10 from Yellowstone Lake. commercial fishermen sport anglersand trout were harvestedby numbers ofcutthroat 1900s, tremendous In theearlyandmid- Resources, Yellowstone NationalPark. biologist withtheCenterfor technicians, andMahonyisfisheries Doepke andErtelare fisheries Physiology, UniversityofWyoming. Department ofZoologyand Park anddoctoralstudent, for Resources, Yellowstone National Bigelow isfisheriesbiologist,Center [email protected] or307/344-2281. State University. Hecanbereached at Department ofEcology, Montana National Parkandaffiliate professor, Center forResources, Yellowstone Koel issupervisoryfisheriesbiologist, Daniel L.Mahony Brian D.Ertel Philip D.Doepke Patricia E.Bigelow Todd M.Koel introduced fishes

feature YELLOWSTONE NATIONAL PARK ARCHIVES PARK NATIONAL YELLOWSTONE ABSTRACT and impactstobearsanglers Yellowstone cutthroat trout decline Nonnative laketrout result in could beessentialtomaintaintheintegrityofGreaterYellowstone Ecosystem. bilis important consumerspecies,includingthethreatenedgrizzlybear( declined, indicatingthatnettingefforts maybeimpactingthepopulation.Asseveral were removed.Thecatchperuniteffort and averagelengthofspawninglaketrouthave using upto16kmofneteachday, May–October. From1994–2004,>100,000laketrout in 2004.To suppresslaketrout,theNationalParkServiceinitiatedagillnettingprogram anglers hasalsodeclined,fromtwofishcaughtperhourin1998tolessthanonefish/h reductions, indicatingcascadinginteractionsinthefoodwebofthissystem.Successby has declinedatspawningstreams1989–2004,andmirroredthatofthecutthroattrout trout atClearandBridgecreekshavedeclined>90%inthepast5years.Activitybybears whirling disease),anddroughtconditions.Numbersofupstreammigratingcutthroat lake trout( Yellowstone Lakeanditstributarystreamshavebeenaffected bytheintroductionof During thepastdecade,Yellowstone cutthroattrout( ), usethispopulationasanenergysource,preservationofremainingcutthroattrout ing forwaystopropagateanddistribute the U.S.FishCommissionbeganlook- Yellowstone Riveringreatnumbers,and the lake,itstributarystreams,and American Angler. suchas periodicals national andlocalnewspapers,aswell the fisherywaswidelypublicizedin as theworld'sfirstnationalparkin1872, lishment ofYellowstone NationalPark (Doane 1871).Shortlyaftertheestab- lake areafortheirbeautyandabundance they werenotedbyearlyexplorersofthe 1995), andhistoricalsignificance, 1995), economic(GresswellandLiss great ecological(SchulleryandVarley of Yellowstone Lake.Thesefishhave Introduction ( adfluvial Yellowstone cutthroattrout trout populationintheworldis Salvelinus naymaycush Oncorhynchus clarkibouvieri The largestinlandcutthroat Anglers beganvisiting Forest andStream ) population ), invasionby and tributaries haveremainedgeneticallypureduetoisolation 2004). ThecutthroattroutofYellowstone Lakeandits cutthroat troutsubspecies(Behnke2002;Koeletal. sion withnonnativerainbowtrout( trout, populationshavebeencompromisedbyintrogres- elsewhere inthenaturalrangeofYellowstone cutthroat 1988). angling regulations(Varley 1983;GresswellandVarley operations andimplementationofincreasinglyrestrictive the mid-1900sresultedinclosureofegg-taking Evidence ofacutthroattroutpopulationdeclineduring after thecreationofNationalParkService(NPS). forvisitorsuntil1919,just cially fishedtoprovidefood a greatamountofanglingpressure,andwerecommer- Gresswell 1988).Thecutthroattroutalsoweresubjectto Yellowstone NationalPark(Varley 1981;Varley and removed foruseinotherwaters,mostlyoutside 1900 to1956,over818millioncutthroattrouteggswere facility onthenorthshoreofYellowstone Lake.From was thedevelopmentofafederally-operatedfishculture North America(Varley andSchullery1998).Theresult the cutthroattroutofYellowstone Laketolocationsacross Of the124tributaries flowingintothelake,68 have largest lakeabove2,000melevation inNorthAmerica. throat trout.At34,000ha,Yellowstone Lakeisthe variety ofenvironmentalconditions forthenativecut- the IntermountainWest, 1998–2004(Cooketal.2004). al., inpress),andadroughtthathas persistedthroughout Myxobolus cerebralis namaycush; stressors, includingnonnativelaketrout( tion hasrecentlybeenexposedtothreeotherpotential makes themextremelyvaluable.However, thepopula- near Canyon(Figure1).Thegeneticpurityofthesefish River located25kmdownstreamfromthelakeoutlet provided bytheLowerandUpperfallsofYellowstone In streamsthroughoutYellowstone NationalParkand Yellowstone Lake and itsdrainagesprovideagreat Myxobolus cerebralis Fisheries |www.fisheries.org |vol30no11 Kaeding etal.1996),theexoticparasite Oncorhynchus clarkibouvieri (the causeofwhirlingdisease;Koelet Ursus arctoshorri- O. mykiss (the causeof ) orother Salvelinus ) of PAT BIGELOW introduced fishes 11 Myxobolus Yellowstone feature ). whirling disease Figure 1. Figure Lake and tributary drainages within National Park, Yellowstone with locations of cutthroat spawning migration trout fall trout traps, cutthroat netting assessment sites, spawning trout cutthroat visual survey streams, and sites sentinel fry exposure (blue = negative and = positive for the red of presence cerebralis An aggressive gillnetting An aggressive targets spawning program on Yellowstone lake trout Lake during September each year. ), bald ), and many other avian ), and many Ursus arctos horribilis Ursus arctos halus p River Clear Creek Clear Pelican Creek Yellowstone (to lake inlet) (to lake Haleaeetus leucoce Haleaeetus Nonnative lake trout would not be a suitable ecologi- Nonnative lake trout would east east ArmArm Arm SouthSouth South and terrestrial species use cutthroat trout as an energy species use cutthroat and terrestrial 1986; Gunther 1995; Schullery source (Swenson et al. 1995). and Varley Lake trout in the Yellowstone cal substitute for cutthroat inaccessible to most consumer system because they are to occupy greater depths within species. Lake trout tend trout. Lake trout remain the lake than do cutthroat Lake at all life stages and they do not within Yellowstone streams, as do cutthroat trout. typically enter tributary systems, suggests that intro- similar Evidence from other, result in the decline of cutthroat duced lake trout will 1974; trout (Cordone and Franz 1966; Dean and Varley Behnke 1992). Bioenergetics modeling suggests that an Lake will average-sized mature lake trout in Yellowstone eagle ( tion and recovery of the cutthroat trout. This subspecies trout. of the cutthroat recovery tion and of the Greater for maintaining the integrity is important natu- the most intact Ecosystem, arguably Yellowstone in the lower 48 ecosystem remaining rally-functioning Grizzly bear ( United States. M. ArmArm SouthSouth Main Basin Bridge ), lake Fishing and possi- Richardsonius Lake Rapids Behnke 2002). Lower Falls LeHardy Upper Falls BreezeBreeze Wyoming ChannelChannel M. cerebralis, West River Thumb Catostomus catostomus CreekCreek ), redside shiners ( SolutionSolution Yellowstone Canyon Yellowstone N. P. (from lake outlet) Rhinichthys cataractae; Idaho Whirling disease exposure site SpawningSpawningSpawning migration migration migrationtraptraptrap FallFallFall netting netting netting assessmentassessmentassessment site site site SpawningSpawningSpawning visual visual visualsurveysurveysurvey streamstreamstream WhirlingWhirling disease disease exposure exposure sitesite Grant Island Montana BasinBasin , the park is placing a high priority on preserva- Carrington Bridge Creek GeyserGeyser ), and lake trout are also present in the lake sys- ), and lake trout are also Couesius plumbeus Contemporary research points to nonnative species as Contemporary research cerebralis the greatest threat to cutthroat trout of the Intermountain the greatest threat to cutthroat et al. (Gresswell 1995; Kruse et al. 2000; Dunham West the recent invasions by lake trout and 2004). With tem due to introductions. chubs ( balteatus November 2005 | www.fisheries.org | Fisheries November 2005 | www.fisheries.org been used by spawning cutthroat trout (Jones et al. 1987; et al. trout (Jones cutthroat by spawning been used features occur et al. 1997). Geothermal Gresswell et al. Lake (Morgan much of Yellowstone throughout these many it is the presence of 2003), but unfortunately, that may increase the variable environments unique and by nonnative and of this system to invasion vulnerability such as lake trout, exotic species bly others in the future. Prior to Euroamerican bly others in the Lake cutthroat trout manipulation, the Yellowstone 10,000 years (since glacial existed for approximately with only one other fish species, recession) in sympatry the longnose dace ( suckers ( longnose Now, COLLIN CHRISTIANSEN COLLIN 12 trout ormore eachyear. consume 41cutthroat has thepotentialto trout inYellowstone Lake suggests thateachlake Bioenergetics research introduced fishes feature Annually, approximately4,000 anglers(5%ofallanglers) and speciessizesoffishcaught byanglersisobtained. al. 1980).Informationonthewaters fished,timespent, upon purchasingaspecialusepermit forfishing(Joneset in anupstreamdirectiononceeachweekfromMay each tributary, andthestandardizedreachesarewalked Figure 1).Spawningreacheswereinitiallydelineatedon Lake andGrant(Reinhart1990;Reinhartetal.1995; along thewesternsideofYellowstone Lakebetween ducted annuallysince1989on9–11tributarieslocated for cutthroattroutandbearactivityhavebeencon- throughout Yellowstone Lake(Jonesetal.1977;Figure1). line overnight,inshallowwater(0–5mdepth),at11sites are placedinsetsoffivenetsperpendiculartotheshore- 7.6 mgraduatedmeshpanelsof19,25,32,38,and51mm) Multi-mesh-size (experimental)gillnets(38.0mlength, ducted duringSeptemberofeachyearsince1969. and lengthstructurehavebeenassessedbynettingcon- Yellowstone Lake,cutthroattroutpopulationabundance Vancouver, 550BB). BritishColumbia,Model counters havebeenused(PulsarElectronics,LTD., North trapped andmanuallyenumerated.Sincethen,electronic Yellowstone Laketospawn.Until1998,allfishwere counted atpermanentweirsastheymoveupstreamfrom 1). FromMay–Julyeachyear, thecutthroattroutare Bridge Creeksince1945and1999,respectively(Figure counting upstream-migratingadultsatClearCreekand monitored thecutthroattroutspawningpopulationby population growth. and (3)describeresultsofeffortstosuppresslaketrout and size,(2)examinetheimpactsonbearsanglers, are to(1)describetrendsincutthroattroutabundance Yellowstone Lakeecosystem.Specifically, ourobjectives rent statusofNPSeffortstopreservetheintegrity Ruzycki 1997;Bigelowetal.2003). of thecutthroattroutinthissystem(Mahonyand long-term laketroutremovalprogramfortheprotection trout withinYellowstone Lake.Theeffortshaveledtoa determine thespatialandtemporaldistributionoflake panel (McIntyre1995),theNPShasusedgillnettingto Following theguidanceofalaketroutexpertadvisory consume 41cutthroattroutperyear(Ruzyckietal.2003). Methods Cutthroat troutspawningvisualsurveys. Cutthroat troutfallnettingassessment. The overallpurposeofthispaperistodescribethecur- Cutthroat troutspawningmigrationtraps. distributed toallanglers assessed viaareportcard effort andsuccesshasbeen mation. bear sightings. trout, freshtracks,and/or scat, partsofconsumed by notingthepresenceof grizzly bearsisestimated bears ( weekly activitybyblack are counted,andthe observed cutthroattrout through July. The Angler reportcardinfor- U. americanus Since 1979,angler Visual surveys The NPShas Within ) and aquaria foranadditional90daysat10–13 July–September. Followingtheexposures,frywereheldin days posthatch)wereexposedfora10-dayperiod, galvanized wiremesh),60–80cutthroattroutfry(25–50 cage (1mheight0.5diameterandconstructedof5mm fry exposures,1998–2003(Koeletal.,inpress).Ineach outlet) wasdeterminedbyuseofcutthroattroutsentinel tributaries andtheYellowstone River(lakeinletandlake of chain reaction(PCR)techniquetoconfirmthepresence (Andree etal.2002)followedbythenestedpolymerase toexamineforthepresenceofmyxospores method occurred initiallybythepepsintrypsindigest(PTD) removal program,1998–2003.Screeningofthesefish talities fromthefallnettingassessmentandlaketrout by examinationofjuvenileandadultcutthroattroutmor- of night. and killdepositedeggsinshallowwaters(<5mdepth)at trofishing wasalsousedtoremovespawninglaketrout any cutthroattroutbycatch.In2004,boat-mountedelec- short duration(typicallyoneday)toreducemortalityof depth), large-meshgillnet(51–70mmmeshsize)setsof (Figure 1),weregillnettedusingshallow-set(0–20m Carrington Island,GeyserBasin,andSolutionCreek locations targetedinthefall,includingBreezeChannel, early October, whenspawningistypicallycompleted.The each year. Theremovalprogramtargetsthesefishuntil gregating nearknownspawninglocationsinlateAugust were inplacefishingforlaketrout. season (lateMay-lateOctober),upto16kmofgillnet length each),typicallyfor>7days.Duringtheopenwater length, setingangsofsixcontiguousnets(1800mtotal water typically40–65mdeep.Gillnetswere300 mm meshsize)gillnetswereplacedonthelakebottomin a lesserextent,thesouthernarms.Small-mesh(19–44 have expandedoutwardtoincludethemainbasinand, years oftheprogram,however, laketroutandgillnetting have beenfoundtobehighest(Figure1).Throughthe trated intheWest Thumb,wherelaketroutdensities 1994. Spatially, thegillnetlocationshavebeenconcen- to suppresslaketroutinYellowstone Lakeeachyearsince park's fisheriesprogram. have voluntarilycompletedandreturnedcardstothe each fryexamined(Baldwinetal.2000). scale of0(noinfection)to5(mostseverefor determine severityofinfection.Severitywasrankedona conducted onfryfrom presence of technique (Andreeetal.1998)wasusedtotestforthe asite developmentpriortobeingsacrificed.ThePCR Abundance Trends inCutthroat Trout at ClearCreek. Only 3,161cutthroattroutascended trout wereobservedincountsofupstream-migrating fish angler harvestregulationsforYellowstone Lakecutthroat M. cerebralis M. cerebralis The prevalenceandseverityof Whirling diseaseprevalenceandseverity. The maturelaketroutofYellowstone Lakebegincon- Lake troutremovalprogram. Impacts ofhistoricalegg-takingoperations andliberal Fisheries |www.fisheries.org |vol30no11 M. cerebralis (Andree etal.1998). within Yellowstone Lakewasdetermined M. cerebralis . Histologicalexaminationwas Gillnetting hasbeenused M. cerebralis -positive exposuresto o The prevalence C toallowpar- within 15 Clear Creek in 1954, just two years prior to and has decreased by over 99% since counts the cessation of fish culture operations on began in 1999 (when 2,363 cutthroat trout Figure 3. Length-frequency distributions of cutthroat Yellowstone Lake (Varley and Schullery ascended the stream to spawn). The decline trout collected during the fall netting assessment on 1998; Figure 2). With angling restrictions, was also evident in results of the fall netting Yellowstone Lake with total number of trout <325 mm (n ) and >325 mm (n ), 1997–2004. the number rebounded during the 1960s and assessment, where an average of 15.9 cut- 1 2 1970s to 70,105 cutthroat trout in 1978 throat trout per net were caught in 1994, and (Jones et al. 1979). Although there was vari- only 6.1 per net were caught in 2002 (Figure ation among years, the increasing trend in 2). Prior to 2003, the reduction in catch by 60 50 1997 cutthroat trout abundance within the fall netting program averaged 11% per n = 245 n = 363 40 1 2 Yellowstone Lake was also indicated by the year since 1994, the year lake trout were first 30 discovered in Yellowstone Lake. During fall netting assessment. An average of 10.0 20 fish per net were caught by this assessment in 2003–2004, however, the fall netting assess- 10 1969, and 19.1 fish per net were caught in ment provided some of the first indications 0 1984 (Figure 2). that the cutthroat trout population may be

60 100 150 200 250 300 350 400 450 500 550 Since the late 1980s, however, there has responding positively to efforts to remove 50 1998 n = 176 n = 369 been a significant decline in the Yellowstone nonnative lake trout. An average of 7.4 fish 40 1 2 Lake cutthroat trout population. The num- per net were caught in 2003, and 7.9 fish per 30 ber of upstream-migrating cutthroat trout net were caught in 2004. 20 counted at Clear Creek was 1,438 during 10 0 2004 (Figure 2). This count was down from Trends in Cutthroat Trout

3,432 in 2003, and 6,613 in 2002, and was 60 100 150 200 250 300 350 400 450 500 550 1999 Length 50 the lowest count since 1954. The fish count- n = 131 n = 315 ing station operated on Bridge Creek, a small 40 1 2 northwestern tributary, indicated that only a Length-frequency data from the fall net- 30 single fish migrated upstream during 2004 ting program from 1997 to 2004 indicated an 20 (Figure 2). The number of spawning cut- increase in length (>325 mm) and reduction 10 0 throat trout in recent years has declined by in numbers of adult cutthroat trout in more than 50% annually in Bridge Creek, Yellowstone Lake (Figure 3). In 2004, fewer 60 100 150 200 250 300 350 400 450 500 550 50 2000 n = 79 n = 317 40 1 2 Figure 2. Number of upstream-migrating cutthroat trout counted at Clear Creek (1945–2004) 30 20 and Bridge Creek (1999–2004) spawning migration traps (A), and mean number of cutthroat 10 trout collected per net during the fall netting assessment on Yellowstone Lake, 1969–2004 (B). 0 Note differences in scale between Clear Creek and Bridge Creek data.

60 100 150 200 250 300 350 400 450 500 550 50 2001

Migrating Cutthroat Trout (x 10 n = 133 n = 230 ) 1 2

3 40 80 25 30

Clear Creek Bridge Creek Creek Upstream Bridge 70 20 20 10 60 A 0 Number of Cutthroat Trout of Cutthroat Number

60 100 150 200 250 300 350 400 450 500 550 50 15 50 2002 n = 178 n = 158 40 40 1 2 30 10 30 20 20 10 5 0 Clear Creek Upstream 10

60 100 150 200 250 300 350 400 450 500 550 2003 0 0 2 50 ) Migrating Cutthroat Trout (x 10 (x Trout Cutthroat Migrating n = 207 n = 200 40 1 2 30 20 20 Fall netting assessment 10 0 15

B 60 100 150 200 250 300 350 400 450 500 550 50 2004 n1 = 239 n2 = 167 10 40 30 20 5 10 0 Mean Cutthroat Trout Per Net Per Trout Mean Cutthroat 0 100 150 200 250 300 350 400 450 500 550 1945 1955 1965 1975 1985 1995 2005 Total Length (mm)

November 2005 | www.fisheries.org | Fisheries 13 14 length ofcutthroat trout caughtbyanglers from Yellowstone Lake,1979–2004. Figure 5. sideofYellowstonethe western LakebetweenandGrant,1989–2004. bears observedduringweeklyspawningvisualsurveysof9–11tributarieslocatedalong Figure 4. introduced fishes Mean Trout Per Hour Mean Cutthroat Trout Per Visit 10 20 30 40 50 60 70 80 0.0 0.5 1.0 1.5 2.0 2.5 Angler-reported catchrateforcutthroat trout andlaketrout, andtheaverage Mean numberofcutthroat trout andmeanactivitybyblackbearsgrizzly Cutthroat trout catch trout Cutthroat 0 9819 9620 2004 2000 1996 1992 1988 9818 9619 9419 2002 1998 1994 1990 1986 1982 1978 feature M. cerebralis due tothelownumbersoflaketroutandincidence (Figure 1),whichmayactasrefugesforcutthroattrout collected inthesouthernarmsofYellowstone Lake years (2002–2004).Manyofthesejuvenileshavebeen cutthroat trout(100–325mm)hasbeennotedinrecent Despite this,anapparentincreaseinnumbersofjuvenile Clear Creekwereinthissizerange(Jonesetal.1993). cutthroat troutsampledinspawningtributariessuchas lected comparedtoearlieryears.Historically, most fish betweenthelengthsof325and425mmwerecol- Consumers Influence onCutthroat Trout during spawningvisualsurveysof9–11tributaries average numberofcutthroattroutobservedeachweek bears inthelakearea(ReinhartandMattson1990).The significant sourceofenergyforblackbearsandgrizzly Yellowstone Laketributarieshistoricallyhaveservedasa Upstream-migrating cutthroattroutwithin Lake trout catch trout Lake in theseareas(seebelow;Koeletal.2004). Trout Cutthroat trout length trout Cutthroat Bear

350 375 400 425 450 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9

Mean Length (mm) Length Mean Mean Bear Activity Per Visit Per Activity Bear Mean tant food sources,thestatusofgrizzlybearpopulation tant food Mattson andMerrill2002).Despitethisthreattoimpor- ened bythedeclineofwhitebarkpine( on bearpopulations,asbearsofthisregionarealsothreat- what theoveralleffectofcutthroattroutdeclineswillbe previously reliedoncutthroattrout. whenthey the regionduringspring-spawningperiod Bears mayhavebeenforcedtouseotherenergysourcesin species intothispristineecosystem(Reinhartetal.2001). ofnonnativeandexotic resulting fromtheintroduction a trophic-levelcascade(Spenceretal.1991),possibly activities bybears50timesin1991.Theseresultssuggest visual surveysofspawningcutthroattroutdocumented dent atotalof8timesduring2004.Bycomparison,the bears usedthesetributaries,asbearactivitywasonlyevi- black bearsandgrizzlybears.Itwasapparentthatfew A similartrendwasobservedinuseofthesestreamsby of8weeksin2004. the 9streamssurveyedoveraperiod only 35cutthroattroutwereseenonspawningreachesof have beenobservedinrecentyears(Figure4).Infact, (1989–2004) hasdeclinedtothepointwherefewtrout Decade afterDiscovery Lake Trout Removal Program a slow thedispersalof drainage wascompletelyclosedin2004anattemptto released unharmed.AnglingonthePelicanCreek have requiredthatcutthroattroutbeimmediately from 2.5in1979to0.32003.Since2001,regulations number ofcutthroattroutcaughtperhourhasdeclined cerebralis Pelican Creek(Figure1),thelaketributarywhere trout caughtbyanglershasbeenevenmoresignificanton 448 mmin2004.Thedeclinenumbersofcutthroat Yellowstone Lakehasincreasedfrom370mmin1995to average lengthofcutthroattroutcaughtbyanglerson increased from0.0in1998to0.12004.Inaddition,the 2004 (Figure5),whilelaketroutcaughtperhour throat troutcaughtperhourfrom2.0in1998to0.8 Lake haveexperiencedadeclineinthenumberofcut- angler reportcardssuggestthatanglersonYellowstone Lake. Inrecentyears,estimatesderivedfromreturnsof than one-thirdofvisitinganglersfishedYellowstone tion foranglersfromaroundtheworld.Historically, more decades (Schwartzetal.2002). bears haveexpandedtheirrangeby48%overthelasttwo in 2004(HaroldsonandFreypress).Inaddition,grizzly it waslistedasathreatenedspecies,toatleast431bears has increasedfromanestimated136bearsin1975when ered tobestableincreasing.Thegrizzlybearpopulation in theGreaterYellowstone Ecosystemiscurrentlyconsid- age of5.5laketrout perunitofeffortwerecaught. In night). Catchratehasdeclinedsince 1998,whenanaver- effort units(oneunit=100 mofnetsetoverone 26,634 laketroutwereremoved in 2004using15,781 years toanaverageof10timesthat of1999.Atotal (Figure 6).Thegillnettingefforthas increasedinrecent program hasremoved>100,000 lake troutsince1994 nonnative specieshaveintensified.TheNPSgillnetting in 1994(Kaedingetal.1996),effortstocounteractthis Aside fromlocalizeddisplacementitisnotknown Yellowstone NationalParkhaslongbeenadestina- Since thediscoveryoflaketroutinYellowstone Lake Fisheries |www.fisheries.org |vol30no11 infection hasbeenmostsevere(seebelow).The M. cerebralis to otherparkwaters. Pinus albicaulis; M. 15 introduced fishes Lake Trout CPUE Spawning Lake Trout (#/100 m net/night) Mean Length (mm) 640 600 560 520 480 feature 6 5 4 3 2 1 0 Gillnet Unit Effort Spawning Lake Trout Length CPUE infection. in three important with an average severity (by with an average severity M. cerebralis M. cerebralis M. cerebralis Spawning Lake Trout Number Lake Trout Number A B 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 M. cerebralis, 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 have been most severe in Pelican Creek, have been most severe

5 0 8 6 4 2 0

Number of lake trout removed, gillnet units of effort (1 unit = 100 m of gillnet units of effort removed, Number of lake trout

30 25 20 15 10

Number (x10 Number ) Effort Unit (100 m/night) (x10 m/night) (100 Unit Effort )

3 3

Spawning Lake Trout Lake Spawning Consistent, annual counts of upstream-migrating Consistent, annual counts Gillnet or Number Trout Lake By 2001, cutthroat trout sentinel fry exposures con- trout sentinel By 2001, cutthroat histological examination) of >4 on a scale of 0 (no infec- histological examination) infection; Koel et al. 2004). tion) to 5 (most severe have not been made in Pelican adult cutthroat trout a historic Creek in recent years, but records exist from fish weir that was previously used to enumerate spawning near from 1964 through 1981 (Jones et al. 1982). Netting tributary the location of the historic weir (near the indi- mouth) for upstream-migrating adults in 2002–2004 spawning streams: Pelican Creek, Clear Creek, and the Pelican Creek, spawning streams: near from the lake outlet River downstream Yellowstone in press). Since then, (Figure 1; Koel et al. Fishing Bridge River upstream of Yellowstone sentinel exposures in the other spawning tributaries have the lake inlet and 13 the parasite. The impacts of failed to detect the presence M. cerebralis fry have been observed in recent where few wild-reared trout sentinel fry exposures years (2001–2004). Cutthroat that >90% of the fry were in this tributary have indicated infected with firmed the presence of firmed the adult fish each year suggests that at least some cutthroat that at least suggests fish each year adult initial trout are surviving Figure 6. Figure obtained by the lake trout catch per unit of effort net/night) used, and lake trout gillnetting season, 1994–2004 Lake during the entire on Yellowstone program removal near spawning locations removed lake trout (A). Number and mean length of mature on Yellowstone Carrington Island, Geyser Basin, and Solution Creek) Channel, (Breeze 1996–2004 (B). Lake during late August–early October, spores in M. cerebralis was discovered in Yellowstone Myxobolus cerebralis Avoiding bycatch of cutthroat trout has been a trout has been bycatch of cutthroat Avoiding population has grown and As the lake trout Thumb remain high Lake trout densities in the West November 2005 | www.fisheries.org | Fisheries November 2005 | www.fisheries.org Whirling Disease and Drought as Whirling Disease and Drought Additional Threats 2004, catch per unit effort (CPUE) for lake trout for lake (CPUE) unit effort catch per 2004, higher than that of (1.69) but was slightly remained low 2001–2003. program. Initially, the lake trout removal challenge for trout for every was very high: 6.6 cutthroat the bycatch high bycatch was in netted in 1995. This lake trout densities of cutthroat trout and part due to the higher trout that occurred within lower densities of lake during the mid-1990s. Gillnetting Lake Yellowstone to reduce bycatch while protocols have been improved lake trout. The gillnets are now maximizing removal of deep (40–65 m depth), except consistently set very periods.during lake trout spawning The bycatch has or less for each lake trout net- been 0.1 cutthroat trout ted since 1998. spawning fish have become a expanded in recent years, program. In 2003, an addi- focal point for the removal near tional lake trout spawning location was identified area, Thumb Geyser Basin (Figure 1). This the West Solution along with areas near Carrington Island, since Creek, and Breeze Channel, has been gillnetted caught 1996. The total number of spawning lake trout in 2004 by gillnetting was 2,371 in 2003 and 7,283 lake trout (Figure 6). An additional 1,063 spawning average were removed by electrofishing in 2004. The spawning length of spawning lake trout removed near recent areas has decreased each year (Figure 6). The lake trout decline in the annual lakewide catch rate of of sex- and the annual reduction in the average length that the ually mature fish are positive indications on removal program is exerting measureable mortality this population. and a serious threat to the cutthroat trout. Model simula- cutthroat tions suggested a 60% or greater decline in the trout popu- trout population within 100 years if the lake (Stapp and lation was permitted to grow uncontrolled sug- Hayward 2002). Cutthroat trout abundance indices has gest that a decline of that magnitude (or greater) investigate already occurred. The NPS will continue to new methods to target the lake trout population. In par- using hydroacoustics, underwater cameras, and ticular, NPS is currently delin- high resolution (1 m) bathymetry, eating and characterizing known lake trout spawning Thumb), to predict where areas (all presently in the West new spawning areas may be pioneered in the lake basin. These potential spawning areas will be closely monitored and targeted for lake trout removal if fish begin to use them in the future. Lake in 1998 among juvenile and adult cutthroat trout (Koel et al. in press). Examination of gillnetting mortali- ties has since confirmed the presence of the parasite Lake, with highest prevalence throughout Yellowstone existing in the northern region of the lake, near known infected streams (see below; Koel et al. 2004). Although the widespread presence of this harmful parasite in the lake is disturbing, the discovery of cated that the spawning cutthroat trout population of Pelican Creek, otolith microchemistry, as well as comparative DNA analyses, has pro- which in 1981 was nearly 30,000 fish, has been essentially lost. With a vided evidence that the lake trout origin was Lewis Lake (Stott 2004; drainage area of 17,565 ha, Pelican Creek is the second largest tributary Munro et al. 2005), a lake within the park that was intentionally stocked to Yellowstone Lake in terms of discharge. with lake trout from Lake Michigan in 1890 (Varley 1981). To date, it Drought in the Intermountain West since 1998 may have impacted remains unknown exactly how the lake trout were introduced to cutthroat trout populations due to increased water temperatures and a Yellowstone Lake. reduction in peak stream flows (U.S. Geological Survey unpublished At present, a mandatory kill regulation is in place for all lake trout data for gage 06186500 Yellowstone River at Yellowstone Lake Outlet). caught on Yellowstone Lake, and the NPS asks anglers each year to assist In many cases, flows in tributary streams have become sub-terminal with the lake trout removal effort in this way. The Yellowstone Lake sit- near the lake, flowing through large sand and gravel bars. This discon- uation represents a unique case in which anglers are solicited to fish for nect of tributary streams from the lake has been occurring during lake trout without the desire to preserve the fishery. In NPS requests for mid-summer and fall, when cutthroat trout fry would typically be out- angler support, it is made clear that the goal is removal of as many lake migrating to Yellowstone Lake. Biologists have consistently noted trout as possible and suppression of the population for the purpose of cut- cutthroat trout fry that are stranded in isolated side channels and pools throat trout conservation. To this point a constituency has not in seasonally-disconnected tributaries. Although cutthroat trout have developed requesting the enhancement of the lake trout fishery in existed in the Yellowstone Lake ecosystem since glacial recession Yellowstone Lake. (Behnke 2002), and evolved in the face of great variation in thermal Since lake trout in Yellowstone Lake are known to prey on the native and other environmental regimes, the current drought is occurring dur- cutthroat trout (Ruzycki et al. 2003), the removal of >100,000 lake trout ing a period when the cutthroat trout are also impacted by lake trout has reduced predation on this important population. The lake trout predation and M. cerebralis. removal program on Yellowstone Lake represents a test case for the devel- opment of similar programs to preserve native salmonids in the Conclusions Intermountain West. For example, lake trout removal is currently being experimentally conducted on Lake Pend Oreille in northern Idaho, and Our results identify long-term impacts of lake trout and M. cerebralis is being considered for Lake McDonald of Glacier National Park and on cutthroat trout in Yellowstone Lake and the Greater Yellowstone Swan Lake of northwestern Montana. Ecosystem. Even with the Yellowstone National Park fisheries program The cumulative effects of lake trout and M. cerebralis have put dedicated to the preservation and recovery of the Yellowstone Lake cut- stress on the Yellowstone Lake cutthroat trout population during a throat trout population, it appears to be in peril. In addition, two period of intense drought in the Intermountain West. The cutthroat important cutthroat trout consumers, the black bear and grizzly bear trout population size of this system was once considered to be in the (icons for Yellowstone National Park and highly sought by millions of vis- millions: however, current abundance indices suggest that only a itors each year), are using cutthroat trout spawning streams much less fraction of that population exists today. The prospects of lake trout frequently. Yellowstone National Park anglers, a third of which fish control and rehabilitating historical cutthroat trout abundance are Yellowstone Lake, also have experienced a significant reduction in catch. yet to be achieved. Relatively low CPUE and an annual decrease in Of great interest to park managers is the timing and original source of the size of sexually mature fish are indicators that the removal pro- lake trout that were illegally introduced to Yellowstone Lake. Research gram is exerting pressure on this lake trout population. A continued on the microchemistry (Sr:Ca ratios) of otoliths has suggested that a lake focus on lake trout removal will be required into the future if cut- trout introduction likely occurred in the late 1980s (Munro et al. 2005). throat trout are to persist in Yellowstone Lake at a level allowing the These results suggest that lake trout existed in Yellowstone Lake for at overall integrity of the Greater Yellowstone Ecosystem to be main- least five years prior to being reported to the NPS by an angler. The tained. TODD KOEL

The National Park Service Freedom on Yellowstone Lake.

16 Fisheries | www.fisheries.org | vol 30 no 11 References

Andree, K. B., E. MacConnell, and R. P. Hedrick. 1998. A nested polymerase chain reaction for the detection of genomic DNA of Myxobolus cerebralis in rainbow trout Oncorhynchus mykiss. Diseases of Aquatic Organisms 34:145-154. Andree, K. B., R. P. Hedrick, and E. MacConnell. 2002. A review of the approaches to detect Myxobolus cerebralis, the cause of salmonid whirling disease. Pages 197-211 in J. L. Bartholomew and J. C. Wilson, eds. Whirling disease: reviews and current topics. American Fisheries Society Symposium 29, Bethesda, Maryland. Baldwin, T. J., E. R. Vincent, R. M. Silflow, and D. Stanek. 2000. Myxobolus cerebralis infection in rainbow trout (Oncorhynchus mykiss) and brown trout (Salmo trutta) exposed under natural stream conditions. Journal of Veterinary Diagnostic Investigation 12:312-321. Behnke, R. J. 1992. Native trout of western North America.histicated American Fisheries Society Monograph 6, Bethesda, Maryland. vers to _____. 2002. Trout and salmon of North America. The Free Press, New York. sults. Bigelow, P. E., T. M. Koel, D. Mahony, B. Ertel, B. Rowdon, and S. T. Olliff. 2003. Protection of native Yellowstone cutthroat ts have relied trout in Yellowstone Lake, Yellowstone National Park, Wyoming. Technical report NPS/NRWRD/NRTR-2003/314. National Park nnovations in Service, Water Resources Division, Fort Collins, Colorado. acoustic trans- Cook E. R., C. Woodhouse, C. M. Eakin, D. M. Meko, and D. W. nized the way Stahle. 2004. Long-term aridity changes in the western United arch. Micro to States. Science 306:1015-1018. e digital iden- Cordone, A. J., and T. C. Franz. 1966. The Lake Tahoe sport fishery. California Fish and Game 52:240-274. keep track of Dean, J., and J. D. Varley. 1974. Fishery management investigations, hic levels. Yellowstone National Park. Bureau of Sport Fisheries and Wildlife, technical report for 1973. Yellowstone National Park, Wyoming. Doane, G. C. 1871. Senate letter from the Secretary of War communicating the Yellowstone Expedition of 1870. 41st Congress, 3rd session, executive document 51. U. S. Government Printing Office, Washington, D.C. Dunham, J. B., D. S. Pilliod, and M. K. Young. 2004. Assessing the consequences of nonnative trout in headwater ecosystems in western North America. Fisheries 29(6):18-26. Gresswell, R. E. 1995. Yellowstone cutthroat trout. Pages 36-54 in M. K. Young, technical ed. Conservation assessment for inland cutthroat trout. U.S. Forest Service General Technical Report RM-GTR-256. Gresswell, R. E., and W. J. Liss. 1995. Values associated with management of Yellowstone cutthroat trout in Yellowstone National Park. Conservation Biology 9:159-165. Gresswell, R. E., W. J. Liss, G. L. Larson, and P. J. Bartlein. 1997. Influence of basin-scale physical variables on life history migration) characteristics of cutthroat trout in Yellowstone Lake. North ysis American Journal of Fisheries Management 17:1046-1064. Gresswell, R. E., and J. D. Varley. 1988. Effects of a century of human influence on the cutthroat trout of Yellowstone Lake. American Fisheries Society Symposium 4:45-52. Gunther, K. 1995. Grizzly bears and cutthroat trout: potential ems Inc.) impacts of the introduction of nonnative lake trout to -4000 Yellowstone Lake. Information paper BMO-8. Bear Management Office, Yellowstone Center for Resources, Yellowstone National vemco.com Park, Wyoming. Haroldson, M. A., and K. Frey. In press. Grizzly bear mortalities. In C. C. Schwartz and M. A. Haroldson, eds. Yellowstone grizzly bear investigations: annual report of the Interagency Grizzly Bear Waves in Study Team, 2004. U. S. Geological Survey, Bozeman, Montana. c Telemetry Jones, R. D., J. D. Varley, D. E. Jennings, S. M. Rubrecht, and R. E. Gresswell. 1977. Fishery and aquatic management program in Yellowstone National Park. U. S. Fish and Wildlife Service, technical report 1976, Yellowstone National Park, Wyoming.

November 2005 | www.fisheries.org | Fisheries 17 Jones, R. D., R. E. Gresswell, D. E. report 1986, Yellowstone National Park, Kruse, C. G., W. A. Hubert, and F. J. Jennings, D. C. Lentz, S. M. Rubrecht, Wyoming. Rahel. 2000. Status of Yellowstone J. S. Vandeventer, and J. D. Varley. Jones, R. D., G. Boltz, D. G. Carty, L. R. cutthroat trout in Wyoming waters. 1979. Fishery and aquatic management Kaeding, D. L. Mahony, and S. T. North American Journal of Fisheries program in Yellowstone National Park. Olliff. 1993. Fishery and aquatic Management 20:693-705. U.S. Fish and Wildlife Service, technical management program in Yellowstone Mahony, D. L., and J. R. Ruzycki. 1997. report 1978, Yellowstone National Park, National Park. U.S. Fish and Wildlife Initial investigations towards the Wyoming. Service, technical report 1992, development of a lake trout removal Jones, R. D., R. E. Gresswell, D. E. Yellowstone National Park, Wyoming. program in Yellowstone Lake. Pages 153- Jennings, S. M. Rubrecht, and J. D. Kaeding, L. R., G. D. Boltz, and D. G. 162 in R. Hamre, ed., Wild Trout VI. Varley. 1980. Fishery and aquatic Carty. 1996. Lake trout discovered in Trout Unlimited and Federation of Fly management program in Yellowstone Yellowstone Lake threaten native Fishers, Ft. Collins, Colorado. National Park. U.S. Fish and Wildlife cutthroat trout. Fisheries 21(3):16-20. Mattson, D. J., and T. Merrill. 2002. Service, technical report 1979, Koel, T. M., D. L. Mahoney, K. L. Kinnan, Extirpations of grizzly bears in the Yellowstone National Park, Wyoming. Jones, R. D., P. E. Bigelow, R. E. C. Rasmussen, C. J. Hudson, S. contiguous United States, 1850-2000. Gresswell, and R. A. Valdez. 1982. Murcia, and Billie L. Kerans. In press. Conservation Biology 16:1123-1136. Fishery and aquatic management Myxobolus cerebralis in native cutthroat McIntyre, J. D. 1995. Review and program in Yellowstone National Park. trout of the Yellowstone Lake ecosystem. assessment of possibilities for protecting U.S. Fish and Wildlife Service, technical Journal of Aquatic Animal Health. the cutthroat trout of Yellowstone Lake report 1981, Yellowstone National Park, Koel, T. M., J. L. Arnold, P. E. Bigelow, P. from introduced lake trout. Pages 28-33 Wyoming. D. Doepke, B. D. Ertel, and D. L. in J. D. Varley and P. Schullery, eds. The Jones, R. D., D. G. Carty, R. E. Gresswell, Mahony. 2004. Yellowstone Fisheries and Yellowstone Lake crisis: confronting a C. J. Hudson, and D. L. Mahony. 1987. Aquatic Sciences: annual report, 2003. lake trout invasion. A report to the Fishery and aquatic management National Park Service YCR-NR-2004- Director of the National Park Service. program in Yellowstone National Park. 03, Yellowstone Center for Resources, Yellowstone Center for Resources, U. S. Fish and Wildlife Service, technical Yellowstone National Park, Wyoming. Yellowstone National Park, Wyoming.

Acknowledgements Lake trout removal was enhanced through a three-year NPS Wildlife Service Bozeman Fish Health Laboratory; the U.S. Natural Resource Challenge, Natural Resource Preservation Geological Survey Western Fisheries Research Center in Seattle; Program grant, and subsequent NPS base funding. Lynn Kaeding, the Montana Department of Fish, Wildlife and Parks; and the Jeffrey Lutch, John McIntyre, S. Thomas Olliff, James Ruzycki, and Wyoming Game and Fish Department, we have been able to James Selgeby were instrumental in the early development of the learn a great deal about whirling disease in the Yellowstone lake trout removal program. Cutthroat trout monitoring and lake Lake ecosystem. Information on annual cutthroat trout spawn- trout removal would not occur without the dedicated assistance ing migration surveys and bear use of Yellowstone Lake of many seasonal technicians and volunteers, and we are grate- tributaries was provided by Kerry Gunther of Yellowstone ful for their support. Special thanks to Barbara Rowdon for her National Park's Bear Management Office and Daniel Reinhart, leadership on Yellowstone Lake, 2001–2004. Funding for whirling Patrick Perrotti, and Eric Reinertson of the park's disease research has been provided by the National Partnership Resource Management and Visitor for the Management of Wild and Native Coldwater Fisheries, Protection Division. Montana Water Center, U.S. Fish and Wildlife Service, Whirling Disease Foundation, and NPS Rocky Mountain Ecosystem Studies Unit. Through collaboration with the Department of Ecology, Montana State University; U.S. Fish and TODD KOEL

Columbine Creek, a large spawning tributary on the eastern shore, disconected from Yellowstone Lake due to drought conditions in July 2004.

18 Fisheries | www.fisheries.org | vol 30 no 11 Morgan, L. A., W. C. Shanks III, D. A. Lovalvo, S. Y. Johnson, W. J. Stephenson, K. L. Pierce, S. S. Harlan, C. A. Finn, G. Lee, M. Webring, B. Schulze, J. Duhn, R. Sweeney, and L. Balistrieri. 2003. Exploration and discovery in Yellowstone Lake: TrackTrack youryour fishfish withwith results from high-resolution sonar imaging, seismic reflection profiling, and submersible studies. Journal of Volcanology and thethe newest,newest, mostmost Geothermal Research 122:221-242. Munro, A. R., T. E. McMahon, and J. R. Ruzycki. 2005. Natural advancedadvanced acousticacoustic chemical markers identify source and date of introduction of an exotic species: lake trout (Salvelinus namaycush) in Yellowstone tracking receiver Lake. Canadian Journal of Fisheries and Aquatic Sciences 62:79- tracking receiver 87. Reinhart, D. P. 1990. Grizzly bear habitat use on cutthroat trout availableavailable today.today. spawning streams in tributaries of Yellowstone Lake. M.S. thesis. Montana State University, Bozeman. Reinhart, D. P., and D. J. Mattson. 1990. Bear use of cutthroat trout spawning streams in Yellowstone National Park. International Conference on Bear Research and Management 8:343-350. Reinhart, D. P., S. T. Olliff, and K. A. Gunther. 1995. Managing bears and developments on cutthroat spawning streams in Yellowstone Park. Pages 161-169 in A. P. Curlee, A. M. Gillesberg, and D. Casey, eds. Greater Yellowstone predators: ecology and conservation in a changing landscape. Proceedings of the 3rd biennial conference on the Greater Yellowstone Ecosystem, Northern Rockies Conservation Cooperative, Jackson, Wyoming. Reinhart, D. P., M. A. Haroldson, D. J. Mattson, and K. A. Gunther. 2001. Effects of exotic species on Yellowstone's grizzly bears. Western North American Naturalist 61:277-288. Ruzycki, J. R., D. A. Beauchamp, and D. L. Yule. 2003. Effects of introduced lake trout on native cutthroat trout in Yellowstone Lake. Ecological Applications 13:23-37. Schullery, P., and J. D. Varley. 1995. Cutthroat trout and the Yellowstone Lake ecosystem. Pages 12-21 in J. D. Varley and P. Schullery, eds. The Yellowstone Lake crisis: confronting a lake trout invasion. A report to the director of the National Park Service. Yellowstone Center for Resources, Yellowstone National VEMCO proudly Park, Wyoming. Schwartz, C. C., M. A. Haroldson, K. A. Gunther, and D. Moody. introduces the VR100 2002. Distribution of grizzly bears in the Greater Yellowstone Ecosystem, 1990-2000. Ursus 13:203-212. Acoustic Tracking Receiver. Spencer, C. N., B. R. McClelland, and J. A. Stanford. 1991. Shrimp stocking, salmon collapse, and eagle displacement: cascading ou can count on the VR100 to get the job done - interactions in the food web of a large aquatic system. Bioscience Ywhether you’re actively tracking large pelagic fish 41:14-21. Stapp, P., and G. D. Hayward. 2002. Effects of an introduced or conducting presence/absence passive monitoring. piscivore on native trout: insights from a demographic model. The VR100 has a flexible systems architecture with Biological Invasions 4:299-316. 8MB of non-volatile internal memory, GPS positioning Stott, W. L. 2004. Molecular genetic characterization and comparison and precise timing, USB link to PC or laptop, and field of lake trout from Yellowstone and Lewis Lake, Wyoming. U. S. installable software upgrades. Other features include: Geological Survey research completion report for project 1443-IA- 15709-9013 to the National Park Service, Yellowstone National 4 Simultaneous, multi-frequency reception and Park, Wyoming. Swenson, J., K. L. Alt, and R. L. Eng. 1986. Ecology of bald eagles detection tracking algorithms in the greater Yellowstone ecosystem. Wildlife Monographs 95, 4 Wide dynamic range allowing multi-tag reception The Wildlife Society, Bethesda, Maryland. without gain adjustment Varley, J. D. 1981. A history of fish stocking activities in Yellowstone 4 Splash proof case with marine grade connectors National Park between 1881 and 1980. Information paper 35. 4 Coded and continuous tags Yellowstone National Park, Wyoming. 4 Operation frequency 10-400kHz _____. 1983. The use of restrictive regulations in managing the wild salmonids in Yellowstone National Park, with particular reference to the cutthroat trout, Salmo clarki. Pages 145-156 in J. M. Walton VEMCO (a division of AMIRIX Systems Inc.) and D. B. Houston, eds. Proceedings of the Olympic wild fish Tel: 902-852-3047 Fax: 902-852-4000 conference. Peninsula College, Fisheries Technology Program, Port Angeles, Washington. www.vemco.com Varley, J. D., and R. E. Gresswell. 1988. Ecology, status, and management of the Yellowstone cutthroat trout. American Fisheries Society Symposium 4:13-24. Making Waves in Varley, J. D., and P. Schullery. 1998. Yellowstone fishes: ecology, Making Waves in history, and angling in the park. Stackpole Books, Acoustic Telemetry Mechanicsburg, Pennsylvania.

November 2005 | www.fisheries.org | Fisheries 19 Smallmouth Bass Introductions in Maine: History and Management Implications

This article traces the early introductions of nonnative smallmouth bass (Micropterus dolomieu) in Maine from 1868 to 1898, as documented primarily in reports of the Maine Commissioners of Fisheries for those years. Early handling and transportation efforts were crude and primitive by today’s standards, but by trial and error they were fisheries history eventually effective. The concepts of using donor waters and public participation were ABSTRACT developed to the point that introductions could not be controlled by the state, a situ- ation that continues today. The commissioners were adamant that bass should be

feature introduced in suitable waters, and should not be introduced in salmonid waters because of possible detrimental effects on native fauna. Despite their wishes, many in unauthorized introductions have been made in the past 100 years, including many salmonid waters. Although it is virtually impossible to prohibit all illegal bass introduc- tions, expanded educational programs and more stringent laws and law enforcement should aid in this effort.

Introduction use this information as a basis to make observations on current efforts to limit the spread of bass in Maine waters, Illegal introductions of nonnative smallmouth bass especially given that Maine waters appear to be warming. (Micropterus dolomieu) have become rampant in Maine over the past 20 years. Introduction of any fish Methods Kendall Warner species into Maine waters is illegal without a permit from the Commissioner of Inland Fisheries and As sources of information on early bass introductions, Warner is a retired Wildlife. Since 1986, the number of smallmouth bass I searched annual reports of the Maine Commissioners of fisheries management waters has increased by 26 due to illegal stockings Fisheries (1867–1898) and Maine Sportsman supervisor who worked (Jordan 2001), despite the implementation of stiff (1893–1895). I also reviewed writings by Henshall at the Maine Department civil (now criminal) penalties imposed by the Maine (1891–1898), Prime (1892), Cheney (1896), Hodgkins Inland Fisheries and legislature and educational efforts by the Maine Wildlife, Bangor. He can Department of Inland Fisheries and Wildlife et al. (2003), Whittier et al. (1997), Jordan (2001), and be contacted at (MDIFW). These recent efforts of maintaining strict Jackson (2002). The primary data that I sought were [email protected] control over the introduction of fish species known to dates and locations of early authorized introductions, . be harmful to native fishes have been unsuccessful. methods used, current distribution, spread of authorized However, any notion that historic bass introductions introductions, and effects on native fish fauna. in Maine were ever under complete state authority is not substantiated by historical distribution records. Results The purpose of this article is to review introduction of smallmouth bass (hereafter referred to as bass) in Maine as Initial establishment a basis for increased understanding of how this fish has, and is, affecting Maine’s fish communities. This article As early as 1868 the Maine commissioners began to will document the types of waters in which authorities give serious thought to introducing a new fish to Maine’s conducting initial releases wanted to release bass, their fauna that would provide sport fisheries in waters con- concerns about bass effects on native fisheries, whether or taining only “worthless” perch (Perca flavescens), sunfish not their concerns were justified, and whether the poten- (Lepomis spp.), and pickerel (Esox niger; Foster and tial range of bass in Maine will continue to expand. I will Atkins 1869). The commissioners considered sources and species of bass, methods of transportation, both into and within the state, sizes of bass, and choice of waters to be stocked. They were adamant that bass not be stocked in salmon and trout waters, as stated by Stanley (1882:15), “It should never be introduced into any waters where there are trout, or from whence it can gain access to trout streams. For ponds whose stock of trout has been exhausted by poachers who murder the fish on their spawning beds, and where no fish but yellow perch, bream, and pickerel are left, it is invaluable.” The first introductions of bass in Maine waters occurred in 1868 (Foster and Atkins 1869), when seven waters in central and southern Maine received this new game fish. Waters and dates of subsequent

20 Fisheries | www.fisheries.org | vol 30 no 11 authorized bass introductions from 1868 to 1881 are I shall take occasion in this report to advise the listed in Table 1. encouragement of private enterprise in the field of Because earlier attempts in transporting bass fry from fish culture, and the limitation of the State’s New Hampshire failed, small numbers of older fish from agency to such operations as seem to be beyond Newburg, New York, were used for initial stockings the reach of private effort. It will be sufficient here Table 1. Early known (Atkins 1870). to say that in my opinion another season would introductions of The commissioners selected waters for most early bass enable the commissioner to do all in the way of smallmouth bass in Maine introductions, but non-sanctioned introductions were introducing black bass that the State ought to do. waters, listing those with current principal fisheries inevitable when they began to consider public participa- It will be seen that six colonies of them are already for bass and those tion in the distribution process, as stated by Atkins established in the State. As soon as they have mul- currently managed for (1870): tiplied in these waters they can be transported to coldwater gamefish.

Water Town County Year Current principal Current introduced bass fishery management Duck Pond (Highland Lake) Falmouth Cumberland 1868 No Warmwater Fitts Pond Clifton Hancock 1868 No Coldwater Newport Pond (Sebasticook Lake) Newport Penobscot 1868 Yes Warmwater Phillips Pond (Phillips Lake,Lucerne Lake) Dedham Hancock 1868 Yes Coldwater Cochnewagon Pond Monmouth Kennebec 1868 Yes Combination Cobbosseecontee Lake (Cobbbossee Lake) Winthrop Kennebec 1868 Yes Warmwater Whitney Pond Canton Oxford 1868 — — Dexter Pond (Wassookeag Lake) Dexter Penobscot 1869 Yes Coldwater Himes Pond Brewer Penobscot 1875 — — Eddington Pond (Davis Pond) Eddington Penobscot 1875 Yes Warmwater Stewart’s Pond Corinna Penobscot 1875 — — Pushaw Pond (Pushaw Lake) Glenburn Penobscot 1875 Yes Warmwater Madigan Pond Houlton Aroostook 1875 — — Long Pond (Caribou, Egg, Long ponds) Lincoln Penobscot 1875 Yes Warmwater China Pond (China Lake) China Kennebec 1875 Yes Combination Snow Pond (Messalonskee Lake) Belgrade Kennebec 1875 Yes Combination Goose Pond Hartland Somerset 1875 — — Lovewell’s Pond Fryeburg Oxford 1875 Yes Combination Trip (p) Pond Poland Cumberland 1875 Yes Combination Bryant’s Pond (Christopher Lake) Woodstock Oxford 1875 No Combination Sabattus Pond Greene Androscoggin 1875 Yes Warmwater Taylor’s Pond Auburn Androscoggin 1875 Yes Combination Crotched Pond Bridgton Cumberland 1875 — — Worthly Pond Peru Oxford 1876 Yes Combination Ellis River Pond (Roxbury Pond) Roxbury Oxford 1876 Yes Warmwater Brettun’s Pond Livermore Androscoggin 1876 Yes Combination Jennie Pond Mexico Oxford 1876 — — Bean Pond Turner Androscoggin 1876 No Warmwater North Pond Sumner Oxford 1876 Yes Combination Great Pond (Long Pond) Belgrade Kennebec 1876 Yes Combination Meddybemps Lake Alexander Washington 1877 Yes Combination LaCoote Pond Vanceboro Washington 1877 No Warmwater Sidney Pond Vassalboro Kennebec 1877 — — Hermon Pond Hermon Penobscot 1877 Yes Warmwater Little Bear Pond Dexter Penobscot 1877 — — Halfmoon Pond Prospect Waldo 1877 Yes Coldwater Mattaceunk Lake (Mattaseunk Lake) Mattawamkeag Penobscot 1877 No Coldwater Goose Pond Dedham Hancock 1877 Yes Warmwater Reeds Pond Dedham Hancock 1877 — — Thompson Lake Oxford Oxford 1877 No Coldwater Norway Lake (Pennesseewassee Lake) Norway Oxford 1877 Yes Combination Songo Pond Albany Oxford 1877 Yes Combination Woodbury Pond (Purgatory Pond) Litchfield Kennebec 1877 Yes Warmwater Sand Pond (Tacoma Lake) Litchfield Kennebec 1877 Yes Combination Little Purgatory Pond Litchfield Kennebec 1877 Yes Warmwater Gardner’s Pond (Gardner Pond) Wiscasset Lincoln 1881 Yes Warmwater Hosmer Pond Camden Knox 1881 Yes Warmwater Kezars Pond (Kezar Lake) Lovell Oxford 1881 Yes Coldwater Heald Pond Lovell Oxford 1881 No Warmwater Cushman Pond Lovell Oxford 1881 No Combination Little Pushaw Lake Corinth Penobscot 1881 Yes Warmwater

November 2005 | www.fisheries.org | Fisheries 21 various others at a comparatively small expense. It bass introductions. Few of the early commissioners might be desirable to establish nurseries at two or had biological training, however, and there were little three other points convenient of access, and leave basic data available on water quality or fish popula- their further dissemination to private effort. tions. Charles G. Atkins was the exception; he was a self-taught fish-culturist and naturalist. For the most Sporting camp owners, politicians, and town officials part, choice of waters for early bass introductions was were recruited to secure bass from some of the seven orig- probably based on the “reputation” of certain waters inal introductions (donor waters) into other candidate in lieu of biological background data. Those waters waters. (Table 1), and most subsequent introductions renowned for their salmon and trout fisheries were were not controlled by the commissioners. The next fisheries history avoided. The problem of unauthorized bass introduc- problem was in securing bass from donor waters for fur- tions began when bass distribution authority was ther introductions, but because initial efforts to hire relinquished to the public. Even the commissioners people to angle bass from “donor waters” for further intro- prior to 1898 admitted that unauthorized bass intro-

feature ductions were largely unsuccessful, the commissioners ductions were “substantial.” decided to undertake this effort themselves. This same report showed that the commissioners were Historical effects on other fish species enamored with bass and took pains to justify their views: Diverse public opinions on bass introductions We do not deem the bass more destructive to the were recognized by the commissioners as early as young of other fishes than the trout, much less so 1898 in the following observations (Carleton et al. than the white perch or the yellow perch, for both 1899:13): the trout and the black bass feed largely on flies….For stocking ponds infested with pickerel, There are various opinions held by the citizens of we know of no fish equal to the black bass. It breeds this State relative to this fish. In some localities more rapidly than any other fish….The young they are held in great favor, while in others no con- grow with wonderful rapidity….Their flesh when demnation is too severe to be hurled at them. They cooked has the same intense white color as that of are very destructive to the pickerel and usually suc- the halibut, is very juicy and sweet in flavor. It is a ceed in practically exterminating them in waters very hard fighter when hooked. where they have placed, and the pickerel being an inveterate enemy to the trout and land-locked By 1878, the authorities were gratified with their suc- salmon, the black bass have in this respect been of cess in bass introductions, but they apparently began to great practical benefit, and besides they are a very run out of funds for this effort: gamy fish to catch, and at certain seasons of the There has been a wide call for black bass from all year are as good a pan fish as there is. We have not parts of the State….We were obliged to close our for many years, however, stocked any of the waters operations in distributing bass, not from having of Maine with black bass, and the laws for their filled all our orders, but from want of funds. protection have all long since been repealed, with (Stillwell and Stanley 1878). one exception. Further observations on both this growing popularity Current status of original introductions of bass fishing as well as examples of harmful effects to other fishes were noted in early issues of Maine Sportsman A total of 51 waters received “authorized” introduc- (1893–1895): tions of bass in Maine from 1868 to 1881. Of these, 40 waters (80.4%) could still be identified in the current The Winthrop Budget gives a brief account of the MDIFW Lake Inventory File 2001. The remaining 11 stocking of the Kennebec Lakes with bass, and of waters were probably either private ponds or ponds for the resulting advantages and disadvantages from its which the name has been changed. Original point of view. The Budget says: ‘The first bass (1868–1881) and 2001 distributions (Jordan 2001) are placed in any of the Winthrop lakes were brought compared in Figure 1. It appears that original introduc- here in the summer of 1869 by the late George tions in central and southern Maine have gradually Shepard Page of New York, who put them into the Cobbosseecontee at East Winthrop. As the bass spread out to include many northeastern and southwest- multiply rapidly, the lakes were soon overrun with ern waters, but most northwestern waters and those in them, greatly to the detriment of the other fishing. extreme northern Maine still do not contain bass. Of One thing is certain, the pickerel and white perch these original 40 introductions, 18 waters (45%) are cur- has been decreasing ever since, while the shore rently classified as warmwater fisheries, and an additional fish, formerly used exclusively for bait for perch 14 waters (35%) are managed for combined warmwater have disappeared entirely. and coldwater fisheries. Only 8 waters (20%) are man- aged for strictly coldwater fisheries. Principal bass The Lewiston Sun says that while the trout fisher- fisheries now occur in 32 (80.0%) of the original autho- men are passing much time on Lake Auburn rized introductions. Principal fisheries are those where a with varied success, the anglers of the two cities species is sought after by anglers and makes up a signifi- who like bass fishing are having no end of sport cant portion of the catch, or, if the only sport species on the river. Bass may be taken without diffi- present, contributes to a significant catch. culty above and below the falls. The bass have It appears that the early commissioners were basi- increased at a wonderful rate in the cally “on target” for waters that they authorized for Androscoggin during the last year. There are so

22 Fisheries | www.fisheries.org | vol 30 no 11 Figure 1. Historical and current distribution of smallmouth bass in Maine waters.

Legend ★ Original introductions—1868–1881 o Smallmouth Bass present in lake survey database—2000

November 2005 | www.fisheries.org | Fisheries 23 many of them that they have nearly extermi- either of the bass species had significantly fewer species nated the red-fin minnows that are caught as of cyprinids than lakes lacking bass. bait for pickerel. Pickerel fishermen who used to I requested information on more recent effects of bass get all of their bait from the river a few years ago say introductions on native fish fauna from current Fishery that there are so few there now that they are Division, MDIFW staff. An illegal smallmouth bass intro- obliged to go elsewhere. duction is believed to be the primary cause for a decline in Cheney (1896) identified New York state as the the abundance and size quality of a stocked brook trout original source of bass stocked in Maine waters. He (Salvelinus fontinalis) fishery in Keyes Pond (Sweden, wrote of their spread in New York: Maine). Bass were introduced in the mid-1980s and fisheries history became well established by the early 1990s. This water his- …stocking of small streams and small ponds torically produced brook trout up to 6 lbs., a rarity in with black bass was being prosecuted with southern Maine. In 1988, trap netting resulted in the cap- vigor. There was a black bass craze; waters ture of 17 brook trout over a 6-day period. Trap netting in feature wholly unfit to the fish were planted, trout 1998 over an 8-day period produced only 1 brook trout (F. waters were destroyed for no judgment was Brautigam, MDIFW, pers. comm.). Similarly, yellow perch exercised in the general black bass planting. As and chain pickerel apparently disappeared from Trickey the fruit of these indiscriminate stocking of Pond (Naples) after introduction of smallmouth bass. A bodies of water, we have today hundreds of winter creel survey in 2004 revealed no perch or pickerel ponds that offer no fishing worthy of the name; in the catches (J.C. Pellerin, MDIFW, pers. comm.). and this applies not only to trout waters that In eastern Maine waters, smallmouth bass presence is have been planted with black bass, but to believed to have affected brook trout abundance in at least waters planted with the same fish which before nine waters (R. Brokaw, R.M. Jordan, G. Burr, MDIFW, contained only coarse fish, so called. pers. comm.). According to R. K. Brokaw (MDIFW, pers. Cheney (1896) finally cautioned: comm.), smallmouth bass introductions were a primary cause of cyprinid declines in Beech Hill Pond (Otis), The Commission realizes the importance of Molasses Pond (Eastbrook), and Indian Lake (Whiting). confining black bass to the waters in which Similarly, survey data for Indian Pond (East Outlet, they are now found, and entrusts the planting of the fish only to its own men.…The spread Moosehead Lake) indicates establishment of illegally should be checked, in spite of the desire…to introduced smallmouth bass caused a decline in brook plant the fish in yet new waters, by those who trout angling success, although anglers who have kept vol- do not understand the harm that may come untary records have shifted from those who target brook from it. To all such this is written as a warning.” trout to those who target smallmouth bass. It is expected that brook trout will cease to be the principal fishery in This was not the case in Maine where most bass Indian Pond and Moosehead Lake (P.R. Johnson, introductions after 1881 were left to the public. MDIFW, pers. comm.). Prime (1892:72-73), a fisherman who traveled Smallmouth bass were illegally introduced into widely in New England, wrote of his opinions on bass Umbagog Lake in the 1980s and, in the late 1990s, had introductions: worked their way up the Rapid River to Middle Dam and Fish Commissioners in some of our States have up the Magalloway to Aziscohos Dam. Rapid River sup- laboriously spoiled the fishing in a great many ports a wild brook trout fishery of national significance; waters by introducing these black bass. Pickerel this fishery is threatened by the presence of smallmouth or perch or pumpkin-seeds are more valuable bass. Brook trout fry, which occur along the margins of the for a fish to the farming population than black river, are vulnerable to predation from young bass, which bass, and black bass when placed in a pond will occupy the same niche. Extensive efforts, in terms of destroy all other fish.” research and fund-raising efforts, are being applied in an effort to mitigate the effects of the bass introduction on Effects on other fish species today this fishery, which produced brook trout several pounds in size. Within this same major drainage, brook trout also Negative effects of bass introductions on minnow reproduce in the lower Magalloway River; the presence of species (Cyprinidae) have been mentioned previously. bass is expected to impact reproductive success in that Whittier et al. (1997) found that cyprinid abundance river as well. (F. Bonney, MDIFW, pers. comm.) and diversity in New England waters increased from Smallmouth bass were illegally introduced in south to north, with Maine having 3 to 7 minnow Moosehead Lake and in Moxie Lake in the 1980s. From species constituting 40–80% of species in each lake. these sources, they are spreading downstream into the The most consistent factor related to minnow species Kennebec River and its tributaries. Bass are commonly richness was the presence of littoral predators. caught in the Kennebec Gorge below Harris Dam and Nonnative predators, especially Micropterus spp., have have reduced the abundance of wild brook trout, as deter- been introduced throughout the Northeast: 69% of mined from voluntary angler records. Bass have moved up the sampled lakes had nonnative predators. the Dead River from the Forks to Grand Falls and are Recently, Jackson (2002) found in two case studies expected to impact the wild brook trout fishery there. strong negative associations between small-bodied They also have access to Little Spencer Stream and fishes, primarily cyprinids, and smallmouth bass and Spencer Stream as far as Spencer Gut; these streams, as largemouth bass (M. salmonides). Lakes containing well as a number of smaller tributaries, also contain wild

24 Fisheries | www.fisheries.org | vol 30 no 11 brook trout populations. The dam on the outlet of Spencer Lake has dete- According to Patterson et al. (2001), bass ranked highest of all riorated to the point where bass may soon be able to negotiate it and Maine sportfish in largest number of anglers, most angler days of use, move up into Spencer Lake, an oligotrophic lake that supports brook and most frequently caught species. Smallmouth and largemouth bass trout, lake trout (S. namaycush), and landlocked salmon (F. Bonney, now occur in 613 Maine lakes and ponds, with smallmouths occurring MDIFW, pers. comm.). in 471 of these waters. In 231 waters smallmouths co-exist with large- The impact of illegal smallmouth bass introductions on wild mouth bass, and in 240 waters, smallmouths are the only species of salmonid populations is not well documented. Efforts to do so include bass present (Jordan 2001). the Rapid River initiative, a survey of Cold Stream in 2004 (Cold Jordan (2001) noted that many oligotrophic lakes in Maine are Stream is the primary brook trout spawning tributaries to the commonly managed for coldwater species, but bass populations in Kennebec Gorge), and “spot” electrofishing in 2005 in other these lakes often produce important fisheries. This is reflected in the Kennebec River tributary to determine the extent of their movement fact that 35% of the original authorized introductions are currently through the drainage (F.Bonney, MDIFW, pers. comm.). managed for combination coldwater-warmwater fisheries. Of 613 bass I personally recall seining and trapping great quantities of min- waters in Maine, 216 (35%) are being currently managed for a com- nows from beaches on Raymond Cape, Sebago Lake, in the early bination of coldwater and warmwater sport fisheries. This includes 1930s. White perch (Morone americana) were common then, but bass both smallmouth and largemouth bass. (Jordan 2001). Although bass were rare. Minnow populations were decimated as a result of DDT have undoubtedly caused irreversible impacts to faunal diversity in spraying in the 1960s. Bass populations are now abundant, but min- many waters, they should not be universally condemned in all situa- nows are rare, except in a few localities. tions. Bass are producing superb fisheries in many waters and provide thousands of hours of recreational fisheries. The tendency toward global warming is well documented. Discussion Hodgkins 2002 and Hodgkins et al. 2003 found that ice-out dates in lakes and changes in river flows are good indicators of climate-related The early commissioners were concerned about waters infested changes. These data suggest that in New England, spring biological with “worthless” pickerel, perch, and sunfish that could prove to be a changes were all caused by a common mechanism—temperature boon to Maine’s fisheries by introducing another gamefish—black increases. Effects of these temperature increases on Maine’s fish habi- bass. They apparently gave serious thought to the species of bass, tat are only conjecture. It is relatively certain, however, that some sources, costs, and methods of transportation, both into and within marginal coldwater habitat currently managed for salmonids will be the state. Modes of transportation were crude and primitive by today’s lost, especially waters with abundant competing species. standards, but these were refined by trial and error and became quite Maine law currently prohibits transportation or introduction of effective. any fish within the state without a written permit (MDIFW 2001). The commissioners originated the concept of “donor waters” in There is a maximum $10,000 fine for conviction of an illegal stock- Maine. Once bass populations were established in a few key waters, ing. Operation Game Thief is also offering a $2,000 reward for these waters would serve as sources for further introductions. information leading to the apprehension of anyone responsible for the They also emphasized “public participation” as vital in introducing illegal introduction of fish into any Maine water. Despite these efforts, bass into “suitable” waters. Members of the public, town and political at least 10 illegal fish introductions were confirmed for Maine waters officials, sporting camps, and anglers’ associations were recruited in in 2003. In addition to these deterrents, in waters with over-abundant this effort. Their roles were to aid in collection, transportation, and bass populations causing stunting and/or excessive competition with distribution of bass. Collection efforts by some of these groups were coldwater fisheries, a special regulation (S-13) providing “no size or often fruitless, so the commissioners went out and spent many hours bag limit on bass” may be established. There are currently 62 S-13- catching bass themselves. Candidate waters were sometimes desig- regulations on Maine waters. Some sentiment recently has been nated by the commissioners, but more often decisions were left to the expressed to apply this rule to major combination-management judgment of local people, who usually had no idea of what waters were waters, where good fisheries occur for both bass and one or two cold- actually suitable for bass. The rationale seemed to be “if it had pickerel, water fishes. The rationale seems to be to manage only for coldwater perch, and sunfish, it must be good bass water.” As a result, no one will gamefish, with no restrictions on bass. Although this rule may be ben- ever know the dates of bass introductions into many Maine waters. eficial in some rare situations where nursery area is limited, it should The commissioners were adamant that bass not be introduced in not be applied universally, especially when special rules on bass are salmon and trout waters, as they recognized possible detrimental effects now in effect to protect spawning larger, older bass. on Maine’s native gamefishes. They lost control, however, when they Furthermore, in the opinion of one biologist (D. P. Boucher, allowed the public to take over bass introductions. The ultimate result MDIFW, pers. comm.), the S-13 (no size or bag limit on bass) regula- was many illegal introductions into coldwater lakes and ponds, a prac- tion has no discernable impact on either the bass populations or tice that continues today. salmonid fisheries. In fact, there is now good evidence to suggest that The opinion of the commissioners was shared by at least one bass heavy harvest of stunted bass populations may enhance bass recruit- advocate in the late 1800s. Henshall (1881), an ardent angler for both ment, because intra-specific competition is reduced, and the remaining trout and bass, wrote: bass are physiologically more fit to produce successful broods. MDIFW continues an aggressive campaign to apprehend and pros- That he will eventually become the leading gamefish of ecute those guilty of illegal fish transportation and introductions. America is my oft-expressed opinion and firm belief. This Public awareness of the dangers of these introductions has been result, I think, is inevitable; if for no other reasons, from a force heightened by exposure through the various media. Both of these of circumstances occasioned by climatic conditions and the efforts should be continued and expanded in the future. operation of immutable natural laws, such as the gradual drying Lack of public awareness, however, is still a problem. According to up, and dwindling away of the small Trout streams, and the Jackson (2002): consequent decrease of Brook Trout, both in quality and quan- tity; and by the introduction of predatory fish in waters where There is a clear recognition that introducing nonindigenous the Trout still exists. predators into aquatic systems may have major ecological con-

November 2005 | www.fisheries.org | Fisheries 25 sequences but there appears to be little public awareness about _____. 1898. More about the black bass, Supplement to the book of the the ecological impact....This case of ‘out of sight—out of black bass. Robert L. Clarke and Co., Cincinnati. mind’ may explain that dichotomy in how terrestrial and Hodgkins, G. A. 2002. Historical changes in lake ice-out dates as aquatic systems are considered with respect to introducing indicators of climate change in New England, 1850-2000. major predators. International Journal of Climatology 22: 1819-1827. Hodgkins, G. A., R. W. Dudley, and T. G. Huntington. 2003. Changes Acknowledgements in timing of high river flows in New England over the 20th century. Journal of Hydrology 278:244-252 Jackson, D. A. 2002. Ecological effects of Micropterus introductions: the I thank the following members of the Fisheries and Hatcheries dark side of black Bass. Pages 221-231 in D. P. Phillip and M.S. Division, MDIFW for their assistance in various phases in preparation Ridgeway, eds. Black bass: ecology, conservation, and management. of this article. M. Gallagher, T. C. Obrey, S. Seeback and P. Wick American Fisheries Society Symposium 31, Bethesda, Maryland. assisted in data retrieval and map preparation. F. R. Bonney, F. C. Jordan, R. M. 2001. Black bass assessment. MDIFW 2001. Maine Brautigam, R. K. Brokaw, G. Burr, P. R. Johnson, R.M. Jordan, and J. C. Department of Inland Fish and Wildlife, Augusta. Pellerin provided recent data on effects of smallmouth bass on associ- Maine, State of. 2001. 2001 Inland Fisheries and Wildlife Laws. 12 ated fishes. D. P. Boucher, J. D. McNeish, and P. M. Bourque deserve MRSA Part 10. Chapters 701-721 and 811 as enacted by Public law, grateful thanks for their painstaking peer reviews of various drafts of the Chapter 420 Section 1 and as amended: 103-104 manuscript. W. B. Krohn also reviewed several drafts and provided Patterson, R. W., D. O. Scroggin, K. J. Boyle, and D. McNeish. 2001. much pertinent literature and advice. I am especially indebted to Diane Maine open water fishing survey, 1999. Staff Paper REP 493, July 2000 Letteney for patiently typing many revisions of the manuscript. (revised January 2001). MDIFW, Augusta. Prime, W. C. 1892. Along New England roads. Harper and Brothers, New References York: 72-73 Stanley, H. O. 1882. Report of the Commissioners of Fisheries and Game, Maine for 1881:15. Atkins, C. G. 1870. Pages 18-21 in Third Report of the Commissioners of Fisheries, Maine—Augusta (1869). Stillwell, E. M., and H. O. Stanley. 1875. Ninth Report of the Carleton, L. T., H. C. Stanley, and C. E. Oak. 1899. Report of the Commissioners of Fisheries, Maine (1875):14-15. Commissioners of Inland Fisheries and Game, Maine (1898):13. _____. 1878. Twelfth Report of the Commissioners of Fisheries, Maine Cheney, A. N. 1896. Report of State of New York Fisheries, Game and (1878):15-16. Forest Commission for 1896: Albany 177-184. Whittier, T. R., D. B. Halliwell, and S. G. Paulson. 1997. Cyprinid Foster, N. W. and C. G. Atkins. 1869. First Report of the Commissioners distributions in Northeast USA lakes: evidence of regional-scale of Fisheries, Maine (1867): 87-88. minnow biodiversity losses. Canadian Journal of Fisheries and Aquatic Henshall, J. A. 1881. Book of the black bass. Clarke and Co., Cincinnati. Sciences 54:1593-1607.

26 Fisheries | www.fisheries.org | vol 30 no 11 column director’s line JAN LUBECK Halibut Cove

Clem Tillion’s house in Halibut Cove

For a while in this past September, it looked The visit to Halibut Cove proved to be a fitting like the city of Anchorage in Alaska was overflow- coda for a once in a lifetime fisheries meeting. ing with fish, fisheries people, and fishing tales. The respect shown by people like Clem for the The annual meeting of the Society attracted more physical and biological surroundings and the than 2,800 scientists and professionals, some 900 themes expressed in the meeting poster, more than the previous record in Portland back in designed by Alaska artist Ray Troll, summarize 1993. With more than 1,900 papers and posters, the mission of AFS: when people respect their 50 symposia, many continuing education courses, natural habitat and its living resources, future and social events galore, the meeting proved once generations will benefit. again that location draws people and when the Lake Placid next year will have a hard act to location is Alaska and the people are fisheries follow but the contrast between mountainous folks, then the combination is a natural. northern New York and the Kenai Peninsula only Many of the attendees took advantage of pre- points out the diversity of the AFS people and the Gus Rassam and post-meeting opportunities to go fishing habitats in which they live and work. Joe Margraf, Anchorage program chair for contributed papers, AFS Executive Director (mostly halibut), hiking, glacier gawking, and, for expressed the great sense of camaraderie exhib- Rassam can be reached those lucky ones, sighting the magnificent wildlife ited at AFS meetings in a recent e-mail: at [email protected]. of Alaska: moose grazing on the roadside, black bears eyeing the stray tourist with curiosity, beluga I often wonder about (and pity) people who whales surfacing for air, otters playing games with view going to scientific meetings—especially onlookers, and eagles watching carefully for the AFS—solely as a venue for listening to talks. For catch of the day. Once the meeting was over, several of the AFS me that’s only one small reason for going. I leaders and some staff took advantage of the invi- went to my first AFS meeting in 1970—the tation of Halibut Cove resident Clem Tillion (AFS centennial meeting—and met so many well- member since 1979!) and his lovely wife Diana to known fisheries scientists. I’ll always remember visit their home overlooking the cove. We drove to that meeting for many reasons, but I can’t tell Homer, calm after the summer tourist season, you about a single talk, although I’m sure I went where Clem picked us up in his boat and con- to many. I’ve been to 30 Annual Meetings since ducted us to a wonderful visit in his home. Diana, who is an artist, gave us a tour of her studio adja- 1970, including the last 26 in a row—I missed cent to their home. She uses octopus ink for many West Yellowstone in 1979 for my daughter’s of her drawings and paintings and concentrates birth. I intend to keep my string intact for as on the natural themes of her surroundings. long as I can into the future.

November 2005 | www.fisheries.org | Fisheries 27 Symposium Sparks Debate: forum economic growth Should the American Fisheries Society adopt a position on economic growth?

“Connections Between Economic Growth and posed in a slideshow by Phil Pister. The position was Fish Conservation,” a symposium held at the 2005 a condensed version of the position advanced by the American Fisheries Society (AFS) conference, included AFS Water Quality eight presentations and a lively concluding discus- Section following the sion. The symposium dovetailed with the “Economic 2004 AFS Annual Growth Forum” of Fisheries Volume 30 and offered Meeting. It contained a chance for AFS members to discuss perhaps the nine precepts or most important issue facing the fisheries profession “whereas” clauses and today: the relationship of economic growth to fish five resulting planks or conservation. Ultimately, the key questions raised “therefore” clauses: were: (1) Should AFS take the position that there is a conflict between economic growth and fish conser- Whereas vation? (2) Should AFS take the position that the (Precepts): establishment of a steady state economy is a more appropriate goal for certain nations than the peren- (1) Economic growth is an increase in the production and consumption nial goal of economic growth? Jerry V. Mead As a professional organization devoted to the of goods and services, and; study and conservation of aquatic ecosystems, AFS (2) Economic growth occurs when Stephen M. Coghlan Jr. may choose to adopt a formal position on economic there is an increase in the product Patricia F. Thompson growth for presentation to the public and the federal of population multiplied by per Mead is a Ph.D. candidate and government, much as the U.S. Society of Ecological capita production and consump- Thompson is a M.S. candidate at tion, and; Economics did in 2003 and the North America the Systems Ecology Laboratory, Section of the Society for Conservation Biology and (3) Economic growth is often and College of Environmental Science The Wildlife Society did in 2004. By reporting the generally indicated by increasing and Forestry, State University of events of the symposium, we hope to stimulate AFS real gross domestic product (GDP) New York, Syracuse. Coghlan is members to investigate the arguments more closely or real gross national product a post-doctoral research fellow and make their opinions known to the Society (GNP), and; and adjunct assistant professor regarding a potential AFS position. (4) Based upon established principles at the Department of Chemistry Our approach is to present a summary of events of physics and ecology, there is a and Physics, Department of as we remember them. We do not attempt to pre- limit to economic growth, and; Environmental Sciences, Arkansas State University, Jonesboro. sent events in chronological order or to recount every (5) A steady state economy is gener- Mead can be contacted at fact or anecdote articulated. Rather, we do attempt ally indicated by stabilized (or [email protected]. to distill the major themes for clarity as objectively as mildly fluctuating) real gross possible. We note that, although we have taken con- domestic product (GDP) or real ventional and ecological economics courses, we are gross national product (GNP), and; trained primarily as ecologists, not as economists. (6) A steady state economy, with a stabilized (or However, we believe that ecologists are essentially mildly fluctuating) product of population multi- “economists of nature.” Ecologists and conventional plied by per capita consumption, is an economists alike study the processes of production alternative to economic growth; and; and consumption, allocation of resources, and distri- (7) A steady state economy, with stabilized (or bution of product. Conventional economists focus on mildly fluctuating) production and consumption these processes as exclusively human endeavors of goods and services, is an alternative to eco- while ecologists tend to deal with a wider scope of nomic growth, and; species. Two groups adopting distinctly different view- (8) The North American economy grows as an points quickly emerged from the symposium integrated whole consisting of agricultural, extractive, manufacturing, and services sectors discussion. We refer to them as “proponents” and that require physical inputs and produce “opponents” of the idea that economic growth con- wastes, and; flicts with fish conservation and that, therefore, a steady state economy (with stabilized population and (9) There is increasing evidence that North American per capita consumption) is more conducive to fish economic growth is having negative effects on conservation than is economic growth. Most of the the long-term ecological and economic welfare of North America and the world. debate centered around a position statement pro-

November 2005 | www.fisheries.org | Fisheries 37 Therefore: approach, as we found more instances of agreement than of disagreement. There was no opposition to precepts 1–3, although pre- (1) There is a fundamental conflict between economic cept 3 led to some elaboration. Although there was no growth and fish conservation based on the ecological disagreement that GDP and GNP were historically entrenched principle of competitive exclusion, and; as indicators of economic growth, proponents and oppo- (2) There is a fundamental conflict between economic nents alike pointed out that economists from a diversity of growth and the ecological services underpinning the philosophical traditions have acknowledged the limitations of human economy (for example, pollination, decomposi- using GDP as an indicator of economic “health” or general tion, climate regulation), and; welfare. Alternative metrics were identified, such as the (3) Technological progress occurs via research and develop- Index of Sustainable Economic Welfare and the Genuine ment that requires funding and the use of natural Progress Indicator, which account for other parameters of resources, has many positive and negative ecological and welfare such as disease rates, infant mortality, and literacy. economic effects, and may not be depended upon to rec- Bhutan’s “Gross Domestic Happiness” was also mentioned. oncile the conflict between economic growth and fish Proponents stressed the fact that, in the vast majority of conservation, and; nations, official national income accounting agencies (such as the U.S. Bureau of Economic Analysis) discount these (4) A steady state economy is a viable, sustainable alternative alternative parameters and indices, concentrating solely on to a growing economy and has become a more appropri- GDP and GNP and conflating economic growth with general ate goal in the larger, wealthier economies of North welfare. America, and; Precept 4 generated the most intense and protracted (5) The long-run sustainability of a steady state economy requires debate. The debate illuminated a major philosophical differ- its establishment at a size that does not breach ecological and ence between conventional or “neoclassical” economics and economic capacity during expected or unexpected supply alternative (e.g., ecological or biophysical) economics. shocks such as droughts and energy shortages. However, despite the tense atmosphere, respect was observed for all viewpoints and equal discussion time was granted for both. Pister also identified four Opponents argued that, in fact, there was no physical “other clauses to consider.” limit to economic growth, because much of economic activ- ity is the result of human ingenuity, creativity, and (1) Because of its negative effects on long-term ecological intelligence, rather than the use of material resources. In and economic welfare, economic growth is an increas- essence, technological progress would always provide substi- ingly dangerous and anachronistic North American goal. tutes for scarce resources and reduce or ameliorate pollution. (2) A steady state economy does not preclude economic They also called attention to an emerging service economy, development, a qualitative process in which different and especially its “information sector,” that would eliminate technologies may be employed and the relative promi- limits to economic growth by dematerializing the economy. nence of economic sectors may evolve. Proponents stressed that the infrastructure associated (3) Upon establishing steady state economies, it would be with service sectors (for example, electricity for computers advisable for North American nations to assist other and universities for research and development) still requires nations in moving from the goal of economic growth to resource inputs and results in waste production. Opponents argued that, as long as the ratio of service sectors (including the goal of a steady state economy, beginning with those information) to natural resource extraction increased, limits nations currently enjoying the highest levels of per capita to growth would be overcome. Proponents then evoked the consumption. ecological principle of trophic levels and claimed that none (4) For many nations with widespread poverty, increasing per of the service sectors, including information, are economi- capita consumption (or, alternatively, more equitable dis- cally relevant in the absence of agricultural, extractive, and tributions of wealth) remains an appropriate goal for the manufacturing sectors to employ the information and other time being, yet the ultimate goal should be the establish- services. Noting that animals (consumers) do not exist with- ment of healthy ecological and social conditions within out plants (producers), they insisted that additional the framework of a steady state economy. agricultural and extractive surplus is required for the growth of manufacturing and service sectors, thereby subjecting the Pister’s final slide announced, “The talks have been given: economy to a size limit determined by the stock of natural let discussion begin!” Discussion began indeed and, before resources or “natural capital.” long, was quite heated. Pister’s show had represented the With all agreeing on the importance of energy to eco- proponent view; opponents rejected the validity of the pro- nomic growth, opponents claimed that the sun provides a posed position statement, with one opponent noting that he virtually limitless source of energy. Proponents responded disagreed with each and every precept. A session moderator that the Earth operates as a system with a fixed rate of solar then projected on-screen each of the “whereas” clauses one energy input, and that energy as well as material is, in fact, a by one so the two groups could identify points of agreement finite resource. They argued, furthermore, that physical laws and disagreement. This turned out to be an illuminating and ecological principles cannot be violated, no matter how

38 Fisheries | www.fisheries.org | vol 30 no 11 November 2005 | www.fisheries.org | Fisheries 39 Continued from page 38. Precept 9 was the other major point of con- tention. Opponents argued that the quality of life intelligent future generations become. The laws has improved immensely, for U.S. and world citi- of thermodynamics hold that matter and energy zens alike, as a result of U.S. economic growth. can neither be created nor destroyed, are con- They pointed to technological developments in served through all transformations, and that health care, disease control, agricultural produc- entropy tends to increase with every transforma- tion, education, and satisfaction of material needs tion. Economic activity, proponents elaborated, and wants. They challenged proponents to state applies energy to natural resources to manufac- how life could be better without economic ture low-entropy goods, but creates high-entropy growth, or why their children should not have an economic growth waste (pollution and heat) in the process. No forum increased standard of living. Without economic physical, biological, or chemical transformation growth, the argument went, we would not have may operate at 100% efficiency, and useful low- the wealth and technology to solve environmental entropy resources are degraded to waste. problems at home and around the world. Has Proponents then asked, if opponents were cor- not, for example, water quality and the amount rect that economic growth could continue of forested land been increasing in the United indefinitely on a planet of fixed size with finite States since the 1970s? resources, would that not be equivalent to main- Proponents agreed that, in many cases, tech- taining a steady state economy on an nology associated with economic growth has ever-shrinking land mass? If so, could we not alleviated much human suffering but has created someday support a trillion dollar economy on the new problems in the process. One proponent was head of a pin? Proponents claimed that economic concerned that her children would have a worse, growth during most of the Industrial Revolution rather than better, existence than her own has been “subsidized” in a sense by solar energy because of environmental degradation, urban fixed over the course of 300 million years (i.e., via sprawl, traffic congestion, wars, and climate the fossilization of fuels), and that fossil fuels, change. The implication was that each of those problems is linked to increasing production and especially petroleum, are becoming scarcer and consumption of goods and services (economic more expensive. growth) and concomitant waste production. One opponent noted that concerns about Proponents also agreed that, at least until very resource scarcity have been expressed since no recently, water quality (as one example of envi- later than Thomas Malthus at the turn of the ronmental health) has improved in the United 19th century, yet we still have been able to main- States, but argued that much of our environmen- tain economic growth. Another opponent tal success has come at the expense of other mentioned the famous bet between Julian Simon countries. Many of the environmental costs asso- and Paul Ehrlich. Ehrlich chose 10 metals and ciated with resource extraction, energy wagered that the prices of each would be higher consumption, and waste production of U.S. eco- in 10 years because of scarcity, whereas Simon, nomic activity are paid by foreign societies, so believing that technology would obviate such while some of our own ecosystems appear to be scarcity, predicted that prices would remain the intact, the damage is out of sight but just as real. same or decrease. Simon won the bet. A propo- Finally, proponents stated that the U.S. economy nent challenged the interpretation of the bet, consumes a disproportionate share of world however, stating that although Simon won per se, resources and produces an equally disproportion- he actually lost in principle because price is not a ate share of world pollution, especially good measure of scarcity for a variety of reasons greenhouse gases, and that much of the resource (e.g., government subsidies). If the energy return extraction in other nations is financed by debt on investment (EROI) had been calculated for with inadequate concern for future generations or foreign citizens. each metal, Simon would have discovered that These were the highlights of the symposium the cheap, abundant metals of the future were discussion, as best we recall. We felt the debates actually of lower grade and required proportion- were healthy and productive, and we hope our ally more energy to extract and process. account is useful to those who missed the sym- On the subject of resource prices and the posium. We conclude that the debates were a Ehrlich-Simon bet, another proponent pointed out classic example of a difference in paradigm, that land, the aggregation of all natural reflecting different backgrounds and areas of resources, has increased in price inexorably for vir- expertise rather than vested interests and tually all of American history. motives. Perhaps it was to be expected that the There were no major disagreements on pre- differences would predominate in the discussion. cepts 5–7, which are simply definitional, nor on What seemed more striking, and most encourag- precept 8, which is standard fare among all ing, was the amount of agreement on most key schools of economic thought. points.

40 Fisheries | www.fisheries.org | vol 30 no 11 Meet a Hutton Scholarship Recipient report HuttonProgram

My name is Michael Freytag and I am 17 years CPR/first aid class with other fish technicians. I got a old. I live in an Inupiaq village in western Alaska CPR card and first aid card, which I think is awe- called Unalakleet. I will be a senior at Frank A. some to have. That trip lasted about three days. The Degnan High School this upcoming school year. For next trip I went with Karen Dunmall and Charlie the summer of 2005, I was accepted as one of the Lean, a fish biologist with the Norton Sound Hutton scholarship recipients. I was paired with two Economic Development Corporation, was to the mentors, Henry Oyoumick, a retired science and Ungalik River, which is located north of Unalakleet. math school teacher, and Karen Dunmall, a fisheries We went there to begin the genetics mapping of biologist working with Kawerak. Kawerak is the chums in Norton Sound. We had to take the axillary regional Native non-profit organization representing processes of the chums and preserve them in vials 20 villages in the Norton Sound area. Henry works at containing ethanol. I thought that was one of the the Native Village of Unalakleet office as their water- coolest things I did so far in my program. We shed coordinator and Karen heads the Fisheries camped at a cabin located at Ungalik River and col- Department for Kawerak in Nome. lected our samples by seining below beaver dams Unalakleet is a coastal town in western Norton and in clear streams. That trip lasted about four Sound in Alaska. The population is approximately days. After that, I went to Nome for habitat assess- 900 year-round residents. Many of the residents ments on the Snake River. I went with two other depend on fisheries for part of their income and oth- people and basically took measurements of the river ers fish for a subsistence lifestyle. Some of the and described what features were on it. That was residents fish commercially for herring, king salmon, for about three days. I really enjoyed working with and crab. We have a fish processing plant located here that generates a lot of jobs for the residents. I would like to thank the American Fisheries Society for accepting me into this program. I was really happy when I found out I was accepted. The first thing I did with my mentors was to have a meeting with them. We discussed what I would be working on through the summer with both of them and talked about my schedule. I started my work here in Unalakleet with Henry. The projects he had planned for me to do here were identifying benthic macro-invertebrates on the rivers and streams near Unalakleet using D-frame nets and preserving a few samples in alcohol-filled vials. He also wanted me to get acquainted with instruments needed to obtain data related to water chemistry. Some of the instruments used were a Hanna combo meter, flow rate meter, a dissolved oxygen indicator, and thermometer. With the macro-invertebrates that I collect, I would then take pictures of them and use them in posters and other projects that children could use to study. Although that was what I was Mentors Henry Oyoumick, a retired science and math school teacher, and Karen Dunmall, proposed to do with Henry for the summer, we a fisheries biologist, with Hutton Scholar Michael Freytag. completed a lot more. We have been working on getting diseased fish sent to a pathologist in Karen on her projects. Then on July 27th, we went Anchorage, Alaska, for analysis. We are planning to seining on the Unalakleet River with Henry for chum Michael Freytag meet with the pathologist in Anchorage to see what genetic samples and that was fun too. procedures and methods are done for the assess- I had a lot of fun this summer. I think this is a Freytag is a senior at ments. Additionally, we are planning to visit a great program for students who are interested in Frank A. Degnan High hatchery located near Elmendorf in Anchorage to the fisheries. In western Alaska, I think a lot of School in Unalakleet, observe the processes within a fish hatchery. students would be interested in doing this if they Alaska. The projects that I worked on with Karen were led in the same direction as I have been. For Note: Funding for travel Dunmall were different than Henry’s projects, which me, this program gave me an extensive prepara- and supplies for Michael made it interesting. Her projects dealt more with tion in what I want to study when I go to college. Freytag was provided by NOAA Cooperative salmon and habitat work. The first thing I did with It gave me a better picture of the fisheries and Agreement NA16FW1272 Karen was to attend a fish technician orientation in how it’s important to villages like Unalakleet and Research and Prevention Nome, Alaska. It was basically going over rules and other communities who depend on fish for subsis- Relative to the 1999 Norton expectations that technicians have to go through tence or commercial use. Again, I would like to Sound Fishery Disaster. For more information about the before they are sent to enumeration camps. We also thank you for accepting me into this program. It Hutton Program see went over boat and gun safety. I also had to take a was a fun summer for me. www.fisheries.org

November 2005 | www.fisheries.org | Fisheries 41