NOAA Technical Memorandum NMFS-NE-193

U. S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

Northeast Fisheries Science Center

Woods Hole, Massachusetts

August 2005

Essential Fish Habitat Source Document:

Longfin Inshore Squid, Loligo pealeii,

Life History and Habitat Characteristics

Second Edition
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.

183. In press.

184. A Large Marine Ecosystem Approach to Fisheries Management and Sustainability. By Frank J. Gable. August 2004. v +

84 p., 38 figs., 10 tables. NTIS Access. No. PB2005-101435.

185. Revised and Updated Edition of F. Bruce Sanford's 1957 "Planning Your Scientific Research Paper." By Jon A. Gibson.

August 2004. x + 36 p., 5 figs., 12 tables.

186. Essential Fish Habitat Source Document: Silver Hake, Merluccius bilinearis, Life History and Habitat Characteristics.

2nd ed. By Meredith C. Lock and David B. Packer. August 2004. v + 68 p., 28 figs., 6 tables. NTIS Access. No. PB2005-101436.

187. In preparation by authors.

188. In preparation by authors.

189. Essential Fish Habitat Source Document: Sea Scallop, Placopecten magellanicus, Life History and Habitat Characteristics.

2nd ed. By Deborah R. Hart and Antonie S. Chute. September 2004. v + 21 p., 6 figs., 2 tables. NTIS Access. No. PB2005-104079.

190. In preparation by authors.

191. Essential Fish Habitat Source Document: Northern Shortfin Squid, Illex illecebrosus, Life History and Habitat

Characteristics. 2nd ed. By Lisa C. Hendrickson and Elizabeth M. Holmes. November 2004. v + 36 p., 13 figs., 1 table. NTIS Access.

No. PB2005-101437.

192. Essential Fish Habitat Source Document: Atlantic Herring, Clupea harengus, Life History and Habitat Characteristics.

2nd ed. By David K. Stevenson and Marcy L. Scott. July 2005. vi + 84 p., 40 figs., 7 tables. NTIS Access. No. PB2005-107567.

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

Carlos M. Gutierrez, 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

August 2005

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-193

Larry D. Jacobson

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

Essential Fish Habitat Source Document:

Longfin Inshore Squid, Loligo pealeii,

Life History and Habitat Characteristics

Second Edition

E-Mail Address: Larry.Jacobson@noaa.gov
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., Nelson et al. 2004a

; Robins

et al. 1991b), 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.

a

Nelson, J.S.; Crossman, E.J.; Espinosa-Pérez, H.; Findley, L.T.; Gilbert, C.R.; Lea, R.N.; Williams, J.D. 2004. Common and scientific names

of fishes from the United States, Canada, and Mexico. 6th ed. Amer. Fish. Soc. Spec. Publ. 29; 386 p. b

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

North Americans. Amer. Fish. Soc. Spec. Publ. 21; 243 p. c

Turgeon, 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. d

Williams, 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.
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 of 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 are 

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 longfin 

inshore squid EFH source document is based on the 

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

McBride, Christine A. Zetlin, and Wallace W. Morse, 

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 ..................................................................................................................................2

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

RESEARCH NEEDS ...........................................................................................................................................................5

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

Tables

Table 1. Summary of life history and habitat characteristics for longfin inshore squid, based on the pertinent 

literature. ............................................................................................................................................................9

Figures

Figure 1. The longfin inshore squid, Loligo pealeii (from Goode 1884). .......................................................................11

Figure 2. Seasonal distributions and abundances of pre-recruit longfin inshore squid collected during NEFSC bottom 

trawl surveys. ...................................................................................................................................................12

Figure 3. Distribution and abundance of pre-recruit longfin inshore squid in Massachusetts coastal waters. ................16

Figure 4. Seasonal distribution and abundance of longfin inshore squid pre-recruits in Narragansett Bay. ...................18

Figure 5. Monthly log10 length frequencies (cm) of longfin inshore squid collected in Long Island Sound...................19

Figure 6. Relative abundance catch/tow and percent occurrence for longfin inshore squid in Long Island Sound ........20

Figure 7. Distribution and abundances of longfin inshore squid in Long Island Sound..................................................21

Figure 8. Seasonal distribution and abundance of longfin inshore squid pre-recruits collected in the Hudson-Raritan 

estuary..............................................................................................................................................................22

Figure 9. Seasonal distributions and abundances of recruit longfin inshore squid collected during NEFSC bottom 

trawl surveys. ...................................................................................................................................................23

Figure 10. Seasonal distributions and abundances of recruit longfin inshore squid in Massachusetts coastal waters. .....27

Figure 11. Seasonal distribution and abundance of longfin inshore squid recruits in Narragansett Bay...........................29

Figure 12. Seasonal distribution and abundance of longfin inshore squid recruits collected in the Hudson-Raritan 

estuary..............................................................................................................................................................30

Figure 13. Distributions of pre-recruit longfin inshore squid and trawls from NEFSC bottom trawl surveys relative to 

bottom water temperature, depth, and salinity. ................................................................................................31

Figure 14. Distributions of pre-recruit longfin inshore squid and trawls in Massachusetts coastal waters relative to 

bottom water temperature and depth................................................................................................................33

Figure 15. Distributions of longfin inshore squid pre-recruits in Narragansett Bay relative to mean bottom water 

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

Figure 16. Distributions of longfin inshore squid pre-recruits in the Hudson-Raritan estuary relative to bottom water 

temperature, depth, dissolved oxygen, and salinity..........................................................................................36

Figure 17. Distributions of recruit longfin inshore squid and trawls from NEFSC bottom trawl surveys relative to 

bottom water temperature, depth, and salinity. ................................................................................................37

Figure 18. Distributions of recruit longfin inshore squid and trawls in Massachusetts coastal waters relative to bottom 

water temperature and depth. ...........................................................................................................................39

Figure 19. Distributions of longfin inshore squid recruits in Narragansett Bay relative to mean bottom water 

temperature and bottom depth..........................................................................................................................41

Figure 20. Distributions of longfin inshore squid recruits in the Hudson-Raritan estuary relative to bottom water 

temperature, depth, dissolved oxygen, and salinity..........................................................................................42

Page 1

INTRODUCTION 

The longfin inshore squid, Loligo pealeii, is a 

schooling species of the molluscan family Loliginidae 

(Figure 1). It is distributed in continental shelf and 

slope waters from Newfoundland to the Gulf of 

Venezuela, and occurs in commercial abundance from 

southern Georges Bank to Cape Hatteras. The fishery 

for longfin inshore squid is managed by the MidAtlantic Fishery Management Council under the 

Atlantic Mackerel, Squid and Butterfish Fishery 

Management Plan, Amendment 8 (MAFMC 1998). 

Within the range of commercial exploitation, the 

population is considered to be a single stock unit. This 

Essential Fish Habitat Source Document provides 

information on the life history and habitat 

characteristics of longfin inshore squid inhabiting the 

Gulf of Maine, Georges Bank, and the Middle Atlantic 

Bight. 

LIFE HISTORY 

See Brodziak (1995) for a brief synopsis of life 

history. More detailed information is provided here. 

EGGS AND LARVAE 

The 1 mm x 1.6 mm eggs are encased in a 

gelatinous capsule as they pass through the female 

oviduct during mating. Each capsule contains 150-200 

eggs (Arnold et al. 1974; Gosner 1978; MAFMC 1998) 

and is about 50-80 mm long and 1 cm in diameter 

(Gosner 1978; Lange 1982; MAFMC 1998). During 

spawning, the male cements bundles of spermatophores 

into the mantle cavity of the female. The jelly is 

penetrated by sperm as the egg capsules pass through 

the oviduct (Black et al. 1987). The egg capsules are 

laid on the bottom in clusters 50-60 cm wide composed 

of hundreds of capsules (Gosner 1978; Griswold and 

Prezioso 1981). Each female lays 20-30 capsules 

(Lange 1982). The number of eggs spawned per female 

has been reported as 950-8,500 (Haefner 1959), 3,500-

6,000 (Summers 1971), 2,500-15,900 (Vovk 1972b), 

and 3,000-6,000 (MAFMC 1998). Development time 

varies from 257 to 642 hrs depending on water 

temperature; 26.7 days to hatching at 12-18o

C, 18.5 

days at 15.5-21.3o

C, and 10.7 days at 15.5-23.0oC 

(Summers 1971). 

Larvae of the longfin inshore squid are referred to 

as paralarvae (Young and Harman 1988). Little is 

known about them because they are planktonic, being 

found in the water column near the surface (McMahon 

and Summers 1971), and require special sampling 

techniques. Larvae 2-4 mm in length have been caught 

in the Gulf of Maine (Bigelow 1924). 

JUVENILES AND SUBADULTS 

There are two juvenile stages. ‘Juvenile’ is the 

stage after the paralarval stage and before the ‘subadult’ 

stage. The subadult stage is before maturity, when 

morphological characteristics of adults are attained 

(Young and Harman 1988). The shift from inhabiting 

surface waters to a demersal lifestyle occurs at 45 mm 

(Vecchione 1981). Off Martha’s Vineyard, the juvenile 

life stage lasts about 1 month. Subadults migrate by 

November to the outer shelf areas where they remain 

until March (Summers 1968a, b). Subadults are 

thought to overwinter in deeper waters along the edge 

of the continental shelf (Black et al. 1987). Young-ofthe-year (subadults) are found with adults in midsummer bottom trawl catches (Summers 1968a, b). 

Juveniles and subadults grow quickly, with growth rates 

dependent on temperature (Hatfield et al. 2001). 

Sexual maturity is first reached at about 8-12 cm 

(Macy 1980; Brodziak and Hendrickson 1999). The 

length at which 50% of individuals are sexually mature 

(L50) is 16-20 cm, depending on season and location 

(Brodziak 1995; Macy and Brodziak 2001; Hatfield and 

Cadrin 2002). 

ADULTS 

Historically, the lifespan of longfin inshore squid 

was believed to be 1-2 years (Summers 1971; Lange 

1982). However, Brodziak and Macy (1996), using 

statolith aging, demonstrated exponential growth and a 

lifespan of less than 1 year. 

Longfin inshore squid reach sizes greater than 40-

50 cm mantle length (ML), although most are less than 

30 cm (Vecchione et al. 1989; Brodziak 1995). They 

are sexually dimorphic – males grow more rapidly and 

reach larger size at age than females (Brodziak 1995). 

Growth depends on temperature (Hatfield et al. 2001) 

and is highest for individuals hatched during winter 

(Macy and Brodziak 2001). Longfin inshore squid 

migrate offshore during late autumn and overwinter in 

warmer waters along the edge of the continental shelf; 

they return inshore during the spring and early summer 

(MAFMC 1998). Mature individuals enter inshore 

waters before immature ones (Macy 1982). Off 

Massachusetts, larger individuals migrate inshore in 

April-May while smaller individuals move inshore in 

the summer (Lange 1982). Longfin inshore squid form 

large schools based on size prior to feeding (Macy 

1980) and make diurnal vertical migrations up into the 

water column at night (MAFMC 1998). This 
Page 2 

movement may be associated with the pursuit of food 

organisms such as euphausiids. 

REPRODUCTION 

Brodziak and Macy (1996), Macy and Brodziak 

(2001), and Hatfield and Cadrin (2002) show that 

longfin inshore squid spawn year round with seasonal 

and geographic peaks that vary among years and 

geographic areas (Lange and Sissenwine 1980). Most 

eggs are spawned in May and hatching occurs in July 

(Summers 1971). Spawning has been reported from 

August to September in the Bay of Fundy (Stevenson 

1934), from May to August in New England waters 

(Summers 1971; Macy 1980), and from late spring to 

early summer in the Middle Atlantic (Lange and 

Sissenwine 1983; Black et al. 1987). Mesnil (1977) 

reported that spawning on the Scotian Shelf and 

Georges Bank occurs during early spring and late 

summer. Spawning south of Cape Hatteras may also be 

important (Hatfield and Cadrin 2002). 

Spawning has been reported in the Gulf of Maine 

in Cobequid Bay and Massachusetts Bay (Bigelow 

1924), the Bay of Fundy (Stevenson 1934), Minas 

Basin (Cohen 1976), along the eastern coast of Nova 

Scotia in St. Margaret’s and Terrence bays (Dawe et al.

1990), on Georges Bank (Mesnil 1977), and in the 

Middle Atlantic in Narragansett and Delaware bays 

(Haefner 1959; Griswold and Prezioso 1981). 

Based on recent research, reproductive biology and 

behavior is complicated for longfin inshore squid. 

Visual and chemical cues regulate competition among 

males for females on spawning grounds (Buresch et al. 

2003). Females may lay multiple clutches over periods 

of up to several weeks (Maxwell and Hanlon 2000; 

King et al. 2003). Eggs in the same capsule from a 

single female may have multiple fathers from multiple 

spawning events and females appear to store sperm 

from spawning events for later use (Buresch et al. 

2001). 

FOOD HABITS 

The diet of the longfin inshore squid changes with 

size; small immature individuals feed on planktonic 

organisms (Vovk 1972b; Tibbetts 1977) while larger 

individuals feed on crustaceans and small fish 

(Vinogradov and Noskov 1979). Cannibalism is 

observed in individuals larger than 5 cm (Whitacker 

1978). Studies by Vovk and Khvichiya (1980) and 

Vovk (1985) showed that juveniles 4.1-6 cm long fed 

on euphausiids and arrow worms, while those 

6.1-10 cm fed mostly on small crabs, but also on 

polychaetes and shrimp. Adults 12.1-16 cm long fed on 

fish (clupeids, myctophids) and squid larvae/juveniles, 

and those > 16 cm fed on fish and squid (Vovk and 

Khvichiya 1980; Vovk 1985). Fish species preyed on 

by longfin inshore squid include silver hake, mackerel, 

herring, menhaden (Langton and Bowman 1977), sand 

lance, bay anchovy, menhaden, weakfish, and 

silversides (Kier 1982). Maurer and Bowman (1985) 

demonstrated the following seasonal and 

inshore/offshore differences in diet: in offshore waters 

in the spring, the diet is composed of crustaceans 

(mainly euphausiids) and fish; in inshore waters in the 

fall, the diet is composed almost exclusively of fish; 

and in offshore waters in the fall, the diet is composed 

of fish and squid. 

PREDATION 

Many pelagic and demersal fish species, as well as 

marine mammals and diving birds, prey upon juvenile 

and adult longfin inshore squid (Lange and Sissenwine 

1980; Vovk and Khvichiya 1980; Summers 1983). 

Marine mammal predators include longfin pilot whale, 

Globicephala melas, and common dolphin, Delphinus 

delphis (Waring et al. 1990; Overholtz and Waring 

1991; Gannon et al. 1997). Fish predators include 

bluefish, sea bass, mackerel, cod, haddock, pollock, 

silver hake, red hake, sea raven, spiny dogfish, angel 

shark, goosefish, dogfish, and flounder (Maurer 1975; 

Langton and Bowman 1977; Gosner 1978; Lange 

1980). 

GEOGRAPHICAL DISTRIBUTION 

Longfin inshore squid occur from Newfoundland 

to the Gulf of Venezuela, however, the principal 

concentrations exploited in the United States occur 

from Georges Bank to Cape Hatteras (Brodziak 1995). 

Longfin inshore squid are generally found at water 

temperatures of at least 9o

C (Lange and Sissenwine 

1980). The population makes seasonal migrations that 

appear to be related to bottom water temperatures; they 

move offshore during late autumn to overwinter along 

the edge of the continental shelf and return inshore 

during the spring and early summer (MAFMC 1998). 

When inshore waters are coldest during winter and 

early spring, the population concentrates along the outer 

edge of the continental shelf. The inshore movement to 

the shelf areas takes place when water temperatures are 

rising (Black et al. 1987) and begins in the south and 

proceeds north along the coast (MAFMC 1998). A 

northerly extension of the range has been noted in 

summer (Black et al. 1987). 

The terms ‘pre-recruit’ (unexploited sizes) and 

‘recruit’ (exploited sizes) are often used in reference to 
Page 3

longfin inshore squid. Exploitation begins at a 

minimum mantle length of about 9 cm. Thus, prerecruits are < 8 cm and recruits are > 9 cm. 

EGGS AND LARVAE 

The egg and larval stages of longfin inshore squid 

were not sampled by the Northeast Fisheries Science 

Center (NEFSC) Marine Resources Monitoring, 

Assessment and Prediction program (MARMAP) 

offshore ichthyoplankton surveys. 

PRE-RECRUITS 

The NEFSC bottom trawl surveys [see Reid et al. 

(1999) for details] captured longfin inshore squid prerecruits during all seasons (Figure 2; note that winter 

and summer distributions are presented as presence 

only data, precluding a discussion of abundances.). In 

winter, pre-recruits were captured from Cape Hatteras 

to Nantucket Shoals, although most were found south 

of Long Island. They were generally found offshore 

and concentrated toward the 200 m isobath. They were 

distributed a little farther inshore in the southern part of 

the range, presumably due to warmer water 

temperatures. In the spring, the distribution extended 

farther to the south, with high concentrations south of 

Cape Hatteras, and farther to the north, with high 

numbers in southern New England and some catches on 

Georges Bank and the Scotian Shelf. Higher 

concentrations were found near the 200 m isobath. In 

summer, they were concentrated nearshore, with a few 

found on central Georges Bank. In autumn, longfin 

inshore squid were distributed along the coast of Maine, 

in Massachusetts Bay, and from Georges Bank to south 

of Cape Hatteras from nearshore to the 200 m isobath, 

with some of the highest concentrations found 

nearshore. This presumably indicates the beginning of 

the offshore migration. 

The spring and fall distributions and abundances of 

pre-recruits around coastal Massachusetts, based on 

Massachusetts inshore bottom trawl surveys [see Reid 

et al. (1999) for details], are shown in Figure 3. In the 

spring, high concentrations occurred south of Cape Cod 

and around Martha’s Vineyard and Nantucket Island. 

Low numbers were found in and around Cape Cod Bay, 

and none were captured north of Cape Cod. Much 

higher numbers of pre-recruits were found in the fall. 

High concentrations were found in Buzzards Bay, 

around Martha’s Vineyard and Nantucket Island, 

throughout Cape Cod Bay, in Massachusetts Bay, and 

north and south of Cape Ann. The lower numbers of 

pre-recruits in inshore waters in the spring was most 

likely due to the survey occurring prior to the main part 

of the inshore migration. 

The seasonal distributions and abundances of 

prerecruits in Narragansett Bay, based upon the 

1990-1996 Rhode Island bottom trawl surveys, are 

shown in Figure 4. In winter, very few were caught, and 

they were only found at one station near the entrance to 

the Bay. Catches increased slightly in spring, and were 

highest during summer and autumn. This pattern 

corresponds to inshore migrations beginning in early 

spring. 

The distributions and abundances of both prerecruit and recruit longfin inshore squid in Long Island 

Sound from April to November 1986-1994, based on 

the Connecticut Fisheries Division bottom trawl 

surveys, are shown in Figure 5, Figure 6, and Figure 7. 

The following description of their distributions relative 

to depth and bottom type is taken almost verbatim from 

Gottschall et al. (2000). 

Longfin inshore squid taken in the survey ranged 

from 2-40 cm mantle length (Figure 5), with the largest 

squid present in May and June. Squid were rarely 

observed in April (4% occurrence), but from May 

through November they were commonly taken 

throughout the Sound. The percent occurrence varied 

little during these months, ranging from 63% in July to 

81% in September (Figure 6D). Abundance remained 

stable through late spring and summer (Figure 6A), and 

then increased dramatically in fall when squid ranging 

in size from 2-12 cm recruited to the trawl. 

Although squid were commonly encountered 

throughout Long Island Sound in late spring, they were 

most abundant east of Stratford Shoal, particularly in 

depths > 18 m on the transitional and sand bottom 

(Figure 6B and C) of the Mattituck Sill and the adjacent 

portion of the Central Basin (Figure 7). In addition, 

they were concentrated in Niantic Bay. In contrast, 

longfin inshore squid appeared to be more dispersed in 

summer. In fall, when small squid were abundant, they 

were distributed throughout the Sound, but were more 

abundant in the Central and Western Basins. During the 

fall generally, abundance tended to increase with depth 

and was highest over mud bottom, with abundance over 

transitional and sand bottoms ranking second and third 

respectively. Although the abundance of squid was very 

low in November, they were still commonly 

encountered throughout the Sound (65% occurrence). 

Abundance was similar over all bottom types but, as in 

the fall period, abundance tended to increase with depth 

(Gottschall et al. 2000). 

Longfin inshore squid pre-recruits were captured in 

the Hudson-Raritan estuary during spring, summer, and 

fall (Figure 8). They were found almost exclusively in 

the eastern portion of the bay and were collected in the 

highest numbers in the summer and autumn. 
Page 4 

RECRUITS 

NEFSC bottom trawl surveys captured longfin 

inshore squid recruits during all seasons (Figure 9; 

again note that winter and summer distributions are 

presented as presence data, precluding a discussion of 

abundances.). Their seasonal distributions are nearly 

identical to that of pre-recruits and illustrate the spring 

and summer inshore and the autumn offshore 

migrations. 

The distribution of longfin inshore squid recruits in 

waters off Massachusetts was almost identical to that of 

pre-recruits, although the overall number of recruits 

was much lower (Figure 10). 

Recruits were caught during all seasons in 

Narragansett Bay (Figure 11). Catches were low in 

winter, increased somewhat in spring, and were highest 

during summer and autumn. This pattern corresponds to 

inshore migrations beginning in spring. 

The distributions and abundances of both prerecruits and recruits in Long Island Sound were 

discussed previously. 

Longfin inshore squid recruits were captured in the 

Hudson-Raritan estuary during spring, summer, and fall 

(Figure 12). They were found mostly in the eastern 

portion of the bay; the highest catches occurred in 

summer and autumn. 

The 1988-1999 Virginia Institute of Marine 

Science (VIMS) trawl surveys of Chesapeake Bay 

suggests that recruit longfin inshore squid (> 12 cm) 

appeared in their catches primarily in April, with a few 

in May, and most likely were limited to sites around the 

Bay mouth and eastward (Geer 2002). 

HABITAT CHARACTERISTICS 

Information on the habitat characteristics and 

preferences of the longfin inshore squid are 

summarized in Table 1. 

EGGS AND LARVAE 

Egg masses are commonly found attached to rocks 

and small boulders on sandy/muddy bottom and on 

aquatic vegetation, such as Fucus sp., Ulva lactuca, 

Laminaria sp. and Porphyra sp. (Arnold et al. 1974; 

Griswold and Prezioso, 1981; Summers 1983). The 

eggs are demersal, are generally laid in waters < 50 m 

deep (Bigelow 1924; Griswold and Prezioso 1981; 

Lange 1982), and are found at temperatures of 10-23o

C 

(McMahon and Summers 1971) and salinities of 30-32 

ppt (McMahon and Summers 1971). 

The larvae are pelagic near the surface (McMahon 

and Summers 1971; McConathy et al. 1980) and occur 

at temperatures of 10-26o

C and salinities of 31.5-34.0 

ppt (Vecchione 1981). Surface waters are important to 

hatchlings and larvae and individuals move deeper as 

they grow older (Vecchione 1981). Longfin inshore 

squid larvae were common in ichthyoplankton samples 

across a wide range of depths and areas (Vecchione et 

al. 2001). 

PRE-RECRUITS 

Juveniles inhabit the upper 10 m of the water 

column over water 50-150 m deep (Mercer 1969; Vovk 

and Khvichiya 1980; Brodziak and Hendrickson 1999). 

They are found at surface water temperatures of 10-

26o

C (Vecchione 1981; Brodziak and Hendrickson 

1999) and salinities of 31.5-34.0 ppt (Vecchione 1981). 

Longfin inshore squid move up (nighttime) and down 

(daytime) in the water column on a daily (diel) basis 

(Hatfield and Cadrin 2002) but the importance of offbottom habitat is unknown because sampling has been 

primarily with bottom trawls. Diel migration patterns 

depend on squid size and season (Hatfield and Cadrin 

2002). 

Distributions of pre-recruits relative to bottom 

water temperature, depth, and salinity based on spring 

and fall NEFSC bottom trawl surveys are shown in 

Figure 13. During the spring surveys, pre-recruits were 

found in a temperature range of 4-21oC, with the 

majority at about 8-14oC. They were found over a depth 

range of 1-400 m, and a salinity range of 31-36, with 

most found at 34-36 ppt. During the fall the pre-recruits 

were found over a wider temperature range of 6-28oC, 

with peaks in abundance between roughly 10-19oC. 

Their depth range during that season was between 1-

400 m, with the majority found above about 60 m. 

Their salinity range was between 29-36 ppt, with the 

majority at 32-33 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 14. In 

the spring, the pre-recruits were found at a temperature 

range of 5-17oC, with most at 10-14oC. Their depth 

range was from 6 m to a depth of approximately 65 m; 

the majority were found between 6-25 m. In the fall 

they were found over a wider temperature range of 5-

22oC, with bimodal peaks at about 8-10oC and a larger 

one 16-20oC. Their depth range during fall was between 

1-85 m, with the majority found between about 6-35 m. 

In the Narragansett Bay bottom trawl survey, prerecruits were found at depths ranging from 10-110 feet 

(3-367 m) (Figure 15). In winter the few pre-recruits 

caught were taken at 90 feet (27 m), in summer and 

spring most were caught at 20-40 feet (6-12 m) and 
Page 5

100-110 feet (30-34 m), and in autumn most were 

caught at 100 feet (30 m). Pre-recruits were collected 

at temperatures ranging from 9-25o

C. They were 

collected at temperatures of 10oC in winter, from 9-

16o

C in spring, from 11-25oC with most at 19oC in 

summer, and from 13-23o

C with most at 20oC in 

autumn. 

In the Hudson-Raritan estuary, pre-recruits were 

collected at temperatures ranging from 11-24o

C, but 

most were taken at 16-21oC. They were also collected 

at depths of 15-75 ft (~5-23 m), with most at 30 ft (9 m) 

and 45-50 ft (14-15 m), and salinities of 20-33 ppt, with 

the highest catch at 30 ppt. They were found at 

dissolved oxygen levels of 5-10 mg/L, with most at 7-8 

mg/L (Figure 16). Longfin inshore squid require 

oxygen concentrations greater than 4 mg/L (Howell and 

Simpson 1994). 

The distributions and abundances of both prerecruit and recruit squid in Long Island Sound relative 

to depth and bottom type, based on surveys by 

Gottschall et al. (2000), were discussed previously in 

Geographic Distribution: Pre-recruits. 

RECRUITS 

Adult longfin inshore squid inhabit the continental 

shelf and upper continental slope to depths of 400 m 

(Vecchione et al. 1989), but depth varies seasonally. In 

spring they occur at depths of 110-200 m (Serchuk and 

Rathjen 1974; Lange and Sissenwine 1980), in summer 

and autumn they inhabit inshore waters as shallow as 6-

28 m (Summers 1968a, b; Serchuk and Rathjen 1974; 

Gosner 1978; Howell and Simpson 1994), and in winter 

they inhabit offshore waters to depths of 365 m (Lange 

1982). They are found on mud or sand/mud substrate 

(Howell and Simpson 1994), at surface temperatures 

ranging from 9-21o

C, and bottom temperatures ranging 

from 8-16o

C (Summers 1969; Lux et al. 1974; 

Serchuck and Rathjen 1974; Lange and Sissenwine 

1980; Macy 1980; Brodziak and Hendrickson 1999). 

Adults, like juveniles, migrate up and down in the water 

column in response to light conditions and the 

importance of off-bottom habitat is unknown. 

Distributions of recruits relative to bottom water 

temperature, depth, and salinity based on spring and fall 

NEFSC bottom trawl surveys are shown in Figure 17. 

During the spring, recruits were found in a temperature 

range of 4-21oC, with the majority at about 7-13oC. 

They were found over a depth range of 1-400 m, and a 

salinity range of about 30-36, with most found at 34-35 

ppt. During the fall the pre-recruits were found over a 

wider temperature range of 6-28oC, with a peak 

between about 10-15oC. Their depth range during that 

season was between 1-400 m, with the majority found 

above about 70 m. Their salinity range was between 30-

37 ppt, with most at 32-33 ppt. 

Around Massachusetts in the spring, the recruits 

were found at a temperature range of 6-17oC, with most 

at 10-13oC (Figure 18). Their depth range was from 

about 1 m to approximately 50 m, with the majority 

found between 6-20 m. As with the pre-recruits, the 

recruits in the fall were found over a wider temperature 

range of 5-22oC, with bimodal peaks at about 8-10oC 

and a larger one 16-20oC. Their depth distribution 

during fall was similar to that of the pre-recruits (range 

of 1-85 m, with the majority found between about 6-35 

m). 

In Narragansett Bay, recruits were found at depths 

ranging from 10 to 120 ft (3-37 m) (Figure 19). In 

winter the few recruits caught were taken at 90-100 ft 

(27-30 m). In summer and spring they were taken at 

depths ranging from 10-120 ft (3-37 m). In spring, 

about 40% were caught at 100-110 feet (30-34 m), with 

another 20% found at 70 ft (21 m), while in summer, 

the majority were caught at 100-110 ft. In autumn, most 

were caught at 90-100 feet (27-30 m). Recruits were 

taken at temperatures ranging from 7-25o

C (Figure 19). 

Seasonally they were collected at 7-10oC in winter, 

with almost all caught at 10o

C; at 9-16oC in spring, with 

most at 9-13oC; at 9-25oC in summer, with most at 18-

21oC; and at 11-23oC in autumn, with a peak at 15oC. 

In the Hudson-Raritan estuary, recruits were 

collected at temperatures ranging from 9-24o

C, but 

most were at 16-17oC (Figure 20). They were also 

collected at depths of 10-75 ft (~5-23 m), with most at 

50 and 60 ft (~15-18 m), and salinities of 20-33 ppt, 

with the highest catch at 30 ppt. They were found at 

dissolved oxygen levels of 5-11 mg/L, with most at 7-8 

mg/L. Longfin inshore squid require oxygen 

concentrations greater than 4 mg/L (Howell and 

Simpson 1994). 

RESEARCH NEEDS 

Human impacts may be significant on sandy 

bottom habitats used by inshore longfin squid for 

their eggs. However, little information is available 

on egg habitat locations, seasonal occurrence, 

sediment characteristics, and depth or water 

chemistry. This type of information might be 

useful for designating marine reserves, seasonal 

closed areas, and other measures. 

Additional information about use of off-bottom 

habitat and vertical distribution of inshore longfin 

squid in the water column is needed for stock 

assessment and management. This is because a 

substantial portion of the inshore longfin squid 

stock may be unavailable to bottom trawl surveys 

that are used to track abundance. 

Information about distribution of inshore longfin 

squid in deepwater off the continental shelf and 

south of Cape Hatteras would be useful because 
Page 6 

bottom trawl surveys do not reach these areas and 

an unknown portion of the stock is resident there 

(NEFSC 2002). 

More information on growth rates and maturity are 

needed from geographically and temporally diverse 

studies. 

The commercially exploited population from Cape 

Hatteras to Georges Bank, inshore and offshore 

and in all seasons, is considered a single stock unit. 

More information is needed on stock structure, 

including gene flow and levels of genetic 

differentiation among geographic areas. 

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Waring, G.T., P. Gerrior, P.M. Payne, B.L. Parry, and 

J.R. Nicolas. 1990. Incidental take of marine 

mammals in foreign fishery activities off the 

northeast United States, 1977-1988. Fish. Bull. 

(U.S.) 88: 347-360. 

Whitaker, J.D. 1978. A contribution to the biology of 

Loligo pealei and Loligo plei (Cephalopoda, 

Myopsida) off the southeastern coast of the United 

States. M.S. thesis, College of Charleston. 

Charleston, SC. 164 p. 

Young, R.E. and R.F. Harman. 1988. “Larva,” “paralarva,” and “subadult” in cephalopod terminology. 

Malacologia 29: 201-207. 
Page 9

Table 1. Summary of life history and habitat characteristics for longfin inshore squid, based on the pertinent literature. 

This table is essentially the same as that used in the first longfin inshore squid EFH source document (Cargnelli et al. 

1999); more recent studies have not been added. 

Life Stage Size and Growth Habitat Substrate Temperature Salinity 

Eggs 1 Incubation time varies 

with temperature: 26.7 d 

at 12-18o

C, 18.5 d at 

15.5-21.3o

C, and 10.7 d 

at 15.5-23.0oC. 

Eggs generally in shallow 

waters, < 50 m and near 

shore. 

Egg masses are 

commonly found on 

sandy/mud bottom; 

usually attached to 

rocks/boulders, pilings, 

or algae such as Fucus, 

Ulva lactuca, Laminaria

and Porphyra sp. 

Eggs found in waters 10-

23o

C; usually > 8oC. 

Optimal development at 

12o

C. 

Found at 30-32 ppt. 

Larvae 2 Paralarvae range in size 

from 1.4-15 mm ML 

(mantle length). 

Growth rates slower for 

winter-hatched animals 

than spring-hatched. 

Found in coastal, surface 

waters in spring, summer, and 

fall. Hatchlings found in 

surface waters day and night. 

Move deeper in water column 

as they grow larger. 

 

Found at 10-26o

C (at 

lower temperatures found 

at higher salinities). 

Found at 31.5-34.0 

ppt. 

Juveniles 3 Size ranges from 

approximately 15 mm - 8 

cm. 

At 6-8 cm sexual size 

dimorphism is evident, 

before offshore 

migrations occur. 

Growth rates of youngof-the-year are 12-38 

mm/month. 

Inhabit upper 10 m at depths 

of 50-100 m on continental 

shelf. Found in coastal 

inshore waters in spring/fall, 

offshore in winter. Migrate to 

surface at night. 

Ontogenetic descent: at 45 

mm, chromatophores are 

concentrated on dorsal rather 

than ventral surface, 

indicating a change from 

inhabiting surface waters to 

demersal lifestyle. 

 

Found at 10-26o

C (at 

lower temperatures found 

at higher salinities). 

Juveniles prefer warmer 

bottom temperatures and 

shallower depths in fall 

than adults. 

Found at 31.5-34.0 

ppt. 

Adults 4 Smallest size at maturity 

8 cm ML; most are > 10 

cm ML. 

Males grow faster than 

females and attain larger 

sizes; larger sizes at 

higher latitudes. 

Growth is rapid, faster in 

warm months (1.5-2.0 

cm/month) than in cold 

months (0.4-0.6 

cm/month). Life span is 

< 1 year. Maximum size 

and age are ~50 cm ML, 

3 yrs. 

Range from Newfoundland 

south to Cape Hatteras, on 

continental shelf and upper 

slope. Most abundant from 

Gulf of Maine to Hatteras. 

March-October: inshore, 

shallow waters up to 180 m. 

Winter: offshore deeper 

waters, up to 400 m on shelf 

edge. 

Most abundant at bottom 

during the day; move upwards 

at night. Generally found at 

greater depths and cooler 

bottom temperatures in the 

fall than juveniles. 

Importance of off-bottom 

habitat poorly understood. 

Mud or sandy mud. Found at surface 

temperatures ranging 

from 9-21o

C and bottom 

temperatures ranging 

from 8-16o

C. 

1

 Bigelow (1924); McMahon and Summers (1971); Arnold et al. (1974); Griswold and Prezioso (1981); Lange (1982); Summers (1983); Dawe et al.

(1990). 2 McMahon and Summers (1971); McConathy et al. (1980); Vecchione (1981); Nesis (1982); Vovk (1983); Young and Harman (1988). 3

 Summers (1968a, b); Mercer (1969); Macy (1980); Vovk and Khvichiya (1980); Vecchione (1981); Young and Harman (1988); Brodziak and 

Henderson (1999). 4

 Haefner (1964); Summers (1968a, b, 1969, 1971, 1983); Rathjen (1973); Lux et al. (1974); Serchuk and Rathjen (1974); Cohen (1976); Mesnil 

(1977); Gosner (1978); Sissenwine and Bowman (1978); Lange (1980, 1982); Lange and Sissenwine (1980); Macy (1980); Nesis (1982); Vecchione 

et al. (1989); Dawe et al. (1990); Howell and Simpson (1994); Brodziak and Macy (1996); Brodziak and Henderson (1999). 
Page 10

Table 1. Cont’d. 

Life Stage Prey Predators Spawning Notes 

Eggs 1 N/A 

 

Most eggs are spawned in May, 

hatching occurs in July. 

Fecundity ranges from 

950-15,900 eggs per female. 

Eggs are demersal. Enclosed in 

a gelatinous capsule containing 

up to 200 eggs. Each female 

lays 20-30 capsules. Laid in 

masses made up of hundreds of 

egg capsules from different 

females. 

Larvae 2 Primary prey are copepods. 

 

"Paralarvae" defined as stage 

after hatching when 

cephalopods are pelagic. 

Tentacles are non-functional at 

< 15 mm. 

Juveniles 3 Primary prey varies with size: 

< 4.0 cm: plankton, copepods; 

4.1-6.0 cm: euphausiids, 

arrow worms; 

6.1-10.0 cm: crabs, 

polychaetes, shrimp. 

Cannibalism observed in 

specimens larger than 5 cm 

ML (small Illex illecebrosus 

were found in 49 of 322 

Loligo stomachs).

Many pelagic and demersal fish 

species as well as marine 

mammals and birds. 

 

Changes in habitat as the squid 

grows are indicated by changes 

in the diet. 

Adults 4 Fish prey includes silver hake, 

mackerel, herring, menhaden, 

sand lance, bay anchovy, 

menhaden, weakfish, and 

silversides. Invertebrate prey 

includes crustaceans 

(Crangon, Palaeomonetes sp.) 

and squid. 

15 cm adults can eat fish up to 

half their mantle length. At 

16-25 cm, consume more fish 

and less crustaceans as growth 

increases; > 25 cm, more 

squid than fish eaten; and > 

30 cm, almost exclusively 

squid. 

Predators include many fishes 

(bluefish, sea bass, mackerel, 

cod, haddock, pollock, hakes, 

sea raven, goosefish, flounder, 

dogfish, angel sharks, skates), 

pilot whale (Globicephala 

melas) and common dolphin 

(Delphinus delphis), and diving 

birds. 

Spawning occurs on Scotian 

Shelf, Georges Bank, Gulf of 

Maine, and from Nantucket 

Shoals to Cape Hatteras in 

shallow waters, 10-90 m, from 

April-November (New England: 

May-August; Bay of Fundy: 

Aug-September). Georges 

Bank: two broods - early spring 

and late summer. Spring spawn: 

hatch in June, mature over 

winter. Summer spawn: hatch in 

fall, mature in 2nd winter. 

Mating occurs during inshore 

migration in spring. Mortality 

occurs after first spawning. 

Loligo form schools according 

to size class prior to feeding. 

Oxygen requirement > 4 ml/l. 

Larger individuals migrate 

earlier (April-May) than smaller 

ones. 

1

 Haefner (1959); Summers (1971); Vovk (1972b), Arnold et al. (1974); Gosner (1978); Griswold and Prezioso (1981); Lange (1982); Nesis (1982); 

Lange and Sissenwine (1983). 2

 Vecchione (1981); Vovk (1983); Young and Harman (1988). 

3

 Vovk (1972b, 1985); Tibbetts (1977); Whitaker (1978); Vinogradov and Noskov (1979); Vovk and Khvichiya (1980); Vecchione (1981). 4

 Stevenson (1934); Summers (1969, 1971); Vovk (1972a, 1985); Rathjen (1973); Maurer (1975); Cohen (1976); Langton and Bowman (1977); 

Mesnil (1977); Tibbetts (1977); Gosner (1978); Vinogradov and Noskov (1979); Lange (1980, 1982); Lange and Sissenwine (1980, 1983); Macy 

(1980); Griswold and Prezioso (1981); Kier (1982); Summers (1983); Maurer and Bowman (1985); Dawe et al. (1990); Waring et al. (1990); 

Overholtz and Waring (1991); Howell and Simpson (1994); Brodziak and Macy (1996); Gannon et al. (1997). 

 
Page 11

Figure 1. The longfin inshore squid, Loligo pealeii (from Goode 1884). 
Page 12

Figure 2. Seasonal distributions and abundances of pre-recruit longfin inshore squid collected during NEFSC bottom 

trawl surveys. 

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

only. 
Page 13

Figure 2. Cont’d. 

Based on NEFSC spring bottom trawl surveys (1968-2003, all years combined). Survey stations where pre-recruits were 

not found are not shown. 
Page 14

Figure 2. Cont’d. 

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

presence only. 
Page 15

Figure 2. Cont’d. 

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

found are not shown. 
Page 16

Figure 3. Distribution and abundance of pre-recruit longfin inshore squid in Massachusetts coastal waters. 

Based on spring Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Survey stations where 

pre-recruits were not found are not shown. 
Page 17

Figure 3. Cont’d. 

Based on fall Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Survey stations where prerecruits were not found are not shown. 
Page 18

Figure 4. Seasonal distribution and abundance of longfin inshore squid pre-recruits in Narragansett Bay. 

Based upon the 1990-1996 Rhode Island bottom trawl surveys. The numbers shown at each station are the average catch 

per tow rounded to one decimal place [see Reid et al. (1999) for details]. 
Page 19

Figure 5. Monthly log10 length frequencies (cm) of longfin inshore squid collected in Long Island Sound, based on 

106,925 squid taken in 771 tows between 1987 and 1994. Source: Gottschall et al. (2000). 
Page 20

Figure 6. Relative abundance (geometric mean catch/tow) catch/tow and percent occurrence (proportion of samples in 

which at least one individual was observed) for longfin inshore squid in Long Island Sound, by month, month and 

bottom type, and month and depth interval. Source: Gottschall et al. (2000). 
Page 21

Figure 7. Distribution and abundances of longfin inshore squid in Long Island Sound, based on the finfish surveys of 

the Connecticut Fisheries Division, 1986-1994. Source: Gottschall et al. (2000). Circle diameter is proportional to the 

number of squid caught, and is scaled to the maximum catch (indicated by “max=” or “max>”); the largest circle 

represents a tow with a catch of > 2,500 squid. Collections were made with a 14 m otter trawl at about 40 stations chosen 

by stratified random design. 
Page 22

Figure 8. Seasonal distribution and abundance of longfin inshore squid pre-recruits collected in the Hudson-Raritan 

estuary. 

Based on NEFSC Hudson-Raritan trawl surveys, January 1992 – June 1997 [see Reid et al. (1999) for details]. 
Page 23

Figure 9. Seasonal distributions and abundances of recruit longfin inshore squid collected during NEFSC bottom trawl 

surveys. 

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

only. 
Page 24

Figure . Cont’d. 

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

found are not shown. 
Page 25

Figure 9. Cont’d. 

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

presence only. 
Page 26

Figure 9. Cont’d. 

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

found are not shown. 
Page 27

Figure 10. Seasonal distributions and abundances of recruit longfin inshore squid in Massachusetts coastal waters. 

Based spring Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Survey stations where 

recruits were not found are not shown. 
Page 28

Figure 10. Cont’d. 

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

recruits were not found are not shown. 
Page 29

Figure 11. Seasonal distribution and abundance of longfin inshore squid recruits in Narragansett Bay. 

Based upon the 1990-1996 Rhode Island bottom trawl surveys. The numbers shown at each station are the average catch 

per tow rounded to one decimal place. 
Page 30

Figure 12. Seasonal distribution and abundance of longfin inshore squid recruits collected in the Hudson-Raritan 

estuary. 

Based on NEFSC Hudson-Raritan trawl surveys, January 1992 – June 1997. 
Page 31

Longfin Squid

NEFSC Bottom Trawl Survey

Spring 1968 - 2003

Pre-recruits (<=8 cm)

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=12514

Occurrence N=2260

Catch N=675377

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=436

Catch N=157137

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=10879

Occurrence N=1967

Catch N=546317

Figure 13. Distributions of pre-recruit longfin inshore 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 longfin inshore squid occurred and medium bars show, within each interval, the percentage of the total number 

of longfin inshore squid caught. Note that the bottom depth interval changes with increasing depth. 
Page 32

Longfin Squid

NEFSC Bottom Trawl Survey

Fall 1963 - 2003

Pre-recruits (<=8 cm)

0

5

10

15

20

25

30

35

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=14186

Occurrence N=6018

Catch N=2369271

0

10

20

30

40

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

Salinity (PPT)

Percent

Trawls N=2272

Occurrence N=1252

Catch N=568694

0

2

4

6

8

10

12

14

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

Bottom Temperature (°C)

Percent

Trawls N=12207

Occurrence N=5079

Catch N=1927730

Figure 13. 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 longfin inshore squid occurred and medium bars show, within each interval, the percentage of the total number of 

longfin inshore squid caught. Note that the bottom depth interval changes with increasing depth. 
Page 33

Longfin Squid

Massachusetts Inshore Trawl Survey

Spring 1978 - 2003

Pre-recruits (<=8 cm)

0

5

10

15

20

25

30

35

40

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=756

Catch N=30502

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=746

Catch N=30469

Figure 14. Distributions of pre-recruit longfin inshore 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 longfin inshore squid occurred and 

medium bars show, within each interval, the percentage of the total number of longfin inshore squid caught. 
Page 34

Longfin Squid

Massachusetts Inshore Trawl Survey

Fall 1978 - 2003

Pre-recruits (<=8 cm)

0

5

10

15

20

25

30

35

40

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=1947

Catch N=789693

0

5

10

15

20

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=1862

Catch N=751665

Figure 14. 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 longfin inshore squid occurred and 

medium bars show, within each interval, the percentage of the total number of longfin inshore squid caught. 
Page 35

Figure 15. Distributions of longfin inshore squid pre-recruits in Narragansett Bay relative to mean bottom water 

temperature and bottom depth. 

Based on Rhode Island trawl surveys, 1990-1996. Open bars represent the proportion of all stations surveyed, while 

solid bars represent the proportion of the sum of all catches. 
Page 36

Figure 16. Distributions of longfin inshore squid pre-recruits in the Hudson-Raritan estuary relative to bottom water 

temperature, depth, dissolved oxygen, and salinity. 

Based on NEFSC Hudson-Raritan estuary trawl surveys, 1992-1997, all seasons and years combined. Open bars 

represent the proportion of all stations surveyed, solid bars represent the proportion of the sum of all standardized 

catches. 
Page 37

Longfin Squid

NEFSC Bottom Trawl Survey

Spring 1968 - 2003

Recruits (>=9 cm)

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=3244

Catch N=252814

0

10

20

30

40

50

60

70

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

Salinity (PPT)

Percent

Trawls N=2270

Occurrence N=599

Catch N=40951

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=2861

Catch N=213009

Figure 17. Distributions of recruit longfin inshore 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 longfin inshore squid occurred and medium bars show, within each interval, the percentage of the total number 

of longfin inshore squid caught. Note that the bottom depth interval changes with increasing depth. 
Page 38

Longfin Squid

NEFSC Bottom Trawl Survey

Fall 1963 - 2003

Recruits (>=9 cm)

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=14186

Occurrence N=7410

Catch N=621412

0

10

20

30

40

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

Salinity (PPT)

Percent

Trawls N=2272

Occurrence N=1459

Catch N=112554

0

5

10

15

20

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

Bottom Temperature (°C)

Percent

Trawls N=12207

Occurrence N=6267

Catch N=519224

Figure 17. 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 longfin inshore squid occurred and medium bars show, within each interval, the percentage of the total number of 

longfin inshore squid caught. Note that the bottom depth interval changes with increasing depth. 
Page 39

Longfin Squid

Massachusetts Inshore Trawl Survey

Spring 1978 - 2003

Recruits (>=9 cm)

0

5

10

15

20

25

30

35

40

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=982

Catch N=51384

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=959

Catch N=50884

Figure 18. Distributions of recruit longfin inshore 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 longfin inshore squid occurred and 

medium bars show, within each interval, the percentage of the total number of longfin inshore squid caught. 
Page 40

Longfin Squid

Massachusetts Inshore Trawl Survey

Fall 1978 - 2003

Recruits (>=9 cm)

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=1376

Catch N=37754

0

5

10

15

20

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=1325

Catch N=36545

Figure 18. 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 longfin inshore squid occurred and 

medium bars show, within each interval, the percentage of the total number of longfin inshore squid caught. 
Page 41

Figure 19. Distributions of longfin inshore squid recruits in Narragansett Bay relative to mean bottom water temperature 

and bottom depth. 

Based on Rhode Island trawl surveys, 1990-1996. Open bars represent the proportion of all stations surveyed, while 

solid bars represent the proportion of the sum of all catches. 
Page 42

Figure 20. Distributions of longfin inshore squid recruits in the Hudson-Raritan estuary relative to bottom water 

temperature, depth, dissolved oxygen, and salinity. 

Based on NEFSC Hudson-Raritan estuary trawl surveys, 1992-1997, all seasons and years combined. Open bars 

represent the proportion of all stations surveyed, solid bars represent the proportion of the sum of all standardized 

catches. 
Publishing in NOAA Technical Memorandum NMFS-NE

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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 Division

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

important species or habitat topics; proceedings and collected papers of scientific meetings; and indexed and/or annotated bibliographies.

All issues receive internal scientific review and most issues receive technical and copy editing.

Northeast Fisheries Science Center Reference Document -- This series is issued irregularly. The series typically includes: data reports

on field and lab studies; progress reports on experiments, monitoring, and assessments; background papers for, collected abstracts of,

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

or copy editing.

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

Reference Document, or to subscribe to the Resource Survey Report, either contact the NEFSC Editorial Office (166 Water St., Woods

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

 MAIL