life history and population characteristics of striped bass in atlantic canada

Transactions of the American Fisheries Society 124:477-507. 1995€> Copyright by the American Fisheries Society 1995

Life History and Population Characteristics ofStriped Bass in Atlantic Canada

ROGER A. RULIFSONDepartment of Biology, and Institute for Coastal and Marine ResourcesEast Carolina University, Greenville, North Carolina 27858-4353, USA

MICHAEL J. DADSWELLDepartment of Biology, Acadia UniversityWolfiille, Nova Scotia BOP 1X0. Canada

Abstract.—Populations of striped bass Morone saxatilis occur in three regions of Atlantic Canada:the St. Lawrence River and estuary in Quebec; the Gulf of St. Lawrence from Chalcur Bay throughNorthumberland Strait; and rivers in New Brunswick and Nova Scotia draining into the Bay ofFundy. At least nine rivers are known or believed to sustain spawning populations: the St. LawrenceRiver (where the spawning stock may be extirpated); the Nepisiguit River in Chaleur Bay; theTabusintac, Miramichi, Kouchibouguac, and Richibucto rivers in the western Gulf of St. Lawrence;the Saint John and Annapolis rivers in the outer Bay of Fundy; and the Shubenacadie-Stewiackcriver system in the inner Bay of Fundy. Historically, commercial striped bass landings in AtlanticCanada were lower than those of the U.S. eastern seaboard, the largest landings being made inNew Brunswick. Angling catches are substantial but difficult to verify. Spawning occurs in tidalstreams several weeks after ice leaves the system. Adults exhibit regional and long-distance summermigration but seek freshwater refugia for overwintering. Studies of growth, population age struc-tures, comparative total instantaneous mortality rates, meristic and morphomctric characters, andparasites, combined with tag returns, strongly suggest that Bay of Fundy rivers, specifically theAnnapolis, Shubenacadie, and Saint John, contain stocks that are mixed with U.S. fish in mostyears. Bay of Fundy fish apparently participate in the striped bass coastal migration on the easternseaboard, whereas populations in Gulf of St. Lawrence rivers appear to be localized. Anecdotalinformation from commercial and angling catches suggests more spawning-overwintering popu-lations occur than are known to science. Periodic population assessments should be conducted toprovide the information necessary for knowledgeable management of the species in eastern Canada.

Striped bass Morone saxatilis is one of the most conducted in recent years to identify rivers of or-important commercial and recreational species igin for individuals of the U.S. coastal migratoryalong the Atlantic coast of North America. Pop- stock (see review by Waldman et al. 1988; Wirginulations north of Cape Hatteras, North Carolina, et al. 1993). Most researchers have not consideredexhibit anadromy by spawning in fresh or brackish the possibility that Canadian populations contrib-waters in the spring and then returning to coastal ute to the mixed stock migrating along the U.S.ocean waters. Juveniles tend to stay in brackish coast. However, results of studies involving tag-coastal waters until 1-2 years of age, then enter ging, parasites, meristic and morphometric char-the ocean and participate in coastal migrations, acters, and biochemical genetics suggest that Ca-Populations south of Cape Hatteras to Florida and nadian and U.S. striped bass populations mix;in the Gulf of Mexico to Texas are considered to however, the extent of this mixing is undeterminedbe endemic and riverine (Mcllwain 1980). Life (Melvin 1978; Dadswell et al. 1984;Hogans 1984;history aspects of southern striped bass popula- Boreman and Lewis 1987; Rulifson et al. 1987;tions were reviewed by Rulifson et al. (1982a, Harris and Rulifson 1988; Waldman et al. 1988).1982b); mid-Atlantic populations were reviewed Striped bass inhabiting Canadian coastalby Setzler et al. (1980). streams and lakes and coastal marine waters have

Knowledge about northern populations of provided important commercial and recreationalstriped bass declines at the U.S.-Canada border, fisheries in Atlantic Canada since early Europeaneven though the species' range extends through settlement (Bayne 1930; Rogers 1936). Renewedthe Bay of Fundy and the Gulf of St. Lawrence to interest in striped bass in Atlantic Canada has co-the St. Lawrence River (Scott and Scott 1988). incided with closure of the commercial and someConsiderable U.S. research, employing a variety recreational fisheries for Atlantic salmon Salmoof biochemical and genetic techniques, has been salarin the 1980s and 1990s, the prospect of tidal

477

478 RULIFSON AND DADSWELL

61°

FIGURE 1.—The three regions of eastern Canada with striped bass populations: the St. Lawrence River andestuary, the Gulf of St. Lawrence-Northumberland Strait, and the Bay of Fundy. (P.E.I, is Prince Edward Island.)

power development in the inner Bay of Fundy(Gordon and Dadswell 1984), recognition of thespecies* potential for aquaculture (Peterson 1991),and restoration programs for specific populations(Beaulieu 1985; Peterson 1991). In October 1986,the New Brunswick Department of Natural Re-sources and Energy recommended that striped bassbe declared a sport fish and that commercial fishingbe prohibited (Savoie et al. 1990). Recent regionaldocuments have described life history aspects andcurrent management practices of striped bass inQuebec, New Brunswick, and Nova Scotia (Beau-lieu 1985; Chaput and Randall 1990; Jessop 1990;Hogans and Stokesbury 1991; Hooper 1991; Mel-vin 1991; Peterson 1991); however, much of thelife history information for Canadian stocks hasyet to be published.

The objectives of our study were to describe thelife history characteristics of striped bass popu-lations in Atlantic Canada, to present a hypothesisconcerning stock interactions during local andlong-distance migrations, and to estimate the pos-sible influence of U.S. fish on Canadian stocks.This information will be useful in developing fu-ture research plans for stock assessment, restora-tion, and management in both countries. Much of

the information comes from our unpublished re-search, but we also relied on mimeographed un-published manuscripts of other workers, federaland provincial in-house reports, university grad-uate theses and dissertations, and interviews withlocal commercial and sport fishermen, conserva-tion officers, and fishery biologists.

Striped Bass in Atlantic CanadaGeographic Distribution

Striped bass occur in three regions of AtlanticCanada: the St. Lawrence River and estuary inQuebec; the Gulf of St. Lawrence from ChaleurBay through Northumberland Strait, specificallyrivers of eastern New Brunswick; and the Bay ofFundy from the St. Croix River in New Brunswickto the Tusket River in southern Nova Scotia (Fig-ure 1). No spawning populations of striped basshave been located on Prince Edward Island or CapeBreton to date. At least nine river systems areknown or believed to sustain spawning popula-tions: the St. Lawrence River; the Nepisiguit Riverin northern New Brunswick; the Tabusintac, Mir-amichi, Kouchibouguac, and Richibucto rivers inthe western Gulf of St. Lawrence; the Saint John

STRIPED BASS IN ATLANTIC CANADA 479

and Annapolis rivers in the outer Bay of Fundy;and the Shubenacadie-Stewiacke river system inthe inner Bay of Fundy.

Commercial FisheryLandings of striped bass in the Canadian Mar-

itimes (New Brunswick and Nova Scotia) are con-siderably lower than those reported for the U.S.eastern seaboard. Commercial harvest of stripedbass was primarily from bycatch in fisheries forother anadromous species, particularly Americanshad Alosa sapidissima, although directed fisherieswere present at some locations, such as the Bel-leisle Bay area of the Saint John River and in theMiramichi region. Greatest reported landings inthe Maritimes were about 50 metric tons (mt) in1945 and 48 mt in 1981 (Melvin 1991; Figure 2).Quantitative changes in landings both within Can-ada and between Canada and the USA were oftenout of phase, suggesting a different recruitmentprocess, different annual migratory events, orboth.

Within the Maritimes, New Brunswick had thegreatest landings. Commercial harvest of stripedbass was primarily from the Saint John River, prin-cipally from Belleisle Bay of the Saint John Riverestuary. Most of the remainder was landed in theMiramichi River region of eastern New Bruns-wick. Gear types included weirs, traps, and gillnets. Harvest in some areas was year-round, in-cluding an under-ice gill-net fishery (Jessop 1991).A winter (December-March) bow-net fishery forstriped bass was traditional in the Richibucto areaof northeast New Brunswick; typical landingswere 1-3 mt (Jessop and Swetnam 1978). NewBrunswick striped bass catches have been sporadicthroughout the years; catch records are not com-plete and often conflict because of overlapping orexclusion of reporting areas. Moderately largecatches occurred between 1875 and the early1890s, and peaked at about 55 mt in 1888 (Rogers1936). The commercial fishery, concentrated in theSaint John River, was closed from 1892 to 1894to protect the stock and was closed again in 1978(Jessop 1991). Annual landings in the Saint JohnRiver estuary averaged less than 4.5 mt between1895 and 1955; harvest then peaked at 48 mt in1959; landings exceeded 9 mt in four of the yearsbetween 1958 and 1970, after which catches de-clined sharply (Figure 3). Striped bass commerciallanding records for the Saint John estuary do notfollow the general trend for other landings in theMaritimes (Figure 2).

In Nova Scotia, the greatest reported commer-

St. Lawrence R.

O•M

O

§

Belleisle BaySt. John River

20-

1930 1950 1970 1990

FIGURE 2.—Commercial landings of striped bass inAtlantic Canada and along the U.S. Atlantic coast since1930. Quebec landings are exclusively from the St. Law-rence River; Maritimes landings are from New Bruns-wick (exclusive of Belleisle Bay) and Nova Scotia; Bel-leisle Bay was the major fishery of the Saint John River,New Brunswick. United States landings are in thousandsof metric tons.

cial landings of striped bass were in Hants andColchester counties (Caddy and Chandler 1976),the area surrounding the inner Bay of Fundy. Land-ings in the Shubenacadie and Stewiacke riverswere primarily from a spring American shad gill-net fishery, whereas harvests in Cobequid Bay andMinas Basin were from a summer and fall weirfishery. Little or no commercial landings havecome from the Annapolis River area since the mid-1940s because commercial fishing was severelyrestricted by regulations (Jessop 1991). Shuben-acadie Lake, near Halifax, is the headwater of the


1875 80 90 1900

FIGURE 3.—Commercial landings of striped bass in the Saint John River estuary. New Brunswick (mainly BelleisleBay), from first fisheries statistics records in 1875 unt i l closure of the fishery in 1978. Data are in metric tons(thousands of kilograms).

Shubenacadie River and sustained a small com-mercial fishery of up to 0.8 mt from the mid-1880sto about 1911 (Jessop 1991). Small commercialfisheries were located on the Atlantic coast ofNova Scotia in the Isaac's Harbor-St. Mary's Riverarea during the late 1800s and early 1900s. Com-mercial Nova Scotia landings peaked at 13 mt in1943 and 15 mt in 1962 (Figure 4A), after whichcatches declined due to low stock abundance andthe eventual elimination of the directed fishery forstriped bass in 1970.

In Quebec, striped bass commercial landingswere from a weir fishery. Landings data date from1920, when 16.1 mt were harvested (Beaulieu1985). Harvest peaks in the St. Lawrence Rivercoincided with the Great Depression (starting in1929), the second World War (1940-1943), andthe postwar period of 1950-1952 (Figure 2). Land-ings of only 0.9 MT were reported in 1963; stockcollapse and the disappearance of striped bassfrom the St. Lawrence River was apparently com-plete by 1966 (Beaulieu 1985). Commercial fish-ing was conducted in the fall in the estuary nearQuebec City and in the Riviere du Loup (Figure5; Vladykov 1945) and was an important com-ponent of inland commercial fishing until the mid-1950s. The estuarine fishery predominated during1954-1956; the inland fishery (Lac St.-Pierre) col-

lapsed between 1953 (11.4 mt) and 1954 (2.7 mt)and stayed between 1 and 3 mt until 1963 (Beau-lieu 1985).

Sport FisheryUntil recently, little attention was given by Ca-

nadian federal and provincial fisheries agencies tomonitoring the striped bass sport fishery becauseinterest was focused on Atlantic salmon and brooktrout Salvelinusfontinalis. After World War II, fish-ing camps for striped bass were popular aroundthe Bay of Fundy, especially on the inner Bay, onthe Annapolis River, and at Reversing Falls at themouth of the Saint John River. The popularity ofthese camps waned in the 1970s as a result of stockdecline and poor catches. A number of them sur-vived, however, and since 1984 fishing interest hasincreased as striped bass have grown more abun-dant.

In New Brunswick, striped bass anglers frequentthe eastern shore from Chaleur Bay south alongthe Gulf of St. Lawrence through the Northum-berland Strait (Figure I) . In the 1960s and 1970s,minor sport fisheries were documented for the Ri-chibucto, Tabusintac, and Miramichi rivers (Wil-son 1958; Dominy 1972). More recently, anglinghas been concentrated in eastern New Brunswickon Miscou Island in the Gulf of St. Lawrence,


1970 1990

FiciURF. 4.—Commercial and recreational landings ofstriped bass in Annapolis County, Nova Scotia. (A)Commercial landings (metric tons) for 1895-1990, pri-marily from shoreline weirs in the Bay of Fundy; weirfishery has been phased out since 1970. (B) Seasonalangler catches (total fish per day) estimated from fishinglicense questionnaires, 1966-1988 (closed bars, Jessop1991), and from creel surveys or reported in the Dun-romin fishing contest (open bars) for the Annapolis River(Jessop 1991).

Nepisiquit Bay, and the region between the Mir-amichi and Kouchibouguac rivers (Madden 1984;Bujold 1985). On the Bay of Fundy coast, popularNew Brunswick angling locations are Passama-quoddy Bay, Cumberland Basin, and the SaintJohn River estuary, especially at Reversing Fallsin Saint John Harbour. Popular estuarine anglingsites included the Saint John River and tributaries,especially the Nerepis, lower Kennebecasis, andlower Hammond rivers (Wilson 1958); the Petit-codiac River, the St. Croix River, and the Maga-guadavic River (Dominy 1972).

In Nova Scotia, sport fishing for striped bass isconcentrated around the Bay of Fundy and in the

Northumberland Strait and Cape Breton. Popularareas along the north shore of Minas Basin andCobequid Bay include Five Islands (mouths ofHarrington and Five Islands rivers) and the estu-aries of the Economy and Bass rivers. Along thesouth shore of the same embayment, sport fishingactivity is concentrated in the Shubenacadie-Ste-wiacke estuary (Jessop and Vithayasai 1979), theNoel Shore, and the Gaspereau River estuary.Along the Atlantic coast, angling occurs in theestuaries of the Mira, Sissiboo, Chebogue, Argyle,Tusket, LaHave, and St. Mary's rivers (Jessop1991).

Perhaps the most popular and well-documentedstriped bass sport fishery in Atlantic Canada is inthe Annapolis River. Catch by hook and line (es-timated from provincial fishing license question-naires) peaked in the 1970s but since has declined(Figure 4B). Questionnaire returns probably un-derestimated the actual catch because no licensewas required for angling striped bass in the tidalwaters, where most of the catch occurred (Jessop1991). Creel surveys conducted in the 1970s and1980s documented the importance of the sport fish-ery in the Annapolis region (Jessop and Doubleday1976; Jessop and Vithayasai 1979; Jessop 1980;Harris and Rulifson 1988). Of the anglers inter-viewed in 1976, 73% did not reside within 100km, and 18% were from the USA and other coun-tries (Jessop and Vithayasai 1979). The 1983 and1985 records from Dunromin campground (a pop-ular angling site) show that U.S. campers camefrom nearly every state and took about 35% of allcampsites used (Rulifson et al. 1987). A 1987 creelsurvey showed that 3.9% of interviewed fishermenwere from the USA, who came primarily duringJuly and August (Harris and Rulifson 1988).

In Quebec, sport fishing for striped bass wasvery popular in August and September and wasconcentrated near Quebec City and in the OuelleRiver (Figure 5). Organized fishing trips werecommon, and from 1954 to 1965 a fishing tour-nament was held annually (Beaulieu 1985). Vla-dykov (1945) reported that some anglers were ableto catch 50-80 striped bass in one trip during 1944.

Life History Aspects by RegionSt. Lawrence River

Historically, the St. Lawrence River striped basspopulation was an important component of the res-ident fish community (Bergeron and Brousseau1975, 1981). The population was considered en-demic, based on tag returns from 1946 to 1962


Overwinter

Spawning

Nursery

Movement

FIGURE 5.—Overwintering sites, possible spawning sites, nursery areas, and seasonal movements of striped bassin the St. Lawrence River and estuary. Quebec; R. denotes river.

(Magnin and Beaulieu 1967). The natal spawningpopulation apparently was decimated in the early1960s (Bergeron and Brousseau 1983) and waspossibly extinct by 1966 (Beaulieu 1985), al-though occasional anecdotal sport catches were re-ported (Lebel 1983). The species was declared en-dangered in Quebec province in 1984 (Beaulieu1985). Distribution was reported from the mouthof the St. Lawrence River at Riviere du Loup toSorel (Lac St.-Pierre), and occasionally to Mon-treal (Figure 5; Cuerrier et al. 1946; Vladykov andMcAllister 1961; Bergeron and Brousseau 1975).

Adult striped bass exhibited a fall migration up

the St. Lawrence River to overwintering areas nearTrois-Rivieres (Lac St.-Pierre; Figure 5), near pos-sible spawning grounds (Vladykov 1946, 1947;Beaulieu 1962), although no study of spawninghabitats was ever conducted (Magnin and Beaulieu1967). Though no commercial fishing was allowedfrom 1 December through 31 May, striped basswere taken as bycatch in Lac St.-Pierre by netfishing under the ice (Beaulieu 1962). Larvae werereported in the mouths of rivers near Louisevilleon the north bank of Lac St.-Pierre (Beaulieu1985). Deterioration of spawning habitat wasthought to have been caused by construction of the


60 r

Z

LL

10

200 280 360 440 520 600 680

FORK LENGTH (mm)

CCHICD 800-

600-

40O-

20O-

100-

B 6.0

X</> 4.0

2.0-

2 ~- 0.62

2 4 6 8 10 12AGE (yrs)

2 4 6 8 10 12 14 16 18 20AGE (y rs )

FIGURE 6.—Population characteristics of striped bass in the St. Lawrence River and estuary, Quebec. (A) Fork-length-frequency of fish captured in weirs near Riviere Ouelle in the St. Lawrence estuary. (B) Age-frequency offish in weirs at Riviere Ouelle. (C) Total natural mortality rate (Z) of St. Lawrence River fish, based on log^transformations of age-frequency data. Data are from Brousseau (1955).

St. Lawrence Seaway (1954-1959), island con-struction for the International World's Fair of 1967in Montreal (1963-1964), and creation of SternsIsland from dredge spoil in 1965 (Beaulieu 1985).Spawning season was probably from mid-May intomid-June; most adults left the area by June, exceptfor late newcomers (Vladykov and Brousseau1957).

Young striped bass migrated downstream to the

St. Lawrence estuary as far as Riviere Ouelle (Vla-dykov 1947), primarily in the Portneuf-Neuvillesector (north bank) and the Ouelle estuary (Figure5; Beaulieu 1985). In July, age-0 fish, 20-25 mmfork length (FL), were found near Neuville (Vla-dykov and Brousseau 1957). By September, age-0 striped bass averaging 75 mm were observedfrom the Saint John-Port Joli river estuary nearQuebec City to the Riviere Ouelle (Beaulieu


100

80

QIL-i—i—i—i—i—i—,—i—< i . i . i . i . i

Age

FIGURE 7.—Growth rate comparisons of striped bass from the regions of Atlantic Canada: the St. Lawrence Riverand estuary (Magnin and Beaulieu 1967), the Gulf of St. Lawrence (Hogans and Melvin 1984), the Saint JohnRiver (Jessop 1980), and the Annapolis River (Dadswell 1976). All growth estimates, except that for the St. LawrenceRiver population, most likely involve fish of mixed origins. Vertical lines indicate sample ranges; symbols aremean fork lengths at age.

1985). In the fall, 200-280-mm fish were in thelower river near the Riviere Ouelle, pursuingyoung American shad and alewives Alosa pseu-doharengus (Beaulieu 1962). Young striped basswere often caught as bycatch in the fishery forrainbow smelt Osmerus mordax (Vladykov 1945).

The population structure of St. Lawrence Riverstriped bass was well documented. Sampling in1954 indicated a bimodal length frequency distri-bution (Figure 6) with peaks of individuals at 260mm and at 420 mm (Brousseau 1955). Growth ofSt. Lawrence River striped bass was slower andmaturity was later than those of other populations

in Canada (Figure 7). The weight-length relation-ship of St. Lawrence River fish reflected slowergrowth (Table 1). Magnin and Beaulieu (1967) re-ported age-0 fish were about 50 mm in July andAugust; by fall, fish averaged nearly 100 mm (Table2). Juveniles were about 175 mm by the secondsummer, 225 mm by fall, 300 mm in the third sum-mer, and 325 mm in the third fall. The oldest fishsampled in the St. Lawrence population was age19, but most fish sampled were ages 1-7 (Table 2).The largest fish recorded was 882 mm FL and theheaviest specimen weighed 10.6 kg (Table 3). TheBrody growth coefficient (K), however, was greater


TABLE 1.—Relationships of weight (WO to fork length (L) for striped bass populations in eastern Canada. Sexes arc:C = combined, M = male, F = female. See footnotes to sample size (N) for length and weight units used in theequations.

River system

St. LawrenceRiver and estuary

Tabusintac

Kouchibouguac

Kouchibouguac,

Year

Prc- 1967

1975-19771975

19781983-1984

198319831978

Sex

C

CC

CCMFC

N

2,I77b

108C

36*

hddd

\6&

Equation3

LogW = 3.0137(logL) - 4.8509

Gulf of St. LawrenceLoglV = 3.322<logL) - 2.319

Log W= 2.85 1 (log/.) -4.48

LogW = 3.14(log/,) - 5.355LogW=2.7l(logL)- 1.31LogW=3.4l(logL) - 2.45LogW=2.8l(logL)- 1.41LogW= -5.164(Xlog/-) -f- 3.1801

r?

0.97

0.99

0.99

0.980.920.740.710.99

Source

Magnin and Beaulieu (1967)

Melvin (1978)

Adapted from Patriquinand Butler (1975)

Hogans and Melvin (1984)Hogans and Melvin (1984)Hogans and Melvin (1984)Hogans and Melvin (1984)Jessop and Swetnam (1978)

Kouchibouguacis.and Richibucto

Annapolis

Shubcnacadic Lake

Shubenacadie River

Stewiacke-Shubenacadie

Saint John

1976

197819761987

1975-1977

19761976

1992

19751973-19751975-1977

CFCFCMFCCCCMF

CCC

ISO*125d

239*27*1

164h

,9h89h

85«43<«18d

37*20"10*

357<168J

64C

Bay of FundyLog\V = 3.1332(log/L)LoglV =LoglV =LoglV =LoglV =LoglV =Log IV =

Log IV =LoglV =LoglV =LoglV =LoglV =LoglV =

Log IV =LogW =LoglV =

3.l319(logL)2.9973(logt)3.23(log£.) -2.94(logL) -3.l6(logL)-2.96(log/.) -

2.976(log/,) -

- 5.1730-5.1834-4.91565.354.785.424.84

- 1.799

3.09578(logL) - 4.9683.23(logt) -3.358(log£.) -2.968(logL) -2.939(IogL) -

3.l4(logL)-2.94(logL) -2.95<Xlog/,) -

5.24

- 5.880-4.815- 4.643

4.90310.92

- 1.773

0.980.980.980.990.980.920.98

0.98

0.980.99

0.960.920.77

0.960.97

Jessop and Vithayasai (1979)Jessop and Vithayasai (1979)Jessop (1980)Ray e ta l . (1984)Harris (1988)Harris (1988)Harris (1 988)

Melvin (1978)

Jessop and Vithayasai (1979)Ray et al. (1984)

Rulifson and Jessop (this paper)Rulif'son and Jessop (this paper)Rulifson and Jessop (this paper)

Melvin (1976)Dadswell (1976)Adapted from Melvin (1978)

a All logs are to base 10.b Millimeters, kilograms.c Centimeters, grams.d Centimeters, kilograms.

than for fish from most other rivers discharging intothe Gulf of St. Lawrence (Table 3), which indicatedrelatively fast growth. Based on Magnin and Beau-lieu 's (1967) data, we estimated (Ricker 1971) thatthe total mortality rate (Z) of St. Lawrence Riverstriped bass was 0.62 (Figure 6C). Vladykov andBrousseau (1957) reported that age at maturity ofSt. Lawrence River fish was earlier for males thanfor females. Age-2 males averaged 269 mm; a "por-tion" of age-3 males (322 mm) was sexually ma-ture, and "most" were mature by age 4, averaging385 mm FL. None of the age-2 (269 mm) or age-3 (328 mm) females sampled from 1944 to 1957were mature. First maturity was noted among age-4 females (396 mm), and most females were matureby age 5 (469 mm) and age 6 (515 mm; Vladykovand Brousseau 1957).

Gulf of St. Lawrence—Northumberland Strait

Historically, striped bass inhabiting coastal wa-ters of northeastern New Brunswick, Nova Scotia,and Prince Edward Island were important membersof the resident fish community and provided smallbut important commercial and recreational fish-eries to Maritime residents. Striped bass frequentshoreline coves and small streams, but spawninghas been documented in only four rivers (Figure8). No spawning grounds have been identified onPrince Edward Island or Cape Breton. The warmshallow waters of Northumberland Strait betweenthe south shore of Prince Edward Island and north-ern shores of New Brunswick and Nova Scotia arein sharp contrast to the surrounding cold NorthAtlantic waters (Scott and Scott 1988); this tem-


TABLE 2.—Lengths at age of striped bass populations in eastern Canada. Asterisks on years indicate that mean lengthsrepresent observed lengths at capture; otherwise, means represent back-calculated lengths. Parenthetic numbers arepercentage age distributions of fish in the samples; empty cells mean that no fish were in that age-class. Sexes are: C= sexes combined; M = males; F = females.

River systemand year

St. Lawrence Riverand estuaryPre-1967

Bathurst Harbour1978-1983*

1986*

Miramichi1984*

Kouchibouguac1978*

1983-1984

Kouchibouguac.Kouchibouguacis,and Richibucto1978*

Mean fork length (mm) in

Sex

C

C

MFC

C

C

C

C

M

F

N

2,696

255

8685120

43

106

303

174

24

33

1

97(26.0)

145

2

216(26.9)

160(1.7)

315

224(12.9)

235(20.1)

3 4

317 388(15.9) (12.1)

Gulf of St

351 429(10.2) (45.5)(20.9) (61.6)(9.4) (38.8)

437(17.5)

373 406

369 441(7.5) (22.6)339 406(16.2) (42.6)

354 440(23.0) (14.4)349 438(58.3) (20.8)352 439(30.3) (12.1)

5

433(9.6)

6

481(5.2)

7

529(3.1)

age-class:

8

567(0.7)

9

580(0.2)

10 11

674(O.I)

12

Lawrence

516(20.0)(7.0)(5.9)483(16.7)

531

510(60.4)467(23.8)

510(36.8)

(16.7)516(39.4)

599(8.6)(2.3)(7.1)536(13.3)

579

553(5.7)527(4.6)

553(3.4)

553(9.1)

643(7.1)(5.8)(14.1)574(11.7)

566

691(2.4)(1.2)(4.7)612(15.8)

610

721(0.4)

(2.4)657(15.8)

688

732(0.4)(1.2)

747(4.2)

739(3.8)

739(2.3)665(4.2)764(91)

815(1.6)

(5.9)758(3.3)

869(3.1)

(9.4)

Bay of FundyAnnapolis

1972

1976*

1978*

1987

Gaspereau1976*

Shubenacadic1976*

1992*

Saint John1971-1972

1973-1975*

1975-1977*

C

M

F

C

C

C

C

C

C

M

F

C

M

F

C

C

177

54

54

150155130

167

19

44

37

20

10

357

46

149

177

62

164

164

167

239

125

127

125

150(0.6)

247

246

250

346

273(36.8)

246(18.2)258(5.4)

209

212

208

312(2.3)428(4.8)

332 404(9.0) (20.9)326 396(13.0) (24.1)340 412(3.7) (11.1)

505(0.6)420(0.8)

463 534(2.4) (26.9)

355 437(42.1) (15.8)

333 441(36.4) (13.6)410 474(10.8) (16.2)402 474(10.0) (30.0)

305 392(7.0) (33.3)310 396(6.5) (43.5)309 395(9.4) (22.8)458 488(11.7) (23.2)509 566(40.3) (11.3)

460(26.0)449(27.8)470(29.6)582(9.0)490(0.8)589(28.1)

499(9.1)553(29.7)558(40.0)

474(37.3)462(32.6)472(37.6)558(15.8)626(9,7)

515(14.7)493(18.5)536

(11. 1)625(18.7)637(1.5)654(12.6)

535(5.3)

610(9.1)

535(14.3)515(13.0)537(21.5)663(9.0)679(6.5)

575(10.7)545(3.7)589(16.7)677(12.9)678(3.1)721(4.2)

620(13.6)660(10.8)672(10.0)648(20.0)

583(1.7)528

591(2.0)677(6.8)678(4.8)

606(7.3)580(5.6)616(11.1)783(5.8)712(14.6)111(3.0)

775(2.7)

775(10.0)

634(1.4)606

652(2.0)726(4.0)838(3.2)

644(4.5)596(3.7)668(7.4)782(7.7)750(16.9)828(7.2)

728(5.4)710(5.0)745(10.0)

691(1.7)685

691(2.0)797(3.4)810(1.6)

684(3.4)629(1.8)686(9.3)828(9.7)783(13.8)875(1.8)

735(5.4)750(5.0)720(10.0)

717(1.1)723

733(0.7)815(6.2)647(1.6)

715(1.7)578(1.8)

830(7.7)784(10.8)916

741(1.7)761(4.3)753(2.0)826(4.5)888(6,5)

765(1.7)

868(8.4)826(14.6)953

864(8.1)

864(30.0)

850(3.4)819(1.6)


TABLE 2.—Extended.


St. LawrenceRiver and estuary

Pre-1967

Bathurst Harbour1978-1983*

1986*

Miramichi1984*

Kouchibouguac1978*

1983-1984

Kouchibouguac,Kouchibouguacis.and Richibucto

1978*

Annapolis1972

1976*

1978*

1987

Gaspereau1976*

Shubcnacadic1976*

1992*

Saint John1971-1972

1973-1975*

1975-1977*

Sex

C

C

MFC

C

C

C

C

M

F

C

M

F

C

C

C

C

C

C

M

F

C

M

F

C

C

Mean fork length (mm) in age-class:

13 14 15 16 17 18 19

736 823 819 736 888(<0.1) (0.1) (<0.1) (<0.1)

Gulf of St Lawrence

884 787(0.4) (0.4)

(1.2) (1.2)

Bay of Fundy

862 879 964 1,028 1,002 1,010(5.8) (4.5) (5,2) (1.3) (1.3) (1.3)

901 938 955 975 1,013 1,035(5.4) (3.8) (3.1) (5.4) (2.3) (2.3)

987 1.020 1,042 1,067 1,087 1,104 1,095(0.6) (0.6) (1.2) (1.2) (3.0) (2.4) (1.8)

895(5.4)

895(20.0)

930 1,025 1,080 1,008 1.037(2.3) ( I . I ) (0.6) (2.8) (1.7)

965 1,050 1,060 1.090(1.6) (3.2) (1.6) (1.6)

20 21 Source

Magnin and Beaulieu(1967)

Madden (1984)

Meagheret al. (1987)

Watling(I985)

Jessop and Swetnam(1978)

Hogans and Melvin(1984)

Jessop and Swetnam(1978)

Williamson (1974)

Jessop and Vithayasai(1979)

1 ,050 Jessop ( 1 980)(0.8)1,128 1.156 Harris (1988)(1.2) (1.8)

Jessop and Vithasayai(1979)

Jessop and Vithayasai(1979)

Rulit'son and Jessop(this paper)

Williamson (1974)

1,030 Dadswell(1976)(0.6)

Adapted from Melvin(1978)


TABU- 3.—Observed and calculated biological variables for selected striped bass populations in eastern Canada.

Population

St. Lawrence RiverBathurst HarbourKouchibouguac RiverShubenacadie RiverAnnapolis River

All fishAll fishFemales

Saint John River

N

2.69637530380

13067889

249

Maximumobserved

forklength(mm)

882935537940

1.054l.287h

I.333C

Maxi-mum

observedweight

(kg)10.67.5

12.5

19.222.3h

22.328.6C

Cal-culated

maximumfork

length(mm)

904968883

1.200

1.1901,2601.4001,125

Brodygrowthcoeffi-cient(/Oa

0.160.160.120.12

0.080.110.090.10

Totalmortal-

ity(Z)a

0.610.430.500.23

0.15

0.19

Maxi-mum

observedage

(years)

19146

13

2021

21

Source

Magnin and Beaulieu (1967)Madden (1984)Hogans and Melvin (1984)Rulifson and Jessop (this paper)

Jessop(l980)Harris (1988)Harris (1988)Dads well (1976)

a Calculated for this study.h A 21-year-old striped bass of probable U.S. origin, based on scale characteristics.c Value given is for 21-year-old striped bass of probable U.S. origin. A second 21-year-old fish of probable Canadian origin was much

smaller (1.090 mm FL; 17.2 kg). Origin determinations were based on scale characteristics.

perature contrast may impede long-distance mi-gration by striped bass into or out of this region.

At least seven rivers draining to the Gulf of St.Lawrence are known as overwintering sites foradult striped bass (Rulifson et al. 1987; four areshown in Figure 8). Whether these overwinteringfish are active participants in the spring spawningis unknown. In many cases, the larger adults areseldom or never observed in the river after the icebreaks up, which suggests that they may migratedownstream and leave in late winter. In the Nepi-siguit Riven large striped bass are angled from Sep-tember through November during the fall uprivermigration, but overwintering areas are undocu-mented (Madden 1984). In the Tabusintac River,residents report that fish of 16-18 kg are harvestedby bow-net and handline fisheries as they move intobays and upstream in late fall; these fish are neverobserved by local fishermen in the spring, eitherleaving or spawning. In the Miramichi River, res-idents report that striped bass averaging 1.4-1.8 kgmigrate upstream in late fall; largest individuals areabout 13 kg. These fish apparently migrate down-river in mid-January; commercial fishermen settheir nets for fish leaving the river, and a bow-netfishery through the ice occurs in early spring. Fish-ermen refer to the overwintering fish as "black-backs" because their dorsal coloration is muchdarker than the "greenback" striped bass migratingupstream in the spring. The Black and Napan riversadjacent to the Miramichi River also have fall up-stream migrations offish up to 18 kg in si/e. Spring-time coloration differences between overwinteringand upstream migrants are often noted in these riv-ers, as well. Hogans and Melvin (1984) found that

the Kouchibouguac River spawning populationoverwinters in the upper freshwater portion of theriver from December to March (Figure 8). TheKouchibouguac and Richibucto rivers are connect-ed by a common shallow lagoon protected from theGulf of St. Lawrence by barrier islands, so the twopopulations are probably closely related and mayoverwinter in a similar manner. Local fishermenreport that a run of larger fish starts in August andcontinues into the fall, at which time sportfishingis most intense. Historically, a winter (December-March) bow-net fishery through the ice was tradi-tional; overwintering striped bass were dipped fromshallow waters by 2-m-diameter nets through largeholes cut in the ice (Jessop and Swetnam 1978).

The exact timing of striped bass spawning ac-tivity in Gulf of St. Lawrence rivers is not welldocumented, but it may occur shortly after iceleaves the river. The pattern involves downstreammigration of overwintering fish as the ice is break-ing up, and because riverine waters warm quickly,the fish spawn in May and early June (Hogans andMelvin 1984, Meagher et al. 1987). Spawning inthe Kouchibouguac River occurs at temperaturesof 12-14.5°C and lasts about 3 d (Hogans andMelvin 1984). Both sexes in spawning conditionwere observed migrating upstream in the Mir-amichi River at 12-14°C and in the RichibuctoRiver at 16-16.5°C (Rulifson et al. 1987). Onenoteworthy exception to spring spawning may bethe Tabusintac River, where local fishermen insistthat spawning occurs in the late summer-early fall;large adults of 16-18 kg, with running milt andeggs, are caught migrating upstream. This phe-nomenon, if correct, would represent the only fall-

STRIPED BASS IN ATLANTIC C A N A D A 489

7' Gulf ofSt. Lawrence

FIGURE 8.—Overwintering sites, known and possible spawning sites, nursery areas, and seasonal movements ofstriped bass in the western Gulf of St. Lawrence; P.E.I, is Prince Edward Island. Heavy arrows show springmovements; light arrows show fall movements.

spawning population known. More information isneeded on the Tabusintac population to determinewhether these fish are fall spawners or are ap-proaching maturity for overwintering.

Spawning habitats of the Gulf of St. Lawrencepopulations appear similar among the rivers (Figure8). Spawning activity is usually just above the headof tide (Hogans and Melvin 1984). Madden (1984)reported that spawning in the Nepisiguit River takesplace at about 4 km upstream from the river mouthand farther but that age-0 fish had not been observedfor 30 years. Spawning is thought to occur at Cain's

Point bridge in the Tabusintac River (approximately25 km upstream of Tabusintac Bay) and in the Mir-amichi River above Newcastle (approximately 40km upstream of Miramichi Bay). Spawning in theKouchibouguac River occurs 5-6 km upstream ofKouchibouguac Bay just downstream of the over-wintering site (Hogans and Melvin 1984). The bot-tom is cobble-sized shale rubble covered by a layerof mud. Eelgrass Zostera marina is the dominantsubmerged vegetation. Waters are clear but darklystained, and there is one dominant daily tide 1-1.5m in amplitude; water currents are 0.25-0.5 m/s at


mid-flood or mid-ebb tides (Rulifson et al. 1987).The Richibucto and Kouchibouguac rivers are ge-omorphologically similar (Jessop and Swetnam1978).

Nursery grounds for young striped bass in Gulfof St. Lawrence rivers are primarily in the estuariesand protected brackish-water lagoonal areas alongthe coast (Figure 8). Historical nursery groundsinclude the Nepisiguit estuary and Bathurst Har-bour, where fishermen targeting rainbow smelt Os-merus mordax commonly catch juvenile and sub-adult striped bass 40-200 mm FL (Madden 1984;Bujold 1985; Meagher et al. 1987). Fish up to 240mm FL are caught in traps for American eels An-guilla rostrata in the lower Tabusintac River dur-ing May; fingerlings appear in the area in the fall(Rulifson et al. 1987). In the Miramichi River, fin-gerlings 75-100 mm FL appear in October belowNewcastle and are captured in Miramichi Bay dur-ing fall as bycatch among Atlantic tomcod Micro-gradus tomcod (McKenzie 1959) and during winterin nets set for smelt (McKenzie 1964). Juvenilesare found in Kouchibouguac Bay during summerand fall, then reenter freshwater to overwinter.Age-0 striped bass frequent brackish waters andthe estuary throughout the year, regardless of theseason (Hogans and Melvin 1984).

No long-term data exist to adequately describepopulation trends in rivers of the Gulf of St. Law-rence and Northerumberland Strait. After limitedsampling, Melvin (1978) reported that the Tabu-sintac River population in 1977 was made up pri-marily of age-0 fish; the remainder were ages 2-4; no older fish were present (Figure 9A). Age-frequency distributions for the KouchibouguacRiver population were determined in 1978 and1983-1984 (Figure 9B, C, D); age-classes 4 and5 dominated, and no fish were older than age 6(Table 2). Jessop and Swetnam (1978) reported afew age-10 fish in their study. The age-class struc-ture of the Kouchibouguac River population mayshift seasonally (Figure 9E, F). A March 1983 sur-vey of the overwintering Kouchibouguac Riverpopulation indicated a prevalence of age-3 males;none was more than 6 years of age. In April-May1983, samples of males still ranged from age 3 toage 6, but age-4 fish were most abundant. In thefall and winter of 1983-1984, samples of malesranged from age 4 to age 6, and age-5 fish weremost abundant. The age structure of female Kou-chibouguac River striped bass also exhibited a sea-sonal shift. Samples of females in March were ages4-5, dominated by age-4 fish; by April-May, fe-males aged 4-6 were represented, but age-5 fish

dominated. In fall-winter 1983-1984, samples offemales aged 5-6 were represented but age-6 fishdominated (Hogans and Melvin 1984; Figure 9D).The largest fish sampled was 527 mm FL (Table3). Based on Hogans and Melvin's (1984) data, weestimate the total mortality rate (Z) of the Kouchi-bouguac population at 0.50 (Table 3). Growth ofKouchibouguac fish is greatest in the third year,and length at age is greater than in the St. LawrenceRiver population but less than that of the BathurstHarbour and Miramichi River populations (Table2). Fish in Bathurst Harbour are heavier at agethan Miramichi River fish (Table 4).

The age at maturity of Gulf of St. Lawrence-Northumberland Strait populations is not well doc-umented. Hogans and Melvin (1984) reported thatnearly all age-4 males and age-5 females sampledwithin the Kouchibouguac River were mature,which is about 1 year earlier than Bay of Fundypopulations. Mean length at maturity was 340-530mm (Melvin 1978; Hogans and Melvin 1984). Fe-cundity of age-5 females (N = 7) was estimatedat 98,286 (SD, 13,828) eggs; the only age-6 femaleexamined contained 121,000 eggs.

Striped bass in eastern New Brunswick exhibita northward summer migration after spring spawn-ing, then return in late fall to overwintering sites(Figure 8). Tagging studies by Hogans and Melvin(1984) and observations by Madden (1984) andothers indicated that fish of mixed origin moveslowly northward, frequenting coastal areas,coves, and estuaries. Striped bass enter MiramichiBay in June and early July. By late summer stripedbass enter Nepisiguit Bay, where sport fishing ismost successful in September. Southward migra-tion occurs rather quickly in late fall as fish moveback toward overwintering grounds.

Ba v of FundyAt least three major rivers support spawning

populations of striped bass in the Bay of Fundy:the Shubenacadie and Annapolis rivers in NovaScotia, and the Saint John River in New Brunswick(Figure 10). Smaller rivers and coastal streams of-ten contain striped bass, but no evidence exists forreproducing populations in smaller streams.

Overwintering of Bay of Fundy striped bass issimilar in timing to that of the Gulf of St. Lawrencepopulations. Fish move into Minas Basin and Cob-equid Bay (the inner Bay of Fundy) during Sep-tember through November to ascend the Shuben-acadie River for overwintering in ShubenacadieLake (Melvin 1978; Rulifson et al. 1987; Figure10). A commercial winter under-ice fishery was


100 B

1001

ZUJDo

0 1 2 3 4 5 6 7 8 0 1 2 3 4 5 6 7 8

0 1 2 3 4 5 6 7 8

100

20

6 ' 2 3AGE (y rs )

FIGURE 9.—Age frequencies of striped bass populations from Gulf of St. Lawrence rivers: (A) Tabusintac River(T), 1978, research gill nets; (B) Kouchibouguac River (K), 1983, research gill nets; (C) Kouchibouguac River,1978, winter hoop-net fishery; (D) Kouchibouguac River, 1983, hoop net fishery; (E) fish at overwintering site(W) during March 1983; (F) fish on spawning grounds (S) during April-May 1983. Data are from Melvin (1978)and Hogans and Melvin (1984).

active in Shubenacadie Lake at the turn of thecentury, and 178 striped bass were caught in fish-ing derbies during 1984-1986 (Jessop 1991). Oth-er adults may overwinter in deep (6-9-m) holeswithin the Stewiacke River (Rulifson et al. 1987).In the Annapolis River, large adults move upstreamthrough the Annapolis Royal causeway to upriverareas in October (Harris and Rulifson 1988). His-torical overwintering grounds for Annapolis Riverfish are unknown, but at the present time fish arethought to congregate in the estuary downstreamfrom Bridgetown. In New Brunswick, striped bassmove into the Saint John River estuary and up-stream in September to overwinter primarily in

Belleisle Bay but also in Washademoak Lake andprobably other deep sections of the estuary (Dads-well 1976).

Spawning activity in Bay of Fundy rivers is inMay and June, depending on the river system. TheShubenacadie and Stewiacke rivers are tidallydominated; the highly turbid waters warm ratherquickly but temperatures decrease rapidly on eachtidal cycle as cooler inner Bay of Fundy waterspush upstream, forming a tidal bore. Adult fishmigrate downstream from Shubenacadie Lake be-ginning the first week in May and arrive at theconfluence of the Shubenacadie and Stewiacke riv-ers (near the head of the salt wedge) by the third


TABLE 4—Weights (kg) at capture of aged striped bass from populations in eastern Canada. Sexes are: C = combined,M = male, F = female.


Mean weight (kg) in age-class:Sex 8 10 11 12 13

Bathurst Harbour1978-1983

Miramichi1984

Kouchibouguac1978

Kouchibouguac,Kouchibouguacis,and Richibucto

1978

C 243

C 43

C 106

C 174M 24F 33

Gulf of St Lawrence

0.50 1.00 1.73 2.41 3.04 3.54 4.32 4.64 5.% 6.59 6.36

0.18 0.59 0.68 1.68 2.27 2.27 3.23 4.54

0.67 1.20 1.88 2.23 5.48

0.16 0.58 1.20 1.88 2.230.54 1.18 1.560.56 1.15 1.88 2.23

Bay of Fundy

5.484.205.90

Annapolis1976

1978Gaspercau

1976Shubenacadie

1976

1992

Saint John1973-1975

1975-1977

C

C

C

ccMF

C

C

150

124

19

44

372010

177

62

0.34

0.21

0.16

0.30 0.40

1.20

0.80

0.60

0.901.00

1.43

1.92

1.60

0.92

1.35

1.49

1.331.33

1.70

2.53

2.34

1.41

1.94

1.952.05

2.55

3.44

2.89

3.35

2.60

3.28

3.53

4.40

3.63

3.62

3.56

3.836.363.66

4.35

4.35

4.83

4.08

7.045.237.04

5.46

8.18

5.50

4.79

5.91

5.46

7.88

7.50

6.68 6.97

5.45 5.88

5.30

5.36

7.75 8.40

6.47 9.58

8.22

6.32

11.67

11.67

9.80

7.04

7.71

7.98

10.20

10.20

11.74

11.70

week in May. These downstream migrants bear thesame black dorsal coloration noted on fish in theMiramichi, Black, and Napan rivers in the Gulf ofSt. Lawrence. At the same time, greenback stripedbass from seaward begin their upstream migrationin April and meet the downstream migrants by thethird week in May. Males in spawning conditionare found at 14°C, and ripe females are commonwhen temperature reaches 16°C. The major spawn-ing activity occurs in the Stewiacke River, about5 km upstream of the confluence with the Shu-benacadie River. Spawning starts in the fourthweek in May and, weather dependent, continuesto about 20 June (Meadows 1991). In 1992, eggswere collected in the Stewiacke River at temper-atures from 15.5 to 22.0°C (Table 5); in 1994, themajor spawning activity occurred when water tem-peratures reached 18°C (K. Tull and Rulifson, un-published data). Spawning in the Annapolis Riverstarts in late May and continues through June atwater temperatures from 15 to 24.4°C (Williams1978; Parker and Doe 1981; Williams et al. 1984).Peak egg production was observed after suddentemperature drops (Parker and Doe 1981); how-

ever, rapid temperature drops to 15-16°C resultedin a temporary cessation of spawning (Williams1978). Spawning in the Saint John River was inMay, beginning at about 11.5°C (Table 5), andmaximum activity was observed at 13.5°C (Dads-well 1976).

Spawning occurs near the head of tide for allthree Bay of Fundy river systems but the habitatsare quite different. Historically, spawning in theShubenacadie River was located from the conflu-ence of the Shubenacadie and Stewiacke rivers up-stream to Shubenacadie Village (km 25-34 fromthe confluence), but little if any spawning has oc-curred in the Shubenacadie River in recent years.In the Stewiacke River spawning occurs in a 6-kmstretch beginning about 3 km upstream of the con-fluence of the Shubenacadie and Stewiacke rivers(approximately 28 km upstream of Cobequid Bay).Both habitats are similar—tidally dominated, witha tidal bore greater than 10 cm in height on in-coming tides. At high tide, the areas are deep,relatively wide, silty, and brackish. In 1992 and1994, eggs were collected in this area at salinitiesfrom 0.0 to 22.8%o (Table 5) and at dissolved ox-


TABLE 4.—Extended.


Mean weight (kg) in age-class:

Sex 14 15 16 17 18 19 20 21 Source

Gulf of St. LawrenceBathurst Harbour

1978-1983Miramichi

1984Kouchibouguac

1978Kouchibouguac,

Kouchibouguacis.and Richibucto

1978

Annapolis1976

1978Gaspereau

1976Shubenacadie

19761992

Saint John1973-19751975-1977

C 7.50

C

C

CCF

Bay of Fundy

C 8.38 11.88 14.53 12.04 13.18

C 10.11 10.49 12.14 13.84 13.48 15.65

CMP

C 12.15 17.20 15.30 15.52

C 13.85 18.00 16.80

Madden (1984)

Walling (1985)

Jessop and Swetnam (1978)

Jessop and Swetnam (1978)

Jessop and Vithayasai (1979)Jessop (1980)

Jessop and Vithayasai (1979)

Jessop and Vithayasai (1979)

Rulifson and Jessop (this paper)

14.20 Dadswell(1976)

Adapted from Melvin (1978)

ygen levels of 6.6-13.8 mg/L (Rulifson, Wood,and Tull, unpublished data). At low tide, both areasbecome narrow, shallow channels lined by mudand sand flats (Rulifson et al. 1987). The Shuben-acadie-Stewiacke population is the only one doc-umented, throughout the species' range, as suc-cessfully reproducing in a tidal bore river. In theAnnapolis River, the primary spawning area is lo-cated above Bridgetown (km 32-40, measuredfrom the Annapolis River causeway; Williams etal. 1984). Before the Annapolis Tidal GeneratingStation began operation in August 1984, this areawas about 7 km upstream from the salt wedgewhere the stream was approximately 30 m wideand consistently 1.5-2 m deep. The bottom wasmainly sand interspersed between basalt and gran-ite rocks and boulders (Williams et al. 1984). Dis-solved oxygen in these waters during the spawningseason is 8.9-9.8 mg/L, pH is 6.6-7.4, hardnessis 20 mg/L, turbidity is 0.7 Jackson turbidity units,and alkalinity is 10 mg/L (Parker and Doe 1981).Historically, it is likely that the Annapolis Riverpopulation spawned successfully in a tidal-bore-dominated system, but the construction of a cause-

way at Annapolis Royal (Figure 10) eliminated thetidal bore phenomenon. In the Saint John River,spawning occurs in tributaries of Belleisle Bay,approximately 64 km above Reversing Falls(Dadswell 1976). The main spawning area, his-torically, was at the head of tide around Hart Islandabove Fredericton, about 140 km upstream fromthe river mouth and 65 km upriver from the limitof saltwater penetration (Adams 1873), but it isnow believed that spawning no longer occurs atthis site (Jessop 1990). Spawning may also occurin the Kennebecasis River (Hooper 1967; Melvin1991), a river similar to those in the Kouchiboug-uac region (Metcalf et al. 1976), which has darklystained, relatively clear waters of pH 6.5, littlesediment load and a gravel substrate.

Information on nursery grounds of Bay of Fundystriped bass is scattered and difficult to compare.For the Shubenacadie-Stewiacke river system, lar-vae were collected in 1919 by A. H. Leim (un-published archived notebooks) in the tidal regionof the lower Shubenacadie River, and age-0 fishhave been collected from the lower ShubenacadieRiver and Cobequid Bay (Figure 10). Some of the


FIGURE 10.—Overwintering sites, known and possible spawning sites, nursery areas, and seasonal movementsof striped bass in the Bay of Fundy region: Annapolis and Shubenacadie rivers, Nova Scotia, and the Saint JohnRiver, New Brunswick.

best evidence for striped bass reproduction in Can-ada was Leim's Shubenacadie River study in 1919.Age-0 striped bass were captured by seine betweenJune and August, during which time they grewfrom 12 mm to 80 mm total length (Figure 11;Dadswell et al. 1984). Age-0 fish 80-100 mm FLwere collected on the north shore of Cobequid Bayin August and September of 1985 (Rulifson et al.1987). Although striped bass spawning was ob-served in the Annapolis River before and after thetidal power project, only three larvae (Williams1978) and no age-0 fish have been collected in thesystem (Williams et al. 1984). In 1985 and 1986,juveniles 200-240 mm FL were caught by beachseine in the estuary above the Annapolis causewayduring a survey for juvenile Alosa species (Stokes-bury 1987). Nursery grounds in the Saint JohnRiver are undetermined.

The population structures of striped bass in theAnnapolis and Saint John rivers are similar, butboth have been quite different from the structuresof other Atlantic Canada populations since the

mid-1970s in that old fish represent a substantialportion of the population (Table 2). Surveys byPenney (1973), Williamson (1974), Jessop (1976),and others indicate that Annapolis River stripedbass have undergone considerable changes in sizeand age structure since the mid-1970s. Williamson(1974) reported that bass caught by anglers in1971-1972 ranged in age from 2 to 12, most fishbeing ages 4-6 (Figure 12). The 1972 populationhad an average age of 6, an average length of 558mm FL, and an average weight of 2.7 kg (Jessop1980). In 1975, Jessop and Doubleday (1976)found a substantial increase in the ages repre-sented; fish ranged from age 1 to age 17, and age-10 and age-12 fish were the most abundant amongolder fish (Figure 12). The average age had in-creased to 7.8, the average length was 660 mm,and the average weight was 3.9 kg. The trend toolder and larger fish continued the following year:average age was 9.4 years, average length was 763mm and average weight was 5.9 kg. By 1978, noyoung age-classes were present; the age range was


TABLE 5.—Water temperature (°C) and spawning activity, indicated by egg deposition per unit of sampling time, ofstriped bass in the salt-wedge-dominated Annapolis River in 1976 (Williams 1978) and 1979 (Parker and Doe 1981),the freshwater-dominated Belleisle Bay of the Saint John estuary in 1975 (Dadswell 1976), and the tidal bore-dominatedStewiacke River in 1992 (Rulifson and Wood, this paper).

Annapolis River. 1976

Date

May 3031

June 123456789

10I I1213141516171819

°C

17.019.218.416.017.518.018.519.019.520.022.021.021.518.516.015.517.519.619.519.721.0

Eggsper

10 min

15

446

1,428363

0

2

892

317

Annapolis River, 1979

Date

May 24253031

Jun 123456789

101112131415161718192021

°C

13.511.515.015.015.015.015.514.516.016.016.015.015.016.017.018.517.516.517.018.019.522.021.019.019.0

Eggsper

24 h

4510831

37349

1.305174

1,2012,8401.231

5125.450

510

9464

1,539259

1010

2236,005

24,98800

Belleisle Bay, 1975

Date

May 121314151617181920

°C

11.511.512.513.515.014.016.014.013.0

Eggsper

24 h

014034866142523345I I0

Date

May 2729

Jun 1

2

34

5678

9

Stewiacke River, 1992

Tide

EbbHighEbbLowEbbLowEbbHighEbbLow ebbEbbLowEbbLowLowHighLowMid ebbLowHighMid ebb

Salin-ity

(%o)

0.011.72.10.00.30.11.2

17.514.5

1.91.50.63.1O.I1.6

18.02.0

15.70.0

21.79.6

°C

14.712.414.013.718.218.417.514.615.516.518.118.816.316.517.616.420.519.122.016.719.6

Eggsper

5 min

0000

14783480

5444000

815I0

34325

40

4-18 or more (Figure 12), the average age was10.9 years, the average length was 703 mm, andthe average weight was 6.4 kg. The most abundantages had shifted to 8-10 years, representing thesame year-classes that had formed the largest pro-portion of the population in 1975. Jessop (1980)concluded that recruitment failure was responsiblefor the shifts in age structure of the Annapolispopulation. In 1987, another creel survey was con-ducted (Harris and Rulifson 1988), which revealedan age range of 3-21 years; over 60% of the in-dividuals were ages 4-6 (Figure 12). The averageage had decreased to 6.3 years, the average lengthto 703 mm, and the average weight to 4.2 kg.Aging remnants of the cohorts that had been abun-dant in 1978 and a deficiency of young year-classeswere evident (Figure 12). Weight frequencies ofstriped bass entered in the Dunromin Campgroundfishing contest each year at Annapolis Royal in-dicated that relatively large numbers of small fish(1-3.5 kg), probably of ages 3-7, were caught in1982, 1983, and 1986 (Figure 13). The dominanceof a second peak in weight frequency of 9-12-kgfish in 1982 was reflected in subsequent years

(through 1986) and may have been caused by amajor recruitment event. Harris (1988) concludedthat some recruitment in the Annapolis River oc-curred since the last creel survey but that relativelylarge numbers of age-3 fish may have recruitedfrom adjacent systems (e.g., the Shubenacadie-Stewiacke watershed) or regionally (e.g., AtlanticCanada and New England). Distributions of stripedbass age indicate that total mortality (Z) is lowerfor Bay of Fundy populations (0.15-0.23) than forpopulations in the Gulf of St. Lawrence (Table 3),which may indicate a high portion of older U.S.migrant fish.

The population structure of Saint John Riverstriped bass was examined from 1967 to 1976, buthas not been examined since. In 1967, Hooper(1967) reported fish ages of 1-9 years; over 50%of the catch was from the 1964 (age-3) year-class(Figure 14). In 1971-1972, Williamson (1974)found few age-1 and age-2 fish; the age structurehad expanded to age 13, and the 1966 and 1967age-classes (ages 4 and 5) dominated the catch. In1974, Dadswell (1976) found that fish ages were1-16, and several prominent age-classes were ev-

496 RUL1FSON AND DADSWELL

80

60

40

20

Striped BassShubenacadie River

1919

(Leim unpub. data)

n=1195

Hatch ?

May I June I July ' August '

FIGURE 11.—Growth (total length) of juvenile striped bass in the Shubenacadie River estuary during the summerof 1919, based on unpublished data collected by A. H. Leim (Dadswell et al. 1984).

ident (Figure 14). In 1975, a similar age range wasobserved, but the catch was dominated by age-3fish, and the year-classes of 1969 and 1966 wereprominent. In 1976, Melvin (1978) found a fairlyuniform distribution over ages 3 to 8, and age 11(1965 year-class) was prominent. This obviouslack of consistency in prominent year-classes sug-gested either sampling problems or mixing ofstocks. Tag returns from concurrent tagging stud-ies showed that mixed stocks were present; sometags were returned from as far away as Long Island(Dadswell 1976).

Age-frequency data from inner Bay of Fundyvary among locations and over time. Fish collectedfrom the Shubenacadie River spawning run in1976 (Jessop and Vithayasai 1979) were of ages2-7; age-3 fish (1973 year-class) were the mostabundant and the remaining age-classes were insimilar modest abundance (Figure 15A). Sixteen

years later, a much expanded age-frequency dis-tribution was evident for the spring 1992 spawningrun (Figure 15D). Ages ranged from 2 to 13, andage-5 fish were most abundant (Rulifson and Jes-sop, this paper). A broad age-class structure wasapparent in Shubenacadie Lake sampling con-ducted during October 1977 (Figure 15B); fish pre-paring for overwintering were primarily age 2, butseveral other age-classes were represented at lowabundance, including age-9 and age-11 fish (Mel-vin 1978). Fish younger than age 2 were not col-lected, most likely the result of gear selectivity.Sampling of the Minas Basin weir fishery duringthe summer and fall of 1985 (Rulifson et al. 1987)reflected the use of Minas Basin and Cobequid Bayas nursery grounds by immature fish. All stripedbass collected by a commercial weir in Minas Ba-sin were recorded; age-classes were estimatedbased on length and weight. The population ranged


r n r r n rn» ii ii ii ii ii ii I L »2 4 6 8 10 12 14 16 1$ 20

o1—-*4 6 8 16 12 i4 16 1g 20

Age (y r )

FIGURE 12.—Age-frequency distributions of striped bass angled in the Annapolis River between 1971 and 1987.Dotted lines connect possibly related cohorts. Data are from Williamson (1974), Jessop and Doubleday (1976),Jessop (1980), and Harris (1988).

from age 0 through age 5, and over half the fishwere age 1 (Figure 15C). Age-0 fish were col-lected, but the brush-type commercial weirs maynot retain them in proportion to their abundance,and the smallest individuals may serve as prey inreceding waters. The absence of fish older thanage 5 coincided with reduced striped bass stocksthroughout the range.

Striped bass from the Annapolis and Saint Johnrivers grow more quickly than fish from the St.Lawrence River (Figure 7) but more slowly thanKouchibouguac River fish (Table 1) and 2-3 cm/year less than U.S. populations. Females in theAnnapolis River are consistently longer at age thanmales; Saint John River females are shorter at agethan males until age 4, at which time they exhibitgreater growth. For both populations, fastestgrowth was exhibited in the third year of life,

which is similar to the Gulf of St. Lawrence pop-ulations. Shubenacadie River fish are larger andheavier at age than other Bay of Fundy populations(Tables 2, 4).

The only information on age of maturity andfecundity at age for Bay of Fundy populations isfrom the Saint John River. Williamson (1974) ex-amined 179 fish collected in May-August 1971and 1972. In that study, 34% of the males weremature: 25% of age-3 fish and 84% of age-5 fish.Only 24% of all females were mature: none at age3, 20% at age 4, 21% at age 5, 82% at age 6, and100% at age 7. No trend of increasing number ofeggs with fish size was evident. Williamson (1974)concluded that the full spawning potential of thesefish was not reached, especially for age-4 and age-5 females. The average number of eggs per femalewas 313,600 at age 5, 72,427 at age 6, 825,504 at


1982

2 4 6

WeightFIGURE 13.—Weight-frequency distributions of

striped bass angled in the Annapolis River and enteredin the annual Dunromin Campground fishing contest(Harris 1988). Dotted line connects a possibly relatedrecruitment pulse from the mid-1970s.

age 7, 742,976 at age 9, 526,848 at age I I , and1,064,320 at age 14. These fecundity estimates forSaint John River striped bass are higher than thevalues reported for Gulf of St. Lawrence fish (Ho-gans and Melvin 1984) but much lower than forstriped bass of similar ages from southern U.S.populations (Olsen and Rulifson 1992).

In general, local migrations are similar amongthe Bay of Fundy populations. Adults migratedownstream in the spring after spawning and enterocean waters, where their destinations are unde-termined. Subadults may remain within their riversystems or venture along the Bay of Fundy coast-line. In the Saint John River, summer upstreammigrations of feeding subadults (<2 kg) once ex-tended as far as the junction of the Aroostook Riv-er, about 320 km from the Saint John mouth (Jes-sop 1991). This migration now is limited by theMactaguac hydroelectric dam above Fredericton,150 km from the river mouth. In the ShubenacadieRiver, subadults and adults travel downstream toenter the Cobequid Bay, the easternmost extensionof the inner Bay of Fundy (Cobequid Bay). Largerfish leave the area, but smaller fish (<400 mm FL)remain to feed. In the fall, an unknown proportionof each population reenters estuaries to migrateupriver to overwintering sites (Figure 10). In theAnnapolis River, adults must exit the estuarythrough the turbines of the Annapolis Tidal PowerGenerating Station, then return to the river in thefall through causeway floodgates.

Striped bass migrating through Bay of Fundywaters move along shallow shorelines and pursuesmall fish prey into the mouths of tidal streams onflood tides. Young nonmigratory striped bass areprimarily benthic feeders, whereas larger fish arepiscivores (Rulifson and McKenna 1987). Thisfeeding activity results in a seasonal change inlipid composition and fat deposition from summerinto autumn (Gallagher et al. 1989).

Long Distance MigrationBased on tagging studies over 41 years (Vla-

dykov and Brousseau 1957; Magnin and Beaulieu1967; Rulifson et al. 1987; Table 6), the followinglong-range migratory patterns of Canadian stripedbass are apparent. For each population, a group oflocal fish migrate up and down the estuaries andinto regional waters during the summer. The St.Lawrence population was endemic and did not ex-hibit long-distance migration. In the Gulf of St.Lawrence and Bay of Fundy populations, largerfish of unknown origin participate in the fall up-


40

20

Oi

40

20

1967Saint John River

Trap Nets= 57.11. :

I o

1971-72Trap 8 Gill Nets

1n = 357$ = 149

20r 1974Gill Nets (A l l meshes)

^°ri974 _ n f ° 4

f • I I - • • rl° • I •

20

Gill Nets1975 I Gl" N6t

I I " = 5?• Mil • •20 P1976

Gil l Netsi.i.... i8 10

Age (Years)12 14 16

FIGURE 14.—Age-frequency distributions of striped bass captured in the Saint John River estuary during fisheriesinvestigations between 1967 and 1976 (after Dadswell et al. 1984).

stream migration and overwinter in freshwaterhabitats of eastern Canada.

Some mixing of Canadian and U.S. populationsis evident, but the proportion of the exchange isgenerally unknown. Boreman and Lewis (1987)reported that striped bass tagged by American Lit-toral Society members along the U.S. eastern sea-board as far south as New Jersey have been re-captured in the Canadian Maritimes. Potomac Riv-er (Maryland) and Hudson River (New York) re-leases have been recaptured in the Annapolis Riverand in Minas Basin (Table 7). Striped bass taggedin Nova Scotia and New Brunswick waters havebeen recaptured as far south as New Jersey, Del-aware, Virginia, and North Carolina (Table 7;Nichols and Miller 1967).

Striped bass tagged in eastern Canada are ca-pable of migrating long distances in a matter ofdays. Williamson (1974) released a striped bass inWestfield, New Brunswick, in September 1972 thatwas recaptured in the Black Stone River, Rhode

Island, in October, 36 d later. A striped bass re-leased in Minas Basin in September 1985 was re-captured 45 d later at Newport, Rhode Island, adistance of 800 km representing an average mi-gration rate of 18 km/d (Rulifson et al. 1987).

Long-distance exchange is not limited to Bay ofFundy waters but also occurs to an unknown extentwith fish from the Gulf of St. Lawrence. Hogansand Melvin (1984) tagged striped bass under theice of the Kouchibouguac River in early March1983; one tagged fish was recaptured 33 d later inthe Wye River, Maryland. It covered a distance ofapproximately 1,700 km, an average migration rateof 51 km/d (Table 7).

Stock IdentificationSeveral studies have been undertaken to deter-

mine whether striped bass populations in AtlanticCanada are genetically separate or mix with U.S.fish and to ascertain possible introgression of U.S.fish into Canadian stocks in some years. Several


80

60

40-

20-

080 -,

60-

40-

20-

0

> 80~U4) 60-

crJJ 40-U.

20-

0

60-

40-

20-

B

Shubenacadie R.1976n-44

Shubenacadie L1977n-85

16

Minas Basin1985n-919

Shubenacadie R.1992n«4l

0 2 4 6 14 16Age (yr)

FIGURE 15.—Age-frequency distributions of stripedbass captured in the inner Bay of Fundy: (A) 1976, an-gled in Shubenacadie River (Jessop and Vithayasai1979); (B) 1977, gillnetted at the overwintering site inShubenacadie Lake (Melvin 1978); (C) 1985, caught bycommercial brush weirs in Minas Basin (after Rulifsonet al. 1987); (D) 1992, commercially gillnetted onspawning grounds of the Shubenacadie River (Rulifsonand Jessop, this paper).

types of analysis have been used: meristic andmorpheme trie, parasite community, blood serum,and mitochondrial DNA (mtDNA).

Only a few meristic characters are commonamong the several studies of Canadian popula-

tions; these include soft ray counts of the dorsal,anal and pectoral fins; a character index (CI), thesum of the dorsal, anal, and pectoral fins raycounts; and lateral line scale counts (Table 8).None of the meristic characters can uniquely iden-tify each Canadian population. The ranges of me-ristic characters are similar among populations(Table 8). Melvin (1978) statistically analyzedthese characters and concluded that the lateral linescale count separated Saint John River striped bassfrom Shubenacadie River and Tabusintac Riverfish, which had similar counts. Melvin (1978) hy-pothesized that introgression of U.S. fish into thestriped bass populations of the Saint John and Shu-benacadie rivers during his study may have influ-enced the results. To test this hypothesis, Melvinexamined Chesapeake Bay fish for meristic char-acters; Chesapeake fish had a CI count and analand pectoral fin ray counts similar to those for fishcollected from the Saint John and Shubenacadierivers. Fish collected from the Tabusintac River,however, were significantly (P < 0.01) differentfrom Chesapeake Bay fish for all meristic char-acters except pectoral soft fin rays.

Melvin (1978) also used several morphometricrelationships in an attempt to discriminate amongpopulations in the Saint John, Shubenacadie, andTabusintac rivers (Table 9). Results indicated thatseveral of the relationships could separate the pop-ulations independently, but there was no consis-tency of multiple variables in separating the threepopulations.

Biochemical methods have been used to sepa-rate Canadian from U.S. striped bass. Melvin(1976, 1978) used protein electrophoresis withoutsuccess to distinguish fish collected from the Ta-busintac, Shubenacadie, and Saint John rivers.Schill and Dorazio (1990) were able to distinguishnorthern striped bass (Stewiacke, Miramichi, Ri-chibucto, Tracadie and Hudson rivers) from south-ern striped bass (Chesapeake Bay and RoanokeRiver) based on antigenic differences in blood se-rum albumin. Wirgin et al. (1993) used restrictionfragment length polymorphism (RFLP) analysis ofmtDNA collected from juvenile (nonmigratory)striped bass to separate populations. Differencesin frequencies of mtDNA length variants (sizepolymorphisms) separated Shubenacadie Riverfish from Miramichi River and Tabusintac Riverfish, and results from all three Canadian sourceswere different from those of the U.S.-spawned At-lantic coastal migratory stock. Wirgin et al. (1993)supported Melvin's (1978) conclusion that Gulf ofSt. Lawrence fish may be isolated from Bay of


TABLE 6.—Summary of striped bass tagging programs in New Brunswick and Nova Scotia.

Number of fish

Tagging area

Bathurst Harbour

Kouchibouguac River

Annapolis River

Minas Basin

Cobequid Bay

Saint John River

Southwest Nova Scotia

Year

198619781983

19751976198119821987

1985

1986

1985

1986

19661969

1972-1975

1966

Tagged

3978

597

944371557

453

I I

710

257

noKM)70

200

Re-covered

40

02454

131

4

112

26

81013

13

U.S.origin

00100002

1

0

3

0

012

3

Tag typea

Roy FD-68BC

ALSFloy FT-I

Floy FD-67Floy FD-67Floy FT-IFloy FT-IFloy FT-I

Floy FT-I

Floy FT-I

Floy FT-I

Roy FT-1

ALSALSALS

ALS

Source

Meagheretal. (1987)Melvin (1978)Hogans and Melvin (1984)

Jessop(1980)Jessop(1980)Dadswell et al. (1984)Dadswelletal. (1984)Harris (1988)

Rulifson and McKenna(this paper)

Rulifson and Langille(this paper)

Rulifson and McKenna(this paper)

Rulifson and Langille(this paper)

Moss (1971)Moss (1971)Williamson(l974);

Dadswell (1976)Moss (1971)

1 ALS is American Littoral Society spaghetti tag.

Fundy populations by the cold waters off CapeBreton and the north Atlantic Ocean.

The community of parasites in a striped bassmay assist in determining its range of ocean mi-gration or site of origin. Hogans (1984) examinedthe parasite communities of striped bass 3-6 yearsold collected from the Kouchibouguac River inSeptember and October 1979. Results suggestedthat several fish either were of U.S. origin or wereCanadian fish that had visited U.S. waters. Sixparasite species were present, three of which wereof freshwater origin: Homalometron pallidum,Neoechinorhynchus rutili, and Philometra rubra(Table 10). The remaining three parasites were ofmarine origin: Lepocreadium setiferoides, Ste-phanostomum tenue, and Echinorhynchus gadi.Three species—L setiferoides, S. tenue, and P. ru-bra—had not been reported for striped bass, or anyother fish species, in Canadian waters but havebeen reported from striped bass in Chesapeake Bay(Paperna and Zwerner 1976). Linton (1901) hadreported E. gadi from striped bass collected in theGulf of Maine. The remaining parasite specieswere new host records (Hogans 1984). Additionalevidence for long-distance migration was found inMay 1983, when Hogans (1985) reported that ap-proximately 20% of striped bass 4-6 years oldcollected from the Kouchibouguac, Kouchiougua-

cis, and Black rivers were infected with Ergasiluslabracis (Table 10). Also, two male striped bass(ages 4 and 5) collected in the same season (1983)from Minas Basin (inner Bay of Fundy) were in-fected with E. labracis. Hogans's (1985) obser-vations extended the range of E. labracis into Ca-nadian waters, possibly a result of striped bassmigrating between Canada and coastal mid-Atlan-tic waters. Paperna and Zwerner (1976) had re-ported this parasite on striped bass in the PotomacRiver, and Wilson (1932) found E. labracis onstriped bass caught off Woods Hole, Massachu-setts. Additional evidence for long-distance mi-gration was found in the parasite communities ofstriped bass collected from the Bay of Fundy in1985. Rulifson et al. (1987) examined parasites of80 striped bass from Minas Basin, of which 72%were infected with one or more taxa of parasites.Nematodes were most prevalent followed by acan-thocephalans and trematodes; few fish were par-asitized with the crustacean species of isopods,copepods, and branchiurans (Table 10). Nematodeinfestation was greater for locally produced age-0 fish shorter than 94 mm FL (88%) than for thewhole population (60%), which could include non-infected fish from southern waters. Acanthoceph-alans were found in 22.5% of all fish age 1 andolder, whereas no acanthocephalans or trematodes


TABLE 7.—Long-distance migration of striped bass in Atlantic Ocean coastal waters between the USA and Canada.

Tagging site Release date Recapture site3 Recapture dateDays at

large Source

Tagged in the United StatesMaryland

Potomac River

Nanticoke RiverNew York

Hudson River

Feb 1959Apr 1960

Apr 1961Apr 14, 1973

Apr 1, 1986UnknownUnknownUnknown

Walton, Minas Basin, NSBear River, Annapolis

basin, NSAnnapolis Royal, NSReversing Falls, NB

Sep 1959Jul 1960

Jul 1961

Oct 25, 1976

Bear River, NS Aug 6, 1986Annapolis River, NS Summer 1987Annapolis River, NS Summer 1987Annapolis River, NS Summer 1987

Tagged in Canada

159 Nichols and Miller (1967)99 Nichols and Miller (1967)

127 Nichols and Miller (1967)

1,279 J. Booneb

128 J. Waldmanc

Unknown Harris (1988)Unknown Harris (1988)Unknown Harris (1988)

New BrunswickDarlings Lake

Westfield RiverReversing FallsKouchibouguac

RiverNova Scotia

Cheboque River

Annapolis River

Minas Basin

Cobequid Bay

Jun 5, 1969

Sep 12, 1972Aug 7, 1973Mar 3. 1983

Aug 12, 1966Sep 4. 1966Jul 13, 1969Jul 13, 1969Aug 29, 1966Jul 14, 1975Aug 5, 1987

Oct 12, 1987Sep 19, 1985Oct 14, 1985Jun 28, 1985Jul 2, 1985

Jul 4, 1985Oct 17. 1985

Montauk. NY

Black Stone River, RISouthampton, NYWye River, MD

Patcong Creek, NJIndian River, DESakonnet River, RILong Beach Island, NJRockingham, NCPotomac River, VAOld Lymc, Connecticut

River, CTLong Island Sound, NYPrices Neck, Newport, RILavallette, NJAssateague Beach, VAShetucket River, Norwich,

CTThames River, Norwich, CTHerring Cove. Cape Cod. MA

Nov 19, 1969

Oct 23, 1972Nov 5, 1973Apr 12, 1983

Jul 15, 1967Mar 13, 1967Jun 10, 1970Apr 22. 1970Jun 24, 1967Mar 25, 1976Oct 28, 1987

Nov 9, 1989Nov 3, 1985Mar 15. 1986Apr 2, 1988May 3, 1986

Apr 19. 1986Jun 22, 1986

167

369033

33717834228330825548

39245

1521,009

305

299248

American Littoral Society5

(1976)Williamson(1974)Dadswell(l976)Hogans and Melvin (1984)

Moss (1971)Moss (1971)Moss (1971)Moss (1971)Moss (1971)ALS (1976)d

Harris (1988)

Harris (1988)Rulifson et al. (1987)Rulifsonet al. (1987)Rulifson (this paper)Rulifsonet al. (1987)

Rulifson et ai. (1987)Rulifson etal. (1987)

u Geographical abbreviations are: CT. Connecticut; DE, Delaware; MA. Massachusetts; MD, Maryland; NB, New Brunswick; NC, NorthCarolina; NJ, New Jersey; NS, Nova Scotia; NY, New York; RI, Rhode Island; and VA, Virginia.

h Maryland Department of Fish and Game, personal communication.c Hudson River Foundation, personal communication.d Returns of the American Littoral Society (ALS) spaghetti tags have been reported in that Society's publication The Underwater Naturalist.

were found in age-0 fish. Parasitic crustaceans arecommon in U.S. waters (Paperna and Zwerner1976) but were rare in Minas Basin fish. Resultsof the three Canadian studies suggest that ratherthan a range extension of the parasites, the infectedfish were parasitized during long-distance migra-tion to the eastern U.S. seaboard or the parasitesoccurred in striped bass that had migrated to Can-ada from U.S. populations.

DiscussionGeneral conclusions can be drawn about the life

history of Canadian stocks of striped bass fromthe information available. Growth of Canadian

striped bass is slower than that of U.S. fish; amongCanadian populations, greatest growth rates occurin the Gulf of St. Lawrence, perhaps a result ofthe warmer coastal habitat provided by North-umberland Strait (Scott and Scott 1988). Age ofmaturity in Canada is about 1 year older than inthe USA. Maximum size and age of Canadianstriped bass are less than those of U.S. populations;fish from Gulf of St. Lawrence populations havethe lowest maximum age and total length but thehighest total mortality. Recruitment success ap-pears to be intermittent in Canadian rivers andoften does not coincide with that of U.S. stocks.

Overwintering of striped bass in ice-covered


TABLE 8..—Ranges of selected mcristic characters of striped bass sampled from eastern Canada. Hudson River andChesapeake Bay data are presented for comparative purposes. Character index (CI) is the sum of the dorsal, anal, andpectoral fin rays (mean ± SD). Sexes are combined.

Samplingarea

Tubusintac RiverMinas Basin-

Cobequid BayAnnapolis RiverShubenacadic LakeSaint John River

Hudson River

Chesapeake Bay

Soft fin rays

Year

1975-19771985

1971-19721975-19771971-19721975-19771936-1949

1807-1949

N

10879

2885

10562

242

237

Dorsal

10-1211-13

10-1211-1210-1211-129-12, 14

10-14

Anal

9-1210-11

10-12

10-119-11

10-12

Pectoral

30-3929-36

28-3632-3831-3530-3828-35

30-37

CI

55.3 ± 1.8656.1 ± 0.28

56.6 ± 1.62

56.3 ± 1.9354.0 ± 1.64

56.3 ±1.41

Lateralline

scales

54-6352-66

56-62

53-69

Source

Me! v i n < 1978)Rulifsonetal. (1987)

Williamson(1974)Melvin(l978)Williamson(1974)Melvin(l978)Raney and de Sylva

(1953)Rancy and de Sylva

(1953)

fresh orestuarian waters is common in these north-ern latitudes. Overwintering fish begin their down-stream journey before ice leaves the streams, andthey are met by striped bass migrating upstreamfrom the sea. Spawning occurs just upstream ofthe salt wedge (except perhaps in the Saint JohnRiver) in late May and June, several weeks afterthe ice has disappeared. Larvae and age-0 fish usebrackish waters as nursery areas; the adults returnto coastal ocean waters.

Striped bass in coastal waters of the Gulf of St.Lawrence are probably of local origin, but somefish of U.S. origin may occur; parasite communitystudies suggest that some fish have resided in mid-Atlantic waters for an undetermined period. Ingeneral, the striped bass found in Gulf of St. Law-rence waters represent Canadian fish of mixed lo-cal stocks that migrate along coastal areas of east-ern New Brunswick northward to Chaleur Bay dur-

ing the summer. These fish then exhibit a quicksouthward migration in late summer and move up-stream in September and October for overwinter-ing. Bay of Fundy populations are mixed with fishof U.S. origin in most years.

Extensive upstream migration of large fish beginsin late summer and continues through October inthe Saint John, Annapolis, and Shubenacadie rivers.In the Saint John River, fish that had moved up-stream to feed move downstream to overwinter. Ma-jor overwintering grounds include Belleisle Bay ofthe Saint John estuary and Shubenacadie Lake inNova Scotia. Spring migrations in the ShubenacadieRiver involve upstream migrants from ocean watersand downstream overwintering migrants from Shu-benacadie Lake. Striped bass frequent inner Bay ofFundy shores in May and early June as they moveto sea, then return to rivers in the overwinteringmigration in late August through October.

TABLE 9.—Ratios of selected morphometric characters of striped bass populations in the Saint John estuary, Shuben-acadie Lake, and Tabusintac River (from Melvin 1978). Ratios are presented as ranges; means ± SDs are given inparentheses.

Ratio

Body depth : fork length

Head length : fork length

Eye diameter : head length

Snout length : head length

Jaw length : head length

Caudal peduncle : body depth

Saint John estuary(N = 62)

0.17-0.29(0.227 ± 0.021)

0.20-0.30(0.256 ± 0.016)

0.08-0.16(0.122 ± 0.017)

0.26-0.35(0.298 ± 0.020)

0.27-0.42(0.349 ± 0.028)

0.32-0.48(0.397 ± 0.032)

Shubenacadic Lake(N = 85)

0.21-0.26(0.246 ± 0.010)

0.24-0.28(0.262 ± 0.007)

0.10-0.16(0.139 ± 0.012)

0.25-0.32(0.281 ± 0.013)

0.26-0.42(0.356 ± 0.020)

0.33-0.42(0.377 ± 0.015)

Tabusintac River(N = 108)

0.21-0.29(0.254 ± 0.013)

0.25-0.31(0.276 ± 0.009)

0.11-0.25(0.205 ± 0.038)

0.23-0.32(0.274 ± 0.017)

0.29-0.38(0.347 ± 0.018)

0.31-0.42(0.358 ± 0.018)


TABLE 10.—Parasites of striped bass in the Canadian Maritimes. Capture sites Kouchibouguac River (KR) and MinasBasin (MB).

Parasite

TrcmatodaHomalometron pallidiumLepocreadium setiferoidaPStephanostomitm tenue0

"Black spot"Acanthocephala

Echinorhynchus gadiNeoechinorhynchus rutiliUnidentified

NematodaPhilometra rubrcPPhilometridae

CrustaceaErgasilus labracisLernaea sp.cArgulus sp.Idata ballica

Fishcapture

site

KRKRKRMB

KRKRMB

KRMB

KRb

MBMBMB

Infectionsite

StomachIntestineIntestineExternal

IntestineIntestineIntestine

Body cavityBody cavity:

all fishfish <94 mm FL

Gill archGill, mouthExternalLiver

Prevalence(percent)

58.852.941.016.2

41.070.522.5

82.3

60.088.0

20.31.36.21.3

Intensity(number/fish)

Mean

2.53.02.77.6

2.77.24.6

21.8

16.337

1I

Range

1-71-91-61-40

1-61-131-12

13-36

2-54

Source

Hogans (1984)Hogans (1984)Hogans (1984)Rulifson et al. (1987)

Hogans (1984)Hogans (1984)Rulifson et al. (1987)

Hogans (1984)Rulifson et al. (1987)

Hogans (1985)Rulifson etal. (1987)Rulifson ct al. (1987)Rulifson and McKenna

(this paper)u Reported otherwise only from Chesapeake Bay.hTwo male striped bass collected from Minas Basin in the same season (1983) were also infected with E. labracis (Hogans 1985).c Lcmaea sp. of Rulifson et al. (1987) may have been Ergasilus labracis.

Tag returns indicate that striped bass tagged inU.S. waters overwinter both with Bay of Fundyfish (freshwater) and with Gulf of St. Lawrencefish (estuaries); the natal origin of these fish isunknown. The incidence of U.S. fish mixing withCanadian stocks probably fluctuates with the pop-ulation size of the U.S. coastal migratory stockand the prevailing water temperatures in the Gulfof Maine, Bay of Fundy, and Gulf of St. Lawrence.

Information concerning striped bass populationsand life history in eastern Canada remains incom-plete. Before substantial management progress canbe made, an in-depth, wide-ranging investigationis required that would examine these populationsconcurrently and provide for extensive tagging ofstriped bass from all populations. Anecdotal in-formation from commercial fishing and anglingsuggest there are more populations than reportedhere, particularly in Nova Scotia.

AcknowledgmentsWe thank the students, commercial fishermen,

anglers, fisheries officers, and scientists who freelyshared their information on striped bass stocks inCanada. We especially thank S. McKenna and ourgraduate students G. Melvin, P. Harris, K. Stokes-bury, K. Tull, and L. Paramore. B. Jessop and W.Hogans shared their unpublished data with us and

helped with invaluable comments. Editorial com-ments of R. Bradford and C. S. Manooch on thedraft manuscript were greatly appreciated. Most ofour research was conducted in conjunction withother projects funded by the Canada Departmentof Fisheries and Oceans, Parks Canada, the Na-tional Science and Engineering Council, the U.S.National Marine Fisheries Service, and the TidalPower Corp.

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Received August 22, 1993Accepted December 28, 1994

life history and population characteristics of striped bass in atlantic canada

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