BiolOgiCal Report 82(11.87) TR EL-82-4 

January 1989 

Species Profiles: Life Histories and 

Environmental Requirements of Coastal Fishes 

and Invertebrates (North Atlantic) 

WINTER FLOUNDER 

7 

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n-‘” ---‘---- - __..__ 

-_ ._ .*.-~-.‘-_c’~ .__ 

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:* 

Fish and Wildlife Service 

U.S. Department of the Interior 

Coastal Ecology Group 

Waterways Experiment Station 

U.S. Army Corps of Engineers 
a 
Biological Report 82(11.87) 

TR EL-82-4 

January 1989 

Species Profiles: Life Histories and Environmental Requirements of Coastal 

Fishes and Invertebrates (North Atlantic) 

WINTER FLOUNDER 

Jack Buckley 

Massachusetts Cooperative Fishery Research Unit 

Department of Forestry and Wildlife Management 

University of Massachusetts 

Amherst, MA 01003 

Project Officer 

David Moran 

U.S. Fish and Wildlife Service 

National Wetlands Research Center 

1010 Gause Boulevard 

Slidell, LA 70458 

Performed for 

Coastal Ecology Group 

Waterways Experiment Station 

U.S. Army Corps of Engineers 

Vicksburg, MS 39180 

and 

U.S. Department of the Interior 

Fish and Wildlife Service 

Research and Development 

National Wetlands Research Center 

Washington, DC 20240 
This series may be referenced as follows: 

U.S. Fish and Wildlife Service. 1983-19 . Species profiles: life histories 

and environmental requirements of coasts fishes and invertebrates. U.S. Fish 

Wildl. Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers, TR EL-82-4. 

This profile may be cited as follows: 

Buckley, J. 1989. Species profiles: life histories and environmental requirements of coastal fishes and invertebrates (North Atlantic)--winter flounder. 

U.S. Fish Wildl. Serv. Biol. Rep. 82(11.87). U.S. Army Corps of Engineers, TR 

EL-82-4. 12 pp. 
PREFACE 

This species profile is one of a series on coastal aquatic organisms, 

principally fish, of sport, commercial, or ecological importance. The profiles 

are designed to provide coastal managers, engineers, and biologists with a brief 

comprehensive sketch of the biological characteristics and environmental 

requirements of the species and to describe how populations of the species may be 

expected to react to environmental changes caused by coastal development. Each 

profile has sections on taxonomy, life history ecological role, environmental 

requirements, and economic importance, if applicable. A three-ring binder is 

used for this series so that new profiles can be added as they are prepared. 

This project is jointly planned and financed by the U.S. Army Corps of Engineers 

and the U.S. Fish and Wildlife Service. 

Suggestions or questions regarding this report should be directed to one of 

the following addresses. 

Information Transfer Specialist 

National Wetlands Research Center 

U.S. Fish and Wildlife Service 

NASA-Slide11 Computer Complex 

1010 Gause Boulevard 

Slidell, LA 70458 

or 

U.S. Army Engineer Waterways Experiment Station 

Attention: WESER-C 

Post Office Box 631 

Vicksburg, MS 39180 

iii 
CONVERSION TABLE 

Multiply EY 

millimeters (mm) 0.03937 

centimeters (cm) 0.3937 

meters (m) 3.281 

meters (m) 0.5468 

kilometers (km) 0.6214 

kilometers (km) 0.5396 

To Obtain 

inches 

inches 

feet 

fathoms 

statute miles 

nautical miles 

square meters (m*) 10.76 square feet 

square kilometers (km2) 0.3861 square miles 

hectares (ha) 2.471 acres 

liters (1) 0.2642 gallons 

cubic meters (m3) 35.31 cubic feet 

cubic meters (m3) 0.0008110 acre-feet 

milligrams (mg) 0.00003527 ounces 

grams (9) 0.03527 ounces 

kilograms (kg) 2.205 pounds 

metric tons (t) 2205.0 pounds 

metric tons (t) 1.102 short tons 

kilocalories (kcal) 3.968 British thermal units 

Celsius degrees ("C) 1.8('(Z) + 32 Fahrenheit degrees 

inches 

inches 

feet (ft) 

fathoms 

statute miles (mi) 

nautical miles (nmi) 

U.S. Customary to Metric 

25.40 

2.54 

0.3048 

1.829 

1.609 

1.852 

millimeters 

centimeters 

meters 

meters 

kilometers 

kilometers 

square feet (ft2) 0.0929 square meters 

square miles (mi*) 2.590 square kilometers 

acres 0.4047 hectares 

gallons (gal) 3.785 liters 

cubic feet (ft3) 0.02831 cubic meters 

acre-feet 1233.0 cubic meters 

ounces (oz) 28350.0 milligrams 

ounces (oz) 28.35 grams 

pounds (lb) 0.4536 kilograms 

pounds (lb) 0.00045 metric tons 

short tons (ton) 0.9072 metric tons 

Metric to U.S. Customary 

British thermal units (Btu) 0.2520 kilocalories 

Fahrenheit degrees (OF) 0.5556 ("F - 32) Celsius degrees 

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ACKNOWLEDGMENTS 

I am grateful for reviews by Wendy Gabriel, National Marine Fisheries 

Service, Woods Hole, Massachusetts, and Arnold Howe, Massachusetts Division of 

Marine Fisheries, Sandwich, Massachusetts. 

W’ 

vi 
Figure 1. Winter flounder. 

WINTER FLOUNDER 

NOMENCLATURE/TAXONOMY/RANGE MORPHOLOGY/IDENTIFICATION AIDS 

Scientific name . . . . Pse;dopleuronectes americanus (Walbaum 

Preferred common name . . . . Winter 

flounder (Figure 1) 

Other common names . . . . . Blackback 

flounder, lemon sole, black flounder 

Class . . . . . . . . Osteichthyes 

Order . . . . . . , Pieuronectiformes 

Family . . . . . . . . Pleuronectidae 

Geographic range: Winter flounder are 

found primarily in estuarine and 

coastal waters along the Atlantic 

coast of North America from 

Newfoundland to Georgia (Leim and 

Scott 1966), except for off-shore 

populations on Georges Bank and 

Nantucket Shoal (Figure 2; Bigelow 

and Schroeder 1953). 

The winter flounder, one of the 

right-eyed flounders, is oval-shaped 

and thick-bodied; the caudal fin and 

peduncle are broader than those of 

other North Atlantic flounders. The 

anal fin is highest at its midpoint 

and is preceded by a short sharp 

spine. The dorsal fin (60-76 rays) 

originates opposite the anterior edge 

of the eye, and is about equal in 

height along its length. The mouth is 

small, not gaping to the eye. The 

left (under) half of the jaw is armed 

with a series of close-set incisors; 

the right (upper) half has only a few 

teeth. 

The winter flounder, like other 

flatfishes, varies in color, depending 

1 
I 

I 

I 

I 

I 

I 

I 

I 

I 

I 

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i CANADA \ 

; PORTLAND 

ATLANTIC OCEAN 

Coastat distribution 

KILOMETERS 

GEORGES BANK 

Figure 2. Distribution of winter flounder in the North Atlantic. 

2 
largely on the color of the surrounding substrate. Most adults tend to 

be reddish brown, olive-green, or 

blackish. Smaller fish generally 

are paler than larger fish. The 

blind side is white and, toward the 

edge, translucent or occasionally 

yellowish. 

SEPARATION FROM OTHER 

RIGHT-EYED FLATFISHES 

Compared with the yellowtail flounder, Limanda ferruginea, the winter 

flounder has a much straighter lateral 

line, a less concave dorsal head profile, and fewer fin rays. 

The winter flounder lacks the mucous 

pits that are conspicuous on the left 

(blind) side of the head of the witch 

flounder (Glyptocephalus cynoglossus); 

it also has three times as many dorsal 

rays as the witch flounder. - 

The scales between the eyes are 

smooth in the smooth flounder (Liopsetta putnami), but rough in the wlnter flounder. Between the two, the 

winter flounder also has the greater 

number of anal fin rays. 

Several morphological characteristics that distinguish larvae of winter 

flounder from those of the other 

flounders common in the western north 

Atlantic were given by La Roche 

(1980). 

REASONS FOR INCLUSION IN THE SERIES 

By virtue of its abundance in estuarine and nearshore waters, the winter 

flounder is one of the most important 

commercial and sport fishes in the 

Northeastern United States. In 

Massachusetts, it is considered a 

major contributor to the commercial 

and sport fisheries (Pierce and Howe 

1977). 

LIFE HISTORY 

Spawning 

The winter flounder spawns in coastal waters as early as December in the 

Southern United States and as late as 

June in Canada. Typically, eggs are 

deposited over a sandy substrate at 

depths of 2 to 80 m (Bigelow and 

Schroeder 1953). 

Most spawning takes place at salinities of 31 to 32.5 ppt in inshore 

waters, and on Nantucket Shoal and 

Georges Bank at slightly higher salinities (32.7 to 33 ppt, respectively; 

Bigelow and Schroeder 1953). Water 

temperature during spawning is usually 

between 0 and 3 OC but may be as high 

as 6 OC (Bigelow and Schroeder 1953). 

The winter flounder spawns at slightly 

higher temperatures on Georges Bank 

than in inshore waters (Lux et al. 

1970). 

The stage of maturity of the winter 

flounder is largely governed by size 

rather than age. Flounders grow 

faster and mature at a younger age in 

the south than in the north. In Newfoundland, males mature at age VI and 

females at age VII (Kennedy and Steele 

1971); in New York, winter flounder 

mature at age II or III (Perlmutter 

1947). 

The fecundity of winter flounder 

reported by Bigelow and Schroeder 

(1953) ranged from 0.5 to 1.5 million 

eggs per female. Saila (1961) 

reported that in Rhode Island waters 

193,000 eggs were produced by a fish 

249 mm total length (TL) and 1.34 million by a fish 428 mm TL. In the 

Weweantic Estuary in Massachusetts, 

numbers of eggs ranged from 435,000 

for a fish 350 mm TL to 3.3 million 

for a fish 450 mm TL (Topp 1967). 

In Newfoundland, Kennedy and Steele 

(1971) reported a fecundity range 

from 99,000 eggs for a fish 220 mm 

TL to 2.6 million for a fish 440 mm 

TL (mean = 0.59 million eggs at 

a mean length of 340 mm TL). The 

3 
following equations for estimating 

fecundity on the basis of weight have 

been published: 

log F = 2.3894 + 1.2403 log W 

(Kennedy and Steele 1971) 

pigment cells dividing the postanal 

portion of the body. At a water temperature of 3.9 C the larvae were 

about 5 mm TL, and the yolk sac was 

absorbed in 12 to 14 days. La Roche 

(1980) provides a detailed description 

of larval development. 

log F = 0.0697 + 1.0659 log W 

(Topp 1967) 

log F = 2.6712 + 1.1383 log W 

(Saila 1961) 

where F = fecundity in thousands of 

eggs and W = total weight in grams. 

Winter flounder undergo a rapid 

metamorphosis at a much smaller size 

than other flatfishes of the North 

Atlantic region (Bigelow and Schroeder 

1953). Their metamorphosis is complete when the larvae are 8 to 9 mm TL 

(Laurence 1975); this transformation 

took 80 days at a water temperature of 

5 OC and 49 days at 8 OC. No metamorphosis was evident at 2 OC (Bigelow 

and Schroeder 1953). 

Winter flounder eggs are demersal, 

adhesive, and 0.74 to 0.85 mm in diameter (Bigelow and Schroeder 1953). 

They have no oil globule when deposited, but acquire one later (Breder 

1924). Incubation time was 15 to 18 

days at 2.8 to 3.3 OC (Bigelow and 

Schroeder 1953), 25 days at 3 OC, and 

7 days at 12 to 14 OC (Rogers 1976). 

Incubation time was inversely related 

to water temperature and salinity 

(Rogers 1976). 

In aquaria, winter flounder larvae 

engage in upward swimming bouts and 

then sink to the bottom where they 

remain for a short time (Sullivan 

1915; Bigelow and Schroeder 1953). 

The larvae of other flatfish species 

are more pelagic. Winter flounder 

larvae are continuous, visual, daylight feeders that cease feeding at 

night (Laurence 1977). 

Juveniles 

Winter flounder eggs seem to be most 

abundant in water with a salinity of 

10 to 30 ppt; at salinities below 5 

ppt or above 40 ppt, some embryos survive, but are usually deformed (Rogers 

1976). The optimal salinity for egg 

survival is 15 to 35 ppt. 

Many embryos become inviable or abnormal at temperatures below freezing 

(-1.8 to 0 "C) and temperatures above 

10 OC (Williams 1975). The optimum 

water temperature range for survival 

is 0 to 10 OC (Williams 1975). 

After metamorphosis, winter flounder 

are benthic and seldom lose contact 

with the substrate. Most juveniles 

spend much of their first 2 years in 

or near shallow natal waters, where 

they move in response to extreme heat 

or cold (Topp 1967). After metamorphosis, the juveniles prefer a substrate of sand or sand and silt 

(Clayton et al. 1978). Older juveniles in estuaries gradually move seaward as they grow larger (Mulkana 

1966). 

Larvae Adults 

In studies by Bigelow and Schroeder The seasonal movements of winter 

(1953) and La Roche (1980), winter flounder differ between populations 

flounder larvae were 2.4 to 3.5 mm TL north and south of Cape Cod. A 5-year 

at the time of hatching. A major tagging study by Howe and Coates 

characteristic of the newly hatched (1975) showed that winter flounder 

larvae was the broad vertical band of north of Cape Cod moved about only 

4 
locally in inshore waters, while those 

south of Cape Cod dispersed more than 

3 mi offshore in a southwesterly 

direction. Adults from Martha's Vineyard and coastal populations from 

south of Cape Cod mixed in Nantucket 

Sound (Pierce and Howe 1977). 

Water temperature seems to be the 

most important environmental factor 

determining seasonal distribution 

(McCracken 1963). In Rhode Island, 

adult winter flounder lived in cooler 

offshore waters during summer and in 

shallow inshore waters in winter and 

early spring (Saila 1961). In Newfoundland, winter flounder remained in 

shallow water during summer as long as 

food was available and water temperatures did not exceed 15 'C (Van 

Guelpen and Davis 1979). Temperature 

is a less important factor in the distribution of'juveniles, which tolerate 

higher temperatures than adults 

(Pearcy 1962). 

Indications are that a local population is defined by fish inhabiting 

several adjacent estuaries (Pierce and 

Howe 1977). Although a large percentage of winter flounder in a tagging 

study were recaptured at or near the 

original locations ’ 

(1961) repo??s??!hat the Sam;! b%? 

ing area is not always reoccupied each 

season. On a larger geographic scale, 

there is evidence that winter flounder 

north and south of Cape Cod and from 

Georges Banks compose three separate 

groups (Lux et al. 1970; Pierce and 

Howe 1977). 

GROWTH CHARACTERISTICS 

Growth Rate 

The rate of growth of the winter 

flounder is rapid until age V or VI 

and then decreases, 

males (Lux 1973). 

particularly in 

After the first 2 

years, females grow faster than males 

(Briggs 1965; Lux 1973; Howe and 

Coates 1975). An exception is in Newfoundland, where the growth rates of 

the sexes are similar (Kennedy and 

Steele 1971). 

The growth rate also differs between 

fish from areas relatively close geographically. Lengths of flounder at 

the same age were significantly different among certain bays on Long Island (Lobe11 1939; Poole 1966). 

Flounder grow to a larger size in the 

Georges Bank population than in inshore populations (Bigelow and 

Schroeder 1953). According to Berry 

et al. (1965), there is no typical 

growth rate for the winter flounder 

because the populations may be exposed 

to different rates of exploitation or 

live under different environmental 

conditions. In addition, the extended 

spawning period (up to 4 months) can 

make comparisons difficult between age 

groups and locations. 

Lux (1973) gave the following von 

Bertalanffy growth equations for winter flounder from Georges Bank: 

Length-Weight Relationships 

The length-weight 

published for adults 

presented in Table 1. 

THE FISHERY 

relationships 

and larvae are 

Commercial and Recreational 

The winter flounder supports valuable commercial and sport fisheries in 

the coastal waters of New England. 

The total commercial catch in the five 

coastal New England States was 15,500 

metric tons (t) in 1983 (U.S. Department of Commerce 1983). From 1935 to 

1980, the annual commercial landings 

in New England ranged between 6,000 

and 15,000 t. The otter trawl is the 

principal fishing gear. 

The winter flounder is a highly 

valued sport species because it is 

5 
Table 1. Published length-weight relationships for adult and larval winter 

flounder. 

Equation 

Adults 

Location Source 

10g,Ow = 3.138 10g,OL-5.239 

where W = g, L = mm 

Georges Bank Lux (1969) 

logloW = 3.1441 logloL-2.072 (female) 

log,oW = 2.9833 log,oL-1.9041 (male) Newfoundland Kennedy and 

where W = g, L = cm Steele (1971) 

Larvae 

logloW = 4.769 logloL-1.347 Laboratory-reared Laurence (1979) 

where W = mg, L = mm 

seasonally abundant in nearshore areas 

and easily captured from boat or 

shore. In New England the sport catch 

has been reported to surpass the commercial catch in some years (Deuel 

1973). 

Population Dynamics 

The age and size of winter flounder 

recruited into the fishery varies with 

the location and the type of fishery. 

Briggs (1965) reported that flounder 

recruited into the sport fishery at 

South Shore Bay, Long Island, were 

from 200 to 260 mm TL. In Nova 

Scotia, recruits into the commercial 

fishery were 3 to 4 years old and 

weighed an average of 363 g (Dickie 

and McCracken 1955). In Narragansett 

Bay, Rhode Island, winter flounder 

were fully recruited into the commercial catch at age III (250 mm TL; 

Saila et al. 1965). 

Estimated natural mortality rates of 

winter flounder ranged from 50% to 54% 

and total annual mortality (natural 

and fishing) ranged from 72% to 78% 

(Poole 1969). Total annual mortality 

rates estimated by Berry et al. (1965) 

on the basis of age composition for 

two different Long Island populations 

were 56% for males and 65% for females 

6 

in one population, and 51% for males 

and 58% for females in the other. The 

instantaneous mortality rates of 

winter flounder in Nova Scotia were 

0.321 (natural) and 0.475 (fishing) 

(Dickie and McCracken 1955). South 

of Cape Cod, Howe et al. (1976) 

reported instantaneous mortality rates 

of 0.1125 (natural) and 0.2445 

(fishing). 

Two important factors affecting mortality are translocation of larvae out 

of the estuary by drift (Pearcy 1962) 

and predation (Dickie and McCracken 

1955). Jeffries and Johnson (1974) 

reported that winter flounder abundance in Narragansett Bay may be partially governed by annual or seasonal 

changes in climate. Because each 

population does not usually disperse 

beyond local waters, the degradation 

of an estuary may have a drastic 

effect on the abundance of recruits in 

nearby coastal waters. 

ECOLOGICAL ROLE 

Food Habits 

Larvae begin to feed 2 to 3 weeks 

after they hatch. They first feed on 

copepods and phytoplankton, but as 

W” 

P 
they reach metamorphosis, their diet 

is composed of copepod nauplii, small 

polychaetes, nemerteans, and ostracods. For detailed descriptions of 

the food habits of larval and juvenile winter flounder, see Pearcy 

(1962). Laurence (1977), who studied 

the effects of food density on larval 

growth and survival, reported that the 

larvae died from starvation in 2 weeks 

at prey (nauplius) densities of 

tO.l/ml; critical prey density was 

about 0.5/ml. Plankton density influenced survival more than it did 

growth. Laurence (1977) demonstrated 

that the density of prey was probably 

the most important factor affecting 

survival. 

Adult winter flounder fed largely on 

organisms of three phyla: Annelida, 

Cnidaria, and Mollusca. In the study 

by Langton and Bowman (1981), the percentages of composition (numbers) of 

prey in flounder stomachs were as follows: Annelida 27% (mostly polychaete 

worms), Cnidaria 26%, Anthozoa 25%, 

Mollusca 16%, and Hydrosoa 4%. The 

composition varied among geographic 

locations. Tyler and Dunn (1976) 

reported that the maintenance ratio 

was 7.9 Cal/g. Detailed studies of 

the food of adult winter flounder were 

made' by Langton and Bowman (1981), 

Wells et al. (I973), Kennedy and 

Steele (1971), Olla et al. (1969), 

Mulkana (1966), and Frame (1973). 

Feeding Behavior 

Winter flounder primarily feed visually and only during daylight (Olla et 

al. 1969; MacDonald 1983). In the Bay 

of Fundy, those in nearshore waters 

usually fed in the intertidal zone 

(Wells et al. 1973). They moved 

inshore about 2 h after low tide and 

returned to the sublittoral zone about 

2 h before the next low tide (Tyler 

1971b). 

When feeding, the winter flounder 

lies motionless with its head raised 

off the bottom, braced by the dorsal 

fin. When a prey is sighted, the fish 

remains motionless, pointing toward 

the prey, and then lunges forward and 

downward to capture it. If no prey is 

sighted, the fish moves to a new location, changing position from four to 

five times per minute (Olla et al. 

1969). 

Comoetition 

The winter flounder has relatively 

few competitors for food and space. 

In many estuaries it is the most abundant demersal species (Richards 1963; 

Oviatt and Nixon 1973). The highly 

productive estuarine and coastal habitats it occupies, combined with its 

omnivorous food habits, tend to reduce 

competition. Jeffries and Johnson 

(1974) suspected that the early spawning and the short period of time to 

metamorphosis permit the larvae to 

reach the juvenile stage before potential competitors enter the bays and 

estuaries. 

Predators 

Adult winter flounder are the prey 

of many of the larger estuarine and 

coastal Dredators such as StriDed bass 

(Morone 'saxatilis), bluefish 

mus saltatrTx), goosefish 

GricanuS),spiny dogfish 

acanthias), oyster toadfish 

(Pomato- 

(Los 

(&Z-E 

($E 

tau), and sea raven ,(nHderni;;i;;;l;;; 

americanus) (Dickie 

1955; Grosslein and Azarovitz 1982). 

Predation is a major cause of mortality in larval and juvenile winter 

flounder. The larvae were heavily 

preyed upon by the small hydromedusa 

t~g'~~a,"ubu,ho,s,a,t~~eartchYat196t2h Tyler 

great 

cormorant (Phalacrocorax carbo), the 

great blue heron (Ardea herodias), and 

the osprey (Pandionhaliaetus) are 

also predators of winter flounder. 

Parasites 

The microsporidian parasite Glugea 

hertwigi is most common and may cause 

high mortality among winter flounder 

7 
less than 30 mm long (TL) (Mulkana 

1966). Klein-MacPhee (1978) provided 

a detailed list of the principal parasites of the winter flounder. 

ENVIRONMENTAL REQUIREMENTS 

Water Temperature 

Winter flounder are commonly found 

in water temperatures of 0 to 25 OC. 

Olla et al. (1969) reported that winter flounder fed at water temperatures as high as 22 OC, but burrow 

into the bottom at higher temperatures. McCracken (1963) gave a preferred temperature range of 12 to 

15 OC. Huntsman and Sparks (1924) 

reported a maximum temperature tolerance of about 30 OC. Under controlled 

conditions, winter flounder can acclimate to higher temperature regimes; 

for example, Everich and Gonzalez 

(1977) reported that the critical 

thermal maximum increased from 26 to 

32 OC as the acclimation temperature 

increased from 4 to 23 OC. An extended period of unusually hot weather 

caused heavy mortality in coastal waters of Long Island Sound (Nichols 

1918). Juvenile winter flounder tend 

to be more tolerant of high temperatures than adults. 

Salinity 

Adult winter flounder commonly live 

in salinities of 5 to 35 ppt (Bigelow 

and Schroeder 1953). Extremes in salinity may lower egg and larval survival and hatching success (see the 

section on eggs and larvae). 

Contaminants 

In a study in the Weweantic River, 

Massachusetts, chlorinated hydrocarbon insecticides and their breakdown 

products (DDT, DDE, heptachlor, heptachlor epoxide, and dieldrin) were 

found in various tissues of the winter 

flounder (Smith and Cole 1970; Smith 

1973). Concentrations of DDT, DDE, 

and heptachlor epoxide were highest in 

ripening ovaries. Agricultural runoff 

was the major source of the contaminants (Smith and Cole 1970). Topp 

(1967) reported that this contamination caused high mortality in the 

Weweantic River. 

In studies of the effects of silver 

on the eggs and larvae of winter 

flounder, Klein-MacPhee et al. (1984) 

found that concentrations of silver 

greater than 54 ug/l sometimes caused 

high mortality of the eggs and yolksac larvae, and that exposure to 92 

ug/l significantly increased egg mortalities. In contrast, Voyer et al. 

(1982) reported that silver in concentrations up to 166 ug/l did not 

increase egg mortality. 

Disease 

About 14% of the winter flounder 

examined from the New York Bight had 

fin erosion (Ziskowski and Murchelano 

1975). It is not known if the disease 

is infectious or noninfectious, but it 

is not usually fatal. Although the 

precise cause of fin rot erosion is 

not known, its high incidence in association with high sediment contamination suggests that contact of the fins 

with toxic sediment is an important 

factor in the development of the 

disease (Sherwood 1982). 

The microsporidian Glugea hertwigi, 

found in the digestive tract of winter 

flounder, was described by Stunkard 

and Lux (1965). The incidence of 

infection in samples ranged from 54% 

in Martha's Vineyard to zero on 

Georges Bank (Stunkard and Lux 1965). 

. 
LITERATURE CITED 

Berry, R.J., S.B. Saila, and D.B. 

Horton. 1965. Growth studies of 

winter flounder Pseudopleuronectes 

americanus in Rhode Island. Trans. 

Am. Fish. Sot. 94(3):259-264. 

habits, metabolism and food utilization. Ph.D. Thesis. University 

of Massachusetts, Amherst. 109 pp. 

Bigelow, H.B., and W.C. Schroeder. 

1953. Fishes of the Gulf of Maine. 

U.S. Fish Wildl. Serv. Fish. Bull. 

53. 577 pp. 

Breder, C.M. 1924. Some embryonic 

and larval stages of the winter 

flounder. Bull. U.S. Bur. Fish. 

38:311-315. 

Grosslein, M.D., and T.R. Azarovitz. 

1982. Fish distribution. MESA New 

York Bight Atlas Monogr. 15. New 

York Sea Grant Institute, Albany, 

N.Y. 182 pp. 

Howe, A.B., and P.G. Coates. 1975. 

Winter flounder movements, growth, 

and mortality off Massachusetts. 

Trans. Am. Fish. Sot. 104(1):13-19. 

Briggs, P. 1965. The sports 

fisheries for winter flounder in 

several bays of Long Island, N.Y. 

Fish Game 12(1):48-70. 

Howe, A.B., P.G. Coates, and D. 

Pierce. 1976. Winter flounder 

estuarine year-class abundance, mortality, and recruitment. Trans. Am. 

Fish. Sot. 105(6):647-657. 

Clayton, G., C.F. Cole, S.A. Murawski, 

and J.D. Parrish. 1978. Common 

marine fishes of coastal Massachusetts. Univ. Mass. Inst. Manage. 

Environ. Publ. R 76-16. 231 pp. 

Deuel, D.G. 1973. 1970 salt-water 

angling survey. U.S. Dep. Comm., 

NOAA, Natl. Mar. Fish. Serv. Stat. 

6200. 54 PP. 

Dickie, L.M., and F.D. McCracken. 

1955. Isopleth diagrams to predict 

equilibrium yields of a small flounder fishery. J. Fish. Res. Board 

Can. 12:187-209. 

Huntsman, A.G., and M.I. Sparks. 

1924. Limiting factors for marine 

animals. 3. Relative resistance to 

high temperatures. Contrib. Can. 

Biol. New Ser. 2:97-114. 

Jeffries, H.P., and W.C. Johnson. 

1974. Seasonal distribution of bottom fishes in the Narrangansett Bay 

area: seven year variation in the 

abundance of winter flounder (Pseudopleuronectes americanus). J. 

Fish. Res. Board Can. 31:369-372. 

Everich, D., and J.G. Gonzalez. 1977. 

Critical thermal maxima of two species of estuarine fish. Mar. Biol. 

(Berl.) 41:141-146. 

Kennedy, V.S., and D.H. Steele. 1971. 

The winter flounder (Pseudopleuronectes americanus in Long Pond, 

Conception Bav. Newfoundland. J. 

Fish.' Res. Board Can. 28:1153-1165. 

Klein-MacPhee, G. 1978. Synopsis of 

biological data for the winter 

Frame, D.W. 1973. Biology of young flounder, Pseudopleuronectes ameriwinter flounder Pseudopleuronectes canus (Walbaum). NOAA Tech. Rep. 

americanus (Walbaum): Feeding NMFS Circ. 414. 43 pp. 

9 
Klein-MacPhee, G., J.A. Cardin, and 

W.J. Berry. 1984. Effects of 

silver on eggs and larvae of the 

winter flounder. Trans. Am. Fish. 

Sot. 113:247-251. 

Langton, R.W., and R.E. Bowman. 1981. 

Food of eight northwest Atlantic 

pleuronectiform fishes. NOAA Tech. 

Rep. SSRF 749. 

La Roche, W.A. 1980. Development of 

larval smooth flounder, Liopsetta 

putnami, with a redescription of 

develoDment of winter flounder. 

Pseudopleuronectes americanus (farnil 

1Y Pleuronectes). Fish. Bull. 

78(4):897-910. 

Laurence, G.C. 1975. Laboratory 

growth and metabolism of the winter 

flounder (Pseudopleuronectes americanus) from hatching through metamorphosis at three temperatures. 

Mar. Biol. (Berl.) 32(3):223-229. 

Laurence, G.C. 1977. A bioenergetic 

model for the analyses of feeding 

and survival potential of winter 

flounder (Pseudopleuronectes americanus) larvae during the period from 

hatching to metamorphosis. U.S. 

Natl. Mar. Fish. Serv. Fish. Bull. 

75(3):529-549. 

Laurence, G.C. 1979. Larval lengthweight relationship for seven species of northwest Atlantic fishes 

reared in the laboratory. U.S. 

Natl. Mar. Fish. Serv. Fish. Bull. 

76(4):890-895. 

Leim, A-H., and W.8. Scott. 1966. 

Fishes of the Atlantic coast of Canada. Fish. Res. Board Can. Bull. 

155. 485 pp. 

Lobell, M.J. 1939. A biological survey of the salt waters of Long Island, 1938. 

fishes. 

Report on certain 

Winter flounder (PseudogA;;rOrctesN y americanus). 28th 

I. No.e:4:63-96. 

Conserv. Dep., Part 

Lux, F.E. 1969. Length-weight relationships of six New England flatfishes. Trans. Am. Fish. sot. 

98(4):617-621. 

Lux, F.E. 1973. Age and growth of 

the winter flounder, Pseudopleuronectes americanus, on Georges 

Bank. Fish. Bull. 71(2):505-512. 

Lux, F.E., A.E. Peterson, and R.F. 

Hutton. 1970. Geographic variation 

in fin ray numbers in winter flounder, Pseudopleuronectes americanus! off Massachusetts. Trans. 

Am. Fish. Sot. 99(3):483-488. 

MacDonald, J.S. 1983. Laboratory 

observations of feeding behavior of 

the ocean pout (Macrozoarces americanus) and winter flounder (Pseudopleuronectes americanus) with reference to niche overlap of natural 

populations. Can. ’ J. Zool. 

61(3):539-546. 

McCracken, F.D. 1963. Seasonal movements of the winter flounder, Pseudo leuronectes americanus 

K&ZZXZZ. J tish. i(es?z 

Can. 20(2):551-585: 

Mulkana, M.S. 1966. The growth and 

feeding habits of juvenile fishes in 

two Rhode Island estuaries. Gulf 

Res. Rep. 2:97-167. 

Nichols, J.T. 1918. An abnormal winter founder and others. Copeia 

55:36-39. 

Olla, B.L., R. Wicklund, and S. Wilk. 

1969. Behavior of winter flounder 

in a natural habitat. Trans. Am. 

Fish. Sot. 98(4):717-720. 

Oviatt, C.A., and S.W. Nixon. 1973. 

The demersal fish of Narrangansett 

Bay: an analysis of community 

structure, distribution and abundance. Estuarine Coastal Mar. Sci. 

1:361-378. 

Pearcy, W.G. 1962. Ecology of an 

estuarine populat ion of winter 

. 

10 
flounder Pseudopleuronectes americanus (WaTbaum). Bull. 

Oceanogr. Collect. Yale Univ 

B‘ingham 

. 18(l). 

78 pp.- 

Perlmutter, A. 1947. The b lackback 

flounder and its fishery in New 

England and New York. Bull. Bingham 

Oceanogr. Collect. Yale Univ. ll(2). 

92 PP. 

Pierce, D.E., and A.B. Howe. 1977. A 

further study on winter flounder 

group identification off Massachusetts. Trans. Am. Fish. Sot. 

106(2):131-139. 

Poole, J.C. 1966. Growth and age of 

winter flounder in four bays of Long 

Island. N.Y. Fish Game 

13(2):206-220. 

Poole, J.C. 1969. A study of winter 

flounder mortality rates in Great 

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Fish. Sot. 98(4);611-617. 

Richards, S.W. 1963. The demersal 

fish populations of Long Island 

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Rogers, C.A. 1976. Effects of temperature and salinity on the survival of winter flounder embryos. 

U.S. Natl. Mar. Fish. Serv. Fish. 

Bull. 74:52-58. 

Saila, S.B. 1961. Study of winter 

flounder movements. Limnol. Oceanogr. 6:292-298. 

Saila, S.B., D.B. Horton, and R.J. 

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fisherv in Rhode' Island. J. Fish. 

Res. Board Can. 22:945-954. 

Sherwood, M.J. 1982. Fin erosion, 

liver condition, and trace contaminant exposure in fishes from 

three coastal regions. In G.F. 

Meyer, ed. Ecological stressand the 

New York Bight: science and 

management. Estuarine Research 

Foundation, Columbia, S.C. 

Smith, R. 1973. Pesticide residues 

as a possible factor in larval winter flounder mortality. Pages 173- 

180 in Proceedings of a workshop in 

egg,Tarval, and juvenile stages of 

fish in Atlantic coast estuaries. 

NOAA Tech. Publ. No. 1. 

Smith, R.M., and C.F. Cole. 1970. 

Chlorinated hydrocarbon insecticide 

residues in winter flounder. Pseudopleuronectes americanus, from the 

Weweantic River Estuary, Massachusetts. J. Fish. Res. Board Can. 

27(12):2374-2380. 

Stunkard, H.W., and F.E. Lux. 1965. A 

microsporidian infection of the 

digestive tract of the winter flounder, Pseudopleuronectes americanus. B' 1 

m71-38;o ’ 

Bull. (Woods m 

Sullivan, W.E. 1915. Description of 

the younq staqes of the winter 

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Sot. 44:125-136. 

Topp, R.W. 1967. An estimate of fecundity of the winter flounder, 

Pseudopleuronectes americanus. J. 

Fish. Res. Board Can. 25(6):1299- 

1302. 

Tyler, A.V. 1971a. Periodic and resident components in communities of 

Atlantic fishes. J. Fish. Res. 

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Tvler. A.V. 1971b. Suraes of winter 

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Fish. Res. Board Can. 28(11):1717- 

1732. 

Tyler, A.V., and R.S. Dunn. 1976. 

Ration, growth, and measures of 

somatic and organ condition in relation to meal frequency in winter 

11 
flounder, Pseudopleuronectes americanus, with hypothesis regag 

mation homeostasis. J. Fish. 

Res. Board Can. 11:933-953. 

U.S. Department of Commerce. 1983. 

Fishery statistics of the United 

States. Washington, D.C. 

Van Guelpen, L., and C.C. Davis. 

1979. Seasonal movements of the 

winter flounder (Pseudopleuronectes 

americanus) in two contrasting inshore locations in Newfoundland. 

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Voyer, R.A., J.A. Cardin, J.F. 

Heltsche, and G.L. Hoffman. 1982. 

Viability of embryos of the winter 

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Toxicol. 2:223-233. 

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Tyler. 1973. Intertidal feeding of 

winter flounder, 

ronectes americanus, 

Fundy. J 

in _f 

30:1374-1378. 

Fish. Res. Board Can. 

Williams, G.C. 1975. Viable embryogenesis of the winter flounder 

(Pseudopleuronectes americanusie;;ot; 

18 15 "C 

33(l)%-74. * 

Mar. Blol. ( 

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1975. Fin erosion in winter flounder. Mar. Pollut. Bull. 6(2):26-29. 

. 

12 
DOCUMENTATION l- REWRT NO. 

~- 

2. 3. Recipient’s Accession NO. 

PAGE Biological Report 82(11.87)* 

I 

4. Titla and Subtitle 9. Report Date 

Species Profiles: Life Histories and Environmental Requirements 

of Coastal Fishes and Invertebrates (North Atlantic)-- 

Winter Flounder 

" *uthor(s) Jack Buckley 

9. PerforminS Organization N8ma and Address 

11. Contract(C) or Grant(G) NO. 

-___ -.-. 

12. Spoonwring OrSwnir8tion Name and Address 

U.S. Department of the Interior Coastal Ecology Group 

Fish and Wildlife Service Waterways Experiment Station 

Research and Development U.S. Army Corps of Engineers 

National Wetlands Research Center P.O. Box 631 

Washington, DC 20240 Vicksburg, MS 39180 

15. Supplementary Notes 

I *U.S. Army Corps of Engineers Report No. TR EL-82-4 

1S. Abstract (Limit: 200 words) 

Species profiles are literature summaries of the taxonomy, life history, and environmental requirements of coastal fishes and aquatic invertebrates. They are designed to 

assist with environmental impact assessments. From 1935 to 1980, the annual commercial 

landings of winter flounder in New England ranged from 6,000 to 15,000 t; the sport 

catch exceeded the commercial catch in some years. Winter flounder are found in waters 

with temperatures of 0 to 25 OC and they usually spawn at 0 to 3 OC. Reported 

fecundities are 0.5 to 1.5 million eggs per female. Metamorphosis from larva to juvenile 

is complete in 49-80 days, depending on temperature. Juveniles remain in or near shallow 

natal waters for much of their first 2 years. Adults of some populations move more than 

3 miles offshore to cooler waters in summer. These adults live in shallow inshore waters 

in winter and early spring. Adult winter flounder feed largely on annelids, cnidariids, 

and mollusks. 

17. Document Analysis a. O*scripton 

Estuaries Fishes 

Fisheries Life cycles 

Feeding habits Temperature 

Salinity 

Contaminants 

b. IdwWRen/OpwwEndcd Terms 

Winter flounder, Environmental requirements 

Pseudopleuronectes americanus Ecological role 

c. COSATI Field/Group 

1S. Availability Statement 

Release Unlimited 

19. Sccuroty Class (This Report) 21. No. of PaSea 

Unclassified 12 

-- 

20. Security Class (This Page) 22 Price 

Unclassified 

See ANSI-239.18) OPTIONAL CORM 272 (C’i 

(Formerly NTIS-35) 

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the Interior has responsibility for most of our nationally owned 

public lands and natural resources. This includes fostering the 

wisest use of our land and water resources, protecting our fish 

and wildlife, preserving the environmental and cultural values of our 

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our energy and mineral resources and works to assure that their 

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administration. 

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