NOAA Technical Memorandum NMFS-NE-189

U. S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

Northeast Fisheries Science Center

Woods Hole, Massachusetts

September 2004

Essential Fish Habitat Source Document:

Sea Scallop, Placopecten magellanicus,

Life History and Habitat Characteristics

Second Edition
166. Report on the Third Northwest Atlantic Herring Acoustic Workshop, University of Maine Darling Marine Center,

Walpole, Maine, March 13-14, 2001. By William L. Michaels, ed. & co-conven., and Philip Yund, co-conven. December 2001. iv +

18 p., 14 figs., 2 app. NTIS Access. No. PB2003-101556.

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. In press.

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

September 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-189

Deborah R. Hart and Antonie S. Chute

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

Essential Fish Habitat Source Document:

Sea Scallop, Placopecten magellanicus,

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).

a

Robins, 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. b

Robins, 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. 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. 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. g

McEachran, 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 sea scallop 

EFH source document is based on the original by David 

B. Packer, Luca M. Cargnelli, Sara J. Griesbach, and 

Sandra E. Shumway, with a foreword by Jeffrey N. 

Cross (Packer 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 ..................................................................................................................................5

HABITAT CHARACTERISTICS .......................................................................................................................................5

ACKNOWLEDGEMENTS..................................................................................................................................................6

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

Tables 

Table 1. Relative abundance of egg, larval, juvenile, and adult sea scallop, Placopecten magellanicus, in 

New England bays and estuaries by salinity zone............................................................................................12

Table 2. Summary of sea scallop habitat parameters based on the pertinent literature..................................................13

Figures 

Figure 1. The sea scallop, Placopecten magellanicus (photograph by Dann Blackwood, USGS)..................................16

Figure 2. Generalized life cycle of the sea scallop, from Stewart and Arnold (1994).....................................................17

Figure 3. Map of Georges Bank indicating the five sub-regions and the clockwise flow of the residual current. ..........18

Figure 4. Distribution of sea scallop spawning beds off the northeast coast of North America......................................19

Figure 5. Distribution and abundance of sea scallops collected during NEFSC scallop surveys during summer 1979-

1993. ................................................................................................................................................................20

Figure 6. Distribution and abundance of sea scallops collected during NEFSC scallop surveys during summer 1994-

2003. ................................................................................................................................................................21

Page 1

INTRODUCTION 

The Atlantic sea scallop, Placopecten magellanicus

(Gmelin), is a bivalve mollusk of the family Pectinidae 

and order Ostreoida (Figure 1). It occurs on the 

continental shelf of the northwest Atlantic from the 

north shore of the Gulf of St. Lawrence south to Cape 

Hatteras, North Carolina. The Atlantic sea scallop is 

one of the most economically important species in the 

northeast United States, and supports the most valuable 

wild scallop fishery in the world. The 2003 U.S. 

landings of 25,476 MT (meats) had an ex-vessel value 

of about $229,000,000. The sea scallop fishery is 

managed under the New England Fishery Management 

Council’s Sea Scallop Management Plan (NEFMC 

2003). This Essential Fish Habitat source document 

provides information on the life history and habitat 

characteristics of sea scallops inhabiting U.S. waters in 

the Gulf of Maine, Georges Bank, and the Middle 

Atlantic Bight. 

LIFE HISTORY 

Besides the account given here, the life history of 

the sea scallop is summarized in NEFSC (2004), 

Stewart and Arnold (1994), Mullen and Moring (1986), 

and Shumway et al. (in prep.). An overview of scallop 

biology and the early history of the fishery may be 

found in Bourne (1964). The life cycle of the sea 

scallop is depicted in Figure 2. 

EGGS 

Sea scallops are among the most fecund of 

bivalves, producing gametes starting in the first or 

second year and eventually producing up to 270 million 

eggs (Langton et al. 1987). The average diameter of 

spawned eggs is 66.81.6 m (Langton et al. 1987). 

After fertilization, the eggs are slightly heavier than 

seawater and probably remain on the sea floor until 

they develop into the first free-swimming larval stages. 

LARVAE 

The first two larval stages of the sea scallop, 

trochophore and veliger, are pelagic. The larvae remain 

planktonic for over a month after hatching (Posgay 

1982). The larvae drift with water currents, but can 

also move independently on a small scale. Veliger 

larvae have been observed to move vertically at 

velocities of 1-1.5 mm s-1, upward by ciliary action and 

downward by sinking with the valves closed (Manuel et 

al. 2000). Tremblay and Sinclair (1990b) found that in 

well-mixed areas, larvae were distributed evenly 

through the water column, while in stratified areas they 

aggregated above the pycnocline. Manuel et al. (2000) 

observed that veligers were able to detect and react to 

small temperature gradients. In stratified areas, the 

veliger’s position relative to the thermocline may be 

beneficial for dispersion, growth, and feeding. 

During the planktonic stage, the shell, eye spots, 

and foot develop (Naidu 1991). A description of larval 

development in the laboratory has been published by 

Culliney (1974) and growth of larval scallops in the 

laboratory has been studied by Hurley et al. (1986). 

Duration of the larval stages is shorter at higher 

temperatures and where food is more plentiful. 

Identification of planktonic sea scallop larvae in 

the wild has previously been described (Tremblay et al.

1987) but the horizontal distribution of larval scallops is 

still largely unknown. Since scallop larvae are 

planktonic, larval transport is influenced by the flow of 

currents in and around spawning areas. 

Sea scallops on Georges Bank are thought to be 

self-sustaining because many of the larvae from this 

region are retained in the Georges Bank gyre (see 

Figure 3) long enough for metamorphosis to be 

complete (Tremblay et al. 1994). Good year classes on 

Georges Bank are associated with tight autumnal gyres 

that tend to retain larvae on the Bank and poorer year 

classes are associated with loose gyres (Posgay 1950). 

McGarvey et al. (1993) found a correlation between 

egg production on the Northern Edge and Peak area of 

Georges Bank and recruitment on Georges Bank, 

possibly due to the high percentage of spawning stock 

biomass that was located in the Northern Edge and 

Peak during the study period. 

Tremblay and Sinclair (1986) reported on the 

horizontal distribution of larval scallops within the Bay 

of Fundy. They found that there was some transport of 

larvae within the Bay by residual currents but that most 

of the larvae either remained in, or were returned to, the 

area of major spawning. They were further able to 

demonstrate that although there was larval dispersal 

beyond the Bay of Fundy, no large-scale exchange 

occurred between Georges Bank and the Scotian Shelf. 

SPAT 

At the end of their pelagic existence, the larvae 

enter the pediveliger stage with the development of a 

foot and byssus gland which secretes threads that are 

used to attach to hard surfaces (Culliney 1974). 

Spatfall (the settling of larval scallops to the bottom), 

and the period immediately following, is thought to be 

particularly important in the formation of scallop beds 
Page 2 

(Posgay 1953) and in determining year class size 

(Bourne 1965; Caddy 1975). Settlement is assumed to 

occur by mid-December on Georges Bank (Thouzeau et 

al. 1991a). The transition to the benthos of the freeswimming larvae occurs at a size of about 0.25 mm 

shell height and is accompanied by drastic changes in 

diet, morphology, and locomotory ability. Mortality 

during natural settlement may be as high as when spat 

are reared artificially (Bourne 1965; Culliney 1974). 

When scallop spat settle, they are extremely 

delicate and do not survive on shifting sand bottoms 

(Merrill and Edwards 1976). Those that land on 

sedentary branching plants, animals or on any other 

hard surface that offers freedom of shell movement may 

have a distinct survival advantage (Larsen and Lee 

1978). 

The availability of suitable surfaces on which to 

settle seems to be a primary requirement for scallop 

survival. Pediveligers show a thigmotactic settling 

response to shell fragments and small pebbles; 

settlement occurs predominantly on the undersides of 

these objects (Caddy 1968; Culliney 1974). Spat also 

settle on navigation buoys (Merrill and Edwards 1976), 

on the red alga Rhodomela conferroides (Naidu 1970), 

on hydroids such as Hydrallmania, and on amphipod 

tubes (Larsen and Lee 1978). Spat settlement varies 

with depth and water turbulence: numbers of spat 

generally increase with increasing depth, but this 

relationship is less evident with increasing water 

turbulence (Pearce et al. 1998). 

A close association exists between the bryozoan, 

Eucratea loricata (= Gemelleria of Baird 1953), and 

post-larval scallops. Eucratea may contain large 

numbers of 2-5 mm juvenile scallops (Baird 1953; 

Caddy 1972). The scallops detach themselves from the 

bryozoan when about 4-5 mm in shell height and then 

attach themselves to the shells of larger scallops (Dow 

1956). 

Pediveligers may delay metamorphosis for a 

considerable time until suitable physical substrates for 

settlement are encountered; however, the effects of this 

delay on the scallops are not known (Culliney 1974). 

JUVENILES AND ADULTS 

Juvenile scallops (5-12 mm shell height) leave the 

original substrate on which they have settled and attach 

themselves by byssus to shells and bottom debris (Dow 

and Baird 1960). As the young scallops grow older, 

they lose their byssal attachment (Caddy 1972). 

Scallops are relatively active until they are about 80 

mm shell height, they swim to escape predation (Baird 

1954) and disturbances such as commercial dredging 

(Caddy 1968). While swimming, young scallops can be 

carried long distances by currents (Baird 1954). The 

maximum size for frequent swimming was found to be 

110 mm (4-6 yrs old). 

There is no evidence of mass migrations by 

scallops. The movements of sea scallops are usually 

localized, and random or current-assisted. Numerous 

tagging experiments have shown that once aggregations 

of adults are formed, they remain fixed (Baird 1954; 

Dickie 1953; Naidu 1970; Schick 1979). Posgay 

(1981) reported that tagged scallops came primarily 

from down-current areas on Georges Bank. While 80% 

of the tagged returns had traveled < 3 km and 97% had 

traveled < 16 km, a few individuals were reported to 

have traveled more than 48 km in 2+ years. Melvin et 

al. (1985) reported primarily circular movements of 

tagged scallops on Georges Bank corresponding to the 

clockwise gyre on the Bank (Figure 3), and while 85% 

of the tagged returns had moved < 15 km, several had 

moved > 50 km. 

REPRODUCTION 

The sexes in sea scallops are separate and can be 

distinguished by the color of the gonads (red for 

females, white for males) as they ripen prior to 

spawning. Hermaphroditism is known to occur, but is 

not common (e.g., Merrill and Burch 1960; Worms and 

Davidson 1986). Mature gametes have been observed 

in females as young as one year (Langton et al. 1987) 

and scallops have been reported to spawn during their 

second year (Naidu 1970; MacKenzie 1979). However, 

significant egg production may not occur until age 4. 

Fecundity is directly related to shell height and 

maximum egg production is not reached until several 

years after maturity. It is estimated that female scallops 

produce 1-270 million eggs per individual (Langton et 

al. 1987); by age 4 (85-90 mm) a female will release 

about two million eggs. Gonad output (egg number) is 

greater in scallops from shallow water (10-20 m), 

where the food supply is typically greater and 

temperatures higher than in scallops from deep water 

(170-180 m) (MacDonald and Thompson 1986; 

MacDonald et al. 1987; Barber et al. 1988). There is 

evidence of latitudinal differences in fecundity. 

MacDonald and Thompson (1988) found that scallops 

from New Jersey were more fecund that those from 

locations further north, although variation along a depth 

gradient on a microgeographic scale may be as great or 

greater than variation on a latitudinal scale (MacDonald 

and Thompson 1985a, b; 1988). 

Shumway et al. (1988) summarized the 

gametogenic cycle of sea scallops from Maine as 

follows. During January, gametogenesis has already 

reached the early developmental stage; energy reserves 

are at their lowest level (Robinson et al. 1981), and 

energy must be mobilized from the accumulated 

reserves. During spring, gametogenesis is underway, 
Page 3

gonad size increases, feeding begins with coincidental 

spring phytoplankton blooms, and energy reserves 

begin to accumulate. During summer (June-August), 

food is plentiful, gametes are ripening and energy is 

derived from spring storage and from food intake 

(Robinson et al. 1981). Spawning takes place in 

September/October and the animals enter a 

reproductively quiescent or rest period. Barber et al.

(1988) found that primary oogenesis in sea scallops 

from Boothbay Harbor, Maine, was initiated in 

February, secondary oogenesis in March, and 

vitelogenesis after June. Spawning and reabsorption of 

mature ova was evident in September and to a greater 

extent in October, after which the animals underwent a 

period of recovery (December/January). 

Spawning generally occurs synchronously when 

males extrude sperm and the females release eggs en 

masse into the water, but it may occur over a more 

protracted period of time depending on environmental 

conditions. It has been suggested that year-class 

strength may correlate with the degree of spawning 

synchrony, rather than fecundity per se (Langton et al.

1987). 

A major annual spawning period occurs during 

late summer to fall (August to October) (Parsons et al.

1992a) although spring or early summer spawning can 

also occur, especially in the Mid-Atlantic (Barber et al.

1988; DuPaul et al. 1989; Schmitzer et al. 1991; 

Davidson et al. 1993; Almeida et al. 1994; Dibacco et 

al. 1995). The timing of spawning can vary with 

latitude, starting in summer in southern areas and in fall 

in the northern areas. MacKenzie et al. (1978) reported 

that off the coast of North Carolina and Virginia, 

spawning generally occurred as early as July and that 

further north on the Mid-Atlantic shelf spawning 

occurred in August. However, there are exceptions to 

this pattern. MacDonald and Thompson (1988) report 

that scallops off of New Jersey spawned up to two 

months later than scallops from Newfoundland 

(September-November versus late August-early 

September). They found no clearly identifiable 

latitudinal trends in the timing of spawning. A biannual 

spawning cycle on the Mid-Atlantic shelf has been 

reported south of the Hudson Canyon, with spawning 

occurring both in the spring and fall (DuPaul et al.

1989; Schmitzer et al. 1991; Davidson et al. 1993). 

Kirkley and DuPaul (1991) found that spring spawning 

in the Mid-Atlantic is the more predictable and 

dominant spawning event, while fall spawning is minor, 

temporally irregular, and sometimes does not occur. 

Schmitzer et al. (1991) also reported that the spring 

spawning was of longer duration and the scallops 

showed greater fecundity than in the fall. 

North of the Hudson Canyon there is generally a 

single annual spawning event starting in late summer or 

early fall. However, there are some reports of biannual 

spawning (spring and fall) in the Gulf of Maine and 

Georges Bank, with the fall spawning being dominant 

(Barber et al. 1988; Almeida et al. 1994, DiBacco et al.

1995). On Georges Bank fall spawning generally 

occurs in late September or early October (Posgay and 

Norman 1958; MacKenzie et al. 1978; McGarvey et al.

1992; DiBacco et al. 1995). In Cape Cod Bay, 

spawning occurs in late September and early October 

(Posgay 1950). In the Gulf of Maine spawning occurs 

in August and September (Drew 1906; Welch 1950; 

Baird 1953; Culliney 1974; Robinson et al. 1981; 

Barber et al. 1988). In the Bay of Fundy the spawning 

period extends from late July to November (Stevenson 

1936; Dickie 1955; Beninger 1987; MacDonald and 

Thompson 1988; Dadswell and Parsons 1992). 

Scallops beds generally spawn synchronously in a 

short time, going from completely ripe to completely 

spent in less than a week (Posgay and Norman 1958; 

Posgay 1976). “Dribble spawning” over an extended 

time period has been reported in scallops from 

Newfoundland coastal waters (Naidu 1970) and 

possibly in the Gulf of Maine (Langton et al. 1987) and 

in New Jersey in June and July (MacDonald and 

Thompson 1988). A rapid temperature change, the 

presence in the water of gametes from other scallops, 

agitation, or tides may trigger scallop spawning 

(Parsons et al. 1992a). 

FOOD HABITS 

Sea scallops are suspension filter feeders, using 

currents created by cilia on the gills to move and filter 

water containing suspended particulate material. Their 

diet primarily consists of phytoplankton and 

microzooplankton (such as ciliated protozoa), but 

particles of detritus can also be ingested, especially 

during periods of low phytoplankton concentrations 

(Bordon 1928; Stevenson 1936; Shumway et al. 1987; 

Grant and Cranford 1991). Dissolved organic matter 

(absorbed through the tissues) has been suggested as an 

additional minor source of nutrition, particularly for 

scallop larvae (Marshall and Lee 1991). Palp-pedal 

feeding (using the ciliated end of the foot to bring 

organic matter from biofilms to the labial palps) as well 

as DOM absorption may also be used by postsettlement scallops, during the time that feeding 

structures on the gill develop (Veniot et al. 2003). 

Juvenile scallops can ingest up to 7 mg of food per 

liter filtered, but at very high concentrations of food or 

suspended detritus, the rate of feeding (clearance rate) 

is reduced (Bacon et al. 1998). Current velocities of 

more than 10 cm/sec also reduce feeding activity in 

juveniles (Wildish and Saulnier 1992). In adult 

scallops, feeding is inhibited at current velocities 

exceeding 25 cm/sec or more, and when scallops have 

been feeding heavily (Pilditch and Grant 1999a). 

Elevated concentrations of inorganic suspended 

material and clay-sized particles can interfere with 

feeding, but the presence of low concentrations (< 0.5 
Page 4 

mg/L) of inorganic particulate matter in the diet may be 

important in enabling sea scallops to efficiently utilize 

phytoplankton cells (Cranford and Gordon 1992). Sea 

scallops have the ability to select higher-quality food 

particles for ingestion and create pseudofeces from 

rejected matter (Bacon et al. 1998). 

Dietary components vary depending on geographic 

location. Sea scallops in coastal areas and bays 

encounter more seaweed and seagrass detritus and may 

be exposed periodically to significant amounts of 

resuspended inorganic material. Offshore scallops feed 

mainly on phytoplankton and resuspended organic 

material. Phytoplankton appears necessary to meet 

scallop energetic demands, although seaweed detritus 

may be an important food supplement in nearshore 

environments (Grant and Cranford 1991). 

Shumway et al. (1987) documented seasonal 

changes in the gut contents of sea scallops. Two 

scallop populations were compared, one in shallow 

water (20 m depths) and one in deep water (180 m). A 

total of 27 species of algae, ranging in size from 10-350 

m were identified, plus a number of miscellaneous 

items including pollen grains, ciliates, zooplankton 

tests, and detrital material, and bacteria. Benthic and 

pelagic food species were equally represented in the 

shallow water population; however, benthic species 

outnumbered pelagic species in the deep-water scallops. 

In both populations, seasonal variations in food items 

occurred and coincided with bloom periods of 

individual algal species. The gut contents generally 

reflected the available organisms in the surrounding 

habitat, indicating that sea scallops are opportunistic 

filter feeders which take advantage of both benthic and 

pelagic food. Cranford and Hill (1999) report that 

ingestion rates of sea scallops in Nova Scotia are 

highest in spring and fall; spring and fall are the periods 

of highest and lowest food concentrations, respectively, 

during the year. These periods of maximum ingestion 

are driven by physiological needs of the scallops rather 

than the concentration of food particles in the water. 

GROWTH 

Larval growth rates of 3.1-3.5 m/day (Gallager et 

al. 1996) and 4.3-4.8 m/day (Manuel et al. 1996b) 

have been reported in mesocosm studies. Culliney 

(1974) reported that development to straight-hinge 

veliger took 4 days at 12-18o

C, development to 

pediveliger took 28 days at 15o

C, and settlement 

occurred in 35 days. The duration of the larval stage on 

Georges Bank is 40-60 days (Thouzeau et al. 1991a). 

The growth rate of post-larval (< 2 mm) scallops in 

Passamaquoddy Bay was estimated to be 0.03-0.06 

mm/d, with the growth rate increasing with size 

(Parsons et al. 1993b). A laboratory study of postlarval development at 14o

C found growth rates between 

0.008-0.028 mm/d, depending on diet (Milke et al.

2004). Wildish and Saulnier (1992) reported juvenile 

growth rates of 0.047-0.199 mm/d at 1.2-7.6o

C, and 

adult growth rates of 0.02-0.121 mm/d in the 

laboratory. 

Adult growth rates show considerable variation 

among populations. Some of the highest growth rates 

have been observed on Georges Bank and in Port-auPort Bay (Stewart and Arnold 1994). Scallops from the 

Gulf of St. Lawrence generally have slower growth 

rates than Gulf of Maine and Bay of Fundy scallops 

(Chouinard and Mladenov 1991). Growth generally 

decreases with depth (Schick et al. 1988; Thouzeau et 

al. 1991a). A study by Pilditch and Grant (1999b) 

indicated that food concentration was more important 

than temperature in controlling growth rates for 2-yearold scallops. Estimates of von Bertalanffy growth 

coefficients for Georges Bank sea scallops range from 

141.8-152.5 mm for the mean maximum shell height 

L, and 0.26-0.34/y for the growth coefficient K

(Thouzeau et al. 1991a), while the only published MidAtlantic growth equation has parameters L = 151.8 and 

K = 0.30 (Serchuk et al. 1979). 

PREDATION 

Scallop larvae are planktonic; thus they are 

potentially preyed upon by filter feeders and planktonic 

carnivores (Langton and Robinson 1990). 

Predation on juvenile and adult sea scallops 

decreases with shell height (Stokesbury and 

Himmelman 1995). Teleost predators of scallops 

include Atlantic cod (Medcof and Bourne 1964), 

wolffish (Medcof and Bourne 1964; Nelson and Ross 

1992), eel pouts such as ocean pout (Klein-MacPhee 

and Collette 2002), sculpins (Caddy 1968, 1973), 

American plaice (Medcof and Bourne 1964; Naidu and 

Meron 1986; Robert et al. 1986), winter flounder 

(Caddy 1968, 1973), and yellowtail flounder (Naidu 

and Meron 1986). 

Invertebrate predators include crabs (Cancer spp.) 

and lobsters (Homarus americanus) that crush or chip 

away the shells (Elner and Jamieson 1979; Stokesbury 

and Himmelman 1995; Nadeau and Cliche 1998; Wong 

and Barbeau 2003). Infestation by the boring 

polychaete Polydora websteri, abundant off eastern 

North America, can weaken the shell and increase 

vulnerability to lobsters (Bergman et al. 1982). A 

number of sea star species, including Leptasterias 

polaris, Astropecten americanus, Crossaster papposus, 

Asterias forbesi, and Asterias vulgaris, have been 

observed to consume sea scallops (Medcof and Bourne 

1964; Naidu and Scalpen 1979; Franz and Worley 

1982; Barbeau and Scheibling 1994a, b; Nadeau and 

Cliche 1998; Wong and Barbeau 2003). Medcof and 

Bourne (1964) reported that small juvenile scallops 
Page 5

were ingested whole by Crossaster papposus. 

Astropecten americanus is also holophagous (Franz and 

Worley 1982), and appears to be a predator of juvenile 

sea scallops in the Mid-Atlantic Bight. Naidu and 

Scalpen (1979) reported that juvenile Asterias vulgaris

were the principal predators of scallop spat in spat traps 

in Newfoundland. A textured substrate such as shell or 

pebble significantly reduced the predation rate of 

Asterias vulgaris on 11-15 mm juvenile scallops in an 

experimental setting (Wong and Barbeau 2003). 

GEOGRAPHICAL DISTRIBUTION 

The sea scallop is distributed on the Atlantic 

continental shelf of North America from the Strait of 

Belle Isle, Newfoundland, to Cape Hatteras, North 

Carolina (Posgay 1957). In the United States, sea 

scallops are located on Georges Bank, in the MidAtlantic Bight, and in the Gulf of Maine. 

Sea scallops typically occur at depths ranging from 

18-110 m, but may also occur in waters as shallow as 2 

m in estuaries and embayments along the Maine coast 

and in Canada (Serchuk et al. 1982; Naidu and 

Anderson 1984). In southern areas, scallops are 

primarily found at depths between 45-75 m, and are less 

common in shallower water (25-45 m) due to high 

temperature (Bourne 1965). Although sea scallops are 

not common at depths greater than about 110 m, some 

populations have been found as deep as 384 m (Merrill 

1959), and deep-water populations at 170-180 m have 

been reported in the Gulf of Maine (Barber et al. 1988; 

Schick et al. 1988). 

Sea scallops often occur in aggregations called 

beds. Beds may be sporadic (perhaps lasting for a few 

years) or essentially permanent (e.g., commercial beds 

supporting the Georges Bank fishery) (Figure 4). The 

highest concentration of many permanent beds appears 

to correspond to areas of suitable temperatures, food 

availability, substrate, and where physical 

oceanographic features such as fronts and gyres may 

keep larval stages in the vicinity of the spawning 

population (Thouzeau et al. 1991a, b; Tremblay and 

Sinclair 1992). 

EGGS AND LARVAE 

Eggs and larvae were not enumerated by the 

Northeast Fisheries Science Center (NEFSC) Marine 

Resources Monitoring, Assessment and Prediction 

(MARMAP) program. Until recently sea scallop larvae 

could not even be identified in the wild (Tremblay et al.

1987). 

However, the NOAA Estuarine Living Marine 

Resources (ELMR) program has identified the presence 

of eggs and larvae in a number of New England bays 

and estuaries (Table 1; Jury et al. 1994). 

JUVENILES AND ADULTS 

Summer NEFSC sea scallop surveys sample from 

Georges Bank to Cape Hatteras (Figure 5 and Figure 6, 

NEFSC 2004). The Gulf of Maine is not routinely 

surveyed. In the Mid-Atlantic Bight, from Long Island 

to Cape Hatteras, scallops occur mostly between 40 and 

70 m, with the densest concentrations near Hudson 

Canyon and off of Delaware Bay. Further north, the 

highest concentrations occur in the Great South 

Channel and in the eastern portion of Georges Bank, 

areas that traditionally have provided large harvests 

(MacKenzie et al. 1978; Serchuk et al. 1979; Posgay 

1981). 

In more recent years, scallop density has increased 

markedly in the three groundfish closed areas that have 

been closed to scallop fishing for most of the time since 

December, 1994 (Figure 6; Murawski et al. 2000, 

NEFSC 2004), and in the Mid-Atlantic Bight, 

especially off of New Jersey, Delaware, and Maryland. 

The increases in the Mid-Atlantic Bight are due to a 

combination of increased recruitment, reduced fishing 

mortality, and the rotational closure of an area south of 

Hudson Canyon for a three year (1998-2001) period 

(NEFSC 2004). 

The NOAA ELMR program has identified the 

presence of juvenile and adult sea scallops in a number 

of New England bays and estuaries (Table 1; Jury et al.

1994). 

In the Gulf of Maine, non-stratified surveys and 

data from commercial catches indicate scallop beds 

occur in inshore areas from Penobscot Bay and 

eastward, as well as in Cape Cod Bay. Offshore beds 

occur on Jeffreys Basin, Cashes Ledge, Fippinnies 

Ledge, Jeffreys Ledge, near Three-Dory Ridge, 

Toothaker Ridge, off Jonesport and Machias bays, and 

on Stellwagen Bank (Serchuk and Rak 1983; Serchuk 

and Wigley 1984). The largest offshore catches in the 

Gulf of Maine have traditionally occurred on Jeffreys 

Ledge and Fippinnies Ledge (Serchuk and Rak 1983; 

Serchuk and Wigley 1984). 

HABITAT CHARACTERISTICS 

The habitat characteristics of sea scallops are 

summarized in Table 2. 
Page 6 

LARVAE 

In laboratory studies, larvae were viable at 

temperatures of 12-18oC (mass mortalities occurred at 

higher temperatures), and at salinities as low as 10.5 

ppt, although salinities ranging from 16.9-30 ppt were 

preferred (Culliney 1974). In nature, larvae initially 

settle on gravelly sand or shell fragments (Langton and 

Robinson 1990), or filamentous animals and plants, 

many of which colonize the backs of adult scallops 

(Stokesbury and Himmelman 1995). Settling larvae 

often actively select substrates covered with a biofilm 

(Parsons et al. 1993a). Prior to settlement, larvae 

circulate with residual currents ranging from 6-25 cm/s 

and tend to concentrate in the upper 10 m of the water 

column (Tremblay and Sinclair 1990a; Parsons et al.

1992b). 

JUVENILES 

In nature juvenile scallops are mainly found on 

gravel, small rocks, shells and silt (Thouzeau et al.

1991a; Parsons et al. 1992b), or attached to branching 

bryozoans, hydroids or algae (Stokesbury and 

Himmelman 1995). Gravel bottoms have higher 

juvenile and adult scallop abundances than other 

substrates (Thouzeau et al. 1991a; Kostylev et al.

2003). In laboratory studies, juveniles have been 

maintained successfully at 1.2-15o

C (Manuel and 

Dadswell 1991; Wildish and Saulnier 1992; Barbeau 

and Scheibling 1994b; Kleinman et al. 1996). Frenette 

and Parsons (2001) determined the best conditions for 

survival for juveniles < 35 mm were temperatures 

below 18o

C and salinities > 25 ppt. The optimal current 

velocity for feeding and growth is 10 cm/s; currents 

stronger than 10 cm/s inhibit feeding (Wildish and 

Saulnier 1992). Stronger currents may be experienced 

in nature (e.g., up to 25 cm/s in Passamaquoddy Bay), 

but can be avoided by feeding during periods of slack 

water (i.e., low and high tide). Maximum densities of 

juveniles on Georges Bank have been found at depths 

of 62-91 m (Thouzeau et al. 1991a). 

ADULTS 

Adult sea scallops are generally found on firm 

sand, gravel, shells and rock (MacKenzie et al. 1978; 

Langton and Robinson 1990; Thouzeau et al. 1991a; 

Stewart and Arnold 1994). Other invertebrates 

associated with scallop beds include sponges, hydroids, 

anemones, bryozoans, polychaetes, mussels, moon 

snails, whelks, amphipods, crabs, lobsters, sea stars, sea 

cucumbers, and tunicates (Kenchington 2000). 

Adult scallops experience optimal growth at 

temperatures of 10-15o

C; temperatures above 21oC are 

lethal (Stewart and Arnold 1994); spawning occurs at 

9.0-11.2o

C (MacKenzie et al. 1978). Adults prefer full 

strength seawater, and salinities of 16.5 ppt or lower are 

lethal (Stewart and Arnold 1994). Scallops require 

some water movement (optimal growth occurs at 10 

cm/s) for feeding, oxygen uptake, and removal of waste 

(Wildish and Saulnier 1992; Stewart and Arnold 1994). 

ACKNOWLEDGEMENTS 

We would like to thank the authors of the first 

edition of this report, David B. Packer, Luca M. 

Cargnelli, Sara J. Griesbach, and Sandra E. Shumway, 

for providing the foundation of this document. This 

document also benefited from comments from Fred 

Serchuk. 

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Page 12

Table 1. Relative abundance of egg, larval, juvenile, and adult sea scallop, Placopecten magellanicus, in New England 

bays and estuaries by salinity zone. 

Based on Estuarine Living Marine Resources (ELMR) data in Jury et al. (1994). 

Eggs Larvae Juveniles Adults 

T M S T M S T M S T M S

Passamaquoddy Bay R A R A R A R A 

Englishman/Machias Bays A R A R A R A 

Narraguagus Bay A R A R A R A 

Blue Hill Bay A R A R A R A 

Penobscot Bay R A R A R A R A 

Muscongus Bay C C C C 

Damariscotta River C C C C 

Sheepscot River C C C C 

Kennebec/Androscoggin 

Rivers 

 R R R R 

Casco Bay C C C C 

Saco Bay R R R R 

Wells Harbor 

Great Bay R C C 

Merrimack River 

Massachusetts Bay C C C C 

Boston Harbor na na R R 

Cape Cod Bay A A A A 

Salinity zone: T = tidal fresh, M = mixing zone, S = seawater. 

Relative abundance: H = highly abundant, A = abundant, C = common, R = rare, blank = not present, na = no data 

available. 
Page 13

Table 2. Summary of sea scallop habitat parameters based on the pertinent literature. 

Life Stage Size and Growth Habitat Substrate 

Eggs 1 Average diameter of spawned eggs: 

66.8+1.6 μm. 

Average wet weight of spawned eggs: 

1.6 x 10-7 g. 

Fertilized eggs are slightly heavier than 

seawater and probably remain on the sea floor 

as they develop into the first free-swimming 

larval stages. 

Larvae 2 Larval length 0.06-0.3 mm. 

Larvae grow through 4 larval stages in 4-6 

weeks (on Georges Bank), then settle to the 

bottom (assumed to occur by midDecember); maximum larval settlement in 

Passamaquoddy Bay occurs in late 

September. 

Development times: 4 days to straight 

hinge veliger at 12-18o

C; 28 days to 

pediveliger at 15o

C, 40 days at 12-16oC; 35 

days to settlement. Pediveligers delayed 

metamorphosis until suitable substrates 

were found. 

Growth rates: 3.1-3.5 μm/day (Trinity Bay, 

Newfoundland), 4.3-4.8 μm/d 

(Passamaquoddy Bay and Georges Bank). 

Fall density on Georges Bank ranged 120-

1500/m2

. 

Vertical distribution: in mixed areas, larvae 

distributed evenly through water column; in 

stratified areas, larvae aggregated above 

pycnocline. Larvae capable of detecting and 

using thermocline. 

Vertical migration: in response to tidal, solar 

cues; 

Passamaquoddy Bay veligers generally found 

shallower than Georges Bank veligers. 

Larvae settle in areas of gravelly sand, shell 

fragments or on hydroids, bryozoans and 

sponges; select substrates covered with a 

biofilm. 

In laboratory studies, larvae settle on 

undersides of pebbles, shell, and glass 

fragments. Artificial substrates have been 

successful as settlement substrate. 

Juveniles 3 Size ~1-80 mm shell height. 

Average shell height at age: 1-14.4 mm, 2-

44.5 mm (50.8 at 66-70 m deep, 33.8 mm 

at 81-85 m). 

Average shell height decreases with 

increasing depth. 

Growth of Bay of Fundy scallops peaks in 

July and is best predicted by temperature. 

Growth rate in one study: 0.047-0.199 

mm/day shell height. 

Maximum density of 1 and 2 year old scallops 

(0.88-1.65 juveniles/m2

) found on northeast 

Georges Bank at depths of 62-91 m; age 1 

were less dispersed than age 2 and mainly 

located on a gravel-pebble deposit in the 

northern half of the Bank. 

In a Passamaquoddy Bay study, 76% of the 

scallops moved an average of 3.3 m in 4 

months. 

Mainly found on gravel, small rocks, shells, 

and among branching animals and plants 

that permit attachment of juveniles. 

Adults 4 Size: approximately > 80 mm, up to 170 

mm shell height. 

Average shell height at age on Georges 

Bank: 3-72.0 mm, 4-89.5 mm, 5-101.7 mm, 

6-112.0 mm, 7-121.0 mm. 

Growth rates of Bay of Fundy scallops in 

lab: 0.020-0.121 mm/d shell height. 

Wide distribution on offshore banks and 

coastal waters from Newfoundland to Cape 

Hatteras; from low tide level to ~100 m line; 

generally shallower in northern populations. 

On Georges Bank, 4 spatially separated subpopulations (Northern Edge and Northeast 

Peak, Southeast Part, Great South Channel 

east, Great South Channel west) are linked by 

larval transport; largest concentrations on 

Northern Edge and Northeast Peak at 37-100 

m. In Maine, mostly in shallow water (< 20m), 

but also on Fippenies Ledge and Jeffreys 

Ledge (56-84 m), and some also observed in 

very deep water (~175m). 

In Mid-Atlantic Bight, largest concentrations 

from Hudson Canyon south to off of Delaware 

Bay; collected at depths of 27-80 m (average 

55 m). 

Generally found in seabed areas with firm 

sand, gravel, shells and cobble substrate. 

Typically abundant in areas with low levels 

of inorganic suspended particulates (fine 

clay size particles). 

Spawning 

Adults 5

Some scallops mature as early as 30 mm, 

but most mature at 90 mm (in 

Passamaquoddy Bay). 

Scallops 50-90 mm (shell height) spawn 

first, followed by those > 90 mm (on 

Georges Bank). 

Reproductive effort increases with age and 

size. 

See Adults. See Adults. 

1

 Langton et al. (1987). 2

 Culliney (1974); Beninger (1987); Langton and Robinson (1990); Tremblay and Sinclair (1990a, b, 1992); Thouzeau et al. (1991a); McGarvey et al.

(1992); Parsons et al. (1993b); Stokesbury and Himmelman (1995); Gallager et al. (1996); Manuel et al. (1996a, b, 1997); Robinson et al. (1996); 

Manuel et al. (2000). 3

 Thouzeau et al. (1991a); Parsons et al. (1992b); Wildish and Saulnier (1992); Stewart and Arnold (1994); Stokesbury and Himmelman (1995); 

Kleinman et al. (1996). 4

 Mackenzie et al. (1978); Langton and Robinson (1990); Thouzeau et al. (1991a); McGarvey et al. (1992, 1993); Wildish and Saulnier (1992); Stewart 

and Arnold (1994); Barber et al. (1988); NEFSC (2004). 5

 Posgay and Norman (1958); Mackenzie et al. (1978); MacDonald and Thompson (1988); Kirkley and DuPaul (1991); Schmitzer et al. (1991); 

McGarvey et al. (1992); Parsons et al. (1992a). 
Page 14

Table 2. Cont’d. 

Life Stage Temperature Salinity Currents Prey 

Eggs 1 Eggs maintained in the laboratory 

at 13-17o

C hatched 32 days post 

fertilization. 

Larvae 2 Laboratory reared larvae from 

New Hampshire were viable at 12-

18o

C; mass mortalities when 

reared at 19o

C. 

Duration of larval stages shorter at 

higher temperatures. 

One study shows that larval 

abundance on Georges Bank peaks 

when thermal stratification is high. 

Laboratory reared 

larvae from New 

Hampshire viable at 

salinities as low as 10.5 

ppt (at 15o

C for 42 h), 

although behavior was 

abnormal (lack of 

swimming). Swimming 

and other behaviors 

appeared normal within 

16.9-30 ppt. 

Larvae circulate with residual currents 

ranging from 6-25 cm/s. 

In Bay of Fundy there is some transport 

of larvae via residual currents, but most 

larvae remain near area of major 

spawning. 

Larval exchange influenced by duration 

and depth of planktonic drift, gyre 

strength, and weak cross-isobath flow. 

Differences in vertical distribution may 

be related to differences in horizontal 

transport by currents. 

Feed on phytoplankton, and 

microscopic animals; also 

dissolved organic matter, 

detrital particles and bacteria. 

Juveniles 3 Juveniles successfully maintained 

at temperatures of 1.2-15oC in 

laboratory. High exfoliation of 

epithelial cells at 21o

C; damage to 

gills, mantles, and gonads at 

exposure to 14.7o

C followed by 

21o

C. Optimum survival at 

temperatures less than 18o

C. 

Consumption rate by predators, 

rate of scallop movement, and 

growth rate increases with 

temperature. 

For optimum survival, 

salinities > 25 ppt. 

Optimum velocity for growth is 10 cm/s; 

> 10 cm/s can result in reduced feeding 

rate; but this can be avoided in areas with 

strong currents by feeding during periods 

of slack water (high and low tide). 

Juveniles from Georges Bank drift with 

currents to the south as they grow older. 

Byssal attachment by scallops < 12 mm 

shell height may be an adaptation to 

strong currents. 

Feed on suspended particles, 

primarily phytoplankton. Newly 

settled juveniles may also feed 

on biofilms. 

Adults 4 Optimal growth at 10-15o

C; lethal 

temperature: > 21o

C. 

Optimal survival at 

salinities approaching 

full-strength seawater; 

lower lethal threshold: 

16.5 ppt. 

Water movement critical for dispersal, 

replenishment of suspended food 

particles, supply of oxygen and removal 

of waste products. 

Adults drift with tidal currents (on 

Georges Bank, 39% moved toward the 

southeast, 30% to the Northwest). A 

1989 study showed southward drift of 

adults. 

Strong tidal currents (> 25 cm/s) inhibit 

feeding; fluctuations in scallop clearance 

rates are related to flow velocity. 

Filter feeders on phytoplankton 

(mostly diatoms) and other 

suspended organic particulates 

from the water; gut contents 

reflect available organisms in 

the immediate environment. 

Stomach contents of scallops 

(90-140 mm shell height) from 

shallow and deep study sites in 

Maine: 27 species of algae 

ranging in size from 10-350 μm, 

pollen grains, ciliates, 

zooplankton tests, detrital 

material, and bacteria. 

Spawning 

Adults 5

Spawning in nature occurs at 

temperatures ranging from 6.5-

16o

C. Georges Bank: typically 9-

11.2o

C. Isle of Shoals, New 

Hampshire: 14-16o

C; spawning 

triggered in lab when temperature 

was raised abruptly from 5 to 

10o

C, or from 12 to 15oC. 

Mid-Atlantic Bight: spawning 

coincided with decreasing and 

stable temperatures in 

winter/spring and increasing 

temperatures in fall; 6.5o

C off 

Long Island; 11o

C off 

Chincoteague, Virginia. Spawning 

may be delayed by low summer 

temperatures. 

 Tidally related spawning cue in 

Passamaquoddy Bay; spawning occurs 

just prior to spring tides. 

 

1

 Karney (1996). 

2

 Culliney (1974); Tremblay and Sinclair (1986, 1990a, b, 1992); Thouzeau et al. (1991a); Tremblay et al. (1994); Manuel et al. (1996a, 1997). 3

 Shumway et al. (1987); Manuel and Dadswell (1991); Thouzeau et al. (1991a); Wildish and Saulnier (1992); Manuel and Dadswell (1993); Barbeau 

and Scheibling (1994b); Stewart and Arnold (1994); Kleinman et al. (1996); Potter et al. (1997); Frenette and Parsons (2001). 4

 Posgay (1981); Shumway et al. (1987); Thouzeau et al. (1991a); Wildish and Saulnier (1992); Stewart and Arnold (1994); Cranford et al. (1998); 

Pilditch and Grant (1999a). 

5

 Culliney (1974); MacKenzie et al. (1978); Schmitzer et al. (1991); Parsons et al. (1992a). 
Page 15

Table 2. Cont’d. 

Life Stage Predators Spawning Notes 

Eggs 1 Predation by a variety of bottom 

predators. 

See spawning adults Fecundity ranges from 1-270 million eggs/ 

individual. Egg production is positively 

correlated with age and shell height 

(exponentially). 

Larvae 2 Scallop larvae potentially 

preyed on by filter feeders and 

larval and adult planktivores. 

Early larval stages (trochophore and veliger) 

are pelagic, swim freely. 

Passamaquoddy Bay larvae reach the veliger 

stage faster than larvae on Georges Bank; a 

shortened trochophore stage may allow earlier 

vertical migration and control of horizontal 

movement (strong tides in this area make 

larvae vulnerable to being advected away). 

No large scale exchange between Georges 

Bank and the Scotian Shelf; self-seeding 

likely for the Great South Channel and 

Northeast Peak; simulations indicate 

significant larval exchange between Northeast

Peak, Southern Flank, and Great South 

Channel. 

Juveniles 3 Predators include Atlantic cod, 

American plaice, yellowtail and 

winter flounder, ocean pout, 

Atlantic wolffish, crabs, 

lobsters, and sea stars. 

The distribution of juveniles is determined by 

the hydrodynamic regime during the larval 

stage and by differential post-larval mortality 

at settlement. 

In laboratory studies, growth rate and 

metabolic condition increased with 

temperature; swimming frequency did not 

affect growth, but increased mass of adductor 

muscle. Somatic production (i.e., growth) 

increases to age 5 then levels off. 

Adults 4 Predators of adults include 

Atlantic wolffish, lobsters, and 

sea stars. Adults damaged by 

dredging and trawling activity 

have been shown to fall prey to 

crabs, starfish, lobster, and 

groundfish (winter flounder and 

sculpins). 

See spawning adults Sea scallop populations on Georges Bank 

have traditionally been the largest, densest, 

and most intensively exploited of the species, 

though very high densities and catches have 

occurred recently in the Mid-Atlantic Bight. 

In a 1989 study, the Northern Edge (2.91 

g/m2) and Northeast Peak (1.32 g/m2) have 

the highest biomass (meat weight) density 

(other areas on Georges Bank much lower, 

0.29-0.44). By area, the Northeast Peak has 

the highest biomass (20599 mt) followed by 

the Northern Edge (4291 mt). 

Sex ratios: Georges Bank, 1:1; Mid-Atlantic 

Bight, 1.4:1 (male: female, 1975 studies). 

Spawning 

Adults 5

 Georges Bank: spawning in late Sept.-early Oct; 

reports of a semi-annual reproductive cycle on 

northeast Georges Bank: spring spawning in May-June 

and fall spawning in Sept.-Oct.; fall is larger, more 

synchronized and consistent among years, spring is 

more protracted, erratic. 

Maine: in shallow sites, spawning in Sept.-Oct.; deep 

water sites, possible minor spawning in May, and more 

abrupt spawning in Oct. (1 month later than in shallow 

sites). 

Passamaquoddy Bay: relatively consistent 

reproductive cycle with major spawning in late Julylate Sept. (peaked Aug.); similar to spawning period in 

southeast Newfoundland and areas of Gulf of Maine. 

New Jersey: mostly Sept.-Nov., some in June-July. 

Mid-Atlantic Bight: Semi-annual gametogenic cycle; 

spring (May/June) and fall (Nov.) spawning; spring 

spawning stronger, more predictable than fall (longer 

duration and greater fecundity in spring). 

Gulf of Maine: deep-water scallops produced 

fewer eggs than shallow-water scallops; 

lowered fecundity reduces the probability that 

the deep-water population is self-sustaining. 

Mid-Atlantic Bight: the significance of the 

spring spawning event to recruitment 

processes uncertain. In spring, deep water 

scallops initiated gamete production about 1 

month later than shallow water scallops. 

Reproduction and size attained by individuals 

are largely controlled by local environmental 

conditions. 

Fecundity and production greater, but 

longevity reduced in New Jersey scallops 

(avg. longevity 1-10 yrs), than in those from 

New Brunswick and Newfoundland (avg. 1-

12 yrs). 

1

 Langton et al. (1987); Stewart and Arnold (1994). 2

 Tremblay and Sinclair (1986, 1992); Langton and Robinson (1990); Tremblay et al. (1994); Manuel et al. (1996a). 3

 Elner and Jamieson (1979); Langton et al. (1987); Thouzeau et al. (1991a); Barbeau and Scheibling (1994a, b); Kleinman et al. (1996). 4

 MacKenzie et al. (1978); McGarvey et al. (1992, 1993); Stewart and Arnold (1994); NEFSC (2004). 5

 Posgay and Norman (1958); Culliney (1974); MacKenzie et al. (1978); Beninger (1987); Barber et al. (1988); MacDonald and Thompson (1988); 

DuPaul et al. (1989); Kirkley and DuPaul (1991); Schmitzer et al. (1991); Dadswell and Parsons (1992); McGarvey et al. (1992); DiBacco et al.

(1995). 
Page 16

Figure 1. The sea scallop, Placopecten magellanicus (photograph by Dann Blackwood, USGS). 
Page 17

Figure 2. Generalized life cycle of the sea scallop, from Stewart and Arnold (1994). 
Page 18

Figure 3. Map of Georges Bank indicating the five sub-regions and the clockwise flow of the residual current. 

From McGarvey et al. (1993). 
Page 19

Figure 4. Distribution of sea scallop spawning beds off the northeast coast of North America. 

From Shumway et al., in prep. 
Page 20

Figure 5. Distribution and abundance of sea scallops collected during NEFSC scallop surveys during summer 1979-

1993. 
Page 21

Figure 6. Distribution and abundance of sea scallops collected during NEFSC scallop surveys during summer 1994-

2003. 

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