--..

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S.... -. -. , .*': , .

...

Coastal Ecology GroupFish and-Wildlife Service Waterways Experiment Station

U.S. Department of the Interior U.S. Army Corps of Engineers

Biological Report 82(11.87)TR EL-82-4January 1989

Species Profiles: Life Histories and Environmental Requirements of CoastalFishes and Invertebrates (North Atlantic)

WINTER FLOUNDER

by

Jack BuckleyMassachusetts Cooperative Fishery Research UnitDepartment of Forestry and Wildlife Management

University of MassachusettsAmherst, MA 01003

Project OfficerDavid Moran

U.S. Fish and Wildlife ServiceNational Wetlands Research Center

1010 Gause BoulevardSlidell, LA 70458

Performed forCoastal Ecology Group

Waterways Experiment StationU.S. Army Corps of Engineers

Vicksburg, MS .39180

and

U.S. Department of the InteriorFish and Wildlife ServiceResearch and Development

National Wetlands Research CenterWashington, DC 20240

This series may be referenced as follows:

U.S. Fish and Wildlife Service. 1983-19 . Species profiles: life historiesand environmental requirements of coastal fishes and invertebrates. U.S. FishWildl. 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.ments of coastalU.S. Fish Wildl.EL-82-4. 12 pp.

Species profiles: life histories and environmental require-fishes and invertebrates (North Atlantic)--winter flounder.

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

PREFACE

This species profile is one of a series on coastal aquatic organisms,principally fish, of sport, commercial, or ecological importance. The profilesare designed to provide coastal managers, engineers, and biologists with a briefcomprehensive sketch of the biological characteristics and environmentalrequirements of the species and to describe how populations of the species may beexpected to react to environmental changes caused by coastal development. Eachprofile has sections on taxonomy, life history, ecological role, environmentalrequirements, and economic importance, if applicable. A three-ring binder isused 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 Engineersand the U.S. Fish and Wildlife Service.

Suggestions or questions regarding this report should be directed to one ofthe following addresses.

Information Transfer SpecialistNational Wetlands Research CenterU.S. Fish and Wildlife ServiceNASA-Slidell Computer Complex1010 Gause BoulevardSlidell, LA 70458

or

U.S. Army Engineer Waterways Experiment StationAttention: WESER-CPost Office Box 631Vicksburg, MS 39180

iii

CONVERSION TABLE

Multiply

millimeters (mm)centimeters (cm)meters (m)meters (m)kilometers (km)kilometers (km)

square meters (m2

)

square kilometers (km2 )hectares (ha)

liters (1)cubic meters (m

3)

cubic meters (m3 )

milligrams (mg)grams (g)kilograms (kg)metric tons (t)metric tons (t)

kilocalories (kcal)Celsius degrees ('C)

inchesinchesfeet (ft)fathomsstatute miles (mi)nautical miles (nmi)

square feet (ft 2 )square miles (mi 2 )acres

gallons (gal).cubic feet (ft 3 )acre-feet

ounces (oz)ounces (oz)pounds (Ib)pounds (Ib)short tons (ton)

Metric to U.S. Customary

BY0.039370.3937

3.2810.54680.62140.5396

10.760.38612.471

0.264235.310.0008110

0.000035270.035272.205

2205.01.102

3.968

1.8( 0 C) + 32

U.S. Customary to Metric

25.402.540.30481.8291.6091.852

To Obtain

inchesinchesfeetfathomsstatute milesnautical miles

square feetsquare milesacres

gallonscubic feetacre-feet

ouncesouncespoundspoundsshort tons

0.09292.5900.4047

3. 7850.02831

1233.0

28350.028.350.45360.000450.9072

0.25200.5556 (OF - 32)

British thermal unitsFahrenheit degrees

millimeterscentimetersmetersmeterskilometerskilometers

square meterssquare kilometershectares

literscubic meterscubic meters

milligramsgramskilogramsmetric tonsmetric tons

kilocaloriesCelsius degrees

British thermal units (Btu)Fahrenheit degrees (OF)

iv

CONTENTS

Page

PREFACE .. TABL.. ............................ ............ ............ ..... vCONVERSION TABLE ......................................................... vACKNOWLEDGMENTS........................................................... vi

NOMENCLATURE/TAXONOMY/RANGE ............................................... 1MORPHOLOGY/IDENTIFICATION AIDS ............................................. 1SEPARATION FROM OTHER RIGHT-EYED FLATFISHES ............................... 3REASONS FOR INCLUSION IN SERIES ........................................... 3LIFE HISTORY .............................................................. 3

Spawning ................................................................. 3Eggs ..... ............................................................... 4Larvae .................................................................. 4Juveniles ...................................................... 4Adults .................................................................. 4

GROWTH CHARACTERISTICS .................................................... 5Growth Rate ............................................................. 5Length-Weight Relationships .............................................. 5

THE FISHERY ............................................................... 5Commercial and Recreational .............................................. 5Population Dynamics ..................................................... 6

ECOLOGICAL ROLE ........................................................... 6Food Habits .............. I ............................................... 6Feeding Behavior ........................................................ 7Competition ............................................................. 7Predators ............................................................... 7Parasites ............................................................... 7

ENVIRONMENTAL REQUIREMENTS ................................................ 8Water Temperature ........................................................ 8Salinity ................................................................ 8Contaminants ............................................................ 8Disease ................................................................. 8

LITERATURE CITED .......................................................... 9

v

ACKNOWLEDGMENTS

I am grateful for reviews by Wendy Gabriel, National Marine FisheriesService, Woods Hole, Massachusetts, and Arnold Howe, Massachusetts Division ofMarine Fisheries, Sandwich, Massachusetts.

vi

Figure 1. Winter flounder.

WINTER FLOUNDER

NOMENCLATURE/TAXONOMY/RANGE

Scientific name . . .. Pseudopleuro-nectes americanus (Walbaum)

Preferred common name . . .. Winterflounder (Figure 1)

Other common names ....... Blackbackflounder, lemon sole, black flounder

Class ..... .......... OsteichthyesOrder ........ PleuronectiformesFamily ......... Pleuronectidae

Geographic range: Winter flounder arefound primarily in estuarine andcoastal waters along the Atlanticcoast of North America fromNewfoundland to Georgia (Leim andScott 1966), except for off-shorepopulations on Georges Bank andNantucket Shoal (Figure 2; Bigelowand Schroeder 1953).

MORPHOLOGY/IDENTIFICATION AIDS

The winter flounder, one of theright-eyed flounders, is oval-shapedand thick-bodied; the caudal fin andpeduncle are broader than those ofother North Atlantic flounders. Theanal fin is highest at its midpointand is preceded by a short sharpspine. The dorsal fin (60-76 rays)originates opposite the anterior edgeof the eye, and is about equal inheight along its length. The mouth issmall, not gaping to the eye. Theleft (under) half of the jaw is armedwith a series of close-set incisors;the right (upper) half has only a fewteeth.

The winter flounder, like otherflatfishes, varies in color, depending

1

/ CANADA

JI

CL.

'U

z

/ MAINE

PORTLAND

A TL ANTIC OCEAJ

Coastal distributionMILES

0 so 100

0 so 100

KILOMETERS

GEORGES BANK

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

v

2

largely on the color of the surround-ing substrate. Most adults tend tobe reddish brown, olive-green, orblackish. Smaller fish generallyare paler than larger fish. Theblind side is.white and, toward theedge, translucent or occasionallyyellowish.

SEPARATION FROM OTHERRIGHT-EYED FLATFISHES

Compared with the yellowtail floun-der, Limanda ferruginea, the winterflounder has a. much straighter lateralline, a less concave dorsal head pro-file, and fewer fin rays.

The winter flounder lacks the mucouspits that are conspicuous on the left(blind) side of the head of the witchflounder (Glyptocephalus cynoglossus);it also has three times as many dorsalrays as the witch flounder.

The scales between the eyes aresmooth in the smooth flounder (Liop-setta putnami), but rough in the win-ter flounder. Between the two, thewinter flounder also has the greaternumber of anal fin rays.

LIFE HISTORY

Spawning

The winter flounder spawns in coast-al waters as early as December in theSouthern United States and as late asJune in Canada. Typically, eggs aredeposited over a sandy substrate atdepths of 2 to 80 m (Bigelow andSchroeder 1953).

Most spawning takes place at salin-ities of 31 to 32.5 ppt in inshorewaters, and on Nantucket Shoal andGeorges Bank at slightly higher salin-ities (32.7 to 33 ppt, respectively;Bigelow and Schroeder 1953). Watertemperature during spawning is usuallybetween 0 and 3 'C but may be as highas 6 'C (Bigelow and Schroeder 1953).The winter flounder spawns at slightlyhigher temperatures on Georges Bankthan in inshore waters (Lux et al.1970).

The stage of maturity of the winterflounder is largely governed by sizerather than age. Flounders growfaster and mature at a younger age inthe south than in the north. In New-foundland, males mature at age VI andfemales at age VII (Kennedy and Steele1971); in New York, winter flounder-mature at age II or III (Perlmutter1947).

The fecundity of winter flounderreported by Bigelow and Schroeder(1953) ranged from 0.5 to 1.5 millioneggs per female. Saila (1961)reported that in Rhode Island waters193,000 eggs were produced by a fish249 mm total length (TL) and 1.34 mil-lion by a fish 428 mm TL. In theWeweantic Estuary in Massachusetts,numbers of eggs ranged from 435,000for a fish 350 mm TL to 3.3 millionfor a fish 450 mm TL (Topp 1967).In Newfoundland, Kennedy and Steele(1971) reported a fecundity rangefrom 99,000 eggs for a. fish 220 mmTL to 2.6 million for a fish 440 mmTL (mean = 0.59 million eggs. ata mean length of 340 mm. TL). The

Severaltics thatflounderfloundersAtlantic(1980).

morphological characteris-distinguish larvae of winterfrom' those of the othercommon in the western northwere given by La Roche

REASONS FOR INCLUSION IN THE SERIES

By virtue of its abundance in estua-rine and nearshore waters, the winterflounder is one of the most importantcommercial and sport fishes in theNortheastern United States. InMassachusetts, it is considered amajor contributor to the commercialand sport fisheries (Pierce and Howe1977).

3

following equations for estimatingfecundity on the basis of weight havebeen published:

log F = 2.3894 + 1.2403 log W(Kennedy and Steele 1971)

log F = 0.0697(Topp 1967)

log F = 2.6712(Saila 1961)

+ 1.0659 log W

+ 1.1383 log W

where F = fecundity in thousands ofeggs and W = total weight in grams.

Winter flounder eggs are demersal,adhesive, and 0.74 to 0.85 mm in diam-eter (Bigelow and Schroeder 1953).They have no oil globule when depos-ited, but acquire one later (Breder1924). Incubation time Was 15 to 18days at 2.8 to 3.3 0C (Bigelow andSchroeder 1953), 25 days at 3 0C, and7 days at 12 to 14 'C (Rogers 1976).Incubation time was inversely relatedto water temperature and salinity(Rogers 1976).

Winter flounder eggs seem to be mostabundant in water with a salinity of10 to 30 ppt; at salinities below 5ppt or above 40 ppt, some embryos sur-vive, but are usually deformed (Rogers1976). The optimal salinity for eggsurvival is 15 to 35 ppt.

Many embryos become inviable or ab-normal at temperatures below freezing(-1.8 to 0 'C) and temperatures above10 *C (Williams 1975). The optimumwater temperature range for survivalis 0 to 10 'C (Williams 1975).

Larvae

In studies by Bigelow and Schroeder(1953) and La Roche (1980), winterflounder larvae were 2.4 to 3.5 mm TLat the time of hatching. A majorcharacteristic of the newly hatchedlarvae was the broad vertical band of

pigment cells dividing the postanalportion of the body. At a water tem-perature of 3.9 0C the larvae wereabout 5 mm TL, and the yolk sac wasabsorbed in 12 to 14 days. La Roche(1980) provides a detailed descriptionof larval development.

Winter flounder undergo a rapidmetamorphosis at a much smaller sizethan other flatfishes of the NorthAtlantic region (Bigelow and Schroeder1953). Their metamorphosis is com-plete when the larvae are 8 to 9 mm TL(Laurence 1975); this transformationtook 80 days at a water temperature of5 0C and 49 days at 8 *C. No metamor-phosis was evident at 2 'C (Bigelowand Schroeder 1953).

In aquaria, winter flounder larvaeengage in upward swimming bouts andthen sink to the bottom where theyremain for a short time (Sullivan1915; Bigelow and Schroeder 1953).The larvae of other flatfish speciesare more pelagic. Winter flounderlarvae are continuous, visual, day-light feeders that cease feeding atnight (Laurence 1977).

Juveniles

After metamorphosis, winter flounderare benthic and seldom lose contactwith the substrate. Most juvenilesspend much of their first 2 years inor near shallow natal waters, wherethey move in response to' extreme heator cold (Topp 1967). After meta-morphosis, the juveniles prefer a sub-strate of sand or sand and silt(Clayton et al. 1978). Older juve-niles in estuaries gradually move sea-ward as they grow larger (Mulkana1966).

Adults

The seasonal movements of winterflounder differ between populationsnorth and south of Cape Cod. A 5-yeartagging study by Howe and Coates(1975) showed that winter floundernorth of Cape Cod moved about only

4

locally in inshore waters, while thosesouth of Cape Cod dispersed more than3 mi offshore in a southwesterlydirection. Adults from Martha's Vine-yard and coastal populations fromsouth of Cape Cod mixed in NantucketSound (Pierce and Howe 1977).

Water temperature seems to be themost important environmental factordetermining seasonal distribution(McCracken 1963). In Rhode Island,adult winter flounder lived in cooleroffshore waters during summer and inshallow inshore waters in winter andearly spring (Saila 1961). In New-foundland, winter flounder remained inshallow water during summer as long asfood was available and water tem-peratures did not exceed 15 'C (VanGuelpen and Davis 1979). Temperatureis a less important factor in the dis-tribution of'juveniles, which toleratehigher temperatures than adults(Pearcy 1962).

Indications are that a local popula-tion is defined by fish inhabiting.several adjacent estuaries (Pierce andHowe 1977). Although a large percent-age of winter flounder in a taggingstudy were recaptured at or near theoriginal tagging locations, Saila(1961) reported that the same breed-ing area is not always reoccupied eachseason. On a larger geographic scale,there is evidence that winter flounder.north and south of Cape Cod and fromGeorges Banks compose three separategroups (Lux et al. 1970; Pierce andHowe 1977).

GROWTH CHARACTERISTICS

Growth Rate

The rate of growth of the winterflounder is rapid until age V or VIand then decreases, particularly inmales (Lux 1973). After the first 2years, females grow faster than males(Briggs 1965; Lux 1973; Howe andCoates 1975). An exception is in New-foundland, where the growth rates of

the sexes are similar (Kennedy andSteele 1971).

The growth rate also differs betweenfish from areas relatively close geo-graphically. Lengths of flounder atthe same age were significantly dif-ferent among certain bays on Long Is-land (Lobell 1939; Poole 1966).Flounder grow to a larger size in theGeorges Bank population than in in-shore populations (Bigelow andSchroeder 1953). According to Berryet al. (1965), there is no typicalgrowth rate for the winter flounderbecause the populations may be exposedto different rates of exploitation orlive under different environmentalconditions. In addition, the extendedspawning period (up to 4 months) canmake comparisons difficult between agegroups and locations.

Lux (1973) gave the following vonBertalanffy growth equations for win-ter flounder from Georges Bank:

male I 550 [1 - -O.37(t-O.05)j

female 1 630 [1 - 0.31(t-0.05)

Lenqth-Weiqht Relationships

The 1publishedpresented

ength-weightfor adults

in Table 1.

relationshipsand larvae are

THE FISHERY

Commercial and Recreational

The winter flounder supports valu-able commercial and sport fisheries inthe coastal waters of New England.The total commercial catch in the fivecoastal New England States was 15,500metric tons (t) in 1983 (U.S. Depart-ment of Commerce 1983). From 1935 to1980, the annual commercial landingsin New England ranged between 6,000and 15,000 t. The otter trawl is theprincipal fishing gear.

The winter flounder is a highlyvalued sport species because it is

5

Table 1.flounder.

Published length-weight relationships for adult and larval winter

Equation Location Source

Adults

logj 0W = 3.138 log1oL-5.239 Georges Bank Lux (1969)where W = g, L = mm

logj 0W = 3.1441 logloL-2.072 (female)

logjoW = 2.9833 log 1oL-1.9041 (male) Newfoundland Kennedy andwhere W = g, L = cm Steele (1971)

Larvae

logjoW = 4.769 log1 oL-1. 347 Laboratory-reared Laurence (1979)where W = mg, L = mm

seasonally abundant in nearshore areasand easily captured from boat orshore. In New England the sport catchhas been reported to surpass the com-mercial catch in some years (Deuel1973).

Population Dynamics

The age and size of winter flounderrecruited into the fishery varies withthe location and the type of fishery.Briggs (1965) reported that flounderrecruited into the sport fishery atSouth Shore Bay, Long Island, werefrom 200 to 260 mm TL. In NovaScotia, recruits into the commercialfishery were 3 to 4 years old andweighed an average of 363 g (Dickieand McCracken 1955). In NarragansettBay, Rhode Island, winter flounderwere fully recruited into the commer-cial catch at age II1 (250 mm TL;Saila et al. 1965).

Estimated natural mortality rates ofwinter.flounder ranged from 50% to 54%and total annual mortality (naturaland fishing) ranged from 72% to 78%(Poole 1969). Total annual mortalityrates estimated by Berry et al. (1965)on the basis of age composition fortwo different Long Island populationswere 56% for males and 65% for females

in one population, and 51% for malesand 58% for females in the other. Theinstantaneous mortality rates ofwinter flounder in Nova Scotia were0.321 (natural) and 0.475 (fishing)(Dickie and McCracken 1955). Southof Cape Cod, Howe et al. (1976)reported instantaneous mortality ratesof 0.1125 (natural) and 0.2445(fishing).

Two important factors affecting mor-tality are translocation of larvae outof the estuary by drift (Pearcy 1962)and predation (Dickie and McCracken1955). Jeffries and Johnson (1974)reported that winter flounder abun-dance in Narragansett Bay may be par-tially governed by annual or seasonalchanges in climate. Because eachpopulation does not usually dispersebeyond local waters, the degradationof an estuary may have a drasticeffect on the abundance of recruits innearby coastal waters.

ECOLOGICAL ROLE

Food Habits

Larvae begin to feed 2 toafter they hatch. They firstcopepods and phytoplankton,

3 weeksfeed onbut as

6

they reach metamorphosis, their dietis composed of copepod nauplii, smallpolychaetes, nemerteans, and ostra-cods. For detailed descriptions ofthe food habits of larval and juve-nile winter flounder, see Pearcy(1962). Laurence (1977), who studiedthe effects of food density on larvalgrowth and survival, reported that thelarvae died from starvation in 2 weeksat prey (nauplius) densities of<0.1/ml; critical prey density wasabout 0.5/ml. Plankton density in-fluenced survival more than it didgrowth. Laurence (1977) demonstratedthat the density of prey was probablythe most important factor affectingsurvival.

Adult winter flounder fed largely onorganisms of three phyla: Annelida,Cnidaria, and Mollusca. In the studyby Langton and Bowman (1981), the per-centages of composition (numbers) ofprey in flounder stomachs were as fol-lows: Annelida 27% (mostly polychaeteworms), Cnidaria 26%, Anthozoa 25%,Mollusca 16%, and Hydrosoa 4%. Thecomposition varied among geographiclocations. Tyler and Dunn (1976)reported that the maintenance ratiowas 7.9 cal/g. Detailed studies ofthe food of adult winter flounder weremade' by Langton and Bowman (1981),Wells et al. (1973), Kennedy andSteele (1971), Olla et al. (1969),Mulkana (1966), and Frame (1973).

Feeding Behavior

Winter flounder primarily feed visu-ally and only during daylight (Olla etal. 1969; MacDonald 1983). In the Bayof Fundy, those in nearshore watersusually fed in the intertidal zone(Wells et al. 1973). They movedinshore about 2 h after low tide andreturned to the sublittoral zone about2 h before the next low tide (Tyler1971b).

When feeding, the winter flounderlies motionless with its head raisedoff the bottom, braced by the dorsalfin. When a prey is sighted, the fish

remains motionless, pointing towardthe prey, and then lunges forward anddownward to capture it. If no prey issighted, the fish moves to a new loca-tion, changing position from four tofive times per minute (Olla et al.1969)..

Competition

The winter flounder has relativelyfew competitors for food and space.In many estuaries it is the most abun-dant demersal species (Richards 1963;Oviatt and Nixon 1973). The highlyproductive estuarine and coastal habi-tats it occupies, combined with itsomnivorous food habits, tend to reducecompetition. Jeffries and Johnson(1974) suspected that the early spawn-ing and the short period of time tometamorphosis permit the larvae toreach the juvenile stage before po-tential competitors enter the bays andestuaries.

Predators

Adult winter flounder are the preyof many of the larger estuarine andcoastal predators such as striped bass(Morone saxatilis), bluefish (Pomato-mus saltatrix), goosefish (T-•s

americanus), spiny dogfish (Squalusacanthias), oyster toadfish (Opsanustau), and sea raven (Hemitripterusamericanus) (Dickie and McCracken1955; Grosslein and Azarovitz 1982).

Predation is a major cause of mor-tality in larval and juvenile winterflounder. The larvae were heavilypreyed upon by the small hydromedusaSaria tubulosa (Pearcy 1962). Tyler(1-971a) reported that the greatcormorant (Phalacrocorax carbo), thegreat blue heron (Ardea heFo-dis), andthe osprey (Pand-io haliaetus) arealso predators of winter flounder.

Parasites

The microsporidian parasite Glugeahertwigi is most common and may causehigh mortality among winter flounder

7

less than 30 mm long (TL) (Mulkana1966). Klein-MacPhee (1978) provideda detailed list of the principal para-sites of the winter flounder.

ENVIRONMENTAL REQUIREMENTS

Water Temperature

Winter flounder are commonly foundin water temperatures of 0 to 25 OC.Olla et al. (1969) reported that win-ter flounder fed at water tempera-tures as high as 22 0C, but burrowinto the bottom at higher tempera-tures. McCracken (1963) gave a pre-ferred temperature range of 12 to15 0C. Huntsman and Sparks (1924)reported a maximum temperature toler-ance of about 30 0C. Under controlledconditions, winter flounder can accli-mate to higher temperature regimes;for example, Everich and Gonzalez(1977) reported that the criticalthermal maximum increased from 26 to32 *C as the acclimation temperatureincreased from 4 to 23 0C. An extend-ed period of unusually hot weathercaused heavy mortality in coastal wa-ters of Long Island Sound (Nichols1918). Juvenile winter flounder tendto be more tolerant of high tempera-tures than adults.

Salinity

Adult winter flounder commonly livein salinities of 5 to.35 ppt (Bigelowand Schroeder 1953). Extremes in sa-linity may lower egg and larval sur-vival and hatching success (see thesection on eggs and larvae).

Contaminants

In-a study in the Weweantic River,Massachusetts, chlorinated hydrocar-bon insecticides and their breakdown

products (DDT, DDE, heptachlor, hep-tachlor epoxide, and dieldrin) werefound in various tissues of the winterflounder (Smith and Cole 1970; Smith1973). Concentrations of DDT, DOE,and heptachlor epoxide were highest inripening ovaries. Agricultural runoffwas the major source of the con-taminants (Smith and Cole 1970). Topp(1967) reported that this contamina-tion caused high mortality in theWeweantic River.

In studies of the effects of silveron the eggs and larvae of winterflounder, Klein-MacPhee et al. (1984)found that concentrations of silvergreater than 54 pg/l sometimes causedhigh mortality of the eggs and yolk-sac larvae, and that exposure to 92pg/l significantly increased egg mor-talities. In contrast, Voyer et al.(1982) reported that silver in con-centrations up to 166 pg/l did notincrease egg mortality.

Disease

About 14% of the winter flounderexamined from the New York Bight hadfin erosion (Ziskowski and Murchelano1975). It is not known if the diseaseis infectious or noninfectious, but itis not usually fatal. Although theprecise cause of fin rot erosion isnot known, its high incidence in asso-ciation with high sediment contamina-tion suggests that contact of the finswith toxic sediment is an importantfactor in the development of thedisease (Sherwood 1982).

The microsporidian Glugea hertwigi,found in the digestive tract of winterflounder, was described by Stunkardand Lux (1965). The -incidence ofinfection in samples ranged from 54%in Martha's Vineyard to zero onGeorges Bank (Stunkard and Lux 1965).

8

LITERATURE CITED

Berry, R.J., S.B. Saila, and D.B.Horton. 1965. Growth studies ofwinter flounder Pseudopleuronectesamericanus in Rhode Island. Trans.Am. Fish. Soc. 94(3):259-264.

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.and larvalflounder.38:311-315.

1924.stages

Bull.

Some embryonicof the winter

U.S. Bur. Fish.

The . sportster flounder inng Island, N.Y.70.

Briggs, P. 1965.fisheries for wintseveral bays of LoFish Game 12(1):48-7

Clayton, G., C.F. Cole, S.A'.,Murawski,and J.D. Parrish. 1978. Commonmarine fishes of coastal Massa-chusetts. Univ. Mass. Inst. Manage.Environ. Publ. R 76-16. 231 pp.

habits, metabolism and food utili-zation. Ph.D. Thesis. Universityof Massachusetts, Amherst. 109 pp.

Grosslein, M.D., and T.R. Azarovitz.1982. Fish distribution. MESA NewYork Bight Atlas Monogr. 15. NewYork 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. Soc. 104(1):13-19.

Howe, A.B., P.G. Coates, and D.Pierce. 1976. Winter flounderestuarine year-class abundance, mor-tality, and recruitment. Trans. Am.Fish. Soc. 105(6):647-657.

Huntsman, A.G., and M.I. Sparks.1924. Limiting factors for marineanimals. 3. Relative resistance tohigh temperatures. Contrib. Can.Biol. New Ser. 2:97-114.

Jeffries, H.P., and W.C. Johnson.1974. Seasonal distribution of bot-tom fishes in the Narrangansett Bayarea: seven year variation in theabundance of winter flounder (Pseu-dopleuronectes americanus). J.Fish. Res. Board Can. 31:369-372.

Kennedy, V.S., and D.H. Steele. 1971.The winter flounder (Pseudopleu-ronectes americanus in Long Pond,Conception Bay, Newfoundland. J.Fish. Res. Board Can. 28:1153-1165.

Klein-MacPhee, G. 1978. Synopsis ofbiological data for the winterflounder, Pseudopleuronectes ameri-canus (Walbaum). NOAA Tech. Rep.NMFS Circ. 414. 43 pp.

Deuel, D.G. 1973.angling survey.NOAA, Natl. Mar.6200. 54 pp.

1970 salt-waterU.S. Dep. Comm.,Fish. Serv. Stat.

Dickie, L.M., and F.D. McCracken.1955. Isopleth diagrams to predictequilibrium yields of a-.small floun-der fishery. J. Fish. Res. BoardCan. 12:187-209.

Everich, D., and J.G. Gonzalez. 1977.Critical thermal maxima of two spe-cies of estuarine fish. Mar. Biol.(Berl.) 41:141-146.

Frame, D.W. 1973. Biology of youngwinter flounder Pseudopleuronectesamericanus (Walbaum): Feeding

9

I

Klein-MacPhee, G., J.A. Cardin, andW.J. Berry. 1984. Effects ofsilver on eggs and larvae of thewinter flounder. Trans. Am. Fish.Soc. 113:247-251.

Langton, R.W., and R.E. Bowman. 1981.Food of eight northwest Atlanticpleuronectiform fishes. NOAA Tech.Rep. SSRF 749.

La Roche, W.A. 1980. Development of'larval smooth flounder, Liopsettaputnami, with a redescription ofdevelopment of winter flounder,Pseudopleuronectes americanus (fami-ly Pleuronectes). Fish. Bull.78(4):897-910.

Laurence, G.C. 1975. Laboratorygrowth and metabolism of the winterflounder (Pseudopleuronectes ameri-canus) from hatching through meta-morphosis at three temperatures.Mar. Biol. (Berl.) 32(3):223-229.

Laurence, G.C. 1977. A bioenergeticmodel for the analyses of feedingand survival potential of winterflounder (Pseudopleuronectes ameri-canus) larvae during the period fromhatching to metamorphosis. U.S.Natl. Mar. Fish. Serv. Fish. Bull.75(3):529-549.

Laurence, G.C. 1979. Larval length-weight relationship for seven spe-cies of northwest Atlantic fishesreared in the laboratory. U.S.Natl. Mar. Fish. Serv. Fish: Bull.76(4):890-895.

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rel a-flat-

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Lux, F.E. 1973. Age and growth ofthe winter flounder, Pseudopleu-ronectes americanus, on GeorgesBank. Fish. Bull. 71(2):505-512.

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10

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11

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12

qn,,, .Ini50272-1.,

REPORT DOCUMENTATION IL REPORT NO. 2-. 1. RCipiaft's Acession No.

PAGE Biological Report 82(11.87)*4. Title and Subtitle S. Report Date

Species Profiles: Life Histories and Environmental Requirements January 1989of Coastal Fishes and Invertebrates (North Atlantic)--Winter Flounder

7. Adthor(s) Jack Buckley 6. Perfor•ning Orgenizatlon Rapt. Na.

9. Performing Organization Name and Address 10. Project/Task/Wortk Unit No.

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

(C)

(G)12. Sponsoring Organization Name and Address

U.S. Department of the Interior Coastal Ecology Group I& Type o Report Priod Covered

Fish and Wildlife Service Waterways Experiment StationResearch and Development U.S. Army Corps of EngineersNational Wetlands Research Center P.O. Box 631 14.

.Washington, DC '20240 Vicksburg, MS 39180

1I. Supplementary No"es*U.S. Army Corps of Engineers Report No. TR EL-82-4

-I&. Abstract (Limit: 200 worde)

Species profiles are literature summaries of the taxonomy, life history, and environ-mental requirements of coastal fishes and aquatic invertebrates. They are designed toassist with environmental impact assessments. From 1935 to 1980, the annual commerciallandings of winter flounder in New England ranged from 6,000 to 15,000 t; the sportcatch exceeded the commercial catch in some years. Winter flounder are found in waterswith temperatures of 0 to 25 'C and they usually spawn at 0 to 3 'C. Reportedfecundities are 0.5 to 1.5 million eggs per female. Metamorphosis from larva to juvenileis complete in 49-80 days, depending on temperature. Juveniles remain in or near shallownatal waters for much of their first 2 years. Adults of some populations move more than3 miles offshore to cooler waters in summer. These adults live in shallow inshore watersin winter and early spring. Adult winter flounder feed largely on annelids, cnidariids,and mollusks.

17. Document Analysis a. Descriptors

Estuaries Fishes SalinityFisheries Life cycles ContaminantsFeeding habits Temperature

b. Identlfief/Op•en-End*d Terms

Winter flounder, Environmental requirementsPseudopleuronectes americanus Ecological role

C. COSATI Feld/Group

I& Asailebillty Statement 19. Securty Class (This Report) 21. No. of Pages

Release Unlimited Unclassified 12. Security Class (This Page) ZL Price

Unclassified I(See ANSi-739.18) OPTIONAL FORM Z7Z (4-77)

(Formerly NTIS-33)DeIa•rt"Wet of Commemce

TAKE PRIDEiniAmerica

DEPARTMENT OF THE INTERIORU.S. FISH AND WILDLIFE SERVICE

As the Nation's principal conservation agency, the Department of the Interior has respon-sibility for most of our .nationally owned public lands and natural resources. This includesfostering the wisest use of our land and water resources, protecting our fish and wildlife,preserving thsenvironmental and cultural values of our national parks and historical places,and providing for the enjoyment of life through outdoor recreation. The Department as-sesses our energy and mineral resources and works to assure that their development is inthe best interests of all our people. The Department also has a major responsibility forAmerican Indian reservation communities and for people who live in Island territories underU.S. administration.