BobertH. Gray, RH G ray & Assoclates,2867 Troon Ct.. Rich and Washington99352 E mail rhgray@ixnetcom com ano

DennisO. Dauble, Pacific Northu/esi Nationa Laboratory P O Box999 Richand. Wash nqton 99352

SomeLife HistoryCharacteristics of Cyprinidsin the HanfordReach, Mid-ColumbiaRiver

Abstract

Cvprinids arc thc non abundanl nonsalmonidfamily of in the Hanford Reach of the Columbia Ri\,er and collectivelv conrprise> 50tZ ofthe communil!. The nonhern pikeminnow (P4chot:heilusoregon,n \irl. ruil!iLle\hiner LRt(rrrrdrorl"r ,ar.drrrr). peamouth(MJ kx:heilustaurinusl.and chisclmouth(Acr.r.l.,i/rr a/aracrrrs)are common. aDdcommon carp (Ctpr.t r.! cd,?i.r. scveral speciesof dacc (Rlilii.lth\: tdtarudae, R. fulcdtur, R. r,.u/rr). rench (nr.a rin..r). and goldfish (Cd,"drrrllr drld!.is) also occur.Cyprinid spawningis concentratedin lale spring and early summerat water tempcraturesranging fron 10 to l6'C. Of thc mosl abuDdantcyprinids. northernpikeminnow, atuiD the largestsizc at maruriry and rcdside shiner rhe smallest. Although allspecics areomDivorousfor at lcasl paft oftheir lifecycle. northernpikeminnow is piscivorousat sizes> 250 mm fbrk leDgth\!hile chisclmc'ulhiDgest mainly pcdph]ton.

Introduction repeatedbecause of reductionsi[ water use de- velopmentandrecent restrictions on samplinggear The Hanfbrd Reach remains unique as the last imposedby environmentalregulations such as the unimpounded,though regulated, sfetch ofthe Co- EndangeredSpecies Act. lumbia River in the United Statesupsteam of BonnevilleDam. The area supports impodant sport This paper describesthe biology of the Cy fisheries for steelhead(a),?.orft) nchus m)-kiss), prinidac. the most abundantand diverse tamily smallmouth bass (Micropterus tblomieu), and, offishes in the Hanford Reach(Gray and Dauble white sturgeon(A c ip e n s e r tansmont dnus), a\d. 1977a).The datawere largely derivedfrom stud containsthe last naturalmain-stem spawning area resthat asscssedpotential impacts ofpower plant fbr fall chinook salmon (O. tshab\tscha, Cray operationson the NLquaticenvironment (Gray and and Dauble 1977a,Dauble and Watson l997). Dauble1976b. 1971b, c,1978,1979b, c, Pageet While therehave been numerous studies ofanadro- al. 197,1,1982). Native cyprinid speciesdiscussed nous fishesin the Hanford Reach(Becker 1973, in detail are the lorthern pikeminnow (P cio- 1990,Dauble et al. 1978.1980, l989. Daubleand chei I us oregone ns is),redside shiner (Rlc/rzrrdsoairu Watson 1997). Iife history information for resi- balteatus), peamotth (My kr heilus caurinus), and, dent nonsalmonidsis limited to smallmouthbass chiselmouth (Acroci eilus ttlumce us).Linited data (Hendersonand Foster 1956,Montgomery et al. arealso presented on dace(RhlnlcAlhys spp.) and 1980), the sandroller (Pertn2sis transnro tana. the non-nativecommon carp (Ctprinus ccLrpio). Gray and Dauble 1976a, 1979a),rwo speciesof Life history aspectspresented include relative sucket(Cotostomus (:olumbiunus, C. nwc rocheilus, abundance,distribution. age and growth. spa$,n- Dauble1980. 1986), and white sturgeon (Haynes ing characteristics,juvenile emergenceand tim- et al. 1978,Haynes and Gray 1981,Crass and ing, habitat use.and diet. Gray 1982).Howevel power-plantsiting and other studiesconducted oyer the past severaldecades StudyAreas/Materials and Methods in the mid-ColumbiaRiver (rcviewedin Becker 1990) provide important intbrmation on other The ColumbiaRiver adjoinsthe U.S. Department abundantresident fishes. These forage tish are of Energy Hanford Site beginning about l0 km important in the proper tunctioning of thc Co- (6 rniles)north ofRichland. Washington.between lumbia River's food web, both as sourcesoffood river kilometers (km) 550 and 629 (river miles and aspredators. It is doubtful that thesetypes of 3,12and 395) (Figure l). The inclusivearea of comprehensivemonitoring studieswill ever be river is called the Hanford Reach.The Hantbrd

122 NorthwestScience, Vol. 75, No. 2.2001

O 2001 bJ_rheNonhq.n Slsntifir A\sociation All rilhrsrescrred i I

Riverkm 576-579 MidriverSite

Riverkm 557€06 DownriverSil€

Richland

Figure L , U- S. Depannenl of Energ,vHanford Reser!ation.and fish sampling locations. reachhas no tributarystreams, althoughthe Snake sampledat various intervalsfrom 1973 through andYakima Rivers enterat river km 530 and 539, 1982. The upriver site was sampled at weekly respectively.Over the lastseveral decades, power- intewals from April 1973to June 1974 and dur- plantsiting studies, evaluations ofreactor discharge ing May throughJune 1975.The down river srte impacts,and other environmental assessments have wassampled at intervalsof aboutthree weeks lrom beenconducted. These efforts have requiredex September1974 through August 1978. The mid- tensivefish samplingprimarily at threeareas (Fig- river sitewas sampledmonthly from March 1981 ure 1). The areas,located at river km 605-613 throughApril 1982. (upriver site), river km 576-579 (mid-river site), Thereare several differences in physicalhabitat and river km 557 to 566 (downriver site), were among the three sites.The upriver site consists

Hanford ReachCyprinids 123 of a straight and deep channel with no islands, composedof two 61 cm (24in) diametertunnels, hastr maximum depthof l0 m, andfew shoreline eachabout 3 m (10 tt) long and setin pairs so the inundationsare presenl The bottom consists l gely mouth of one tunnel facedthe mouth of the sec- of bouldersand cobble >10 cm in diarneter The ond. Each tunnel was supportedby 6 hoopswith rnid-river site includes a main channel and two throatsattached on the first and third hoop. The large backwatcr areas.The botton is lairly uni tunnelswere connectedwith a lead,6l cm (2,1 form consistingof cobblc, a t'cw boulders.some in) high and 6 m (20 ft) long. Mesh size of the gravel,and cemented material. The channelat thc throats,netting surroundingthe hoops and con- downstrcamsite is shallower wider and contains nectinglead. was 1.25cm (0.5in) squaremesh. islands; adjoining it are sloughs and backwater Backpackond boat electoshocker:Backpack areas.The bottom variestiom sandto cobbleand electroshockingwas conducted using a Snith-Root bouldersand is often cementedas outcroppings Type VII Electrofisher At each station,a single ofupper Ringold-typematerial (Gray andDauble 50-metertransect was sampledat depthsless than lq77br. Addil iona ll). thc'downrirer.iteis unin- a meter Also, a Smith-RootType VI Electrofisher terfacearea under the influenceof both the back- powered by a 240-volt gcncrator was mounted water cffcct of the upper end of McNary Dam on a boat and usedto samplenear shore lbr adult and the free-flowing Columbia River Thus, the fish.Asingle 500m transectwas sanrpled at depths downrivel sitehas grcaterhabitat complexity than upto3m. the othersites. Hook and llre: Trotlines were set along the Severaltypes of sampling gear and methods shorelineand in mid-channel.Trotlines consisted were usedto suit variousareas and habitattypes, of a 9 m (30 ft) long, 0.6 cm (l/4 in) diameter' and to compensatefor gear selectivity. Those Iead mainline and 10 altemating No. 2 and 2/0 pedinent to this discussioninclude: hooks.Hooks baitedwith worms or small pieces Gill nets: GlIl nets were set from shore into offlsh were attachedto the mainline at 1 m (3 ft) the river or were setoffshore perpendicular to the intcrvalsby a0.3m (l fi) long,60Jbtest monofila- cuffent in deeperwatcr. Nets were about 2 m (6- ment leaderand swivel. Trotlines were set in the 8 ti) deep and composedof panels of different aftemoon and retrieved thc next morning about merh.it e rungingin 1.25cm (0.5in) increment' l9 h later.Hook and line samplingwith artillcial from 1.25-10cm (0.5-4 in) squaremesh. Nets lurcs was underlakenwhen time perrnitted. variedin lengthfrom 7.5 37.5m (25-125ft). Net Also, some fish specieswere collected from length was dependent on location and the trayeling screensof a water intake structure hydrogeologicalconditions ofthe river at the lime at ColumbiaRiver km 608 (Gmy et al. 1975,Page of the set.Gill nets wcrc sct in the atternoonand et al. 1975,Gray andDauble 1977a), and cylin- retrievedthe next moming about 19 h later. ddcal traps\r,ere set on the river bottom Trummel nets: Trammel nets werc set from in theintale forehry. A fterr olleLtiun. rpecies \\ erc shoreinto the river. Nets were about 2 n (6-8 ft) identified using standardtaxonomic keys (i.e.. deepand composedof 6-m (20-ft) panelsof dif'- Schultz1936. Carl et al. 1967.Eddy 1969,Bond ferentmesh size ranging in 1.25cm (0.5in) in 1973). crementsfrom 2.5-10cm (1-4-in)square mesh. Collectedfish werecounted and catch per unit Trammel nets u'ere 30m (100 ft) Iong, r'ere set cftbrr (CPUE) was determined.Fish CPUE with in thealtemoon and retricred thc rrertmorning gill net,trammel net, hoop net, and beach scine about 19 h later. was calculatedas fbllows: gill and trammel nct Beachseines: Beach seining was doneat sev- CPUE wasthe averagenumber offish caughtper cral locations on both sidesof the river at each ovemightset (about l9 h) of 30 m (100ft) ofnet: studysite. Seining encompassed about 30 m (100 hoop net CPUE was the avcragenumber of fish jrl ft) of rh,'reline er\'hlu\rlion. The .eincuas caughtper 24 h set; beachseine CPUE was the 1.2 m (4 ft) deep,9 m (30 ft) long and consisted averagenumber of fish caught per seine haul: (1/8 of 0.32cm in) mesh. trotline CPUE was the averagenumber of fish Hoop nets: Hoop netswere usually set paral- caughtper ovemight set(about l9 h) of l0 baited Iel to the cunent.checked every l6-18 hrs,and hooks;hookand line CPUE wasthe averagenum rotatedamong various locations. Hoop netswere ber of fish caughtper man hour fished.

12:l Gray and Dauble Fish were measured(fork length IFLI to the However. data were not always dircctly compa- nearestmm). weighed(gonads were removed and rablebecause ofdifferences in samplinggear and $eighedseparately to thenearest0.l g). andsexed. durationof the studiesat eachsite. For example, Reproductivestatus (ripe, gravid, spent).includ- while gitl and trammel nets and trotlines were ing presenceor absenceof secondaLrysex char usedat thc upriverand downriver sites,they were acteristicsand percent gonad to body weight,was not usedat the mid-river site. Electrofishingwas determined.Sorne stomach contents were identi- usedin lateryears at the downriver site and at the fied macroscopically.Additionalty, the upper di- mid-river site, but not at the upriver site. Only gestivetract and storrach of sclectedspecimens hoop nets and beach seineswere used 1br one were removed and placed in 10c/rfonnalin for completeannual cycle at all three sites. subsequentnicroscopic examination. Food organ- ismswere identitled to thelowest taxon possible Resultsand Discussion andthe numberof stomachscontaining each taxon was recorded.Two indiceswere usedto cvaluate RelativeAbundance Visual $'ere diet. estimates made of the fraction Up to 411species of fish representing13 farrilies eachfood item contributcdto the total volume of occurin theColumbia Rivcradjoining the Hanford gut (Hill 1962,Windell contents 1970).Empty Site(Gray and Dauble 1977a, Page et. al. 1982). $'ere stomachs not included in the calculations. Ten speciesrepresent the family in The numberol'stomachs in which a taxon oc- the HanfordReach: northern pikeminnow, redside cuned was also recordedand expressedas a per- shiner, peamouth. chiselmouth, common carp. centageof the total number of stomachsexam- longnose.feopard aLnd speckleddace (R. cataractae, ined (frcqucncy of occurrence). R.falutns, R. oscuLur),tench ( finca rarca)and gold- To determinethe age of flsh, scalesfrom the frsh(Carassitrsuurulru). Exceptfbr leoparddace. left sideabove the lateral linc posteriortothe dorsal tench,and goldfish, all specieswere collectedat fin basewere viewed with a microfiche projcctor all sites.The combinedcatch of cyprinidscom- (DaubJeand Gray 1977)at a magnificationof32X. prisedor er 50i ol'rhetolal lor ull threesire.. Scaleradii and annuli measurenentswere made Nothem pikeminnowand redside shiner were alongthc longitudinalaxis through the scalefo the mostabundant (CPUE andrelative abundance cus.Thc rclationshipbetween fork lengthand scale index)cyprinids at theupriver site while peamouth (Tesch radii was thendctcrmined fbr eachspecies and northempikeminnow were mostabundant at 1970)and used to estimateannual growth of in- the downriver site (Table 1, Gray and Dauble (DeVries dividual fish by back calculation and 1977a).Nothem pikeminnowand peamouth were Fric 1996). the most abundantcyprinids collectedat the Length-weightrelationships ofhsh werebased midriver site, where chiselmouthoccurred less on thelogarithmic equation: In W = ln C + n ln L, frequently(Page et al. l982) thanat the other two whcre W is the $,eight in grams, L is the fork sites.Common carp was collected sporadically lengthin millimetersand C andn areempidcally in small numbersthroughout the Hantbrd Reach. derived constants.To dctcnnine if gonad devel- Thethree dace species were collected infrcquently, opmentinfluenced the length weightrelationship. with speckledand longnosc dace being most com seasonalcondition tactors were determinedfor mon;lcoparddaca was only collectedat theuprivcr malurcpikcminnow and chiselmouth liom thenon andmidriver sites.Tench and goldfish. introduced logarithmic fbrm of the length-weight formula: cyprinids, were not collectedat any of the three C = W/L". whcrc C is the condition tactor,W and sitesalthough single individuals were collected L are the sameas above.and n is the regression in oLhersampling er entr outside the three primiul coefficient detennined from thc lcngth-weight sludyareas (Dauble, pers. observation). relationship(Tesch 1970). Data tbr llsh collected at each site were used L fe History Characteristics (e.9., 10dcvelop lif'e history descriptions diet,age Northernpikeminnow and growth, time and duration of spawningand fry emcrgence,etc.) fbr theHanlbrd Reach. Where Nofihem pikeminnow werecdlected by all sam- possiblc.comparisons were madeamong sites. pling methods,were presentthroughout the year

Hanfbrd ReachCyprinids 125 TABLE L Catch-per-unit-eftbft(CPUE) and relativeabundance index (RAI) for cyprinids caughtby gillnct. hoopnetandbeach seine at river km 605-613 (June 1973 thfough May I97'1) and river km 557-566 (October 1974drough September 1975.adapled irom Gray and Dauble 1977a).

CPUE(river km 605-613) CPUElriver km 557-566) Gill Hoop Beach Gjll Hoop Beach Sp((ies Net Net Seine RAI' Net Nc! Seine RAI r

Northern pikeminnow 2.19 0..18 2.98 2.08 2.13 0.28 0.78 1.06 Redsideshiner 0.21 0.99 1.61 r.96 0.84 o.1'7 0.02 0.5.1 0.,19 0.01 o.22 o.24 1.70 0.01 1.59 1.10 Chisclmourh 0.,19 0.01 0.01 o.1'7 2.23 0.03 0.0r 0.70 Common carp 0.15 0.05 0.01 0.09 0.01 0.01 0.29 0.10

Total cyprinids ,1.16 1.,19 7.91 6.91 0.50 3.38 3.60

Other Non-salmonids 0.95 0.l7 2.82 1.31 5.98 0.44 t.'70 2.51 Salmonids I 0t) 0.01 2.55 1.19 0.93 0.0.1 |lJ.85 6.61 rRAl bascdon averageoI lhree annualCPUE valuesfbr eachspecies at eachcollecting sire.

- GILLNEI .-. aEACHSEINE FL MEANFoRKLENGTH,cm

-t =t r FL= ii -ii.ri-a.r , R - 't0 R .=2.93.7 F1. $.2 t R = t1.r41.1

3

2

1

0 S O N D J F M AM J J A S OND J F MAMJJ A S OND J FMAMJ J AS ONDJ FIIIAIIIJJ A lqt1 1975 t!76 19n i978

Figure 2. Monthly calch-peFunit-etrof(CPUE) andsize (range and meanfor peakcatches) of northcmpileminnow (P4,croclcilrr .)'"Sonsrrir) collecledby gillnel andbeachseiDe iDthe Hanfbrd Reach,Columbia Riverat the do$ nri ver sanpl ing sile (rivermile 557 to 566),September 1974 through Augus! 1978. at all sites.and showeda characteristicdecline in near-shoreHanford Reach in net samplesmay numbersfrom Novemberthrough March (Figure reflect offshore movement, movement to the 2). Northem pikeminnow werethe most frequent McNaryPool after spau ning. or the.pecie. may fish collectedby hook and line. Both juveniJes be relatively inactive during low temperaturesin and adultswere collected in net samplesat depths winter Adult northern pikeminnow in lakes re- to 3 m fiom March throughAugust. About twice portedly move to deeper water from October as many.juvenilesoccurred in back\r,aterare s throughApril (Carl et al. 1967). asin the river at the mid-river samplingsite (Page Spawning of northern pikeminnow in the et al. 1982).Juvenile northem pikeminnow were Hanford Reachoccured from late-Junethrough abundantin near shoreareas in July andAugust early August when water temperaturesranged from (Figure 2). The decline in adult numbersin the 14 to 18.5'C. Young-of-the-year were first

126 Grayand Dauble collectedin shorelineareas during eaLrlyJuly, usu- increasedfrom May though Juneor July (mean ally in denseschools along with redside shiner peakvalue = 7.8) anddeclined through July while andpeamouth. The percentgonad to body weight the rutio ofunspent to spentadults declined from (gonadosomaticindex [GSI]) offemales (n = 59) mid-Junethrough earlyAugust (Figure 3a). Mean 100 100 a) b) * 80 80 9li 60 60 6-< 40 40 i6 20 20 o 0 0

16 '16 .F

100 100 c) d) I 80 O '/i 60 E-< 40 >t 20 o 0

16 x- 16 .5

G

l! 'lr{ nI a ,.,Ii'I r I ri April May June July Aug April May June July Aug

Figure 3. Spawningperiods as indicatedb]' unspentto spcntrados and maturity indices (gonadosomaticindices) of: a) northem pikenrinnow (Pqcho(heilus oregone".rlr), b) redside shinet (Richanlsoniut balteetus). c) peamourh (M)ro.reil&r .a!,'lllllJ). and d) chiselmoulh (AcrocheilLrsaluta.ex.l) collecGd in the Hanford Reachof the Columbia River ar $c downriver sampling sire (rivcr mile 557 lo 566).

HunfordReach Cyprinids 127 GSI of males(n = l7) increasedthrough June or 9.7t 1.2(n = 6. Jan-Mar),9.2 t I .0 (n = 33,Apr eady July (mean peak value = 4.7). Sexual di- Jun),and 9.4 t 0.7 (n = 21. Jul Sep).Those tbr morphismin the tbrm ofspawning colorat;onwas maleswere 9.0 1 1.0(n = I 1.Oct-Dec), 8.3 t 0.5 apparenton ripe malesin July with smalltubercles (n = 4, Jan-Mar),8.9 t 0.9 (n = 38,Apr Jun).and presenton the dorsalportion of the headand body 8.8 t L4 (n = l0, Jul Sep).Seasonal differences of somemales and a dark lateralband prcsenton in conditionfactor due to golad developmentwerc spawningmales. not reflectedin the data. Nonhern pikeminnowspawning takes place Annual growth incrementsof northem pike- from Junethrough July in westemMontana (Hill minnow were >50 mm though ages 3 and de- 1962),Idaho (Jeppson | 957,Beamesderfer 1992), clined after age,1(Table 2). Annual growth rates and the lower Columbia River dninage (Patten declinedalter maturity (ages6 l1). Growth ratcs andRodman 1967). Up-river movements may be of pikeminnow in the Columbia River appearto a precursorto spawning(Beamesderfer 1992, be similar to those describedin ldaho (Jeppson Martinelli and Shively 1997).Other pre spawn- and Platts 1959),but greaterthan thoserepofted 'swarming" "chasing" ing activitiesinclude and in Montana(Hill 1962). (Patten of femalesby males and Rodman 1967) We collected northern pikeminnow up to l5 and arc apparentlytriggered by water tempera- yca$ old, with individuals>330 mm being al- tures (Beamesderferl992). Eggs are gelatinous mostexclusively females which is consistentwith andadhesive and are deposited over gravel or rocks data from the lower Columbia and SnakeRivers (Jeppson alongthe shoreline 1957.Carletal.1967). (Puker et al. 1995).Norrhem pikeminnow report- Hatching occurs in about a week at l5-20'C edly maturein 5 to 6 yeals(Carl et al. 1967,Brown (Jeppson 1957,Carl. et. al. 1967,Bell 1973). 1971,Bell 1973)or 7 years(10) with malespos- The mathematicalrclationship between length sibly maturingearlier than females (Beamesderfer and weight of northernpikeminnow >80 mm FL 1992).ln thelower Colurnbia and Snake Rivers, (n = 238) was describedby the fomula: ln W = t-emalenorthem pikeminnow apparcntly grow -11.6 + 3.0 In length.That tbr male no hern fasterafter age 5.1ive longer. and may havelower pikeminnow > 240 mm FL (n = 9,1)was ln W = mortality mtes than males (Pafker et a1.,1995). -10.42+ 2.82 In length(S,r = I 0.10) and for Our data show t'emalenorthern pikemlnnow in f'cmales>250 mm FL (n = 107) was lnW= -12.2+ the Hanford Rcachrnature in 5 to 6 yearsat about 3.13ln length(S,. t 0.12).Mean conditionfac- 32 cm FL. Males matureabout 1 year carlier at,1 torsfbr femaleswere 9.2 + 0.5(n = 13.Oct-Dec), to 5 yearswhen they are about 25 cm FL. These

TABI-E 2. Mcan back calculaledlbrk lengthsand incrementsof gfowth (sexescombincd) for northcm pikeminnow (P^ ch.7.,{"lrlr r)ngrr?rrrj) collected.rtthe downriler sanpling site.

Back-CalculatedFork Lensths l ml at EachYear ol I-ile AgcCroup Number 1 l0 ll

1 29 50 2 22 98 J t1 58 I 1,1 161 57 110 165 216 5 1.1 55 |2 l1). 211 266 6 l3 5l l2l 182 231 2/0 309 'I tl 88 137 l9l 232 211 3l I 8 99 156 206 250 195 333 365 9 IJ 58 r07 157 206 255 291 331 211 406 tL) 5 50 92 113 r80 225 278 310 36'1 399 '128 .l 50 98 1:19 202 211 186 317 3:L1 3T9 '115 .1.11

Total t11

Grand A\cragc 5.1 106 162 2l I 253 292 32) 165 198 122 .1,11

Incfement ofGrowth 51 52 56 ,19 12 39 30 .13 33 2,t 19

l28 Grayand Daublc agesarc about two yearsearlier lor femalesand Adult northem pikeminnow are known to be one year carlicr for malesthan thosereponed for opportunisticand highly predaciouson smaller nonhcm pikeminnowfrom ldaho'sSt. Joe's River lish includingtheir orln species,other cyprinids (Beamesderfer1992). (Schultz193-5). yellow perch(Hill 1962).srick- (Godtiey (sculpins, Macroscopicexamination of gut contentsfrom leback 1955),bottomlish Carl young 471 northem pikeminnow >25 mm showedthat et al. 1967.Buchanan 1981). and salmon (Monro adult caddisflies.fish. and crayfishwere the most andtrout 1934,Foerster and Ricker 1941. Ricker 1941,Godfrey 1955.Betl 1973.Brown frequentdietary itcms. Microscopic analyses andMoyle 1981.Buchanan et al. l98l,Rieman showedfish (37%).caddisfly larvac (26%) and et al. 1991,Tabor et al. 1993),especially below crayfish(18%) wcrc &e mostliequently occur- hydroelectricdams or nearrelease sites ofhatch- ring food itemsu,hile fish werethe dominantitem ery-rearedsalmonids (Thompson 1959, Thomp- by volume(32%). followed by crayfish(16%). sonand Tufts 1967,Poe et al. 1991,Vigg ct al. Microscopic analyses of 23 northern pike 199l. Mesa 199,1.Collis et al. 1995.Ward et al. minnowjuvcnilcs (FL. 2.0 to 3.4 cn) collected J995,Shively et al. 1996).They may alsofeed in {ugu't rnLlSeplember al thcdr'$ n.trcrm sitc heavily on crayfish and a variety of othcr itcms shou'edthey mainly feed on insectlarvae. All life includingsurface swarming adult ants, stagesof chironomidswere presentwith pupac water boatmen, caddisfly larvae and mayfly occurringin 707. ofthc stonlachsexamined. Less nymphs(Thompson 1959, Carl et al. 1967,Hill imponantitems included caddisfly lar.rae and adult 1962.Roger et al. 1993,Shively et al. 1996. cicadellidsand Collembola. Burmesderfer1992). Young northern pikeminnow Northcrn pikcminnow preyed on juvenile may cvenfced on plankton(Carl et al. 1967). percids,centrarchids, cyprinids, percopsids. salmo- Redside shiner nids.catosk)mids. and cottids,and thcrc was a trendtoward increasedpredation on crayfishf'rom Redsideshiner showed a sinilar declinein catch April through Septemberand incrcasedsurface as did northem pikeminnow from Novcnlber feedingon caddisf'lyadults during the day in late through March (Figure 4). Redside shiner was summer Whetherthis rellects a prelerencefbr or the mostabundant cyprinid at the upriversite, and simply availability of caddisfliesis unknown. In wascommon at the downrivcr site.Young-of-the- Idaho,rnost feeding apparently occurs dudng the year redsideshiners were collectednear shorein day (Bcamcsdcrfer1992). lateJulyandAugust, and occurred in peaknumbeni

- GILLNET .-. BEACHSEINE Ft uem tonx Lercta,cm FL = 10.7 20

6 R = 11,2-18,0

5 5

R = 2.2.3.9

3 3

2 2

1

0 S O N D J F M Ail|J J A S ON D J F MAMJJ AS ON D J Fi'AMJJASON DJ FMAMJ J A 19f4 1975 1976 19n l97l Figure .1. Monthl,v catch?ef-uniceffo|t (CPUE) and size (rrnge and mean for peak catchet of fedside shiner (R,./m,ifirnr,.! /'dlledllA) collected b) gi11net and beach senrein the HaDfbrdRerch. Columbia River at ihe do$nriler sampling site rri\er nile 557ro 566) Seprenber197,1 through,{ugusr 1918.

Hanfbrd ReachCyprinids 129 from late July through Septemberor October. TABLE 3. Mean back-calculaledfork lenglhsand Incrcments Smallerpeaks in catchesofjuvenile fish occurred ol growth (sexes comb;ned) lbr redside shiner (Richardioniut in May and were likely juveniles from the previ- r.rllirdlllJ) collected ar the do\lnri\er sampling site ous year'sspawning. Redside shiners were col- lected in the traveling screenwash of a power Back CalculatedFork Lengths plant intake at the upriver site and we observed (mm) ar Each Year ofLift largc schoolsof adults in front of the traveling Agc Croup Number screensin late July and August 13750 Redside shiner spawning occured from late 22652 84 Junethrough Ju15 al ualertemperJrure{ ranging 3 l8 57 89 r21 from 14.5to 18"C.On one occasionwe found '+ 16 51 92 12.1 l:19 adultsactively spawningin a largeenclosed pool 5 tl 52 E6 113 137 156 otf the main river channelin early July. The GSI 6l52 11 102 l2l t33 l,l3 Total 109 of f'emalesgenerally increasedthrough June or GrandA\,erage 52 8? 120 1.13 r54 earlyJuly (meanpeak value = l0.5,n = ll,range 1,13 lncremen!of crowth 52 35 31 23 ll 11 4.0 to 14.7) while the ralio of unspentto spent adults decreasedfrom early to late July (Figure 3b). The GSI of malespeaked in Juneor early lengthsranging from 12 to 15 cm. Elsewhere, July (mean peak value = = 3.5. n 6, range 2.3 to redsideshine$ reportedlymature in 2 to 3 years 5.4).Spawning coloration wasnoted on both sexes (Brown 1971,Bell 1973)or older(Lindsey and rn early June. The lateral stripe and operculum Northcote 1963).Calculated growrh tbr eachage appearedbright yellow with a reddish tinge in class was greaterthan that reportedby Lindsey mature adults. rurdNorthcote ( I 963) u sing length-fiequency data. Spawning time in other parts of the species We found no individuals over 6 yearsold. mnge repofiedlyvaries with altitudeand geographi- Macroscopicanalyses of55 redsideshiner gut cal location.In westemMontana, spawning takes contentsshowed adult caddisfliesand midgeflies place from April (Weisel through mid-June and were the most frequently occuning items. Mi Newman l95l In Wyoming, ). at higher altitudes, croscopic analysesof l5 guts showed adult the spawning period is delayed and contracted, caddisflies (477o), caddisfly pupae (27a/c)and occuring from late (Simon Junethrough early July larvae(20cl.), and chironomid pupae(20%) were 19,16).ln parts ofBritish Columbia,spawning takes the most common food iterns.Adult caddisflies place fiom early May through late July (Lindsey (38%)and periphyton ( l27o)were the dominant andNofihcote 1963). Elsewhere,includingWash food items by volume and therewas evidenceof ington, spawningoccurs from May throughAu- leedingon residentfish eggs.Redside shiners gust(Carl et al. 1967,Bell 1973).Eggs are dem- ingestedmainly caddisfly adultsand pupaefronr ersaland adhesive and arc depositedover gravel April through Septemberand caddisfly and or rocks(Weisel and Newman 1951)or among midgefly larvae from October through Decem- vegetation(Carl et al. 1967) near shore.Hatch- ber We also microscopicallyexamincd stomach ing at 21 to 23'C occurs 3 to 7 dtrysafrer fefiili- contentsof 15juvenile redside shiner (FL, 2.0to zation(Weisel andNewman 1 951). 3.9cm). Chironomid pupae and pre-emergent adult The mathematicalrelationship betwecntbrk fbrms were the primary fbod items and occurred lengthand weight ofredside shiners >100 mm (n in all of theirstomachs. = I l8) wasdescribed by thetbrmula: lnW= 9.7 Redsideshiners reportedly ingest gammarids. + 2.7In length(S,, = I 0.10).Seasonal condition molluscs,algae andsmallhsh (Weisel ;urd Newman factorswere not determingdbecause of small 1951, Carl et al. I 967),are sometimes cannabalisric samplesizes of matureadults during the late tall on their own eggs and prolarvae (Weisel and and winter. Newman 1951),or appearto be nomelectiveteed Growth was greatestdudng the first year and ers. rngestingmainly aquatic and terrestrial in- back-calculatedlength closely approximatedac- sectsincluding beetles,adult dipteraand dragon- tual length at age I (Tab)e3). Redsideshiners in fly nymphs(Weisel and Newman 1951,Carl et the Hanford Reach matured at ages 3-4 at fork al. 1967).

130 Grayand Dauble Peamauth throughearly July (Schulu 1935).Peamouth eggs areadhesive and deposited over gravelorpebbles Peamouthwere collected in all months except alongthe shoreline(Schultz 1935, Cnrl etal. 1967). Januaryand February.primarily by beachseine (Figure 5). The winter decline in nun'rberswas The mathematicalrelationship between fork consistentat all sites.Relative abundance of adults length and weight of peamouthwas describedby -10.6 increasedin April, peakedin May through July, the fomula: ln W = + 2.8 ln length.That and declinedin August. for males > 200 mm FL (n = 40) was ln W = -8.82+ 2.53ln L (Srx= t 0.15)and that for fe- Peamouthspawn earlier than other cyprinids males> 230mm FL (n = 68) wasln W = 5.01+ in theHanford Relch. Spru ningcoloration was 1.86ln L (S".= t 0.05).Seasonal condition fac- mo\te\ iJenluhen udult numbers began increar- tors tbr peamouthwere not determined ing in April. Spawningcommcnced in mid-May because ofsmall samplesizes during when watertemperatures ranged from 10 to 11"C all but the latespring. and peaked in June when water temperatures Growth ratesof peamouthwere greatest(av- rangedfrom 12 to 15"C.The GSI of f'emales(n = erage incrementfiom 51 to 63 mm) during the 37) increasedfiom late Aprit through mid June first 3 yearsand then declined gradually (Table (mean peak value = 18.5), and declined shnrply 4). Back calculatedgrowth rateswerc similar to thercafietwhile theratio ofunspenttospent adults thosedescribed in Montana(Hill 1962).Peamouth declinedfrom late May through early June(Fig- in the Hanford Reachof the Columbia River mature ure 3c). Ripe males occurredin late May when at 3 to 4 yearsof age when they reach about 25 the GSI peakedat ,1.9,(n = 8) and most males cm FL. Elsewhere,male peamouthmay mature werespent by lateJune. Peamouthjuveniles were earlier than femalesand both sexesmature in 3 first collected near shore in mixed schoolswith to 5 years(Brown 1971,Lux l97l). The maxi- other cyprinids and catostomidsof similar size mum ageofpeamouth we capturedwas estimated in June.Adult peamouthappeared to move off at 7 years. shoreor out of the downriver sampling areaby Macroscopicanalyses ofgut contentsfrom 32 mid summer. peamouth>20 cm FL showed adult caddisflies Spawning in other pans of the speciesrange were the most common tbod item followed by is also reportedto occur in May and June (Hill gastropods,and periphyton.Microscopic analy- 1962,Carl et al. 1967,Bell 1973).However. spawn- sesofgut contentsfrom 3I peamouth(ll4to322 ing in Lake Washingtonmay occur from April mm FL) indicatedperiphyton (77%), snails (,12%),

- GILLNET

ir uen ronxreterx. c.

- FL -?.5 R =.60'eo i'=i:i.u.u [t: l1]*.3

R - 19.&28.3ll ii Fr=za.s j: =273_315 ir=+l R

R - 13,1.213

SON DJ FM AM J J A S OND J F MAIIIJJ AS ONDJ FMAMJ JASONDJ FMAMJ J A t974 1975 1976 19n 1978

Figure 5. Monthly catch pcr unil cflort (CPUE) ard size (range and mean for pcak calches)of peamouthchub lMllrxrlrr:iftrs .drlnrr.rr) collectedb,v gill nct and beachseine in fie Hanford Reach.Columbia Riler at the downriver sampling sitc oiver mile 557 to 566). Seprember197,1through August 1978.

HanfordReech Cl prinids l3l TABLE'1. Mean calcularcdlbrk lengrhsand increnenrs of chironomid pupae(6,1%) and leLrvae(507o) and grorvth (sexcscombined) for peanoulh (Mlld- caddiflylarvae (36%). , n,//r., rr'r,,r,,/.oll(.te,lrr I eonurfl,(r .-nF pling site. Elsewhcre,peamouth feed on aquaticand ter rcstrialinsects, including adult ants.larval dipterans Back-CalculatedFork Lengths and mayfly nymphs.cladocerans, and copepods (mm) at EachYear of Life (Hill 1962,Carl et al. 196?).Larger individuals A.e-Grout Numh(r 2t1561 occasionallyfeed on molluscsbutrarely on smaller I 11 t; fish (Carl et aJ.1967). The speciesis alsothought ) 58 123 to preyon salmoneggs in gravel(Carl et al. 1967). -l 26 60 122 178 .t 1 6l r 2,1 172 220 Chiselmouth 5 t5 7l 121 170211 218 Chiselmouth $'ere collected in all months but '/6 81 136 179218 251 28.1 60 I 1,1 156 199 235 267289 Februaryand March (Figure6). Peakabundance occurredliom May throughNovember or Decem- Toral 83 ber.The specieswas collectedmainly in gill and Crand A\ erage 63 123 1712tl 216 216 289 trammel nets and to a lesserextent by hoop net and beach seine.Two characteristicabundance 63 60 5t 39 33 30 13 peaks gill Growlh in net CPUE were noted in early sum- mer and early falJandmay reflect up and down streamspawning migrations. caddisflylarvae (35%) andrnidgefly pupac ( 167o) Spawningcommenced in lateMay whenmean $ere the most tiecluently occuring items. Per- water temperatureswere about l5oC and peaked iphyton (32ol.)and snails (27%) were the major from early Junc through July $,hen mean water lbod items by volume. Ingestionof adult aquatic temperatureranged from l3 to l8'C. TheCSI of lnsectscoincided with availabilityand was greatest femalesincreased in May and June (mean peak from April through SeptemberIn June 197,+,we value = 7.6) and declined abruptly through lare obser,'edschooling adult peamouth surface I'eeding June while the ratio of unspent to spent adults on adultcaddisflies in the forebayofapowerplant declinedfrom late May thrcughJuly (Figure3d). intake. Microscopic exanination of tbod items Mean GSI of males was highest in May and from l,l juvenilepeamouth (FL, ,+.2to 6.3 cm) Junc(rnean peak value = 3.0).Spawning colora- seinedfiom SeptembertoDecember 1976 showed tion. including dark lateral bandsand a general

.-. BEACHSEINE Fr uent ronx relcrx,"t -"i'u ['i?N.11r,.,..=*.0* i':#:L*.'f \ i':;f,,0 6

?

3

2

I

0 SO N OJ FM AMJ J A S ONDJ F MAMJJ ASOND J FMAMJJASONOJ FMAMJ J A 1974 t975 1976 $n 1978 Figurc 6. Monfhly catch pcr unil eftbfi (CPtjU) and size(range and mean fbr peakcatches) ofchisetn|.ru]l(Acnx.tleih$ dhnaceus) collected by gill ncl and beachseine in the Hrnfbrd Reach.Columbia Rivcr ar the downrivcr sanDlinp sitc (ri!er mile 557 1o566), Septenber 197.1lhrough August 1978.

132 Gray and Dauble reddening offin ray bascswere noted beginning Conmon carp were collected in small nunr- in mid-June.We found no concentrationoljuve bersthrcughout the Hantbrd Reach,but werenot nile chiselrnouthin nearshoreareas, compared to capturedin winter.Peak catches ofcommon carp other cyprinids. although a few were collected generallyoccurred in spring and summer Com- by beachscine in late sLlmmer.Chiselmouth are mon ca4r were observedspawning in backwater' thought to spawnin sl]rallertributary streamsof sloughsin May and Junewhen shorelinevegeta- the Columbia River. Eggs arescattered or depos- tion wasinundated and water temperatures ranged ited in masscsand attachedto stonesor rocks. ftom l3 to 16'C. The GSI offemalespeakcd from Little elseis known concerninglife history ofthe lateApril throughJuly (meanpeak value = 22.,1). speciesinWashington(MoodieandLindsey 1972). The GSI of maleswas usually highest in latcApril = The nathematical relationship between fork (meanpcak value 8.6) and declinedin May and length and weight of chiselmouth> 200 mm FL June.First-year growth of commoncarp appeared (n = 345)was descdbed by the formula:ln W = rapid. Juvenilescollected in Septemberranged - 10.9+ 2.9 ln length(S., = t 0.11). Average con from 6l to 69 mm FL. dition lactor valuesfor fen.taleswere 19.2t 1.4 The mathematicalrelationship betweenfork (n = 50. Oct-Dec),l8.3 I 1.7(n = 5. Jan-Mar). lcngth andweight ofcommon carp > 230 mm FL 19.3t 1.5(n = 28,Apr-Jun). and 17.6t 2.4 (n = (n = 79, bothsexes combined) was described by 29.Jul Sep).Tbose for nraleswcte 18.81 1.9(n the fbrmula:ln W = -8.91+ 2.64ln length(S,* = = 21,OccDcc). 18.812.,1(n= 13,Apr-Jun),and t 0.11).Growth mtes, based on scaleanalyses of I 6.8I 1.,1(n = 12,Jul-Sep). Values for theJuly to common carp 70 to 580 mm FL. appearedmost Septemberinterval $erc lower for both malesand rapid during the first 5 to 6 yearsof age. Maxi- temalessuggesting post spawningcondition. mum age of collected flsh was estimatedat 14 Based on scale analysesof 33 chiselmouth, years(n - 26). Males maturedin 3 to 4 yearsat growth ilppearedmost rapid during the first 3 years about 25 cm FL, while femalesusually maturcd at annualincrements of 60 to 80 mm. Longevity a year later at fork lengths >30 cm. Elsewhere, did not exceed8 years.We estimatedthat both cary arc reportedto matureafter I to 3 yea$ (Brown sexesmature at 4 to 5 yearswhen they are about 1971.Bell 1973),with malesmaturing earlier than 25 cmFL. Chiselmouthin BritishColumbiaspawn females. in 3 to ,1years with malesmatudng slightly ear- Microscopicanalyses ofstomach contents lrom lier thanfemales (Moodie and Lindsey 1972). 19common car? showed periphyton (9,17.), snails Microscopic analysesof 33 chiselmouthgut (41clr),caddisfly larvae (29%) andgmss and seeds contents showed periphyton was the most fre- (297r) $,ercthe most frequently occurring items quently occurring item and nade up the bulk of and indicatedbottom feeding.Periphyton (5137c) naterial ingestedby volume (97%). Caddisfly and adult caddistlies( | 17, ) werethe major food larvae(52clc) and midgefly larvac (21%) were the itemsby volume.Additionally, we obscrvcdcarp nert mostftequentlyoccurring items. By volume. feedingon the bottom in shallownearshorc areas insectlarvae were only presentin traceamounts in spring and feedingheavily on surfacedetritus throughoutthe year.Studics in Bdtish Columbia in pools nearshore throughoul the summer.Else indicatcfiat althougholder chiselmouthingest whcrc. car! are reported to feed on bottom or- large amountsof filamcntous algae.it is not di- ganismsincluding crustaceans, aquatic lan"ae gested(Moodie and Lindsey 1972). Nutrition is andnymphs, worms, snails,algac and other plant derivedprimarily from diatomsand other unicel materials(Carl et al. 1967). lular algaeand srnall invertebratesingested with Three dace species(longnose, leopard, and thefilamcntous algae. speckleddace) were collected at the mid-riversite. primarily by backpackelectroshocker in August Othercyprinlds throughNovember 198L Specklcddacc was the Commoncarp, an introducedspecies (Sterba 1963), most abundanttbllowed by longnoseand leop- and dace (R,4inichrlr-r'rspp.) were collected less ard dace.Dace were collectednear shorc in 1-m frequently than otber cyprinids. Limited life his- (3-ft) deep water with cobble bottom and switt tory data tbr thcsc speciesare discussedbelow. current.

HanfbrdReach Cyprinids l33 We collectedlownurnbers ofjuvenile longnose larvae,while adultsprey upon stonefly,damsellly daceby beachseine fiom May throughOctober. and other insect larvae (Carl et al. 1967).Addi- irndloung-ul-the )crr Jl thedor-lnrirer sitc in tlonally, mayflies,blackflies. fish eggsand algae August.Juvcnile emergcnce in earlyAugust sug- may be eaten(Brown l97l). gestsspawning occuned in July.Elsewhcre, spawn- ing repofiedly takesplace in springor early sum- Summaryand Conclusions mer over gravel in shallow streams:although no nestis built,males guard the spawning area (Carl Tenspecies of cyprinidoccur in dreHanford Reach etal.1q57. Broun l!,71. Bell la7.tr. ofthe ColumbiaRiver: most comnlon are north- Microscopicgut contentanalyses of l9longnose em pikeminnow, redsideshiner, peamouth, and dace(FL,4.0-8.5 cm []0 gutsempty orconrained chiselmouth.Several species of dace,and tench unidentifiableitemsl) showedperiphyton and al- and goldfish also occur Nofihem pikeminnow, gae (14%), zooplankton, Duphnio (33c/c),.dnc) redsideshiner andpeamouth spawn in theHanford nridge fly lawae occurredmost frequently.DaTrlrila Reachtiom late May to early July at warer tem- (33%), periphyton and algae (28olc),and midge peraturesmnging lrom 10 to l6'C. We lbund no fly larvae (1,17c)comprised the bulk of rhe diet evidenceof chisclmouthspawning. Following egg by volume. Other iternsincluded plant material. hatchingand early f'ry development,large num- lepidopterilund caddi\tl) Iarvre.Mi(roscopic gur bersofjuvenile cyprinidsrear together in shal contentanalyses of l l juvenile(FL L8 to 4.2crr) low nearshoreareas of the main channel and longnosedace collectcd fiom May through Oc- sloughsduring late summer and fall. Growth ratcs tober showed chironomid laNae occufled n]ost ofcyprinids arehighest during the first threeyears ftequently (7661,)and comprised 3 | 7r by volume. and decline following maturity. Mosr cyprinids Chironomid pupac occurred in 20% of rhc guts in the Hantbrd reach are omnivorous; northem and corrprised 317oby volumc. Dacerepor.tedly pikeminnowis piscivorousat maturity;while other feedon i[unaturc aquaticinsccts: young eat midge cyprinidsmainly ingest larvae and periphyton.

LiteratureCited Carl, G. C.. W A. Clemcns. and C. C. Lindsey. 1967.The FreshwalerFishesol BririshColunbia. HandbookNo. Beamesderfer.R. C. 1992. Reproduclionand early lile his- 5. BritishColumbia Provincial Museum, Vicroria. B.C. lo^ ol nunhrnl.qll-ul.,h.Pt,, lrth? r -,, q..n.a,^. Collis,K., R. E. Beaty,and B. R. Crain. 1995.Changes in in ldaho s St. Joe Riler. Environmental tsioiogy of catch rate and diet of noahen squardsh associatcd Fishes35:231,2.11. wilh releaseof halche.y-rearcdiuvenilesalmonids in Becker,C. D. 1973.Food and growth paramcrers ofjurenile a Columbia River rcservoir.NorrhAlnerican Joumal of Fisherie! Management chinook salm(rn,Or.orhr.rra' ardllrs.lid. in cen- 15:3,16-357 Crass.D. W.. and R. H. cray. 1982. Snour dimorphism tral Columbia River. l-r.S.Fish and wildliii Ser!ice. in \\'hrtesturgeon (;.,ii(rr(r trun\monk t},e Fishcr]Bullerin l1:387 .l{Xl. lu\) frcm Cohmbia River at Hanford. Washinglon. Fishcries Becker.C. D. 1990.Aquaric Bioenvironnenrat Sludies:The Bulletin80:158 160. HanfordExpedencc I91,1,i1:1. Srudics in Environmenol Daublc. D. D. 19E0.Lile hisrory ol rhe bridgclip sucker in Science:19.Etsc!ier Ne$ Yofk. Nex York. |hecentral Colunbi.r Rive. Transactionsol theAmcri Bell. M. C. 1973. Fisheries t{andbook of Engineering Rc caDFisheries Society 109:92,98. quucmenr\ and Biological Crilcria. Fish-Eng. Res. Daublc.D. D. I 986. I -ile hisroryand ecology of the la+escatc Prog. Corys oi Engineers,\orth Pacific Di!.. Pon- svcker (Cabstomus macp.reil,/s) in rhc Cotumhia land.OR. River.American Midland Naluratist ll6:356 367_ Bond C. E. 1973.Ke!\ to Orelon FreshwarerFishes. Tcch. Dauble.D. D. andR. H. Cray.1977. Microfische rcaderfor B u1l. 5 8. AgriculruralExpcrimenr Station. Orcgon Stare proiecringfi sh lcale irnages.progressive Fish Culturisl Llni!cISil],Cor\allis. 39:75. Uro$n. C. J. D. 1971.Fishe\ of Mont.rnr.Agricullural Ex- Daublc. D. D.. R. H. Cra). R. w. Hanf, and T. iU. posron. perinicntStation. Nlonrana State Uni!c t). Bozeman. 1978. Occurrenceof a u,ild albino chinook salmc'n Montana. lonnfll)n( hus tshdt\)t\.r.r) in rhe Colunbia ltivcr. Norlh\iest Brolvn.L. R.. and P B. Molle. l981.The impaclof squaw Scicrce52:108 109. D:ruble,D. D.. R. H. fish on salnonid populations::r revie$. Nonh Amcri cray. and T. L. Page.1980. Inpoftance ollnsects andzooplanklon in rhedict ()1o,age canJournal of FilhcdesNlanagcment l:10.1-t I L chinook salmon (On.r,/,l.rcrr.r t\hawttschd) rn lhe cenrral Buchamn. D. V. R. lvl. Hooten,and J. R. Nloring. t981. \orlh- Colunbia Rirer. Nodh\\,esrScieDce 5.1:253 258. ern squ.t\rfish( Ph.l,o..lleilrr ortr)r.,rrlr) predation Dauble.l). D. T. L. Pagc and R. W Hant 1989.Spatiat on salnonids in rcclion\ ol thc Willamette Juvenllc di\tribulion of jurcnile salmonids iD the Hanlbrd Rivcr basin. Oregon. Can.rdianJoumal of Fishcries Reach.Colunbia Ri\'cr. U. S. Nationat N{arinc Fish- ardAquatic Sciencesl8:360 16.1. cries Servicc Fishery tsullcrin 87:775 790. 134 Gray andDauble Dauble.D. D.. and D. G. Watson.1997. Slalus of fall chinook Gray,R. H.. andD. D. Dauble.1979b. Fish sludiesnear wNP salnronpofulations in the l1id Columbia River, I 9.18- 1, 2 and 4i Januarythrough Decenber 1977. Pages 1992.No(h Ancrican Joumal ofFisheries Managc 5.1-5.6,1.1r:Cray. R. H.. andT. L. Page.Aquatic Eco mcnl I7:283-300. logical StudiesncarwNP 1.2 and,l, Januarythrcugh Devries. D. R. and R. V Frie. 1996.Deternination of age December 1977. WPPSS Cohmbia River Ecology ard growth.Pages 481-512.In:B. R. Murph]'and D. StudiesVol. 5. Baxelle Pacific Northwest Laborato- W. willis (eds.),Fisheries Techniqucs. AmericanFish- ries, Richland.Washington. eries Society,Bethesda, Nlaryland. Gray.R. H., andD. D. Dauble. 1979c.Fish studiesnear WNP- Eddf. S. E. 1969.The Fresh\LaierFishes.2"d Edilion. Will l, 2 and,l: JanuarythoughAugusl 1978.Pages 5.1- ian C. Bro$n Co.. Dubuque,Iow.t. 5.52. /ll: Gray, R. H., and T. L. Page.Aquatic Eco Focrstcr.R. E., and W E. Ricker 19,11.The effect of reduc logical StudjesnearwNP 1,2 and'1-January through tion ofprcdacious fish on survival ofyoung socke!e August 1978.WPPSS Columbia River EcologySludics salmon at Cultus Lake. Joumal of the FisheriesRe- vol. 6. Battelle Pacific Northwest Laboralorics. searchBoard of Canada5:315-336. Richland.Washington. Godfie). H. 1955.On lhe ecologyof SkeenaRivcr whjtefishes. Cray. R. H., T. L. Page. E. G. Wolf. and M. J. Schneider. Caregonu\ and Prosopiulr. Joumal of the Fishefies 1975. A Study of fish impingementand screenpas ResearchBoard of Canada l2r,l99-5:12. sageat Hanford Ccneraling Project - a progtessre- Cra],.R. H., dD. D. Daublc. 1976a.Nelvdistribulionrecords pon- Baftelle Pacilic No(hwest Laboratories. Richland. andnolcs on life historyand behaviorofthc sandroller washlngton. (Ejgennrannand Eigelxnann). Pe rcop s i s tro n smonknla Halncs. J. M.. and R- H. Gray- 1981.Diel and scasonalmove- Slelis 9:369370. mcnts of \lhite sturgeon(Acipenser transnontunus) R. H.. and D. D. Dauble. 1976b. Synecology of the cray. in themid Columbia riler. Transactionsol theAmeri fish communily ncar Hanford GeneratingProjcct and can Fisheriessocicty 107r275-280. assessmentofplanl operaiions,Pages 5.1 5.56. /r: T. Hayncs,J. M-, R- H. Gray, and J. C. Monlgomcry. 1978.Sea- L. Page.R. H. Gra), and E. G. Wolf. Final Report on sonal movements of while sturgeon, A.?.,ri.rcr. Aquatic Ecologic.rlStudics Conducted at the Hanlbrd transmontanus.in thc nid Columbia river. Fisheries Gereratirg Projcct. I973-197,1,WPPSS Columbia Bulletin79:367 370. Rivcr Ecology StudiesVol. 1. Bailclle Pacific Nofih- Henderson,C., and R. F Foster 1956.Studies ofsmallnrouth west Labora|orics.Richland. Washington. bass(Micrcpterus dolotnieul in thc Columbia lti\er Gray. R. H.. and D. D- Dauble. 1977a.Chccklist and relative nearRichland, WA. Tfdnsactionsof thc Amcncan Fish- abundanceoflish specicsfrom lhe Hanford Reachoi eriesSocicty 36:l l2-127. the Colunbia River Northwest Scierce 51:208 215. Hill. C. W 1962. Observationson the life histories of the Gra], R. H., .tnd D. D. Dauble. 1977b. Synecology of the peamouth (Mr/.,.letfur rcurinus) a d the northern fish connunit! nearWNP 1. 2 and,l and assessmenl squawfish (PO.r.,.,ztl".! r,rer'.)renrir) jn Montana. of surlability ol plant arca for salmonid spaw ng, ProceedingsoI the Montana Academy of Science Pages5.1 5.7l. /r: Graf,R. H., T. L. Page,and E. G. 22:21-11. \\bll Aquatic Ecological Studies Condrcted near Jeppson,P W. 1957.The control of squawfishb) use oi dy wNPl. 2 and :1. Seprember197:l Through Scptem' namite. spot treatment.and reducdon of lakc lcvcls. bcr 1975.wPPsS Columbia Rivcr Ecolog! studies fi'h Culruri'rlq:168 l-1. Vol. 2. Battelle Pacific Norlhwest Laboralories. Propre+:\e W Platts- 1959.Ecology rnd conlrol Richland. Washington. Jeppson.P W.. and S. of the Columbia squawfish in nofihem ldaho lakes. Gra]..R. H., and D. D. Dauble. 1977c.Fish communit! stud FisheriesSociet) 88:197 ies ncar WNP- l, I and:l: Oclobcr 1975through Feb- Transaclionsof theAmerican ruart' 1976.Pages 5.1 5.15. /n: Gray, R. H., and T. L. 201. Page.Aqualic Ecological Studies nearwNP l, 2 and Lindsey. C. C., and T. G. Nolthcote. 1963. Lile hislor)- of with paricu 1, October 1975th-rough Fcbruary 1976.WPPSS Co redsideshinefs Ri.rdldrorir,.! bdl/zatur lumbia Ri!cr Ecolog,vStudiesVol. 3. Brllellc Pacitlc lar referenceto movementsin and out ofSixteen Mile NorthrvestLaboratories. Richland. washington. Ldke streams.Joumal of lhc FisheriesResearch Board Cra,"-.R. H.. and D. D. Dauble. 1978.Fish studie\ neafW)lP- of Canada20:1001 1030. 1.2and,l:Nldrch rhJough l)ecember 1976, Pages 5.1 Martinelli, T. L. and R. P Shi\ely. 1997. Scasonaldistribu- 5.7.1.1n: Cra). R. H.. and T. L. Page.Aquadc Eco !ion. movementsand habilal associadonsofnorlhem -(.enorr.. logical SNdies nearWNP l. 2 and,l. \'Iarch through \qua$lFhir r$o luqerColunrbrJ Rr\(r Decenrber1976, WPPSS Colunbia Rirer Ecology Regulated Rivers: Research and ManagenrentI I :5.13- Sudies Vol. :1.Baitelle Pacilic )ioflhwcs! Laborato- 556. ries. Richlard, $ishington. Mesa, M. C. 1994. Efilcts of multiple acutestressors on tIe Gra], R. H.. and D. D. Dauble. 1979a.Biology of the sand predaloravoidance ability and phlsiology ol iu!enilc rollcr in the central Columbia Rivcr. Transactionsot chinook salmon.Transactions of the American Fish the Aneican FisheriesSociet) 108:646 6,19. eriesSocietv 123r786 793.

Hanford ReachCyprinids 135 MontgoNery,J. C.. D. H Fickieisen, andC. D. Beckel 1980. Schullz, L. P 1936. Keys to the fishes of Washingron.Ore Factorsinluencing smallmouthbassproduction in rhe gon andclosely adjoining regions. Universiry of Wash- Hanlbrd Area, Columbia River. Nofhwest Scicncc ingtonPublications in BioloSy2:103-228. 5,1:2967-305. Shively. R. S.. T. P Poe. and S. T. Sauter t996. Feedinere Moodie.G. E. E., andC. C. Lindsey.1972. Liit historyofa sponseby northcm squawfishto ahatcheryrelease of unique cyprinid lish. the chisclmoul}l 6crc.^heilus juvenilc salmoDidsin the Clearwater River. Idaho. .rftdd.erlr) in Bri!ish Columbia. Syesis5:55 61 Transactionsof rhe American FisheriesSociely Nlunro,J. A. l9l,l. The food of the squawfish.Biological 125:230236. Bo.lrd ofCanada. Pacific ProgressRepori l9:3-.1. Simon, J. R. 1946. \orning Fishes. Bullcrin of the W]'o Page.T. L.. D. D. Dauble.and D. A. Neiuel. 1982.Colun ming cane and Fish Depaflment 4:l-129. bia River aquatic ecological studiesnear the Skagir./ Srerba.C. 1963. Fresh\\'aierfishes of the world, Translated Hanlbrd Nuclear Projccl: linal repon. tsatcllcPacilic and relised by D. W. Tucker Viking kess,Inc., New Northwest Laborabries, Richland.WashingroD. York. Page.T. L., R. H. Gray, and E. G. \\b11e. 1975. Reporr on Tabor It. A.. R. S. Shnelt. and T. P Poe. 1993.Prcdalion on impingement studicsconduded at rhe Hantbrd cen- juvenile salmonidsb) smallmouthbass and nofihem erating Prciect N{ ch and April 1975. Batrelle Pa squar'fishin theColumbia River nearRichland. Wash cilic Nolth\\'est I-abontories.Richland. Washingron_ ington.Nonh AmericanJoumal ofFisheriesManage- Page.t L., E. C \blle. Gray, Ii. H.. and M. J. Schneider ncnl 13:831-838. 197,1.Ecological conparison of ihe Hanford No. I Tesch, F. x'. 1970. Age and growd. In W E. Ricker (ed.), and WNP-2 Silcs L'n the Columbia Ri\,cr Baitelle Fish production in frcsh waters.Blackwcll Scienrific Noft h\\'est.Richland. Washington. Pubiications,Oxford and Edinburgh.Pagcs 93 123. P ker. R. M.. M. P Zimnrerman. and D. L. Whrte. 1995. Thomplon. R. B. 1959.Food of |he squawfishPO.iro.leillr Variability in biologic.rl chrracteriitics of norrhern .)r.?orcrsir (Richardson)iD rhe lowerCotumbia River ,iqualltish in &e lower Columbia and SnakeRivers. U. S. Fishand wildliie Service.Fishcry Bullelin 60: Tmnsactionsof thc American FisherjcsSociet! .13-58. l2,l:3353,16. ThoDpson. R. B. and D. F. Tufh. 1967.Predarion by Dotl] Paltcn.B. G. and D. T. Rodman. 1969.Reproductivc behav- Vardcn and nofihern squawlish on hatchcr] reared iol nf n^nlrrm.qLJ$ tr.h. Pt., h." h ttu,reyn, r,.,i,. sockcye salmon in Lakc Wenarchee.Washingron. Transactionsof theAmerican Fishcries Society 98:108 Transaclionsof the American Fisheries Sociely 96: 1. 121 421 . Poc. T. P, H. C. Hansel. S. Vigg. D. E. Palmcr. and L. A. Vig8. S.. T. P Poe.L. A. Pendergan.aDd H. C. Hanscl. 1991. Pendcrgast.I 991 . Feeding of prcdaceousf-rshes on out Ralesof consumptionofjuvenile salnronidsand al- nigrating juveni lc salnonids in John Day ReseNoir, rcrnate prey fish by nonhern squawfish, walleyes. Colunbia River. Transactionsof the American Fish smallmouih bass. and channel calish in John Day eries Societ] 120:405:120. resenoir. Columbia River.Transacrions of rheAned, Rrcke! $l E. I 9:1I . The consumptionof young sockevesatmon can Fishcies Society 120:421:138. by prcd.rious fish. Journal of the FisheriesResearch W.ud, D. L.. J. H Petersen,and J. Loch. 1995.lndcx o1 pre- Bourdof C:rnada5:291 313. daiion on juvenile salmonidsby nofthem squawlish Ricman. B. E.. R. C. Bcamesderfer,S. Vigg. and T. P Poe. in the lo$cr and middle Colunbia River. and in rhe 1991.Eslimated loss ofJuvcnilc salmonidsro preda lowcrSnatreRirer. Transacdons of theA mcricanFish- don by nodhernsqua$1jsh. walleye\, andsnalinouth erics Society 12,1:32131,1. bassin John Day Reserl'oir.Columbia River Trans $'ciscl,C. F. and H. W. Newman.1951. Breeding habi6. actions of the Anrcrican Fisheries Socicry 120:,1,18- developmcnl and early life hisrory of Ri.lafdr rri?rs 158. 7rar.,.t&.r.anorthwestern minnow' Copeia3i187 19.1. Schullr. L. P 1935. Ihe !pa$ring habits of the chub. \4indell. J. T. 1970. Food analysisand rate of digcslion. 1, (ed.). MrL)cheilus rcririnu.r a fo|age fish of some value. W E. Ricker Fish producrion in fresh $aters. Transactionsof the American FishcriesSocietv Soc. Blackrcll Scientific Pubiicalions,Oxford aDd 65:l11 1.17. Edinburgh.Pages 197-203.

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136 Grayand Dauble