NOAA Technical Memorandum NMFS-NE-191

U. S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

Northeast Fisheries Science Center

Woods Hole, Massachusetts

November 2004

Essential Fish Habitat Source Document:

Northern Shortfin Squid, Illex illecebrosus,

Life History and Habitat Characteristics

Second Edition
167. Assessment and Characterization of Salt Marshes in the Arthur Kill (New York and New Jersey) Replanted after a Severe

Oil Spill. By David B. Packer, ed. December 2001. x + 228 p., 54 figs., 58 tables, 6 app. NTIS Access. No. PB2004-106067.

168. U.S. Atlantic and Gulf of Mexico Marine Mammal Stock Assessments – 2001. By Gordon T. Waring, Janeen M. Quintal, and

Steven L. Swartz, eds., Phillip J. Clapham, Timothy V.N. Cole, Carol P. Fairfield, Aleta Hohn, Debra L. Palka, Marjorie C. Rossman, U.S.

Fish and Wildlife Service, and Cynthia Yeung, contribs. December 2001. vii + 310 p., 43 figs., 54 tables, 2 app. NTIS Access. No. PB2002-

104350.

169. U.S. Atlantic and Gulf of Mexico Marine Mammal Stock Assessments – 2002. By Gordon T. Waring, Janeen M. Quintal, and

Carol P. Fairfield, eds., Phillip J. Clapham, Timothy V.N. Cole, Lance P. Garrison, Georgia Department of Natural Resources, Aleta A. Hohn,

Blair G. Maise, Wayne E. McFee, Debra L. Palka, Patricia E. Rosel, Marjorie C. Rossman, U.S. Fish and Wildlife Service, Frederick W. Wenzel,

and Cynthia Yeung, contribs. September 2002. vii + 318 p., 42 figs., 56 tables, 4 app. NTIS Access. No. PB2003-104167.

170. Interaction of Shelf Water with Warm-Core Rings, Focusing on the Kinematics and Statistics of Shelf Water Entrained

within Streamers. By Ronald J. Schlitz. March 2003. v + 35 p., 26 figs., 3 tables. NTIS Access. No. PB2005-100158.

171. Length-Weight Relationships for 74 Fish Species Collected during NEFSC Research Vessel Bottom Trawl Surveys. By

Susan E. Wigley, Holly M. McBride, and Nancy J. McHugh. March 2003. vi + 26 p., 1 fig., 3 tables. NTIS Access. No. PB2003-106486.

172. Variability of Temperature and Salinity in the Middle Atlantic Bight and Gulf of Maine Based on Data Collected as

Part of the MARMAP Ships of Opportunity Program, 1978-2001. By Jack W. Jossi and Robert L. Benway. March 2003. vi + 92

p., 74 figs., 5 tables. NTIS Access. No. PB2004-100835.

173. Essential Fish Habitat Source Document: Barndoor Skate, Dipturus laevis, Life History and Habitat Characteristics. By

David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano. March 2003. v + 23 p., 14 figs., 1 table. NTIS Access. No. PB2003-104257.

174. Essential Fish Habitat Source Document: Clearnose Skate, Raja eglanteria, Life History and Habitat Characteristics.

By David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano. March 2003. v + 50 p., 25 figs., 2 tables. NTIS Access. No. PB2003-

104260.

175. Essential Fish Habitat Source Document: Little Skate, Leucoraja erinacea, Life History and Habitat Characteristics. By

David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano. March 2003. v + 66 p., 27 figs., 2 tables. NTIS Access. No. PB2003-104259.

176. Essential Fish Habitat Source Document: Rosette Skate, Leucoraja garmani virginica, Life History and Habitat

Characteristics. By David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano. March 2003. v + 17 p., 11 figs., 1 table. NTIS Access.

No. PB2003-104258.

177. Essential Fish Habitat Source Document: Smooth Skate, Malacoraja senta, Life History and Habitat Characteristics. By

David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano. March 2003. v + 26 p., 15 figs., 1 table. NTIS Access. No. PB2003-104256.

178. Essential Fish Habitat Source Document: Thorny Skate, Amblyraja radiata, Life History and Habitat Characteristics. By

David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano. March 2003. v + 39 p., 18 figs., 2 tables. NTIS Access. No. PB2003-104255.

179. Essential Fish Habitat Source Document: Winter Skate, Leucoraja ocellata, Life History and Habitat Characteristics.

By David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano. March 2003. v + 57 p., 25 figs., 2 tables. NTIS Access. No. PB2003-

104254.

180. Variability in Blood Chemistry of Yellowtail Flounder, Limanda ferruginea, with Regard to Sex, Season, and Geographic

Location. By Renee Mercaldo-Allen, Margaret A. Dawson, Catherine A. Kuropat, and Diane Kapareiko. September 2003. vi + 20 p., 1

fig., 10 tables. NTIS Access. No. PB2004-107074.

181. In press.

182. U.S. Atlantic and Gulf of Mexico Marine Mammal Stock Assessments – 2003. By Gordon T. Waring, Richard M. Pace, Janeen

M. Quintal, Carol P. Fairfield, and Katherine Maze-Foley, eds., Nicole Cabana, Phillip J. Clapham, Timothy V.N. Cole, Gregory L. Fulling,

Lance P. Garrison, Aleta A. Hohn, Blair G. Maise, Wayne E. McFee, Keith D. Mullin, Debra L. Palka, Patricia E. Rosel, Marjorie C. Rossman,

Frederick W. Wenzel, and Amy L. Whitingham, contribs. May 2004. vii + 287 p., 47 figs., 58 tables, 4 app., index. NTIS Access. No.

PB2004-106431.

Recent Issues in This Series:
U. S. DEPARTMENT OF COMMERCE

Donald L. Evans, Secretary

National Oceanic and Atmospheric Administration

Vice Admiral Conrad C. Lautenbacher, Jr., USN (ret.), Administrator

National Marine Fisheries Service

William T. Hogarth, Assistant Administrator for Fisheries

Northeast Fisheries Science Center

Woods Hole, Massachusetts

November 2004

This series represents a secondary level of scientific publishing. All issues employ

thorough internal scientific review; some issues employ external scientific review.

Reviews are -- by design -- transparent collegial reviews, not anonymous peer

reviews. All issues may be cited in formal scientific communications.

NOAA Technical Memorandum NMFS-NE-191

Lisa C. Hendrickson and Elizabeth M. Holmes

National Marine Fisheries Serv., 166 Water St., Woods Hole, MA 02543

Essential Fish Habitat Source Document:

Northern Shortfin Squid, Illex illecebrosus,

Life History and Habitat Characteristics

Second Edition
Editorial Notes on "Essential Fish Habitat Source Documents"

Issued in the NOAA Technical Memorandum NMFS-NE Series

Editorial Production

For "Essential Fish Habitat Source Documents" issued in the NOAA Technical Memorandum NMFS-NE series, staff

of the Northeast Fisheries Science Center's (NEFSC's) Ecosystems Processes Division largely assume the role of staff of

the NEFSC's Editorial Office for technical and copy editing, type composition, and page layout. Other than the four covers

(inside and outside, front and back) and first two preliminary pages, all preprinting editorial production is performed by,

and all credit for such production rightfully belongs to, the staff of the Ecosystems Processes Division.

Internet Availability and Information Updating

Each original issue of an "Essential Fish Habitat Source Document" is published both as a paper copy and as a Web

posting. The Web posting, which is in "PDF" format, is available at: http://www.nefsc.noaa.gov/nefsc/habitat/efh.

Each issue is updated at least every five years. The updated edition will be published as a Web posting only; the

replaced edition(s) will be maintained in an online archive for reference purposes.

Species Names

The NMFS Northeast Region's policy on the use of species names in all technical communications is generally to

follow the American Fisheries Society's lists of scientific and common names for fishes ( i.e., Robins et al. 1991aa

, bb),

mollusks (i.e., Turgeon et al. 1998c

), and decapod crustaceans ( i.e., Williams et al. 1989d), and to follow the Society for

Marine Mammalogy's guidance on scientific and common names for marine mammals (i.e., Rice 1998e

). Exceptions to this

policy occur when there are subsequent compelling revisions in the classifications of species, resulting in changes in the

names of species ( e.g., Cooper and Chapleau 1998f

; McEachran and Dunn 1998g).

aRobins, C.R. (chair); Bailey, R.M.; Bond, C.E.; Brooker, J.R.; Lachner, E.A.; Lea, R.N.; Scott, W.B. 1991a. Common and scientific names

of fishes from the United States and Canada. 5th ed. Amer. Fish. Soc. Spec. Publ. 20; 183 p.

bRobins, C.R. (chair); Bailey, R.M.; Bond, C.E.; Brooker, J.R.; Lachner, E.A.; Lea, R.N.; Scott, W.B. 1991b. World fishes important to

North Americans. Amer. Fish. Soc. Spec. Publ. 21; 243 p. cTurgeon, D.D. (chair); Quinn, J.F., Jr.; Bogan, A.E.; Coan, E.V.; Hochberg, F.G.; Lyons, W.G.; Mikkelsen, P.M.; Neves, R.J.; Roper, C.F.E.;

Rosenberg, G.; Roth, B.; Scheltema, A.; Thompson, F.G.; Vecchione, M.; Williams, J.D. 1998. Common and scientific names of aquatic

invertebrates from the United States and Canada: mollusks. 2nd ed. Amer. Fish. Soc. Spec. Publ. 26; 526 p. dWilliams, A.B. (chair); Abele, L.G.; Felder, D.L.; Hobbs, H.H., Jr.; Manning, R.B.; McLaughlin, P.A.; Pérez Farfante, I. 1989. Common

and scientific names of aquatic invertebrates from the United States and Canada: decapod crustaceans. Amer. Fish. Soc. Spec. Publ. 17;

77 p.

e

Rice, D.W. 1998. Marine mammals of the world: systematics and distribution. Soc. Mar. Mammal. Spec. Publ. 4; 231 p. f

Cooper, J.A.; Chapleau, F. 1998. Monophyly and interrelationships of the family Pleuronectidae (Pleuronectiformes), with a revised

classification. Fish. Bull. (Washington, DC) 96:686-726. gMcEachran, J.D.; Dunn, K.A. 1998. Phylogenetic analysis of skates, a morphologically conservative clade of elasmobranchs

(Chondrichthyes: Rajidae). Copeia 1998(2):271-290.
Page iii

PREFACE TO SECOND EDITION

One of the greatest long-term threats to the viability of 

commercial and recreational fisheries is the continuing 

loss of marine, estuarine, and other aquatic habitats.

Magnuson-Stevens Fishery Conservation and

Management Act (October 11, 1996)

The long-term viability of living marine resources

depends on protection of their habitat.

NMFS Strategic Plan for Fisheries Research

(February 1998)

The Magnuson-Stevens Fishery Conservation and 

Management Act (MSFCMA), which was reauthorized

and amended by the Sustainable Fisheries Act (1996), 

requires the eight regional fishery management councils 

to describe and identify essential fish habitat (EFH) in 

their respective regions, to specify actions to conserve 

and enhance that EFH, and to minimize the adverse 

effects of fishing on EFH. Congress defined EFH as 

“those waters and substrate necessary to fish for

spawning, breeding, feeding or growth to maturity.”

The MSFCMA requires NOAA Fisheries to assist the 

regional fishery management councils in the

implementation of EFH in their respective fishery

management plans.

NOAA Fisheries has taken a broad view of habitat 

as the area used by fish throughout their life cycle. Fish 

use habitat for spawning, feeding, nursery, migration, 

and shelter, but most habitats provide only a subset of 

these functions. Fish may change habitats with changes 

in life history stage, seasonal and geographic

distributions, abundance, and interactions with other

species. The type of habitat, as well as its attributes and 

functions, are important for sustaining the production of 

managed species.

The Northeast Fisheries Science Center compiled 

the available information on the distribution,

abundance, and habitat requirements for each of the

species managed by the New England and Mid-Atlantic

Fishery Management Councils. That information is

presented in a series EFH species reports (plus one

consolidated methods report). The EFH species reports 

are a survey of the important literature as well as

original analyses of fishery-independent data sets from 

NOAA Fisheries and several coastal states. The species 

reports are also the source for the current EFH

designations by the New England and Mid-Atlantic

Fishery Management Councils, and understandably

have begun to be referred to as the “EFH source

documents.”

NOAA Fisheries provided guidance to the regional 

fishery management councils for identifying and

describing EFH of their managed species. Consistent 

with this guidance, the species reports present

information on current and historic stock sizes,

geographic range, and the period and location of major 

life history stages. The habitats of managed species are 

described by the physical, chemical, and biological

components of the ecosystem where the species occur.

Information on the habitat requirements is provided for 

each life history stage, and it includes, where available, 

habitat and environmental variables that control or limit 

distribution, abundance, growth, reproduction,

mortality, and productivity.

The initial series of EFH species source documents

were published in 1999 in the NOAA Technical

Memorandum NMFS-NE series. Updating and review 

of the EFH components of the councils’ Fishery

Management Plans is required at least every 5 years by 

the NOAA Fisheries Guidelines for meeting the

Sustainable Fisheries Act/EFH Final Rule. The second

editions of these species source documents were written 

to provide the updated information needed to meet

these requirements. The second editions provide new

information on life history, geographic distribution, and 

habitat requirements via recent literature, research, and 

fishery surveys, and incorporate updated and revised 

maps and graphs. This second edition of the northern

shortfin squid EFH source document is based on the 

original by Luca M. Cargnelli, Sara J. Griesbach, and 

Christine A. Zetlin, with a foreword by Jeffrey N. Cross 

(Cargnelli et al. 1999).

Identifying and describing EFH are the first steps 

in the process of protecting, conserving, and enhancing 

essential habitats of the managed species. Ultimately, 

NOAA Fisheries, the regional fishery management

councils, fishing participants, Federal and state

agencies, and other organizations will have to cooperate 

to achieve the habitat goals established by the

MSFCMA.
Page iv
Page v

Contents

INTRODUCTION........................................................................................................................................................................ 1

LIFE HISTORY........................................................................................................................................................................... 1

GEOGRAPHICAL DISTRIBUTION......................................................................................................................................... 3

HABITAT CHARACTERISTICS.............................................................................................................................................. 4

RESEARCH NEEDS................................................................................................................................................................... 6

ACKNOWLEDGMENTS ........................................................................................................................................................... 6

REFERENCES CITED................................................................................................................................................................ 6

Tables

Table 1. Summary of life history and habitat parameters for northern shortfin squid Illex illecebrosus........................12

Figures

Figure 1. The northern shortfin squid, Illex illecebrosus (from Goode 1884)....................................................................14

Figure 2. Hypothetical migration path of the northern shortfin squid, Illex illecebrosus..................................................15

Figure 3. Spawning area of northern shortfin squid (encircled) during late May...............................................................16

Figure 4. Percent by weight (g) of the major prey items in the diet of northern shortfin squid........................................17

Figure 5. Distribution and abundance of northern shortfin squid from Newfoundland to Cape Hatteras........................18

Figure 6. Seasonal distributions and abundances of pre-recruit northern shortfin squid collected during NEFSC 

bottom trawl surveys..............................................................................................................................................19

Figure 7. Distribution and abundance of pre-recruit northern shortfin squid in Massachusetts coastal waters...............23

Figure 8. Seasonal distributions and abundances of recruit northern shortfin squid collected during NEFSC bottom 

trawl surveys...........................................................................................................................................................24

Figure 9. Distribution and abundance of recruit northern shortfin squid in Massachusetts coastal waters......................28

Figure 10. Distributions of pre-recruit northern shortfin squid and trawls from NEFSC bottom trawl surveys relative to 

bottom water temperature, depth, and salinity......................................................................................................29

Figure 11. Distributions of pre-recruit northern shortfin squid and trawls in Massachusetts coastal waters relative to 

bottom water temperature and depth. ....................................................................................................................31

Figure 12. Distributions of recruit northern shortfin squid and trawls from NEFSC bottom trawl surveys relative to 

bottom water temperature, depth, and salinity......................................................................................................33

Figure 13. Distributions of recruit northern shortfin squid and trawls in Massachusetts coastal waters relative to 

bottom water temperature and depth. ....................................................................................................................35

Page 1

INTRODUCTION

The northern shortfin squid, Illex illecebrosus

(Figure 1), is a highly migratory species of the family 

Ommastrephidae. Distributed across a broad

geographic area, I. illecebrosus is found in the

northwest Atlantic Ocean between the Sea of Labrador 

and the Florida Straits (66oN to 29o30'N; Roper et al.

1998). Throughout its range of commercial

exploitation, from Newfoundland to Cape Hatteras,

North Carolina, the population is considered to

constitute a single stock (Dawe and Hendrickson 1998).

The southern stock component (inhabiting U.S. waters) 

is managed by the Mid-Atlantic Fishery Management 

Council, in accordance with the Atlantic Mackerel,

Squid and Butterfish Fishery Management Plan

(MAFMC 1998), and the northern stock component

(inhabiting waters between Newfoundland and Nova 

Scotia) is assessed and managed by the Northwest

Atlantic Fisheries Organization (Hendrickson et al.

2002). Both stock components are managed based on 

an annual quota.

This Essential Fish Habitat Source Document

provides information on the life history and habitat 

characteristics of northern shortfin squid. Data sources 

and methodologies used to prepare this document are 

described in Reid et al. (1999).

LIFE HISTORY

The life history characteristics of northern shortfin 

squid have been reviewed by Black et al. (1987), Perez 

(1994), and O’Dor and Dawe (1998). Most of the

supporting studies have been based on the northern

component of the stock. Like many squid species, I.

illecebrosus lives for less than one year, has a high 

natural mortality rate, and exhibits a protracted

spawning season whereby overlapping “microcohorts” 

enter the population throughout the year and exhibit 

variable growth rates (Caddy 1991; Jackson 1994). The

life cycle is comprised of oceanic and neritic

components. During spring, squid migrate onto the

continental shelf between Newfoundland and Cape

Hatteras. During late autumn, squid migrate off the

continental shelf, presumably to a winter spawning site 

(Black et al. 1987). The seasonal proportion of squid 

residing beyond the continental shelf is unknown

because this habitat is not sampled during seasonal U.S. 

and Canadian bottom trawl surveys. Little is known

about either the habitat of mature individuals

(particularly females) or the winter habitat of the

species.

The life cycle (Figure 2) proposed by Black et al.

(1987) remains hypothetical because several aspects

remain unknown. These include the location of the 

winter spawning site, migration patterns between

northern and southern stock components, the autumn 

spawning migration route, and what fraction of the

stock inhabits waters beyond the continental shelf.

New life history information regarding population 

structure, spawning location, lifespan, and age and size 

at maturity are described herein. This new information 

is based on data collected on the U.S. shelf during a 

stratified, random bottom trawl survey of the population 

(Hendrickson 2004).

EGGS AND PARALARVAE

Illex illecebrosus egg masses have never been

collected in the wild (O’Dor and Dawe 1998) but have 

been described from laboratory spawning events. The 

gelatinous egg balloons are 0.5 to 1.0 m in diameter and 

contain between 10,000 and 100,000 eggs (Durward et

al. 1980). Females can produce multiple egg masses 

(Durward et al. 1978). Mature eggs are ovoid, ranging 

from 0.9 x 0.6 to 1.0 to 0.8 mm in size, and weigh 

between 200 and 250 μg (Durward et al. 1980).

Laboratory studies indicate that hatching occurs in 

16 days at 13°C, 12 days at 16°C, and 8 days at 21°C;

normal embryonic development requires water

temperatures of at least 12.5°C (O’Dor et al. 1982b).

Paralarvae may remain within the remnants of the egg 

mass to utilize the nutrients as a food source (Durward 

et al. 1980). In the laboratory, paralarvae hatch at 

approximately 1.1 mm mantle length (ML) (Durward et

al. 1980), then enter a transitional stage at

approximately 5.0 mm ML, followed by a juvenile

stage at about 7.0 mm ML (Hatanaka 1986).

Based on the distribution of paralarvae, it is

hypothesized that the Gulf Stream serves as the primary 

transport mechanism for egg masses and paralarvae 

(O’Dor 1983; Rowell et al. 1985a). Paralarvae have

been collected during all seasons (Roper and Lu 1979), 

from south of Cape Hatteras to as far north as the tail of 

the Grand Bank (Dawe and Beck 1985; Hatanaka et al.

1985). Paralarvae are most abundant in February and 

March, in the nutrient-rich waters of the Gulf

Stream/Slope Water convergence zone; above the

thermocline at temperatures greater than 13°C

(Hatanaka et al. 1985).

I. illecebrosus paralarvae hatched in the laboratory 

were 1.10 to 1.25 mm ML (O’Dor et al. 1986). Illex sp. 

hatchlings have only been collected in waters south of 

Cape Hatteras (35.5oN) and during February through 

March (Dawe and Beck 1985; Rowell et al. 1985a).

However, species identification of Illex paralarvae is 

problematic, particularly if caught south of New Jersey, 

due to the difficulty in distinguishing between

paralarvae of I. illecebrosus and two sympatric Illex

species (Vecchione and Roper 1986).
Page 2

JUVENILES AND ADULTS

Onset of the juvenile stage at 8 to 10 mm ML is

indicated by a separation of the proboscis into a pair of 

tentacles (Roper and Lu 1979). Juveniles collected in 

surveys conducted in the Gulf Stream and continental 

slope waters during February through May ranged in 

size from 10 to 94 mm ML (O’Dor 1983). During late 

spring, juveniles migrate onto the continental shelf

between Nova Scotia and Cape Hatteras (O’Dor 1983; 

Black et al. 1987).

Juveniles caught on the continental shelf in late 

May ranged in size from 34 to 68 mm ML. Gonadal

development began at about 64 mm ML in males and at

74 mm ML in females. Males attained 50% maturity at 

a smaller size and older age than females, but these 

differences were not statistically significant. Size- and 

age-at-maturity increased with latitude and were

correlated with decreases in water temperature

(Hendrickson 2004). Mean size at maturity may also 

vary inter-annually (Coelho and O’Dor 1993). In

inshore Newfoundland waters, the percentage of mature 

males and male gonadosomatic indices were

significantly higher for squid hatched in May than

during March or April. Although females do not

become mature in inshore Newfoundland waters, those 

hatched in May were more mature than those hatched in 

March or April (Dawe and Beck 1997). Captive

females matured within 40 to 60 days (O’Dor et al.

1977).

In Nova Scotian and Newfoundland waters, males 

mature at a faster rate than females and are believed to 

emigrate during autumn from continental shelf fishing 

areas before females (Black et al. 1987). Evidence for 

this phenomenon is a seasonal decline in the percentage 

of males collected on the Scotian Shelf during some 

years (Amaratunga 1980a). However, a reduction in the 

proportion of males, which tend to be small individuals, 

can also result from cannibalism by larger females

(O’Dor and Dawe 1998). During late autumn, nearlymature squid migrate from all continental shelf fishing 

areas (Hurley 1980; Black et al. 1987), presumably to 

spawn. Most mature females have been collected from 

the U.S. shelf (Hendrickson 2004), but four have also 

been recorded from waters off the coast of

Newfoundland and Nova Scotia (Dawe and Drew

1981).

Age estimation, accomplished by counting daily

growth increments in the statoliths, has been validated 

for I. illecebrosus (Dawe et al. 1985; Hurley et al.

1985). Increment counts of statoliths from mated

females caught in the Mid-Atlantic Bight indicate a

lifespan of about 115 to 215 days (Hendrickson 2004).

Squid inhabiting warmer waters of the Mid-Atlantic

Bight exhibit faster rates of growth and maturation, and 

possibly a shorter lifespan, than squid from the northern 

stock component (Hendrickson 2004). The species may 

achieve a maximum size of 35 cm ML and 700 g, with 

females achieving larger sizes than males (Hendrickson 

1998; O’Dor and Dawe 1998).

The terms recruit and pre-recruit are used herein to 

describe geographical distributions and habitat

characteristics for the exploited and unexploited

portions of the stock, respectively. Exploitation occurs 

at a minimum mantle length of 10 cm ML, the

approximate length at which individuals migrate onto 

the continental shelf (O’Dor 1983; Hendrickson et al.

1996). Pre-recruits and recruits are thus defined as

individuals ≤ 10 cm ML and ≥ 11 cm ML, respectively.

REPRODUCTION

Mating and spawning have only been observed in 

captivity. I. illecebrosus is a semelparous, terminal

spawner whereby spawning and death occur within

several days of mating (O’Dor 1983). Mature females

collected on the U.S. continental shelf had mated with 

as many as four males (Hendrickson 2004).

Until recently, few mature females and only two 

mated individuals have been recorded (Dawe and Drew 

1981). However, back-calculations of hatch dates based

on statolith age analyses indicate that spawning occurs 

during October through June (Dawe and Beck 1997; 

Hendrickson 2004).

A winter spawning area, located off the east coast 

of Florida in the vicinity of the Blake Plateau (Figure 2;

Black et al. 1987), has been inferred based on: the

presence of Illex sp. hatchlings along the north wall of 

the Gulf Stream during January and February (Dawe 

and Beck 1985; Rowell et al. 1985a; Vecchione and 

Roper 1986; Coelho and O’Dor 1993); the offshore

migration of adults in late autumn (Black et al. 1987); 

and the presence of minimum water temperatures

required for hatching (O’Dor et al. 1982b; Trites 1983; 

Rowell and Trites 1985).

The only confirmed spawning area is located in the 

Mid-Atlantic Bight, at depths of 113 to 377 m, where a 

large number of mated females were collected between 

39o10 ́N and 35o50 ́N during late May (Figure 3;

Hendrickson 2004). This spawning area overlaps

spatially with the fishing grounds of the directed fishery 

(Hendrickson et al. 1996). Spawning may also occur 

offshore in the Gulf Stream/Slope Water frontal zone, 

where paralarvae have been collected (O’Dor and Balch 

1985; Rowell et al. 1985a), or south of Cape Hatteras 

during winter where Illex sp. hatchlings have been

collected (Dawe and Beck 1985). Previous reports of 

mated females consist of three individuals that were 

caught south of Georges Bank (Dawe and Drew 1981).
Page 3

DIET

Trophic relationships between I. illecebrosus and 

other marine species are described by Dawe and

Brodziak (1998). Northern shortfin squid feed primarily 

on fish and crustaceans, but cannibalism of small

individuals (most likely males) by larger females also

occurs, particularly during autumn (Squires 1957;

Froerman 1984; Maurer and Bowman 1985; Dawe 

1988). An ontogenetic shift in diet from a

predominance of crustaceans to a predominance of fish 

and squid is evident in squid from both stock

components (Maurer and Bowman 1985; Dawe 1988).

Fish prey consists of the early life history stages of 

Atlantic cod, Arctic cod and redfish (Squires 1957,

Dawe et al. 1997), sand lance (Dawe et al. 1997),

mackerel and Atlantic herring (O’Dor et al. 1980a;

Dawe et al. 1997), and haddock and sculpin (Squires 

1957). Illex also feed on adult capelin (Squires 1957; 

O’Dor et al. 1980a; Dawe et al. 1997) and longfin

inshore squid, Loligo pealeii (Vinogradov 1984).

Illex exhibit diel vertical migrations (Roper and

Young 1975; Brodziak and Hendrickson 1999) and

both juveniles (Arkhipkin and Fedulov 1986) and adults 

feed primarily at night in the upper layers of the water 

column (Maurer and Bowman 1985). On the U.S. shelf

in the spring, I. illecebrosus primarily consume

euphausiids, whereas fish and squid were the dominant 

prey in the summer and fall. I. illecebrosus 6-10 cm and 

26-30 cm in size eat mostly squid, while 11-15 cm Illex

eat mostly crustaceans and fish, and individuals 16-20

cm eat mostly crustaceans (Maurer and Bowman 1985).

Illex gut content data collected during Northeast 

Fisheries Science Center (NEFSC) bottom trawl

surveys (Link and Almeida 2000) were combined

across seasons to compute the percent composition of 

major prey categories (Figure 4). For both pre-recruits

(92%) and recruits (57%), a majority of the gut contents 

consisted of well-digested prey. Pre-recruit prey types 

that could be identified consisted of crustaceans (3%) 

and fish (3%). The diet of recruits consisted of

cephalopods (30%), crustaceans (including euphausiids, 

7%), and fish (6%).

PREDATION AND MORTALITY

Numerous species of pelagic and benthic fishes

prey on Illex, including bluefin tuna (Butler 1971),

silver hake and red hake (Vinogradov 1972). Other fish 

predators include bluefish (Maurer 1975; Buckel 1997), 

goosefish (Maurer 1975; Langton and Bowman 1977), 

fourspot flounder (Langton and Bowman 1977),

Atlantic cod (Lilly and Osborne 1984), sea raven

(Maurer 1975), spiny dogfish (Templeman 1944;

Maurer 1975), and swordfish (Langton and Bowman 

1977; Stillwell and Kohler 1985; Scott and Scott 1988).

Mammalian predators include pilot whales (Squires

1957; Wigley 1982) and the common dolphin (Major 

1986). Seabird predators include shearwaters, gannets, 

and fulmars (Brown et al. 1981). Northern shortfin

squid are known to exhibit a variety of defense

mechanisms that may reduce predation, such as

camouflage coloration (O’Dor 1983), schooling

behavior, jetting, and ink release (Major 1986).

MIGRATION

Northern shortfin squid are highly migratory. An 

individual tagged off Newfoundland was recaptured off 

the coast of Maryland, more than 1,000 miles away 

(Dawe et al. 1981b). A hypothetical, annual migration 

route (Figure 2, from Black et al. 1987) has been

constructed based on seasonal squid distribution

patterns observed in bottom trawl surveys of the U.S. 

and Canadian continental shelves, concentrations of

hatchlings in offshore waters south of Cape Hatteras 

during winter (Dawe and Beck 1985; Hatanaka et al.

1985; Rowell et al. 1985a), and suggestions that the 

neutrally-buoyant egg masses and paralarvae are

rapidly transported northeastward by the Gulf Stream 

current (Trites 1983).

Seasonal distribution patterns in Illex abundance

suggest that annual migrations off the U.S. shelf in

autumn and onto the shelf in spring occur

simultaneously along the entire length of the shelf edge 

rather than over the shelf in a gauntlet pattern

(Hendrickson 2004; also see Geographical Distribution

below). Tagging studies have demonstrated a

southeastward migration of individuals from

Newfoundland during autumn (Dawe et al. 1981b).

However, the migration patterns between northern and 

southern stock components remain unknown and the 

offshore fraction of the population is not well

understood because NEFSC surveys do not extend

beyond the edge of the continental shelf and few

stations are sampled in the deepest survey strata (185 to 

366 m).

GEOGRAPHICAL DISTRIBUTION

Illex illecebrosus utilizes oceanic and neritic

habitats and adults are believed to undergo longdistance migrations between boreal, temperate and subtropical waters. Data from U.S. and Canadian seasonal 

bottom trawl surveys (1975 to 1994) indicate that

northern shortfin squid are distributed on the

continental shelf of the U.S. and Canada, between

Newfoundland and Cape Hatteras, North Carolina

(Figure 5). The species is present in the Gulf of St.
Page 4

Lawrence, along the western edge of the Grand Bank, 

and along the western shore of Newfoundland, but are 

most abundant on the U.S. and Scotian Shelf.

Paralarvae and juveniles inhabit the Gulf Stream-slope

water interface, located off the continental shelf of the 

U.S. and Canada, and juveniles also occur on the U.S. 

continental shelf. Adults have primarily been collected 

on the shelf due to sampling depth limitations of U.S. 

and Canadian bottom trawl surveys.

The southernmost limit of the range of I.

illecebrosus is difficult to identify because of its cooccurrence with I. coindetii and I. oxygonius.

Distinguishing between the three species is difficult

given the high degree of interspecific and intraspecific 

variability in morphological characters (Roper and

Mangold 1998; Roper et al. 1998).

EGGS AND PARALARVAE

Egg masses have never been collected in nature

(O’Dor and Dawe 1998). Paralarvae have been

collected during all seasons (Roper and Lu 1979), but 

predominately during January and February in the

nutrient-rich waters of the Slope Water-Gulf Stream

frontal zone, from south of Cape Hatteras to as far north 

as the Grand Bank (Dawe and Beck 1985; Hatanaka et

al. 1985). The Gulf Stream has been hypothesized as 

the primary mechanism for the transport of egg

balloons and paralarvae toward the Grand Bank (Trites 

1983) based on its northeastern trajectory and rapid

current (about 100 km/day).

PRE-RECRUITS

NEFSC bottom trawl surveys [see Reid et al.

(1999) for details] have captured pre-recruits during all 

seasons (Figure 6; note that winter and summer

distributions are presented as presence/absence data,

precluding a discussion of abundances). In winter, the 

occurrence of pre-recruits is very low and distributed 

along the shelf edge between Cape Hatteras and

Georges Bank. In the spring, pre-recruits are

concentrated in low densities along the shelf edge,

including the southern Scotian Shelf; the highest

densities are found off of Cape Hatteras. By summer, 

pre-recruits have migrated onto the continental shelf

and are distributed throughout all depths, primarily in

the Mid-Atlantic Bight and along the coast of Maine. In

autumn, pre-recruits begin their return migration to

waters off the shelf. During fall, pre-recruits are present 

at depths greater than 60 m, between Georges Bank and 

Cape Hatteras, and are most abundant along the shelf 

edge. Low densities are also present in the Gulf of

Maine.

The autumn distribution and abundance of prerecruits around coastal Massachusetts, based on

Massachusetts inshore bottom trawl surveys [see Reid 

et al. (1999) for details], is shown in Figure 7. During 

all of the spring surveys conducted between 1978 and 

2003, only 16 individuals were collected at five stations 

in Massachusetts coastal waters. During the fall

surveys, pre-recruits were present at very low densities 

at 28 stations located primarily off Cape Ann and

northern Cape Cod Bay (at depths of 20 to 60 m); a few 

higher concentrations were found east of Cape Cod and 

near Nantucket Island.

Few (N=30) northern shortfin squid were caught 

during seasonal surveys of Narragansett Bay between 

1990 and 1996. Illex were captured only during the 

summer and at three stations. Individuals ranged in size 

from 4 to 11 cm ML.

ADULTS

NEFSC bottom trawl surveys indicate similar

seasonal distributions of recruits and pre-recruits. (See 

Figure 8 for recruits; again note that winter and summer 

distributions are presented as presence/absence data,

precluding a discussion of abundances). Recruit

abundance during spring and autumn appears to be

greater than that of pre-recruits, but this is partially due

to differences in catchability between the two size

groups. The occurrence of recruits on the U.S.

continental shelf is lowest during winter and

concentrated along the shelf edge (at depths around 366 

m) between Georges Bank and Cape Hatteras. By

spring, recruits are still only present at low densities 

and concentrated near the shelf edge, extending to south 

of Cape Hatteras, where some of the highest densities 

are found. Recruits occur both inshore and throughout 

the continental shelf during the summer between the 

Gulf of Maine and Cape Hatteras. During autumn, 

recruits begin to migrate back offshore and south, as 

indicated by their high concentrations at depths

between 60 and 366 m.

Recruit abundance was low in Massachusetts

coastal waters during 1978-2003, particularly during

the spring, when only a few recruits were collected at 

four stations. During the fall, recruits were collected at 

more stations and at higher densities. Abundance was 

highest in the waters off Cape Ann and the northern

portion of Cape Cod Bay (Figure 9).

HABITAT CHARACTERISTICS

Habitat characteristics of northern shortfin squid 

are summarized by life history stage in Table 1.
Page 5

EGGS AND PARALARVAE

Based on lab studies, egg balloons probably occur

at water temperatures between 12.5°C, the minimum

temperature required for successful embryonic

development (O’Dor et al. 1982b), and 26°C (Balch et

al. 1985). Egg masses are neutrally-buoyant and

probably occur in midwater near the pycnocline (O’Dor 

and Balch 1985).

Illex sp. paralarvae have been collected at water 

temperatures from 5 to 20°C (Vecchione 1979; O’Dor 

1983; Dawe and Beck 1985; Hatanaka et al. 1985; 

Vecchione and Roper 1986), with maximum

abundance, in the Gulf Stream, at temperatures greater 

than 16.5°C (Hatanaka et al. 1985) and salinities

ranging from 35 to 37 ppt (Vecchione 1979; O’Dor 

1983; Dawe and Beck 1985; Vecchione and Roper 

1986). Paralarvae exhibit diel vertical migrations and 

are more abundant in the upper layer of the water at 

night and in deeper water during the day (Hatanaka et

al. 1985).

JUVENILES

During the spring, epipelagic juveniles migrate

from oceanic to neritic waters as they grow. Juveniles

have been collected from continental slope waters at 

temperatures from 14.3 to 16.3°C (Fedulov and

Froerman 1980; Perez 1994), at temperatures above 

16°C in the Gulf Stream (Perez 1994), and at

temperatures from 5 to 6°C on the Scotian Shelf in

spring (Perez 1994). During late May, juveniles (34 to 

68 mm ML) were collected along the southeast flank of 

Georges Bank, at depths of 140 to 260 m, where surface 

and bottom temperatures were 10.6°C and 9.9°C,

respectively (Hendrickson 2004). Juveniles have been 

collected at salinities of 34 to 37 ppt (Vecchione 1979; 

Amaratunga et al. 1980b; Fedulov and Froerman 1980; 

Rowell et al. 1985a). South of Cape Hatteras, squid 7 

to 10 cm ML are most abundant during spring

(Whitaker 1980).

Distributions of pre-recruits relative to bottom

water temperature, depth, and salinity based on spring 

and fall NEFSC bottom trawl surveys from the Gulf of 

Maine to Cape Hatteras are shown in Figure 10. During 

the spring surveys, pre-recruits occur in deep water and 

are most abundant at depths ranging from 101 to 300 m, 

bottom temperatures of 11 to 14oC and at salinities of 

35 to 36 ppt. During the fall surveys, pre-recruits occur

in greater abundance across a broader depth range, and

in a wider range of temperatures and salinities. During 

autumn, juveniles are most abundant at bottom

temperatures of 10 to 13oC and salinities of 32 to 35 

ppt.

The spring and autumn distributions of pre-recruits

in Massachusetts coastal waters relative to bottom

water temperature and depth based on Massachusetts 

inshore bottom trawl surveys are shown in Figure 11.

As observed in the NEFSC spring surveys, pre-recruits

are distributed offshore during spring; only 16

individuals were collected at five stations in

Massachusetts coastal waters (< 80 m deep), most at 

temperatures of 11oC and at a depth of 11 to 15 m.

During the fall, pre-recruits were present at very low 

densities at only 28 stations and were most abundant at 

depths of 31 to 55 m, and were found over at bottom 

temperatures of 6 to 10oC.

ADULTS

Adults have been captured at temperatures ranging

from -0.5 to 27.3°C (Whitaker 1980), salinities of 30 to 

36.5 ppt (Palmer and O’Dor 1978), and at depths

ranging from the surface to 1000 m or more (Whitaker 

1980), depending on the time of year (see Migrations 

above). In summer, on the eastern U.S. continental

shelf, adults are most abundant at depths of 100 to 200 

m (Bowman 1977; Grinkov and Rikhter 1981) and are 

not generally found in waters shallower than 18 m. In 

the fall and winter, adults migrate offshore, and have 

been found at 100 to 945 m (Amaratunga et al. 1980a; 

Felley and Vecchione 1995). However, there is little 

information on the offshore component of the

population, which may be found at depths greater than 

1000 m (O’Dor and Dawe 1998).

Distributions of recruits relative to bottom water 

temperature, depth, and salinity based on NEFSC

spring and fall bottom trawl surveys are shown in

Figure 12. During the spring, recruits are found over a 

temperature range of 4 to 20°C, but are most abundant 

at 10 to 14°C. Recruits are found over a depth range of 

11 to 500 m, but are most abundant at 121 to 400 m. 

The majority of recruits occur at a salinity of 35 ppt in 

the spring. In the autumn, recruits were found over a 

bottom temperature range of 4 to about 21°C, and are

most abundant at 8 to 13°C. Recruit abundance

increased with depth between 31 and 140 m and

reached a secondary peak at 201 to 300 m. During

autumn, recruits occur over a salinity range from 31 to 

36 ppt.

The spring and autumn distributions of recruits in

Massachusetts coastal waters relative to bottom water 

temperature and depth based on Massachusetts inshore 

bottom trawl surveys are shown in Figure 13. The few 

adults caught in the spring occurred at temperatures of 

10 to 13°C and at depths of 11 to 15 m, 26 to 30 m, and 

41 to 45 m. Recruits were caught at higher densities in 

the fall and were found over a temperature range of 4 to 

15°C, with most recruits found between 7 and 9°C.
Page 6

Recruits were caught over a depth range of 11 to 85 m, 

with most found between 41 and 75 m.

RESEARCH NEEDS

Additional research is needed to better understand 

the life cycle of this species. In particular, recruitment 

patterns and the exchange of squid between the

northern and southern stock components remain

unknown, along with the migration routes from fishing 

areas during autumn. U.S. research surveys do not

include waters deeper than 366 m, so it remains

unknown what fraction of the stock resides offshore and 

whether spawning occurs there. In addition, the

location of the winter spawning area has not been

confirmed.

ACKNOWLEDGMENTS

The authors thanks Jason Link for summarizing

data from the NEFSC food habits database and Jackie 

Riley and Claire Steimle for library assistance. We are 

also grateful to John McCarthy for preparing the

histogram graphs and to David Packer for his editorial 

comments and organization of the document.

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Table 1. Summary of life history and habitat parameters for northern shortfin squid Illex illecebrosus.

Life Stage Size and Growth Habitat Substrate Temperature Salinity

Eggs 1

Eggs protected in 

gelatinous masses 

ranging in diameter 

from 30 - 100 cm. In 

lab studies, females 

produced 10,000 -

400,000 eggs.

Egg masses have not been collected 

in the wild. It is hypothesized that 

egg masses are transported 

northeasterly in the Gulf Stream 

current based on distribution of 

paralarvae and temperatures required 

for hatching. 

Egg masses are 

pelagic; do not attach

to substrate (based 

on lab-induced

spawning events).

Egg incubation lasts 16 

days at 13°C, 12 days at 

16°C and 8 days at 21°C;

normal development 

requires at least 13°C.

Egg masses have 

greater density than 

seawater; possibly 

become neutrally -

buoyant in colder, 

higher-density

water (shown in lab 

studies).

Paralarvae 2

Hatchling:

1.0-1.1 mm

Paralarvae:

1.2-5.0 mm 

Transitional:

5.1-6.9 mm

Size at hatch: 1.0-1.1

mm ML. 

Paralarvae have nonfunctional tentacles; 

body not yet elongated. 

Rhynchoteuthion (Type 

C1

) larval stage ends 

when proboscis splits 

into 2 tentacles. Mantle 

length increases during 

migration from Gulf 

Stream to continental 

shelf. Only find 

hatchlings south of 

Cape Hatteras.

Offshore, along continental shelf edge 

from surface waters t o 360 m. 

Hatching occurs at inshore boundary 

of Gulf Stream, 9 to 16 days after 

spawning. Paralarvae have been 

found during winter and spring off 

tail of Grand Bank. Abundant in late 

February -March in Gulf Stream/slope 

waters above the thermocline. The 

convergence of Gulf Stream and slope 

water creates an area of high 

productivity that is beneficial to 

young for feeding and growth. 

Undergo diel vertical migrations; 

greater abundance during the day at 

50-100 m.

Found from 

5-20°C; maximum 

abundance at 

temperatures > 16.5°C.

Found at salinities 

ranging from 35-37

ppt.

Prerecruits 3

≤ 10.0 cm

Separation of proboscis 

into tentacles indicates 

onset of juvenile stage. 

Larger juveniles found 

from east to west, 

indicating westward 

movement on 

continental shelf with 

growth. Growth is 

approximately 1.5 

mm/day.

Winter: in Gulf Stream/slope water 

interface.

Spring: begin migration onto U.S. 

continental shelf (Georges Bank to 

south of Cape Hatteras).

Summer: occur throughout U.S. shelf 

(Georges Bank to Cape Hatteras).

Fall: migrate off U.S. shelf (Georges 

Bank to south of Cape Hatteras).

Undergo diel vertical migrations; 

greater abundance near the bottom 

during the day.

Gulf Stream: > 16°C;

slope water: < 16°C;

surface: 14-21°C;

continental shelf: 5-6°C

in spring.

Pre-recruits on U.S. shelf 

most abundant at bottom 

temperatures > 10°C and 

surface temperatures 

14.6-20.5°C.

Found at salinities 

ranging from 34-37

ppt.

Recruits 4

≥ 11 cm

Can reach size of 35 cm 

ML.

Life span less than a 

year; (215 days for 

squid in U.S. shelf 

waters). Males and 

females grow about 1 

mm/day. Females

generally larger than 

males.

Range from Labrador to south of 

Cape Hatteras; most abundant at 

depths of 100-150m.

Winter: low abundance along edge of 

U.S. shelf; presumably in warmer 

waters offshore and south of Cape 

Hatteras.

Spring: begin migration onto U.S. 

shelf (Georges Bank to south of Cape 

Hatteras) and Scotian Shelf.

Summer: occur throughout U.S. shelf 

(Gulf of Maine to Cape Hatteras), 

Scotian Shelf and inshore 

Newfoundland waters.

Fall: migrate off continental shelf of 

U.S. and Canada; presumably to 

spawn.

Increase in squid size with depth in 

autumn.

Over various 

sediment types, 

including sand-silt or 

"Sambro sand" 

(sediment between 

banks and edges of 

basins on Scotian 

Shelf, as well as 

along edge of 

continental shelf, 

100-300m). Avoid 

areas inhabited by 

anemones.

Illex ≥ 11 cm found at 

bottom temperatures 

ranging from 3.5-15.0°C

(surface > 20°C), most 

abundant at bottom 

temperatures of 5-

10.0°C. Maturation may 

be enhanced by high

temperatures but not 

initiated by it.

Generally found at 

30-36.5 ppt. 

1 Durward et a l. (1978); O’Dor and Durward (1979); Durward et al. (1980); O’Dor et al. (1980b, 1982b, 1986); O’Dor (1983); O’Dor and Balch (1985); Rowell et al.

(1985a); Perez (1994). 2 Vecchione (1979); Amaratunga (1980a); Durward et al. (1980); O’Dor (1983); Trites (1983); Dawe and Beck (1985); Hatanaka et al. (1985); Rowell and Trites (1985); 

Rowell et al. (1985a); Vecchione and Roper (1986); Young and Harman (1988); Mann and Lazier (1991); Perez (1994). 3 Squires (1957); Vecchione (1979); Amaratunga (1980a); Amaratunga et al. (1980b); Fedulov and Froerman (1980); Dawe et al. (1981a); Coelho (1985); Rowell et al.

(1985a); Black et al. (1987); Nigmatullin (1987); Perez (1994); Dawe and Beck (1997); Brodziak and Hendrickson (1999). 4 Frost and Thompson (1933); McLellan et al. (1953); Squires (1957, 1967); Templeman (1966); Mercer (1973a, b); Mercer and Paulmier (1974); Bowman (1977);Mesnil

(1977); O’Dor et al. (1977, 1980a); Amaratunga et al. (1978, 1980a); Lange (1978); Lux et al. (1978); Palmer and O’Dor (1978); Amaratunga and McQuinn 1979; 

Fedulov and Froerman (1980); Hurley (1980); Whitaker (1980); Dawe and Drew (1981); Dawe et al. (1981b); Grinkov and Rikhter (1981); Lange and Johnson (1981); 

Scott (1982); Wigley (1982); Amaratunga 1983; Waldron (1983); Roper et al. (1984); Coelho (1985); Dawe and Beck (1985, 1997); Rowell et al. (1985b); Vecchione et

al. (1989); Laptikhovsky and Nigmatullin (1993); Perez (1994); Felley and Vecchione (1995); Brodziak and Hendrickson (1999); Hendrickson (2004).
Page 13

Table 1. Cont’d.

Life Stage Prey Predators Spawning Notes

Eggs 1

Spawning has bee induced in the 

lab, but not observed in the wild.

Lab studies indicate that egg 

masses are spawned pelagically.

Eggs that are presumably spawned in 

Gulf Stream waters can hatch in 

northern shelf waters > 12.5oC 

(transported by Gulf Stream at rate of 

7 km/hr); can also hatch in warm Gulf 

Stream waters.

Paralarvae 2

Hatchling:

1.0-1.1 mm

Paralarvae:

1.2-5.0 mm 

Transitional:

5.1-6.9 mm

Hatchlings may spend early 

life in remains of egg mass to 

utilize the nutrients for food. 

Yolk-sac not especially large; 

food must be adequate to 

sustain hatchling during this 

stage of rapid growth and 

increased metabolism.

Gulf Stream may be important mode 

of transportation for paralarvae 

throughout range in NW Atlantic; 

initially flows northeastward along 

shelf, off Cape Hatteras, then flows 

easterly and creates eddies in which

young are transported westward into 

slope waters.

Prerecruits 3

≤ 10.0 cm

Primarily feed on crustaceans 

(euphausiids) at night near 

the surface; also consume 

nematodes and fish.

Gulf Stream presumably transports 

juveniles northward; hydrographic 

variability in this system may explain 

annual abundance differences.

Recruits 4

≥ 11 cm

Visual predators; feeding rate 

reduced in highly turbid 

waters. Feed primarily on 

fish and are cannibalistic 

(larger females cannibalize 

smaller males, increased in 

autumn). Fish prey includes 

juvenile Atlantic cod, 

mackerel, redfish, sand lance, 

Atlantic herring, and adult 

capelin. Seasonal/ontogenetic 

diet shifts, during spring 

(offshore): euphausiids; 

during summer-fall (inshore): 

fish and squid. 

Many pelagic and 

benthic fishes feed 

heavily on Illex,

including bluefin tuna 

and silver and red hakes. 

Other fish predators 

include shark and 

dogfish species, fourspot 

flounder, Atlantic cod, 

swordfish, bluefish, 

goosefish, and sea raven. 

Mammalian predators 

include common dolphin 

and pilot whales. Avian 

predators include 

shearwaters, gannets, 

and fulmars.

Spawning likely pelagic and 

occurs during October-July.

Late May: mated females 

indicate spawning area (113-377

m) in Mid-Atlantic Bight; 

overlaps with fishing grounds.

Winter: presumably spawn 

during December-March in the 

Gulf Stream and/or south of 

Cape Hatteras where Illex sp.

hatchlings were collected.

Lab studies indicate females 

mate, spawn once (may release 

multiple egg masses), then die 

within a week. Mated females

collected in Mid-Atlantic Bight 

indicate females may mate with 

as many as four males. 

Diel vertical migrations: more 

abundant on bottom at dawn/dusk and 

day than at night; feed primarily at 

night before sunrise near surface or 

mid-water. Migrate to bottom or 

deeper waters during daytime. Change 

color to camouflaged pattern when 

resting on bottom to reduce risk of 

predation by benthic species.

1 O’Dor et al. (1980b, 1982a, 1986); O’Dor (1983); Rowell et al. (1985a); Perez (1994). 2 Durward et al. (1980); Trites (1983); Rowell and Trites (1985); O’Dor et al. (1986); Vecchione and Roper (1986); Csanady and Hamilton (1988); Mann and Lazier (1991); 

Perez (1994). 3 Amaratunga et al. (1980b); Dawe et al. (1981a); Coelho (1985); Arkhipkin and Fedulov (1986). 4 Templeman (1944); Squires (1957, 1966, 1967); Vinogradov (1970, 1972, 1984); Butler (1971); Mercer and Paulmier (1974); Maurer (1975); Langton and Bowman 

(1977); Bennett (1978); Durward et al. (1978); Hirtle (1978); Ennis and Collins (1979);Froerman (1979); Vinogradov and Noskov (1979); Amaratunga (1980b, 1983); 

Amaratunga et al. (1980a); Fedulov and Froerman (1980); Hurley (1980); Lange and Sissenwine (1980); O’Dor et al. (1980a, b); Brown et al. (1981); DeMont (1981); 

Hirtle et al. (1981); Wigley (1982); O’Dor (1983); Lily and Osborne (1984); Dawe and Beck (1985, 1997); Maurer and Bowman (1985); Nicol and O’Dor (1985); O’Dor 

and Balch (1985); Rowell et al. (1985a); Stillwell and Kohler (1985); Major (1986); Scott and Scott (1988); Vecchione et al. (1989); Laptikhovsky and Nigmatullin 

(1993); Perez (1994); Dawe et al. (1997); Brodziak and Hendrickson (1999); Hendrickson (2004).
Page 14

Figure 1. The northern shortfin squid, Illex illecebrosus (from Goode 1884).
Page 15

Figure 2. Hypothetical migration path of the northern shortfin squid, Illex illecebrosus.

From Black et al. (1987).
Page 16

Figure 3. Spawning area of northern shortfin squid (encircled) during late May.

Based on the distribution of mated females (Hendrickson 2004).
Page 17

Diet Composition of Major Prey Items

Pre-recruits

0

20

40

60

80

100

Well Digested

Prey

Cephalopods Misc.

Crustaceans

Euphausiids Misc. Fish

Percent Diet Composition (g)

Recruits

0

15

30

45

60

Well Digested

Prey

Cephalopods Misc.

Crustaceans

Euphausiids Misc. Fish

Percent Diet Composition (g)

Figure 4. Percent by weight (g) of the major prey items in the diet of northern shortfin squid.

From specimens collected during NEFSC bottom trawl surveys from 1973-2001 (all seasons). For details on NEFSC diet 

analysis, see Link and Almeida (2000).
Page 18

Figure 5. Distribution and abundance of northern shortfin squid from Newfoundland to Cape Hatteras.

Based on research trawl surveys conducted by Canada (DFO) and the United States (NMFS) from 1975-1994

(http://www-orca.nos.noaa.gov/projects/ecnasap/ecnasap_table1.html).
Page 19

Figure 6. Seasonal distributions and abundances of pre-recruit northern shortfin squid collected during NEFSC bottom 

trawl surveys.

From NEFSC winter bottom trawl surveys (1981-2003, all years combined). Distributions are displayed as 

presence/absence only.
Page 20

Figure 6. Cont’d.

From NEFSC spring bottom trawl surveys (1968-2003, all years combined). Survey stations where pre-recruits were not 

found are not shown.
Page 21

Figure 6. Cont’d.

From NEFSC summer bottom trawl surveys (1969-1995, all years combined). Distributions are displayed as 

presence/absence only.
Page 22

Figure 6. Cont’d.

From NEFSC fall bottom trawl surveys (1967-2003, all years combined). Survey stations where pre-recruits were not 

found are not shown.
Page 23

Figure 7. Distribution and abundance of pre-recruit northern shortfin squid in Massachusetts coastal waters.

From fall Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Survey stations where prerecruits were not found are not shown.
Page 24

Figure 8. Seasonal distributions and abundances of recruit northern shortfin squid collected during NEFSC bottom 

trawl surveys.

From NEFSC winter bottom trawl surveys (1981-2003, all years combined). Distributions are displayed as 

presence/absence only.
Page 25

Figure 8. Cont’d.

From NEFSC spring bottom trawl surveys (1968-2003, all years combined). Survey stations where recruits were not 

found are not shown.
Page 26

Figure 8. Cont’d.

From NEFSC summer bottom trawl surveys (1969-1995, all years combined). Distributions are displayed as presence

absence only.
Page 27

Figure 8. Cont’d.

From NEFSC fall bottom trawl surveys (1967-2003, all years combined). Survey stations where recruits were not found 

are not shown.
Page 28

Figure 9. Distribution and abundance of recruit northern shortfin squid in Massachusetts coastal waters.

From fall Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Survey stations where recruits

were not found are not shown.
Page 29

NEFSC Bottom Trawl Survey

Spring/Pre-recruits (<= 10 cm)

0

5

10

15

20

25

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=10879

Occurrence N=340

Catch N=3659

0

5

10

15

20

25

30

1-10

11-2021-3031-4041-5051-6061-7071-8081-90

91-100

101-120121-140141-160161-180181-200201-300301-400401-500

>500

Bottom Depth (m)

Percent

Trawls N=12514

Occurrence N=382

Catch N=4816

0

10

20

30

40

50

60

24 25 26 27 28 29 30 31 32 33 34 35 36 37 38

Salinity (PPT)

Percent

Trawls N=2270

Occurrence N=102

Catch N=1645

Figure 10. Distributions of pre-recruit northern shortfin squid and trawls from NEFSC bottom trawl surveys relative to 

bottom water temperature, depth, and salinity.

Based on NEFSC spring bottom trawl surveys (temperature and depth: 1968-2003, all years combined; salinity: 1991-

2003, all years combined). Light bars show the distribution of all the trawls, dark bars show the distribution of all trawls 

in which northern shortfin squid occurred and medium bars show, within each interval, the percentage of the total 

number of northern shortfin squid caught. Note that the bottom depth interval changes with increasing depth.
Page 30

NEFSC Bottom Trawl Survey

Fall/Pre-recruits (<= 10 cm)

0

5

10

15

20

25

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=11467

Occurrence N=818

Catch N=8405

0

2

4

6

8

10

12

14

16

1-10

11-2021-3031-4041-5051-6061-7071-8081-90

91-100

101-120121-140141-160161-180181-200201-300301-400401-500

>500

Bottom Depth (m)

Percent

Trawls N=13433

Occurrence N=1009

Catch N=10382

0

5

10

15

20

25

30

35

24 25 26 27 28 29 30 31 32 33 34 35 36 37 38

Salinity (PPT)

Percent

Trawls N=2272

Occurrence N=198

Catch N=1224

Figure 10. Cont’d.

Based on NEFSC fall bottom trawl surveys (temperature and depth: 1967-2003, all years combined; salinity: 1991-2003,

all years combined). Light bars show the distribution of all the trawls, dark bars show the distribution of all trawls in 

which northern shortfin squid occurred and medium bars show, within each interval, the percentage of the total number 

of northern shortfin squid caught. Note that the bottom depth interval changes with increasing depth.
Page 31

Massachusetts Inshore Bottom Trawl Survey

Spring/Pre-recruits (<= 10 cm)

0

20

40

60

80

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=2407

Occurrence N=5

Catch N=16

0

20

40

60

80

100

1-5

6-10

11-1516-2021-2526-3031-3536-4041-4546-5051-5556-6061-6566-7071-7576-8081-85

Bottom Depth (m)

Percent

Trawls N=2482

Occurrence N=5

Catch N=16

Figure 11. Distributions of pre-recruit northern shortfin squid and trawls in Massachusetts coastal waters relative to 

bottom water temperature and depth.

Based on spring Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Light bars show the 

distribution of all the trawls, dark bars show the distribution of all trawls in which northern shortfin squid occurred and 

medium bars show, within each interval, the percentage of the total number of northern shortfin squid caught.
Page 32

Massachusetts Inshore Bottom Trawl Survey

Fall/Pre-recruits (<= 10 cm)

0

5

10

15

20

25

30

35

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=2244

Occurrence N=28

Catch N=64

0

5

10

15

20

25

30

1-5

6-10

11-1516-2021-2526-3031-3536-4041-4546-5051-5556-6061-6566-7071-7576-8081-85

Bottom Depth (m)

Percent

Trawls N=2338

Occurrence N=28

Catch N=64

Figure 11. Cont’d.

Based on fall Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Light bars show the 

distribution of all the trawls, dark bars show the distribution of all trawls in which northern shortfin squid occurred and 

medium bars show, within each interval, the percentage of the total number of northern shortfin squid caught.
Page 33

NEFSC Bottom Trawl Survey

Spring/Recruits (>= 11 cm)

0

5

10

15

20

25

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=10879

Occurrence N=770

Catch N=9321

0

5

10

15

20

25

30

1-10

11-2021-3031-4041-5051-6061-7071-8081-90

91-100

101-120121-140141-160161-180181-200201-300301-400401-500

>500

Bottom Depth (m)

Percent

Trawls N=12514

Occurrence N=874

Catch N=10504

0

20

40

60

80

24 25 26 27 28 29 30 31 32 33 34 35 36 37 38

Salinity (PPT)

Percent

Trawls N=2270

Occurrence N=120

Catch N=2119

Figure 12. Distributions of recruit northern shortfin squid and trawls from NEFSC bottom trawl surveys relative to 

bottom water temperature, depth, and salinity.

Based on NEFSC spring bottom trawl surveys (temperature and depth: 1968-2003, all years combined; salinity: 1991-

2003, all years combined). Light bars show the distribution of all the trawls, dark bars show the distribution of all trawls 

in which northern shortfin squid occurred and medium bars show, within each interval, the percentage of the total 

number of northern shortfin squid caught. Note that the bottom depth interval changes with increasing depth.
Page 34

NEFSC Bottom Trawl Survey

Fall/Recruits (>= 11 cm)

0

5

10

15

20

25

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=11467

Occurrence N=4617

Catch N=104051

0

5

10

15

20

1-10

11-2021-3031-4041-5051-6061-7071-8081-90

91-100

101-120121-140141-160161-180181-200201-300301-400401-500

>500

Bottom Depth (m)

Percent

Trawls N=13433

Occurrence N=5404

Catch N=134252

0

10

20

30

40

50

24 25 26 27 28 29 30 31 32 33 34 35 36 37 38

Salinity (PPT)

Percent

Trawls N=2272

Occurrence N=776

Catch N=14760

Figure 12. Cont’d.

Based on NEFSC fall bottom trawl surveys (temperature and depth: 1967-2003, all years combined; salinity: 1991-2003,

all years combined). Light bars show the distribution of all the trawls, dark bars show the distribution of all trawls in 

which northern shortfin squid occurred and medium bars show, within each interval, the percentage of the total number 

of northern shortfin squid caught. Note that the bottom depth interval changes with increasing depth.
Page 35

Massachusetts Inshore Bottom Trawl Survey

Spring/Recruits (>= 11 cm)

0

5

10

15

20

25

30

35

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=2407

Occurrence N=4

Catch N=6

0

10

20

30

40

50

60

1-5

6-10

11-1516-2021-2526-3031-3536-4041-4546-5051-5556-6061-6566-7071-7576-8081-85

Bottom Depth (m)

Percent

Trawls N=2482

Occurrence N=4

Catch N=6

Figure 13. Distributions of recruit northern shortfin squid and trawls in Massachusetts coastal waters relative to bottom 

water temperature and depth.

Based on spring Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Light bars show the 

distribution of all the trawls, dark bars show the distribution of all trawls in which northern shortfin squid occurred and 

medium bars show, within each interval, the percentage of the total number of northern shortfin squid caught.
Page 36

Massachusetts Inshore Bottom Trawl Survey

Fall/Recruits (>= 11 cm)

0

5

10

15

20

25

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=2244

Occurrence N=287

Catch N=2069

0

5

10

15

20

25

30

1-5

6-10

11-1516-2021-2526-3031-3536-4041-4546-5051-5556-6061-6566-7071-7576-8081-85

Bottom Depth (m)

Percent

Trawls N=2338

Occurrence N=294

Catch N=2085

Figure 13. Cont’d.

Based on fall Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Light bars show the 

distribution of all the trawls, dark bars show the distribution of all trawls in which northern shortfin squid occurred and 

medium bars show, within each interval, the percentage of the total number of northern shortfin squid caught.
Publishing in NOAA Technical Memorandum NMFS-NE

Manuscript Qualification

This series represents a secondary level of scientific

publishing in the National Marine Fisheries Service (NMFS).

For all issues, the series employs thorough internal scientific

review, but not necessarily external scientific review. For most

issues, the series employs rigorous technical and copy editing.

Manuscripts that may warrant a primary level of scientific

publishing should be initially submitted to one of NMFS's

primary series (i.e., Fishery Bulletin, NOAA Technical Report

NMFS, or Marine Fisheries Review).

Identical, or fundamentally identical, manuscripts should

not be concurrently submitted to this and any other publication

series. Manuscripts which have been rejected by any primary

series strictly because of geographic or temporal limitations

may be submitted to this series.

Manuscripts by Northeast Fisheries Science Center

(NEFSC) authors will be published in this series upon approval

by the NEFSC's Deputy Science & Research Director. Manuscripts by non-NEFSC authors may be published in this series

if: 1) the manuscript serves the NEFSC's mission; 2) the

manuscript meets the Deputy Science & Research Director's

approval; and 3) the author arranges for the printing and binding

funds to be transferred to the NEFSC's Research Communications Unit account from another federal account. For all

manuscripts submitted by non-NEFSC authors and published

in this series, the NEFSC will disavow all responsibility for the

manuscripts' contents; authors must accept such responsibility.

The ethics of scientific research and scientific publishing

are a serious matter. All manuscripts submitted to this series

are expected to adhere -- at a minimum -- to the ethical guidelines

contained in Chapter 1 ("Ethical Conduct in Authorship and

Publication") of the CBE Style Manual, fifth edition (Chicago,

IL: Council of Biology Editors). Copies of the manual are

available at virtually all scientific libraries.

edition of the United States Government Printing Office Style

Manual. That style manual is silent on many aspects of scientific

manuscripts. NEFSC publication and report series rely more on the

CBE Style Manual, fifth edition.

For in-text citations, use the name-date system. A special

effort should be made to ensure that the list of cited works contains

all necessary bibliographic information. For abbreviating serial

titles in such lists, use the guidance of the International Standards

Organization; such guidance is easily accessed through the various

Cambridge Scientific Abstracts’ serials source lists (see http://

www.public.iastate.edu/~CYBERSTACKS/JAS.htm). Personal communications must include date of contact and full name and mailing

address of source.

For spelling of scientific and common names of fishes, mollusks, and decapod crustaceans from the United States and Canada,

use Special Publications No. 20 (fishes), 26 (mollusks), and 17

(decapod crustaceans) of the American Fisheries Society (Bethesda,

MD). For spelling of scientific and common names of marine

mammals, use Special Publication No. 4 of the Society for Marine

Mammalogy (Lawrence, KS). For spelling in general, use the most

recent edition of Webster’s Third New International Dictionary of

the English Language Unabridged (Springfield, MA: G.&C.

Merriam).

Typing text, tables, and figure captions: Text, tables, and

figure captions should be converted to WordPerfect. In general,

keep text simple (e.g., don't switch fonts and type sizes, don't use

hard returns within paragraphs, don't indent except to begin

paragraphs). Also, don't use an automatic footnoting function; all

notes should be indicated in the text by simple numerical superscripts, and listed together in an "Endnotes" section prior to the

"References Cited" section. Especially, don't use a graphics

function for embedding tables and figures in text.

Tables should be prepared with a table formatting function.

Each figure should be supplied both on paper and on disk, unless

there is no digital file of a given figure. Except under extraordinary

circumstances, color will not be used in illustrations.

Manuscript Preparation

Organization: Manuscripts must have an abstract, table

of contents, and -- if applicable -- lists of tables, figures, and

acronyms. As much as possible, use traditional scientific

manuscript organization for sections: "Introduction," "Study

Area," "Methods & Materials," "Results," "Discussion" and/

or "Conclusions," "Acknowledgments," and "References Cited."

Style: All NEFSC publication and report series are

obligated to conform to the style contained in the most recent

Northeast Fisheries Science Center

Operations, Management & Information Services Staff

Research Communications Branch

Editorial Office

Manuscript Submission

Authors must submit one paper copy of the double-spaced

manuscript, one disk copy, and original figures (if applicable).

NEFSC authors must include a completely signed-off "NEFSC

Manuscript/Abstract/Webpage Review Form." Non-NEFSC authors who are not federal employees will be required to sign a

"Release of Copyright" form.

Send all materials and address all correspondence to: Jon A.

Gibson (Biological Sciences Editor), NMFS Northeast Fisheries

Science Center, 166 Water Street, Woods Hole, MA 02543-1026.
National Marine Fisheries Service, NOAA

166 Water St.

Woods Hole, MA 02543-1026

Publications and Reports

of the

Northeast Fisheries Science Center

The mission of NOAA's National Marine Fisheries Service (NMFS) is "stewardship of living marine resources

for the benefit of the nation through their science-based conservation and management and promotion of the health

of their environment." As the research arm of the NMFS's Northeast Region, the Northeast Fisheries Science

Center (NEFSC) supports the NMFS mission by "conducting ecosystem-based research and assessments of living

marine resources, with a focus on the Northeast Shelf, to promote the recovery and long-term sustainability of these

resources and to generate social and economic opportunities and benefits from their use." Results of NEFSC

research are largely reported in primary scientific media (e.g., anonymously-peer-reviewed scientific journals).

However, to assist itself in providing data, information, and advice to its constituents, the NEFSC occasionally

releases its results in its own media. Currently, there are three such media:

NOAA Technical Memorandum NMFS-NE -- This series is issued irregularly. The series typically includes: data reports of longterm field or lab studies of important species or habitats; synthesis reports for important species or habitats; annual reports of overall

assessment or monitoring programs; manuals describing program-wide surveying or experimental techniques; literature surveys of

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and/or summary reports of scientific meetings; and simple bibliographies. Issues receive internal scientific review, but no technical

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Resource Survey Report (formerly Fishermen's Report) -- This information report is a quick-turnaround report on the distribution

and relative abundance of selected living marine resources as derived from each of the NEFSC's periodic research vessel surveys of the

Northeast's continental shelf. There is no scientific review, nor any technical or copy editing, of this report.

OBTAINING A COPY: To obtain a copy of a NOAA Technical Memorandum NMFS-NE or a Northeast Fisheries Science Center

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Hole, MA 02543-1026; 508-495-2228) or consult the NEFSC webpage on "Reports and Publications" (http://www.nefsc.noaa.gov/

nefsc/publications/).

ANY USE OF TRADE OR BRAND NAMES IN ANY NEFSC PUBLICATION OR REPORT DOES NOT IMPLY ENDORSEMENT.

MEDIA

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