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Racey (2012): Temperature and Maternal Effects on Hatch Rate and Length at Hatch of Hybrid Bass (White Bass × Striped Bass) Larvae, North American Journal of Aquaculture, 74:3, 283-288 To link to this article: http://dx.doi.org/10.1080/15222055.2012.672372 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. 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NorthAmericanJournalofAquaculture74:283–288,2012 (cid:1)C AmericanFisheriesSociety2012 ISSN:1522-2055print/1548-8454online DOI:10.1080/15222055.2012.672372 ARTICLE Temperature and Maternal Effects on Hatch Rate and Length × at Hatch of Hybrid Bass (White Bass Striped Bass) Larvae S.E.Lochmann*andK.J.Goodwin Aquaculture/FisheriesCenter,UniversityofArkansasatPineBluff,1200NorthUniversityDrive, MailSlot4912,PineBluff,Arkansas71601,USA C.L.Racey ArkansasGameandFishCommission,2NaturalResourcesDrive,LittleRock,Arkansas72205,USA 2 1 0 2 r e b m e pt e S Abstract 5 Broodstockselectionisanapproachthatcouldimproveaspectsofhybridbass(whitebassMoronechrysops × 2 4 stripedbassM.saxatilis)production.Weexaminedincubationtemperatureandmaternaleffectsonmodalperiodof 2 3: incubation(h),hatchrate,andlengthathatch.Gametesfromthreetofourwhitebassdamsandonestripedbasssire 2 wereusedtoproducehybridseachweekfor4weeks.Eggsfromeachdamwereincubatedat14,16,18,and20◦C. at Theperiodofincubation,hatchrate,andlengthathatchweredeterminedforlarvaefromeachdam × temperature ] s combination.Therelationshipbetweenincubationperiodandincubationtemperaturewasexponential,andwarmer e ri temperaturesresultedinprogressivelyshorterincubationperiodsandless-variablehatchingduration(i.e.,theperiod e h fromfirsthatchtolasthatch).Theeffectsofincubationtemperatureanddamexplainedapproximatelyequalamounts s Fi ofthevariabilityinlengthathatch.Damweightdidnotexplainasignificantamountofvariabilityinmodalincubation Of period,hatchrate,oraveragelengthathatch.Selectionforincreasesinlengthathatchcouldofferimprovements nt tothehybridbassindustrybyimprovingsurvivalinpondsorbyeliminatingtheneedforrotifers atfirstfeeding e intanks.Thebenefitsofproducinglarvaethatarelargerathatchshouldbecomparedwiththeeffortandcostofa m rt breedingprogramthatselectsforlengthathatch. a p e D [ y b d de The global seafood market is worth more than US$170 × in production or reductions in production costs are crucial to a o 109 annually, but the USA is experiencing a $9 × 109 annual developingthehybridbassindustry. nl w seafoodtradedeficit(FAO2009).AnincreaseinU.S.aquacul- Broodstockselectionisanapproachthatcouldimproveas- o D ture production will be necessary to reverse this trade imbal- pects of hybrid bass production. Garber and Sullivan (2006) ance. The industry producing hybrid bass (white bass Morone outlinedthepotentialbenefitsofselectivebreedingforthehy- chrysops × striped bass M. saxatilis) is an important compo- brid bass industry. Categories of traits that could be selected nent of U.S. aquaculture production. The striped bass and its for improvement include growth, health, morphology, and re- hybrids ranked fifth in finfish production in the most recent production.Thebenefitsoffastgrowthordiseaseresistancein U.S.aquaculturesurveyconductedbytheNationalAgricultural a culture setting are obvious. Larval length at hatch is another Statistics Service (NASS 2006). However, production in 2009 traitthatcouldaffectculture. fell below 4 million kg for the first time since 2000 (Turano Largerlarvaewouldhaveawiderpreysizespectrumatfirst 2010).Onepossiblereasonforthedeclineinproductionisthe feeding, and they would also be better able to avoid predation steady 2.7% increase in production costs experienced by the in earthen ponds (Paradis et al. 1996). Bosworth et al. (1997) hybridbassindustrysince1997(Turano2010).Improvements examinedtraitsofwhitebass,stripedbass,andhybrideggsand *Correspondingauthor:[email protected] ReceivedApril4,2011;acceptedJuly11,2011 PublishedonlineMay24,2012 283 284 LOCHMANNETAL. larvaeina3 × 3dialleliccrossstudy.Stripedbasslarvaewere White bass were examined periodically before the predicted larger athatch thanwhitebass,and hybrids wereintermediate ovulation time. Ripe white bass dams were weighed and then in length at hatch (Bosworth et al. 1997). Furthermore, there strip-spawnedintoindividualplasticcontainers.Theweightof was a significant positive striped bass maternal effect (relative dams ranged from 0.4 to 1.3 kg during the study. Milt from to white bass maternal effects) on length at 12 h posthatch onestripedbassandasmallamountofwaterwereaddedtothe and 7 d posthatch (dph) and on the percentage of larvae that container,andthegameteswerethoroughlymixed.Eggswere consumed brine shrimp Artemia spp. at 7 dph (supporting the placedinMcDonaldhatchingjarsandtreatedwitha150-mg/L link between length at hatch and feeding ability). Lochmann solutionoftannicacidfor10min(HodsonandHayes1989)to etal.(2009)demonstratedamaternaleffectonlengthathatch reduceadhesionandclumping.Afterthetannicacidtreatment, ofhybridbasslarvaebutwereunabletodeterminewhetherthe eggswererinsedwithwellwaterfor30min.Portableaerators effectwasphenotypicorgenotypic.Thereisevidencethatsome wereusedtokeeptheeggsinsuspensionduringtransportationto larvaltraitshaveatleastapartialphenotypicbasis.Larvaefrom theAquaculture/FisheriesCenterattheUniversityofArkansas, striped bass dams less than 4.5 kg had smaller mouths at first PineBluff. feedingthandidlarvaefromlargerdams(Zastrowetal.1989). AttheAquaculture/FisheriesCenter,eggswereincubatedin Nevertheless, these studies suggest length at hatch might be recirculationsystems(MarineBiotech,Boston,Massachusetts). increasedthroughaselectivebreedingprogram. Fourseparaterecirculationsystemseachhadfour63-Lacrylic 2 1 In the Lochmann et al. (2009) study, there was also a aquaria; each aquarium contained a single 6.8-L McDonald 0 r 2 significanteffectoftheincubationperiod(h)onlengthathatch. hatching jar and a larval collector sieve (300-µm mesh). Each e b Larvaeincubatedatcoolertemperatureshatchedlaterandwere recirculationsystemhadanactivatedcarbonfilter,acartridgefil- m e larger at hatch but had smaller yolk and oil volumes at hatch. ter,abiofilter,andanultravioletsterilizer.Waterflowedthrough ept This observation suggests that length at hatch or length at the filters, into a hatching jar, into a larval collector, into an S 5 yolk absorption can be increased through the manipulation of aquarium,andbacktothefilters.Waterinthefourrecirculation 4 2 incubationtemperature.Morganetal.(1981)demonstratedthat systems was maintained at 14, 16, 18, and 20◦C. The num- 3:2 stripedbasslarvaefromeggsincubatedat22–24◦Cwerelonger ber of eggs in four 1-mL samples was determined by manual at 2 at 24 h posthatch than larvae from eggs incubated at cooler counting (mean ± SD = 1,623 ± 31 eggs/mL). Each dam ] temperatures. Conversely, Peterson et al. (1996) found that was represented once at each incubation temperature by plac- s rie striped bass larvae were shorter at initial feeding when reared ing 4,000 eggs/dam (number determined volumetrically) into he at18◦Cthanwhenrearedat14–16◦C.Reducingtheincubation hatchingjarsineachrecirculationsystem. s Fi temperatureandincreasingtheincubationperiodtomanipulate Eggs or larvae were checked every 6 h until hatching was f O lengthathatchcouldnegativelyaffecthatchrate.Wewantedto observedandevery3hthereafter.Whenlarvaewereobserved nt examinetheimportanceofincubationtemperatureforinfluenc- in a larval collector, the incubation period (i.e., the length of e m inghatchrateandlengthathatch.Wealsowantedtocompare time [h] from fertilization to appearance in the collector) was rt a theincubationtemperatureeffectversusthematernaleffecton recordedforthoselarvae.Thecollectorswereemptiedevery3 p De variabilityinlengthathatch.Theobjectiveofthisstudywasto hafterinitiationofhatchinguntilhatchinghadceased.Larvae [ y quantify the variation in hybrid bass hatch rates and length at were removed from the collector, euthanized with tricaine b d hatchamongincubationtemperaturesandamongdams. methanesulfonate (MS-222; FINQUEL, Argent Laboratories, e d Redmond,Washington),andphotographedwithaSpotInsight a o METHODS Color digital camera (Model 3.2.0; Diagnostic Instruments, nl w Whitebasseggsandstripedbassmiltwereusedtoproduce Sterling Heights, Michigan) that was mounted on a dissecting o D hybridsatKeoFishFarm,Keo,Arkansas,inspring2005.White microscopeandinterfacedwithacomputer.Fivetofifteenlarvae bassdamscamefromtheArkansasRiverandwereamixof(1) fromeachdam × temperaturecombinationwerephotographed wildfishthatwerespawned in2004,heldinponds forayear, duringeach 3-hperioduntilapproximately 30–50 larvaefrom and spawned again in 2005 (dams A–F; weeks 1 and 2); and eachdam × temperaturecombinationhadbeenphotographed. (2) wild fish that were caught early in 2005 and held for 1–3 Unphotographed larvae from each 3-h period were preserved months before spawning (dams G–N; weeks 3 and 4). Striped andlaterenumeratedtodeterminethemodalincubationperiod basswerethefarm-raisedprogenyoffishfromLakeOuachita, andhatchrates(i.e.,percentofeggshatching)foreachdamat Arkansas.Captivefishwerefedanadlibitumrationoffathead eachincubationtemperature.Standardlengthandfinfoldwidth minnowPimephalespromelasandgoldenshinersNotemigonus were measured for each larva with Image-Pro Plus software crysoleucas.Gametesfromthreetofourwhitebassdamswere version4.5.1.22(MediaCybernetics,SilverSpring,Maryland). crossedwithgametesfromonestripedbasssireeachweekfor4 Temperaturewasmeasuredwithamercurythermometerev- weekstoproducehybrids.Differentstripedbasssiresandwhite ery6hduringincubation.Dailywatersamplesweretakenfrom bassdamswereusedeachweek. each recirculation system. Dissolved oxygen (DO) was mea- Finalmaturationandovulationwereinducedwithinjections suredwithaYSIModel52oxygenmeter,andpHwasmeasured of human chorionic gonadotropin (Rees and Harrell 1990). withanOrionModel290Ameter.Hardnesswasmeasuredwith TEMPERATUREANDMATERNALEFFECTSONHYBRIDBASSLARVAE 285 Hachreagents(Hach,Loveland,Colorado),andtotalammonia is the fixed effect of temperature j, (TIME × TEMP) is the ij nitrogen(TAN)wasdeterminedbymeansoftheNesslermethod interactionamongtimeandtemperaturegroups,DAM(TIME) ki and a Hach DR/2000 spectrophotometer. We report only the is the random (genetic) effect of dam k nested in week i, and un-ionized ammonia based on the formula for the un-ionized ε is the random residual (Bosworth et al. 1997; Wang et al. ijkl fractionofTAN(Emersonetal.1975). 2006). Wedeterminedwhethermodalincubationperiodandhatch The effects of time, temperature, and the time × temper- ratewererelatedtoincubationtemperature.Weusedafactorial ature interaction were tested against the mean square of the analysis of variance (ANOVA; GLM procedure in the Statis- DAM(TIME) term(Zar1999).Toexaminetheeffectofdam ki tical Analysis System [SAS]; SAS Institute 1990) to analyze weightonlengthathatch,wedeterminedtheaveragelengthat the hatch data. Time (i.e., week of the study) and temperature hatchoflarvaefromeachdam × temperaturecombination.We were fixed effects, and dam weight was a covariate (i.e., ran- usedafactorialANOVA(GLMprocedureinSAS)inwhichtime dom effect). Modal incubation period and hatch rate were the and temperature were fixed effects, dam weight was a covari- response variables. The effects of time and temperature were ate, and average length at hatch was the response variable. As tested against the mean square of the time × dam weight before,theeffectsoftimeandtemperatureweretestedagainst interaction term and temperature × dam weight interaction themeansquareofthetime × damweightinteractiontermand term,respectively(Zar1999).Hatchrateswerearcsine–square- temperature × dam weight interaction term, respectively. An 2 1 root transformed to normalize and homogenize the residuals. αlevelof0.05wasusedforallstatisticaltests. 0 r 2 The relationship between incubation temperature and incuba- e b tionperiodforallhatchedlarvaewasexaminedwithlinearand m e exponential least-squares regression models (REG and NLIN RESULTS ept proceduresinSAS). Water quality was similar between temperature treatments S 5 Weexaminedtheeffectoftemperatureanddamonlengthat (i.e., recirculation systems) and between weeks of the study. 2 4 hatch.Length-at-hatchdatawereexaminedwithamixed-effects MeanDOwasconsistentlyatorabove7.6mg/L(Table1).The 2 3: nested ANOVA (MIXED procedure in SAS). The following pHofeachtreatmentaveraged7.2–7.5duringthestudy.Water at 2 modelwasusedtoanalyzelengthathatch: hardness averaged 188 mg/L or higher in each treatment dur- ] ing each week. The un-ionized portion of TAN averaged less s rie Yijkl =µ+TIMEi +TEMPj +(TIME×TEMP)ij than0.008mg/Lthroughoutthestudy.Themaximumobserved e sh +DAM(TIME)ki +εijkl, un-ionized fraction of TAN was 0.02 mg/L. Mean tempera- Fi tures were always within a few tenths of a degree of target f O whereY isanindividualobservationoflengthathatch,µis temperatures and varied little within a treatment during any ent the overiajklll mean, TIMEi is the fixed effect of week i, TEMPj week. m rt pa TABLE 1. Average(±SD)waterqualitymeasuresindifferenttreatments(i.e.,recirculationsystems)duringeachweekofthestudytodeterminetemperature De andmaternaleffectsontheincubationperiod,hatchrate,andlengthathatchofhybridbass(femalewhitebass × malestripedbass). [ by Treatment(nominal Dissolved ed Week temperature,◦C) oxygen(mg/L) pH Waterhardness(mg/L) Actualtemperature(◦C) d a nlo 1 14 8.0(0.7) 7.4(0.1) 202(131) 14.3(0.5) w 16 8.0(0.6) 7.4(0.3) 197(126) 16.1(0.3) o D 18 7.9(0.4) 7.4(0.1) 194(124) 18.1(0.2) 20 7.7(0.4) 7.5(0.1) 188(126) 19.9(0.2) 2 14 8.8(0.3) 7.4(0.2) 265(13) 14.3(0.5) 16 8.7(0.4) 7.4(0.1) 295(18) 15.7(0.3) 18 8.2(0.4) 7.4(0.1) 261(15) 18.2(0.3) 20 7.6(0.4) 7.4(0.1) 269(8) 20.2(0.3) 3 14 8.9(0.6) 7.4(0.1) 264(9) 14.3(0.4) 16 8.6(0.5) 7.3(0.2) 271(7) 15.9(0.4) 18 8.1(0.5) 7.5(0.1) 261(8) 17.8(0.3) 20 7.7(0.5) 7.4(0.1) 269(8) 20.2(0.2) 4 14 9.3(0.3) 7.3(0.2) 286(12) 13.9(0.3) 16 8.8(0.5) 7.2(0.2) 289(5) 15.9(0.2) 18 8.6(0.5) 7.4(0.2) 277(11) 17.8(0.2) 20 8.3(0.5) 7.4(0.2) 282(9) 20.1(0.2) 286 LOCHMANNETAL. DISCUSSION The relationship between incubation temperature and incu- bationperiodforhybridbasswasdifferentfromtherelationship reported for striped bass. Rees and Harrell (1990) presented a linear relationship between incubation period and incubation temperature (Figure 1). The exponential model provided the best fit to our data. At a low temperature (∼14◦C), the Rees and Harrell (1990) model would underestimate the period of incubation for hybrid bass by almost 24 h. At temperatures of 19–20◦C,theReesandHarrell(1990)modelslightlyoveresti- mates the incubation period. When spawning and hatching of hybridbassareextendedbeyondthetypicalspawningseasonor FIGURE1. Relationshipbetweenincubationtemperatureandmodalincuba- ifincubationoccursatwarmerorcoolertemperaturesthanthose tionperiod(h)ofhybridbass(femalewhitebass × malestripedbass;solid typifyingthespawningseason,therelationshippresentedhere line).Theformularepresentsthemodelthatprovidedthebestfittothedata islikelytoprovideabetterestimateoftheincubationperiod. fromthisstudy.ThedashedlinerepresentstherelationshipreportedbyRees Incubation temperature did not affect hatch rate of hybrid 2 andHarrell(1990). 1 bassacrosstherangetestedinthisstudy.Theresultsofourstudy 0 2 r were similar to those reported by Geist et al. (2006) for Chi- e b nook salmon Oncorhynchus tshawytscha. In that study, initial m e The average hatching duration (period from first hatch to incubationtemperaturesrangedfrom13.0◦Cto16.5◦C.Lower ept lasthatch)rangedfrom19.1 ± 7.2h(mean ± SD)at20◦Cto initial incubation temperature increased the incubation period S 5 58.1 ± 7.5 h at 14◦C. Modal incubation period ranged from but did not influence the hatch rate of Chinook salmon (Geist 2 4 41 to 106 h and averaged 65.5 ± 21.2 h over the course of et al. 2006). Conversely, incubation temperature did affect the 2 3: thestudy.Modalincubation periodvaried significantlyamong hatchratesofstripedbass(Morganetal.1981)andwhiteperch ] at 2 tdeammpewraetiugrhetse(fFfe=ct9w.8e6r,ednfo=t s3i,gPni=fic0a.n0t4.6T).hTehmeotidmeel eefxfpelcatinanedd Mpeorarotunreearmanegreicsa(n1a2–(M28o◦rCgaannadn1d0R–2as4i◦nC1,9r8es2p)eacctrivoesslyw).idTehetemre-- s e ri 98% of the variability in modal incubation period (i.e., ex- lationshipswerequadratic,andhatchratewasoptimalatabout he plainedvariability=1−sumofsquares[SS] /SS ).The 18◦Cforstripedbassand14.1◦Cforwhiteperch.Awidertem- s error model Fi majority of the explained variability was due to temperature perature range in our study would probably have resulted in Of (SS /SS = 0.90). An exponential model (Figure 1) reduced hatch rates outside of the typical incubation tempera- nt besttemfipt themordeellationship between incubation temperature and turerange.Thereappearedtobenophenotypiceffectofwhite e m incubationperiod(i.e.,hadthehighestr2value).Asincubation bassdamweightonthehatchrateofhybrids.Ourobservations rt pa temperatureincreased,theincubationperioddecreasedandthe aresomewhatsimilartothoseofSecoretal.(1992),whofound e D variabilityintheincubationperiodalsodecreased.Hatchrates noeffectofdamweightoneggsizeorembryosurvivalinstriped [ y ranged from 10.5% to 64.2% during the study. The effects of bass. b d time, temperature, and dam weight were not significant in the Incubation temperature affected length at hatch of hybrid e d factorialANOVAthatexaminedvariabilityinhatchrate. bass.Asignificantinfluenceofincubationtemperatureonlength a nlo Individual standard length at hatch ranged from 2.42 to athatchwaspreviouslyreportedforstripedbass(Morganetal. w 3.34 mm. Length at hatch varied among times (F = 6.59, 1981)andwhiteperch(MorganandRasin1982).Theseresults o D df = 3, P = 0.010) and among temperatures (F = 3.79, aredissimilartothoseofGeistetal.(2006),whoreportednoef- df = 3, P = 0.047). Length at hatch was greatest at the fectofinitialtemperatureonlengthathatchofChinooksalmon. lowest incubation temperature. The time × temperature Lochmannetal.(2009)observedlongerincubationperiodsand interaction was not significant; however, the random effect largerlarvaeathatchwhenhybridbasswereincubatedatcooler of DAM(TIME) was significant (F = 24.00, df = 10, P < temperatures. Morgan et al. (1981) reported the opposite ef- ki 0.001). The model explained 33% of the variability in length fect of incubation temperature on striped bass; in their study, at hatch (1 − SS /SS = 0.33). Approximately equal stripedbasslarvaethatwereincubatedatwarmertemperatures error model amountsofthemodelvariabilitywereexplainedbytemperature exhibitedlongerlengthsat24hposthatch.Formoronids,incu- (SS /SS =0.20)andDAM(TIME) (SS /SS bationtemperatureappearstobecriticalfordetermininglength temp model ki dam[time] model =0.18).Least-squaresmeansforlengthathatchvariedamong athatch.Therefore,experimentstoexaminetheefficacyofmax- damsbyasmuchas9.2%.Averagelengthathatchamongdam imizinglengthathatchthroughabroodstockselectionprogram × temperaturecombinationsrangedfrom2.80 ± 0.07to3.18 shouldpayparticularattentiontoconsistencyoftheincubation ± 0.07 mm (mean ± SD). The effects of time, temperature, environment.Otherwise,theinfluenceofdifferencesinincuba- and dam weight were not significant in the factorial ANOVA tiontemperaturemightconfoundtheinterpretationofmaternal examiningvariabilityinaveragelengthathatch. effectexperiments. TEMPERATUREANDMATERNALEFFECTSONHYBRIDBASSLARVAE 287 Hybrid bass eggs are susceptible to saprolegniasis (fungal differences in sire, differences in spawning period, and differ- infection caused by Saprolegnia spp.) and low DO levels dur- ences in duration of captivity for dams. A unique striped bass ingincubation.Longerincubationtimessubjectthedeveloping sire was spawned in each of the 4 weeks of the study. Hence, embryos to greater risk. Nevertheless, under the conditions of differences in sire could help explain our observations. Wang our study, increasing the length at hatch by reducing incuba- et al. (2006) reported a significant striped bass sire effect on tion temperature was not detrimental to hatch rate. Producers length of juvenile hybrid bass. Bosworth et al. (1997) did not couldmaximizelengthathatchbyreducingincubationtemper- explicitlymentionthesignificanceoftheeffectofmalegenetic ature without reducing embryo survival. However, Lochmann group or individual male nested in another effect. Paternal ef- etal.(2009)reportedthatyolkandoilglobulevolumesathatch fectinfluencedgrowthratebutnothatchrateorlengthathatch were smaller in larvae with longer incubation times. Reduced of winter flounder Pseudopleuronectes americanus (Fraboulet yolkvolumesathatchwouldprobablydecreasethetimetoyolk etal.2009).Inaddition,timeinourstudyrepresentedspawns absorption, may result in little change in larval length at yolk that occurred at different periods during the spawning season absorption,andcouldinfluencethepointatwhichhybridbass under potentially different conditions and also represented the havefunctionalmouths.Thetransitiontoexogenousfeedingis useofwhitebassdamsthatwereheldincaptivityfordifferent acriticalperiodinlarvaldevelopment.Hybridbasslarvaeheld lengthsoftimebeforespawning.Wehesitatetospeculateabout at 18◦C are stocked into ponds or offered live feed in tanks at thesignificanteffectoftimesinceitwasnotouroriginalintent 2 1 4–5dph,coincidentwithyolkabsorption.Decreasingthedura- toexaminedifferencesamongsiresoramongperiodsduringthe 0 r 2 tionoftheyolksacstagealsodecreasesthetimeavailableforthe spawningseason.Asireeffectonlengthathatchwouldbecome e b developmentofexogenousfeedingbehavior.Thesepossibilities apparent during a diallelic cross study, and we have already m e shouldbeexploredbeforethetemperature(andthusincubation emphasizedtheimportanceofsuchastudy. ept period)ismanipulatedtoalterlengthathatch. Futureworkexaminingdamandsireeffectsandheritability S 5 Generally, length at hatch of hybrid bass from our study ofthelength-at-hatchtraitiscritical.Ourresultssuggestthata 2 4 was similar to results reported by Bosworth et al. (1997) and modest measure of variability in length at hatch is influenced 2 3: Lochmannetal.(2009).TheeffectofDAM(TIME)ki wassig- byamaternaleffect.Selectionforrapidgrowthordiseaseresis- at 2 nificant,indicatingamaternaleffectonlengthathatch.Least- tance would undoubtedly benefit the hybrid bass industry. We ] squaresmeansinthisstudyvariedslightlymorethanthe6.5% suggestthatselectionforimprovementinlengthathatchcould s rie variation in mean length at hatch reported by Lochmann et al. also offer improvements to the industry. Larvae that are larger e h (2009).Highvariabilityinmeanlengthathatchamongdamsin- at hatch may survive better in ponds or may be able to feed s Fi creasesthelikelihoodthatabroodstockselectionprogramcould onlargerpreyitemsatfirstfeeding;thus,itmaybepossibleto f O improvethistrait.Bosworthetal.(1997)foundasignificantma- producehybridbassthatdonotrequirerotifersatfirstfeeding. nt ternaleffectamonggeneticgroups(stripedbass,whitebass,and Eliminating the need for rotifers at first feeding would make e m hybrids).Theyreportedfavorablestripedbassmaternaleffects tankproductionmoreeconomicalandcouldleadtoyear-round rt a on length at 12 h posthatch and on ability to ingest live feed. productionoffingerlings.Thebenefitsofproducinglarvaethat p De However,Bosworthetal.(1997)didnotreportwhethertheran- arelargerathatchmustbecomparedwiththeeffortandcostof y [ dom effect of individual dam nested within the time × dam abreedingprogramthatselectsforthistrait. b d genetic group interaction was significant. Wang et al. (2006) e d did report a significant maternal effect of white bass dams on a o lengthofjuvenile(<20g)hybridbass. ACKNOWLEDGMENTS nl w Thematernaleffectonlengthathatchobservedinourstudy We thank Adam Fuller, Anita Kelly, Alf Haukenes, and o D could be phenotypic, genotypic, or both. Dam weight did not two anonymous reviewers for their useful comments on the influenceaveragelengthathatch,whichsuggeststhatthemater- manuscript. We are grateful to Mike Freeze and Mike Clark naleffectonlengthathatchmightbegenotypic.Iftheobserved (KeoFishFarm)forprovidingeggsforthestudy.Thisproject differences in length at hatch were due to phenotypic mater- wassupportedbyfundingfromtheEvans–AllenProgramofthe nal effects, then selection would not lead to improvements in U.S.DepartmentofAgriculture. lengthathatch.Acontrolledstudythatemploysadiallelicde- signsimilartothatusedbyWangetal.(2006)wouldprovidethe REFERENCES necessarydataforcalculatingtheheritabilityoflengthathatch. 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Bartley a , Eric J. Wagner a & Randall W. Oplinger a a Utah Division of Wildlife Resources, Fisheries Experiment Station, 1465 West 200 North, Logan, Utah, 84321, USA Version of record first published: 05 Jun 2012. To cite this article: Matthew S. Bartley, Eric J. Wagner & Randall W. Oplinger (2012): Conservation Aquaculture of Northern Leatherside Chub and Effects of Temperature on Egg Survival, North American Journal of Aquaculture, 74:3, 289-296 To link to this article: http://dx.doi.org/10.1080/15222055.2012.686007 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. 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NorthAmericanJournalofAquaculture74:289–296,2012 (cid:1)C AmericanFisheriesSociety2012 ISSN:1522-2055print/1548-8454online DOI:10.1080/15222055.2012.686007 ARTICLE Conservation Aquaculture of Northern Leatherside Chub and Effects of Temperature on Egg Survival MatthewS.Bartley,EricJ.Wagner,*andRandallW.Oplinger UtahDivisionofWildlifeResources,FisheriesExperimentStation,1465West200North,Logan, Utah84321,USA 2 1 0 2 Abstract r e Wepresent4yearsofdatathatrefineaquacultureprotocolsforthenorthernleathersidechubLepidomedacopei, b m aspeciesofconservationconcernintheIntermountainWest.Experimentsexaminedlifehistorytraits(ageatfirst e pt spawningandthermallimitstoegghatchingsuccess)andaquaculturetechniques(brooddensity,spawningsubstrate e typeandsurfacearea,andfeedingmethodsforfry).Testsshowedthatleathersidechubcanreproduceasearlyas S 5 age2.Multiplespawnsperfemaleduringayearwerealsodocumented.Survivalofeggswascomparedatincubation 6 2 temperaturesof18.4,23.0,24.6,and26.8◦C.Eggsat18.4◦Chadthehighestsurvivaltohatching(54.0%);eggsat26.8◦C 2 had significantly lower survival (1.5%). Egg survival at 23.0◦C and 24.6◦C (32–33%) was significantly lower than 23: survivalat18.4◦C.Aquacultureexperimentsshowedthatthemeantotalnumberofeggsproduceddidnotsignificantly at differbetweenbrooddensitiesof8.4(1,246 ± 1,236eggs[mean ± SD])or16.8(2,224 ± 1,600eggs)fish/m3.Studies s] showedthatleathersidechubpreferredspawningovernaturalcobblesubstratetospawningovermarblesubstrate. rie More eggs were recovered from a three-substrate tray treatment (1,350 cm2) than from a single tray treatment he (450cm2).FrygivenbrineshrimpArtemiaspp.withprobioticbacteriaorfedwithanautomated,morecontinuous s Fi drip feeder did not show any advantages in growth over time. Juveniles at rearing densities of 800, 1,700, and f 3,400fish/m3didnotdiffersignificantlyingrowthrates,deformities,ormortalities.Thisresearchprovidesgeneral O nt guidelinesforrearingnorthernleathersidechubandsomeadditionalinformationonthespecies’lifehistory. e m rt a p e D [ y Therecoveryofthreatenedorendangeredspecieshasbeen andLentsch1998;Johnsonetal.2004).Recentpopulationsur- b d and continues to be an important aspect of fisheries manage- veys have found reductions in abundance and distribution of e d ment. Threats to these species include dewatering, increasing northern leatherside chub, resulting in part from water with- a nlo averageseasonaltemperatures,nonnativefish,habitatdegrada- drawalsandthepresenceofpredatorybrowntrout(Walseretal. w tion, and a host of other issues (Brouder and Scheurer 2007; 1999;WilsonandBelk2001;BelkandJohnson2007).Northern o D Walseretal.1999).Oneofthebarrierstomanagementisalack leathersidechubarecurrentlyconsidereda“speciesofconcern” ofknowledgeaboutbasiclifehistorytraits.Alackofaquacul- inUtah. turetechniquesforrearingrarespeciescanalsobeabarrierto In an effort to preclude listing this species as threatened or species recovery. An understanding of basic life history traits endangered,aninteragencyrecoveryteamhasidentifiedhatch- andmethodsfortheaquacultureofaspeciescanaidinconser- erysupplementationaspartofarecoveryplan.However,basic vation efforts for imperiled species (Rakes et al. 1999; Sarkar lifehistoryinformationonnorthernleathersidechubonwhich etal.2006). propagation protocols can be based is limited, though recent NorthernleathersidechubLepidomedacopei(formerlyGila effortshaveprovidedsomedata(Johnsonetal.1995;Siglerand copeiorSnyderichthyscopei)isasmallcyprinidspeciesfound Sigler1996;WilsonandBelk1996,2001;Billmanetal.2008a, in the Bonneville Basin and the upper Snake River drainage 2008b).Inthisstudywehadtwoobjectives:(1)toprovidead- of the western United States (Sigler and Sigler 1987; Wilson ditional life history data on northern leatherside chub such as *Correspondingauthor:[email protected] ReceivedJanuary31,2011;acceptedSeptember12,2011 PublishedonlineJune5,2012 289 290 BARTLEYETAL. ageatfirstspawningandupperthermallimitsforsuccessfulegg mentswereused,with44–45eggsperreplicate.Theharvested incubation, and (2) to provide better protocols for aquaculture eggs were treated with 60 mg/L copper sulfate solution for 2 of the species. The aquaculture tests explored effects of brood minat18.8◦C,andrinsedthreetimesbeforebeingplacedinthe density, fry density, spawning substrate area and composition, incubation chamber. The incubation chambers were made by automateddripfeeders,andprobioticfeedsforfry. cuttinga32-mm-diameterPVCpipeintoa5–cm-longsection. Thesectionwascappedwithmosquitomeshnettingsecuredby rubber bands. This chamber was then placed (without temper- METHODS ingoracclimation)inaplastictub(11L)suppliedcontinuously The following studies were performed at the Fisheries Ex- withwater(about1.6L/min)atthedesiredtemperature.Toma- perimentStation,Logan,Utah.Thesetestswereconductedover nipulatewatertemperature,weusedimmersionheaterstoheat 4 years (2006, 2007, 2009, and 2010). Adult leatherside chub waterinaseparatetank.Waterfromthistankwasmixedwith were collected from Deadman Creek (October 2004) and Yel- nonheatedwaterfromthesamesource,andvariouswatermix- lowCreek(August2005),tributariestoMillCreekintheupper ingratios(warm/cold)wereusedtomanipulatetemperaturesin sectionoftheBearRiverdrainage,SummitCounty,Utah(2007 theplastictubs.Forthefirst4daysofincubation,theeggswere studies).FishwerecollectedagaininJune2008(YellowCreek disinfectedbyplacingthewholechamberinto60mg/Lcopper only)forthe2009and2010studies. sulfatesolutionfor2min.Thedeadeggsandhatchedfryinthe 2 1 All the spawning occurred indoors in fiberglass tanks chamberwerecountedafter7d,whenhatchingwascompleted. 0 r 2 (0.9m × 1.8m × 33cmofwaterdepth)andreceived1.7–2.0 Age at first observed spawning.—In 2006, 36 northern e b L/min of hatchery well water. During winter (before the start leatherside chub were placed into a single fiberglass trough m e ofthespawningseason),fishweremaintainedattemperatures (244 × 61 × 30 cm) to determine the earliest age in which ept between 16.5 and 16.8◦C. At the beginning of each spawn- leatherside chub can spawn. At this time the fish were 1 year S 5 ing season (in late March or April), warmer well water was old, derived from hatchery spawning events the previous year. 6 2 addedtoincreasetemperaturesto18.8◦C.Thistemperaturewas Thistankwassuppliedwithtwotraysofsubstrate,smallcobble 2 3: maintained throughout each spawning season (March through (see above) or medium cobble (3.5 cm diameter) mixed with ] at 2 Saecputrermenbte,rt)h.eAou0t.2p5u-t ohfpwsuhbicmhewrsaibsledipreucmtepdwthaosuugshedathoocrirzeoante- ypeolrltoiowngolfasassbmeaaldlesr(1tr.o8ucgmhd(1ia2m2e×ter3).5T×he1ta8nckma)lssoucspoenntadiendedona s rie tal polyvinylchloride (PVC) pipe with perforations to create a aPVCframeabovethebottomofthetroughtocreatebothcover e h laminarflow.Arangeofvelocitieswascreatedineachtankby and a riffle area where the small cobble tray was located (see s Fi puttingasheetofplasticinthecenterofthetank,perpendicular Billmanetal.2008aforfurtherdescription).Thesubstratewas f O tothefloor,butatanangletothewalls(seeBillmanetal.2008a checked twice a week between April and September by using nt for full description). The resulting velocities provided higher theprocedurementionedabove.Thistankwasalsomonitored e m flow areas for the spawning substrate and occasional use and in2007(whenfishwereage2),usingthesameprotocol. rt a exercisebythefluvialleathersidechubaswellasrestingareas Broodstockdensity.—Thisstudytookplaceduringthe2007 p De oflowervelocity.Automatedfeeders(Eheim“Feed-Air”digital and 2009 seasons. Egg production (total eggs per season and [ y automatic feeder and Rena LG100) were programmed to feed numberofspawningevents)wascomparedbetweentwobrood- d b Tetraminflakes4timesperday,at3%oftotalbodyweightper stockdensitytreatments:either4or5(lowdensity;8.4/m3)or de day (about 13 g per week). Phillips full-spectrum lights were 9or10spawners(highdensity,16.8/m3)pertank.Broodstock a o setontimerstomatchtheoutdoorphotoperiod. densitytreatmentswerelimitedbytheavailabilityofbothadult nl w Spawningsubstrates(mediumcobbleormarbles,depending fishandtankspace.Thelow-andhigh-densitytreatmentswere o D ontheexperiment)weremadeavailabletothefish.Whensub- replicatedbyusingtwotankspertreatmentinbothseasons,pro- stratewasscreenedforeggs,itwasdonebyremovingaplastic vidingfourreplicatespertreatmentforthestatisticalanalysis. tray(15 × 30 × 3cm)containingsubstrate,rinsingitwithwell Spawningsubstratetests.—Twosubstrateexperimentswere water,andthenexaminingboththerinsewaterandthesubstrate conducted, one comparing the preference for either cobble or for eggs. If any eggs or fry were present, their numbers were colored marble substrate, and the second comparing two sub- determined.Allhandlingofeggswaswithabulbpipette.After strate surface-area treatments. In the first test (2007 spawning eachscreening,thesubstratewasreplacedwithafreshtraycon- season),fishchosebetweentwospawningsubstrates:smallcob- tainingdisinfected(1200mg/Lbenzethoniumchloridesolution, ble(21–48mminsize,mean31mm)orcoloredglassmarbles >15minexposure)substrate.Eggcheckswereperformed2–5 (mixof1.5-and2.5-cm-diametermarbles).Fivespawningtanks times per week, concluding when no more signs of spawning (replicates)wereused,fourofwhichwerealsopartofthebrood- activitywereobserved. stockdensityexperiment;thefifthtankhad14broodfishpresent. Egg incubation temperature.—In 2010, survival to hatch Thesubstrateswereplacedinaplastictray(15 × 30cm × 3 wascomparedamongfourtemperatures:18,22,24,and26◦C. cmtall),and1-cm(barmeasure)meshwasplacedintheplastic Eggs for the experiment were all derived from a single spawn traytocreatea“falsefloor”inthetraysthatcollectedeggs.The (18.8◦C).Threereplicatesforeachofthefourtemperaturetreat- false floor was intended to make egg collection easier and to
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