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Fws/Usace - Species Profile: Winter Flounder
ML072060561
Person / Time
Site: Oyster Creek
Issue date: 01/01/1989
From: Buckley J, Moran D
Univ of Massachusetts - Amherst, US Dept of Interior, Fish & Wildlife Service, US Dept of the Army, Corps of Engineers
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Office of Nuclear Reactor Regulation
Davis J NRR/DLR/REBB, 415-3835
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References
TR EL-82-4 82(11.87)
Download: ML072060561 (21)


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--..' --- --- .... .-* _ --- .- --~S.... -. -. , .*': , ....Coastal Ecology Group Fish 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-82-4 January 1989 Species Profiles:

Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (North Atlantic)WINTER FLOUNDER by 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 coastal 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.ments of coastal U.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 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-Slidell 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 millimeters (mm)centimeters (cm)meters (m)meters (m)kilometers (km)kilometers (km)square meters (m 2)square kilometers (km 2)hectares (ha)liters (1)cubic meters (m 3)cubic meters (m 3)milligrams (mg)grams (g)kilograms (kg)metric tons (t)metric tons (t)kilocalories (kcal)Celsius degrees ('C)inches inches feet (ft)fathoms statute 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 BY 0.03937 0.3937 3.281 0.5468 0.6214 0.5396 10.76 0.3861 2.471 0.2642 35.31 0.0008110 0.00003527 0.03527 2.205 2205.0 1.102 3.968 1.8(0 C) + 32 U.S. Customary to Metric 25.40 2.54 0.3048 1.829 1.609 1.852 To Obtain inches inches feet fathoms statute miles nautical miles square feet square miles acres gallons cubic feet acre-feet ounces ounces pounds pounds short tons 0.0929 2.590 0.4047 3. 785 0.02831 1233.0 28350.0 28.35 0.4536 0.00045 0.9072 0.2520 0.5556 (OF -32)British thermal units Fahrenheit degrees millimeters centimeters meters meters kilometers kilometers square meters square kilometers hectares liters cubic meters cubic meters milligrams grams kilograms metric tons metric tons kilocalories Celsius degrees British thermal units (Btu)Fahrenheit degrees (OF)iv CONTENTS Page PREFACE .. TABL.. ............................

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.....v CONVERSION TABLE .........................................................

v ACKNOWLEDGMENTS...........................................................

vi NOMENCLATURE/TAXONOMY/RANGE

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1 MORPHOLOGY/IDENTIFICATION AIDS .............................................

1 SEPARATION FROM OTHER RIGHT-EYED FLATFISHES

...............................

3 REASONS FOR INCLUSION IN SERIES ...........................................

3 LIFE HISTORY ..............................................................

3 Spawning .................................................................

3 Eggs ..... ...............................................................

4 Larvae ..................................................................

4 Juveniles

......................................................

4 Adults ..................................................................

4 GROWTH CHARACTERISTICS

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5 Growth Rate .............................................................

5 Length-Weight Relationships

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5 THE FISHERY ...............................................................

5 Commercial and Recreational

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5 Population Dynamics .....................................................

6 ECOLOGICAL ROLE ...........................................................

6 Food Habits ..............

I ...............................................

6 Feeding Behavior ........................................................

7 Competition

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7 Predators

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7 Parasites

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7 ENVIRONMENTAL REQUIREMENTS

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8 Water Temperature

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8 Salinity ................................................................

8 Contaminants

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8 Disease .................................................................

8 LITERATURE CITED ..........................................................

9 v 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.

vi Figure 1. Winter flounder.WINTER FLOUNDER NOMENCLATURE/TAXONOMY/RANGE Scientific name .... Pseudopleuro-nectes americanus (Walbaum)Preferred common name .... Winter flounder (Figure 1)Other common names .......Blackback flounder, lemon sole, black flounder Class ..... ..........

Osteichthyes Order ........ Pleuronectiformes 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).MORPHOLOGY/IDENTIFICATION AIDS 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

/CANADA JI CL.'U z/ MAINE PORTLAND A TL ANTIC OCEAJ Coastal distribution MILES 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 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 floun-der, Limanda ferruginea, the winter flounder has a. much straighter lateral line, a less concave dorsal head pro-file, 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 (Liop-setta putnami), but rough in the win-ter flounder.

Between the two, the winter flounder also has the greater number of anal fin rays.LIFE HISTORY Spawning The winter flounder spawns in coast-al 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 salin-ities of 31 to 32.5 ppt in inshore waters, and on Nantucket Shoal and Georges Bank at slightly higher salin-ities (32.7 to 33 ppt, respectively; Bigelow and Schroeder 1953). Water temperature during spawning is usually between 0 and 3 'C but may be as high as 6 'C (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 New-foundland, 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 mil-lion 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 Several tics that flounder flounders Atlantic (1980).morphological characteris-distinguish larvae of winter from' those of the other common in the western north were given by La Roche REASONS FOR INCLUSION IN THE SERIES By virtue of its abundance in estua-rine 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).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)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 of eggs 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 (Breder 1924). Incubation time Was 15 to 18 days at 2.8 to 3.3 0 C (Bigelow and Schroeder 1953), 25 days at 3 0 C, and 7 days at 12 to 14 'C (Rogers 1976).Incubation time was inversely related to water temperature and salinity (Rogers 1976).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 sur-vive, but are usually deformed (Rogers 1976). The optimal salinity for egg survival is 15 to 35 ppt.Many embryos become inviable or ab-normal at temperatures below freezing (-1.8 to 0 'C) and temperatures above 10 *C (Williams 1975). The optimum water temperature range for survival is 0 to 10 'C (Williams 1975).Larvae In studies by Bigelow and Schroeder (1953) and La Roche (1980), winter flounder larvae were 2.4 to 3.5 mm TL at the time of hatching.

A major characteristic of the newly hatched larvae was the broad vertical band of pigment cells dividing the postanal portion of the body. At a water tem-perature of 3.9 0 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.

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 com-plete when the larvae are 8 to 9 mm TL (Laurence 1975); this transformation took 80 days at a water temperature of 5 0 C and 49 days at 8 *C. No metamor-phosis was evident at 2 'C (Bigelow and Schroeder 1953).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, day-light feeders that cease feeding at night (Laurence 1977).Juveniles 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 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 (Mulkana 1966).Adults The seasonal movements of winter flounder differ between populations north and south of Cape Cod. A 5-year tagging study by Howe and Coates (1975) showed that winter flounder 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 Vine-yard 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 New-foundland, winter flounder remained in shallow water during summer as long as food was available and water tem-peratures did not exceed 15 'C (Van Guelpen and Davis 1979). Temperature is a less important factor in the dis-tribution of'juveniles, which tolerate higher temperatures than adults (Pearcy 1962).Indications are that a local popula-tion is defined by fish inhabiting.

several adjacent estuaries (Pierce and Howe 1977). Although a large percent-age of winter flounder in a tagging study were recaptured at or near the original tagging locations, Saila (1961) reported that the same breed-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, particularly in males (Lux 1973). After the first 2 years, females grow faster than males (Briggs 1965; Lux 1973; Howe and Coates 1975). An exception is in New-foundland, where the growth rates of the sexes are similar (Kennedy and Steele 1971).The growth rate also differs between fish from areas relatively close geo-graphically.

Lengths of flounder at the same age were significantly dif-ferent among certain bays on Long Is-land (Lobell 1939; Poole 1966).Flounder grow to a larger size in the Georges Bank population than in in-shore 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 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 1 published presented ength-weight for adults in Table 1.relationships and larvae are THE FISHERY Commercial and Recreational The winter flounder supports valu-able 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. Depart-ment 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.flounder.Published length-weight relationships for adult and larval winter Equation Location Source Adults logj 0 W = 3.138 log 1 oL-5.239 Georges Bank Lux (1969)where W = g, L = mm logj 0 W = 3.1441 logloL-2.072 (female)logjoW = 2.9833 log 1 oL-1.9041 (male) Newfoundland Kennedy and where W = g, L = cm Steele (1971)Larvae logjoW = 4.769 log 1 oL-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 com-mercial 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 commer-cial catch at age II1 (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 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 mor-tality 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 abun-dance in Narragansett Bay may be par-tially 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 after they hatch. They first copepods and phytoplankton, 3 weeks feed on but as 6 they reach metamorphosis, their diet is composed of copepod nauplii, small polychaetes, nemerteans, and ostra-cods. For detailed descriptions of the food habits of larval and juve-nile 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<0.1/ml; critical prey density was about 0.5/ml. Plankton density in-fluenced 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 per-centages of composition (numbers) of prey in flounder stomachs were as fol-lows: 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. (1973), Kennedy and Steele (1971), Olla et al. (1969), Mulkana (1966), and Frame (1973).Feeding Behavior Winter flounder primarily feed visu-ally 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 loca-tion, changing position from four to five times per minute (Olla et al.1969)..Competition The winter flounder has relatively few competitors for food and space.In many estuaries it is the most abun-dant demersal species (Richards 1963;Oviatt and Nixon 1973). The highly productive estuarine and coastal habi-tats it occupies, combined with its omnivorous food habits, tend to reduce competition.

Jeffries and Johnson (1974) suspected that the early spawn-ing and the short period of time to metamorphosis permit the larvae to reach the juvenile stage before po-tential competitors enter the bays and estuaries.

Predators Adult winter flounder are the prey of many of the larger estuarine and coastal predators such as striped bass (Morone saxatilis), bluefish (Pomato-mus saltatrix), goosefish americanus), spiny dogfish (Squalus acanthias), oyster toadfish (Opsanus tau), and sea raven (Hemitripterus americanus) (Dickie and McCracken 1955; Grosslein and Azarovitz 1982).Predation is a major cause of mor-tality in larval and juvenile winter flounder.

The larvae were heavily preyed upon by the small hydromedusa Saria tubulosa (Pearcy 1962). Tyler (1-971a) reported that the great cormorant (Phalacrocorax carbo), the great blue heron (Ardea heFo-dis), and the osprey (Pand-io haliaetus) 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 para-sites 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 win-ter flounder fed at water tempera-tures as high as 22 0 C, but burrow into the bottom at higher tempera-tures. McCracken (1963) gave a pre-ferred temperature range of 12 to 15 0 C. Huntsman and Sparks (1924)reported a maximum temperature toler-ance of about 30 0 C. Under controlled conditions, winter flounder can accli-mate to higher temperature regimes;for example, Everich and Gonzalez (1977) reported that the critical thermal maximum increased from 26 to 32 *C as the acclimation temperature increased from 4 to 23 0 C. An extend-ed period of unusually hot weather caused heavy mortality in coastal wa-ters of Long Island Sound (Nichols 1918). Juvenile winter flounder tend to be more tolerant of high tempera-tures than adults.Salinity Adult winter flounder commonly live in salinities of 5 to.35 ppt (Bigelow and Schroeder 1953). Extremes in sa-linity may lower egg and larval sur-vival and hatching success (see the section 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) were found in various tissues of the winter flounder (Smith and Cole 1970; Smith 1973). Concentrations of DDT, DOE, and heptachlor epoxide were highest in ripening ovaries. Agricultural runoff was the major source of the con-taminants (Smith and Cole 1970). Topp (1967) reported that this contamina-tion 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 pg/l sometimes caused high mortality of the eggs and yolk-sac larvae, and that exposure to 92 pg/l significantly increased egg mor-talities.

In contrast, Voyer et al.(1982) reported that silver in con-centrations up to 166 pg/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 asso-ciation with high sediment contamina-tion 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).8 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. 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 larval flounder.38:311-315.

1924.stages Bull.Some embryonic of the winter U.S. Bur. Fish.The .sports ter flounder in ng Island, N.Y.70.Briggs, P. 1965.fisheries for wint several bays of Lo Fish Game 12(1):48-7 Clayton, G., C.F. Cole, S.A'.,Murawski, and J.D. Parrish. 1978. Common marine 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. University of Massachusetts, Amherst. 109 pp.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. Soc. 104(1):13-19.

Howe, A.B., P.G. Coates, and D.Pierce. 1976. Winter flounder estuarine 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 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 bot-tom fishes in the Narrangansett Bay area: seven year variation in the abundance 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 of biological data for the winter flounder, 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-water U.S. Dep. Comm., Fish. Serv. Stat.Dickie, L.M., and F.D. McCracken.

1955. Isopleth diagrams to predict equilibrium yields of a-.small floun-der fishery. J. Fish. Res. Board Can. 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 young winter flounder Pseudopleuronectes americanus (Walbaum):

Feeding 9 I 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.Soc. 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 development of winter flounder, Pseudopleuronectes americanus (fami-ly Pleuronectes).

Fish. Bull.78(4):897-910.

Laurence, G.C. 1975. Laboratory growth and metabolism of the winter flounder (Pseudopleuronectes ameri-canus) from hatching through meta-morphosis 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 ameri-canus) 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 length-weight relationship for seven spe-cies 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.B. Scott. 1966.Fishes of the Atlantic coast of Can-ada. Fish. Res. Board Can. Bull.155. 485 pp.Lobell, M.J. 1939. A biological sur-vey of the salt waters of Long Is-land, 1938. Report on certain fishes. Winter flounder (Pseudo-leuronectes americanus).

Annu. Rep. N.Y. Conserv. Dep., Part I. No. 14:63-96.Lux, F.E. 1969. Length-weight tionships of six New England fishes. Trans. Am. Fish.98(4):617-621.

rel a-flat-Soc.Lux, F.E. 1973. Age and growth of the winter flounder, Pseudopleu-ronectes 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 floun-der, Pseudopleuronectes ameri-canus, off Massachusetts.

Trans.Am. Fish. Soc. 99(3):483-488.

MacDonald, J.S. 1983. Laboratory observations of feeding behavior of the ocean pout (Macrozoarces ameri-canus) and winter flounder (Pseudo-pleuronectes americanus) with ref-erence to niche overlap of natural populations.

Can. J. Zool.61(3):539-546.

McCracken, F.D. 1963: Seasonal move-ments of the winter flounder, Pseu-doeleuronectes americanus, on t-e Atlantic coast. J. Fish. Res. Board 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.ter founder and 55:36-39.An abnormal win-others. Copeia Olla, B.L., R. Wicklund, and S. Wilk.1969. Behavior of winter flounder in a natural habitat. Trans. Am.Fish. Soc. 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 abun-dance. Estuarine Coastal Mar. Sci.1:361-378.

Pearcy, W.G.estuarine 1962. Ecology of an population of winter 10 A flounder Pseudopleuronectes ameri-canus (Walbaum).

Bull. B1ng-ham Oceanogr.

Collect. Yale Univ. 18(1).78 pp.Perlmutter, A. 1947. The blackback flounder and its fishery in New England and New York. Bull. Bingham Oceanogr.

Collect. Yale Univ. 11(2).92 pp.Pierce, D.E., and A.B. Howe. 1977. A further study on winter flounder group identification off Massachu-setts. Trans. Am. Fish. Soc.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 South Bay, New York. Trans. Am.Fish. Soc. 98(4):611-617.

Richards, S.W. 1963. The demersal fish populations of Long Island Sound. Bull. Bingham Oceanogr.

Yale Univ. 18(2). 101 pp.Rogers, C.A. 1976. Effects of temp-erature and salinity on the survi-val 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. Ocean-ogr. 6:292-298.

Saila, S.B., D.B. Horton, and R.J.Berry. 1965. Estimates of the the-oretical biomass of juvenile winter flounder, Pseudopleuronectes ameri-canus (Walbaum), required for a Tish-ery in Rhode Island. J. Fish.Res. Board Can. 22:945-954.

Sherwood, M.J. 1982. Fin erosion, liver condition, and trace con-taminant exposure in fishes from three coastal regions. In G.F.Meyer, ed. Ecological stress7nd the New York management.

Foundation, Bight: science and Estuarine Research Columbia, S.C.Smith, R. 1973. Pesticide residues as a possible factor in larval win-ter flounder mortality.

Pages 173-180 in Proceedings of a workshop in egg,7arval, 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, Pseudo-eleuronectes americanus, from--th-e eweantic River tuary, Massachu-setts. 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 floun-der, Pseudopleuronectes ameri-canus. Biol. bull. (Woods Hole)Pi--71-387.

Sullivan, W.E, 1915. Description of the young stages of the winter flounder (Pseudopleuronectes ameri-canus Walbaum. Trans. Am. Fish.Soc. 44:125-136.

Topp, R.W. 1967. An estimate of fe-cundity of the winter flounder, Pseudopleuronectes americanus.

J.Fish. Res. Board Can. 25(6):1299-1302.Tyler, A.V. 1971a. Periodic and res-ident components in communities of Atlantic fishes. J. Fish. Res.Board Can. 28:935-946.

Tyler, A.V. 1971b. Surges of winter flounder Pseudopleuronectes ameri-canus into the intertidal zoneT F.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 rela-tion to meal frequency in winter 11 flounder, Pseudopleuronectes ameri-canus, wit hypothesis regarding population 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 in-shore locations in Newfoundland.

Trans. Am. Fish. Soc. 108(1):26-37.

Voyer, R.A., J.A. Cardin, J.F.Heltsche, and G.L. Hoffman. 1982.Viability of embryos of the winter flounder Pseudopleuronectes ameri-canus exposed to mixtures of cadmium and silver in combination with se-lected fixed salinities.

Aquat.Toxicol. 2:223-233.

Wells, B., D.H. Steele, and A.V.Tyler. 1973. Intertidal feeding of winter flounder, Pseudopleu-ronectes americanus, in the Bay of Fundy. J.. Fish. Res. Board Can.30:1374-1378.

Williams, G.C. 1975. Viable embryo-genesis of the winter flounder (Pseudopleuronectes americanus) from-1. 8 to 1.5 -C. Mar_. BolT. (Berl.)33(1): 71-74.Ziskowski, J., and R.A. Murchelano.

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

12 qn,,, .Ini 50272-1., 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 1989 of Coastal Fishes and Invertebrates (North Atlantic)--

Winter Flounder 7. Adthor(s)

Jack Buckley 6.

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 Station Research and Development U.S. Army Corps of Engineers National 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 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 'C and they usually spawn at 0 to 3 'C. 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. Descriptors Estuaries Fishes Salinity Fisheries Life cycles Contaminants Feeding habits Temperature

b.

Terms Winter flounder, Environmental requirements Pseudopleuronectes 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) of Commemce TAKE PRIDE iniAmerica DEPARTMENT OF THE INTERIOR U.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 includes fostering 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.

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