Distribution of Pelagic Fishes in Sheepscot River-Back River Estuary, Wiscasset, Maine" (1973)

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Marine Sciences Faculty Scholarship School of Marine Sciences

1-1-1973 Distribution of Pelagic Fishes in Sheepscot River- Back River Estuary, Wiscasset, Maine C. W. Recksiek

James McCleave University of Maine - Main, [email protected]

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Repository Citation Recksiek, C. W. and McCleave, James, "Distribution of Pelagic Fishes in Sheepscot River-Back River Estuary, Wiscasset, Maine" (1973). Marine Sciences Faculty Scholarship. 38. https://digitalcommons.library.umaine.edu/sms_facpub/38

This Article is brought to you for free and open access by DigitalCommons@UMaine. It has been accepted for inclusion in Marine Sciences Faculty Scholarship by an authorized administrator of DigitalCommons@UMaine. For more information, please contact [email protected]. Distribution of Pelagic Fishes in the Sheepscot River-Back River Estuary, Wiscasset, Maine

CONRADW. RECKSlEK AND JAMES D. McCnEAvE Department of Zoology, University of Maine, Orono, Maine 04473

ABSTRACT Fifteen speciesof pelagic fishes were collectedin 156 gill net sets at eight locationsin the SheepscotRiver-Back River estuary, Wiscasset,Maine, June 1970 through December 1971. Highestcatches occurred June through August. Only the rainbowsmelt is a year-roundresident. Differencesin abundancein space and time are apparently related to temperature. During the summer,alewives, blueback herring, and Atlantic menhadenwere most abundantin the relatively warm Back River estuary, while Atlantic herring, Atlantic mackerel, and spiny dogfish were most abundant in the more oceanic SheepscotRiver estuary. Prolonged near-freezingtempera- turesapparently limit the time pelagicfishes spend in the estuaryand limit the number of species which can inhabit it. It is hypothesizedthat the distribution of pelagic specieswhich exhibited preferencesfor colderwater, suchas Atlantic herring, would be most affectedby artificial warming of the surfacewaters of the Back River estuary,if a new atomic poweredgenerating plant were allowed to dischargeheated effluent directly into it.

INTRODUCTION stant, temperature usually represents the This study was designedto determinethe master factor controlling . . . distribution. In coastal areas and estuaries distribution relative seasonalabundance of pelagic fishes in the Sheepscot River-Back River estuary, depends increasingly on both factors, with Wiscasset,Maine, and to evaluate the effects varying degreesof dominance of either one." of samplinglocation, temperature, and salinity The effects of hydrographic conditionsupon distribution of North Atlantic fishes have been on the occurrenceof principal species. This investigationis part of an effort by several treated extensivelyin a symposium(ICNAF, 1965). The influence of temperature on the investigators to provide basic ecological information prior to the completionof a 855- distribution of fishes was discussedby Brett megawatt atomic electric power plant. (1970). Hela and Laevastu (1962) sum- Fish distribution studies from North Amer- marized known temperature optima for several ican east coast inshore areas and estuaries species. The relation of estuarine organisms have dealt with both pelagic and demersal to salinity is reviewed by Pearse and Gunter components. Seasonal occurrence of fishes (1957) and Gunter (1961). has been studied in a Georgia salt marsh DESCRIPTION OF STUDY AREA (Dahlbergand Odum, 1970), in shorehabitats near Beaufort, North Carolina (Tagatz and Stickney(1959) describedthe hydrography Downloaded by [University of Maine] at 11:04 22 December 2011 Dudley, 1961), on Virginia's eastern shore and geographyof the SheepscotRiver estuary (Richardsand Castagna,1970), Chesapeake and its relation to the Kennebec River estuary. Bay and York and PamunkeyRivers (Mass- The Back River estuaryis separatedfrom the mann, 1962), in the Delaware River estuary SheepscotRiver by Westport Island (Fig. 1). (de Sylva, Kalber, and Shuster, 1962), the A narrow causeway connectsthe island and surf waters of Long Island (Schaefer, 1967), the mainland. The atomic power plant is the Mystic River estuary (Pearcy and Rich- located near sampling station G2 where Back ards, 1962), near WoodsHole, MassachusettsRiver widens into Montsweag Bay. Except (Lux and Nichy, 1971), in the Annisquam for a navigable channel from HockomockBay River-GloucesterHarbor (Massachusetts) to the causewayand from near G3 into Mont- coastalsystem (Jerome, Chesmore, and Ander- sweagBrook, MontsweagBay is shallow and son, 1969), and PassamaquoddyBay (Tyler, almost exposed at low tide. 1971). Kinne (1964) statedthat: "In the The flood tide into Back River and Mont- open ocean where salinities are rather con- sweag Bay flows south through the narrow 541 542 TRANS. AMER. FISH. SOC.,1973, NO. 3

causewayopening and north through Hocko- mockBay. Tidal currentsfrom Berry Island (G1) to aroundG2 are sufficientlyweak to HIGHWAYpermit seasonal icing over. At G3 strongtidal currents follow the channel. Near G4 tidal currentsare mild. Localtidal currents,created by large eddies,are irregular and unpredict- able. At G5 the currents are powerful and

UDY follow the channel. Near G6 in Cross River AREA tidal currentdirections vary greatlywith tidal stage. o 1

KILOMETERS MATERIALS AND METHODS Sampling stations were selected near and at increasingdistances from the power plant BERRY to determinethe distributionof pelagicfish G1 POWER fauna; however station selection was influenced by tidal currents and weather considera- tions. StationsG1, G2, G3, and G7 are in the BackRiver estuary,and stationsG4, G5, G6, andG8 are in the SheepscotRiver estuary (Fig. 1). Pelagic fish were sampled with cross- channel, anchored gill nets fished from a bow roller on a 5-m fiberglassskiff (Boston Whaler). The gill nets were of three mono- filamentnylon panels, each panel 2.4 m (8 ft) x 15.2 m (50 ft) with 5.1 cm (2 in), 1.3 cm (•A in), 2.5 cm (1 in) bar mesh.The sinking gill net was suspendedso as to fish 1.9 m (6 ft) below the surface. The efficiency of gill nets is reduced in strong currents and winter weather conditions prevented regular sampling in open areas. Thus south Montsweag Bay and the main channelof the lower SheepscotRiver estuary were not sampledbecause of strong currents. Six stations (G1-G6) were sampledthrough- Downloaded by [University of Maine] at 11:04 22 December 2011 out the study. G8 was exposed to winter weather, and served only as a supplemental station during the summer of 1971. G7 was sampled the same summer. Regular sampling was conductedfrom June 1970 to December 1971 during neap tides after full moon. During the summer,samples were collectedfortnightly on the neap tides. Two consecutivenights (winter) or three consecutivenights (summer) were neededto fish all stations. The first station net was set at about 1500 EST and was retrieved at about F•cuaE 1.--The SheepscotRiver-Back River estuary, Wiscasset,Maine, showinggillnetting stations 0700 EST the next morning. Remainingnets G1 throughG8. Contourdepth: 3 m. were set and retrieved as soon as possible RECKSIEK AND McCLEAVE--PELAGIC FISH DISTRIBUTION IN MAINE 543

thereafter. Samplingwas suspendedin Back TABLE 1.--Percentage compositionby numbers and weights o] the pelagic fish catchesin Sheepscot River from Januarythrough March 1971 and River-Back River estuary, June 1970 through De- in the SheepscotRiver in January 1971 due cember 1971 to ice conditions. Percent Percent A water sample was collectedat a depth of total of total numbers weight of 1.9 m at the beginning and conclusionof of catch of catch eachset by meansof a Meyer sampler. The Alewife (Alosa pseudoharengus) 41.5 21.9 two sampleswere mixed and the salinity deter- Blueback herring (Alosa aestivalis) 23.1 15.3 Atlantic herring (Clupea harengus) 21.9 3.1 mined by hydrometer. A continuousrecord Rainbow smelt (Osmerus mordax) 5.7 1.5 Atlantic menhaden (Brevoortia tyrannus) 3.6 16.4 of temperature was obtained from a thermo- Atlantic mackerel (Scornbet scombrus) 2.0 3.4 Spiny dogfish (Squalus acanthias) 1.4 34.8 graph suspendedat a depth of 1.8 m. American shad (Alosa sapidissima) 0.4 0.6 The catches were fixed in 10% formalin for Striped bass (Morone saxatilis) 0.2 2.7 ButterfishSilverhake (Peprilus(Merluccius triacanthus)bilinearis) ] at least 10 days and preservedfor study in 0.3 50% 2-propanol. Striped bass and spiny dog- Hickory shad (Alosa mediocris) Pollock (Pollachius vireos) fish were studied fresh. The numbers and BluefishWhite perch(Pomatomus (Motone saltatrix)americana) i 0.2 total weight of each specieswere recorded. The 1.3-cm and 2.5-cm mesh selected specific sizesof Atlanticherring, alewife,and blueback RESULTS AND DISCUSSION herring. These specieswere separatedon the Fish Catches in Sheepscot basisof size into two groups and the numbers and Back Rivers and weights recorded. Catches,expressed as both numbers and This study yielded 9,840 individualsof 15 weights,of thosespecies which were frequently pelagicspecies in 156 gill net sets (Table 1).x encounteredat G1, G2 and G4 were treated Six speciesof demersal fishes, outside the statistically. Catchesat the other stationswere scopeof this study, were occasionallycaught not used for quanitative comparisonssince in gill nets. tidal currents were believed to have decreased Most of the common pelagic fish speciesof the efficiency of the gill nets. The experi- the SheepscotRiver-Back River estuary were mental model was a two-factor randomized encounteredMay throughNovember, with the complete-blockdesign. Analysis of variance largestcatches occurring June through August was employedin testing the null hypotheses (Fig. 2). With the exceptionof rainbowsmelt that the effects of sampling location (factor and perhapswhite perch, they leave the estu- 1, 3 levels; G1, G2, G4), months (factor 2, ary during the winter months. 7 levels; Junethrough December), interaction Differences between the Sheepscot River betweenmonths and samplinglocations, and and Back River faunas were particularly strik- year (blocks,2 levels; 1970, 1971) were non- ing during the summer (Fig. 2). Catchesin significant. In summer, when two samples the Back River estuary, in terms of both

Downloaded by [University of Maine] at 11:04 22 December 2011 were collectedper month, only thosecollected numbers and weights, were dominated by during the neap tides after full moon were alewivesand blueback herring. Atlantic men- usedin this analysis.Duncan's new multiple- haden contributedmarkedly to the weight of range test was used to test for significant the catches. In the Sheepscot River estuary differences among main effect levels when Atlantic herring were dominant numerically and spiny dogfish dominated by weight. At interaction was nonsignificant. When interac- G6 in Cross River spiny dogfish even dom- tion wassignificant, as wasthe caseonly with inated numerically. Large catchesof Atlantic numbers and weights of Atlantic menhaden, mackerel at G8 in the lower SheepscotRiver Duncan's test was performed separatelyfor estuary in summer suggestthat this fish may each samplinglocation to detectdifferences in play an important role in the ecologyof the time. Analysesof like designwere conducted lower estuary. for summer data with catches during fortnightly neaptides in 1970 and 1971 (factor 2, x A table of catch numbers and weights for all eight stationsand all sampling periods is available from 6 levels;June through August). the junior author upon request. 544 TRANSß AMER. FISHß SOC., 1973, NO. 3 12t •l Spinydogfish Bluebackherringt300

-200

4 -100

300- Alewife American shad -6

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100 - -2 ß"'""..:'"'".•:'"'

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40-601 Atlantic menhaden,.: Atlantic herring.?•9'-2OO

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FZCURE2.--Seasonal abundance of the major pelagic fish speciesat one station in Back River estuary (G2 ), one in the Upper SheepscotRiver estuary(G4 - ß '), and one in the lower SheepscotRiver estuary(G6 ---), June 1970to December1971. RECKSIEK AND McCLEAVE--PELAGIC FISH DISTRIBUTION IN MAINE 545

The statistical analyses support the infer- TaSLE Z--Significant di//erences in mean numbers o/ pelagic fish collectedby gillnetting at stations ence that the summertime pelagic fish fauna G1, G2, and G4 in the SheepscotRiver-Back River of the Back River estuaryis distinctfrom that estuary, Wiscasset,Maine as determinedby Dun- of the Sheepscot.Analyses of weightsof fishes can'snew multiple-rangetest (P < .05). Data /or 1970 and 1971 and /or monthly samplingperiods caught gave essentially the same results as June throughDecember, and [ortnightly sampling those for numbers. For all speciesanalyzed, periodsJune throughAugust combined. Means not with the exceptionof rainbow stock, catches underscoredby the same line are significantly diJJerent at the two stations,G1 and G2, in the Back River estuary were never significantly differ- Species Monthly Fortnightly

ent from each other (Table 2). During the Spiny dogfish G4 G1 Gg G4 G1 G2 summer,significantly more alewives,blueback Blueback herring G2 G1 G4 Gg G1 G4 herring, and Atlantic menhadenwere cap- Alewife G2 G1 G4 Gg G1 G4 American shad G1 G2 G4 G1 G4 G2 tured in Back River than at G4 in the Sheep- Atlantic menhaden Duncan's test G1 G2 G4 not performed scot River estuary. Significantly more spiny Atlantic herring G4 G1 G2 G4 G2 G1 dogfish and Atlantic mackerel were collected Rainbow smelt G2 G4 G1 G4 G2 G1 at G4 in the Sheepscotthan at the Back River Atlantic mackerel G4 G1 • G4 G1 G2 estuary stations. Differences in the catch of Atlantic herring were not statistically signifi- if the link betweensalinity and distribution is cant, yet 75% of all Atlantic herring collected direct (Holliday, 1971). Some inferencescan were from G4 in the SheepscotRiver estuary. be drawn from our study on the roles of Differences betweenspecies composition of temperature and salinity on the abundancein fishes caught in the Back River and Sheepscot time and spaceof severalpelagic fishes in the River estuaries became less distinct through estuary. The effects on fish distribution of the fall and during the spring. During this artificially high temperatures,which would time Atlantic herring appearedmore frequently result from unrestricted power plant opera- at the Back River stations,and Atlantic men- tion, can also be hypothesized for some of haden were caught in the SheepscotRiver the dominant pelagic species. estuary in the late spring. About 99% of the spiny dogfish were col- For the speciesanalyzed, differences in lected at water temperaturesbetween 8 C and catches between 1970 and 1971, with the 16 C. On a catch-per-net-setbasis preferences exceptionof American shad, were nonsignifi- for temperaturesbetween 10 C and 14 C (Fig. cant. Significantly more American shad were 4) and salinities above 28•o (Fig. 5) are caught during 1971 than 1970. evident.Jensen (1966) concludedfrom survey The stationsin the SheepscotRiver estuary data that "dogfish in the Northwest Atlantic were more saline than those in the Back River prefer bottom water between 7.2 C and 12.8 estuary (Fig. 3). During the summer G8 in C." Leim and Scott (1966) statedthat spiny the lower Sheepscotand G6 in Cross River dogfish do not appear until the temperature

Downloaded by [University of Maine] at 11:04 22 December 2011 were the coldest,followed by G4 and G5 near reaches 5.6 C and leave if it exceeds 15.6 C. Wiscasset,followed by stations in the Back Their virtual exclusion from surface waters River-MontsweagBay area (Fig. 3). In the of the Back River estuary suggeststhat these winter the temperatureregime reversed,i.e., waters constitute a hostile environment the Back River stations were coldest followed throughoutthe year: too cold in winter, too by G5 and G4 in turn followed by G6. warm in summer, and not saline enough in spring. In the lower SheepscotRiver estuary Role o[ Temperatureand Salinity where salinitiesare high throughoutthe year, The effects of temperature on fish distribu- dogfish are found only when temperatures tion have been emphasized(Hela and Lae- exceed about 8 C. vastu, 1962; Kinne, 1964; de Sylva, 1969) Atlantic herring show a distinct preference often to the near exclusionof salinity. How- for temperaturesbetween 10 C and 16 C (Fig. ever, the latter may be a most important factor 4). Since they were captured over a wide in distributionof estuarineorganisms (Gunter, range of salinities (Fig. 5), temperature is 1961), but it is often difficult to determine apparently a more important factor influenc- 546 TRANS. AMER. FISH. SOC., 1973, NO. 3

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I I I I I I I I I I I I I I I I I I I I d d A S O N D d F M A M d d A 5 O N D RECKSIEK AND McCLEAVF_,•PELAGIC FISH DISTRIBUTION IN MAINE 547

ing distribution. The scarcity of Atlantic 20 C) and they were not caught at low tem- herring in the Back River estuary during the peratures (Fig. 4). All three specieswere summer implies that their distribution is prac- caught over a wide range of salinities (Fig. tically limited to temperaturesunder 16 C. 5), but peak catcheswere in the 225• to 265• Exposure for 14 hr to temperatures above range. All three were more abundant in the 23 C were lethal to yearling Atlantic herring Back River estuary than in the Sheepscot acclimatedto 15 C (Blaxter, 1960). Exposure during the summermonths, which suggestsa for 48 hr to 21.2 C caused 50% mortality in preferencefor warmerwater. Graham(1956) Atlantic herring held at 8 C-11 C (Brawn, reporteda tolerancelimit (the water tempera- 1960). Stickney(1969), in a laboratorystudy ture extreme permitting 50% survival after of orientation of juvenile Atlantic herring to 40 hours) of 23 C in yearling alewives ac- temperature and salinity, establisheda tem- climated to 9 C. Trembley (1960) found an perature preferenceof 12 C in fish acclimated upper tolerance limit range of 26.7 C to 32.2 to 15C. B•rzing (1949) found that most C. Marcy, Jacobson,and Nankee (1972) re- small Atlantic herring, in the Gulf of Riga, cently reported field studies which suggest were caught between 8 C and 12 C. In this that the upper natural temperature limit of study Atlantic herring were relatively scarce young shad, and probably blueback herring at all stationsduring August,and thosecaught as well, may be about 30 C. Alewives,blue- were most frequently in the cooler Sheepscot back herring, and shad were caught in about River estuary. Under natural conditions the the same relative percentages over the same Back River estuary during the warmer months temperature intervals, and since over 90% is a marginal habitat for Atlantic herring. were caught above 14 C and in the warmer Any pronouncedwarming of the Back River (summer) Back River estuary, it is apparent estuary will likely exclude Atlantic herring that they should tolerate more artificial warm- from the area. ing of the habitat than species more typical An optimum temperature range of 12 C to of the SheepscotRiver estuary. Since no gill 14 C has been reported for Atlantic mackerel nets were set in water above 20 C, a definite (Dannevig, 1955), and the highest tempera- preference range cannot be stipulated, yet ture at which they commonly occur is 20 C 16 C to 20+ C seems reasonable. (Bigelow and Schroeder, 1953). Although Most Atlantic menhadenwere caught in the Atlantic mackerel were commonly caught at Back River estuary during the summer sug- temperatures of 18 C to 20 C in this study, gestinga preferencefor warmer water (Fig. most were caught in the Sheepscot River 4). Their distribution seemsless influenced estuary, not in Back River. This tendency by salinity (Fig. 5). Reinties (1960) stated would probably be strengthened during the that the movement of Atlantic menhaden north warmer months should temperaturesrise over in spring and south in autumn along the 20 C. Sette(1950) concluded that watercolder Atlantic coast coincided with the seasonal shift than 7 C or 8 C forms a temperature barrier of the 10 C isotherm. Only about 7% of the Downloaded by [University of Maine] at 11:04 22 December 2011 to northward springtime advance of the Atlan- Atlantic menhadenin this study were caught tic mackerel contingent which summers in the below 10 C. Although they were caught over Gulf of Maine. Our observations substantiate a wide range of temperatures(Fig. 4), there this conclusion as we caught no mackerel at were peaks in abundancein August and Sep- temperaturesbelow 9 C. tember in the Back River estuary (Fig. 2). Bluebackherring, alewives,and American During July through Septembertemperatures shad all showed a peak in catch-per-net-set in Back River were consistently within the at the highesttemperatures sampled (18 C to 15 C to 20 C preferred temperature range re-

F•½ua•.&--Average salinity and temperatureduring gillnetting at stationsG1, G2, G3 ( ), G4, G5 (- ß ß), and G6 (---) in the SheepscotRiver-Back River estuary,Wiscasset, Maine, June 1970 through December1971. Stationssignificantly different (Duncan'snew multiple-rangetest, P < .05) drawn with different symbol. 548 TRANS. AMER. FISH. SOC., 1973,NO. 3

Spiny Blueback 5O dogfish herring -- 40

10 • I I., , ,' ,

Alewife American 5O shad 40

Atlantic Atlantic 50 a. menhaden herring 4O

Rainbow Atlantic 50 smelt mackerel 40

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0 4 8 12 16 20 4 8 12 16 20 TEMPERATURE' øC

FzcuaE4.--Percentages of averagecatches (per gill net set) per 2 C temperatureinterval for the major pelagic fish speciesof the SheepscotRiver-Back River estuary,June 1970to December1971. All stationsand dates combined.

ported for menhadenby Goode (1879). A of Atlantic menhaden are difficult to predict. preferencefor 15 C to 20+ C water seemscon- It seems unlikely that moderate increases firmed by this study, but since nets were not would alter their distribution in the estuary. set at temperatures above 20 C, effects of Rainbow smelt were collected in relatively artificially warmed water on the distribution small numbers over the widest temperature RECKSIEK AND McCLEAVE•PELAGIC FISH DISTRIBUTION IN MAINE 549

Spiny Blueback --60 dogfish herring --50

--40

•-30

-20

Alewife American 30 shad --20

•o•

r• Ld Atlantic 1 Atlantic --30•. menhaden herring 1 --20

Rainbow Atlantic --40

smelt mackerel --30

14 18 22 26 30 14 18 22 26 30 SALINITY ø/oo

FIGURE5.--Percentages of averagecatches (per gill net set) per2%0 salinity interval for the majorpelagic fish speciesof the SheepscotRiver-Back River estuary,June 1970to December1971. All stationsand dates Downloaded by [University of Maine] at 11:04 22 December 2011 combined.

range (0 C to 20 C, Fig. 4) and the widest Comparison With Other Studies salinity range (145• to 315c, Fig. 5) in both Although direct comparisonsof the results the SheepscotRiver and Back River estuaries. of this study with thoseof othersare difficult Since a tolerance limit of 21.5 C to 28.5 C due to dissimilaritiesof gear, somegeneraliza- depending on acclimation temperature has been reported by Huntsman and Sparks tions can be made. Tyler (1971) concluded (1924), it seems unlikely that a moderate from his trawling study of the fish community tem•rature increase will adversely affect in a deepportion of PassamaquoddyBay, New smelt distribution in Back River. Brunswick,that "in temperate regions, in- For the other pelagic species,the outcome shore, deepwater fish communities are made of any artificial warming cannot be predicted up of regular, summer, and winter periodic due to their scarcity in the gill net catches. components. There are also numerous occa- 550 TRANS. AMER. FISH. SOC., 1973, NO. 3

sional species. Spring and fall seasonsare Yet if a sampling schemesimilar to the one transitional with regard to fish fauna. Forma- in the present study had been used, more tion of temporal componentsis largely related speciesmay have qualified. to temperatureregime,... At latitudes where Evidently, prolonged near-freezingtempera- there is relatively greater annual temperature tures rather than yearly temperature range fluctuation, there are proportionately more most severely limit the occupation of tem- speciesin the temporal componentsand fewer perate estuariesby pelagic fish. Virtually all in the regular component." The results of this the pelagic fish leave the SheepscotRiver- study support Tyler's conclusionsthat forma- Back River estuaryDecember through March. tion of temporal componentsis largely related The number of species which are able to to temperatureregime, and that with greater occupythe estuary is likewise affected. annual temperature fluctuation, more species are in the temporal component than in the ACKNOWLEDGMENTS regular component. The pelagic fish fauna in We thank Mrs. Shirley Recksiek who as- the SheepscotRiver-Back River estuary,where sisted in the field and helped complete the annual temperature variation is greater than manuscript; and Dr. Hugh H. DeWitt and in nearby PassamaquoddyBay, is wholly tem- Messrs. Mason D. Bryant, Paul C. Jensen, poral with the exception of rainbow smelt William J. Hauser, Timothy E. Targert, and (Fig. 2). In addition,there is no species,with StephenM. Fried for helpful suggestionsand the possible exception of Atlantic herring, aid in the field. Researchfunds were provided which can be termed a "winter periodic." It by the Maine Yankee Atomic Power Company. would be more properly termed a "spring/fall

periodic" in this estuary. LITERATURE CITED Norden (1966) and Fox and Mock (1968), in their studies of seasonal occurrence of fishes Bf.RZlN•, B. 1949. Dber temperaturebedingteTier- wanderungenin der Ostsee. Oikos 1: 29-33. in two Louisiana bays, encounteredannual Cited in I. Hela and T. Laevastu, Fisheries hy- temperatureranges of 29 C (6 C to 35 C) and drography.Fishing News (Books) Ltd.• London. p. 27. 25 C (10 C to 35 C), respectively.These ranges Bigsnow, H. B., ANDW. C. Scnao•.D•.a. 1953. Fishes are not much greater than the range in the of the Gulf of Maine. U.S. Fish Wildl. Serv., Back River estuary (Fig. 3). Fox and Mock Fish. Bull. 53: 1-577. (1968) in Barataria Bay, found the number BLAXT•.a,J. H.S. 1960. The effectof extremesbf temperatureon herringlarvae. J. Mar. Biol. Ass. of fish and fish speciesto be lowest during U.K. 39: 605-608. Cited in P. L. Altman and winter and highest during summer. Norden's D. S. Dittmer (eds.), Environmental biology. Fed. Amer. Soc. Exp. Biol., Bethesda,Maryland. (1966) data from Vermilion Bay exhibitedthe p. 73. same trend. He stated that "water tempera- BaxwN, V. M. 1960. Temperature tolerance of un- ture, rather than salinity, appears to exert a acclimated herring (Clupea harengus L.). J. greater influence on the seasonal movements Fish. Res. Bd. Canada 17: 721-723. Ba•.TT,J. R. 1970. Temperature.Animals. Fishes. of fishesin and out of Vermilion Bay." Tem- Functionalresponses. 516-560 p. In O. Kinne Downloaded by [University of Maine] at 11:04 22 December 2011 poral components dominated the Louisiana (ed.), Marine ecology. Vol. 1, Part 1. John pelagic fish faunas which were generally more Wiley & SonsLtd., London. DanLa•.aG,M.D., aN• E. P. O•u•a. 1970. Annual diversethan thoseof the presentstudy. cycles of species occurrence, abundance, and Richardsand Castagna(1970), in a beach diversityin Georgia estuarinefish populations. seine and trawl survey in inlet, mid-channel, Amer. Midi. Natur. 83: 382-392. DANNY.rIG, A. 1955. Mackerel and sea temperature. inshore beach, and tidal creek habitats of Measurements--21April to 15 May, 1952. Prak- Virginia's eastern shore, recorded tempera- tiske Fiskeforsok,1952. Arsber. NorgesFisk. 5: 64-67. Cited in I. Hela and T. Laevastu, Fish- tures from 4.0 C to 31.5C. The temporal eries hydrography.Fishing News (Books) Ltd., component dominated the pelagic fauna. London. p. 21. Several pelagic speciesencountered could be Fox, L. S., A• W. R. Mock:, Ja. 1968. Seasonal occurrence of fishes in two shore habitats in properly termed winter periodics. Catches Barataria Bay, Louisiana. Proc. Louisiana Acad. were heaviest during the summer months. $ci. 31: 43-53. They classifiedonly one pelagic fish, the bay Goo•., G.B. 1879. The natural and economical historyof the Americanmenhaden. U.S. Comm. anchovy (Anchoa mitchilli) as a resident. Fish and Fish., Pt. 5, Rep. Comm.1877 (Append- RECKSIEK AND McCLEAVE--PELAGIC FISH DISTRIBUTION IN MAINE 551

age A): 1-529. Cited in J. W. Reintjes. 1969. linities, and fishes collectedduring trawl surveys Synopsisof biologicaldata on the Atlantic men- of ChesapeakeBay and York and Pamunkey haden, Brevoortia tyrannus. U.S. Fish Wildl. Rivers 1956-1959. Virginia Inst. Mar. Sci. Spec. Serv., Food Agr. Organ. Fish. Synop. No. 42, Sci. Rep. 27: 1-3. Circ. 320. p. 5. NORDEN,C. R. 1966. The seasonaldistribution of GRAHAM,J. J. 1956. Observationson the alewife, fishes in Vermilion Bay, Louisiana. Wisconsin Pomolobuspseudoharengus (Wilson), in fresh Acad. Sci. Arts Letters 55: 119-137. water. Pub. Ontario Fish. Res. Lab., LXXIV, PEABeY,W. G., AND S. W. RICHARDS.1962. Dis- Univ. Toronto Biol. Ser. No. 62, 43 p. Cited in tribution and ecology of fishes of the Mystic P. L. Altman and D. S. Dittmer (eds.), Environ- River estnary,Connecticut. Ecology43: 248-259. mental biology. Fed. Amer. Soc. Exp. Biol., PEARSE,A. S., AND G. GUNTER. 1957. Salinity, p. Bethesda,Maryland. p. 78. 129-157. In J. W. Hedgepeth(ed.), Treatiseon GUNTER,G. 1961. Somerelations of estuarineorgan- marineecology and palcoecology,Vol. 1. Ecology. isms to salinity. Limnol. Oceanogr.6: 182-190. Mere. Geol. Soc. America 67. HELA,I., ANDT. LAEVASTU.1962. Fisherieshydrog- REINTJES,J.W. 1969. Synopsisof biological data raphy. Fishing News (Books) Ltd., London. on the Atlantic menhaden,Brevoortia tyrannus. 119 p. U.S. Fish Wildl. Ser., Food Agr. Organ. Fish. HOLLIDAY,F .G.T. 1971. Salinity. Animals. Fishes, Synop. No. 42, Circ. 320. 30 p. p. 997-1083. In O. Kinne (ed.), Marine ecology. RICHARDS,C. E., AND M. CASTAGNA.1970. Marine Vol. I. Part 2. fishes of Virginia's eastern shore (inlet and HUNTSMAN,A. G., ANDM. I. SPARKS.1924. Limit- marsh, seaside waters). Chesapeake Sci. 11: ing factorsfor marine animals. 3. Relative resis- 235-248. tance to high temperatnres. Contr. Can. Biol. SETTE,O. E. 1950. Biology of the Atlantic mackerel Fish., Nova Scotia, 2: 97-114. Cited in P. L. (Scomber scombrus) of North America. II. Altman and D. S. Dirtmet (eds.), Environmental Migrations and habits. U.S. Fish Wildl. Serv., biology. Fed. Amer. Soc. Exp. Biol., Bethesda, Fish. Bull. 51: 251-358. Maryland. p. 77. SCHAEFER,R. H. 1967. Speciescomposition, size INTERNATIONAL COMMISSION FOR THE NORTHWEST and seasonal abundance of fish in the surf waters ATLANTIC FISHERIES. 1965. ICNAF environmen- of Long Island. New York Fish Game J. 14: tal symposimn. Spec. Pnb. No. 6. Headquart. 1-46. Comm.,Dartmonth, Nova Scotia,Canada. 914 p. STICKNEY,A. P. 1959. Ecology of the Sheepscot JENSEN,A.C. 1966. Life historyof the spiny dog- River estuary. U.S. Fish Wildl. Serv. Spec. Sci. fish. U.S. Fish Wildl. Serv., Fish. Bull. 65: Rep., Fish. 309: 1-21. 527-579. 1969. Orientationof juvenile Atlantic her- JEROME,W. C., JR., A. P. CHESMORE,AND C. O. ring(Clupea harengus harengus L.) totempera- ANDERSON,JR. 1969. A study of the marine ture and salinity. In Food Agr. Organ.Fish. Rep. resources of the Annisquam River-Gloucester 2: 323-342. Harbor coastalsystem. Monogr. Ser. No. 3. Mas- SYLYA,D. P. DE, F. A. KALBER,JR., AND C. N. sachusettsDiv. Mar. Fish. 62 p. SHUSTER,JR. 1962. Fishes and ecological con- KINNE, O. 1964. The effects of temperatnre and ditions in the shore zone of the Delaware River salinity on marine and brackish water animals. estuary,with notes on other speciescollected in II. Salinity and temperature salinity combina- deeper water. Inf. Ser. Univ. Delaware, Publ. tions. Oceanogr.Mar. Biol. Ann. Rev. 2: 281- 5:164 p. 339. TAGATE,M. E., ANDD. L. DUDLEY. 1961. Seasonal LEIM, A. H., ANDW. B. SCOTT. 1966. Fishesof the occurrence of marine fishes in four shore habitats Atlantic Coastof Canada. Fish. Res. Bd. Canada, near Beaufort,North Carolina,1957-1960. U.S. Bull. 155: 1-485. Fish Wildl. Serv. Spec. Sci. Rep., Fish. 390: Lux, F. E., AND F. E. NICHY. 1971. Number and 1-19. lengths,by season,of fishescaught with an otter TREMBLEY,F.J. 1960. Researchproject on effects trawl near Woods Hole, Massachusetts,Septem- of condenserdischarge water on aqnatic life. ber 1961 to December 1962. U.S. Dep. Comm. Inst. Res., Lehigh Univ., Prog. Rep., 1956-1959.

Downloaded by [University of Maine] at 11:04 22 December 2011 Spec. Sci. Rep., Fish. 622: 1-15. Cited in P. A. Krenkel and F. L. Parker (eds.), MARCY, B.C., JR., P.M. JACOBSON,AND R. L. Biologicalaspects of thermal pollution. Vander- NANKEE. 1972. Observations on the reactions bilt Univ. Press,Nashville, Tennessee.p. 238. of young American shad to a heated effluent. TYLER,A. V. 1971. Periodic and resident compo- Trans. Amer. Fish. Soc. 101: 740-743. nents in communities of Atlantic fishes. J. Fish. MASSMANN,W. H. 1962. Water temperatnres, sa- Res. Bd. Canada 28: 935-946.