Atlas of the Biology and Distribution of the Squids Illex Illecebosus and Loligo Pealei in the Northwest Atlantic

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Atlas of the biology and distribution of the squids Illex illecebrosus and Loligo pealei in the Northwest Atlantic

C.A.P. Black, T.W. Rtmell and E.G. Dame

and Oceans et Océans Canada Canadian Special Publication of Fisheries and Aquatic Sciences 100

Fisheries & Oteùns- LIBRARY

Atlas of the biology and distribution of th teff. ?. .4 1998 squids IIlex illeeebrosus and Loligo pealei in the Northwest Atlantic BIBLIOTHÈQUE Pêches & Ocdtins

G.A.P. Black and T.W. Rowell Department of Fisheries and Oceans Scotia-Fundy Region Biological Sciences Branch P.O. Box 550 Halifax, Nova Scotia B3J 2S7 and E.G. Dawe Department of Fisheries and Oceans Newfoundland Region Science Branch P.O. Box 5667 St. John's, Newfoundland A 1 C 5X1

Department of Fisheries and Oceans Ottawa 1987 0 Minister of Supply and Services Canada 1987

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Correct citation for this publication:

Black, G. A. P., T. W. Rowell and E. G. Dawe. 1987. Atlas of the biology and distribution of the squids Illex illecebrosus and Loligo pealei in the Northwest Atlantic, Can. Spec. Publ. Fish. Aquat. Sci. 100: 62p. Contents

List of Tables iv

List of Figures iv Abstract/Résumé

Preface vi

Introduction 1

Data Presentation 1

Classification 2

Biology 3

Life History Synopsis 3

Distribution 3

Interactions with Other Species 4

Growth 4

Reproduction 5

Maturity 5

Fecundity and Spawning 5

Early Life Stages 6

Physiology 6

Fishery 7

History 7

Trends 7

Catches 7

Regulations 8

Utilization 8

Future Prospects 8

Short Term 8

Long Term 9

Acknowledgments 9

References 59 List of Tables

1. Progression of the feeding pattern of IIlex illecebrosus and Loligo pealei. 10

2. Northwest Atlantic fish predators of squid (IIlex illecebrosus or Loligo pealei ). 10

3. Nominal catches (tons) of short-finned squid (IIlex illecebrosus) in the Northwest Atlantic from 1963 to 1985. 11

List of Figures

1. Drawings of IIlex illecebrosus and Loligo pealei. 2

2. Map of NAFO Subareas and Divisions. 12

3. Hypothetical migration path of Illex illecebrosus off the Northeastern U.S.A. 13

4. Interannual distribution patterns of Illex illecebrosus off the Northeastern U.S.A. 14

5. Interannual distribution patterns of Loligo pealei off Eastern Canada. 15

6. Interannual distribution patterns of Illex illecebrosus. 16

7. Seasonal distribution patterns of Illex illecebrosus. 18

8. Seasonal distribution patterns of Loligo pealei. 22

9. Monthly distribution patterns of Illex illecebrosus. 26

10. Monthly distribution patterns of Loligo pealei. 28

11. Distribution patterns of Illex illecebrosus in relation to bottom temperature. 29

12. Distribution patterns of Loligo pealei in relation to bottom temperature. 32

13. Distribution patterns of Illex illecebrosus in relation to maturity stage. 34

14. Nominal world catches of squid. and all fish, and shellfish in marine and freshwater. 40

15. Nominal Canadian catches of squid in the Northwest Atlantic. 40

16. Nominal catches of Illex illecebrosus in the Northwest Atlantic, and biomass estimates. 41

17. Interannual distribution patterns of nominal catches of squid from NAFO Subareas. 42

18. Monthly distribution patterns of Illex illecebrosus - International Observer Program sampling. 46

19. Management areas and boundaries relevant to squid fisheries. 58

iv Abstract

Black, G. A. P., T. W. Rowell, and E. G. Dawe. 1987. Atlas of the biology and distribution of the squids Illex illecebrosus and Loligo pealei in the Northwest Atlantic. Can. Spec. Publ. Fish. Aquat. Sci. 100: 62 p.

The biology, distribution, and fisheries of the short-finned squid (Illex illecebrosus) and the long-finned squid (Loligo pealei) in the Northwest Atlantic are summarized for the Northwest Atlantic Fisheries Organization Convention area. The primary synopses are presented graphically (using maps and charts) as a visual representation of distributional patterns, and fisheries statistics. In consequence of its importance within the Canadian Fisheries Conservation Zone, greatest emphasis has been given to description of the short-finned squid and its fishery.

Résumé

L'article résume sommairement la biologie, la distribution et la pêche de l'encornet (Illex illecebrosus) et du calmar (Loligo pealei) à l'intérieur des zones de pêches de l'Organisation des Pêches de l'Atlantique Nord-ouest. On a utilisé une présentation gra- phique par cartes et diagrammes pour i4strer les principaux sommaires des patrons de distribution et des statistiques de pêche. A cause de son importance à l'intérieur de la Zone Canadienne de Conservation des Pêches, nous avons accordé une attention spéciale à l'encornet et à sa pêche. Preface

Over the years, the Biological Sciences Branch and its several predecessors have col- lected vast amounts of resource information in order to gain an understanding of the state of the biological resources and the factors which influence their abundance and well- being. This information exists in various forms from handwritten reports in filing cabi- nets, to sophisticated, but unpublished data bases, through to assorted published reports on one or another aspect. Unfortunately, not enough is available in readily understandable and comprehensive form suitable for use by the fishing industry, the general public and our own Department.

In an attempt to improve on this situation, the Biological Sciences Branch has undertak- en a program to display, in graphic form, resource information presented with relevant but deliberately spare supporting text. The ultimate intention, or goal, is to make available as much as is feasible of our accumulated information through production of a series of resource atlases. To accomplish this, a position of Fisheries Analyst was created with the incumbent working with scientists within the Branch and elsewhere, to seek out and as- semble the data, followed by analysis, interpretation and the choice of the most appropri- ate means of display.

Since conveying resource information graphically has not yet reached the status of a science, the first incumbent, Mr. G.A.P. Black, has been faced with a development task of considerable and daunting proportions. Many tools are available with many different ways of analyzing and representing the data; major adaptations and further development were required to fulfill our intentions. Additionally, although many atlases displaying resource information are extremely beautiful examples of the cartographers' craft, not all are supported by data prepared with sufficient rigour to make them reliable; viewers can be seduced by the beauty of the presentation and not be aware of or overlook the shallow- ness of the data. To avoid this trap, we decided at the outset that in this series of atlases, the emphasis should be placed on objectivity, quantitative representations, comprehen- siveness, simplicity and reliability, as well as good looks. To this end, Mr. Black has spent much time in assembling the data and developing the computer programs for analysis, and determining the best way to present the data, as well as to store it electronically in a form which permits ready access and updating.

We chose the squids, il/ex illecebrosus and Loligo pealei, as the species to serve as guinea pigs in developing the art and to determine the extent to which we could fulfill the objectives stated above. It is my belief that Mr. Black has been successful in devising a system which meets these objectives. In the course of preparing this atlas and the sequel on the scallop, Placopeeten magellanicits, he has together with the co-authors illustrated very well how this system functions in presenting resource information graphically. The two atlases, one on squids and the other on scallops, bring to a close the strictly experimental phase in which the foundation for the steady production of a continuing series of resource atlases has been laid.

James E. Stewart Director Biological Sciences Branch Scotia-Fundy Region

vi Introduction Data Presentation

Canadians have been fishing for squids since the late Figures showing the spatial relationship of squids to rel- 1800s and, since the development of a significant offshore evant biological and abiotic parameters are presented in ac- fishery in the early to mid-1970s, have actively participated cordance with the guidelines of Bertin (1981). The maps in the management of this valuable renewable resource. At are presented at small scale (using a Mercator projection) in its peak in 1979, the Canadian squid fishery was significant groups, intending to convey a graphic representation rather in relation to other Canadian fisheries, and to the interna- than a cartographic product, emphasizing contrasts or simi- tional squid fisheries market. In consequence, a coordinat- larities. ed international research effort was initiated in 1980 under the aegis of the Northwest Atlantic Fisheries Organization The choice of a presentation format involved the weigh- (NAFO) to provide the biological information for an effec- ing of contradictory objectives, including: clarity, simplici- tive management regime. ty, resolution, precision, and uniformity. Two mechanisms were used to balance these objectives: discrete data scaling, By virtue of their geographic distribution, most squids and definition of the sampling area. caught in Canadian waters are short-finned squid (IIlex illecebrosus), which form the major component of the squid The data values used in developing the figures are dis- fisheries and, for which the majority of the research data is cretely scaled using logarithmic transformations to provide available. Although long-finned squid (Loligo pealei) is a constant ratio of circle (symbol) size. The scaling varies regularly seen over the more western areas of the Scotian between figures so as to minimize secant (overlapping) cir- Shelf, only rarely are significant numbers present. cles as much as possible. In the legend, the numbers brack- eting a given circle, are the data group limits within which Specific critical aspects of the life cycle of I. illecebro- all data values are represented by the same sized circle. sus remain to be confirmed by observation. The location of spawning is still unknown, although there is evidence that On the maps, the area of sampling is indicated by an un- the major spawning areas are south of Cape Hatteras and derlying grey shading. This defines the spatial area within probably in the Gulf Stream Frontal Zone over the Blake which all sampling occurred for the given criteria and, as Plateau. Limited evidence also suggests that the species is such, any extrapolation .of squid abundance or variability passively and (or) actively migratory over long distances outside these areas is hypothetical. (Florida to Newfoundland and return). These factors, coupled with the extreme recruitment variability and un- The data for a number of the figures were obtained from known predator/prey impacts, argue strongly for fuller research surveys directed towards both the assessment of study and careful international management. squid and groundfish stocks, using bottom trawl random stratified surveys. The results are presented as kg per tow, Included in this report is information on: the biology of or number of squid per tow (1.75 nautical miles or approxi- both I. illecebrosus and L. pealei; the distribution of I. illec- mately 30 minutes). While the variation in the effective ebrosus and the northerly extent of the distribution of L. catch-rate of the gear between the vessels cannot be elimi- pealei; the spatial distribution of I. illecebrosus in relation nated, at the scale used in these figures, the data presenta- to the abiotic factors, time of year and sea-bottom tempera- tion is not significantly altered by this variation in sampling ture; and variation in these distributions in relation to the methods. maturation of both males and females throughout their life- span. Additionally, world squid catches are compared to In order that the figures be comprehensible, much de- world catches of all fish and shellfish species; squid catch- tailed information had to be.aggregated. In most cases this es, by statistical unit areas are tabled and mapped for recent was done using more than one parameter (e.g. time and ge- years; and the spatial relationship between squid directed ographic area). The reader is cautioned to consider that catch and bycatch are mapped on a monthly basis for a while these computer-generated maps are based on actual number of years. data, they do not represent absolute limits on the real spatial distribution and biological variability of the species. This report is intended to provide a concise yet scientifi- cally rigorous textual and visual synopsis of the biological and fisheries information available for I. illecebrosus and L. pealei in the Northwest Atlantic, concentrating on the Ca- nadian fishing region. The report has been prepared for use by the industry, fisheries managers, and interested lay read- ers.

For a more detailed review of the biology of I. illecebrosus and L. pealei, the reader is also referred to O'Dor (1983) and Summers (1971), respectively. Note that un- less otherwise specified, IIlex refers to IIlex illecebrosus and Loligo refers to Loligo pealei.

1 Classification Common Name: Short-finned squid' Long-finned squid

Class: Cephalopoda Cuyier,1798 Subclass: Coleoidea Bather,1888 Order: Teuthoidea Naef,1916

Suborder: Oegopsida d'Orbigny, 1845 Myopsida d'Orbigny, 1845 Family: Ommastrephidae Steenstrup, 1857 Loliginidae Steenstrup, 1861 Subfamily: Illicinae Posselt, 1890 Genus: Illex steenstrup, 1880 Lol i go Steenstrup, 1880 Species: illecebrosus (LeSueur, 1821) pealei LeSueur, 1821

17-35 cm. mantle length

Ventral Dorsal Ventral Dorsal view view view view Illex Loligo illecebrosus pealei

Figure 1. General representation of the dorsal and ventral view of Illex illecebrosus and Loligo pealei as adults (adapted from Roper et al., 1984). Note that although the scale is approximate, Loligo pealei are larger as adults.

I The English word "squid" is a dialectal variant of "squirt" and refers to the activity of the siphon during movcinent.

2 Biology Life History Synopsis Distribution

There are four closely related species of IIlex in the The NAFO Subareas and Divisions are shown in Figure western Atlantic: 11lex illecebrosus, I. oxygonius, I. coinde- 2 (page 12). The life cycle of Illex is generally understood tii, and I. argentbuts, whose morphological similarity make with exception of the exact location(s) of spawning (Figure species separation of adults difficult and, at present, impos- 3, page 13). Evidence suggests that the major spawning sible for larval and juvenile stages (Roper et al., 1969). Il- event occurs south of Cape Hatteras during December lex oxygonius, the sharptail shortfin squid, occurs in the through January in the Gulf Stream/Slope Water frontal western Atlantic from Chesapeake Bay to southern Florida zone (Rowell et al., 1985a; Rowell and Trites, 1985). Sec- and the southeastern Gulf of Mexico and in the eastern At- ondary and tertiary spawning events occur in spring and lantic in the Gulf of Guinea (Roper et al., 1984). 11lex coin- summer and in some years are important contributors to the detii, the broadtail shortfin squid, occurs in the western At- resource biomass in the more southern shelf areas (NAFO lantic from southeast Florida throughout the Gulf of Subareas 5&6). Neutrally buoyant egg niasses apparently Mexico and Caribbean Sea and in the eastern Atlantic from drift passively in the subsurface region of maximum salini- Norway to the Gulf of Guinea including the Mediterranean ty of the Gulf Stream (O'Dor and Balch, 1985; Rowell and and Black Sea (Roper et al., 1984). 11lex argentinus, the Ar- Trites, 1985). Hatching occurs in about 9-16 days (O'Dor gentine shortfin squid, is found in the western Atlantic et al., 1982). The resultant larvae and juveniles apparently along the coast of Argentina and around the Falkland Is- continue to drift northward with the Gulf Stream and dur- lands (Hatanaka et al., 1985). ing the summer the juvenile Illex move onto the Shelf and into shallow water off northern New England, Nova Scotia 11lex illecebrosus is a pelagic cephalopod with an appar- and Newfoundland. On the Shelf, males undergo earlier ent life span of about 1-1.5 years. Spawning is believed to maturation than females and appear, based on sex ratios, to occur only once and the spawned animals probably die migrate south during the fall in advance of the females shortly after. The current hypothesis is that most of the (Lange and Sissenwine, 1983). Fully mature females have population spawns south of Cape Hatteras, over the Blake only rarely been encountered (Dawe and Drew, 1981). Evi- Plateau, in late autumn and early winter (in water masses dence for a long-distance spawning migration includes tag- having characteristics midway between Continental Edge ging data for a squid released in Newfoundland and report- and Yucatan Straits Water) (Rowell et al., 1985a; Rowell ed captured off Maryland, covering 2,027 km in 107 days and Trites, 1985). The egg masses are believed to be neu- (Dawe et al., 1981a). trally buoyant when released, and are transported north by the Gulf Stream (Rowell et al., 1985a; O'Dor and Balch, Loligo spawn in the spring through summer in shallow 1985). The distribution is related to water masses and tem- water, primarily between Cape Cod and Cape Hatteras perature regimes. Adults are observed over the continental (Lange and Sissenwine, 1983). Loligo produced from the shelf from the southeastern U.S. to Newfoundland and early spawning period may mature early and spawn during Labrador during the summer and autumn. During autumn the late summer of the following year, otherwise they the maturing adults migrate southward to the spawning are- spawn during the spring and summer of their second year, as. with some surviving and spawning at about two years of age. They overwinter in deeper waters along the Shelf- Loligo pealei, the second but much less important spe- edge. cies in terms of relevance to Canadian fisheries, is related to two other loliginids of commercial interest in the west- Both species are distributed as far south as Florida; Loli- ern Atlantic, Loligo (Doryteuthis) plei and Lolliguncula bre- go is also found in the northern Gulf of Mexico, although vis. 2 Loligo (Doryteuthis) plei, the arrow squid, occurs pre- both species appear to be most abundant north of Cape Hat- dominantly south of Cape Hatteras, in the Gulf of Mexico, teras (Voss and Brakoniecki, 1985; Whitaker, 1980). The and apparently as far south as northern Argentina. Lolli- more northerly distribution of Illex, relative to Loligo, in the guncula brevis, the brief squid, occurs in coastal and estu- Georges Bank - Gulf of Maine area, is evident from the arine waters, and has been observed from Maryland to Rio USA bottom trawl surveys (Figure 4 & 5, pages 14-15) de la Plata, Brazil (Voss, 1956). (Whitaker, 1980), and the combined U.S.A. and Canadian bottom trawl surveys (Figure 6, pages 16-17). The general Loligo pealei is a pelagic cephalopod distributed from seasonal distribution of Illex north of Cape Hatteras is hint- Canadian waters to the upper Gulf of Mexico and Cuba. ed at by the distribution of catches from the combined bot- Loligo pealei is generally found in very low abundance on tom trawl surveys of the Northeast Fisheries Center (USA), the Scotian Shelf and then only on the the more western ar- and Scotia-Fundy, Gulf, and Newfoundland Regions (Can- eas and in the Bay of Fundy (Rowell, 1986). Spawning oc- ada) (Figure 7, pages 18-21). The pattern of a northerly ex- curs during the late spring and summer south of Cape Cod. tension of range during summer and fall is also evident for Loligo has a life span of 1-2 years. Adults are found pre- Loligo (Figure 8, pages 22-25). For both species, especially dominantly from Cape Hatteras to Nantucket Shoals. Illex, a seasonal increase in abundance is evident. Although the catch rates between the regions and research vessels are 2 All are members of the family Loliginidae. not standardized, the overall pattern is clear. Seasonal

3 catches by bottom trawl research surveys vary from year to As both IIlex and Loligo grow, the diet changes from one year. Aggregated monthly plots from 1970 to 1980 bottom dominated by crustaceans to one composed largely of fish trawl surveys indicate a general movement of IIlex onshore and squid (Table 1, page 10) (Amaratunga, 1983; Amara- and northward during the summer and a retreat southward tunga et al., 1979; Froernian, 1984; Maurer and Bowman, during the fall (Figure 9, pages 26-27), and a similar 1985; Vinogradov, 1984; Vovk, 1983). Illex illecebrosus, monthly distribution of Loligo for the northeastern U.S.A. obtained from groundfish surveys in the Gulf of Maine to (Figure 10, page 28). the Cape Hatteras area during the spring, summer, and autumn of 1979-80 fed primarily on squid (56% of stomach The onshore movement of squid to the Shelf areas takes contents by weight), while Loligo fed primarily on fish place during spring and early summer when water tempera- (56.2%) (Maurer and Bowman, 1985) , Scotian Shelf sam- tures are increasing, and there is evidence that distribution pling of Illex indicates that feeding frequency is reduced as is influenced by bottom-water temperature (Scott, 1978). the season progresses (Amaratunga et al., 1979). The detailed distribution of squid catches in relation to the bottom-water temperature from research trawl surveys sug- Illex was found to be the almost exclusive diet of the gests, however, that more than temperature influences the long-finned pilot whale (Globicephala inelaena) in New- distribution of IIlex (Figure 11, pages 29-31) and Loligo foundland waters, with the two arriving and departing the (Figure 12, pages 32-33). Bottom trawl research surveys region annually, almost simultaneously (Mercer, 1975). A during July 1970-1980 did. not demonstrate a relationship population of 50,000 pilot whales was estimated to con- between depth or bottom-water temperature and mean sume 166,000-249,000 tonnes during 100 days of residen- catch rate (Mohn, 1981); however examination of squid cy in inshore Newfoundland waters. distribution patterns during 1980-83 indicated the highest concentrations on the Scotian Shelf were in the western Fish predators of Mex and Loligo (Table 2, page 10) in- area where bottom-water temperature exceeded 6°C. De- clude common demersal species such as cod, haddock, pol- spite this, high concentrations of IIlex are regularly found in lock, red and silver hakes and many others (Lange, 1980). areas of low bottom-water temperature, which appears to In years of high abundance, Illex was a common prey item be a preference rather than a requirement (Rowell et al., for cod (especially larger cod) around Newfoundland, al- 1985b). Migration inshore, to and over the continental though not as significant as either capelin or sand lance shelf, appears linked to food availability. (Lilly and Osborne, 1984).

Interactions with Other Species Growth

The trophic relationships of IIlex (Ennis and Collins, Estimates of the growth rate of Illex vary considerably. 1979) and Loligo (Vovk, 1974) are similar. The food of Growth estimates from size data are affected by immigra- both species consists of primary, secondary, and tertiary tion and emigration within the sampling region. Squid sam- consumers, while both are prey species, mainly at the pled from Newfoundland catches between May and Octo- fourth trophic level. ber doubled in mantle length and exhibited an average 6- fold weight increase over the 5 month period (a 1.0-4.0 cm. A Scotian Shelf study of the interactions of IIlex with increase in mantle length/month) (Squires, 1957). Mesnil cod, haddock, pollock, silver hake, yellowtail flounder, red (1977) reported growth rates of 1.6 cm/month for males hake, redfish, and environmental factors (location, depth, and 1.9 cm/month for females. Von Bertalanffy growth salinity, surface and bottom-water temperature) indicated curves indicate sexual, areal, and interannual variability in IIlex (and silver hake) catch rates were simultaneously pos- growth (Amaratunga, 1980). itively correlated with bottom-water temperature, salinity, sediment particle size, and depth (Waldron, 1983). Feeding studies indicate a food conversion efficiency of 35%-51% (O'Dor et al., 1979). Refined measures of aver- In the coastal waters of eastern Newfoundland and Lab- age daily feeding rate (5.2% of body weight), average daily rador, a study of Ille.x- predation on cod, capelin, and her- growth rate (1.3%), and average daily feeding rate for ring suggested a possible predation effect (significant nega- maintenance (1.8%) have been reported, where the daily tive correlation) between squid abundance and capelin feeding rate varied from 0%-15% of body weight apparent- (NAFO Div. 2 & 3K), assuming squid prey on age-group 0 ly due to behavioural interaction in the school (Hirtle et al., and 1 capelin (Dawe et al., 198 1 h). Additionally, in New- 1981). Additionally, assimilation efficiency has been esti- foundland inshore waters, cod have been found to be the mated at 0.86 (Wallace et al., 1981). most important fish prey on the northeast coast, with redfish the only fish prey species observed from south coast Loligo growth has been estimated at 1.7-2.0 cm/month samples (Dawe et al, 1983). Fish were identified through during the summer period, ranging to 0.4-0.6 cm/month otoliths found in the stomachs of the squid sampled. An in- over the winter (Mesnil, 1977). Growth estimates of Loligo crease in the incidence of cannibalism, as the season from the Gulf of Mexico were similar to those from north- progressed, appeared to be related to a decrease in the ern areas (Hixon et al., 1981). availability of other prey species.

4 Reproduction Fecundity and Spawning

Maturity Mating behavior of IIlex has been neither fully observed or described, although the few brief episodes of courtship A consensus is lacking as to which of the several maturi- behavior observed between captive squid suggest a pattern ty-stage scales used is most valid in representing the pro- similar to that seen in Loligo (see below) but possibly less gression of maturation in IIlex and Loligo pealei. (Juanico, pronounced or more secretive (O'Dor et al.,1980; O'Dor, 1983). Different maturity-stage indices are used for each 1983). Observation on fully mature captive males suggests sex. Among the various scales, that for IIlex females is rela- they may accumulate several hundred spermatophores in tively objective, being based on the ratio of nidamental advance of mating. Mating is triggered by the presence of gland length (NGL) versus mantle length (ML) (Durward fully mature females, and results in the implantation of et al., 1979). NGL/ML ratios have been found to corre- male spermatophores in the female mantle cavity. Individu- spond reasonably well with the histological condition of al males may mate with more than one female (O'Dor et the ovaries (Coelho et al., 1982). The scale for males is al., 1980). Mating may take place well before spawning. subjective, and there is little doubt that its application by During the spawning of IIlex illee'ebrosus, eggs from the various investigators may be inconsistent (Mercer, 1973a; oviducts, sperm from the implanted spermatophores, and Durward et al., 1979; Amaratunga and Durward, 1979). jelly from the nidamental glands, are mixed with seawater and exuded slowly into a spherical gelatinous low density The apparent earlier maturation of the male IIlex may be egg mass (O'Dor et al., 1980). The density of the egg mass the artifact of a truncated male maturity index scale. As is determined by the density of the eggs (with a specific spermatogenesis is apparently continuous from an early de- gravity of 1.10), the gelling agent, and the constituent sea- velopmental stage, three other reproductive features have water. Because the bulk of the mass is seawater, the mass is been suggested as maturation indicators; spermatophore almost neutrally buoyant and may become neutrally morphology (Voss, 1983), spermatophore number (O'Dor buoyant if the surrounding water increases in density et al., 1980), and degree of hectocotylization (Schuldt, (through a decrease in temperature or increase in salinity) 1979). Samples of IIlex from St. Margaret's Bay (Nova as generally occurs with increasing depth in near-surface Scotia), Newfoundland, and a captive population (Dalhou- waters. The egg-mass buoyancy, rate of temperature equili- sie University) suggest that relative hectocotylization is an bration, and terminal velocity affect the sinking rate. The unreliable index of maturity, but may be an indicator of en- ecological significance of this is that the spawning location vironmental conditions experienced during development (yet to be discovered), must provide a physical environ- (Coelho and O'Dor, 1984). Accidental loss of spermato- ment that results in a sinking rate such that the egg mass re- phores may invalidate spermatophore counts as an ade- mains within a suitable temperature regime throughout the quate index. A preliminary maturity-stage index compris- incubation period. Females may produce more than one egg ing 9 stages based on reproductive/ecological scales has mass, and still not be completely spawned out (eggs have been proposed to provide further differentiation (Burukov- been observed in the ovary of dead females after spawning) sky et al., 1984). (Durward et al., 1980).

Bottom trawl surveys of IIlex on the Scotian Shelf be- Egg masses obtained in laboratory spawnings contained tween 1979 and 1983 indicated differences in the monthly up to 100,000 ova, spaced about 1-2 cm apart and were 0.5- and spatial distribution of maturity stages (Figure 13, page 1.0 m in diameter (Durward et al., 1980; O'Dor and Balch, 34-39). Analysis of these data for the 1980-83 period indi- 1985). Although egg counts indicate a potential of about 5 cated that maturation varied from year to year, and may be x 105 ova per female, true or functional fecundity may be associated with the considerable variation in between-year well below this, since fully mature males may, by the and within-year size distributions. Length frequencies placement of their spermatophores, stimulate premature showed a general increase in modal length of the primary spawning in females (O'Dor et al., 1980). Less than half of cohort from July to September, with secondary and tertiary the spawned eggs survived the incubation period under the cohorts of smaller IIlex in the western regions of the Shelf laboratory conditions. during September and October being relatively more abundant in low biomass years (Rowell et al., 1985b). Growth When Loligo are held in captivity, they usually school as curves for 1977-1979 showed general yearly similarities, a group. However when mating behaviour is induced, the with the males maturing earlier and at smaller mean length males become increasingly excited, and begin darting than females (Amaratunga, 1980). about, challenging other males, and attempting to separate individual females from the school. A preference is shown Only seven female IIlex in advanced stages of maturity for particular females. When a male is courting a female, a are known to have been captured, all in the period 1968- median arm is held upwards in an "S" shape, while the 1980 (Dawe and Drew, 1981). Captures were reasonably chromatophores in front of his eyes are expanded to form a evenly spread from Cape Hatteras to Bay of Islands, New- dark brown patch. Challengers may displace males, result- foundland and provide no indication of probable spawning ing in the largest or strongest males coupling with the most area(s). desirable females. During copulation the male, flashing his chromatophores swims beside and slightly below the fe-

5 male. The male grasps the female with his arms around the Physiology mantle, and pulls himself forward so that his arms are close to the mantle opening. The male uses his hectocotylus (a The respiration and swimming performance of Illex has modified arm) to reach inside his own mantle and pick up a been studied experimentally (Webber and O'Dor, 1985). /i- bundle of spermatophores from the penis. The spermato- lex has a negative buoyancy of 3%-4%. While swimming phore bundle is then "cemented" to the inside of the fe- (tail first), the body is angled upwards by the jet to maintain males mantle or oviduct; the male releases the female. The its position vertically, and the squid uses 6 times more en- entire process is consummated in 5-20 seconds. During ergy than sockeye salmon per unit distance travelled for spawning, the female produces a jelly-like egg capsule speeds of 1 and 2 body lengths/second. The rate of 02 con- from the oviduct. The egg capsule is implanted with sperm sumption increases exponentially with swimming speed re- released from the transferred spermatophores. The female sulting in a critical swimming speed of 1.6 - 2.2 body takes the egg capsule in her arms so that the sticky end, lengths/sec. (62-88 cm/sec.). The most economical speed of containing no eggs; faces outwards. The egg capsule is transport was estimated to be 50-70 cm/sec. The long- commonly fastened to seaweeds such as Fucus, or if possi- distance migration of a tagged Illex (sec Distribution above) ble, attached to an existing egg mass, by pushing the sticky represented an average speed of at least 22 cm/sec. end of the egg capsule into the centre of the egg mass (Ar- nold et al., 1974). Squids have proven to be useful experimental animals for investigating neurophysiology and vision. The giant Early Life Stages axon found in squid is sufficiently large to permit dissec- tion and facilitate the conduct of experiments in neurophys- As an aid in identifying and aging egg masses, a devel- iology. The large and well developed eyes of squids indi- opmental staging scheme for in vitro and in situ spawned cate a high degree of dependence on or use of visual 11lex was developed by O'Dor et al. (1982) based on previ- information. As a result, the physiology and biochemistry ous work on IIlex coindetii (Naef, 1923) and Todarodes pa- of vision generally have been studied using the eye and op- cificus (Hayashi, 1960). In their study, O'Dor et al. (1982) tic lobes of Loligo (Arnold et al., 1974). Squid behavior demonstrated that embryonic development failed at tem- when attacking jigs (sec below) and their apparently great peratures below 12.5°C, while the development rate at ability in net avoidance (O'Dor, 1983) can be associated 21°C was nearly twice that at I2.5°C. with a strong response to visual stimuli.

The larvae of IIlex have been described as Rhyncho- teuthion type C', on the basis of morphological differences between a small number of larvae sampled from Cape Hat- teras to Georges Bank (Roper and Lu, 1978; Vecchione, 1978). 3 Subsequently, a large sample of juveniles between the Gulf Stream and the edge of the Canadian continental shelf indicated the Gulf Stream contributed to the distribution of the early life stages (Amaratunga et al., 1980; Fedu- lov and Froerman, 1980; Rowell et al., 1985a; Dawe and Beck, 1985a,b). Additionally, the occurrence of 0.8-1.0 mm larvae in the Gulf Stream at a latitude of 31.5° N, suggested that spawning probably occurs south of Cape Hatte- ras (Rowell and Trites, 1985; Trites and Rowell, 1985). 4

3 Rhynchoteuthion was used to describe these larvae, referring to 4 Rhynchoteuthion Type B' larvae (probably Ommastrephes) the fused tentacles forming a proboscis (Rhynchos - snout were found during the survey and overlap the distribution of Type [Greek]). The proboscis later separates into distinct tentacles dur- C (believed to be Illex illecebrosits). The spawning location can- ing the transitional stage. The final separation is used as the enter- not be inferred exactly as the larvae were being transported by the ion for definition of the change from larval to juvenile stages. Gulf Stream at a maximum rate of about 120 kinAlay.

6 Fishery Trends History The 1960's saw an increase in the total world landings of Squid jigging (and trap fisheries to a lesser degree) for all marine species, which leveled off after 1971. The annual Illex developed off Newfoundland during the late 1800's increase in the world catch since 1970 has been 1%-2% (Hurley, 1980). Catches from these fisheries occur during (Worms, 1983). 6 Squid landings accounted for only about the summer months, peaking in September, and historically 1.5% of the world landings throughout the 1970's (Figure were used primarily for bait in the longline cod fishery 14, page 40). (Squires, 1957). Through the 1970's, markets for squid for human consumption in Japan, the Soviet Union and Spain The size of world-wide commercially important squid expanded or opened up for Illex (Dawe, 1981). Squid jig- stocks has been estimated as high as 49 million tonnes (Li- ging remains an important fishing method when squid are pinski, 1977) in comparison to 1985 world landings of available inshore, particularly in Newfoundland (Mercer, squids, cuttlefishes and octopuses of 1,672,033 tonnes 1973b). Squid jigging was initially conducted solely by (FAO yearbook, 1987). hand from small inshore boats. The lure or jig is about 70 - 120 mm long and consists of one or more rings of sharp barbless hooks with an ellipsoid lure above (Hamabe et al., Catches 1982). 5 The jig is rhythmically jerked upwards through the water to attract the squid and ensure that it remains on the Peak catches for NAFO Subareas 2, 3, 4, and 5+6 took- hook. Automated squid jiggers are now used, especially on place in 1974, 1979, 1979, and 1976 respectively (Figures larger vessels, to permit the setting of a larger number of 15-17, pages 40-45). Catches vary by more than an order of lines simultaneously. magnitude and may be taken variously as an index of availability of squid in the fishing areas, as a reflection of vary- Squid fishing in the U.S.A. (Maine to N. Carolina) being market demand, and somewhat more tenuously, as a gan in the late 1800's as incidental catches of both Illex and possible indicator of overall abundance. Research survey Loligo in the summer otter trawl fisheries and the inshore data show that the peak catches in Subareas 3 & 4 were in- Massachusetts trap fisheries. Catches were used as bait for dicative of greater abundance over these areas of the Shelf, cod fishing, to supply small fresh fish markets or, failing but the declining U.S. catches in Subarea 5 & 6 after 1976 these uses, was discarded (Lange, 1980). are not indicative of reduced abundance over those areas. Rather they reflect a management regime which restricts The Soviet distant-water fleet first reported squid catch- catch levels. es as part of the trawl fishery in 1964, and the Japanese started reporting in 1967. The fishery began off the U.S.A. The great variation in catch results in a fishery, which in in the late 1960's and subsequently along the Scotian Shelf its poor years has little economic value, but in its best years in the early to mid 1970's (Lange and Sissenwine, 1983). It (such as 1979) has been among the most valuable in Cana- is interesting to note that although the distant-water fleets da (comparable to scallops and lobsters when the value of also fish the Grand Banks, little of the Subarea 3 (New- the international market is considered). The International foundland) catch is taken offshore. Even in the peak year Observer Program of the Department of Fisheries and of 1979, the bulk of the fishing in Newfoundland was by Oceans has sampled both domestic and foreign fishing inshore trap and jig. The reverse was true on the Scotian fleets since 1979. Although observers are not able to sam- Shelf and off the U.S.A. where the greatest part of the fish- ple all catches, the data are representative of the commer- ery took place along the Shelf-edge. Bottom trawling, mid- cial fishery. The yearly variation in catches, and the geo- water trawling, and squid jigging have been used by the graphically restricted directed fishing zone of Subarea 4 distant-water fleets. Bottom trawling is the most economi- (the Small Mesh Gear Line) are shown in Figure 18 (pages cally efficient, but, as the quality standards for processing 46-57). squid are high, the larger catches made by trawlers may result in wastage when the entire catch can not be processed By-catches of Illex and Loligo by the distant-water fleets before it spoils (Wiseman, 1982). While squid jigging re- (primarily the U.S.S.R. and Poland) began to be reported in sults in the best quality product, large offshore squid- 1964, and became increasingly significant when fishing for jigging vessels may not be able to fish other species when, silver hake, red hake, mackerel, and herring commenced as in the northwest Atlantic, the season is short (Wiseman, and necessitated regulatory control (see below). 1982).

5 The colour of the jig can apparently be determined by the squid Observation on large schools, indicates that the squid hold off un- at a distance of 3 metres at 5 metres depth. At night, the stem of til one individual makes an attack, which initiates simultaneous at- the jig is visible at 1 metre, where attacking squid have been ob- tacks by many others. served to pause momentarily, close to about 60 cm and then attack. While squid do not struggle noticeably as the jig is hauled in, an adequate rate of retrieval is required to prevent the squid 6 The 1980 total world landings were over 72 million tons. from disentangling its arms from the hooks.

7 Regulations ment of this regulation, in conjunction with TACs and bycatch restrictions for both species. In 1980, in response to Management and regulation of squid resources was ini- previous catches in excess of the existing TACs, a TAC of tiated in 1974 by the International Commission for the 150,000 t was established to protect against fishing mortali- Northwest Atlantic Fisheries (ICNAF). 7 Distant-water fish- ty above 40% in years of moderate abundance (Northwest ing fleets began reporting squid in 1964 (U.S.S.R., 4 t), and Atl. Fish. Organ. Sci. Coun. Rep., 1980). The reduction in by 1973 reported catches totaling 45,090 t (Lange and Sis- fishing effort in subsequent years suggests that the fishery senwine, 1983). A preemptive quota set the total allowable is self-regulating in years of low abundance (Northwest catch (TAC) of Loligo (only) at 71,000 t for 1974. At that Atl. Fish. Organ. Sci. Coun. Rep., 1984). time the catch statistics did not separate IIlex and Loligo, but the majority of the catches off the U.S.A. were of Loli- go. The Illex fishery was just developing. In 1976, separate Utilization TACs were established in the northeastern U.S.A. (Subare- as 5 & 6) of 30,000 t for Illex, and 44,000 t for Loligo. Sub- Utilization of squid as bait has been important since the areas 2-4 were assigned Illex TACs of 15,000 t (Int. Comm. beginning of the Newfoundland fishery over a century ago. Northwest Atl. Fish. Redbook, 1975). Both Illex and Loligo A small dried squid industry, with export for human con- were managed as single stocks throughout their distribution sumption to China, existed in 1910 and lasted until 1948. ranges. The extension of jurisdiction by Canada and the This market was revived in 1978 (Hurley, 1980). The pri- U.S.A. in 1977 resulted in the formation of 2 management mary utilization of squid from the northwest Atlantic since regimes. 1976 has been for human consumption as whole, cleaned and slit (mantle), or as tubes (mantle) in a variety of forms After the U.S.A. Fishery Conservation and Management — fresh, frozen, dried, semi-dried, or half-skinned (Wise- Act became effective in 1977, the U.S.A. withdrew from man, 1982). ICNAF and assumed management responsibility for the fishery resources within 200 miles of its coast. The Nation- Japan is the most important market for squid; an estimat- al Marine Fisheries Service (NMFS) and the South Atlan- ed 600,000 t are available for annual consumption in its do- tic, Mid-Atlantic, and New England Fishery Management mestic market (2.8 kg per capita) (Wiseman, 1982). Other Councils jointly developed a Fisheries Management Plan, markets include Spain, Hong Kong, and eastern Europe; the which established the TACs for Illex and Loligo at 30,000 t Canadian market is tiny. and 44,000 t, respectively. 'Fishing windows' 8 were established to regulate the foreign squid fisheries in time and place (Figure 19, page 58). In the U.S.A., both Illex and Lo- ligo are currently managed on an April 1-March 31 fishing Future Prospects year (Lange, 1984a; Lange, 1984b). Short Term In 1977, Canada assumed management responsibility for the fisheries resources within a Fisheries Conservation With catches of Illex varying by as much as a factor of Zone extending 200 miles off its coast. Canada first solicit- 60 in individual years between 1969 and 1979 in NAFO ed advice from ICNAF (and subsequently NAFO) on the Subareas 2-4 (Table 3, page 10), there is no prospect for a management of Illex in 1974. In 1977, the TAC in Subareas stable fishery. In contrast, the fishery in Subareas 5 & 6, 3 & 4 was set at 25,000 t. For Subareas 4VWX, the by- while showing the same trends and fluctuations, is much catch of juvenile haddock associated with the silver hake more stable. The population dynamics of Illex are not yet fishery prompted the introduction in 1979, of the Small well understood, thus the effects of increased fishing effort Mesh Bottom Trawl Fisheries Regulations, which provided are not yet clearly separable from other factors which con- for the use of codends under 130 mm inside a restricted tribute to the fluctuations. zone extending seawards from the shelf-edge, from April 15 to November 15 (Figure 19, page 58). Subsequently, the The apparent resilience of cephalopods to intensive fish- fishing season was further restricted to a July 1 - November ing has been linked to their alternative life strategy (lower 15 period. A multispecies management regime for both II- fecundity, reduced number of spawnings, larger eggs with lex and silver hake is implemented through the enforce- presumably a lower mortality rate) in comparison to that of

7 ICNAF was established in 1950 to promote the conservation 8 'Fishing windows' are areas along the northeastern USA, over and optimum utilization of the fishery resources of the Northwest the continental slope, designated for foreign fishing within speci- Atlantic area within a framework of international cooperation by fied fishing seasons. participating countries. The successor to ICNAF, NAFO was ap- proved January 1, 1979 by Canada, Cuba, the E.E.C., G.D.R., Ice- land, Norway and the U.S.S.R. based on a similiar objective of multilateral cooperation.

8 longer-lived bony fish. Their relatively low fecundity may short-term (months) projections of likely abundance and result in significant abrupt declines in stock size when the availability to the fishery. parent stock is overfished (Caddy, 1983). In the short term there is no prospect for predicting stock levels of IIlex even Squid are part of the food chain at all trophic levels. a few months in advance of the fishery: the life span of the Their interactions with other commercially important spe- species is too short and knowledge of the biotic and abiotic cies remains important from both the biological and fisher- factors influencing spawning success and survival of the ies viewpoint. Failure to understand their role in the fisher- early life stages (larvae and juveniles) is too rudimentary. ies ecology of the Shelf may seriously hamper effective In years of high abundance, the economic value of the spe- management of other fisheries resources. cies may be extremely high and the ecological impacts are likely to be highly significant to other fish stocks .

Catches of Loligo are likely to remain relatively unim- Acknowledgments portant in Canada's fisheries, although in certain years commercially significant concentrations occur over the The data used to generate the maps and diagrams were western areas of the Scotian Shelf, on Georges Bank and in obtained from a variety of sources. Bottom trawl ground- the Bay of Fundy. As with IIlex, there is at present no pros- fish surveys and dedicated squid surveys conducted by pect of predicting stock levels in these areas. Canada, and the U.S.A. (see below). The data set entitled the Scotia-Fundy Region's Squid Program contains data from the Canadian surveys and in addition, results provided Long Term from cooperative international squid surveys conducted with the Union of Soviet Socialist Republics, France and A greater understanding of the biology of the squids 11- Japan. The value of the considerable efforts by the many lex and Loligo is required to define their population dynam- scientists and personnel who planned and conducted these ics. As stock size is so variable, more reliable predictors of surveys is greatly appreciated; we thank all those who par- year-class strength may assist the industry by allowing ticipated.

Data sources used for analysis within this report.

0 CV ■-o oo (--q en 7t. oo oo oo oo oo 00 CT CT CT CT a, CT CT CT CT CT \ CT CT CT CT ON

Research Surveys: U.S.A. Groundfish • • • • • • • • • • • • • • • • Gulf Region Groundfish • • • • • • • • • • • • 0 0 0 0 Scotia-Fundy Region Groundfish • • • • • • • • • • • • • • • • Scotia-Fundy Region Squid O0 0 0 0 0 0 • • • • • • • • 0 Newfoundland Region Squid O 0 0 0 0 0 0 0 • • • • • • 0 0 Canadian Int, Obs. Program O0 0 0 0 0 0 0 0 • • • • • • • • data available 0 data unavailable

9 Table 1. Generalized progression of the feeding pattern of Illex and Loligo. Derived from Amaratunga et al. (1979), Amaratunga (1983), Vovk (1983), Froerman (1984), and Maurer and Bowman (1985).

small juveniles — chaetognaths and copepods predominate in diet of Illex and Loligo respectively — euphausids, amphipocla, and other small crustacea important — as Illex grows, chaetognaths become less important', and euphausids predominate

large juveniles — crustacean component is the most important, with euphausids predominating — fish, particularly myctophids, and cephalopods become important items in diet

adults — squid and fish dominate, relative importance varies with availability

Table 2. Northwest Atlantic fish predators of the squids Illex illecebrosus and (or) Loligo pealei. Source: Mid-Atlantic Fishery Management Council (Anon., 1978) and Lange (1980).

Alewife Rainbow smelt American john dory Red hake Atlantic bonito Redfish Atlantic croaker Roughtail stingray Atlantic silverside Roundnosed grenadier Atlantic angel shark Sand tiger Atlantic tomcod Scup Barndoor skate Sea raven Barrelfish Silver hake Bigeye thresher shark Skipjack tuna Black sea bass Smooth dogfish Bluefin tuna Spiny dogfish Bluefish Spotted hake Butterfish Striped bass Clearnose skate Summer flounder Cod Swordfish Fourspot flounder Tautog Goosefish Thorny skate Haddock Threespine stickleback Hickory shad Thresher shark Lancetfish Tilefish Little skate Toadfish Longhorn sculpin Weakfish Mackerel White hake Mackerel shark White marlin Night shark White perch Northern pilot whale White shark Northern searobin Windowpane flounder Offshore hake Winter skate Opah Witch flounder Oyster toadfish Yellowfin tUllit Pollock

10 Table 3. Nominal catches (tonnes) of short-finned squid (IIlex illecebrosus) in the Northwest At- lantic from 1963 to 1985. Source: Northwest Atl. Fish. Organ. Sci. Counc. Rep., 1980, 1986.

Year Subarea 2 Subarea 3 Subarea 4 Subarea 5+6

1963 0 2,199 103 1,210 1964 0 10,408 369 193 1965 0 7,831 433 563 1966 0 5,017 201 1,562 1967 0 6,907 126 2,662 1968 0 9 47 4,948 1969 0 21 65 2,802 1970 0 111 1,274 2,453 1971 0 1,607 7,299 4,036 1972 0 26 1,824 14,713 1973 2 620 9,255 15,178 1974 31 17 389 16,653 1975 0 3,764 13,993 13,790 1976 0 11,254 30,510 24,936 1977 6 32,748 47,199 24,883 1978 7 41,369 52,688 17,568 1979 1 88,832 73,259 17,341 1980 1 34,779 34,826 17,864 1981 0 18,061 14,142 15,574 1982 0 11,164 1,744 18,188 1983 0 0 421 11,623 1984 0 368 404 9,876 1985 9 404 269 6,069

9 Preliminary data.

11 ; Boundaries of Subareas Greenland Boundaries of Divisions 200 m Depth contour ë;1>t -5

60 - 2G

55 -

Canada 2J

Atlantic 3K Ocean 50 -

Newfoundland

3M 45 - ,,ep , re 30 U.S.A. 4V

• Capetp 3N 5Z 40 - 6

, 6B 6D 6E 6F 6G 6H

60 35 2

75° 70 65 60 55 50 45 40

Figure 2. Subareas and Divisions of the NAFO Convention Area.

12 Canada

50 •

45 —I

Slope Water

Cape CCie , Gulf Stream U.S.A. :Late fall 40 —I mature adults return Late winterlearly spring larvae and juveniles Sargasso Sea drift with Water Cape Hatteras,, Gulf Stream 35 —1

Suspected spawning 30 —1 ground

25 —I

20° 1 1 1 80° 75 70 65 60 55 50 45

Figure 3. Hypothetical migration path of Illex illecebrosits with passive Gulf Stream transport to the northern feeding grounds and active southward migration to the suspected spawning areas south of Cape Hatteras.

13 45 .. • • • • e• • • • • • • • • • *le«. 40 •••• O. 5. .0 • 1972 35

45 • ••••• ee..eelree ••• lr •• I. ee ....e •. • • •• • elb•• , . • • • .0.•110.0 •5••• 4.. ••.4. E. • • ••••••• . • 400•410 e 10..6.. • •• .•e. • •••••Kecié. 4;;emigare 40 • • id. • one * • •fed •• •••5 • ••• •0 :ir • • • 1975 •

•• IS•. •••••••9 ••0 •• 0•• 14.600 • •••••• I.. 060. toe. ee . giim•• • • ...re -- - 40.•eee••••• •••«•••••••1 •.••0 '1466 •••551.• :›,,,,42•81 • • • • • ...UM*** • 01(0•003 • SIM • mg. e': erie • dre• • m04(10080 •••••00101101. • ••e•••••••■•• •••••••••110 41101000•010** 002040•11.00•1 NS* • weep cm • • *MI ale .0. 600 • *e• • .0 "ffll se •••• P.6110, s. ' 41,0 • SS • .• lb* -• .0 1978 19K

45 •e. •••••• •••••.. . .• e•••••• • Ire,. mg& -11- • • • 40 •Isas • f• • • "fee?

0* •• • 1981 1983 35

45 • • • 75 70 65 60 • • • • • • 1500 • .. • •• • .0..4 ..... • 200 **** • 40 ee• •• 05. ••• —30 .00. 4 •00 • 0.6 • 0.1 kgItow 1984 1985 35° 75° 70 65 60 75 70 65 60

Figure 4. Interannual distribution patterns of Illex illecebrosus off the northeastem U.S.A. and western Scotian Shelf between 1972 and 1985, based on data from U.S.A. spring and fall research surveys.

Area sampled is indicated by shading, and mean catch in kg/tow for each 0.5° square is represented by a solid circle scaled to the catch level.

• ;. ■•,:: .. ..110i.• • • ■ •••e. to II • ise• •• . •egié .•‘•») .• eb 1••111111 le* . ••i•s•leedie • lFInèiledie: '414•*11■ 011Wee'_ . 40 ••••, elle•••C•. *OOP ,- er: - ,4114114 .e*011e '. *gee . •1110 !Is« ,-,• " 0/11 — top' .0801 . •• ..111. 400 •le. *OS •• I. •• 1972 ' :: 1973 •• 1974 35-j 7-- 45

.., ...404, • Ilbe - • 01111011 • mi ••••••••••••

40- ::;DI• . .-.. 4,..10H•11, eat» •ve» , 4010 •40. Oa a É., •011e. : 4. ID» .imme ,•41.• •111 •se .1. 40 es 1975 11 1976 c• 1977 35_ 45

- *•• ■ •*00, -*Se- ...•■ ,, ...... • 5 *OlOgeigNIO**-' ....de.ses... ei_11U•111■11uee , •" .0 u*•1140119"." -: 40 - • •goe , , es •54•580 >es ,gwell •*•e• 09i40 • *elf •lee. 400 heep • At• . •1• 41114 •.0 •, 1978 ,a 1979 1980 35 • q. - • 45

.4.. *••444 .,:•egef • '0 40 . OOOOO ...e 0*4111/110010**. ' • • ** 411•X•BOT" . " 413-1 /-• •«■■ •• Omer. • 11- , • 7 - •;'• • *411 •, e.41.11, ..1.001e..x. • *SO' ••• • Ilt*: ••.. , •• •.. •• *00 ••• 1981 • •gb 1982 e*, 1983 35 45 75 70 65 60

— 1500 ••••00•11108,44Y • -200 •.• eiouteueie*.• .01001140,14*.444 - • 40 -1 • i.e..- * 4 -.7. ••*:44e**0*' — 30 ale • 0,011) •.*.0, 4 •***4 0.6 4.11114 .1110 0.1 kgItow .0111 1984 1985

75° 70 65 60 75 70 65 60

Figure 5. Interannual distribution patterns of Loligo pealei off the northeastern U.S.A. and western Scotian Shelf between 1972 and 1985, based on data from U.S.A. spring and fall research surveys.

Area sampled is indicated by shading, and mean catch in kg/tow for each 0.5° square is represented by a solid circle scaled to the catch level.

15 ▪

• •

1970 1971 40

40 1972 1973

45 • • • • 00000 • • • O O •••11. • .. • ...... see 0. ...••••••.) emreee..e 1974 ..0•..0.0 1975 40 *eq..

45 ,›Iir •• . •800 " 4416. •016 • •.• e•il, • elhe • • •••• .111.4111M . • ..... 11 • •11(440 •10». 1976 1977 40`Lererts__ • 1 70° 65 60 55 50 45 70 65 60 55 50 45

Figure 6. Interannual distributional patterns of Illex illecebrosus off eastern Canada and the northeastern 1500 U.S.A. between 1970 and 1985, based on data from Canadian spring, summer, and fall research surveys, and • 200 • spring and fall U.S.A. research surveys. — 30 4 Area sampled is indicated by by shading, and mean catch in kg/tow for each 0.5° square is represented by a solid circle scaled to the catch level. • 0.6 0.1 kgltow 16

50 • ., .... • -••• ■• : ... S. "« I. • es . ••• * •e. i -.;..s.'... - ...... , ;*Ic...• •,., 45 •• • ... ••••■ ••• ,. iiii ing. • •• ••• - ‘.mr...;••: •• • . • ..... . . • ..4.... ....„-.8....emie eq.. • •* ****** .....•• ..•• • •• ...Hob • «.• • *** - S. .... ••• ••«••••110/1 •• •• ••• ■■ • • 114.• ...:••••,: -::‘••••■ • • • 41è •..... 4‘. leelid 1978 . 4*'*'*..** *** 1979 40 • • .*Willibbe.

50 • • •••• • e ..• i

••. • • • 45 ee 1 *•• -. .. '.. .. , .... • • lob •• ... ... • ...... .. . _• ,.4 .:•85 •• • ••' *O. • • ••••• •:•1> • .. I... , ... • • le,l; ••;111111. 1980 *,41:;!:: 1981 40 .. ..4.».•

45 *WI. •■ • •

• . • - 41/•.1' • F. Ile• • 1982 :.,-..11:*.7• 1983 40 ... ••-• isueree-

te.•....e.. • ,..s.i.‘• - 1984 ....e.e. 1985 40 ' 1••••• . ele•WW" . FI 1 70° 65 60 55 50 45 70 65 60 55 50 45

Figure 6 cont. Interannual distribution patterns of Illex illecebrosits off eastern Canada and the north- — 1500 eastern U.S.A. between 1970 and 1985, based on data from Canadian spring, summer, and fall research 200 • surveys, and spring and fall U.S.A. research surveys. — 30 4 Note: A corresponding figure for Loligo pealei is not presented, as it .1- 0.6 seldom occurs on the Scotian Shelf in significant numbers. 0.1 kgilow 17 50

45 • • SIS • of. • Mee • 0 . .

- 40

' Spring 1970 Summer 1970 Fall 1970 35

45 re•ee .• ......

Summer 1971 Fall 1971 35

' • •• • 45 OO OOO Foe OOOOOO t• • • •••B•• I. • Mal• ee • tee • • .8. • •• • • ... •

a... •geg•ele•• 40 .614 S. toeIt' Summer 1972 Fall 1972 35° 70° 65 60 55 50 45 70 65 60 55 50 45 70 65 60 55 50 45

1000 Figure 7. Seasonal distribution patterns of IIlex illecebrosus off the eastern U.S.A. and • 100 Canada between 1970 and 1980. Based on data from U.S.A. and Canadian research surveys. .- 10 • 1 Area sampled is indicated by shading, and mean catch in kg/tow for each 1° square is represented by a solid circle scaled to catch level. 0.1 kgItow

18 50

Summer 1973 Fall 1973 35

45 riable:r ea. *** COO ernell • "te

ev"-•

Summer 1974 Fall 1974 35

• • 45 tea, • • ••• ‘sti." • • easee.. »I• se. • • • e. • seem.. I. • .8. • • • • •••te•le • • • 4. • •soo, • 145..e. .. eq.) 40 : 5 Spring 1975 Summer 1975 Fall 1975 35° 70° 65 60 55 50 45 70 65 60 55 50 45 70 65 60 55 50 45

— 1000 Figure 7 cont. Seasonal distribution patterns of Illex illecebrosus off the eastem U.S.A. and 100 Canada between 1970 and 1980. Based on data from U.S.A. and Canadian research surveys. 10 1 catch in kg/tow for each 1° square is Area sampled is indicated by shading, and mean 1-- 0.1 kgItow represented by a solid circle scaled to catch level.

19 Spring 1976 Summer 1976 Fall 1976

Spring 1977 Summer 1977 Fall 1977

• • • a. • . (o • • •.0 toe) •8•.0 e•i mr..e.gee.. *eel 0 ******* e el.* •el» • •1 • eee OP 0 * •11. Kale" ...Keefe* • 0 .1.11014000 e 11.18.1M4 ••... *OM. •...... e .. , sal. OD Spring 1978 Summer 1978 Te Fall 1978 I I 70° 65 60 55 50 45 70 65 60 55 50 45 70 65 60 55 50 45

— 1000 Figure 7 cont. Seasonal distribution patterns of Illex illecebrosus off the eastern U.S.A. and 100 Canada between 1970 and 1980. Based on data from U.S.A. and Canadian research surveys. —10 Area sampled is indicated by shading, and mean catch in kg/tow for each 1° square is represented by a solid circle scaled to catch level. 0.1 kgItow

20 • • • ••• et■te■ -0* «ez «•• 0.••••• (ee) • • • w• • Ir.. •esee el, 0 , • .,0400 • ee,e, 01111. • gr. • go• 00i enetleje. reellet000*********04eur• leVeit.111ene te.90•••••■■ •••

;. woo*******

Spring 1979 Summer 1979 r Fall 1979

Spring 1980 Summer 1980 1 i 1 1 ! 70° 65 60 55 50 45 70 65 60 55 50 45 70 65 60 55 50 45

— 1000 Figure 7 cont. Seasonal distribution patterns of IIlex illecebrosus off the eastern U.S.A. and •— 100 • Canada between 1970 and 1980. Based on data from U.S.A. and Canadian research surveys. — 10 1 Area sampled is indicated by shading, and mean catch in kg/tow for each 1° square is 0.1 kgItow represented by a solid circle scaled to catch leve,

21 45

eeeee •••••••"7* - ee.e. • eie • • •• • 0. Oi 35 Fall 1970 45

e.g, f.41mmieee... • • • • 40 •e••*

35 Fall 1971 45

•liegkee• 41 •• le - _

35°-

Spring 1972 Summer 1972 Fall 1972

75° 70 65 75 70 65 75 70 65

—250 Figure 8. Seasonal distribution patterns of Loligo pealei off the eastern U.S.A. and • 3Ø Canada between 1970 and 1980. Based on data from U.S.A. and Canadian research surveys. 5 0.7 Area sampled is indicated by shading, and mean catch in kg/tow for each 0.5° square is represented by a solid circle scaled to catch level. 0.1 kgItow

22 r • • • Ik OM go ellie Ir. •••••••• .6,1( F tee, - 1TZ

• •••••••••84),r 40 !sly 4

Spring 1975 Summer 1975 Fall 1975 1 1 1 1 75° 70 65 75 70 65 75 70 65

—250 Figure 8 cont. Seasonal distribution patterns of Loligo pealei off the eastern U.S.A. and • 30 Canada between 1970 and 1980. Based on data from U.S.A. and Canadian research surveys. • , —e 0.7 Area sampled is indicated by shading, and mean catch in kg/tow for each 0.5° square is 0.1 kgItow represented by a solid circle scaled to catch level.

23 45

„4. •••• •• >-• ■•• •••••••.•*.... 11.00000...... 40 *elm • «090. 0.* 00. 0300 0010 • • 000 00 . •0 35

Spring 1976 Summer 1976 Fall 1976 45

• 1. • • 0.0... 40 .• • ...gm" gm.* em.

• 35

Fall 1977 45

effl ■ 40 ••,...• ",' 047 - 1111 .00 .0 s. 35° •

Spring 1978 Summer 1978 Fall 1978 -r 75° 70 65 75 70 65 75 70 65

—250 Figure 8 cont. Seasonal distribution patterns of Loligo pealei off the eastern U.S.A. and I_le 30 Canada between 1970 and 1980. Based on data from U.S.A. and Canadian research surveys. 5 Area sampled is indicated by shading, and mean catch in kg/tow for each 0.5° square is 0.7 represented by a solid circle scaled to catch level. 0.1 kgItow

24 45

~.\ . ~, ...... " ...... " I..t : ,..~... - ....' \' /'" 1/-""-"'," ... ' ,' .. , .. ..' .. /' .. ,,/ • • all "..,, _ _ ::'4 ~:' J). "" 40 . ....l" O ~••• ·• ••• • ~ . ,, ~ . • • f" ......

~ :' f;. ~:'~; C-:~~-__''\7 ~ .... ) •• j" ...... " ...... ", I ...... --.jlOt,...... ".. .' : ::--i· :~ ,. ... '"' 40 .0,' ...... , .. ' ...... ,. 35" •

75' 70 65 75 70 65 75 70 65

- 250 Figure 8 cont. Seasonal distribution patterns of Loligo pealei off the eastern U.S.A. and ~ 30 Canada between 1970 and 1980. Based on data from U. S.A. and Canadi an research surveys. ~ 5 ~0.7 Area sampled is indicated by shading, and mean catch in kg/tow for each OS square is ~ O.I kg/low represented by a solid circle scaled to catch level.

25 75' 70 65 60 55 50 45 75 70 65 60 55 50 45

— 2000 Figure 9. Monthly distribution patterns of IIlex illecebrosus off the eastern U.S.A. and 200 Canada for the period 1970-80. Based on data from U.S.A. and Canadian research surveys. • —30 Area sampled is indicated by shading, and mean catch in kg/tow for each square is • 0.5° - • J represented by a solid circle scaled to catch level. — 0.7 — 0.1 kgItow

26 50

•• •

40 •

March April 35

• ••• 45 • • *14 .• •.... ■II1L'e " ..*" 5 ca..» e*O • • er" • «lei.' en. • eelleemo.• • -• (1.(1. • elleb• en* ee • •• 40 • 0410 III •

July August 35

- ■ ■ • . eeeee •••• ••igulp._,„_•» • e e• Agee. ' tee .oelb • 40 • .1

November December 35°- 1 75° 70 65 60 55 50 45 75 70 65 60 55 5() 45

— 2000 Figure 9 cont. Monthly distribution patterns of Illex illecebrosus off the eastem U.S.A. and ql±. 200 Canada for the period 1970-80. Based on data from U.S.A. and Canadian research surveys. —• 30 • kg/tow for each 0.5° square is — Area sampled is indicated by shading, and mean catch in • represented by a solid circle scaled to catch level. — 0.7 I— 0.1 kgItow

27 "

January February March

....•. . . 40 :"# .••• ••••• I "

October Nm'cmber December l -,--, 1 1 1 75' 70 " 10 65 7.' 70 " Figure 10. Momhly t.1I~lnblillon pattern!> of 1.0 /il."(1pt. ·lIM off the c:I.)\cm U.S.A. for the - J. ~ 40 ArclI ~nnplcd I~ indktllc.1 b) ~h,l{ l m~ .•LlII.I UIf.:.lIllatch In kgltu\\ for each 0." Mluare l~ • , n..1)re,cnlcd by a »olid circle ~lllcd 10 catch level. -• 0.7 0.1 ~}: ftol\' 2M January February March

',' " ....•. .' ~. -. :#1'" • • ..- ••' .. I

OC lober Novembe r December I I 75' 70 " 75 70 65 70 " Figure 10. Monlhly disuibulion paltems of L(lII,I:(I {Wt/lei off Ihe e:blem U.S.A. f(W Ihe - 300 period 1910-80. Based on cb la from U.S.A. and Can:ulian research " ur"c)"~ .

Arc" ~ lIm pkd is indicmcd by .• hading. and I1Ie:'1I .. :UI:h IJ1 Io. g/low for each O.Y ~uarc i~ . , n.'Prc\emed bY:L solid circle scaled 10 cawh level. -• ""0,7 -'- 0.1 k/(I tolt" 28 ' •.., " " .••...... • . ... , ',' .. .

1.9' C

...... • .. . '. ..' ~ :"~':.':' .-... , . ' .., ":I.or. , .' ..... /

,. ...• • .... -.: ' '0' •

6' · 6.9' C I 75' 10 .5 60 55 50 45 75 10 65 60 " 45

Figurt I I. Distribution of II/e.\' iIIf'Ct'broslIs in rela1ion 10 boilom lcmpcrnlurc. Based on . :0 dala from U.S.A. and Canadian rescaTC'h surveys for the period 1970-80. .!... 60 Area samplt.:d is indicated by shading. and mean catch in kgltow for cachO.S' ~u3re is • 7 rcprc'OCn100 by a solid circle scaled 10 calch level. -' - 0,' 0.1 kgl loll'

• • • doo. • M. 45 .-TH. • • • 01• • el» .• S..) ••• W.. eeeee .04011 • •••• •etto 11•• •441 « • ee.. se • ...... • MI» «.. • • «••• • • .5 ....01..... eeelg tee 111•4»•••••• es. . • 53. ••••• »Oleo. ...ee eel. eeeee •e. 85.11,41. 40 pe •z ..• •• • »op s. ••• .49 •■• • 7° - 7.9 ° C 35 8° - 8.9 ° C

OD 45 • • 041 • • • •• • e eirwreeege • •0.0 .0 .... • • .100a.010 m.c: • lie .01.01..011108. . • .....5.0•/• • 0.0.0.0. . • ee eltellee e-es. le • gge .910 gm '••• •ffle••• r .....00 •s.s. 1104.1ffledffle 40 00•11Wefflinen0...g..s.s• ee..1. tffl, «40 eee • ge. s. 9° - 9.9 ° C . 11 35

45 • eleq, ee •• ."9r- • • • • eeeee • • • •• •we> O* .c.. • • 5 . 5 MO •...... 00.... : • • • slob e...5...51•• 0.. •e...0 .temeesseeemp. • 5••••5111111l• PE•le•••/«.• • 101•41110 40 e•alu•e•elp 04 Me •eg. cum 00 se 35 11° - 11.9 ° C 12° - 12.9 ° C

75° 70 65 60 55 50 45 75 70 65 60 55 50 45

— 5000 Figure 11 cont. Distribution of Illex illecebrosus in relation to bottom temperature. Based on 500 data from U.S.A. and Canadian research surveys for the period 1970-80. 1, — 60 • Area sampled is indicated by shading, and mean catch in kg/tow for each 0.5° square is — represented by a solid circle scaled to catch level. • — 0.8 — 0.1 kgItow

30 50

45 •• • • •••••• .41•01.90 ét• •• 14•1111.11 • seffle•••11 effligeOP, tm.e 40 —MIWilm• I. *se • e•• I.

on, 13° - 13.9°C 14° - 14.9°C 35

45 • • •••• • 06, orm entelleteee• Mole 4010, • 40 s. ffle

. 11 16 0 - 16.9 ° C 35° 15° - 15.9° C Fi

75° 70 65 60 55 50 45 75 70 65 60 55 50 45

— 5000 Distribution of Illex illecebrosus in relation to bottom temperature. Based on Figure 11 cont. I 500 data from U.S.A. and Canadian research surveys for the period 1970-80. • —60 0 Area sampled is indicated by shading, and mean catch in kg/tow for each 0.5° square is — 7 • represented by a solid circle scaled to catch level. — 0.8 0.1 kgItow

31 ° - 2.9 ° C 3° - 4.9°C

5° - 6.9°C 7° - 8.9°C

• •• • • • • • . •boom •beeeeefeee.e.. »111•10••••••••• eque. .s.... e. IMO 001,* ogee •dm. 00•11• *** .e. .1» .9

4: 9 0 - 10.9°C 11° - 12.9°C

75° 70 65 60 75 70 65 60

500 Figure 12. Distribution of Loligo pealei in relation to bottom temperature. Based on • 90 data from U.S.A. research surveys for the period 1970-80. • 16 • Area sampled is indicated by shading, and mean catch in kg/tow for each 0.5° square is 3 represented by a solid circle scaled to catch level. 0.5 0.1 kgItow

32 45

do.

• 00 •e. • •• ine••••••••• • •• ...teemeeeee 2•18•1116•08•111••• •• •rnee ' 40 j,."•••••• •• 66 siage • •••) 66e• ••• «tee r 660 Imo 600 *66 . •• ibe se 00 se 35 • 13° - 14.9 ° C 15 ° - 16.9 ° C 45

..- • se• / ; 60111 • s••• ••• • 40 e• T• ,' ••

00 oè 0* 35

17° - 18.9° C 19° - 20.9° C 45

• or,/ 450 7 s. r

35°- -

21° - 22.9°C 23° - 24.9 ° C

75° 70 65 60 75 70 65 60

500 Distribution of Loligo pealei in relation to bottom temperature. Based on Figure 12 cont. • 90 data from U.S.A. research surveys for the period 1970-80. le— 16 • , Area sampled is indicated by shading, and mean catch in kg/tow for each 0.5° square is — • represented by a solid circle scaled to catch level. — 0.5 0.1 kgItow

33 • •

45 t

January February March 20, 40

45 • , • 066066 • .6. .61100 •••••••• ••••• ■•••

May .Iune 40

• • 45 eft• • ..,114 'erne* *Hi ...esesee..

September 40

November ;001,, ,111. 1«,,- 40 201 1 70° 65 60 70 65 60 70 65 60

—100 Figure 13. Monthly pattem of distribution over the Scotian Shelf of immature male -25 Illex illecebrosus during the period 1970-83. • — u Area sampled is indicated by shading, and mean catch in number of animais per tow for -e— 1.5 each 0.5° square is represented by a solid circle scaled to • catch level. — 0.3 i_ .i #/tow Note: A similar figure is not presented for Loligo pealei in view of the generally limited distribution of this species over the Scotian shelf.

34 45

- Voà

i•J April May June

• 11041.01, *ea

ee •60•egme •000 hwe,

— 100 Figure 13 cont. Monthly pattern of distribution over the Scotian Shelf of maturing male • 25 Illex illecebrosus during the period 1970-83. • —6 6 Area sampled is indicated by shading, and mean catch in number of animals per tow for — 1.5 • each 0.5° square is represented by a solid circle scaled to catch level. —0.3 0.1 #/tow

35 45

zo0 December

70 65

100 Figure 13 cont. Monthly pattern of distribution over the Scotian Shelf of mature male 25 IIlex illecebrosus during the period 1970-83. 0 — 6 Area sampled is indicated by shading, and mean catch in number of animals per tow for - 1.5 • each 0.5° square is represented by a solid circle scaled to catch level. — 0.3 i_ .i #/tow

36 • 45

6.4 January March 200 r 40

• 45 • • els ••e••. ' • ••• • 14 • 4,•1eibtee •c. •.e..ene• OD S.) April May

... -s • .• . • . • • , • , • .. 45 •• • .... ' 6 s. • .s.»•.7• . e• ems • ••••• • • ..Hi is. si. see *0* 600• •••.sie e.• ..•. •••••• •• •1••••• ••••••■•••• 6.••••• •• •• • 11011. s. ••■•••■• imie. 59 ••••

— 100 Figure 13 cont. Monthly pattern of distribution over the Scotian Shelf of immature female • 25 Illex illecebrosus during the period 1970-83. • —6 • , Area sampled is indicated by shading, and mean catch in number of animals per tow for — • each 0.5° square is represented by a solid circle scaled to catch level. — 0.3 — 0.1 #/tow Note: "immature female" includes females with a nidamental gland length / mantle length ratio <0.1 ( see Durward et al., 1979).

37 \

‘‘, 45

May June 40

• •• • • • • • 45 • • .11 se ego» • • • -4 • • • • •• emie .6 e• •••• egmeem.e..16 ego effleeeee

July August September 40

45 • - • 440• ••••• • • • eta OP

40 r October November - December

70 65 60 70 65 60 70 65 60

— 100 Figure 13 conf. Monthly pattern of distribution over the Scotian Shelf of incipient maturing female • 25 lex illecebrosus during the period 1970-83. • —6• Area sampled is indicated by shading, and mean catch in number of animais per tow for • each 0.5° square is represented by a solid circle scaled to catch level. — 0.3 Note: "incipient maturing female" includes females with a nidamental gland length / mantle — 0.1 #/tow length ratio between 0.1 and 0.2 (see Durward et al., 1979).

38 45

June 111.!

• 00000 • • mr eeeee ee• ee .e I. • ••••••• eee 1.à July

. , , October ,..- December

40 —t 1 i 1 70 65 60 70 65 60 70 65 60

100 over the Scotian Shelf of advanced maturing female Figure 13 cont. Monthly pattern of distribution 25 IIlex illecebrosus during the period 1970-83. —• Area sampled is indicated by shading, and mean catch in number of animals per tow for each 0.5° square is represented by a solid circle scaled to catch level. • 0.1 etow Note: "advanced maturing female" includes females with a nidamental gland length / mantle length ratio between 0.2 and 0.35 (see Durward et al., 1979).

1.5 80

I 1.375 All freshwater and marine species

1.25 70

rn 1.125 ) es nn 1 60 of to

0 ns io

0.875 ill (m

cd 0.75

50 ecies

CT' C/) 0.625 All sp

0 1 1 0 1970 1972 1974 1976 1978 1980

Figure 14. Nominal world catches of squid and nominal world catches of all fish and shellfish in marine and freshwater in millions of tonnes. Source: Worms (1983).

Subarea 3

75 —I 1000) x

(tonnes

hes 50 catc id u

sq Subarea 4 1 l 1

ina 1 1

Nom 1 25

1 A I 1/ 11

--- \ ■ --- I • —7 -T i I ' L- 1920 1930 1940 1950 1960 1970 1980

Figure 15. Nominal catches (thousands of tonnes) of Illex illecebrostis from Canadian waters for the period 1920-84. Subarea 3 catches prior to 1955 are unavailable. Although breakdown by species is not given prior to 1967, it is highly unlikely that species other than Illex illecebrosus contributed significantly to the catch. Source: Northwest Atl. Fish. Organ.Sci. Counc. Rep. 1980, 1985.)

40 100 a

80 1 Catch - Subarea 3

• J .:

•1 i". x 1000)

1. es 60 1 v. nn I. to ■ • C • e • hes ( hes tc Catch - Subarea 4 • 40 1 .• l ca • • ina • • / Nom 20 1 Catch - Subarea 5+6

1 . • - • /‘ • *. / • • ./ •

1964 1968 1972 1976 1980 1984

) tow per ber num an

Me Res. surveys - Subarea 4 — I ( tes ima t es nce da n Abu

1964 1968 1972 1976 1980 1984

Figure 16. a Nominal catches (tonnes) of short-finned squid ( Il/ex illecebrosus) in the Northwest Atlantic from 1963 to 1985; b Research survey abundance estimates (mean number per tow) from Canadian and U.S.A. research surveys for the period 1967-85. Source: Northwest At!. Fish. Organ. Sci. Counc. Rep. 1980, 1986.

41 50

35°

75° 70 65 60 55 50 45 40 75 70 65 60 55 50 45 40

60000 Figure 17. Nominal catches (tonnes) of squids (11lex and Loligo) by NAFO Subareas and Division for the years 1964 through 1984. Catches from Subareas 3 & 4 are almost e 6000 entirely IIlex illecebrosus , while those from Subareas 5 & 6 represent a mixture of Illex e illecebrosus and Loligo pealei. Catch levels (total nominal catch) for each Division 700 • (or Subarea) is represented by a solid circle scaled to catch level. 80 • 9 1 tonnes round fresh weight

42 1971

1973

1975

45 40

60000 Figure 17 cont. Nominal catches (tonnes) of squids (IIlex and Loligo) by NAFO I> Subareas and Division for the years 1964 through 1984. Catches from Subareas 3 & 4 6000 are almost entirely IIlex illecebrosus, while those from Subareas 5 & 6 represent a mixture of • 700 11lex illecebrosus and Loligo pealei. Catch levels (total nominal catch) for each Division • (or Subarea) is represented by a solid circle scaled to catch level. 80 • 9 1 tonnes round fresh weight

43 50 •

.44( , i'AP• pîç‘ 45 • • 40 re•id 1111 gli

35 1976 1977 50 /9!pAinle6-1*( 45 • • 40 411i EL)! if20111'w I • 35 1978 e • 1979

45 • • • 40 £611 111111 Agin 6111:1 • 35 1980 g • 1981

75 70 65 60 55 50 45 40 75 70 65 60 55 50 45 40

60000 Figure 17 cont. Nominal catches (tonnes) of squids (Illex and Longo) by NAFO Subareas and Division for the years 1964 through 1984. Catches from Subareas 3 & 4 • 6000 are almost entirely Illex illecebrosus, while those from Subareas 5 & 6 represent a mixture of • Illex illecebrosus and Loligo pealei. Catch levels (total nominal catch) for each Division 700 (or Subarea) is represented by a solid circle scaled to catch level. • — 80 • — 9 1 tonnes round fresh weight

44 1982 1983

75 70 65 60 55 50 45 40

1984

45 40

60000 Figure 17 cont. Nominal catches (tonnes) of squids (11lex and Loligo) by NAFO Subareas and Division for the years 1964 through 1984. Catches from Subareas 3 & 4 — 6000 are almost entirely IIlex illecebrosus, while those from Subareas 5 & 6 represent a mixture of lex illecebrosus and Loligo pealei. Catch levels (total nominal catch) for each Division 700 • (or Subarea) is represented by a solid circle scaled to catch level. 80 • 9

1 tonnes round fresh weight

45 45

Coe 40H January

4Ø-j 1 May ''1

- 45

45 Ine Mae

•• imè

40°4 _zoo ^ September October

70° 65 60 55 70 65 60 55 1980

Figure 18. Monthly distribution of commercial catches of Illex illecebrosus by foreign fleets — 7000 in Canadian waters as determined from International Observer Program sampling for the Ile - 400 • years 1980 through 1985. - 20 • Areas sampled are indicated by shading. Catches in kg/hour per 0.5° square are represented — 1 by circles scaled to catch level. Open circles are used for squares where all sets (tows) caught — 0.1 kglhour Illex only as a bycatch. Solid circles are used for squares where one or more sets caught squid as a "directed catch" (i.e. greater than 50% by weight).

46 45

40 —re-"k March April

7,001, 40 - July

, .7 ,, y ç.„ I November December

70° 65 60 55 7( ) 65 60 55 1980

Figure 18 cont. Monthly distribution of commercial catches of Illex illecebrosus by foreign - 7000 fleets in Canadian waters as determined from International Observer Program sampling for the 400 • years 1980 through 1985. —20 • Areas sampled are indicated by shading. Catches in kg/hour per 0.5° square are represented 1 by circles scaled to catch level. Open circles are used for squares where all sets (tows) caught — 0.1 kglhour Illex only as a bycatch. Solid circles are used for squares where one or more sets caught squid as a "directed catch" (i.e. greater than 50% by weight).

47 45

_I 40 —I January

—

40°- 2.00 September October 1 70° 65 60 55 70 65 60 55 1981

Figure 18 cont. Monthly distribution of commercial catches of Illex illecehrosus by foreign — 7000 fleets in Canadian waters as determined from International Observer Program sampling for the -400 • years 1980 through 1985. — • Areas sampled are indicated by shading. Catches in kg/hour per 0.5° square are represented — 1 by circles scaled to catch level. Open circles are used for squares where all sets (tows) caught — 0.1 kglhour Illex only as a bycatch. Solid circles are used for squares where one or more sets caught squid as a "directed catch" (i.e. greater than 50% by weight).

48 45

March

July

,.„

40° November

—V 70° 65 60 55 70 65 60 55 1981

Figure 18 cont. Monthly distribution of commercial catches of Illex illecebrosus by foreign — 7000 fleets in Canadian waters as determined from International Observer Program sampling for the • 400 years 1980 through 1985. —20• • Areas sampled are indicated by shading. Catches in kg/hour per 0.5° square are represented — by circles scaled to catch level. Open circles are used for squares where all sets (tows) caught — 0.1 kglhour Illex only as a bycatch. Solid circles are used for squares where one or more sets caught squid as a "directed catch" (i.e. greater than 50% by weight).

49 11 1.••••,,

45 • .,

/ ■J\

^ - - 40 January February

rà7-

oCZo-ob œb

\34. , . 1,00 - 7:9C- 40 May June

• ■er

40°- I% "k _200 1- September October

70° 65 60 55 70 65 60 55 1982

Figure 18 cont. Monthly distribution of commercial catches of Illex illecehrosus by foreign —7000 fleets in Canadian waters as determined from International Observer Program sampling for the 400 • years 1980 through 1985. — 20 • Areas sampled are indicated by shading. Catches in kg/hour per 0.5° square are represented — 1 by circles scaled to catch level. Open circles are used for squares where all sets (tows) caught -9— 0.1 kglhour Illex only as a bycatch. Solid circles are used for squares where one or more sets caught squid as a "directed catch" (i.e. greater than 50% by weight).

50 45 „

40 March I April

„

c4.

20c.. 401 July August

70° 65 60 55 70 65 60 55 1982

Figure 18 cont. Monthly distribution of commercial catches of IIlex illecehrosus by foreign — 7000 fleets in Canadian waters as determined from International Observer Program sampling for the le— 400 • years 1980 through 1985. —20 • Areas sampled are indicated by shading. Catches in kg/hour per 0.5° square are represented — by circles scaled to catch level. Open circles are used for squares where all sets (tows) caught — 0.1 kglhour IIlex only as a bycatch. Solid circles are used for squares where one or more sets caught squid as a "directed catch" (i.e. greater than 50% by weight).

51 45

40 —I January February

45 ■

- 40 — ej." May June

September October —V 1 1 1 70° 65 60 55 70 65 60 55 1983

Figure 18 cont. Monthly distribution of commercial catches of !Ilex illecehrosus by foreign — 7000 fleets in Canadian waters as determined from International Observer Program sampling for the le— 400 • years 1980 through 1985. —20 • , Areas sampled are indicated by shading. Catches in kg/hour per 0.5° square are represented — by circles scaled to catch level. Open circles are used for squares where all sets (tows) caught —• 0.1 kglhour Illex only as a bycatch. Solid circles are used for squares where one or more sets caught squid as a "directed catch" (i.e. greater than 50% by weight).

52 45

March

c;,.> nj,„ —ZOO 40 July August

7 I I 40--1-22c- November

70° 65 60 55 70 65 60 55 1983

Figure 18 cont. Monthly distribution of commercial catches of Illex illecebrosus by foreign — 7000 fleets in Canadian waters as determined from International Observer Program sampling for the i_400 years 1980 through 1985. • —20 Areas sampled are indicated by shading. Catches in kg/hour per 0.5° square are represented — 1 by circles scaled to catch level. Open circles are used for squares where all sets (tows) caught —0.1 kglhour Illex only as a bycatch. Solid circles are used for squares where one or more sets caught squid as a "directed catch" (i.e. greater than 50% by weight).

53 45

y 40 _ - January

• eiro

J y ;-* -,.. 200 1- September October 1 70° 65 60 55 70 65 60 55 1984

Figure 18 cont. Monthly distribution of commercial catches of Illex illecehrosus by foreign — 7000 fleets in Canadian waters as determined from International Observer Program sampling for the 400 • years 1980 through 1985. — 20 • Areas sampled are indicated by shading. Catches in kg/hour per 0.5° square are represented — 1 by circles scaled to catch level. Open circles are used for squares where all sets (tows) caught — 0.1 kglhour Illex only as a bycatch. Solid circles are used for squares where one or more sets caught squid as a "directed catch" (i.e. greater than 50% by weight).

54 December

55 1984

Figure 18 cont. Monthly distribution of commercial catches of 11 lex illecebrosus by foreign — 7000 fleets in Canadian waters as determined from International Observer Program sampling for the i-400 • years 1980 through 1985. — 20 • 1 Areas sampled are indicated by shading. Catches in kg/hour per 0.5° square are represented by circles scaled to catch level. Open circles are used for squares where all sets (tows) caught —0.1 kg/ hour IIlex only as a bycatch. Solid circles are used for squares where one or more sets caught squid as a "directed catch" (i.e. greater than 50% by weight).

55 45

- -

40 'e-1" January February

40 May

October 1 1 70 0 65 60 55 70 65 60 55 1985

Figure 18 cont. Monthly distribution of commercial catches of Illex illecebrosus by foreign 7000 fleets in Canadian waters as determined from International Observer Program sampling for the 400 • years 1980 through 1985. —20 Areas sampled are indicated by shading. Catches in kg/hour per 0.5 0 square are represented 1 by circles scaled to catch level. Open circles are used for squares where all sets (tows) caught kglhour Illex only as a bycatch. Solid circles are used for squares where one or more sets caught squid as a "directed catch" (i.e. greater than 50% by weight).

56 45

40°-

70° 65 60 55 70 65 60 55 1985

Figure 18 cont. Monthly distribution of commercial catches of IIlex illecebrosus by foreign — 7000 fleets in Canadian waters as determined from International Observer Program sampling for the • 400 • years 1980 through 1985. —20 • 1 Areas sampled are indicated by shading. Catches in kg/hour per 0.5° square are represented by circles scaled to catch level. Open circles are used for squares where all sets (tows) caught — 0.1 kglhour Illex only as a bycatch. Solid circles are used for squares where one or more sets caught squid as a "directed catch" (i.e. greater than 50% by weight).

57 45 -t

Georges lewt Canadian "Small Mesh Gear Line"

U.S.A. 'fishing windows"

I I I I I 75° 70 65 60

Figure 19. Management areas and boundaries relevant to the squid fisheries (200-mile limit not shown).

58 References Burukovsky, R.N., Yu.M. Froerman, and Ch.M. Nigma- tullin. 1984. Reproductive biology and scale of maturity Anon. 1975. Int. Comm. Northwest At!. Fish. Redbook. stage of reproductive system of female squid (Illex illece- 127 p. brosus). Northwest Ail. Fish. Organ. Sci. Coun. Res. Doc. 84(IX/120): 4 p. Anon. 1978. Final environmental impact statement/ fishery Caddy, J.F. management plan for the squid fishery of the Northwest 1983. The Cephalopods: Factors relevant to Atlantic Ocean. Mid-Atlantic Fishery Management Coun- their population dynamics and to the assessment and management cil. (Room 2115, Federal Building, 300 South New St., Do- of stocks. in Advances in assessment of world ce- ver, Del. 19901-6790). 91-96. phalopod resources. Progrès réalises dans l'évaluation des ressources mondiales de céphalopodes. Progresos realizados en Anon. 1979-80. Northwest At!. Fish. Organ. Sci. Coun. la evaluaci6n mundial de cefal6podos. J. F. Caddy Rep. 189 p. (ed.). FAO Fish. Tech. Pap./FAO, Doc. Tech. Pêches/FAO, Doc. Tec. Pesca 231: 416-452. Anon. 1984. Northwest Atl. Fish. Organ. Sci. Coun. Rep. 126 p. Coelho, M.L., H. Dupouy, and T. Amaratunga. 1982. Some histological observations on gonadal development of Anon. 1987. Yearbook of fishery statistics. Annuaire statis- Illex illecebrosus (Le Sueur). Northwest Atl. Fish. Organ. tique des pêches. Annuario estadistico de pesca. 1985 Sci. Coun. Res. Doc. 82(VI/30): 13 p. catches and landings. FAO, Rome. Vol 60. 463 p. Coelho, M.L., and R.K. O'Dor. 1984. Male sexual features Amaratunga, T. 1980. Growth and maturation patterns of of Illex illecebrosus and maturity staging. Northwest At!. Fish. Organ. the short-finned squid (Illex illecebrosus) on the Scotian Sci. Coun. Res. Doc. 84(IX/101): 22 p. Shelf. Northwest AU. Fish. Organ. Sci. Coun. Res. Doc. 80(11/30): 17 p. Dawe, E.G. 1981. Development of the Newfoundland squid (Illex illecebrosus) fishery and management of the resource. J. Shellfish Res. Amaratunga, T. 1983. The role of cephalopods in the ma- 1(1): 137-142. rine ecosystem. in Advances in assessment of world cephalopod resources. Progrès réalises dans l'évaluation des Dawe, E. G., and P. C. Beck. 1985a. Distribution and size ressources mondiales de céphalopodes. Progresos realiza- of short-finned squid (Illex illecebrosus) larvae in the northwest dos en la evaluaci6n mundial de cefal6podos. J. F. Caddy Atlantic from winter surveys in 1969, 1981, and 1982. (ed.). FAO Fish. Tech. Pap./FAO, Doc. Tech. Pêches/FAO, J. Northwest At!. Fish. Sci. 6(2): 43-45. Doc. Tec. Pesca 231: 379-415. Dawe, E. G., and P. C. Beck. 1985b. Distribution and size of juvenile short-finned squid Amaratunga, T., and R.D. Durward. 1979. Standardiza- (Illex illecebrosus) (Mollus- tion of data collection for the short-finned squid, Illex illec- ca: Cephalopoda) south of Newfoundland during winter. Vie et Milieu ebrosus. Int. Comm. Northwest Atl. Fish. Sel. Pap. No. 5: 35(3/4): 139-147. 37-41. Dawe, E.G., P.C. Beck, H.J. Drew, and G.H. Winters. Amaratunga, T., J.D. Neilson, D.J. Gillis, and L.G. Val- 1981a. Long-distance migration of a short-finned squid (Il- dron. 1979. Food and feeding of the short-finned squid, Il- lei illecebrosus). J. Northwest. At!. Fish. Sci. 2: 75-76. lex illecebrosus, on the Scotian Shelf in 1978. lot. Comm. Northwest At!. Fish. Res. Doc. 79(11/11): 24 p. Dawe, E.G., and H.J. Drew. 1981. Historical records of mature female short-finned squid (Illex illecebrosus) from Amaratunga, T., T. Rowell, and M. Roberge. 1980. the northwest Atlantic and the first record of a mature fe- Summary of joint Canada/U.S.S.R. research program on male captured inshore at Newfoundland. J. Northwest Atl. Fish. Organ. short-finned squid (Illex illecebrosus), 16 February to 4 Sci. 2: 61-65. June 1979: spawning stock and larval survey. Northwest Dawe, E.G., Lilly, Atl. Fish. Organ. Sci. Coun. Res. Doc. 80(11/38): 36 p. G.R. and H.J. Drew. 1983. Predation by short-finned squid (Illex illecebrosus) in Newfoundland inshore waters. Arnold, J.M., W.C. Summers, D.L. Gilbert, R.S. Man- Northwest Atl. Fish. Organ. Sci. Coun. Res. Doc. alis, N.W.Daw, and R.J.Lasek. 1974. A guide to labora- 83(IX/74): 16 p. tory use of the squid Loligo pealei. Marine Biological La- boratory, Woods Hole, Mass. 74 p. Dawe, E.G, G.R. Lilly, and J.A. Moores. 1981 b. An examination of the influence of squid (Illex illecebrosus) on Bertin, J. 1981. Graphics and graphic information- recruitment in several finfish stocks off eastern Newfound- land and Labrador. Northwest Processing. Transi. by W. J Berg and P Scott - Walter de At!. Fish. Organ. Sci. Coun. Res. Doc. Gruyter, New York. 273 p. 81(V1/25): 14 p.

59 Durward, R.D., T. Amaratunga, and R.K. O'Dor. 1979. Juanico, M. 1983. Squid maturity scales for population Maturation index and fecundity for female squid, IIlex il- analysis. in Advances in assessment of world cephalopod lecebrosus (LeSueur, 1821). Int. Comm. Northwest Atl. resources. Progrès réalises clans l'évaluation des ressources Fish. Res. Bull. No. 14: 67-72. mondiales de céphalopodes. Progresos realizados en la ev- aluaci6n mundial de cefalcipodos. J. F. Caddy (cd.). FAO Durward, R.D., E. Vessey, R.K. O'Dor, and T. Amara- Fish. Tech. Pap./FAO, Doc. Tech. Pêches/FAO, Doc. Tec. tunga. 1980. Reproduction in the squid, IIlex illecebrosus: Pesca 231: 341-378. first observations in captivity and implications for the life cycle. Int. Comm. Northwest Atl. Fish. Sel. Pap. No. 6: 7- Lange, A.M.T. 1980. The biology and population dynam- 13. ics of the squids, Loligo pealei (LeSueur) and Illex illecebrosus (LeSueur), from the Northwest Atlantic. M.Sc. the- Ennis, G.P., and P.W. Collins. 1979. Food and feeding of sis, Univ. of Wash., Seattle Wash., 178 p. the short-finned squid, Illex illecebrosus, during its seasonal occurrence in the Newfoundland area. Int. Comm. North- Lange, A.M.T. 1984a. An assessment of the long-finned west Atl. Fish. Sel. Pap. No. 5: 25-29. squid resource off the northeastern United States, autumn 1984. NMFS, NEFC, Woods Hole Lab. Ref. Doc. No. 84- Fedulov, P.P., and Y.M. Froerman. 1980. Effect of abiot- 3'7. 24 p.

ic factors on distribution of young shortfin squids, Illex illecebrosus (LeSueur, 1821). Northwest Atl. Fish. Organ. Lange, A.M.T. 1984b. Status of the short-finned squid (V- Sci. Coun. Res. Doc. 80(VI/98): 22 p. lex illecebrosus) off the northeastern U.S.A., November 1984. NMFS, NEFC, Woods Hole Lab. Ref. Doc. No. 84- Froerman, Y.M. 1984. Feeding spectrum and trophic rela- 38. 20 p. tionships of short-finned squid (11lex illecebrosus) in the Northwest Atlantic. Northwest Atl. Fish. Organ. Sci. Coun. Lange, A.M.T., and M.P. Sissenwine. 1983. Squid re- Studies 7: 67-76. sources of the northwest Atlantic. in Advances in assessment of world cephalopod resources. Progrès réalises dans Hamabe, M., C. Hamuro, and M. Ogura. 1982. Squid l'évaluation des ressources mondiales de céphalopodes. jigging from small boats. FAO Fishing Manuals. Fishing Progresos realizados en la evaluaci6n mundial de ce- News Books Ltd. 74 p. fal6podos. J. F. Caddy (ed.). FAO Fish. Tech. Pap./FAO, Doc. Tech. Pêches/FAO, Doc. Tee. Pesca 231: 21-54. Hatanaka, H., S. Kawahara, Y. Uozumi, and S. Kasaha- ra. 1985. Comparison of life cycles of five ommastrephid Lilly, G.R., and D.R. Osborne. 1984. Predation by Atlan- squids fished by Japan: Todarodes pacifïcus, IIlex illece- tic cod (Gadus morhtta) on short-finned squid (Illex illecebrosus,lex argentinus, Nototodarus sloani sloani and brosus) off eastern Newfoundland and in the northeastern Nototoclarus sloani gouldi. Northwest Atl. Fish. Organ. Gulf of St. Lawrence. Northwest Atl. Fish. Organ. Sci. Sci. Coun. Studies 9: 59-68. Coun. Res. Doc. 84 (IX1 108): 16 p.

Hayashi, S. 1960. Development of the squid, °minas- Lipinski, M. 1977. The place of squids in the biological trephes sloani pacificus (Steenstrup). Nagasaki Daigaku and fishery structure of the world ocean. in Squid sympo- Suisangaktibu Kenkyo Hokoku / Bull. Fac. Fish. Hagasaki sium. June 5-6. 1973, Gydnia. M. Lipinski (Ed.). Trans. Univ. 9: 43-48. from Polish. Foreign Scientific Publications Dept., Nat. Centre for Scientific, Technical and Economic Information. Hirtle, R.M.W., M.E. Demont and R.K. O'Dor. 1981. 173 p. Feeding, growth and metabolic rates in captive short-finned squid, Illex illecebrosus, in relation to the natural popula- Maurer, R.O., and R.E. Bowman. 1985. Food consumption. J. Shellfish Res. 1(2): 187-192. tion of squids (Il/ex illecebrosus and Loligo pealei) off the northeastern United States. Northwest AU. Fish. Organ. Sci. Hixon, R.F., R.T. Hanlon and W.H. Hulet. 1981. Growth Coun. Studies 9: 117-124. and maximal size of the long-finned squid Loligo pealei in the northwestern Gulf of Mexico. J. Shellfish Res. 1(2): Mercer, M.C. 1973a. Sexual maturity and sex ratios of the 181-185. ommastrephid squid, Illex illecebrosus, at Newfoundland (Subarea 3). Int. Comm. Northwest Atl, Fish. Res. Doc. 73/ Hurley, G.V. 1980. Recent developments in the squid, Illex 71: 14 p. illecebrosus, fishery of Newfoundland, Canada. Mar. Fish. Rev. 42(7-8): 15-22. Mercer, M.C. 1973b. Nominal catch of squid in Canadian Atlantic waters (Subareas 2-4), 1920 to 1968. lut. Comm. Northwest Atl. Fish. Res. Doc. 73/73: 10 p.

60 Mercer, M.C. 1975. Modified Leslie-Delury population Rowell, T.W. 1986. Management units and the squid re- models of the long-finned pilot whale (Globicephala rne- sources of the Gulf of Maine, IIlex illecebrosus and Loligo laena) and annual production of the short-finned squid (II- pealei . Can. Atl. Fish. Sci. Adv. Comm. Res. Doc. 86/59: lex illecebrosus) based upon their interaction at Newfound- 21 p. land. J. Fish. Res. Bd. Can. 32(7): 1145-1154. Rowell, T.W., and R.W. Trites. 1985. Distribution of lar- Mesnil, B. 1977. Growth and life cycle of squid, Loligo val and juvenile IIlex (Mollusca: Cephalopoda) in the Blake pealei and IIlex illecebrosus, from the Northwest Atlantic. Plateau region (Northwest Atlantic). Vie et Milieu 35(3/4): Int. Comm. Northwest Atl. Fish. Sel. Pap. No. 2: 55-69. 149-161.

Mohn, R.K. 1981. Indices of abundance and bottom tem- Rowell, T.W., R.W. Trites, and E.G. Dawe. 1985a. Dis- perature relationships for squid in Div. 4VWX. Northwest tribution of short-finned squid (IIlex illecebrosus) larvae Atl. Fish. Organ. Sci. Coun. Res. Doc. 81(IX/99): 10 p. and juveniles in relation to the Gulf Stream frontal zone between Florida and Cape Hatteras. Northwest Atl. Fish. Or- Naef, A. 1923. Die cephalopoden. Fauna Flora Golfo Nap- gan. Sci. Coun. Studies 9: 77-92. oli Monographia, 35(1). plates 9-12. Rowell, T.W., J.H. Young, J.C. Poulard and J.P.Robin. O'Dor, R.K. 1983. Illex illecebrosus. in Cephalopod Life 1985b. Changes in distribution and biological characteris- Cycles Vol. 1. P.R. Boyle (ed.). Academic Press 1983. p. tics of IIlex illecebrosus on the Scotian Shelf, 1980-83. 175-199. Northwest Atl. Fish. Organ. Sci. Coun. Studies 9: 11-26.

O'Dor, R.K., and N. Balch. 1985. Properties of IIlex illec- Schuldt, M. 1979. Contribuckin al conocimiento del ciclo ebrosus egg masses potentially influencing larval oceano- reproductor de IIlex argentious. Monogr. Commun. Invest. graphic distribution. Northwest Atl. Fish. Organ. Sci. Cient., Argent. 10: 110 p. Coun. Studies 9: 69-76. Scott, J.S. 1978. Distribution of squid, /Hex illecebrosus, O'Dor, R.K., N. Balch, E.A. Foy, R.W.M. Hirtle, D.A. on the Scotian Shelf 1970-76. Int. Comm. Northwest Atl. Johnston, and T. Amaratunga. 1982. Embryonic devel- Fish. Sel. Pap. No. 3: 93-96. opment of the squid, IIlex illecebrostts, and effect of temperature on development rates. J. Northwest Atl. Fish. Sci. Squires, H.J. 1957. Squid, Illex illecebrosus (LeSueur), in 3(1): 41-45. the Newfoundland fishing area. J. Fish. Res. Board Cati. 14(5): 693-728. O'Dor, R.K., R.D. Durward, E. Vessey, and T. Amara- tunga. 1979. Feeding and growth rates in captive squid (II- Summers, W.C. 1971. The age and growth of Loligo pea- lex illecebrosus) and the influence of food availability on lei, a population study of the common Atlantic Coast squid. growth in the natural population. Int. Comm. Northwest Biol. Bull. (Woods Hole). 141: 189-201. Atl. Fish. Res. Doc. 79(11/16): 16 p. Trites, R.W., and T.W. Rowell. 1985. Larval and juvenile O'Dor, R.K., E. Vessey, and T. Amaratunga. 1980. Fac- distribution of the short-finned squid (Illex illecebrosus) in tors affecting fecundity and larval distribution in the squid, the Cape Hatteras-Florida Straits area in the December- lex illecebrosus. Northwest Atl. Fish. Organ. Sci. Coun. January period, 1984-1985. Northwest Atl. Fish. Organ. Res. Doc. 80(11/39): 9 p. Sci. Coun. Res. Doc. 85(VI/69): 36 p.

Roper, C.F.E., and C.C. Lu. 1978. Rhynchoteuthion lar- Vecchione, M. 1978. Larval development of Illex (Steen- vae of Ornmastrephid squids of the western North Atlantic, strup, 1880), in the northwestern Atlantic, with comments with the first description of larvae and juveniles of IIlex il- on Illex larval distribution. Proc. Biol. Soc. Wash. 91(4): lecebrosus. Proc. Biol. Soc. Wash. 91(4): 1039-1059. 1060-1075.

Roper, C.F.E., C.C. Lu, and K. Mangold. 1969. A new Vinogradov, V.I. 1984. On feeding of two squid species in species of IIlex from the western Atlantic and distribution the northwest Atlantic. Northwest Atl. Fish. Organ. Sci. aspects of other IIlex sp. (Cephalopoda: Oegopsida). Proc. Coun. Res. Doc. 84(IX/117): 7 p. Biol. Soc. Wash. 83: 295-322. Voss, G.L. 1956. A review of the cephalopods of the Gulf Roper, C.F.E., M.J. Sweeney and C.E. Nauen. 1984. of Mexico. Bull. Mar. Sci. Gulf Caribb. 6: 85-178. FAO species catalogue. Vol. 3. Cephalopods of the world. An annotated and illustrated catalogue of species of interest Voss, G.L. 1983. A review of cephalopod fisheries biology. to fisheries. FAO Fish. Synop., 125 Vol. 3: 277p. Memoirs of the National Museum Victoria, 44: 229-241.

61 Voss, G.L., and T.F. Brakoniecki. 1985. Squid resources Webber, D.M. and R.K. O'Dor. 1985. Respiration and of the Gulf of Mexico and southeast Atlantic coasts of the swimming performance of short-finned squid (IIlex illece- United States. Northwest Atl. Fish. Organ. Sci. Coun. brosus). Northwest At!. Fish. Organ. Sci. Coun. Studies 9: Studies 9: 27-38. 133-138.

Vovk, A.N. 1974. The position of the longfin squid Loligo Whitaker, J.D. 1980. Squid catches resulting from trawl pealei (LeSueur) in the ecosystem. Tr. Atl. Nanchno Issled. surveys off the southeastern United States. Mar. Fish. Rev. Inst. Rybn. Khoz. Okeanogr. 58168-:176. English trans. 42 (7-8): 39-43. Fish. Mar. Ser. Trans. ser. No. 3977. Wiseman, E. 1982. Annex to the Worldwide Fisheries Vovk, A.N. 1983. Food relations of long-finned squid, Lo- Marketing Study. Prospects to 1985. Dept. of Fish. and ligo pealei (LeSueur), in the Northwest Atlantic and its po- Oceans Fish. Econ. Devel. & Mark. Rep.: 109 p. sition in the ecosystem. Northwest Atl. Fish. Organ. Sci. Coun. Res. Doc. 83(IX/83): 20 p. Worms, J. 1983. World fisheries for Cephalopods: a sy- noptic overview. in Advances in assessment of world ce- Waldron, D.E. 1983. Factors influencing Scotian Shelf phalopod resources. Progrès réalises dans l'évaluation des finfish and squid interactions with special reference to sil- ressources mondiales de céphalopodes. Progresos realiza- ver hake. Northwest Atl. Fish. Organ. Sci. Coun. Res. Doc. dos en la evaluaci6n mundial de cefal6podos. J. F. Caddy 83(IX/92): 28 p. (ed.). FAO Fish. Tech. Pap./FAO, Doc. Tech. Pêches/FAO, Doc. Tee. Pesca 231: 1-20. Wallace, I.C., R.K. O'Dor, and T. Amaratunga. 1981. Sequential observations on the digestive process in the squid, IIlex illecebrosus. Northwest Atl. Fish. Organ. Sci. Coun. Studies 1: 65-69.

62 DUE DATE tiCot ab_ot

201-6503 Printed in USA