North American Journal of Fisheries Management 15:680-687, 1995© Copyright by the American Fisheries Society 1995

An Overview of the Smallmouth Bass in Nova ScotiaALAN J. MCNEILL

Nova Scotia Department of Fisheries, Aquaculture and Inland Fisheries DivisionPictou, Nova Scotia BOK JHOt Canada

Abstract.—Introduced to Nova Scotia waters in 1942, smallmouth bass Micropterus dolomieuhave provided an alternative fishery to the more popular brook trout Salvelinus fontinalis in localareas. In recent years, angler interest in smallmouth bass has increased dramatically, and a modesttournament fishery has developed in addition to noncompetitive angling. Since 1971, illegal trans-fers and movements of fish within watersheds have been responsible for approximately doublingthe distribution of this species over the past 15 years. Estimated annual catch and average catchper angler have increased 32-fold over the same time period. Population structure and length-at-age data from biological surveys in 1992 and 1993 indicated that growth was slower than in othernorthern waters. Observations of smallmouth bass nesting activity suggest that nest building andspawning commence when water temperature reaches 16-18°C, which typically occurs at the endof May or first week of June in Nova Scotia, and nest guarding is terminated by early July.Management practices focus on restricting the illegal introduction of smallmouth bass to newwatersheds and sustaining a quality fishery where smallmouth bass are established.

The northern and eastern limits to the nativerange of smallmouth bass Micropterus dolomieuwere bounded by the Appalachian range (Mac-Crimmon and Robbins 1975); thus there are noindigenous populations in New England or theMaritime Provinces of Canada. Smallmouth basswere first introduced in Nova Scotia via NewBrunswick from stock originating in Maine. Thefirst appearance of smallmouth bass in the Mari-time Provinces was in the Chiputniticook Lakeson the New Brunswick-Maine border around 1870

.(Can 1949). It has been suggested that they werestocked from a private pond in Maine by a small-mouth bass angling enthusiast (Catt 1949). Overthe next 50 years they were introduced and estab-lished self-sustaining populations in several wa-tersheds in southwestern New Brunswick, includ-ing the Kennebecasis, Lower St. Johns, Magagua-davic, and St. Croix rivers (Smith 1942; Catt1949). These populations provided the seed stockfrom which all Nova Scotia introductions were de-rived. No records indicate that smallmouth basswere ever introduced directly from their indige-nous range.

The first successful planting occurred in 1942when 107 adults were taken from Wheatons Lake,New Brunswick, to Bunker Lake, Yarmouth Coun-ty, Nova Scotia (Catt 1949). During 1944-1953,10 lakes in Nova Scotia were stocked with small-mouth bass, 6 of which developed sustaining pop-ulations (Catt 1949; Gilhen 1974). After 1953 gov-ernment-sanctioned stocking of smallmouth bassceased until 1967. During that interim, lake surveyrecords and angler reports revealed that unauthor-

ized introductions or natural dispersal within wa-tersheds increased the number of lakes inhabitedby smallmouth bass.

In 1967 and 1968, the Nova Scotia Departmentof Lands and Forests, with the approval of theFederal Department of Fisheries and Oceans(DFO) and the Nova Scotia Light and Power Co.Ltd., introduced smallmouth bass into LumsdenPond, Black River Lake, Gaspereau Lake, andMethals Lake in Kings County (Bain 1993). Theseand other lakes, by virtue of a hydroelectric powerdevelopment, form what is known as the Gasper-eau-Black River Complex and have developedinto the premier smallmouth bass fishery in theprovince. In 1971, Spectacle Lake, Digby County,was stocked, and a successfully reproducing pop-ulation was established in coexistence with chainpickerel Esox niger. The most recent smallmouthbass introduction occurred in 1984, when MurphyLake, Kings County, in the Gaspereau-Black Riv-er Lake Complex was stocked (B. Sabean, NovaScotia Department of Natural Resources, personalcommunication).

Although unauthorized introductions of small-mouth bass are poorly documented, distributionaldata from documented introductions, historicallake survey and creel survey data, and interviewswith outfitters, government biologists, and localanglers indicated an increase in the waters inhab-ited by smallmouth bass during 1942-1993. Hebdaet al. (1990) compiled much of this informationfrom a wide variety of sources, including inter-views with operators of sporting goods stores andtackle supply shops, and an angler survey ques-

680

SMALLMOUTH BASS IN NOVA SCOTIA 681

[̂ Watersheds BLakes]

80

70

60

8 50

0§ 40

8O 30

20

10

1945 1950 1955 1960 1965 1970 1975 1980 1985 1990

YearFIGURE 1.—Occurrence of smallmouth bass in watersheds and lakes in Nova Scotia, 1942-1993.

tionnaire distributed to members of 32 fish andwildlife groups in Nova Scotia.

Reliance on angler reports incorporates error inspecies distributional data. The most common er-ror is species misidentification. In Nova Scotiasmallmouth bass are routinely confused with whiteperch Morone americana, striped bass Morone sax-atilis, and to a lesser degree, yellow perch Percaflavescens. For some years between 1966 and 1992,general fishing license stub returns reported small-mouth bass catches in all 18 counties, even thoughsmallmouth bass have probably never existed in 9of the 18 counties. A second error associated withangler reports is misidentification of the waterbody, which sometimes happens when there aredifferent local and accepted gazetteer names for alake or when several lakes in the same county sharethe same name. Additionally, when angler inter-views are relied on to document the year of anillegal introduction, anglers may recall the yearthey first captured a smallmouth bass, not (unlessthey were personally involved) the year of the ac-tual introduction.

The Nova Scotia Department of Fisheries(NSDOF) adopted the ranking system of Hebda etal. (1990) to record distributional data for small-mouth bass in Nova Scotia. A rank is assigned toeach record of smallmouth bass presence accord-ing to the source of information. Ranks, in orderof decreasing confidence, were (1) confirmed sur-vey data, (2) reports from two or more independent

anglers, and (3) reports from one angler or partyof anglers. No attempt was made to assign a greateror lesser ranking to an angler report based on thecompetence of the individual, although the obser-vations of a knowledgeable and experienced anglermay be as valid as scientific survey data (Hebdaet al. 1990). Because level two ranking has neverbeen repudiated by a scientific survey, these re-ports are considered accurate.

From 1955 to 1979, the majority of "new"smallmouth bass lakes probably resulted from nat-ural colonization through contiguous waters in thesame watershed. Only three new watersheds werecolonized by smallmouth bass, two of them as aresult of government stocking programs. From1980 to 1993, illegal transfers increased the num-ber of watersheds inhabited by smallmouth bassfrom 9 to 23 (Figure 1). With the exception ofMurphy Lake in 1984, the most recent govern-ment-sanctioned introduction occurred in 1971.Since 1971 the number of lakes inhabited by small-mouth bass has approximately doubled, growingfrom 34 to 71.

The FisheryRecreational fishery catch and effort statistics

were compiled from several sources by theNSDOF and (before 1987) by the Nova Scotia De-partment of Natural Resources. Since 1966, vol-untary angler returns of a survey stub attached tothe general fishing license have provided annual

682 McNEILL

total catches by species and county, and since1990, effort by species. License stub return rateshave ranged from 3.4 to 19.1% of total licensesales and averaged 6.6% between 1971 and 1990.The use of voluntary license stub returns may in-corporate bias from nonreporting or underreport-ing by unsuccessful anglers, or overreporting byavid anglers (Claytor and O'Neil 1991). However,if it is assumed that such bias is consistent overtime, the trends evident in the data may be reliableeven if the absolute values of the catch informationare not. Between 1971 and 1990, 30% of respon-dents caught no fish or did not fish, which suggeststhat avid anglers were not the only ones returninglicense stubs.

A second source of catch information for therecreational fishery in Nova Scotia is the Surveyof Sportfishing in Nova Scotia (sportfishing sur-vey), a part of a national and provincial series ofangler surveys published every 5 years, beginningin 1974 (Gumming 1979; DFO 1984, 1988, 1994).In addition to catch by species, the sportfishingsurvey includes information on licensing trendsand angler preferences, effort, motivation, expen-ditures, and investments. Participants in the surveyare chosen at random from the current year's li-cense vendors' stubs and stratified by license typeand nationality (e.g., general or salmon license,Canadian resident, Canadian nonresident, or U.S.nonresident). The survey is coordinated by theDFO and the provincial agency responsible for rec-reational fisheries management. Information iscollected by means of a questionnaire that ismailed about 4 months after the end of the fishingseason; a reminder notice and second question-naire are sent after an additional 30 days. Ques-tionnaire returns average about 70%.

Although an avidity bias (Thompson 1991) maybe present in the sportfishing survey, such bias isprobably less pronounced than in the license stubdata because the survey incorporates random sam-pling. On the other hand, most license stubs arereceived within 2 months of the end of the anglingseason, whereas the sportfishing survey question-naire is generally filled out by respondents 4-6months after the season ends. Fisher et al. (1991)reported that in general, estimates appear to in-crease with the length of recall period. In NovaScotia, total catches calculated from the sportfish-ing surveys have always been higher than thosecalculated from license stubs, although the trendsare similar.

Total estimated smallmouth bass catches re-ported from license stub returns increased during

200

160

160

140

120100

80

6040

20I0

2500

: Smallmouth bass

20 year mean /\

20 year mean

FIGURE 2.—Estimated total catch of smallmouth bassand brook trout by anglers in Nova Scotia, 1966-1992.Estimates were made from voluntary license stub re-turns. The 20-year mean is for 1971-1990.

1966-1992. There was a substantial increase incatch beginning in 1987, and the 1992 catch was4.2 times greater than the 20-year average (1971-1990; Figure 2). In contrast, the estimated catchfor brook trout Salvelinus fontinalis (the most pop-ular game fish in Nova Scotia) decreased after1983 and in 1992 was 36% lower than the 20-yearaverage. In 1992, average catch per angler forsmallmouth bass was four times greater than the20-year average, whereas for brook trout, the av-erage catch per angler was 20% lower than the 20-year average. Although in 1992 the average catchper angler for smallmouth bass was much lowerthan that for brook trout (3.1 versus 16.6), themajority of anglers were targeting brook trout;smallmouth bass were only targeted in certain ar-eas. Angler log books indicated that in most cases,catch per hour for smallmouth bass was consid-erably higher than for brook trout.

Sportfishing survey data indicated an evengreater increase in smallmouth bass catches during1974-1990 (Table 1). The catch for 1990 was 32times that estimated for 1974. In 1974, smallmouthbass ranked eighth in total catches for sport fish,whereas in 1992 only brook trout and rainbowsmelt Osmerus mordax were caught in greaternumbers (Gumming 1979; DFO 1994). Addition-ally, the proportion of smallmouth bass released

SMALLMOUTH BASS IN NOVA SCOTIA 683

TABLE 1.—Numbers of licensed anglers, estimated total catch, estimated retained catch, percent released, and annualaverage catch per angler for smallmouth bass and brook trout from the Nova Scotia sportfishing survey, 1974-1990;NA = data not available.

Year

1974

1980

1985

1990

Licensedanglers

73,419

82,449

78,095

68,276

Number of fish

Species

Smallmouth bassBrook troutSmallmouth bassBrook troutSmallmouth bassBrook iroutSmallmouth bassBrook trout

Caught

13,8001,706,500

NA2.712,503

81,0352,295,695

439,0742,035,523

Retained

NANANA

2.541.68718,007

1,711,87515,482

1,232,482

Percentreleased

NANANA

678259639

Average catchper license

0.223.2NA32,9

1.029.46.4

29.8

by anglers increased from 78% in 1985 to 96% in1990; in contrast, brook trout anglers released 25%of their catch in 1985 and 39% in 1990.

TournamentsThe increased interest in smallmouth bass an-

gling has given rise to a modest tournament fisheryin recent years. A tournament is defined as havingat least 25 participants who compete according toa set of regulations designed to test fishing prow-ess. Winners are ranked according to the totalweight of successfully released fish. The numberof fish allowed for weigh-in is usually six, andpenalties are assessed for dead fish. When mini-mum length limits are enforced, they are either254 or 305 mm total length. Although tournamentsare currently organized by a number of groups,they have similar regulations, and many share thesame sponsors. The number of tournaments hasgrown from a single tournament that has operatedannually since 1982 to 31 events in 1995. Thesetournaments have met with little opposition fromconservation groups or local residents, other thanthe complaints of litter and minor property damagenormally associated with large groups. TheNSDOF has adopted a "hands off" approach totournament fishing, provided general angling reg-ulations are observed.

A number of problems exist with the collectionof catch data and population structure informationfrom smallmouth bass tournaments. Willis andHartmann (1986) reviewed surveys from Shupp(1978) and Duttweiler (1985), both of whom re-ported that a majority (64 and 72%) of respondingstate agencies only occasionally collected datafrom tournaments. Problems associated with sta-tioning employees at a large number of tournamentweigh-ins, compounded by the bias incurred fromlength limits, may account for the languid effort

by state conservation agencies (Willis and Hart-mann 1986). In Nova Scotia, catch per unit effortand structural indices derived from tournamentcatches have provided relative comparisons ofabundance and population structure between lakes,or trends within one lake over time.

Smallmouth Bass Growth and ReproductionSmallmouth bass exhibit a wide range of growth

rates throughout their native and naturalized rangein North America. Length at age of smallmouthbass from various locations has been reported byseveral authors (Scott and Crossman 1973; Chanceet al. 1975; Coble 1975; Garrett 1983; Bennett etal. 1991; Dunsmoor et al. 1991). Notwithstandingsome exceptions, growth of smallmouth bass isinversely related to latitude.

Nova Scotia is the easternmost extent of thenaturalized range of this species and is close tothe northern limit. The average number of frost-free days in Nova Scotia ranges from 110 to 170(Dzikowski 1985) in areas inhabited by small-mouth bass. Nova Scotia lakes can be classified asrelatively unproductive (Alexander et al. 1986).Because of the short growing season and unpro-ductive environmental conditions, smallmouthbass grow slowly in Nova Scotia.

In 1992 and 1993 smallmouth bass were col-lected from Black River Lake (Kings County),Grand Lake (Halifax County), and Elliot Lake(Annapolis County) by trapnetting, gillnetting, andelectrofishing. Age and growth were estimated byscale reading. Smallmouth bass in Black River andGrand lakes exhibited growth similar to or slightlylower than other smallmouth bass in the northeast(Smith 1942; Stone et al. 1954; Turner andMacCrimmon 1970). Fish were 5-6 years old be-fore they reached 300 mm fork length (FL) andtook about 10 years to reach 400 mm. Smallmouth

684 McNEILL

TABLE 2.—Mean fork length (mm) at age for small-mouth bass from three lakes in Nova Scotia. Values inparentheses are sample sizes (left) and standard deviations(right).

Black River Lake Grand Lake Elliot LakeAge 44°56'N, 64°24'W 44°55'N, 63°36'W 44°56'N, 65°11'W

(years) 668 ha 1,841 ha 18 ha

2345678910111213

174(24.20)232 (23, 32)285(10,21)329 (6, 29)350 (2, 2)369 (9, 14)390(4,13)395(2, 11)411 (3, 11)421(1)469(1)

122(5,15)167(11, 15)203 (6, 8)257(16,29)300(16,20)322(17, 17)336(17,21)364 (5, 10)386(5,14)370 (2, 0)

427 (3, 24)

132(8,7)172(12,10)194 (8, 13)218(1)229 (4, 5)244 (3, 9)248 (3, 9)258(1)

bass in Elliot Lake, however, exhibited slowergrowth than any recorded in the literature; the old-est specimen collected was age 9 and only 258 mmFL (Table 2). A mark-recapture population esti-mate done in 1992 in Elliot Lake indicated thedensity of smallmouth bass (all ages) was 254/ha(SD = 94), which suggests that crowding probablycontributed to slow growth.

The reproductive biology of smallmouth bass inNova Scotia, including water temperature atspawning, duration of nest guarding, nest site se-lection, and spawning success, was similar to thatreported by Doan (1940), Turner and Mac-Crimmon (1970), Coble (1975), and Cantin andBromage (1991). In 1969, Bain (1993) collectedinformation from 18 smallmouth bass nests ob-served in Lumsden Pond (Kings County), NovaScotia. Nest building began in late May at a watertemperature of 16°C. Sixteen of the 18 nests ob-served were successful in producing swim-up fry,and nest guarding was terminated by the first weekof July. Seventeen nests were in close associationwith stumps or boulders. Average water depth atthe nest was 104 cm and the mean distance fromshore was 3.4 m. Nest substrate ranged from siltto gravel (Bain 1993).

In 1993 the NSDOF undertook a more detailedstudy of smallmouth bass reproduction in SecondLake (Halifax County). Nesting stage, nest den-sity, water depth, distance from shore, nest cover,and substrate were observed every 2-3 d alongfour shoreline transects from mid-May to mid-July. Eighty-six nests were observed along 1,610m of shoreline. Nesting stage was arbitrarily di-vided into four categories: excavated or newlyconstructed nests without eggs or fry; nests with

eggs present; nests with hatched fry in close as-sociation with the substrate; and nests in whichthe fry were dispersed in the water column, alongshore, or in nearby cover (Figure 3). Newly ex-cavated nests were first observed on June 9 andmany had eggs present, although these nests werenot observed 2 d earlier. Excavation of new nestscontinued through June 18, after which no newnests were observed. Egg deposition peakedaround the middle of June and continued until theend of June; hatching peaked around the thirdweek of June and was complete by early July. Frywere dispersed from the nest and parental guardingended by the first week of July.

The four shoreline transects observed repre-sented different habitats: (1) shallow (gradient<10%) with nesting cover consisting of logs,stumps, or boulders; (2) shallow with no cover;(3) deep (gradient ̂ 10%) with cover; and (4) shal-low, marshy areas (Table 3). Nest density washighest (10.8/100 m) in shallow areas with ade-quate cover and gravel or silt substrate, and lowest(1.7/100 m) in shallow marshy, silty areas wherethe substrate consisted of dense vegetation and de-tritus. These observations are consistent with thoseof Turner and MacCrimmon (1970) and Coble(1975). Mean water depth at the nest site and meandistance from shore were 130 cm and 18.7 m infavorable habitat and 56 cm and 3 m in poor hab-itat. Nesting success, defined as the proportion ofnests observed that produced swim-up fry, rangedfrom 38 to 68% and was not closely related tohabitat; however, the lowest success occurred inpoor habitat. Doan (1940) reported nesting successrates from 0 to 57% in Georgian Bay, Lake Huron,and Reynolds and O'Bara (1991) observed successrates of 13% and 35% over two years in Tennesseestreams.

Selection of a nest site close to some form ofcover has been documented by several studies(Doan 1940; Turner and MacCrimmon 1970; Co-ble 1975). In Second Lake, 76-100% of all nestsobserved in each of the four habitats were locatedclose to boulders, logs or stumps, and occasionallyhuman debris such as submerged boats or tires.Smallmouth bass usually nest over gravel or rockysubstrate, or, more rarely, woody debris or densevegetation (Turner and MacCrimmon 1970; Scottand Crossman 1973). In Second Lake, substrate inall four nesting habitats varied; smallmouth basspreferred a gravel-cobble mixture over mud or silt,followed by boulder or rock, and finally detritusor vegetation.

SMALLMOUTH BASS IN NOVA SCOTIA 685

3 Excavated E3Eggs Present HI Fry Present]I Fry Dispersed —Temperature J

100 -iPCD

10I I I I I I I

9 Jun 15 Jun 21 Jun 27 Jun 3 Jul12Jun 18 Jun 24 Jun 30 Jun 6 Jul

1993FIGURE 3.—Nesting stages by date and water temperature for smallmouth bass in Second Lake, Halifax County,

Nova Scotia, 1993.

DiscussionFor many years after the initial introductions,

smallmouth bass in Nova Scotia were largely un-known to most anglers except in a few localizedareas. Within the last 25 years, and particularlysince the early 1980s, the fishery has undergone aremarkable expansion, and the species is now con-sidered a prime game fish throughout much of the

TABLE 3.—Nesting characteristics for smallmouth bassfrom four habitats in Second Lake (44°47'N, 63°39'W),Nova Scotia, 1993.

Habitat type

Characteristic

Shoreline length (m)

NoCover, cover,shallow shallow

510 360

Cover, Marshy,deep shallow

270 470Nests

Number 55 17 6 8Density (nests/100 m) 10.8 4.7 2.2 1.7% successful 68 65 83 38Mean depth (cm) 130 86 59 56Mean distance to

shore (m) 18.7 5.8 2.7 3

province. The increase in average catch per anglerper year has closely mirrored total estimated catch-es, which suggests a real increase in stock abun-dance rather than elevated catch as a result of in-creased effort.

A number of factors have contributed to thisgrowth, the most notable being the rate at whichsmallmouth bass are colonizing new watersthrough natural migration or illegal transfers. Asa result of these movements, the most compellingconcern is the potential for interspecific impactsupon indigenous species. It is uncertain whetheror not smallmouth bass adversely affect salmonids.Indeed, demonstrating that smallmouth bass haveactually caused or contributed to the decline ofsalmonids is difficult (Fletcher 1991). Catt (1949)reported rapid diminution of brook trout and land-locked Atlantic salmon Salmo salar populations inChiputniticook and Magaguadavic lakes, NewBrunswick, following the introduction of small-mouth bass. Extensive sampling and many anglerinterviews in 1992 failed to record brook trout inElliot Lake, where they were reported to be plen-tiful before the arrival of smallmouth bass. On the

686 McNEILL

other hand, Fletcher (1991) reviewed 10 case his-tories from the northwestern United States andfound no indications of reduced salmonid survivalafter smallmouth bass introductions.

In Nova Scotia, it is against the law to transferany fish species between water bodies, and generalbait provisions prohibit the use of smallmouth bassas bait. Through articles in newspapers and thepopular literature and participation in fishing andoutdoor recreation exhibitions, the NSDOF has at-tempted to educate anglers regarding the potentialadverse effects of the indiscriminate stocking ofsmallmouth bass. A smallmouth bass introductionpolicy was developed in 1992 to address the con-cerns of anglers who must travel considerable dis-tances to reach smallmouth bass waters. Throughcareful selection of watersheds that are suitablefor smailmouth bass but marginal for salmonids,controlled introductions of smallmouth bass maycurtail further indiscriminate stocking. Converse-ly, such introductions may serve to exacerbate theproblem by creating a local stock for illegal trans-fers into neighboring waters.

A second factor in the growth of the smallmouthbass fishery is the dramatic increase in angler par-ticipation. The Canadian Association of Small-mouth Anglers (CASA), founded in Nova Scotiain 1989 by 16 smallmouth bass enthusiasts,claimed over 200 members in 1993. A tackle shopdedicated exclusively to smallmouth bass hasopened in response to the demand. Improved skillsmay also contribute to increased catches, as an-glers new to the sport tailor their techniques to-wards catching smallmouth bass.

The practice of catch and release cannot be over-looked with respect to the smallmouth bass fisheryin Nova Scotia. In 1990, there was a 10-fish limitand no length restriction for smallmouth bass. Ev-idence from angler diaries, license stub returns,and personal observations suggested that most ofthe releases were voluntary and not a result of thelimit. The proportion of smallmouth bass caughtand released in 1990 (96%) was the highest re-corded for any game fish in the sportfishing surveydata for Nova Scotia. The widespread opinion ofanglers that smallmouth bass are not as palatableas brook trout or other game fish may contributeto the release of many fish.

The promotional efforts of state conservationagencies, angling groups such as the Bass AnglersSportsman Society, outdoor writers, and syndicat-ed televised fishing programs have resulted inwidespread voluntary catch and release (Clark1982; Graff 1988). In Nova Scotia, the NSDOF

and CASA have promoted catch and release. Be-cause hooking mortality is relatively low for small-mouth bass (Weidlein 1987; Green et al. 1988),catch and release represents a valuable manage-ment option. Generally, anglers are compliant be-cause they are confident most of the fish they re-lease will survive. Because the relatively newsmallmouth bass fishery in Nova Scotia has de-veloped concurrently with the practice of catch andrelease and because there was no tradition of har-vesting fish for consumption as in the case of othergame fish species, harvest of smallmouth bass islow. However, a substantial trophy smallmouthbass fishery in Nova Scotia is unlikely because ofthe climate and low aquatic productivity.

AcknowledgmentsI thank R. Heighten for assistance in field col-

lections and laboratory analysis, the members ofthe Canadian Association of Smallmouth Anglers,and all anglers, biologists, and others who pro-vided information on tournaments and smallmouthbass distributions. D. MacLean, N. Adams, and R.Bancroft provided critical reviews of the manu-script and gave me considerable latitude in per-forming my other duties during its preparation.

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