc. Rep. FAO 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 United Kingdom FAO 2 086 1 662 2 487 4 711 6 090 2 513 3 606 4 544 4 622 6 902 8 113 5 380 6 211 5 142 5 157 TOTAL 25410 43433 42366 49740 55502 41155 49860 49030 43978 54033 53371 50983 49948 53066 57341 39 O:\scicom\LRC\WGCEPH\REPORTS\2002\WGCEPH02.doc 39

40

Annex 2 (Cont’d) FAO table 5: 1970-2000 differences between FAO and ICES capture statistics for all cephalopods in area 27 (difference = table 4 – table 3) Country Source 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 Belgium Difference 200 100 100 -550000 0000000 -1 Channel Islands Difference 0 0000000 -1-8-1-100-2 0 Denmark Difference 0 0000000 00-2-1001 0 Faeroe Islands Difference 0 00 0000 -11-49-504-2132 -249 -200 -1147 -32 France Difference 7900 11500 740000001 -10000-10 0 Germany Difference 0 0000000 0000000 1 Iceland Difference 0 0000000 0000000 0 Ireland Difference 0 000000-4 0000000 0 Isle of Man Difference 0 0000000 000-1000 0 Italy Difference 0 000000208 0 485 1870 0 575 0 0 0 Japan Difference 0 0010066010 0000000 0 Korea, Republic of Difference 0 0000000 0000000 0 Lithuania Difference 0 0000000 0000000 0 Netherlands Difference 100 0 0 -310000 0000000 0 Norway Difference 0 400000000 11-10011 0 Portugal Difference 2900 4200 5800 89000000 0000000 0 Spain Difference 29000 29000 13700 -31 -28 -1 0 0 -1000000 0 Sweden Difference 0 0000000 0000000 0 Un. Sov. Soc. Rep. Difference 0 0000000 0000000 0 United Kingdom Difference 1400 1700 700 41 -167100 11110-1-8 1 TOT Diff. 41500 46900 27700 8924 -129 0 1 205 -12 430 1363 -2134 326 -201 -1155 -31

Country Source 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 Belgium Difference 1 1 0 -1 -2 -2 -1 0 0 -4 -2 0 0 -1 566 Channel Islands Difference 0 0 5 0 -9 5 0 3 2 1 0 0 -4 -1 35 Denmark Difference 0 -1 0 0 0 0 -1 0 1 1 0 0 0 -1 20 Faeroe Islands Difference 0 -11 0 0 0 0 0 0 0 0 0 0 0 0 0 France Difference 0 0 0 6 12 8 0 13 12 0 21 0 0 20961 24816 Germany Difference 0 0000001 00-10005 Iceland Difference 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 Ireland Difference 0 0 2 0 10 0 0 1 0 -1040 0 1 0 0 322 Isle of Man Difference 0 1 0 0 0 0 0 0 0 0 0 0 0 0 2 Italy Difference 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Japan Difference 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Korea, Republic of Difference 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 Lithuania Difference 0 0 0 0 0 0 241 0 0 0 0 0 0 0 0 Netherlands Difference 0 0 0 0 0 0 0 0 0 0 0 0 0 0 874 Norway Difference 1 -1 1 0 0 0 0 0 0 0 0 0 0 0 0 Portugal Difference 0 0 0 0 0 0 11 0 0 0 0 -1 0 2 12056 Spain Difference 1 2 2 0 0 0 0 0 0 0 0 0 0 0 13487 Sweden Difference 0 0 0 0 0 0 0 4 0 0 0 0 0 0 0 Un. Sov. Soc. Rep. Difference 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 United Kingdom Difference -1 -1 -33 6 7 0 0 51 3 35 14 6 8 4 5157 TOT Diff. 2 -10 -23 11 18 11 251 73 18 -1007 32 6 4 20964 57341 40 O:\scicom\LRC\WGCEPH\REPORTS\2002\WGCEPH02.doc

FAO capture statistics for cephalopods FAO capture statistics for cephalopods in area 27 (Northeast Atlantic) in area 37 (Mediterranean and Black Sea) 90 000 90 000

80 000 80 000

70 000 70 000

60 000 60 000

50 000 50 000

40 000 40 000

30 000 30 000

20 000 20 000

10 000 10 000

0 0 1970 1972 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 1970 1972

FAO capture statistics by group of cephalopods in area 27 FAO capture statistics by group of cephalopods in area 37 (Northeast Atlantic) (Mediterranean and Black Sea) Cuttlefishes Squids Octopuses Cuttlefishes Squids Octopuses Cephalopods nei

40 000 40 000

35 000 35 000

30 000 30 000

25 000 25 000

20 000 20 000

15 000 15 000

10 000 10 000

5 000 5 000

0 0 0 2 4 6 8 0 2 4 6 8 0 2 4 6 8 0 970 972 974 976 978 980 982 984 986 988 990 992 994 996 998 000 197 197 197 197 197 198 198 198 198 198 199 199 199 199 199 200 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2

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ANNEX 3

Visualisation of ICES WGCEPH Fishery Statistics.

Maps of annual landings per Cephalopod resource and per area

15— 10— 5— 0— 5— 10—

All Cephalopods

15 000T 10 000T 5 000T 1 000T

1995 1996 1997 1998 1999 2000 2001

• Map n° 1: Annual Landings of all cephalopod species in the period 1995-2001 (2001 is provisional)

42

15— 10— 5— 0— 5— 10—

Cuttlefish

15 000T 10 000T 5 000T 1 000T

1995 1996 1997 1998 1999 2000 2001

• Map n° 2: Annual Landings of cuttlefish (Sepiidae) in the period 1995-2001 (2001 is provisional)

43

15— 10— 5— 0— 5— 10—

Loliginid Squid

4 000T 2 000T 1 000T 500 T

1995 1996 1997 1998 1999 2000 2001

• Map n° 3: Annual Landings of Common Squid (Loliginidae) in the period 1995-2001 (2001 is provisional)

44

15— 10— 5— 0— 5— 10—

Short Finned Squid

4 000T 2 000T 1 000T 500 T

1995 1996 1997 1998 1999 2000 2001

• Map n° 4: Annual Landings of Short Finned Squid (Ommastraephidae) in the period 1995-2001 (2001 is provisional)

45

15— 10— 5— 0— 5— 10—

Octopods

15 000T 10 000T 5 000T 1 000 T

1995 1996 1997 1998 1999 2000 2001

• • Map n° 5: Annual Landings of Octopods (Eledone spp. and Octopus vulgaris) in the period 1995-2001 (2001 is provisional)

46

15— 10— 5— 0— 5— 10— N N N 2A 2B# 5A N

12N A 5B

N

4A

N

6AB N 4B

N N 7BC 7A N 7F N 4C

N

7GK N 7DE

Year 2000 All cephalopods

N Belgium 8AE Denmark Faroe France Germany Iceland Ireland N 100 T Netherlands Norway N 1 000 T Portugal N Sweden N 10 000 T Spain UK 9AB

Map n° 6: Year 2000 cephalopod landings (all species) per country and per area

47

15— 10— 5— 0— 5— 10— N N N 2A 2B# 5A N

12N A 5B

N

4A

N

6AB N 4B

N N 7BC 7A N 7F N 4C

N

7GK N 7DE

Year 2000 Cuttlefish

N Belgium 8AE Denmark Faroe France Germany Iceland Ireland N 100 T Netherlands Norway N 1 000 T Portugal N Sweden

N Spain 10 000 T UK 9AB

Map n° 7: Year 2000 cuttlefish landings per country and par area

48

15— 10— 5— 0— 5— 10— N N N 2A 2B# 5A N

12N A 5B

N

4A

N

6AB N 4B

N N 7BC 7A N 7F N 4C

N

7GK N 7DE

Year 2000 Loliginid squid

N Belgium 8AE Denmark Faroe France Germany Iceland Ireland N 100 T Netherlands Norway N 1 000 T Portugal N Sweden N 10 000 T Spain UK 9AB

Map n° 8: Year 2000 Long Finned Squid landings per country and par area

49

15— 10— 5— 0— 5— 10— N N N 2A 2B# 5A N

12N A 5B

N

4A

N

6AB N 4B

N N 7BC 7A N 7F N 4C

N

7GK N 7DE

Year 2000 Short Finned Squid

N Belgium 8AE Denmark Faroe France Germany Iceland Ireland N 100 T Netherlands Norway N 1 000 T Portugal N Sweden

N Spain 10 000 T UK 9AB

Map n° 9: Year 2000 Short Finned Squid landings per country and par area

50

15— 10— 5— 0— 5— 10— N N N 2A 2B# 5A N

12N A 5B

N

4A

N

6AB N 4B

N N 7BC 7A N 7F N 4C

N

7GK N 7DE

Year 2000 Octopuses

N Belgium 8AE Denmark Faroe France Germany Iceland Ireland N 100 T Netherlands Norway N 1 000 T Portugal N Sweden

N Spain 10 000 T UK 9AB

Map n° 10: Year 2000 Octopods landings per country and par area

51

ANNEX 4(I)

CEPHALOPOD BIBLIOGRAPHY (2001-02)

Abollo, E., Gestal, C. & Pascual, S., 2001. Anisakis infestation in marine fish and cephalopods from Galician waters: an updated perspective. Parasitology Research, 87(6): 492-499.

Agin, V., Chichery, R. & Chichery, M.P., 2001. Effects of learning on cytochrome oxidase activity in cuttlefish brain. Neuroreport, 12(1): 113-116.

Allcock, A.L., Piatkowski, U., Rodhouse, P.G.K. & Thorpe, J.P., 2001. A study on octopodids from the eastern Weddell Sea, Antarctica. Polar Biology, 24: 832-838.

Anderson R.C. & Wood, J.B., 2001. Enrichment of giant pacific octopuses: Happy as a clam? Journal of Applied Animal Welfare Science, 4(2): 157-168.

Baron, P.J., 2001. First description and survey of the egg masses of Loligo gahi (d'Orbigny, 1835) and Loligo sanpaulensis (Brakoniecki, 1984) from coastal waters of Patagonia. Journal of Shellfish Research, 20(1): 289-295.

Bartol, I.K., 2001. Role of aerobic and anaerobic circular mantle muscle fibers in swimming squid: Electromyography. Biological Bulletin, 200(1): 59-66.

Bartol, I.K., Mann, R. & Patterson, M.R., 2001. Aerobic respiratory costs of swimming in the negatively buoyant brief squid Lolliguncula brevis. Journal of Experimental Biology, 204(21): 3639-3653.

Bello, G., 2001. Dimorphic growth in male and female cuttlefish Sepia orbignyana (Cephalopoda: Sepiidae) from the Adriatic Sea. Helgoland Marine Research, 55(2): 124-127.

Bellido, J.M., Pierce, G.J. & Wang, J., 2001. Application of generalised additive models to reveal spatial relationships between environmental variables and squid abundance in Scottish waters. Fisheries Research 52, 23-40.

Benoit-Bird, K.J. & Au, W.W.L., 2001. Target strength measurements of Hawaiian mesopelagic boundary community animals. Journal of the Acoustical Society of America, 110(2): 812-819.

Berger, D.K. & Butler, M.J., 2001. Octopuses influence den selection by juvenile Caribbean spiny lobster. Marine and Freshwater Research, 52(8): 1049-1053.

Bettencourt, V. & Guerra, A., 2001. Age studies based on daily growth increments in statoliths and growth lamellae in cuttlebone of cultured Sepia officinalis. Marine Biology, 139(2): 327-334.

Bjørke, H., 2001. Predators of the squid Gonatus fabricii (Lichtenstein) in the Norwegian Sea. Fisheries Research, 52: 113-120.

Brown, E.G., Pierce, G.J., Hislop, J.R.G. & Santos, M.B., 2001. Interannual variation in the summer diets of harbour seals Phoca vitulina at Mousa, Shetland (UK). Journal of the Marine Biological Association of the United Kingdom, 81(2): 325-337.

Buresch, K.M., Hanlon, R.T., Maxwell, M.R. & Ring, S., 2001. Microsatellite DNA markers indicate a high frequency of multiple paternity within individual field-collected egg capsules of the squid Loligo pealeii. Marine Ecology Progress Series, 210: 161-165.

Bustamante, P., Cosson R.P., Gallien I., Caurant, F. & Miramand, P., 2002. Cadmium detoxification processes in the digestive gland of cephalopods in relation to accumulated cadmium concentrations. Marine Environmental Research, 53: 227-241.

Cabrera, J.L. & Defeo, O., 2001. Daily bioeconomic analysis in a multispecific artisanal fishery in Yucatan, Mexico. Aquatic Living Resources, 14(1): 19-28.

52

Chiao, C.C. & Hanlon, R.T., 2001. Cuttlefish cue visually on area - Not shape or aspect ratio - of light objects in the substrate to produce disruptive body patterns for camouflage. Biological Bulletin, 201(2): 269-270.

Chiao, C.C. & Hanlon, R.T., 2001. Cuttlefish camouflage: Visual perception of size, contrast and number of white squares on artificial checkerboard substrata initiates disruptive coloration. Journal of Experimental Biology, 204(12): 2119-2125.

Childerhouse, S., Dix, B. & Gales, N., 2001. Diet of new Zealand sea lions (Phocarctos hookeri) at the Auckland islands. Wildlife Research, 28(3): 291-298.

Clancy, C.M.R. & Simon, J.D., 2001. Ultrastructural organization of eumelanin from Sepia officinalis measured by atomic force microscopy. Biochemistry, 40(44): 13353-13360.

Clark, Y.M., Miller, A.M., Mendoza, J.E., Ramirez, T., Rivas, F., Rederford, V.R., Ochoa, G.H. & Robles, L.J., 2001. Myosin III localization and expression in octopus retinas. Molecular Biology of the Cell, 12: 1615.

Clay, J.R. & Shrier, A., 2001. Action potentials occur spontaneously in squid giant axons with moderately alkaline intracellular pH. Biological Bulletin, Marine Biological Laboratory, Woods Hole, 201(2): 186-192.

Collins, M.A., O'Dea, M. & Henriques, C., 2001. A large Cirroteuthis magna (Cephalopoda: Cirroctopoda) caught on the Cape Verde Terrace (North Atlantic). Journal of the Marine Biological Association of the UK, 81(2): 357-358.

Collins, M.A., Yau, C., Allcock, L. & Thurston, M.H., 2001. Distribution of deep-water benthic and bentho-pelagic cephalopods from the Northeast Atlantic. Journal of the Marine Biological Association of the UK, 81(1): 105-117.

Cronin, T.W. & Shashar, N., 2001. The linearly polarized light field in clear, tropical marine waters: Spatial and temporal variation of light intensity, degree of polarization and e-vector angle. Journal of Experimental Biology, 204(14): 2461-2467.

Dai, T., Hong, M., Zheng, G. & Zheng, Y., 2001. Experiment of fishing gear and methods of light-pelagic trawl for squid. Journal of Shanghai Fisheries University, 10(1): 26-30.

Daly, H.I., Pierce, G.J., Santos, M.B., Royer, J., Cho, S.K., Stowasser, G., Robin, J.P. & Henderson, S.M., 2001. Cephalopod consumption by trawl caught fish in Scottish and English Channel waters. Fisheries Research, 52(1-2): 51-64.

Dauphin, Y., 2001. Characteristics of the soluble organic matrices of the aragonitic shells of the three modern cephalopod genera. Neues Jahrbuch Fur Geologie Und Palaontologie-Monatshefte, 2: 103-123.

Davies, A., Gowen, B.E., Krebs, A.M., Schertler, G.F.X. & Saibil, H.R., 2001. Three-dimensional structure of an invertebrate rhodopsin and basis for ordered alignment in the photoreceptor membrane. Journal of Molecular Biology, 314(3): 455-463.

Denis, V. & Robin, J.P., 2001. Present status of the French Atlantic fishery for cuttlefish (Sepia officinalis). Fisheries Research, 52(1-2): 11-22.

Di Beneditto, A.P.M., Ramos, R.M.A., Siciliano, S., dos Santos, R.A., Bastos, G. & Fagundes-Netto, E., 2001. Stomach contents of delphinids from Rio de Janeiro, southeastern Brazil. Aquatic Mammals, 27(1): 24-28.

Di Cosmo, A., Di Cristo, C. & Paolucci, M., 2002. A estradiol-17 beta receptor in the reproductive system of the female of Octopus vulgaris: Characterisation and immunolocalization. Molecular Reproduction and Development, 61(3): 367-375.

Di Cosmo, A., Di Cristo, C. & Paolucci, M., 2001. Sex steroid hormone fluctuations and morphological changes of the reproductive system of the female of Octopus vulgaris throughout the annual cycle. Journal of Experimental Zoology, 289(1): 33-47.

53

Di Cristo, C., Paolucci, M., Iglesias, J., Sanchez, J. & Di Cosmo, A., 2002. Presence of two neuropeptides in the fusiform ganglion and reproductive ducts of Octopus vulgaris: FMRFamide and gonadotropin-releasing hormone (GnRH). Journal of Experimental Zoology, 292(3): 267-276.

Diatta, Y., Clotilde-Ba, F. & Capape, C., 2001. Role trophique du poulpe commun, Octopus vulgaris, chez les elasmobranches de la cote du Senegal (Atlantique oriental tropical) comparaison avec les especes des cotes Tunisiennes (Mediterranee centrale). Acta Adriatica - Institute of Oceanogarphy and Fisheries Split Croatia, 42(1): 77-88.

Dickel, L., Chichery, M.P. & Chichery, R., 2001. Increase of learning abilities and maturation of the vertical lobe complex during postembryonic development in the cuttlefish, Sepia. Developmental Psychobiology, 39(2): 92-98.

Ding, T., & Song, H., 2001. Estimation on the cephalopod biomass in the sea area of middle-northern East China Sea. Journal of fisheries of China, 25(3): 215-221.

Domingues, P.M., Kingston, T., Sykes, A. & Andrade, J.P., 2001. Growth of young cuttlefish, Sepia officinalis (Linnaeus 1758) at the upper end of the biological distribution temperature range. Aquaculture Research, 32(11): 923-930.

Donovan, S.K., Portell, R.W. & Pickerill, R.K., 2001. A shell of the cephalopod Sepia Linne from the coast of Carriacou, Grenadines, Lesser Antilles. Caribbean Journal of Science, 37(1-2): 125-127. dos Santos, R.A. & Haimovici, M., 2001. Cephalopods in the diet of marine mammals stranded or incidentally caught along southeastern and southern Brazil (21- 34 degrees S). Fisheries Research, 52(1-2): 99-112.

Ferrara, F., Fabietti, F., Delise, M., Bocca, A.P. & Funari, E., 2001. Alkylphenolic compounds in edible molluscs of the Adriatic Sea (Italy). Environmental Science & Technology, 35(15): 3109-3112.

Fonseca, V.S.S., Petry, M.V. & Jost, A.H., 2001. Diet of the Magellanic penguin on the coast of Rio Grande do Sul, Brazil. Waterbirds, 24(2): 290-293.

Forsythe, J.W., Walsh, L.S., Turk, P.E. & Lee, P.G., 2001. Impact of temperature on juvenile growth and age at first egg-laying of the Pacific reef squid Sepioteuthis lessoniana reared in captivity. Marine Biology, 138: 103-112.

Gestal, C., Serra, C., Guerra, A., & Pascual, S., 2002. Scratching the sprocyst surface: characterisation of European Aggregata species by Atomic Force Microscope. Parasitology Research, 88: 242-246.

Gestal, C., Guerra, A., Pascual, S., & Azevedo, C., 2002. The life cycle of Aggregata eberthi and A. octopiana (Aplicomplexa, Aggregatidae) from NE Atlantic. European Journal of Protistology, 37: 427-435.

Guerra, A., Perez-Losada, M., Rocha, F. & Sanjuan, A., 2001. Species differentiation of Sepia officinalis and Sepia hierredda (Cephalopoda: Sepiidae) based on morphological and allozyme analyses. Journal of the Marine Biological Association of the United Kingdom, 81(2): 271-281.

Guerra, A., Rocha, F., Gonzalez, A.F. & Buckle, L.F., 2001. Embryonic stages of the Patagonian squid Loligo gahi (Mollusca: Cephalopoda). Veliger, 44(2): 109-115.

Hatfield, E.M.C., Hanlon, R.T., Forsythe, J.W. & Grist, E.P.M., 2001. Laboratory testing of a growth hypothesis for juvenile squid Loligo pealeii (Cephalopoda: Loliginidae). Canadian journal of fisheries and aquatic sciences, 58(5): 845-857.

Hedd, A. & Gales, R., 2001. The diet of shy albatrosses (Thalassarche cauta) at Albatross Island, Tasmania. Journal of Zoology, 253(1): 69-90.

Hernandez-Lopez, J.L., Castro-Hernendez, J.J. & Hernandez-Garcia, V., 2001. Age determined from the daily deposition of concentric rings on common octopus (Octopus vulgaris) beaks. Fishery Bulletin, 99(4): 679-684.

Hurtado, J.L., Montero, P., Borderias, J. & Solas, M.T., 2001. High-pressure/temperature treatment effect on the characteristics of octopus (Octopus vulgaris) arm muscle. European Food Research and Technology, 213(1): 22-29.

54

Hurtado, J.L., Montero, P. & Borderias, R., 2001. Chilled storage of pressurized octopus (Octopus vulgaris) muscle. Journal of Food Science, 66(3): 400-406.

Hyrenbach, K.D., 2001. Albatross response to survey vessels: implications for studies of the distribution, abundance, and prey consumption of seabird populations. Marine Ecology Progress Series, 212: 283-295.

Ishikawa, M., Suzuki, F., Ishida, M., Nagashima, Y. & Shiomi, K., 2001. Identification of tropomyosin as a major allergen in the octopus Octopus vulgaris and elucidation of its IgE-binding epitopes. Fisheries Science, 67(5): 934- 942.

Jabeur, C., Gobert, B. & Missaoui, H., 2000. Typology of the small-scale fishing fleet in the gulf of Gabes (Tunisia). Aquatic Living Resources, 13(6): 421-428.

Jackson, G.D. & Moltschaniwskyj, N.A., 2001. The influence of ration level on growth and statolith increment width of the tropical squid Sepioteuthis lessoniana (Cephalopoda: Loliginidae): an experimental approach. Marine Biology, 138: 819-825.

Jahnke, L.L., Eder, W., Huber, R., Hope, J.M., Hinrichs, K.U., Hayes, J.M., Des Marais, D.J., Cady, S.L. & Summons, R.E., 2001. Signature lipids and stable carbon isotope analyses of octopus spring hyperthermophilic communities compared with those of Aquificales representatives. Applied and Environmental Microbiology, 67(11): 5179-5189.

Jones, G.B., Mercurio, P. & Olivier, F., 2000. Zinc in fish, crabs, oysters, and mangrove flora and fauna from Cleveland Bay. Marine Pollution Bulletin, 41(7-12): 345-352.

Kaehler, S. & Pakhomov, E.A., 2001. Effects of storage and preservation on the delta C-13 and delta N-15 signatures of selected marine organisms. Marine Ecology-Progress Series, 219: 299-304.

Karnik, N.S. & Chakraborty, S.Kr., 2001. Length-weight relationship and morphometric study on the squid Loligo duvauceli (d'Orbigny) (Mollusca / Cephalopoda) off Mumbai (Bombay) waters, west coast of India. Indian Journal of Marine Sciences, 30(4): 261-263.

Kasugai, T., 2001. Feeding behaviour of the Japanese pygmy cuttlefish Idiosepius paradoxus (Cephalopoda: Idiosepiidae) in captivity: evidence for external digestion? Journal of the Marine Biological Association of the United Kingdom, 81(6): 979-981.

Kawabata, A., 2001. Measurement of the target strength of live squid, Todarodes pacificus Steenstrup, in controlled body tilt angle. Bulletin of the Tohoku National Fisheries Research Institute. Shiogama, 64: 61-67.

Kawamura, G., Nobutoki, K., Anraku, K., Tanaka, Y. & Okamoto, M., 2001. Colour discrimination conditioning in two octopus Octopus aegina and O. vulgaris. Nippon Suisan Gakkaishi, 67(1): 35-39.

Kishimura, H., Saeki, H. & Hayashi, K., 2001. Isolation and characteristics of trypsin inhibitor from the hepatopancreas of a squid (Todarodes pacificus). Comparative Biochemistry and Physiology, B, 130(1): 117-123.

Klug, C., 2001. Functional morphology and taphonomy of nautiloid beaks from the Middle Triassic of southern Germany. Acta Palaeontologica Polonica, 46(1): 43-68.

Koueta, N., Boucaud-Camou, E. & Noel, B., 2002. Effect of enriched natural diet on survival and growth of juvenile cuttlefish Sepia officinalis L. Aquaculture, 203(3-4): 293-310.

Koueta, N. & Boucaud-Camou, E., 2001. Basic growth relations in experimental rearing of early juvenile cuttlefish Sepia officinalis L. (Mollusca: Cephalopoda). Journal of Experimental Marine Biology and Ecology, 265(1): 75-87.

Kubo, T. & Saeki, H., 2001. Role of sorbitol in manufacturing dried seafood from heated squid meat. Fisheries science. Tokyo, 67(3): 524-529.

Kulicki, C., Tanabe, K., Landman, N.H. & Mapes, R.H., 2001. Dorsal shell wall in ammonoids. Acta Palaeontologica Polonica, 46(1): 23-42.

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Landman, N.H., Jones, D.S. & Davis, R.A., 2001. Hatching depth of Nautilus pompilius in Fiji. Veliger, 44(4): 333-339.

Lewy, Z., 2002. The function of the ammonite fluted septal margins. Journal of Paleontology, 76(1): 63-69.

Li, X., Wang, Z. & Jiang, X., 2001. The purification of melanin extracted from squid ink. Journal of Shanghai Fisheries University, 10(3): 252-256.

Lieb, B., Altenhein, B., Markl, J., Vincent, A., Van Olden, E., Van Holde, K.E. & Miller, K.I., 2001. Structures of two molluscan hemocyanin genes: Significance for gene evolution. Proceedings of the National Academy of Sciences of the United States of America, 98(8): 4546-4551.

Lipinski, M.R., 2001. Preliminary description of two new species of Cephalopods (Cephalopoda: Brachioteuthidae) from South Atlantic and Antarctic waters. Bulletin of the Sea Fisheries Institute, Gdynia, 152: 3-14.

Liu, H., Yang, H. & Zhang, S., 2001. Preliminary study on seawater salinity and temperature distributions and their relationship with the squid fishing grounds in 175 degree W-170 degree W area of North Pacific. Journal of Shanghai Fisheries University, 10(3): 229-233.

Lordan, C., Collins, M.A., Key, L.N. & Browne E.D., 2001. The biology of the ommastrephid squid, Todarodes sagittatus, in the Northeast Atlantic. Journal of the Marine Biological Association of the United Kingdom, 81(2): 299-306.

Martinez, P., Bettencourt, V., Guerra, A. & Moltschaniwskyj, N., 2001. How temperature influences muscle and cuttlebone growth under food-stress condition on juvenile Sepia elliptica (Mollusca: Cephalopoda). Canadian Journal of Zoology, 78: 1855-1861.

Marvin, L.F., Zatlyny, C., Leprince, J., Vaudry, H. & Henry, J., 2001. Characterisation of a novel Sepia officinalis neuropeptide using MALDI-TOF MS and post-source analysis. Peptides, 22: 1391-1396.

Mathger, L.M. & Denton, E.J., 2001. Reflective properties of iridophores and fluorescent 'eyespots' in the loliginid squid Alloteuthis subulata and Loligo vulgaris. Journal of Experimental Biology, 204(12): 2103-2118.

Messenger, J.B., 2001. Cephalopod chromatophores: neurobiology and natural history. Biological Reviews, 76(4): 473- 528.

Miyazaki, T., Nakahara, M., Ishii, T., Aoki, K. & Watabe, T., 2001. Accumulation of cobalt in newly hatched octopus Octopus vulgaris. Fisheries Science, 67(1): 170-172.

Mori, J., Kubodera, T. & Baba, N., 2001. Squid in the diet of northern fur seals, Callorhinus ursinus, caught in the western and central North Pacific Ocean. Fisheries Research, 52(1-2): 91-98.

Mori, K., & Nakamura, Y., 2001. Migration patterns of the Pacific subpopulation of Japanese common squid Todarodes pacificus, estimated from tagging experiments. Bulletin of the Hokkaido National Fisheries Research Institute. Kushiro, 65: 21-43.

Mouat, B. Collins, M.A. & Pompert, J., 2001. Patterns in the diet of Illex argentinus (Cephalopoda: Ommastrephidae) from the Falkland Islands jigging fishery. Fisheries Research, 52(1-2): 41-50.

Nagai, T., Yamashita, E., Taniguchi, K., Kanamori, N. & Suzuki, N., 2001. Isolation and characterisation of collagen from the outer skin waste material of cuttlefish (Sepia lycidas). Food Chemistry, 72(4): 425-429.

Neethiselvan, N., 2001. A new species of cuttlefish Sepia ramani sp. nov. (Class: Cephalopoda) from Tuticorin Bay, southeast coast of India. Indian Journal of Marine Sciences, 30(3): 185-185.

Neethiselvan, N., Venkataramani, V.K. & Srikrishnadhas, B., 2001. Reproductive biology of the siboga squid Doryteuthis sibogae (Adam) from Thoothukkudi (Tuticorin) coast, southeast coast of India. Indian Journal of Marine Sciences, 30(4): 257-260.

56

Nesis, K.N., 2001. West-Arctic and East-Arctic distributional ranges of cephalopods. Sarsia, 86(1): 1-11.

Norman, M.D., Finn, J. & Tregenza, T., 2001. Dynamic mimicry in an Indo-Malayan octopus. Proceedings of the Royal Society of London Series B-Biological Sciences, 268(1478): 1755-1758.

Norman, M.D. & Finnn, J., 2001. Revision of the Octopus horridus species-group, including erection of a new subgenus and description of two member species from the Great Barrier Reef, Australia. Invertebrate Taxonomy, 15(1): 13-35.

Okamoto, M., Anraku, K., Kawamura, G. & Tanaka, Y., 2001. Selectivity of colour of shelter by Octopus vulgaris and O. aegina under different background colours. Nippon Suisan Gakkaishi, 67(4): 672-677.

Orr, A.J. & Harvey, J.T., 2001. Quantifying errors associated with using fecal samples to determine the diet of the California sea lion (Zalophus californianus). Canadian Journal of Zoology, 79(6): 1080-1087.

Pascual, S., & Guerra, A., 2001. Vexing question on fisheries research: the study of cephalopods and their parasites. Iberus, 19: 87-95.

Pascual, S., Gonzalez, A.F., Gestal, C., Abollo, E. & Guerra, A., 2001. Epidemiology of Pennella sp (Crustacea: Copepoda), in exploited Illex coindetti stock in the NE Atlantic. Scientia Marina, 65(4): 307-312.

Parveen, Z., Large, A., Grewal, N., Lata, N., Cancio, I., Cajaraville, M.P., Perry, C.J. & Connock, M.J., 2001. D- Aspartate oxidase and D-amino acid oxidase are localised in the peroxisomes of terrestrial gastropods. European Journal of Cell Biology, 80(10): 651-660.

Pecl., G., 2001. Flexible reproductive strategies in tropical and temperate Sepioteuthis squids. Marine Biology, 138: 93- 101.

Phillips, K.L., Jackson, G.D. & Nichols, P.D., 2001. Predation on myctophids by the squid Moroteuthis ingens around Macquarie and Heard Islands: stomach contents and fatty acid analyses. Marine Ecology Progress Series, 215: 179- 189.

Piatkowski, U., Heinemann, H. & Pütz, K., 2001. Cephalopod prey of king penguins (Aptenodytes patagonicus) breeding at Volunteer Beach, Falkland Islands, during austral winter 1996. Fisheries Research, 52(1-2): 79-90.

Piatkowski, U., Pierce, G.J. & Da Cunha, M.M., 2001. Impact of cephalopods in the food chain and their interaction with the environment - Preface. Fisheries Research, 52(1-2): 1-1.

Piatkowski, U., Pierce, G.J. & Da Cunha, M.M., 2001. Impact of cephalopods in the food chain and their interaction with the environment and fisheries: an overview. Fisheries Research, 52(1-2): 5-10.

Pierce, G.J., Wang, J.J., Zheng, X.H., Bellido, J.M., Boyle, P.R., Denis, V. & Robin, J.R., 2001. A cephalopod fishery GIS for the Northeast Atlantic: development and application. International Journal of Geographical Information Science, 15(8): 763-784.

Quast, M.J., Neumeister, H., Ezell, E.L. & Budelmann, B.U., 2001. MR microscopy of cobalt-labeled nerve cells and pathways in an invertebrate brain (Sepia officinalis, Cephalopoda). Magnetic Resonance in Medicine, 45(4): 575- 579.

Quetglas, A., Gonzalez, M., Carbonell, A. & Sanchez, P., 2001. Biology of the deep-sea octopus Bathypolypus sponsalis (Cephalopoda: Octopodidae) from the western Mediterranean Sea. Marine Biology, 138(4): 785-792.

Reid, A., 2001. A new cuttlefish, Sepia grahami, sp nov (Cephalopoda: Sepiidae) from Eastern Australia. Proceedings of the Linnean Society of New South Wales, 123: 159-172.

Robles, L.J., Clark, Y.M., Mendoza, J., Ramirez, T. & Rivas, F., 2001. Localization and expression of myosin III in light- and dark- adapted octopus retinas. Investigative Ophthalmology & Visual Science, 42(4): 1970.

57

Rocha, F., Guerra, A. & Gonzalez, A.F., 2001. A review of reproductive strategies in cephalopods. Biological Reviews, 76(3): 291-304.

Rodhouse, P.G., Elvidge, C.D. & Trathan, P.N., 2001. Remote sensing of the global light-fishing fleet: an analysis of interactions with oceanography, other fisheries and predators. Advances in Marine Biology, 39: 261-303.

Roper C.F.E. & Vecchione, M., 2001. Pickfordiateuthis bayeri, a new species of squid (Cephalopoda: Loliginidae) from the western North Atlantic Ocean discovered by submersible. Bulletin of the Biological Society of Washington, 10: 301-310.

Ruiz-Capillas, C., Moral, A., Morales, J. & Montero, P., 2002. Preservation of shelf life of pota and octopus in chilled storage under controlled atmospheres. Journal of Food Protection, 65(1): 140-145.

Ruth, P., Schmidtberg, H., Westermann, B. & Schipp, R., 2002. The sensory epithelium of the tentacles and the rhinophore of Nautilus pompilius L. (Cephalopoda, Nautiloidea). Journal of Morphology, 251(3): 239-255.

Saito, M. & Sugiyama, K., 2001. Major and c-series gangliosides in lenticular tissues: mammals to molluscs. Comparative Biochemistry and Physiology B-Biochemistry & Molecular Biology, 130(3): 313-321.

Saito, M., Kitamura, H. & Sugiyama, K., 2001. Occurrence of gangliosides in the common squid and pacific octopus among protostomia. Biochimica et Biophysica Acta-Biomembranes, 1511(2): 271-280.

Santos, M.B., Clarke, M.R. & Pierce, G.J., 2001. Assessing the importance of cephalopods in the diets of marine mammals and other top predators: problems and solutions. Fisheries Research, 52(1-2): 121-139.

Santos, M.B., Pierce, G.J., Gonzalez, A.F., Santos, F., Vazquez, M.A., Santos, M.A. & Collins, M.A., 2001. First records of Taningia danae (Cephalopoda: Octopoteuthidae) in Galician waters (Northwest Spain) and in Scottish waters (UK). Journal of the Marine Biological Association of the United Kingdom, 81(2): 355-356.

Santos, M.B., Pierce, G.J., Herman, J., Lopez, A., Guerra, A., Mente, E. & Clarke, M.R., 2001. Feeding ecology of Cuvier's beaked whale (Ziphius cavirostris): a review with new information on the diet of this species. Journal of the Marine Biological Association of the United Kingdom, 81(4): 687-694.

Santos, M.B., Pierce, G.J., Reid, R.J., Patterson, I.A.P., Ross, H.M. & Mente, E., 2001. Stomach contents of bottlenose dolphins (Tursiops truncatus) in Scottish waters. Journal of the Marine Biological Association of the United Kingdom, 81(5): 873-878.

Santos, M.B., Pierce G.J., Smeenk, C., Addink, M.J., Kinze, C.C., Tougaard, S. & Herman, J., 2001. Stomach contents of northern bottlenose whales (Hyperoodon ampullatus) stranded in the North Sea. Journal of the Marine Biological Association of the United Kingdom 81, 143-150.

Seibel B.A., & Carlini, D.B., 2001. Metabolism of pelagic cephalopods as a function of habitat depth: A peanalysis using Phylogenetically Independent Contrasts. Biological Bulletin, 201: 1-5.

Seibel B.A., & Walsh, P.J., 2001. Potential Impacts of CO2 injection on deep-sea biota. Science, 294: 319-320.

Shigeno, S., Tsuchiya, K. & Segawa, S., 2001. Embryonic and paralarval development of the central nervous system of the loliginid squid Sepioteuthis lessoniana. Journal of Comparative Neurology, 437(4): 449-475.

Shigeno, S., Kidokoro, H., Tsuchiya, K., Segawa, S. & Yamamoto, M., 2001. Development of the brain in the oegopsid squid, Todarodes pacificus: An atlas up to the hatching stage. Zoological Science, 18(4): 527-541.

Simpfendorfer, C.A., Goodreid, A. & Mcauley, R.B., 2001. Diet of three commercially important shark species from Western Australian waters. Marine and Freshwater Research, 52(7): 975-985.

Sinn, D.L., Perrin, N.A., Mather, J.A. & Anderson, R.C., 2001. Early temperamental traits in an octopus (Octopus bimaculoides). Journal of Comparative Psychology, 115(4): 351-364.

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Smale, M.J., Sauer, W.H.H. & Roberts, M.J., 2001. Behavioural interactions of predators and spawning chokka squid off South Africa: towards quantification. Marine Biology, 139(6): 1095-1105.

Sumbre, G., Gutfreund, Y., Fiorito, G., Flash, T. & Hochner, B., 2001. Control of octopus arm extension by a peripheral motor program. Science, 293(5536): 1845-1848.

Sun, M., Zhang, S. & Qian, W., 2001. Approach on biological characteristics of Ommastrephes bartrami on the central and eastern squid fishing grounds in the north Pacific. Marine fisheries/Haiyang Yuye. Shanghai, 23(1): 21-24.

Thompson, J.T. & Kier, W.M., 2001. Ontogenetic changes in fibrous connective tissue organization in the oval squid, Sepioteuthis lessoniana Lesson, 1830. Biological Bulletin, Marine Biological Laboratory, Woods Hole, 201(2): 136- 153.

Thompson, J.T. & Kier, W.M., 2001. Ontogenetic changes in mantle kinematics during escape-jet locomotion in the oval squid, Sepioteuthis lessoniana. Biological Bulletin, Marine Biological Laboratory, Woods Hole, 201(2): 154- 166.

Tintant, H., Lang, J., Moussa, B., Alzouma, K. & Dikouma, M., 2001. Paleocene nautiloids of Niger. Geobios, 34(6): 629-656.

Tosti, E., Di Cosmo, A., Cuomo, A., Di Cristo, C. & Gragnaniello, G., 2001. Progesterone induces activation in Octopus vulgaris spermatozoa. Molecular Reproduction and Development, 59(1): 97-105.

Tsujita, C.J. & Westermann, G.E.G., 2001. Were limpets or mosasaurs responsible for the perforations in the ammonite Placenticeras? Palaeogeography Palaeoclimatology Palaeoecology, 169(3-4): 245-270.

Vassallo-Agius, R., Watanabe, T., Imaizumi, H., Yamazaki, T., Satoh, S. & Kiron, V., 2001. Effects of dry pellets containing astaxanthin and squid meal on the spawning performance of striped jack Pseudocaranx dentex. Fisheries Science, 67(4): 667-674.

Vassallo-Agius, R., Imaizumi, H., Watanabe, T., Yamazaki, T., Satoh, S. & Kiron, V., 2001. Effect of squid meal in dry pellets on the spawning performance of striped jack Pseudocaranx dentex. Fisheries Science, 67(2): 271-80.

Vecchione, M. & Galbraith, J., 2001. Cephalopod species collected by deepwater exploratory fishing off New England. Fisheries Research, 51: 385-391.

Vecchione, M., Young, R.E., Guerra, A., Lindsay, D.J., Clague, D.A., Bernhard, J.M., Sager, W.W., Gonzalez, A.F., Rocha, F.J. & Segonzac, M., 2001. Worldwide observations of remarkable deep-sea squids. Science, 294(5551): 2505-2506.

Velasco, F., Olaso, I. & Sanchez, F., 2001. The role of cephalopods as forage for the demersal fish community in the southern Bay of Biscay. Fisheries Research, 52(1-2): 65-77.

Villaneuva, M.R.N. & Defeo, O., 2001. Growth of Octopus maya (Mollusca: Cephalopoda) along the coast of Yucatan, Mexico: a long term analysis. Revista De Biologia Tropical, 49(1): 93-101.

Voight, J.R., 2001. The relationship between sperm reservoir and spermatophore length in benthic octopuses (Cephalopoda: Octopodidae). Journal of the Marine Biological Association of the United Kingdom, 81(6): 983-986.

Voight, J.R., 2001. Morphological deformation in preserved specimens of the deep- sea octopus Graneledone. Journal of Molluscan Studies, 67: 95-102.

Von Boletzky, S., Fuentes, M. & Offner, N., 2001. First record of spawning and embryonic development in Octopus macropus (Mollusca: Cephalopoda). Journal of the Marine Biological Association of the United Kingdom, 81(4): 703-704.

Waluda, C.M., Rodhouse, P.G., Podesta, G.P., Trathan, P.N. & Pierce, G.J., 2001. Surface oceanography of the inferred hatching grounds of Illex argentinus (Cephalopoda: Ommastrephidae) and influences on recruitment variability. Marine Biology, 139(4): 671-679.

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Waluda, C.M., Rodhouse, P.G., Trathan, P.N. & Pierce, G.J., 2001. Remotely sensed mesoscale oceanography and the distribution of Illex argentinus in the South Atlantic. Fisheries Oceanography, 10(2): 207-216.

Wright, G.M., Keeley, F.W. & Robson, P., 2001. The unusual cartilaginous tissues of jawless craniates, cephalochordates and invertebrates. Cell and Tissue Research, 304(2): 165-174.

Yamaguchi, Y., Nishinokubi, H. & Yamane, T., 2001. Relationship between current conditions and catch of golden cuttlefish Sepia esculenta in Shimabara Sound. Nippon Suisan Gakkaishi, 67(3): 438-443.

Yokoyama, H. & Masuda, K., 2001. Kudoa sp (Myxozoa) causing a post-mortem myoliquefaction of North-Pacific giant octopus Paroctopus dofleini (Cephalopoda: Octopodidae). Bulletin of the European Association of Fish Pathologists, 21(6): 266-268.

Zheng, G., Zheng, Y., Dai, T., Hong, M. & Huang, G., 2001. The experiment report on the fishing gear and methods of light-pelagic trawl for squid. Marine sciences/Haiyang Kexue. Qingdao, 25(4): 5-8.

Zheng, X.D., Wang, R.C., Wang, X.F., Xiao, S. & Chen, B., 2001. Genetic variation in populations of the common Chinese cuttlefish Sepiella maindroni (Mollusca: Cephalopoda) using allozymes and mitochondrial DNA sequence analysis. Journal of Shellfish Research, 20(3): 1159-1165.

Zielinski, S., Sartoris, F.J. & Portner, H.O., 2001. Temperature effects on hemocyanin oxygen binding in an Antarctic cephalopod. Biological Bulletin, 200(1): 67-76.

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ANNEX 4(II)

CEPHALOPOD GREY LITERATURE (2001-02)

Agnew, D.J., Hill, S.L. & Beddington, J.R., 2001. Using predictions of recruitment strength to manage single-cohort stocks of an annual squid species: Loligo gahi in the Falkland Islands. International Council for the Exploration of the Sea, C.M. 2001/K:32 (poster).

Ahern, J. & Yuta, T., 2001. Apparatus to capture Stoloteuthis leucoptera. New Hampshire Sea Grant. NHU-T-01-002, 50 pp.

Anderson, R.C., 2001. Shells found in octopus-occupied beer bottles. The Festivus, 33(2): 17-19.

Anon., 2001. An update of the fishery for short-finned squid (Illex illecebrosus) in the Newfoundland area during 2000 with descriptions of some biological characteristics. Northwest Atlantic Fisheries Organisation, Dartmouth, NS (Canada). Scientific council research document, no. 01/57, 8 pp.

Anon., 2001. Assessment of subarea 3+4 northern shortfin squid (Ilex illecebrosus) for 2000. Northwest Atlantic Fisheries Organisation, Dartmouth, NS (Canada). Scientific council research document, no. 01/61, 13 pp.

Anon., 2001. Deep-sea fishery off southern Brazil: Recent trends of the Brazilian industry. Northwest Atlantic Fisheries Organisation, Dartmouth, NS (Canada). Scientific council research document, no. 01/117, 21 pp.

Anon., 2001. Cephalopod species captured by deep-water exploratory trawling in the eastern Ionian Sea. Northwest Atlantic Fisheries Organisation, Dartmouth, NS (Canada). Scientific council research document, no. 01/131, 9 pp.

Anon., 2001. Loligo forbesi and Ommastrephid squids by-catches on the Northeastern Ionian slope: Preliminary analysis of stock structure based on exploratory trawling. Northwest Atlantic Fisheries Organisation, Dartmouth, NS (Canada). Scientific council research document, no. 01/144, 6 pp.

Anon., 2001. The effect of oceanographic conditions on the spatial distribution of redfish in the Irminger Sea. Northwest Atlantic Fisheries Organisation, Dartmouth, NS (Canada). Scientific council research document, no. 01/154, 8 pp.

Arkhipkin, A.I., Sirota, A.M., Remeslo, A.V., Polishchuk, I.A. & Middleton, D.A.J., 2001. The influence of seasonal environmental changes on ontogenetic migrations of the squid Loligo gahi around the Falkland Islands. International Council for the Exploration of the Sea, C.M. 2001/K:01.

Bellido, J.M., Pierce, G.J. & Wang, J., 2001. Environmental GIS modelling on the Scottish veined squid Loligo forbesi. International Council for the Exploration of the Sea, C.M. 2001/K:03 (poster).

Biemann, M. & Piatkowski, U., 2001. Amounts and composition of trace elements in the statoliths of loliginid squids and their relationships to environmental variables. International Council for the Exploration of the Sea, C.M. 2001/K:05 (poster).

Caddy, J.F., 2001. A global perspective to fisheries for highly migratory species and their management: The additional standards provided for regional bodies by the code of conduct and the UN fish stocks agreement. Special Report. Inter-American Tropical Tuna Commission, no. 12, pp. 5-20.

Caverivière, A. & Demarcq, H., 2001. Abundance indexes of Octopus vulgaris and the coastal upwelling intensity in the south of Senegal. International Council for the Exploration of the Sea, C.M. 2001/K:33 (poster).

Challier, L., Royer, J. & Robin, J.P., 2001. Variability in age-at-recruitment and juvenile growth in English Channel Sepia officinalis described with statolith analysis. International Council for the Exploration of the Sea, C.M. 2001/K:06 (poster).

Dawe, E.G., Colbourne, E.G., Jones, D., Methven, D.A. & Hendrickson, L.C., 2001. Squids as potential indicator species of environmental or ecosystem change in the Northwest Atlantic Ocean. International Council for the Exploration of the Sea, C.M. 2001/K:07.

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Denis, V., Royer, J., Peries, P., Wang, J., Pierce, G.J., Boyle, P.R. & Robin, J.P., 2001. French and UK bottom trawl fisheries in the English Channel: spatial and temporal patterns for fishing effort and Cephalopod catch and integration of fleet components in the computation of squid and cuttlefish abundance indices. International Council for the Exploration of the Sea, C.M. 2001/K:08 (poster).

Diekmann, R. & Piatkowski, U., 2001. Early life stages of cephalopods in the Sargasso Sea: Distribution and diversity relative to hydrographic conditions. International Council for the Exploration of the Sea, C.M. 2001/K:09.

Domingues, P.M., Sykes, A. & Andrade, J.P., 2001. The use of artemia or mysids as food for hatchlings of the cuttlefish Sepia officinalis; effects on growth and survival throughout the life cycle. Aquaculture 2001: Book of Abstracts, World Aquaculture Society, 143 J.M Parker Coliseum Louisiana State, University Baton Rouge LA 70803 USA, 2001, 191.

Domingues, P.M., Kingston, T., Sykes, A. & Andrade, J.P., 2001. Growth, feeding rates and food conversions of cuttlefish Sepia officinalis fed different prey, from hatchling to adult sizes. Aquaculture 2001: Book of Abstracts, World Aquaculture Society, 143 J.M Parker Coliseum Louisiana State, University Baton Rouge LA 70803 USA, 2001, 192.

Georgakarakos, S., Haralabous, J., Koutsoubas, D., Arvanitidis, C. & Kapantagakis, A., 2001. Modelling and forecasting annual cephalopod catches in Greek waters by applying time-series analysis techniques and artificial neural networks. International Council for the Exploration of the Sea, C.M. 2001/K:11.

Golub, A.N., 2001. Oceanographic factors influencing squid larval distribution in the Southwest Atlantic. International Council for the Exploration of the Sea, C.M. 2001/K:12 (poster).

Herrmann, M., Gonschior, H. & Piatkowski, U., 2001. Hydrographic changes push European common squid Alloteuthis subulata into Kiel Bay, western Baltic Sea, its easternmost area of distribution. International Council for the Exploration of the Sea, C.M. 2001/K:13 (poster).

Jereb, P., Massi, D., Norrito, G. & Fiorentino, F., 2001. Preliminary observations of environmental effects on spatial distribution and abundance of Eledone cirrhosa and Illex coindetii in the Sicilian Channel (Central Mediterranean Sea). International Council for the Exploration of the Sea, C.M. 2001/K:34 (poster).

Jouffre, D., Caverivière, A., Lanco-Bertrand, S. & Gascuel, D., 2001. Virtual population analysis of the Senegalese Octopus vulgaris stock. International Council for the Exploration of the Sea, C.M. 2001/K:35 (poster).

Kang, W., 2001. How do squid and octopuses change colour? Scientific American, 284(5): 100.

Laptikhovsky, V.V., Remeslo, A.V., Nigmatullin, Ch.M. & Polishchuk, I.A., 2001. Recruitment strength forecast in the shortfin squid, Illex argentinus (Cephalopoda: Ommastrephidae) based on satellite SST data, and some consideration on the species’ population structure. International Council for the Exploration of the Sea, C.M. 2001/K:15.

Laptikhovsky, V.V. & Remeslo, A.V., 2001. The Falkland Current as a governor of biological patterns in the shortfin squid, Illex argentinus on the high seas (45-47 S). International Council for the Exploration of the Sea, C.M. 2001/K:16 (poster).

Lord, R., 2001. Masters of camouflage. Guernesy Press: 19.

Lordan, C., 2001. The reproductive biology of Illex coindetii in Irish waters: With an examination of life-cycle in relation to oceanographic features. International Council for the Exploration of the Sea, C.M. 2001/K:17.

Lordan, C., Warnes, S., Cross, T. & Burnell, G., 2001. The distribution and abundance of cephalopod species caught duing demersal trawl surveys west of Ireland and in the Celtic Sea. Irish Fisheries Investigations No 8, Marine Institute, Dublin, 26 pp.

Melzner, F., 2001. Determination of biochemical indices for instantaneous growth in juvenile cephalopods. Diplomarbeit vorgelegt von Frank Melzner, 1-88.

Melzner, F., Forsythe, J.W., Lee, P.G., Clemmesen, C., Piatkowski, U. & Rosenthal, H., 2001. Can RNA/DNA ratios predict recent growth in the European cuttlefish Sepia officinalis (Mollusca: Cephalopoda)? International Council for the Exploration of the Sea, C.M. 2001/K:19.

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Middleton, D.A.J. & Arkhipkin, A.I., 2001. Environmental effects on the distribution and migrations of the squid Illex argentinus (Ommastrephidae) in Falkland Islands waters. International Council for the Exploration of the Sea, C.M. 2001/K:20.

Nigmatullin, Ch.M & Shukhalter, O.A., 2001. The macro-ecosystem variation of helminth fauna in ommastrephid squid Sthenoteuthis oualaniensis. International Council for the Exploration of the Sea, C.M. 2001/K:21.

O'Dor, R.K., 2001. Counting Cephalopods in the Census of Marine Life. International Council for the Exploration of the Sea, C.M. 2001/K:22.

Pierce, G.J. & Boyle, P.R., 2001. Empirical modelling of interannual trends in abundancef squid (Loligo forbesi) in Scottish waters. International Council for the Exploration of the Sea, C.M. 2001/K:24.

Rasero, M., Román, E., Portela, J.M. & Cardoso, J.A., 2001. A contribution to the knowledge of the Cephalopod fauna of Svalbard Islands. International Council for the Exploration of the Sea, C.M. 2001/K:25 (poster).

Rasero, M., Sánchez, F. & Alvarez-Pérez, M., 2001. Geographical and bathymetric distribution of cephalopods in N- NW Spanish shelf and slope: results of the "Demersales 2000" cruise. International Council for the Exploration of the Sea, C.M. 2001/K:27 (poster).

Roberts, M.J., 2001. Chokka squid abundance maybe linked to changes in the Agulhas Bank ecosystem during spawning and the early life cycle. International Council for the Exploration of the Sea, C.M. 2001/K:28.

Royer, J., Peries, P. & Robin, J.P., 2001. Stock assessments of English Channel Loliginid squid: updated depletion methods and new analytical methods. International Council for the Exploration of the Sea, C.M. 2001/K:29.

Santurtun, M., Lucio, P., Quincoes, I. & Artetxe, I., 2001. Cephalopod catches of the Basque fleet in the Northeastern Atlantic waters during the period 1994-2000. International Council for the Exploration of the Sea, C.M. 2001/K:30.

Seixas, S. & Pinheiro, T., 2001. Seasonal variations for the concentrations of trace elements in common octopus (Octopus vulgaris) at Portuguese waters. International Council for the Exploration of the Sea, C.M. 2001/K:36 (poster).

Vecchione, M., 2001. Cephalopods of the continental slope East of the United States. In: Island in the Stream: Oceanography and Fisheries of the Charleston Bump, pp. 153-160, American Fisheries Society Symposium, vol. 25. American Fisheries Society, Bethesda, USA.

Waluda, C., 2001. Environmental influences on the recruitment of Illex argentinus in the Southwest Atlantic. PhD thesis, University of Aberdeen.

Weaver, D.C. & Sedberry, G.R., 2001. Trophic subsidies at the Charleston Bump: food web structure of reef fishes on the continental slope of the Southeastern United States. In: Island in the Stream: Oceanography and Fisheries of the Charleston Bump, pp. 137-152, American Fisheries Society Symposium, vol. 25. American Fisheries Society, Bethesda, USA.

Wood J.B. 2002. What we don't know about nautilus. Tentacle, 10: 22-23.

Xavier, J.C., Rodhouse, P.G. & Croxall, J.P., 2001. New estimates of the Martialia hyadesi stock based on predator diet - implications for South Atlantic squid fisheries. International Council for the Exploration of the Sea, C.M. 2001/K:37 (poster).

Zuur, A.F., 2001. Estimation of common trends in fisheries time-series. International Council for the Exploration of the Sea, C.M. 2001/K:31.

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WORKING DOCUMENT N° 1

ICES Working Group on Cephalopod Fisheries and Life-History Rome, Italy 10-12 April 2002

Suggestions for an experiment on size-selective mortality, especially for species subject to small mesh selection during juvenile aggregations

John F. Caddy Senior Research Fellow, Dept Environmental Science and Technology, Imperial College, University of London And Professor, CINVESTAV, Merida, Mexico

[email protected]

Introduction

There has been considerable speculation recently (e.g. Smith 1994) on the role of fishing in selecting for slow-growing, early-maturing genotypes in the wild population, as a result of the faster-growing components of the population entering the range of action of fishing gear prior to the slower-growing members of the cohort and being differentially fished out. This phenomenon has been described as ‘Lea’s phenomenon’ where back-calculated early growth rates of older fish are found to be slower than for those of juveniles. The question focussed on in this note, which does not present new data or analyses but suggests a reflection on this issue, is whether size selectivity in juvenile fisheries acts in the same selective fashion as for large mesh gear? Also fundemental to this perspective is the idea that fishing effort is not randomly distributed, but habitually is aimed at ‘patches’ of locally abundant recruits, and may drop off for older fish as suggested by Abella et.al. (1997). This question of the possibleselective effect of fisheries on sub-adults seems potentially susceptible to experimental investigation in fisheries on fast-growing species such as squids in juvenile nursery or aggregation areas. Caddy (1991) suggested modelling fishable populations of Illex illecebrosus as transient visitors on the Scotian shelf, and it seems that availability of squids to commercial exploitation may not be throughout life, but during some aggregation for feeding and/or prior to reproductive migrations. This communication poses a question that is potentially available for experimental investigation for migratory species showing this behaviour, including both squids and finfish.

Are there vulnerable aggregations of juveniles?

The early life history of a number of marine fish and invertebrates show nursery aggregations (as described for squids such as Illex and for Merluccius merluccius), and these aggregations may persist for a period of from several weeks to several months. In attempting a preliminary model of the Illex illecebrosus fishery, one may suppose that when exploited, competition for food in the nursery may result in ‘self-thinning’ as a result of selective mortality processes (see e.g. Frechette and Le Faivre (1990) for benthic invertebrates), and hence a rapid divergence of growth rates within the ‘cohort’ with result. (This assumes that the cohort was the result of a single spawning event or coherent in time).

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Figure 1. Three phases in aggregation of Illex illecebrosus on the Scotian Shelf in 1977 (Freehand copy of Fig 1 from Caddy 1979). The point is, that if this ‘model’ is correct, only during Phase II can one reasonably assume the population is ‘closed’ on the fishing grounds. Analyses of the apparent rate of mortality combining Phases I and II are liable to underestimate, while combining Phases II and III may lead to overestimate.

In the situation that selective fishing and (daily ring sampling) of this aggregation occurs, it would be worth exploring what effect this thinning will have on the final growth rate of the survivors of the cohort. This is a question that could be addressed by sequential sampling of the juvenile aggregation for its duration (Phases II – III on the ground), by examination of the daily rings on otoliths or statoliths to see if there are trends in net growth rate of individuals of the same birth date with time. The experimental procedure being proposed would address the question of how does selective fishing on juveniles in a nursery select for the growth rates of survivors by the time they leave the nursery or exploitable aggregation?

Assuming that the cohort is subject to exploitation by fine mesh gear for its period of residence in the nursery or aggregation, (as is the case for example for juvenile aggregations of Mediteranean hake - A. Abella pers. com.) then the combination of a high natural mortality rate (see e.g. Caddy 1991, which results from an aggregation of predators in an area of high vulnerability, and also from a directed fishing mortality rate, could together result in rapid cohort decimation.

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Figure 2: The scenario considered in this note. A cohort of juveniles aggregated on a fishing ground is subject to exploitation followed by dispersal to a wider adult habitat where it is less available to capture.

Selectivity

Most studies on selectivity appear to consider the selection process as similar to that described for large mesh trawl selection, with an ogive such that juveniles enter the range of mesh selectivity progressively, and that they are then are fully available to fishing for the rest of their life. This may not however be the situation where fisheries are with small mesh gear on juveniles that are aggregated but disperse later. Under these circumstances, fishing mortality is also a function of the degree of aggregation. Thus, Ft = qt*at*f , where qt is the catchability coefficient at age t, and at is the availability at this age. (If this age group is not present on the fishing grounds, then although it would be retained by the gear if present, despite qt> 0, it will not be captured).

Most applications of selection ogives also tend to use once-annual cohort-type calculations such as VPA carried out on annual cohorts of species with multi-year life spans. In fact the appropriate time scale for selection processes on juveniles should be shorter (weeks, two-week periods), especially for short-lived and/or fast-growing species, and should take into account (as for the Mediterranean) that juveniles spend some time (weeks, months) in such aggregations. During this time they are particularly vulnerable to capture by fine mesh gear.

There are therefore two factors to consider in selectivity: a) probability of capture as a function of size, and b) the duration of exposure to fishing, (which depends not just on the catchability coefficient), or in other words, changes the availability of the cohort to fishing.

As will be evident from figure 3, though the probability of capture of fast-growing juveniles increases fastest, their duration of exposure to intensive fishing is much shorter, assuming that after they have dispersed from the fishing grounds (upper dotted line), their availability to fishing is much reduced. From geometrical considerations, if Fig 3 is a realistic picture of the situation, slower-growing individuals would suffer a higher mortality rate before leaving the nursery area in situations where an intensive juvenile fishery applies there. (In other words, where the fishing intensity is aggregated over the higher densities and much less so elsewhere, as suggested by Caddy 1975). This speculation assumes the situation differs from that with large mesh gear for species with multiple year classes, where slow-growing individuals enter the range of exploitation of the gear late, are exposed to fishing for the rest of their life, and hence

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there is differentially greater survival of slow-growers selected for by their reduced catchability and shorter duration of exposure to fishing as juveniles.

The question that is being asked here, is whether the combined natural and fishing mortality rates would effectively select for or against faster-growing individuals before they leave the aggregation and disperse elsewhere on the fishing grounds where they could be less vulnerable to both fishing and predation at lower densities (fig 2)? One may expect that in Phase I of fig 1, faster-growing individuals may arrive first on the fishing grounds, (so that the period of relevance to this experiment should begin when all the cohort is available to fishing).

Given that this scenario is theoretically possible and worth testing in the field, we can suppose that:

1) The probability of capture and the fishing rate on the aggregation (Fagg) will be a product of the duration and the intensity of fishing, i.e. Fagg = Σqt.ft.∆t , where the summation term is over the n weeks of stay in an aggregation or nursery area of a group of fish from a cohort with a variable rate of growth. (Note that availability is assumed unity during period ∆t, and zero after it. Note also that n is assumed to vary widely from individual to individual (fig 3) where a range of different growth rates occur in the population).

2) The two factors that will determine the value of Fagg, are the total duration of availability, ∆t on the nursery ground, and qt the selectivity of the gear. In conventional fishery calculations it is normally considered that the latter is the determining factor, since stock assessment calculations are usually performed on an annual basis and the assumption is made that after full selection, the availability to capture by the gear remains high and constant, irrespective of degree of aggregation. This would not necessarily be the case for species showing vulnerable juvenile aggregations, if sequential calculations such as VPA were made (say) at a 1-2 week interval for intervals ∆t(1) , ∆t(2) , ∆t (3) …… ∆t(n).

3) As can be seen from figure 2, although the probability of capture increases faster with size for individual (1) than for the other slower-growing individuals (2, 3,& 4), the duration of high vulnerability on the fishing grounds of the slower- growing juveniles is greater. This conceivably could more than make up for the initially lower q values for slower- growing individuals. This scenario would only be valid of course, if the duration of stay in the nursery habitat is a function of size and not age.

Figure 3: Scenario for a situation where residence in the juvenile aggregation is bracketed by the sizes corresponding to the two horizontal dotted lines, and selectivity by fishing is as shown on the left of the diagram. The question is, what would be the relative survival rate of growth forms 1) to 4), assuming their duration of exposure to intensive fishing is given by Dt(1) to Dt(4). (Note: These symbols are the same as ∆t(1) , ∆t(2) , ∆t (3) …… ∆4).

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An experimental suggestion:

It would be relatively simple to simulate the process described above mathematically, but relatively trivial, since the real question is whether the assumptions as set out above are correct.

The presence of statolith rings in squids, and daily rings in or close to the nucleus of the otolith in juvenile finfish, both seems to offer the possibility however of investigating the relative survival of slow- and fast-growing individuals in exploited aggregations of juveniles. If during the course of the life history stage passed in the nursery or in a fishable aggregation, the back-calculated relationship between otolith/statolith diameter and number of rings shows a positive slope, I suppose one can assume that selection for faster growth is occurring in the exploited nursery stage, (assuming that the back-calculated ages shows that the individuals being compared had approximately the same birth date), and vice versa if the reverse is the case? A comparison of the relative frequency of slow growth in samples taken early on within the aggregation with those at the end of the aggregated phase would therefore be revealing, as would be a comparison of back-calculated juvenile growth rates from the ring spacing on otoliths/statoliths of those adults that survived the fishery on juvenile aggregations. Were these predominantly slow or fast growing as juveniles?

I believe this type of experiment is important for a number of reasons. Principally that too many assumptions on selectivity and growth have been based around fisheries for adult fish, leading to mistaken conclusions on the viability of fisheries for juveniles and adults (See Abella et al. (1997), and Caddy and Abella 1999). The possibility that a positive selection for growth rate could occur as is proposed for investigation here (a reverse ‘Lea’s phenomenon’?), has not to my knowledge been considered in the literature, and seems a critical experiment to carry out at this time when fisheries on spawning adults are coming under re-examination.

Although this paper is not promoting the exploitation of juvenile aggregations, it is noted that there has been insufficient study of the strategy that has begun to be discussed for sedentary invertebrate resources of thinning dense aggregations of recruits. In fact, for finfish resources, conserving some adult spawning biomass in refugia and aiming for a controlled exploitation of the larger juveniles, has proved to be a sustainable exploitation pattern for Mediterranean groundfish even in absence of quotas and complete effort control (see Abella et.al. 1997). If the hypothesis discussed in this paper is correct, paradoxically, thinning of slow-growing individuals in dense aggregations of juveniles subject to food limitation, may be possible and if so could lead to increases in average growth rates. This seems a hypothesis worth further investigation, both for cephalopods and for fisheries on other resource aggregations prior to reproduction.

References

Abella, A.J., J.F. Caddy and F. Serena (1997). Do natural mortality and availability decline with age? An alternative yield paradigm for juvenile fisheries, illustrated by the hake Merluccius merluccius fishery in the Mediterranean. Aquat.Living Resour., 10:257-69

Caddy, J.F. (1975). Spatial model for an exploited shellfish population, and its application to the Georges Bank scallop fishery. J. Fish. Res. Bd. Can., 32: 1305-28.

Caddy, J.F. (1979). Preliminary analyses of mortality, immigration, and emigration of Illex populations on the Scotian Shelf. ICNAF Res. Doc. 79/VI/120 20 p (Mimeo).

Caddy, J.F. (1996). Modelling natural mortality with age in short-lived invertebrate populations: Definition of a strategy of gnomonic time division. Aquat.Living Resour., 9:197-207

Caddy, J.F. (1991). Daily rings on squid statoliths: an opportunity to test standard population models? Squid Age determination Using Statoliths. NTR/ITPP Special publication (Italy): p 53-66.

Caddy, J.F. and A. Abella (1999). Reconstructing reciprocal M vectors from length cohort analysis (LCA) of commercial size frequencies of hake, and fine mesh trawl surveys over the same grounds. Fish. Res. 41: p 169-175.

Frechette M., D. Lefaivre (1990). Descriminating between food and space limitation in benthic suspension feeders using self-thinning relationships. Mar. Ecol. Prog. Ser. 65, p 15-23.

Smith, P.J. (1994). Genetic diversity of marine fisheries resources: possible impacts of fishing. FAO Fish. Tech. Pap 344, 53p.

68

WORKING DOCUMENT N° 2

ICES Working Group on Cephalopod Fisheries and Life-History

Rome, Italy 10-12 April 2002

Influence of oceanographic factors and abundance of Octopus vulgaris in Galician waters (NE Atlantic)

A.F. González, A. Guerra, J. Otero and F.J. Rocha Instituto de Investigaciones Marinas, Consejo Superior de Investigaciones Científicas, Eduardo Cabello 6, 36208 Vigo, Spain

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The aim of this work is to integer biological and fishing data of paralarvae and adults of common octopus, and climate data in order to study the influence of atmospheric-oceanographic parameters in the natural variation of a resource of socio-economic importance in Galicia.

The study area is situated in waters off the Northwestern Spanish coast (Figure 1). The wind regimes, principally in the western Galician coast, favour the upwelling in this area, where a deep-water mass, the south flowing subtropical Eastern North Atlantic Central Water Mass (ENACWst), is upwelled during spring and summer along the Galician coast. This cold water mass greatly increases the primary production and the estuarine circulation.

Sampling methods. A total of 19 research cruises were undertaken during 2000 and 2001. We undertook a zooplankton sampling (oblique hauls) from close the bottom to the surface with a 700 mm diameter Bongo pair nets of 370 m mesh. The bongo net was equipped with a current meter to calculate the volume of water filtered during each haul, which permit estimations of abundance of parlarvae (No m-3). Vertical temperature-salinity profiles were obtained at the beginning and the end of each transects using a CTD. On the other hand, the total water volume comprised in both areas studied (east and west sides of the Cíes Islands) was estimated, using the surface and the average depth, in order to calculate abundance of paralarvae.

Treatment of samples. The zooplankton samples were fixed with 4% buffered formalin during 24 hours and then preserved in 70% alcohol. Paralarvae of Octopus vulgaris were separated at the lab using reference keys and paralarvae of this species.

Adult samplings. Samples of O. vulgaris were obtained from commercial landings in the Galician fishing port of Moaña (Ría of Vigo). A total of 156 specimens (85 males and 71 females) were collected from small-scale trap fishery. Sex and wet body weight (BW) were recorded. Dorsal mantle length (ML) was measured to the nearest mm. Potential fecundity of females in maturity stages 3 and 4 were estimated. Ovary samples were taken from their medial parts and weighed to the nearest 0.01 g. These samples were fixed in 4% formaline and stored in 70% ethanol. The real fecundity was estimated from egg masse obtained under rearing conditions from wild specimens collected in the Ría de Vigo

Fishery and oceanographic data. Annual catches made by the whole Galician fleet for Octopus vulgaris were recorded for the period 1990-2000 based on the official statistics of the Galician Government. Catches obtained in the area studied (Cíes fishing ground) were made by small boats. To evaluate the size of the spawning stock in this area it is necessary to obtain the following parameters: growth rate, natural mortality, fishing mortality, exploitation rate and mortality rate. The estimation of exploitation rate is derived from the values of the instantaneous rate of natural mortality. Once known the biomass of the spawning stock, as well as the number of eggs spawned by the females (real fecundity), we estimated the number of paralarvae that would hatch in the area studied and the mortality rate during the early life stage of the paralarvae.

Information on the intensity and direction of geostrophic winds was derived from daily surface atmospheric pressure data to provide an upwelling index. These data were obtained from the Spanish Meteorological Institute's recording station located in the western Galician coast. To relate the inflow of subtropical Eastern North Atlantic Central water into the continental shelf with the abundance of paralarvae, we used the mean temperature near the sea floor (twenty meters, T 20).

RESULTS

Octopus vulgaris paralarvae

A total of 53 and 54 paralarvae of O. vulgaris were obtained during 2000 and 2001, respectively. The highest number of paralarvae was found during late spring and autumn. Its distribution was different in the study area, being more frequent in the deeper transects, at depths ranging from 85 and 105 m. The size of the paralarvae varied from 1.20 and 2.35 mm ML. During the hauls, we collected newly hatched paralarvae as well as specimens up to 2.35 mm ML, being most frequent the specimens ranging from 1.40 and 1.80 mm ML.

The total volume of water estimated in the study area was 4329x106 m3 and 2730x106 m3 for the outer and inner areas, respectively. On the other hand, the volume of water filtered by the pair bongo net (and the number of paralarvae collected in both areas allowed us to calculate abundances (No of paralarvae/m3) in the inner and outer waters of the area studied. The estimated number of paralarvae present in the Cíes fishing ground was 11.516.476 and 11.265.686 for 2000 and 2001, respectively.

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Concerning the relationship between abundance of paralarvae and oceanographic parameters, the paralarvae of the common octopus are related to the ENACWst. Thus, we observed higher abundances when upwelling events were occurring.

Mortality rate estimation of the paralarvae

The exploitation rate of the common octopus population in the Cíes fishing ground was 57% for the year 2000 (González et al. unpub. data). The potential number of paralarvae available in the area studied during 2000 is estimated as follows: a) The exploitation rate is 57% and the catch is 1186 Mt for year 2000 in the Cíes fishing ground. Therefore, at least 895 Mt of common octopus should be available in the fishery. b) Taking into account that the sex ratio in the fishery is 1:1 (females represent 50% of the animals in the fishery) and considering a conservative average weight for the mature females of 2.5 Kg, the potential total number of spawners would be 179,000. Each female of O. vulgaris spawn an average of 100,000 eggs. Thus, the potential number of hatchlings would be 17900x106. c) Considering that the number of paralarvae in the plankton was 11.5x106, the mortality rate in this early stage of development would be 99.94%.

Relationship between catches for the whole Galician fleet and upwelling index

Among a total of 156 adult specimens of common octopus examined, 85 females and 34 males were mature. Most mature females were found during spring summer and autumn. During summer, 100% of the females were mature. The sex ratio during 2000 and 2001 was 1:1. The monthly visits to the ports revealed that catches are higher during autumn, with a peak centered in October. There are not catches during May due to the closing season established by the Government of Galicia.

The maximum catch in the Galician waters during last decade was observed in 1998. The pattern followed by the catches is similar to that experienced by the upwelling index with a delay of one or two years.

PRELIMINARY CONCLUSIONS

• The waters surrounding the Ría de Vigo is an area of reproduction and spawning of Octopus vulgaris. The spwaning of this species extends all year round with two peaks centred during spring and autumn.

• Spawning is related to the presence of northeastern Atlantic central waters in the shelf during upwelling events.

• The estimated number of paralarvae in the Cíes fishing ground for 2000 and 2001 was 11.5 x106 and 11.3 x106, respectively.

• Mortality of paralarvae was estimated to be 99.94%.

• There is a cyclic recurrence between strength of upwelling and catches.

• Existence of a retard of 1-2 years between upwelling and variation of catches.

• The effort has to be controlled because it affects the availability of spawners and, therefore, on the abundance of paralarvae.

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Ría de Vigo E D C B A

Cíes Islands

Figure 1

72

WORKING DOCUMENT N° 3

ICES Working Group on Cephalopod Fisheries and Life-History

Rome, Italy 10-12 April 2002

RECENT TRENDS IN MEDITERRANEAN CEPHALOPODS CAPTURE PRODUCTION

Patrizia Jereb

Istituto Centrale per la Ricerca scientifica e tecnologica Applicata al Mare (ICRAM) Via di Casalotti 300 – 00166 Rome - ITALY

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INTRODUCTION

Cephalopod catches increased steadily in the last 30 years, from about 1 million Mt to around 3.5 million Mt (Figure 1), confirming a potential development of the fishery hypothesized by different authors (Voss, 1973, Clarke, 1983, Boyle, 1990). With the increasing exploitation of other, more conventional resources, and the subsequent world fishery view marked by over-fishing and decline of finfish stocks, cephalopods gained even more attention, as one of the few remaining group of resources still capable of experiencing increase in landings, at least as for some groups/species and some areas of the world (Caddy and Rodhouse, 1998).

Squid are by far the main representative in the capture/world production, accounting for about 80% of total cephalopod catches: the impressive increase in cephalopod production during the 1980s is mainly due to the discovery of squid resources in the SW Atlantic. Squid are also mainly responsible for landings/catches fluctuations, which, however, are a characteristic of the whole group. Squid stocks also experienced true collapses at least in two well known cases (NW Pacific Todarodes pacificus fishery failure in the 1970s and NW Atlantic Illex illecebrosus fishery failure in the 1980s) and this is thought to have occurred mainly as a consequence of unfavourable environmental conditions (or environmental changes) probably aggravated by heavy fishing pressure (Dawe & Warren, 1993).

The many peculiarities of cephalopod biology, physiology and life cycle which make them more difficult to study and understand in the fishery context than finfish resources (Caddy, 1983, Voss, 1983, Pauly, 1985, Boyle, 1990, Lipinski, 1998) along with the related difficulties to apply to them methods used to assess and manage more “conventional” fisheries (see Pierce and Guerra, 1994 for a review) are among the main reasons responsible for the fact that very few cephalopod fisheries are currently managed.

Caddy (1983) suggested that cephalopod “life-strategy” may guarantee against environmental stress conditions (including those caused by heavy fishing), but other authors afterwards highlighted the susceptibility of cephalopod stocks to overfishing (Rosemberg et al., 1990) and in more recent years, with cephalopod fisheries experiencing further developments due a still growing market demand, parallel concern developed regarding potential for over-exploitation (Pierce and Guerra, 1994, O’Dor and Dawe, 1998, Haimovici et al., 1998, Sanchez et al., 1998).

This resulted in a broad consensus versus a common effort to make good use of the experience and errors of the past in finfish management, in order to avoid similar errors and possible failures in the field of cephalopod exploitation (Lipinski et al., 1998).

The value of research as a key factor towards this goal was also highlighted, especially in the fields of life-cycles clarification, stock structure (genetics), role in the food web and interactions with the environment.

The last topic seems of particular interest within the more general context of climate/environmental global changes, since cephalopods unusual biological characteristics and short life cycles are strongly linked to the environment. Therefore cephalopods are potentially very good “indicators” of changes in environmental conditions, both locally and on a broader scale (O’Dor and Dawe, 1998; Arkhipkin et al., 2001; Bendik, 2001; Dawe et al., 2000, 2001; Jereb et al., 2001; Laptikhowsky and Remelso, 2001; Roberts, 2001).

THE MEDITERRANEAN CASE

Due to the increasing importance of the EU cephalopod market, at present the main world market for this commodity (GLOBEFISH, 2001), it seems even more important to monitor European cephalopod fishery, and a special focus should be posed to the Mediterranean fishery, due to its contribution to European cephalopod production and the peculiarities of both the Mediterranean Sea and the Mediterranean fishery context.

Therefore it seems interesting to put in evidence what was observed during an updating of the available information on Mediterranean and Italian cephalopod resources within a research project currently carried on by ICRAM Institution and briefly discussed within the ICES Working Group Annual Meeting held in Rome (10-12 April, 2002), namely, a decreasing trend in Mediterranean captures production of the last fourteen years (Figure 2; Table 1; FAO, 2002a. Data were analysed with the exclusion of those referring to the Black Sea and the Marmara Sea).

This decreasing trend is also evidenced when analysing captures trends of the three main groups separately (Figure 3), even though this is less evident for the “cuttlefishes” category, which is the one presenting more evident “pulses” or fluctuations along the way.

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A further analysis revealed that Octopus vulgaris and the two Eledone species capture productions are both decreasing among Octopods (Figures 4; 11), as are Loligo species and Todarodes sagittatus among squid (Figure 5) and the Sepiidae/Sepiolidae category among cuttlefishes (Figure 6), while Sepia officinalis maintains an increasing trend among fluctuations.

Even though Italy is the main Mediteranean cephalopod producer (Figures 7-8), therefore in principle the main responsible of the generally decreasing trend, an analysis of the captures by group/species and country (Figure 9-20) was done, at least for those countries which represent noteworthy producers among the Mediterranean ones, as an attempt to identify possible individual trends and fluctuations.

On the whole, and taking into account cephalopod abundance characteristic “fluctuating” scheme, a declining trend is observed in octopods and squid captures, where Todarodes sagittatus, Loligo spp and Eledone spp captures registered in 1999 their lowest value since the ‘70s. For the other species/categories the situation is more articulated, but only Sepia officinalis showed an increasing trend when considering recent years.

An analysis was also done to compare captures in the different Mediterranean areas (Figures 21-24), this revealing a decreasing trend for the Adriatic and the Ionian sectors, while in other areas the situation is more or less “stable”.

In the Adriatic Sea, the pronounced peak which interrupts an otherwise rather continuous trend in the last decade is due to the Italian unusual production of the Sepiidae-Sepiolidae group in 1994 (Figure 33-34). The decreasing trend showed by squid captures is mainly due to T. sagittatus captures decline (Figure 26; 30), while Loligo species, even at low levels compared to the ‘70s, show fluctuations (Figure 26; 30). Fluctuating peaks characterize the Octopods (Figure 27), with both the Eledone species and Octopus vulgaris captures being presently in a declining phase, after the high production of 1990-1992 (Figures 31-32). As for the Sepiidae/Sepiolidae group, a production peak in 1994 (Figures 33- 34) interrupts a slowly declining trend since the ‘80s.

In the Ionian Sea also (Figures 35-44) squid and octopuses captures showed a declining trend, mainly due to Octopus vulgaris and Loligo spp decline. Todarodes sagittatus showed a high production pulse in 1998 which interrupted an otherwise descending trend while in Eledone spp the observed trend reminds of the already mentioned years of unusually high abundance (“plague years”; Boyle, 1997) alternating with years of low abundance. Present situation however, show captures being at the lowest level within the 30 years range analysed. Sepia officinalis again is in the increasing phase, while for the Sepiidae- Sepiolidae group the opposite is true.

In the Aegean Sea captures peaked for the three groups in the late ‘80s (Figure 45), but squid dropped by almost immediately afterwards (Figure 46) and captures are still in a declining phase. Octopus vulgaris (Figure 47) maintained high levels till the recent most years and is now in the descending phase, while Sepia officinalis production by Greece is still in the increasing phase (Figure 48); interestingly enough this is not so for Turkish production, which decreased abruptly after the peaks of 1998-1990.

Also in the Levant basin (Figure 49) Loligo spp, Octopus vulgaris and Sepia officinalis captures experienced a peak in 1988-1991 (Figures 50-52), to go down again and “stabilise” at lower levels, which, however, show an increasing trend for Loligo spp. (Figure 50), while fluctuations characterise the Octopus vulgaris (Figure 51) production and Sepia officinalis is declining but not so the Sepiidae-Sepiolidae category (Figure 52). Turkey is the main cephalopod producer in that area.

The analysis of the Sardinia area (Figures 53-56) indicate a more “stable” situation, with the typical fluctuations characteristic of the group. Perhaps worth mentioning (even considering the low “numbers” in comparison with other situation) the increasing amount of the Tunisian S. offficinalis production (Figure 57).

As for the Balearic area (Figures 58-60), while the octopuses and squid groups show a more or less stable trend in the production of the last decade, the Sepiidae/Sepiolidae are in moderate decline and Sepia officinalis production increased in the last 5-6 years. A closer look to the squid production however (Figure 61) show an slight declining trend in the Loligo spp captures, in spite of the characteristic sporadic high peaks.

As for the Gulf of Lions (Figures 62-64), no trend seems evident, except for a decline in Sepia officinalis production.

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DISCUSSION AND CONCLUSIONS

In spite of the understandable and well known limits/limitations of using FAO captures/production statistics, these are the main world wide references when an historical series of records analysis is needed on a global scale.

Basing on these data, a decreasing “trend” in the Mediterranean cephalopod captures production since 1988 is apparent when analysing global numbers, in spite of the “ups” detectable in 1992, 1994 and 1997. Recent-most data (FAO 2002b, Mediterranean captures production for the year 2000) stands for another “up” (i.e. about 53500 Mt compared to the 50700 reported for 1999), but GFCM updated statistics were not yet available, so they could not be analysed.

A more detailed analysis obviously evidenced a more variable situation, even though a declining trend in Loliginid squid, Todarodes sagittatus and, with a more complex pattern, of the Eledone species, characterises the last decade of captures production data. Also the lower trend of the Sepiidae-Sepiolidae group in more than one area is worth noting, as is the opposite trend shown by Sepia officinalis.

Cephalopods are known for their abundance fluctuations. These are also detectable, for example, in the North East Atlantic production captures, even when analysed per group (Figure 65), were the increasing trend of the last twenty years is evident.

However, as it was commented during the Workshop, the situation framed for the Mediterranean Sea during the last decade by the analysis of FAO data does not seem to fit into such a general scheme.

A throughout analysis of the reported observations was not the objective of this note, first, among others reasons, because it would require the availability of more specific data and information on a national (Italian) scale as well as on a regional level, thus requiring co-operation among the different interested Mediterranean countries.

Therefore, only some comments are hereby expressed, to contribute stimulating a closer attention to Mediterranean cephalopods, a more adequate monitoring and, possibly, better focused research activities towards fisheries resources of such a relevant importance for the Mediterranean economy and the Mediterranean ecosystem.

Caddy and Rodhouse (1998) suggested that, where cephalopods have been heavily fished as in the Mediterranean, “there is evidence that landings now vary as a function of fishing effort and environmental variation”. If this is true, the observed trend is even more interesting.

The European Community is working towards a progressive reduction of fishing effort in the Mediterranean, goal which each member country is asked to contribute to. How this will be obtained, however, is still under discussion, due to the peculiarities of the Mediterranean fishery. In any case, any general information on fishing effort variation would be difficult to relate to the observed trend in cephalopod captures unless ancillary (focused) information were collected.

Other observations recently supported evidence that fisheries productivity increased in the Mediterranean since the ‘70s (see Caddy, 2000 and Caddy and Garibaldi, 2000 for a review) with parallel increase in the piscivores landings (e.g. swordfish, hake) and zooplanktivores (e.g. sardine, anchovy, sprat). In such a scenario, cephalopods species like ommastrephids, for example, would play a very interesting role as prey/predators resources on not irrelevant importance. The effects of consumption of important pelagic squid and fish by predatory fish was recently studied in the Northeastern USA shelf (Overholtz et al., 2000), and the conclusion made that changes in predatory abundance may have important implications for long-term fishery yields of pelagic species. The observed trend in the Todarodes sagittatus captures production seems therefore of peculiar interest.

Environmental variations are occurring on a global scale, and special attention was recently paid to the Mediterranean “tropicalization” (Andaloro & Rinaldi, 1998), a phenomenon which is currently under observation. How this could affect cephalopod species may be a more important topic to investigate than previously thought.

Market pressure of foreign products against Mediterranean ones (i.e. economically competitive) should also be investigated in details, even though a very preliminary analysis of Italian cephalopod trade data in recent years (Figures 66- 67; FAO, 2002c) did not show peculiar trends in relation to this aspect.

Even though contradictory trends may be obtained when comparing biomass estimates computed by scientific surveys to commercial catch data (e.g. Roel and Payne, 1998), the possibility to compare data and observations is nonetheless an essential tool for investigation. Therefore, recent-most MEDITS (Mediterranean International Trawl Survey, a research project funded by the EU and several Mediterranean National Institutions) data reports/comments on a broad

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comparative scale, once available, could help in getting important elements toward a better diagnosis of the observed situation.

In this respect it is noteworthy to mention that a decreasing trend in the abundance of Eledone cirrhosa was reported for the Ligurian and Tyrrhenian Sea (i.e. 20 kg/km2 in 1994, 14,1 kg/km2 in 1996 and 12.2 kg/km2 in 1999) basing on the analysis of MEDITS-IT data (Ardizzone and Belluscio, 1999).

To conclude with, present observations support the need for a better look at the situation of the Mediterranean cephalopod populations.

REFERENCE LIST

Andaloro, F. & Rinaldi, A. (1998). Fish biodiversity change in Mediterranean Sea as tropicalization phenomenon indicator. In: D'Angelo, E.G. and Zanolla, C. (Eds.): Indicators for assessing desertification in the Mediterranean. Rome, ANPA

Ardizzone, G.D. & Belluscio, A. (1999). MEDITS (International bottom trawl surveys in the Mediterranean) Biological Report – Survey Results for Italy – Area M1 – Ligurian and Tyrrhenian Seas.

Arkhipkin, A.I., Sirota, A.M., Remelso, A.V., Polishchuk, I.A. & Middleton, D.A.J. (2001) The influence of seasonal environmental changes on ontogenetic migration of the squid Loligo gahi around the Falkland Island. ICES, C.M. 2001/K:01

Bendik, A.B. (2001) Abundance explosion of the Jumbo squid, Dosidicus gigas, on the high seas of the Peruvian region in relation to anomalous oceanographic patterns. ICES, C.M. 2001/K:04

Boyle, P.R. (1990). Cephalopod Biology in the Fisheries Context. Fisheries Research 8, 303-321.

Caddy, J.F. (1983). The cephalopods: factors relevant to their population dynamics and to the assessment and management of stocks. FAO Fish. Tech. Pap. 231, 416-452.

Caddy, J.F. (2000). Marine catchment basin effects versus impacts of fisheries on semi-enclosed seas. ICES Journal of Marine Science 57, 628-640.

Caddy, J.F. & Garibaldi, L. (2000). Apparent changes in trophic composition of world marine harvests: the perspective from the FAO capture database. Ocean & Coastal Management 43, 615-655.

Caddy, J.F. & Rodhouse, P.G. (1998). Cephalopod and groundfish landings: evidence for ecological change in global fisheries? Review in Fish Biology and Fisheries 8, 431-444.

Clarke, M.R. (1983). Cephalopod biomass. Estimation from predation. Memoirs of the National Museum Victoria 44, 95-107.

Dawe, E.G., Colbourne, E.B. & Drinkwater, K.F. (2000). Environmental effects on recruitment of shorth-finned squid (Illex illecebrosus). ICES Journal of Marine Science 57, 1002-1013.

Dawe, E.G., Colbourne, E.G., Jones, D.D., Methven, D.A. & Hendrickson (2001). Squids as potential indicators species of environmental or ecosystem change in the Northwest Atlantic Ocean. ICES, C.M. 2001/K:07

Dawe, E.G. & Warren, G. (1993) Recruitment of short-finned squid in Northwest Atlantic Ocean and some environmental relationships. J. Cephalopod Biol. 2, 1-21.

FAO (2002) a. Fishstat plus (v.2.30). GFCM (Mediterranean and Black Sea) captures production (1970-1999). Also downloadable at: www.fao.org/fi/statist/fisoft/fishplus.asp

FAO (2002) b. Fishstat plus (v. 2.30). Capture fish production (1970-2000). Also downloadable at: www.fao.org/fi/statist/fisoft/fishplus.asp

FAO (2002) c. Fishstat plus (v. 2.30). Commodities trade and production (1976-1999). Also downloadable at: www.fao.org/fi/statist/fisoft/fishplus.asp

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GLOBEFISH (2001). Cephalopods. Commodity update.

Haimovici, M., Brunetti, N.E., Rodhouse, P.G., Csirke, J. & Leta, R.H. (1998). Illex argentinus. In: Rodhouse, P.G., Dawe, E.G. and O'Dor, R.K. (Eds.), Squid recruitment dynamics. The genus Illex as a model. The commercial Illex species. Influences on variability. FAO Fisheries Technical Paper 376, 27-58.

Jereb, P., Massi, D., Norrito, G. & Fiorentino, F. (2001). Preliminary observations of environmental effects on spatial distribution and abundance of Eledone cirrhosa and Illex coindetii in the Sicilian Channel (Central Mediterranean Sea). ICES, C. M. 2001/K:34

Laptikhovsky, V.V. & Remelso, A.V. (2001). The Falkland Current as a governor of biological patterns in the shortfin squid, Illex argentinus on the high seas (45-47 S). ICES C.M. 2001/K :16

Lipinski, M.R. (1998). Cephalopod life cycles: patterns and exceptions. In: Payne, A.I.L., Lipinski, M.R. and M.A.C. Roeleveld (Eds.). Cephalopod Biodiversity, Ecology and Evolution. S.Afr. J. mar. Sci. 20, 439-447.

Lipinski, M.R., Butterworth, D.S., Augustyn, C.J., Brodziak, J.K.T., Christy, G., Des Clers, S., Jackson, G.D., O'Dor, R.K., Pauly, D., Purchase, L.V., Roberts, M.J., Roel, B.A., Sakurai, Y. & Sauer, W.H.H. (1998). Cephalopod fisheries: a future global upside to past overexploitation of living marine resources? Result of an international workshop, 31 August - 2 September 1997, Cape Town, South Africa. In: Payne, A.I.L., Lipinski, M.R. and M.A.C. Roeleveld (Eds.). Cephalopod Biodiversity, Ecology and Evolution. S.Afr. J. mar. Sci. 20,

O'Dor, R.K. & Dawe, E.G. (1998). Illex illecebrosus. In: Rodhouse, P.G., Dawe, E.G. and O'Dor, R.K. (Eds.), Squid recruitment dynamics. The genus Illex as a model. The commercial Illex species. Influences on variability. FAO Fisheries Technical Paper 376, 77-104.

Overholtz, W.J., Link, J.S. & Suslowicz, L.E. (2000). Consumption of important pelagic fish and squid by predatory fish in the northeastern USA shelf ecosystem with some fishery comparisons. ICES J. Mar. Sci. 57, 1147-1159.

Pauly, D. (1985). Population dynamics of of short-lived species, with emphasison squids. NAFO Sci. Coun. Studies 9, 143-154.

Pierce, G.J. & Guerra, A. (1994). Stock assessment methods used for cephalopod fisheries. Fisheries Research 21, 255- 285.

Roberts, M.J. (2001) Chokka squid abundance maybe linked to changes in the Alghulas Bank ecosystem during spawning and the early life cycle. ICES, C.M. 2001/K:28

Roel, B.A. & Payne, I.L. (1998). Management of the South African chokka squid jig fishery under uncertainty regarding trends in resource abundance. ICES, C.M. 1998/M :16

Rosenberg, A.A., Crombie, J.A., Beddington, J.R. & Kirkwood, G.P. (1990). The assessment of stocks of annual squid species. Fisheries Research 8, 335-350.

Sanchez, P., Gonzalez, A.F., Jereb, P., Laptikhovsky, V.V., Mangold, K.M., Nigmatullin, C.M. & Ragonese, S. (1998). Illex coindetii. In: Rodhouse, P.G., Dawe, E.G. and O'Dor, R.K. (Eds.), Squid recruitment dynamics. The genus Illex as a model. The commercial Illex species. Influences on variability. FAO Fisheries Technical Paper 376, 59-76.

Voss, G.L. (1973). Cephalopod resources of the world. FAO Fish. Cir. 149, 75.

Voss, G.L. (1983). A Review of Cephalopod Fisheries Biology. Memoirs of the National Museum Victoria 44, 229-241.

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TABLE 1 - Mediterranean and Black Sea captures production (Mt) 1970-1999 (FAO, 2002 a) (1/4)

Country area Species 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 Albania Ionian Loligo spp Albania Ionian Octopus vulgaris Albania Ionian Sepia officinalis Algeria Balearic Cephalopoda Algeria Balearic Loligo spp Algeria Balearic Octopodidae Algeria Balearic Sepia officinalis Croatia Adriatic Cephalopoda Croatia Adriatic Loligo spp Croatia Adriatic Octopus vulgaris Croatia Adriatic Sepiidae, Sepiolidae Cyprus Levant Octopodidae 60 50 62 52 45 44 51 60 64 66 58 59 68 188 157 Cyprus Levant Sepia officinalis 66 55 62 52 45 44 45 53 64 57 54 74 70 91 95 Egypt Levant Sepiidae, Sepiolidae 346 384 442 338 304 322 255 337 560 743 356 644 691 990 780 France Gulf of Loliginidae, Lions Ommastrephidae France Sardinia Loliginidae, 92 Ommastrephidae France Gulf of Loligo spp 197 932 1400 1058 199 261 373 315 408 521 369 380 292 372 203 Lions France Sardinia Loligo spp 20 29 15 17 27 17 5 15 8 1 9 France Gulf of Octopus vulgaris 957 1605 1329 1155 1001 925 1038 1061 1064 823 699 Lions France Sardinia Octopus vulgaris 8 6 France Gulf of Sepia officinalis 249 259 256 341 307 196 225 277 300 367 365 264 Lions France Sardinia Sepia officinalis 25 37 42 42 5 3 33 45 47 30 33 32 France Gulf of Todarodes sagittatus 9 8 Lions France Sardinia Todarodes sagittatus 9 13 Gaza Levant Loligo spp Strip(Palestine) Gaza Levant Sepiidae, Sepiolidae Strip(Palestine) Greece Aegean Loliginidae, 447 461 250 300 266 171 137 128 156 177 203 209 223 271 127 Ommastrephidae Greece Ionian Loliginidae, 102 101 32 21 27 48 44 47 27 8 46 52 30 121 Ommastrephidae Greece Aegean Loligo spp 201 220 305 339 417 397 551 483 443 419 429 519 345 Greece Ionian Loligo spp 20 25 54 59 58 51 49 29 70 69 61 110 Greece Aegean Octopodidae 734 808 712 630 735 889 806 739 883 972 1069 917 671 748 611 Greece Ionian Octopodidae 29 25 42 20 41 43 30 48 35 24 41 13 21 86 Greece Aegean Octopus vulgaris 375 344 227 Greece Ionian Octopus vulgaris 52 24 67 Greece Aegean Sepia officinalis 394 372 325 431 450 636 707 679 734 598 709 516 463 518 644 Greece Ionian Sepia officinalis 34 27 28 21 43 51 58 64 46 29 46 81 46 70 Israel Levant Cephalopoda Italy Adriatic Eledone spp 1370 960 1412 966 863 613 557 549 1405 789 687 885 1032 2404 1071 Italy Ionian Eledone spp 490 470 578 1220 1393 1401 1246 1260 1236 1101 862 946 990 1115 772 Italy Sardinia Eledone spp 1220 1310 1469 464 425 598 694 560 493 485 521 201 564 791 881 Italy Adriatic Loligo spp 2117 1830 1658 1231 1672 1637 1426 1721 1579 1680 1653 930 837 879 930 Italy Ionian Loligo spp 626 600 585 1405 1758 1819 1636 1542 1641 1647 1741 2744 1683 1436 2244 Italy Sardinia Loligo spp 2469 2420 2552 1298 1404 1336 1364 1234 862 953 1164 657 608 1215 1466 Italy Adriatic Octopus vulgaris 1756 1234 1822 1028 1095 1113 1155 1260 1136 1071 1095 1344 1444 1202 949 Italy Ionian Octopus vulgaris 1792 1705 2095 4835 4954 5431 4980 4388 4678 4404 5035 5861 5409 4678 5253 Italy Sardinia Octopus vulgaris 4574 4925 5522 2285 2240 2658 3255 3070 3160 2838 2738 1828 2072 3449 2938 Italy Adriatic Sepiidae, Sepiolidae 7016 6850 8760 6341 7503 6614 5039 5297 3733 4234 8773 6533 7224 8392 6660 Italy Ionian Sepiidae, Sepiolidae 2113 1919 2209 4125 4150 3706 3694 3620 3415 3235 3282 1146 1509 2728 3425 Italy Sardinia Sepiidae, Sepiolidae 2952 2851 3310 1551 1559 1500 1933 1532 1124 1556 1515 3276 3831 1675 1568 Italy Adriatic Todarodes sagittatus 452 391 354 517 479 550 698 1412 498 576 903 2315 912 1788 2458 Italy Ionian Todarodes sagittatus 328 315 307 1047 1172 1229 1352 1561 1535 1375 1131 1011 1539 1272 2327 Italy Sardinia Todarodes sagittatus 1563 1535 1615 900 1028 914 975 1074 913 828 837 297 664 860 945 Lebanon Levant Octopus vulgaris Lebanon Levant Sepiidae, Sepiolidae

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Table 1 (continued 2/4) Country area Species 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 Malta Ionian Loligo spp 6 5 3 5 5 3 3 1 1 2 1 Malta Ionian Octopodidae 100 39 34 33 29 28 17 13 8 7 17 22 Sepiidae, Malta Ionian Sepiolidae 2 3 18 16 5 9 10 13 8 5 10 Todarodes Malta Ionian sagittatus Loliginidae, Morocco Balearic Ommastrephidae 1412 Morocco Balearic Octopodidae 226 529 Sepiidae, Morocco Balearic Sepiolidae 23 53 100 64 62 65 47 48 97 174 52 124 98 142 Slovenia Adriatic Loligo spp Slovenia Adriatic Octopus vulgaris Slovenia Adriatic Sepia officinalis Spain Balearic Cephalopoda Spain Gulf of Lions Cephalopoda Loliginidae, Spain Balearic Ommastrephidae 100 124 143 150 26 Spain Balearic Loligo spp 2502 2619 1200 1000 1100 1236 972 1024 1958 1719 1835 980 1125 1192 1200 Spain Gulf of Lions Loligo spp Spain Balearic Octopodidae 4066 4233 5000 4300 4200 4590 5441 4871 3947 3003 4717 4650 6082 6628 5918 Spain Gulf of Lions Octopodidae Spain Balearic Sepia officinalis Spain Gulf of Lions Sepia officinalis Sepiidae, Spain Balearic Sepiolidae 2356 2897 2500 1700 1500 1282 1485 1571 2038 1526 1884 1220 1273 1419 1350 Sepiidae, Spain Gulf of Lions Sepiolidae Todarodes Spain Balearic sagittatus Todarodes Spain Gulf of Lions sagittatus Tunisia Ionian Cephalopoda 1030 1040 1721 1718 1364 Tunisia Sardinia Cephalopoda 39 70 118 91 Tunisia Ionian Eledone spp Tunisia Sardinia Eledone spp Tunisia Ionian Loligo spp 78 73 100 100 104 86 157 301 288 394 375 142 159 133 158 Tunisia Sardinia Loligo spp 9 5 18 10 22 17 6 9 10 12 5 2 Tunisia Ionian Octopus vulgaris 86 176 198 800 600 2082 2161 3248 3619 3208 3978 2799 2202 3290 4812 Tunisia Sardinia Octopus vulgaris 19 27 54 100 100 62 40 64 56 46 44 20 20 38 39 Tunisia Ionian Sepia officinalis 290 239 258 615 435 633 2048 819 1009 1205 985 1277 2214 1689 2433 Tunisia Sardinia Sepia officinalis 46 42 82 100 106 134 125 145 124 113 130 111 160 140 126 Turkey Aegean Loligo spp 25 100 100 50 47 24 5 16 13 49 17 161 138 Turkey Levant Loligo spp 26 2 17 6 5 22 43 78 28 4 152 Turkey Aegean Octopus vulgaris 76 10 10 23 19 12 19 30 72 35 127 218 Turkey Levant Octopus vulgaris 11 9 4 8 5 6 23 50 25 10 10 Turkey Aegean Sepia officinalis 59 46 19 43 28 62 16 162 184 Turkey Levant Sepia officinalis 300 400 300 200 154 170 121 132 217 205 55 12 184 Turkey Black Sea Octopus vulgaris Turkey Marmara Sea Octopus vulgaris 1 1 1 Turkey Black Sea Sepia officinalis Turkey Marmara Sea Sepia officinalis 131 1 Turkey Black Sea Loligo spp 1 Turkey Marmara Sea Loligo spp 3 Yugoslavia SFR Adriatic Loligo spp 234 239 210 191 271 263 207 263 298 300 264 221 228 260 253 Yugoslavia SFR Adriatic Octopus vulgaris 253 218 132 99 98 85 107 133 132 122 107 106 114 113 100 Yugoslavia Sepiidae, SFR Adriatic Sepiolidae 151 157 186 171 206 233 261 208 182 150 213 170 156 188 164 Yugoslavia , Fed. Rep. of Adriatic Loligo spp Yugoslavia , Fed. Rep. of Adriatic Octopus vulgaris Yugoslavia , Fed. Rep. Sepiidae, of Adriatic Sepiolidae 4458 4465 4988 4426 4627 4702 4827 4767 4627 4385 5166 4710 4939 5529 5670 TOTAL 7 8 3 4 5 4 5 3 2 2 0 2 8 2 6

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Table 1 (continued 3/4)

Country area Species 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 Albania Ionian Loligo spp 7 47 34 93 93 Albania Ionian Octopus vulgaris 66 75 59 93 90 Albania Ionian Sepia officinalis 39 33 33 51 51 Algeria Balearic Cephalopoda 70 29 Algeria Balearic Loligo spp 331 250 200 250 250 300 101 53 138 202 240 Algeria Balearic Octopodidae 382 190 185 543 305 Algeria Balearic Sepia officinalis 424 400 350 400 500 600 351 567 619 492 312 Croatia Adriatic Cephalopoda 552 799 455 431 483 443 265 646 Croatia Adriatic Loligo spp 263 330 268 273 233 287 290 170 Croatia Adriatic Octopus vulgaris 155 231 187 193 180 293 435 Croatia Adriatic Sepiidae, Sepiolidae 167 203 118 118 102 151 215 145 Cyprus Levant Octopodidae 101 67 108 119 201 208 170 258 288 474 300 190 199 228 179 Cyprus Levant Sepia officinalis 80 90 92 95 79 108 106 103 174 218 153 111 89 146 140 Egypt Levant Sepiidae, Sepiolidae 640 1030 724 941 631 614 659 787 1067 1031 1097 1365 1370 1152 1449 France Gulf of Loliginidae, Lions Ommastrephidae 112 66 107 86 86 67 43 43 43 France Sardinia Loliginidae, Ommastrephidae 28 France Gulf of Loligo spp Lions 325 311 277 278 244 262 262 261 220 136 150 135 465 465 222 France Sardinia Loligo spp France Gulf of Octopus vulgaris Lions 647 806 659 643 1163 1400 1250 1196 1161 873 1146 706 439 439 1404 France Sardinia Octopus vulgaris France Gulf of Sepia officinalis Lions 209 258 236 213 191 259 193 253 151 130 100 85 85 85 75 France Sardinia Sepia officinalis 30179211111 France Gulf of Todarodes sagittatus Lions 38 74 75 49 11 21 19 87 87 87 France Sardinia Todarodes sagittatus 10 6 23 Gaza Levant Loligo spp Strip(Palestine) 15 23 30 61 60 Gaza Levant Sepiidae, Sepiolidae Strip(Palestine) 41 62 98 144 145 Greece Aegean Loliginidae, Ommastrephidae 209 234 283 282 381 470 443 693 546 586 788 679 472 405 505 Greece Ionian Loliginidae, Ommastrephidae 2722391031021 92 118 133 41 161 170 Greece Aegean Loligo spp 430 461 555 631 707 595 1015 1026 1258 897 794 680 478 321 267 Greece Ionian Loligo spp 47 21 33 47 72 73 49 109 111 204 151 176 145 105 130 Greece Aegean Octopodidae 669 632 515 433 285 425 463 693 747 1048 1352 697 658 605 662 Greece Ionian Octopodidae 28 9 2 12 13 5 26 15 33 118 99 112 82 127 132 Greece Aegean Octopus vulgaris 430 597 605 810 789 1270 2256 3479 2303 2691 2666 2765 2788 1575 1797 Greece Ionian Octopus vulgaris 29 27 26 20 45 86 40 84 153 195 190 154 164 98 113 Greece Aegean Sepia officinalis 808 1173 997 1477 1434 1918 1663 1717 1840 2572 2292 1630 2222 1691 2606 Greece Ionian Sepia officinalis 39 29 29 37 49 78 73 63 101 281 224 189 159 116 126 Israel Levant Cephalopoda 95 87 91 76 68 64 50 50 50 98 76 120 Italy Adriatic Eledone spp 1515 1065 1382 1753 721 862 927 1643 1748 1915 1067 646 524 573 432 Italy Ionian Eledone spp 832 742 525 1071 1130 1146 1553 1202 1081 1228 778 508 590 423 292 Italy Sardinia Eledone spp 1324 941 963 738 619 964 886 787 713 785 655 785 476 510 317 Italy Adriatic Loligo spp 1349 1273 1052 866 847 941 963 830 654 1288 1179 873 1425 599 474 Italy Ionian Loligo spp 2306 2404 2366 2333 2358 2658 3076 2846 2640 3152 3204 2997 1524 1006 820 Italy Sardinia Loligo spp 1575 1773 1934 1559 1019 1144 1093 1052 1078 842 1100 1496 1192 632 615 Italy Adriatic Octopus vulgaris 957 1040 982 847 701 996 2016 2023 2295 1891 1909 1619 1715 1220 1044 Italy Ionian Octopus vulgaris 5318 6548 5291 5728 5976 6886 6102 5636 6320 4627 5051 4555 4513 4716 3012 Italy Sardinia Octopus vulgaris 3144 3408 3911 3779 2468 2281 2390 2641 2950 3143 2797 2883 2679 3542 2943 Italy Adriatic Sepiidae, Sepiolidae 1001 8419 7468 5882 6068 4237 4190 5780 3816 4112 4 5537 3929 3904 3525 3280 Italy Ionian Sepiidae, Sepiolidae 3932 5003 2820 3668 2949 2556 2920 2708 2371 2321 3944 2584 2246 2218 1305 Italy Sardinia Sepiidae, Sepiolidae 2003 2120 2736 3681 1548 1545 1406 1460 1199 1281 1669 1461 1232 1096 1135 Italy Adriatic Todarodes sagittatus 3383 3291 3544 3409 4100 4048 2860 3389 2760 1866 836 910 554 534 579 Italy Ionian Todarodes sagittatus 2736 3333 2669 2685 3182 3179 3521 3519 2893 2918 3076 2857 1375 2983 865 Italy Sardinia Todarodes sagittatus 883 1115 1199 1143 961 988 1037 892 798 953 877 905 685 478 612 Lebanon Levant Octopus vulgaris 25 25 Lebanon Levant Sepiidae, Sepiolidae 25 25 25 20 25 25 25 25 25 25 50 25 50

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Table 1 (continued 4/4) Country area Species 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 Malta Ionian Loligo spp 18 5 1 1 1 1 2 2 2 2 Malta Ionian Octopodidae 38 7 9 6 10 4 2 3 3 4 6 11 11 9 11 Malta Ionian Sepiidae, Sepiolidae 57 4 4 4 6 2 2 1 1 5 3 3 5 Malta Ionian Todarodes sagittatus 2 2 Loliginidae, Morocco Balearic Ommastrephidae 20 13 11 28 63 17 37 40 25 122 32 10 2 2 3 Morocco Balearic Octopodidae 717 492 205 80 98 8 72 78 71 25 69 36 98 68 115 Morocco Balearic Sepiidae, Sepiolidae 157 141 273 81 97 129 153 205 215 139 265 178 162 132 157 Slovenia Adriatic Loligo spp 4 4 4 2 2 4 3 2 Slovenia Adriatic Octopus vulgaris 11 25 6 34 2 7 Slovenia Adriatic Sepia officinalis 12 21 4 10 6 5 18 18 Spain Balearic Cephalopoda 470 347 20 Spain Gulf of Lions Cephalopoda 20 15 Loliginidae, Spain Balearic Ommastrephidae 592 547 747 673 Spain Balearic Loligo spp 928 1099 1040 860 796 750 750 700 700 670 670 486 443 391 997 Spain Gulf of Lions Loligo spp 205 175 210 150 150 100 100 30 30 19 19 14 5 Spain Balearic Octopodidae 7111 6770 4354 4965 5477 5500 5500 5490 5490 5490 5490 5386 4747 4904 5461 Spain Gulf of Lions Octopodidae 1273 1160 96 100 100 210 210 210 210 208 151 230 104 Spain Balearic Sepia officinalis 496 437 474 983 Spain Gulf of Lions Sepia officinalis 3392 Spain Balearic Sepiidae, Sepiolidae 1409 1239 900 789 894 1300 1200 1100 1000 900 800 746 780 834 446 Spain Gulf of Lions Sepiidae, Sepiolidae 133 130 460 Spain Balearic Todarodes sagittatus 51 130 130 102 136 Spain Gulf of Lions Todarodes sagittatus 10 19 17 8 Tunisia Ionian Cephalopoda Tunisia Sardinia Cephalopoda Tunisia Ionian Eledone spp 660 185 305 Tunisia Sardinia Eledone spp 8 67 87 Tunisia Ionian Loligo spp 143 137 233 238 172 116 115 147 166 239 197 187 207 150 182 Tunisia Sardinia Loligo spp 2 15 14 13 43 42 54 48 38 43 43 46 138 166 Tunisia Ionian Octopus vulgaris 5150 4879 8025 12256 9097 6390 6265 7459 3274 1787 1802 3383 5609 3097 2306 Tunisia Sardinia Octopus vulgaris 42 548 300 39 38 27 72 85 76 54 66 77 72 32 43 Tunisia Ionian Sepia officinalis 3808 3765 6410 6040 6499 5825 5689 6825 6156 4948 3300 4085 6212 4595 6264 Tunisia Sardinia Sepia officinalis 197 422 12 151 94 84 190 228 159 173 217 255 267 340 358 Turkey Aegean Loligo spp 183 360 358 1003 3393 2702 180 321 100 374 129 209 188 387 146 Turkey Levant Loligo spp 139 41 40 839 730 452 87 236 280 199 197 149 200 109 210 Turkey Aegean Octopus vulgaris 240 242 406 993 1574 362 299 468 240 495 426 687 758 1334 434 Turkey Levant Octopus vulgaris 11 14 24 52 457 87 37 107 154 145 170 101 233 100 72 Turkey Aegean Sepia officinalis 178 185 261 1793 2279 2059 187 153 224 426 355 240 396 396 217 Turkey Levant Sepia officinalis 78 114 161 1723 3031 2073 162 457 290 290 570 387 492 354 313 Turkey Black Sea Octopus vulgaris 3 3 Marmara Turkey Sea Octopus vulgaris 1 18 30 687 102 33 1 78 19 6 14 9 16 1 Turkey Black Sea Sepia officinalis 50 1 5 1 Marmara Turkey Sea Sepia officinalis 5 8 1084 56 231 12 1 8 17 7 6 Turkey Black Sea Loligo spp 2 2 Marmara Turkey Sea Loligo spp 7 7 1556 78 17 4 5 6 32 4 2 Yugoslav ia SFR Adriatic Loligo spp 353 228 197 205 269 374 214 Yugoslav ia SFR Adriatic Octopus vulgaris 150 154 159 179 180 211 121 Yugoslav ia SFR Adriatic Sepiidae, Sepiolidae 159 154 190 235 214 296 169 Yugoslav ia, Fed. Rep. of Adriatic Loligo spp 4 10 7 13 12 13 13 12 Yugoslav ia, Fed. Rep. of Adriatic Octopus vulgaris 2 6 4 4 8 8 9 9 Yugoslav ia, Fed. Rep. of Adriatic Sepiidae, Sepiolidae 2 6 5 9 10 9 10 10 6572 6840 6771 7694 7247 6783 7171 6544 6867 6254 5926 6013 5428 5048 TOTAL 3 3 3 83346 5 1 1 3 7 6 5 6 9 1 2

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WORKING DOCUMENT N° 4

ICES Working Group on Cephalopod Fisheries and Life-History

Rome, Italy 10-12 April 2002

Experimental study of the effect of enriched frozen diet on survival and growth of juvenile cuttlefish Sepia officinalis L.

by

Dr. Noussithé Koueta Université de Caen Laboratoire de Biologie et Biotechnologies Marines I.B.B.A. 14032 Caen Cedex France [email protected]

Abstract

Juvenile cuttlefish hatched in the laboratory were reared during 10 days with live shrimps enriched in polyunsaturated fatty acids (PUFA), then divided in 4 dietary groups fed respectively on live shrimps, frozen shrimps, fish oil enriched frozen shrimps and fish protein hydrolysate (Gabolysat) enriched frozen shrimps during 20 days. The group fed on fish oil enriched frozen shrimps exhibited a high rate of survival. During the rearing, enriched frozen diet maintained weight increase as in group fed on live shrimps Although the low ration observed in the group fed on fish oil enriched frozen shrimps during 20 days, the food conversion rate was as in other groups and the weight gain was better than in group fed on fish protein hydrolysate (Gabolysat) enriched frozen shrimps. From 20 to 30 days old, relative weight gain was higher and the food conversion rate was better in juvenile cuttlefish of the group fed on fish protein hydrolysate enriched frozen diet.

These data indicate that the quality of the diet enrichment during juvenile cuttlefish rearing depends on the age of the animal PUFA appears very important until 20 days old. They need PUFA during their youngest stage. After that age the proteins are more important in juvenile cuttlefish nutrition.

Keywords: Conversion rate; Diet; Fish oil; Fish protein hydrolysate Frozen diet; Gabolysat; Growth; Juvenile cuttlefish; PUFA; Relative weight gain; Survival; Weight increase.

Introduction

The culture of cephalopods is becoming an interesting area due to their fast growth (Mangold and Boletzky, 1973; Mangold, 1983; Forsythe and Heukelem, 1987), their scientific importance in electrophysiology, behaviour science biomedical and environmental sciences, endocrinology (Agin et al. 2001 Bustamente et al, 2002; Di Cosmo et al. 2002 Kashimura et al.2001 Dickel et al. 2001 Miyazaki et al. 2001; Tosti et al., 2001) an and their commercial value (Navarro and Villanueva, Koueta and Boucaud 2001). Juvenile as well as mature cuttlefish are characterised by a predatory diet but only subadults and adults can be reared on artificial diet.(Castro et al. 1993, Koueta and Boucaud- Camou, 1999). In cephalopods rearing, paralarval and juvenile stage culture remains still difficult (Villanueva et al 2002;). Actually relative growth up animals have been obtained when juvenile cuttlefish were reared with alternative diets but the young animals in these trials were fragile and their growth rate was low (Choe, 1966; Boletzky and Hanlon, 1983; Boletzky, 1989; DeRusha et al., 1989; Koueta and Boucaud-Camou, 1999). The variability in the quality and quantity of food greatly affect early growth

The effectiveness of PUFA enrichment of natural live prey on juvenile cuttlefish growth has been demonstrated (Koueta et al. 2002). But an balanced inert food to rear the early stage could be essential for cephalopod aquaculture The aim of

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this study was to test the effect of enriched inert food on juvenile cuttlefish survival and growth. In order to formulate artificial food according to the age of the animals different quality of diet were used

Materials and methods

1- Eggs origin

All the eggs were laid in the laboratory by sexual mature females trawled of Normandy coast and maintained in a large tank receiving water from the sea.

2- Animals.

The eggs were placed on floating sieves distributed in tank connected to the semi-closed system as previously described (Koueta and Boucaud-Camou, 1999). As hatching lasted several days, hatchlings were placed in the small tanks of 707 cm2 in groups of ten animals according to their age and fed ad libitum on live young shrimps.

3- Rearing system.

Culture system, filtration system, water circulation system, water oxygenation system, light and temperature, stocking and culture densities were as previously described (Koueta and Boucaud-Camou, 1999). The pH was 8, salinity 35.5%0, the concentration of O2 measured with electronic oxymeter was excellent, (8.9-9.7 ppm corresponding to a saturating + rate of 101 to 110%. NH4 measurements using a colorimetric kit was < 0.5mg/l then NH3 concentration < 0.02 mg/l at the pH of rearing. For nitrites and nitrates the rates using a colorimetric kit were respectively <0.1 mg/l and < 10 mg/l. Physical and chemical parameters were maintained constant by avoid surpopulation, dead cuttlefish, dead prey, food remains poor oxygenation, disinfection , washing of bacterial support, or no renewal of sea water.

Before entering the system the natural see water which is used to renew the circuit runs through a system of U.V. lamps with a flow rate of 60 l/h (93% renewal per day). The mechanical filters, which consist of foam and synthetic fibres were cleaned daily only with sea water to avoid lethal osmotic choc on nitrite bacteria. The temperature of sea water is maintained between 19.5°C - 20.5°C by the heating elements in the inferior conditioning tank. The rearing device receives 12 h of light/ 24 h.

4 - Experiment

A total of 160 juvenile cuttlefish of maximum 2 days old were selected measured weighed, and randomly distributed in small tanks. Each tank contained four animals well separated by a thick partition. The cuttlefish were divided into 4 groups of 40 animals receiving respectively young shrimps

The fish oil and fish powder used were provided by Dielen Laboratoires and contained respectively 18% EPA, 12% DHA of total fatty acids, 1mg/g of vitamin A, α-tocophérol (anti oxydant) for the fish oil, 74% of protein, 9% of lipids containing 12.6% EPA and 21% DHA of total fatty acids for the fish powder.

5 - Prey enrichment

Fresh surimi of shrimp were impragnated during 24 hours at 4°C in Dielen laboratoires oil fish then distributed to young Crangon crangon at the rate of 5 g of surimi for 10 g of the live prey. During the first experiment we have used progressively 1 to 5 g of enriched surimi to involve significative growth in juvenile cuttlefish.

The enrichment of Crangon crangon with fish powder was carried up at the rate of 500 mg incorporated in 2 g of fresh shrimp surimi then used as the Crangon crangon diet.

For mysid enrichment, the fish powder was dissolved in sea water at the rate of 250 mg in 5 liters of sea water for 150 mysids ( Mysidopsis almyra and Shistomysis sp).

Compressed air was introduced into enrichment tanks from a number of narrow pipes opening on bubblers and these tanks were cleaned daily to avoid pollution involved by over feeding or the quality of the diets. The live preys (Crangon crangon and mysids) were kept with enriched food during 24 hours.

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6 - Feeding methodology

According to the diet previously decribed, the prey were offered once a day at 10 a.m. The daily ration (maximum ration) for juvenile cuttlefish as observed by Koueta and Boucaud-Camou (2001) was 40 % of animal body weight and involved maximum growth. The daily ration was adjusted according to animal weight after 5 or 10 days during 30 days of rearing

7 - Growth and food conversion rate analysis

The amount of food ingested by the specimens in each container was measured by weighing each day the food remaining in the individual tanks.

The feeding rate (FR) was expressed as FR (% Body weight / day) = (FI / mean Wt x 100), where FI is the weight of food ingested by each animal (w) during the time (t) of the experiment.

The conversion efficiency (%) was calculated as growth weight/weight of food ingested x 100.

Weight and length measurements were as described Koueta and Boucaud-Camou (1999). Weight growth (mg) = Final weight- initial weight.

Length growth (mm) = final length - initial length.

Relative gain of weight (%) = weight growth x 100/ initial weight

Growth rate was expressed as “ Instantaneous relative Growth Rate ” (IGR).

IGR (% Body weight / day) = (Ln W2 – Ln W1) x 100 / t, W2 and W1 are, respectively, final and initial weight (mg) of each animal, and t the duration of the experiment in days.

Speed of weight growth (mg/day) = weight growth/day

Results

1 – Survival (figure 1)

The survival was better in group fed on enriched frozen diet during the first 20 days than in other groups. From 20 to 30 days, the survival was stable in all groups. The difference remains the one observed during the 20 first days. The survival observe n the group fed on live shrimps was low because some competition between live shrimps and juvenile cuttlefish.

2 – Weight increase (figure 2)

The weight increase was high in the groups fed on frozen shrimps and the group fed on fish oil enriched frozen shrimps, but the difference was not significant.

3- Relative weight gain (figure 3).

The relative weight gain were better in the group fed on frozen shrimps and the group fed on fish oil enriched shrimps. After 20 days old the relative weight gain increase in the group fed on fish protein hydrolysate enriched frozen diet than in other groups.

3- Ration on different groups (figure 4)

At 15 days old, the ration was high in the group fed on frozen shrimps than in other groups. After 30 days old the ration was low in the groups fed on fish oil enriched shrimp and the group fed on fish protein hydrolysate enriched frozen shrimp but more in this group.

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4 – Food conversion rate (figure 5)

At 15 days old the food conversion was low in the group fed on frozen shrimp than in other 3 groups. After 30 days old the conversion rate increase in group fed on fish protein hydrolysate enriched shrimp than in 3 other groups. The conversion rate was high in groups fed on fish oil enriched frozen shrimp than in the group fed on live shrimp and the group fed on frozen shrimp.

Conclusion

This experiment confirm ours previous observations that frozen diet can be used to rear juvenile cuttlefish previously fed during the 10 first days of their live with enriched natural diet.

During rearing, frozen diet and enriched frozen diet maintained weigh increase as in group fed on natural diet when juvenile cuttlefish were previously fed on enriched live diet. This data suggest that enrichment are essential to assume well growth when fed juvenile cuttlefish on frozen diet

The relative weight gain and conversion rate were better after 20 days old in the group fed on fish protein hydrolysate and suggest the importance of protein in growth at that age

The ration was high in the group fed on frozen diet but the conversion rate was low during all the rearing. In the group fed on fish oil enriched shrimp, the ration was low but the conversion rate was than in the groups fed on live diet and the group fed on fish protein hydrolysate enriched shrimp during 20 days. These observations suggest that fish oil enrichment is better during that period of live.

The low ration and high conversion rate observe in the group fed on fish protein enriched shrimp after 20 days old suggest that this diet was better for juvenile cuttlefish rearing.

The quality of the enrichment depend on the age of juvenile cuttlefish. Fatty acids appears very important until 20 days old during juvenile cuttlefish survival and growth. They need PUFA (polyunsaturated fatty acids) during their youngest growth. After 20 days old protein are more important for juvenile cuttlefish growth. The impact of the quality of enriched frozen diet on digestive capability of juvenile cuttlefish must be elucidated in order to explain the physiological effect of different diets on digestion and growth.

References

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Boletzky S. V. Et Hanlon R. T., 1983. A review of the laboratory maintenance, rearing and culture of cephalopods mollusks. Memoirs of the national museum, Victoria 44 : 147-187.

Boletzky S. V., 1989. Elevage de céphalopodes en aquarium : acquis récents. Bulletin de la société zoologique de France. 1989. 114. N° 4 : 31-40.

Bustamante P., Cosson Rp, Gallien I., Caurant F. Et Miramand P. 2002. Cadmium detoxification processes in the digestive gland of cephalopods in relation to accumilated cadmium concentrations. Marine Environmental research, 53: 227-241.

CASTRO, B.G., Di MARCO, F.P., Derusha, R.H. And LEE, P.G., 1993. The effects of surimi and pelleted diets on the laboratory survival, growth, and feeding rate of the cuttlefish. Sepia officinalis L. Journal of Experimental Marine Biology and Ecology 170: 241-252.

Choe, S. 1966. On the growth, feeding rates and the efficiency of food conversion for cuttlefishes and squids. Korean Journal of Zoology 9: 72-80.

Dickel L., Chichery M.P. Et Chichery R. 2001. Increase of learning abilities and maturation of the vertical lobe complex during post-embryonic development in the cuttlefish, Sepia. Developmental psychobiology, 39(2): 92- 98.

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Di Cosmo A., Dicristo C. Et Paolucci M. 2002. A estradiol-17 beta receptor in the reproductive system of the female of Octopus vulgaris: Characterization and immunolocalisation. Molecular Reproduction and Development, 61(3): 367-375.

Derusha, R.H., FORSYTHE, J.H., Dimarco, F.P. And. HANLON, R.T., 1989. Alternative diets for maintaining and rearing cephalopods in captivity. Laboratory Animal Science 4, 306-312.

Forsythe, J. W. And Van Heukelem, W.F. 1987. Growth. In: Boyle, P.R. (Ed), Cephalopd Life Cycle, Vol. 2. Academic press, London,pp. 351- 365.

Kishimura H., Saeki H Et Hayashi K. 2001. Isolation and characteristics of trypsin inhibitor from the hepatopancreas of a squid (Todarodes pacificus). Comparative Biochemistry and Physiology,B, 130(1): 117-123.

Koueta.N.And Boucaud-Camou E., 1999. Food intake and growth in reared early juvenile cutlefish Sepia officinalis L. (Mollusca Cephalopoda) J. Exp. Mar. Biol. Ecol. 240: 93-109.

KOUETA.N.And BOUCAUD-CAMOU E. (2001)Basic growth relations during rearing of early juvenile cuttlefish Sepia officinalis L (Mollusca Cephalopoda). J. Exp. Mar. Biol. Ecol., 265 : 75-87

Koueta, N., Boucaud-Camou, E. And Noël, B. (2002) Effects of enriched natural diet on survival and growth of juvenile cuttlefish Sepia officinalis L. Aquaculture. 203 : 293-310

Mangold K. 1983 Octopus vulgaris : In : Boyle, P R (Ed), Cephalopod Life Cycle, vol. 1. Academic Press, London,pp.335-364

Mangold, K.M., And Boletzky .S.V. 1973. New data on reproductive biology and growth of Octopus vulgaris. Mar. Biol. 19, 7-12

Miyazaki T., Nakahara M., Ishii T., Aoki K. Et Watabe T 2001. Accumulation of cobalt in newly hatched octopus Octopus vulgaris. Fisheries Research, 67(1): 170-172.

Navarro, J.C. And Villanueva, R. 2001. Lipid and fatty acid composition of early stages of cephalopods: an approach to their lipid requirements. Aquaculture 183, 161-177

Tosti E., Di Cosmo A., Cuomo A., Dicristo C. Et Gragnaniello G. 2001. Progesterone induces activation in Octopus vulgaris spermatozoa. Molecular Reproduction and Development, 59(1): 97-105.

Villanueva, R., Koueta, N., And Boucaud-Camou, E. (2002) Growth and proteolytic activity of Octopus vulgaris paralarvae at different food rations during first feeding, using Artemia nauplii and compound diets Aquaculture 205 269-286

Figures legends

Figure 1. Effect of enriched frozen diet on survival (%) during juvenile cuttlefish growth.

Figure 2. weight increase(mg) during juvenile cuttlefish growth. Effect of different diets

Figure 3 Effect of different diets on relative weight gain of juvenile cuttlefish during rearing (%)

Figure 4. Comparison of the ration between the group fed on frozen diet and others groups (% of body weight/day)

Figure 5. Changes of the conversion rate in different feeding groups during growth of juvenile cuttlefish (%)

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Figure 1: Effect of enriched frozen diet on survival during juvenile cuttlefish growth

100 98 96

l 94 a v

i 92 v r 90 u 88 % s 86 84 82 80 10 15 20 25 30 age (days) live shrimp frozen shrimp oil fish enriched frozen shrimp fish protein hydrolysate enriched frozen shrimp

Figure 1. Effect of enriched frozen diet on survival (%) during juvenile cuttlefish growth.

Figure 2

180

g) 160 m

( 140 e 120

eas 100 r c

n 80 i 60 ght i 40 e

w 20 0 10 15 20 25 30 age (days) live shrimp frozen shrimp fish oil enriched frozen shrimp fish hydrolysate enriched frozen shrimp

Figure 2. weight increase(mg) during juvenile cuttlefish growth. Effect of different diets

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Figure 3 90 )

% 80 (

in 70 a 60 g t

h 50 g i

e 40

w 30 e v i

t 20 la

e 10 r 0 10 15 20 25 30 age (days) live shrimp frozen shrimp fish oil enriched frozen shrimp fish protein hydrolysate enriched frozen shrimp

Figure 3 Effect of different diets on relative weight gain of juvenile cuttlefish during rearing (%)

Figure 4

35 ay) d / 30 ght i

e 25 w

y 20

15 bod of 10 % 5 on ( i 0

Rat 10 15 20 25 30 age (days) live shrimp frozen shrimp fish oil enriched frozen shrimp fish protein hydrolysate enriched frozen shrimp

Figure 4. Comparison of the ration between the group fed on frozen diet and others groups (% of body weight/day)

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Figure 5:

80 e t a

r 60 n o si

) 40 ver (%

n 20 co

d 0 o o f -20 10 15 20 25 30 age (days) live shrimp frozen shrimp fish oil enriched frozen shrimp fish protein hydrolysate enriched frozen shrimp

Figure 5. Changes of the conversion rate in different feeding groups during growth of juvenile cuttlefish (%)

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WORKING DOCUMENT N° 5

ICES Working Group on Cephalopod Fisheries and Life-History

Rome, Italy 10-12 April 2002

ASSESSMENT OF THE ENGLISH CHANNEL CUTTLEFISH STOCK BASED ON ANALYTICAL METHODS.

Royer J. and Robin J.P.

Université de Caen, Laboratoire de Biologie et Biotechnologies Marines, Esplanade de la Paix,14032 Caen cedex, France [email protected]

Introduction

The English Channel area is the most important fishing ground for cuttlefish in Europe with average landings above 10500 T in the period 1995-1998 (Anonymous, 2001). Landings from this area consist exclusively of Sepia officinalis and the 2-year life cycle of English Channel population has been described by previous authors (Boucaud-Camou et al, 1991; Boucaud-Camou and Boismery, 1991).

Preliminary trials to estimate English Channel cuttlefish stock size have been carried out using depletion methods. However, Dunn's assessments (1999) considered only the UK fishery and Kelly's work (1999) underlined difficulty to obtain abundance indices (CPUE) reliable in juveniles as well as in adults.

This paper presents Cohort Analysis applied to cuttlefish cohorts exploited between 1995 and 1998. Fish market sampling for length frequencies is used to analyse the mixing of annual cohorts. Estimates of recruitment and fishing mortality at age are used to understand the exploitation level of 1995 and 1996 cohorts according to Thomson and Bell simulations.

Materials and Methods

Stock boundaries

The studied stock is limited to the English Channel stricto sensu (ICES Divisions 7D and 7E).. Although cuttlefish are observed in the Southern North Sea (division 4C) and to the West of the English Channel (7H) abundance indices are much lower in these areas than in the English Channel (Denis, 2000). Genetic studies revealed little difference between English Channel and Bay of Biscay populations (Robin et al, 2000) and exchanges can happen on wintering grounds off Finisterre (Denis et Robin, 2001). The rationale for considering an English Channel stock is related to the availability of comprehensive landing statistics on a monthly basis for this area. UK and France monthly landings were collected and they represent more than 99% of the English Channel cuttlefish yield (Anonymous, 2001).

Fishery Data

National data bases were interrogated for records of total cuttlefish landings (kg). Scotland (SOAEFD data base) and England and Wales (CEFAS data base) are referred to in the text as UK data. In the case of France (Centre Administratif des Affaires Maritimes data base) landings are sorted out per commercial category (4 categories based on individual weight) a detail which was also collected. In both countries (UK and France) Fishery Statistics are available on an ICES rectangle basis.

In addition "non-official" records from the French inshore fishery were obtained through the "Comité Régional des Pêches Maritimes". These records mainly concerned catches by cuttlefish traps.

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Biological sampling

Catch structure is based on the sampling of bottom trawlers landings at the Port-en-Bessin fish market and cuttlefish trap landings at Granville fish market. On each sampling day two cuttlefish boxes (25 kg boxes, generally taken among the landings of two different boats) were sampled in each commercial category. Traps catch only adult cuttlefish which belong to commercial category 1.

Length-frequencies were used to distinguish age groups using the Normsep module implemented in the FISAT package (Gayanilo et al, 1995). In each annual cohort, two age groups were considered: the first one representing early-hatched or (fast growing animals) which are recruited in autumn and the second one late-hatched (or slower growing) animals which enter the fishery in spring of the next year. It is worth noting that recruits enter the stock at a similar size (6-8 cm DML) but at a different age for autumn recruits and spring recruits. This decomposition of the catch in numbers by age- groups is not as accurate as in the analysis of Loliginid squid where monthly ages were considered (Royer et al, 2002). This is mainly due to the lack of accurate growth curve available in this cuttlefish population. In Loliginid squid growth curves were derived from statolith ageing (Natsukari and Komine, 1992; Collins et al, 1995) whereas in English Channel cuttlefish growth was only described with modal progression (Medhioub, 1986).

Analytical modelling (Cohort analysis and Thomson and Bell simulations)

The methodology applied to English Channel cuttlefish cohorts is the same as age-based models developed in Loliginid squid populations (Royer et al, 2002).

Cohort analysis is used to back-calculate the history of each cohort (survivors and fishing mortalities). An empirical estimate of natural mortality (M = 0.1) was derived from Caddy's method (1996) and terminal fishing mortalities were optimised iteratively.

Yield and average biomass for the 1995 and 1996 cohorts at various levels of fishing mortality were estimated with the classical Thomson and Bell model (Sparre and Venema, 1998) in which number of recruits and fishing mortalities estimated with cohort analysis are used to simulate yield and biomass. Mean weights-at-age are assumed to be constant whatever the multiplier factor of fishing mortalities.

Results & Discussion

Total catch structure (figure 1).

Monthly landings in numbers by age-group show that the number of juvenile exploited in the first autumn of a cohort life-span is variable (in cohorts 1995-1998 the maximum is 4.3 and the minimum 2.8 million animals, observed in 1995 and 1996 respectively). In the two cohorts that were entirely exploited (1995 and 1996) more than 1/3 of the catch occured during the second autumn. Adult catches, made during the second spring, represented nevertheless a significant proportion of each cohort catches (11 % and 15% in 1995 and 1996 respectively).

Stock size and recruitment

Cohort analysis provides monthly estimates of population numbers by age-group and values for the first exploited month correspond to recruitment (figure 2). The total number of recruits born in 1995 and born in 1996 was estimated at 64 and 47 million animals respectively.

Cohort analysis recruitment estimates can be compared with inter-annual trends in catch rate (CPUE). GLM indices applied to the commercial category of small cuttlefish have been derived from French trawlers catch and effort data (figure 3). (The analysis of trends in trawlers CPUE with GLM techniques was presented to ICES ASC: Denis et al, 2001). In spite of the low number of data points, it is worth noting that both methods give similar results.

Fishing mortalities

Mortality vectors averaged (across all age-groups) by calendar month indicate that fishing mortality peaks in spring in most years but 1996 (figure 4). Selection curves in 1995 and 1996 cohorts (figure 5) underline that fishing mortality in the second spring is much higher than in the first. At this stage, fishing mortalities already suggest that growth over- fishing on these two cohorts is rather unlikely.

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Production curves (figure 6)

Simulated yields according to a range of values for mf (multiplier of fishing mortality) confirm that the observed situation (mf = 1) corresponded to a full exploitation of both 1995 and 1996 cohorts. This is the first diagnostic of the effect of fishing on a European cuttlefish stock.

Future analyses

Although exploitation diagnostics for 1995 and 1996 cohorts do not reveal over-fishing, simulations can be developed to better understand interactions between fishing fleets. In particular, the effect of increased mortality on autumn recruits (which are caught by inshore trawlers) can be studied. The interaction between traps and trawlers catches corresponds also to a situation of conflict. Since cuttlefish traps only catch adults, its analysis will be limited by the lack of a spawning stock / recruitment relationship.

References:

Anonymous, 2001. Report of the Working Group on Cephalopod Fisheries and Life History. ICES CM 2001/G:04, 28pp + Annexes.

Boucaud-Camou E., Koueta N., Boismery J., Medhioub A. 1991. The sexual cycle of Sepia officinalis L. from the Bay of Seine. In Boucaud-Camou E. (Editor) "The cuttlefish", Act. 1st International Symposium on the Cuttlefish Sepia Caen, June 1-3 1989, Centre de Publications de l'Université de Caen, pp. 141-151.

Boucaud-Camou E., Boismery J. 1991. The migration of the cuttlefish (Sepia officinalis) in the English Channel. In: Boucaud-Camou E. Ed. The cuttlefish, Centre de publication de l'Université de Caen, 179-189

Caddy J.F., 1996. Modelling natural mortality with age in short-lived invertebrate populations: definition of a strategy of gnomonic time division. Aquat. Living Resour., 9, 197-207.

Collins, M.A., Burnell, G.M., and Rodhouse, P.G. 1995. Age and growth of the squid Loligo forbesi (Cephalopoda: Loliginidae) in Irish waters. Journal of Marine Biological Association of the United Kingdom, 75(3): 605-620.

Denis V., 2000. Variations spatio-temporelles d'abondance des céphalopodes exploités depuis les côtes atlantiques françaises et influence de parameters environnementaux. Ph D. Thesis, Université de Caen, France, 293 p.

Denis, V., and Robin, J.P. 2001. Present status of the French Atlantic fishery for cuttlefish (Sepia officinalis). Fisheries Research, 52:11-22.

Denis V., Royer J., Périès P., Wang J., Pierce G.J., Boyle P. R., Dunn M. R., Robin J.P. 2001. French and UK bottom trawl fisheries in the English Channel: spatial and temporal patterns for fishing effort and Cephalopod catch and integration of fleet components in the computation of squid and cuttlefish abundance indices. ICES CM 2001/K:08, (poster).

Dunn M., 1999. The exploitaton of selected non-quota species in the English Channel. Ph D. Thesis, University of Porsmouth, K. 326 p.

Dunn M., 1999. Aspects of the stock dynamics and exploitation of cuttlefish, Sepia officinalis (Linnaeus, 1758), in the English Channel. Fish. Res., 40 (3): 277-293

Gayanilo, F.C., Sparre, J.R. and Pauly, D. 1995. The FAO-ICLARM Assessment Tools (FISAT) User's Guide. FAO Computerized Information Series (Fisheries), N° 8. Rome, FAO, 126 p.

Kelly, E. 1999. Assessment of the English Channel Cuttlefish Stock (Sepia officinalis L.) using Depletion Methods based on Biological Data and Commercial Catch Data Collected both in France and the UK. Master thesis, 53 p.

Medhioub A., Etude de la croissance et du cycle sexuel de la seiche (Sepia officinalis L.) des côtes normandes., Thèse de 3ème cycle. Université de Caen, 1986, 117 p.

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Natsukari, Y., and Komine, N. 1992. Age and growth estimation of the European squid, Loligo vulgaris , based on statolith microstructure. Journal of Marine Biological Association of the United Kingdom, 72(2): 271-280.

Robin J.P., Koutsoubas D., Moreno A., Denis V., Arvanitidis C.,Cuhna M., Pereira J., Kotoulas G., 2000. Global patterns of the cuttlefish Sepia officinalis Linnaeus, 1758 (Cephalopoda, Sepiidae) stocks in the Northeastern Atlantic and the Mediterranean: an integrated overview. CIAC 2000 Millenium Cephalopod Conference, Aberdeen July 2000. (poster).

Royer J., Peries P., and Robin J.P. 2002. Stock assessments of English Channel Loliginid squid: updated depletion methods and new analytical methods. ICES J. Mar. Sci., 59, (in press).

Sparre, P., and Venema, S.C. 1998. Introduction to tropical fish assessment. FAO Fisheries Technical Paper, No 306, Rome.

Figures:

Figure 1: English Channel cuttlefish catch structure: monthly catches in numbers by age-group. (95 (1) refers to animals born in 1995 and recruited in autumn 1995 and 95 (2) refers to animals born in 1995 and recruited in spring 1996).

Figure 2: Number of recruits estimated for each age-group with Cohort Analysis back-calculations.

Figure 3: Comparison of inter-annual trends in recruitment described with Cohort Analysis number of recruits per year and with CPUE (inter-annual component of GLM estimates) (in this graph recruits which entered the stock during 1995 belong to the 1995 (2) spring and 1996 (1) autumn age-groups).

Figure 4: Estimates of fishing mortality in each calendar month (averages across cohorts)

Figure 5: Estimates of fishing mortality at age in 1995 and 1996 cohorts.

Figure 6: Simulated yield and mean biomass of Sepia officinalis from the Thomson and Bell model applied to the 1995 and to the 1996 cohorts.

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93(1) 93(2) 94(1) 94(2) 95(1) 95(2) Catch (in millions) 96(1) 96(2) 97(1) 97(2) 98(1) 4.5

4

3.5

3

2.5

2

1.5

1

0.5

0 01/ 04/ 07/ 10/ 01/ 04/ 07/ 10/ 01/ 04/ 07/ 10/ 01/ 04/ 07/ 10/ 95 95 95 95 96 96 96 96 97 97 97 97 98 98 98 98

Figure 1: English Channel cuttlefish catch structure: monthly catches in numbers by age-group. (95 (1) refers to animals born in 1995 and recruited in autumn 1995 and 95 (2) refers to animals born in 1995 and recruited in spring 1996).

Recruitment (millions) 60

50

40

30

20

10

0 95(1) 95(2) 96(1) 96(2) 97(1) 97(2) 98(1) Age-group

Figure 2: Number of recruits estimated for each age-group with Cohort Analysis back-calculations.

95

Cohort Analysis GLM Correlation coefficient r = 0.99 115 1.0

95 0.8

75 0.7

55 0.5

35 0.4

15 0.2 1995 1996 1997 1998

Figure 3: Comparison of inter-annual trends in recruitment described with Cohort Analysis number of recruits per year and with CPUE (inter-annual component of GLM estimates) (in this graph recruits which entered the stock during 1995 belong to the 1995 (2) spring and 1996 (1) autumn age-groups).

Average Fi 1995 1996 1997 1998 0.6

0.5

0.4

0.3

0.2

0.1

0 Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Figure 4: Estimates of fishing mortality in each calendar month (averages across cohorts)

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Fa,i 1995 Cohort 1996 Cohort

1

0.8

0.6

0.4

0.2

0 relative 3 5 7 9 11 13 15 17 19 21 23 age

Figure 5: Estimates of fishing mortality at age in 1995 and 1996 cohorts.

1995 Cohort Yield Biomass 14000

12000

10000

8000 nes n o

T 6000

4000

2000

0 mf 0123

1996 Cohort Yield Biomass 14000

12000

10000

8000 nes n o

T 6000

4000

2000

0 mf 0123

Figure 6: Simulated yield and mean biomass of Sepia officinalis from the Thomson and Bell model applied to the 1995 and to the 1996 cohorts.

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