Adam Molesrard Ryan Scott, Auke Bay LaboratoryA aska F sheres Sc ence Center,Natonal N4ar
ne F sheres
Service NOAA 1T305G ac er HighwayJuneau Alaska9980T8626
Growthand Behaviorof JuvenileAlaskanFlatfishesin the Laboratory
Abstract
Ju\cnilc Prcilic halibut reafed in the laboratorygrc$ a! a laster rate md h.rd a more uriform distibutioD ir behavior reslsthan
jurcnilcsofyello$iinsoleandfocksole.Ju\,'enile)'.cllowfinsoleandrock\olehadsimilar90dgro$lhrates({1.71-0.8ltfUWD)
iIr lhc laborator,"'al 10'C. where.rsPacific halibut grer significanll] iaster (l.29L BWD). Rock sole grew rhe saDreon bolh mud
and sand.despilea kno$n prefefenceforsand substratcin the teld. Yellowfin sole androck solc hadan aggregrteddistribution ll1
&. unk\ as meanrred bv nearest-neighboranallsis rcg.trdlessof the presenceof othcl species,whereasPacific halibut had .r
unifbm distribution. These are thc tirsl obser!alions ofgrolvth and beharior ofjLrlcniles of these\pecies in ihe labora|orr-.
Introduction
Understandinginterspeciticdifferencesin growth
and behavioramongvariousspeciesofjuvenile
flatfishesmay increaseundcrstandingof such
specific differencesas timc spent in nurseryar
eas,vulnerability to predators.types and sizesof
prey itens, intraspccific and interspecific competitiol for tbod and space.and productivity of
the stock.Theseprocesses
are all influencedto
some dcgrce by initial growth (Weatherleyand
Gi111987)andsocialinteractions(Olla ct al. 1996)
of juvenilesof many fish species.Growth also
hastensor delays recruitment of f'latflshesinto
the fishery: larger age-0 t'latfisheshave reduced
predationand faste[maturationthitn smallerage
0 flatfishes (van dcr Veer et al. 199,1).In addition, behaviorcan rcgulatedistribution, feeding.
andpredatoravoidancein many lish species(O11a
et al. 1996).
Intuitively. different speciesmay have different growth mtes and distributionsasmechanisms
forpanitioning the nurserygrounds.Juvenileflatfishesreducecompetitiveinteractionsbetween
speciesthroughdift'erences
in habitat.suchasdcpth
and substrate(Norcrosset al. 1995).Thesediflerencesin field distribution of similar-sizedin
dividuals are probably the result of a complex
interaction of a number of v;Lriablessuch as interspecificdifferenccsin growth ratesandbehavioral interactions.
I Author to whon correspondcncc should be adclfessed.
Email: Adan.Nlole\@noaa.go\'
To betler understandtheseinitial diiTerences
among speciesof Pacific flatfishes.we assessed
growth ratesand spatialdistributionamongnewly
\ e l c d r e c r u i rr\ , f l h r e ec o m m e r c i l l l )i m p u n u n t
speciesof flatllshes.We choseyoung-of-the-year
Pacilichalibut(11?poglos,in.i
yellowtin
slenolepis),
sole(Li nanda asp er), ar\dI ock solc (Lep idop setto
bilineans).The adultsof thesespectesconstitute over half of the flatfish catch in the nofih
eastemPacific Ocean.totaling a high of netfly
300,000metric tons in 1992 (FAO 1995).The
combinedflatfish fisherv is the third largestfishcry in the northeastemPacific Ocean,after walleye pollock (Therqgrd chulutgranna) andPa
ciflc salmon (Ontttrhynchus spp.) (FAO 1995).
Extensive knowledge has been gathered about
adultsoftheseflatfishspecies.butfarlessis known
about the juvenile stage.Becausemany of the
environmentalconditionsthat atlect growth and
behavior cannot be controlled in the field, our
studieswere conduotedin the laboratory.
Following a planktoniclar,'alstage,the recruits
of all three speciessettleout into subtidal nurscry areas(<40 m deep) for the first year or two
(Norcrosset al. 1995)wherc they t'eedlargely on
crustaccans.
Following migration into deeper
watersasadults,the Pacitichalibutgrow to nearly
3 m in length, whereasycllowfin and rock solc
ire typically <,15-55cm.In the studv desoribed
here. *e comparedgrowth ratesand spatial distributions amorg young-of-the-yearilatfish juveniles to determine whether differenceswere
evident as carly as the lirst year of life.
N o r t h w e s tS c i e n c eV. o l . 7 6 ,N o . 1 . 2 0 0 2
41
Methods
Preparaton of Anirna s
Yellowfin sole and rock sole wele obtainedfrom
Auke Bay, Alaska. by 6'mm-mesh beach seine
in June.1994and 1995.Pacitichalibutwerecollectedin 10-30 m depthsin Middle and Kalsin
Baysoff Kodiak lslandby plumb staffbearntruwl
(,1nm mesh)inAugust1994and 1995.All spccinens (46 109mm standardlength, mean69 mm
t 15.9)uere transpoftedlivc 1()the Auke Bay
Laboratoryandheldin flowthrcughseawatertanks
Fish wefe
on a mixed sandand mud substrate.
held lor two months to acclimate to laboratory
conditions.
lndividual fish wereidentifiedfbr repeatedsize
measurements
and behavioralobservationsby
marking juveniles on the ventral surtaceat one
or more of four locations along the lateral margin with Alcian Blue dye (65 mg/ml aqueous
solution) to give each lish a unique mark. This
allowed us to measurcthe growth ntes ln each
fish rather than following the mean changeof a
groupoffish. This dye markingmethod(Thedinga
and Johnson1995)is a fast.external,non-invasjve procedurethat pennits mark [etention even
in rapidlygrowingjuvenile fi shes.Unanesthetized
flatfish wereiet iniected $ith dyc using a Panjet
held 25 mm above the skin surface.Excessdye
was wipcd off. and the fish held in a rccovery
tank lbr I hr to verity dye retention.Alcian Blue
dye docsnot affectjuvenile flatfish growth, alter
tissuestmcture,or beoomeincorporatcdinto internal organs(Thedingaet al. 1997).
GrowthTests
Gro$,th testswerc conductedin 70-L tanks (30 x
60 x 40 cm) underoutdoortanslucentpanelswith
a constantl2 hr/d of supplemcntalfull-spectntm
lighting.Scawater
t'lowrateswerel.4Llmat28%o
salinity and l0'C. suchas is commonly obsen'ed
during late summcr in flatfish nursery areasin
southeastemand southcental Alaska. Temperature was controlledby rgsistivehcaters.mercury
s$'itches,and associatedrelays.
Groups of fish were reared on specific substrates.Each spccieswas testedon its prcfened
sediment.Pacific halibut and rock sole on sand,
yellowfin soleon mud (MolesandNorcross1995).
In addition.rock solewas alsotestedon mud so
that growth for that speciescould be neasured
42
Moles and Scott
on prefened and unprefered sediments.Mud was
gatheredintefiidally.trozenandthawedthreetimes
to kill any organismspresent,then sievedto re
move pafticlcs and macrolhunal prey items organism: largerlhan 63 !m. Sand'6-l :4q Fm
diameter)obtainedfrom a local gravel yard was
sicvcdto yield a meanparticlesizeof 125 pm.
Fish were fed to satiationwith a dail.vration of
commercial blood worms (TubiJe.rtubilex) and,
mysids (M1sir spp.) for two weeks before and
suchas mysids
throughoutthe test.Crustaccans
are a common componentofjuvenile flatfish di
ets,andthe additionofthc bloodwormsenhanccd
the fecding rcsponse.The fish were fed six times
pcr dr1 to reduccfeed wa\le and size rariance
due to possiblecompetitiontbr tbod amongfish
(Briinniis and Alaniira 1993).All fish ate vora
ciouslywhenlbod becameavailable.
Therewerefour treatmentgroups(Pacifichalibut on sand,yellowfin sole on mud, rock sole on
sand,and rock sole on mud), each with 20 tish,
rearcdfor 90 d. Becauseit was only possibleto
obtain 10 distinctivedye marks,it was necessary
to spreadthe 20 testflsh ofeach treatmentamong
two tanks.This gave a final total of 8 tanks of l0
fish each.
Individuallcngths(to nearestrrm) andweights
(in mg) weremeasuredat 0, 30,60, and 90 d on
the same individually-markedflsh. Fish were
weighedlive in a taredbeakerof watcr,thcnquickly
measuredlbr length and returned to the water.
Growth ratesfor length(incrementalgrowth mte)
andweight (specificgrowth rate)were calculated
for each fish (Fonds et al. 1995). Daily length
incfements(dZ) were computedfrom differences
in standardlength (l,) over time as: dL = (1"'0
4.,",.)/t,whcrc I is time in days. Specific growth
rate was computedas C = (ln ly..,r ln lv.,,,,)/tx
100,whereW is body wet weight.Only the 90 d
growth mtes are presentedhere. Differences in
growth ratesbetweenthe four treatmentgroups
werc analyzed using a repeatedmeasuresoneway ANOVA at day 0 and day 90 fbllowed by
Tukey'stestof multiplecomparisons
at P = 0.05.
Spatlal Behavior Tests
The spetial distribution of f'latfisheswas determined using a gdd painted on the bottom of a
4.8-m diameter 18.000 L circular tank divided
into I l8 observation
areas.Waterdepthwas I m.
flow rate 20 L/min (to maintain sufficient oxy-
TABLE 1. Mcan lcnglhs. $eighls, and growlh r:ttes(lSE) ofthree speciesofjuvenile flatfishesrearedin the laboraroryfor 90 d.
ICR= incremenlal grolvth rale as mln increaseper day SGR= specific gfowth rate as % body \\eight pcrda]-, N= 10.
Wilhin ICR or SCR. means with the samesuperscriptare noi significantly different (P<0.05).
lro.k $lc .r -uJ
Y e ' l o $f i n ^ e
90d
L e n g l h( n n )
\ V e i e h t( g )
IGR
SGR
7: (1.61
1 . 1 0( 0 . 1 9 )
0d
90d
90d
0d
(1.1) 8,r(3.9)
9l (1.,r)
ll (,r)
89 (,r)
66 (4)
79(.1)
6',7
I 1.63(0.62) ,r.38(0.71) E.89(1.:1) 1.76(0.79) 6.2(0.9.1) 3.96().8s) 6.9(0.92)
0.2,1(
0.ulf
0.19(0.01)r
0 . 1 5( 0 . 0 r ) b
0 . l 9( 0 . 0 r ) '
1.29(0.01)"
0.El (0.04)"
0.71(0.01)"
0 . 8 2( 0 . 1 0 ) b
gen), salinity 29%o. and,tcmperature 10oC; no
sedimentwas present.Twenty Alcian blue dyemarkedjuveniles (averagelength 80 mm) of the
tcstspccicsand20 unmarkedfish of the sameor
a dift-erentspecieswere chosen randomly and
ir'rroducedto the tank by net. Using the grid, we
measuredthc distanccbetweeneachmarkedfish
and its markednearestneighborat 30 min jntervalsfor20hr allov"'ingusto calculatemelurnearest
neighbormtios over 20 hrperiods foreach group.
Atier 20 hr, the fish were removed and 40 new
fish of the next trial were introduced.All 6 comyellowfin sole/yellowfin
binations(halibut&a1ibut,
sole.rock sole/rocksole,halibut/yellowfin,
halibut/rocksole,andyellou'finsolehocksole)were
testedwith tlve replicatetrials eachfor a total of
l00 testindividualsper species.No fish was used
tbr more than a singletrial to insureindependent
obsen'ations.
The dist ce-to-nearestneighbor method(Clark
and Evans 1954)describedthe intaspecific spa
tial distribution of a given speciesin the presence of other flatfishes.both of the sameand of
anotherspecies.Nearest-neighbor
distanceswere
calculatedbetweenmarkedindividualsofthe same
species,but not of different species.The measureof spacing(R) indicatesthe degreeto which
the distributionofthe sampledeviatesfrom a random distribution (R = 1). The smallcr the value
of R, the more aggregatedthe distribution: the
higher the value. the more unifom the distribu
tion. We examineddifferenccsin nearestneighbor ratiosusingWilcoxonsignedranktests.
tion factorsalsodid not dift'ersignificantlyamong
all tanks and treatmentgroups.
Pacific halibut grew significantly taster than
the othertreatmentgroups(P < 0.001).Daily incrcmentalgrowth rates(mntd) were signitlcantly
greater(P < 0.05) for Pacifichalibut thanfbr rock
sole or yellowfin sole (Table 1). By day 90. Pacific halibut had grown 0.24 mm,/d.26% gre;iter
than rock sole on sand or yellowfin sole. All of
thesefeatment grcupsincreasedin lengthgreater
than rock sole on mud. There was a significant
difference in incrementalgrowth rate but not in
spccific growth rate betweenrock sole on mud
and on sand. Other differences in growth rate
between.pecie.or sub:lralellpes uere not 'ignificant. Yellowfin soleandrock solegrew at nearly
identical ratesthroughoutthe experiment.
During the behaviortests,halibut distributed
themselves
uniformlyR = 1.83-2.00)in the experimental tanks when other specieswere not
present.In contmst,when all ,10fish in the tank
were yellowfin sole, more aggregationoccurred
(R = 0.86 t 0.02). Rock sole were significantly (
(R = 0.71 t 0.01)
P < 0.001)more aggregated
thanyellor'fin sole.Distributionpattemsfor each
speciesdid not changewhen a secondspecieswas
present(Table2). excepttbr a decreasein uniformity ofPacifichalibutwhenrock solewerepresent.
Both yellowfin soleandrock solebecameslightly
TABLE 2. Neafest-neighbordistances(R valucs t SE) lbr
experimentalgroupsof markcdjuvenite Ialiishe s
of the same species(mcasured species) iD the
presenccofan equal sizedgroup ofthe sameor a
difttrent species(inlluencespecies).N= l00fish.
Results
All fish grcw substantiallyover the 90-d test period.Yellowfln soleweightsincreasedl03o/r,rock
soleweightsincreased65% (mud)urd 74olc(sand).
and halibut wcights increased1847o.Differences
in initial size betweenthe specieswere not significantfor lengthor weight.Initial meancondi-
Measured
Species
Yellowfirsole
Rocksole
Pacilichalibul
Yellor'fin
lnfluence SDecies
Rock
Pacilic
halibul
0.86(0.01) 0.66(0.01) 2.00(0.1)
0.79(0.01) 0.71(.01)
L83(0.02)
0.83(0.01) 0.73(0.01) L94(0.02)
JuvenileFlatfish Grorlth and Behavior
more aggregatedin the presenceofthe other spe
cies. but this differencewas significant only for
ycllorvfinsolc(P < 0.01).Thc dislributionof Pacific halibut was not significantly different in the
presenceof yellou'fin sole,but was significantly
less unifbrm in the presenceof rock sole (P <
0.01). Yellowfln soleand rock soledid not alter
their spacingin the presenceof Pacitic halibut.
Discussion
This is the firstlaboratoryestimateofgrowth ntes
follo\\"ing settlenent lbr thesevaluablecommercial species.a periodthat is impoftantto lubsequentsurvival.Flatfishes,as most fishes,grow
most rapidly asjuveniles (Paul et al. 199:1).and
llny reductionin juvenile growth rate prolongs
theiuvenile stage.Mofiality duringthis phaseand
timing of onsctof maturationarc dircctlydctcrnined by fish size(Zijlstra et al. l982, Rqnsdory
prcdationon thc
1993.).As fish sizc incre;iscs,
fish is rcduccdfor somc flatfishes,and in such
casessunival is relateddirectly to growth (Wit
ting rnd Able 1993,van der Veer et al. 1994).
Fish that areableto grow quickly during the early
juvenile stage,such as Pacific halibut.are less
susceptibleto predation and have a competitive
, r , . l r . r n l , r, .grer r ' ' 1 , ' ue r - g r r \ iun g l i . h .
The dillerences in growth between Pacitic
halibut and the other flatfish speciesexaminedin
l h i 5 \ l u d ) a r e i n t e r e . t i n gg.i r e n t h e u n i q u ee n
ergyrequirements
ofPacifichalibut.Pacilichalibut.
rvhich grew rnole rapidly than yellowfin sole or
rock solc in our study,havc grcatcrcnergycxpendituresthan yellowfin sole. The oxygen re
quirementtbr.juvenile
halibutis four timeshigher
thantbr ycllowfin solc.duc to longcrmigrations
andmoreactivefeeding(Paulet al. 1994).Juvenile halibutcornpensate
ibr this high rateby eating relatively largermealsthando adult fish (Paul
et l. I99,1).In our study,grouth ratesof Pacitlc
halibutestimatedby lengthwere 26-'l.greaterthrLn
lbr yellorvlin sole, and the specitic growth rate
ofhalibut overthe90 d testperiodwar 557. greater
than for yellowfin sole.The growth lates of flat
fish in our studyare similarto estimatesofgrowth
in thc lirst ycar using agc and size data lron the
Gulf ofAlaska.Smithct al. (1995)cstimatcdthat
yellowfin sole in the Gulf of Alaska grow 0.5%
BWD in their llrst year and Paul et al. (1994) esrimatcdthatPacifichalibutgrowth is atlcasl0.75%
BWD. Pacific halibutjuveniles grow rapidly. tdMdes and Scott
pling in lengthby age-l (Southward1967).This
initial surgcof growth is an impoftantiactor in
determiningyear classstrengthin Pacific halibut
(Hagenand Quinn 1991).
Despitestrongevidencefrom both laboratory
and lleld studiesthat rock sole have a preference
for sandovcr mud, thc growth ratestbr rock sole
on sandwere not significantlygrcaterthanrock
sole on mud. When given a choice in the laboratory betweena varietyof substrates,.juvenile
rock
solesglectthosesubstrates
oontainingsandnearly
every time. When given a choice betwccn mud
and sand only, rock sole always selectedsatrd
(MolesandNorcross1995).In thetield,juvenile
rock sole are often lbund on sand substrates
(Norcrosset al. 1995).
Pacitlc halibut not only difl-ertrom the other
two speciesin spatialbehavior,but alsohavc diflcrcntfccdingbchaviorandthey engagein teritorial partitioning. Pleuronectidsrely on visual
cuesin diurnal feeding (de Croot l97l) in addiIn our
tion to variouslevelsof chemoreception.
tests,Pacific halibut fed in the water column and
relied heaviJyon visual detectionof prey.Adult
Pacific halibut maintainhomeareasspatiallydistinct fronl thc home areasof other Paciflc halibut
(Phillip Hooge.U.S. Fish and Wildlifc Service.
Clacier Bay, Alaska. personalcommunication).
This is similarto the unifbrmdistributionof the
juvenilePacifichalibutin our study.In contrast,
the demersallyfeedingyellou'finsole and rock
solemay aggregatefor the samercasonotherspecies school.such as protectionfrom predato$.
FcmalcPacifichalibutreachan adult length
of I m within only 8 yr (Trumblect al. 1993)in
conffast to yellowfin sole, which mature at an
averagelength of 28 cm about 9 yr (Wilderbuer
et al. 1992).Rock solearebelievedto havea similar
age and size at 50% maturity as yellowfin sole.
This h'emendousdifference in growth was evidenteven in the young-ot-the-year
in our study.
Significantlygreatergrowthratesas age-Ojuvenileswould allow Pacifichalibutto move offshore
sooner,escapenearshorepredatorsat an earher
agc,andmaturequickly.Additionally.vhen Pacitlc halibutdistributethemselvesunifomly with
respectto eachother.competitionis reduccdand
territory is partitioned.Ratherthan adoptagonis
tic behavior common to such fish as salmonids
(Ryer andOIla 1995)to competefor food, Pacific
halibutmay establishhuntingareasdevoidof other
Paciflc halibut. The idea that some of the differencesin size and distribution of adult flatfishes
are the resuhofdit'felencesin growth and behav
ior of agc-0juvcnilesdeservesadditionalstudy
in the 1leld.
LiteratureCited
Brinnis. E.. and A. Alanlir?l. 1993. lvlonitoring thc fecding
activi!- ofindi!ldual fish $'ith a denand teeding sys
t . m . J o u m a lo l F i r h B i o l o g y , l 2 : 2 0 9 2 1 5 .
. i s t r n c et o n e a r e snt e i g h C l a r k .P J . .a n dH C . E v a n s .I 9 5 . 1 D
bof as a measureof spatial relatiorships ir popula
r i o n s .E c o l 0 g ) 3 5 : , 1 t 5- 1 5 3 .
de Groot. S. J. 1971.On lhc inlcrrclarionshipsbctq ccn mor
pholog! of thc alj entar! ract. food and feeding
behaviouf in flatfi\hes (Pisces: PLeuronectifbnnes).
N e l h e r l u n d Js o u n a l o f S c aR c s c a r c h5 r l l | 1 9 6 .
I-r\O (Food and Agdcultufe Organizatior, U.N.). 1995.Re' r < $ o l r h <n J r e . l $ u r l L l i . h J r \ r c , r u r c . . I n - i n c
fishcrics. FAC) FishcriesCircular 88,1,Ronre.ltaly.
Fonds. Ni.. Nl. Tanaka.and H. $'. vrn der Veer. I995.Feedi n g r n d 8 r o l v t h o f r u v e n i l e J a p a n e s ef l o u n d e r
Pardlichth|s ol^,aft16 tn relatior to rcmpeftture and
lbod suppl]. Nc(hcrlands Joumal of Sea Research
l ' 1 : 1I I l l 8 .
Hagen,P T., and T. J. Quinn ll. 1991.Long lcrm groslh d!
n a ' n i c s o ft o u n e P a c i f i ch a l i b u t e: ! i d e n c e o fI e m p e f a ture-inducedtadaiion.FisheriesReseuch(,Amste n)
1 1 : : 8 31 0 6 .
. 995S
. c d i m e npt r e f e r e n c ien
N l o l c s .A . . a n dB . L . N o r c r o s s 1
ju\cnile Pacific fl arfishes.NethedandsJounnl of Sea
Research
l'1:ll7-181.
Norcross.B. L.. B.,A..Holladay. and F. J. \,lurcr. l995.Nurs
cry area characteristici of pleuronectids in coastal
Alaska. USA. Netherhnds Joumal of Sea Rese.fch
l,l:l6l 175.
. 996.
O l l a . t s .L . . I 1 . W D a v i s .C . H . R y e f ,a n dS . M . S o g a r d 1
Behavioufaldeterminrntsof distribution and sur\ival
i D e a r l y s | . t S e so l w a l l e , " ' ep o l l o c k . I r ? , ' . r t , . r
thdLognmmot a syn$csrs of cxpcrinrcntal studles.
F i s h c r i c sO c e a n o g r a p h5! ( S u p p l .l ) : 1 6 7 1 7 8 .
Paul. A. J., J. l\,1.Paul, and R. L. Smith. 199,1.Eners,"-ard
ration rcquircmcnts of juvenile Pacific halibut
(Hippogios s stenolepis)basedon enefgy consulnp'
rion andlrowrh nos. Jounul olFish Biolog! 1,1iIli23
1l-JlL
Rijn sdorp.A . I). 1993. Relationshipbet$'eenjuveni le grolvth
md the onsei of sexualnraturny offenrale Nonh Sea
pltice. Pleuro|ectet pldtur'.\.r.Cttnadian Joumrl ol'
F i s h c r i c sa n dA q l r a l i cS c i c n c c s5 0 : l 6 l l l 6 : l l .
Acknowledgements
The authors would like to thank Dr. Brenda
Norcross of the University of Alaska Fairbanks
lor hcr rcvicws.advice,andencouragement
in thc
executionof this study.
R y e r C . H . . a n dB . L . O l l a . 1 9 9 5 .T h e i n l l u e n c eo f l b o d d i s iribuiion upon the delelopnrentofaggressiveandcorr
pcrili!c bcha\iour in ju\cnilc chum salnon.
Otftorlt\Ichus k(ta. Joumulol Fish Btulogy ,16:26.1
212.
S n r i t h .R . L . . A . . 1 P
. a u l .a n dJ . N 4 .P a u l . 1 9 9 5 .M i n i m a l r b o d
requirenents lbr tellowlin sole ir Alaska: esiimate\
fromlaboratory biocncrgcrics.Pagcs185 295 /n Pro
ceedings of the International SynposiuIn on Nofih
Pacjfic Flatfish. Alaska Sca Crant ProgramAK SC
95 0,1.Unnersity ol Alask.r F irbanks.A],rsk.r.
Southward, G. NL 1967. Gfowrh of P.rcific halibut. Repon
4 l i l 5 9 . I n l c n i a t i o n aP
l a c i f i c H a l i b u !C o m m i s s i o n .
Seatrle.Wrslingron.
T h e d i n g a J. . F . a n d S . \ V J o h n s o n .1 9 9 5 .R e t e n t i o no f j e t iniec|edmarks oniu!cnilc coho and sockclc salmon.
T r a n s a c t i o n so f t h e A m e r i c a n F i s h e r i e s S o c i e t f
12,+:782
785.
T h e d i n g aJ, . F l .A . M o l e s .a n dJ . T . F u j i o k a .1 9 9 1 .M a r k r e
lcnlion andgrowth ofjcl injeclcdjurcnilc marinc lirh.
F i s h e r )B u l l e t i n( U . S . )9 5 : 6 2 9 6 3 3 .
T r u l n b l e ,R . J . , J . D . N e i l s o n .W R . B o w e r i n g . . l n dD . A .
l\'lcCaughran.D. A. 1991. Atlantic halibut
lHippo(bssus hippogbs$rr) and Paciiic h,rlibut (,ry'
.\reri)l?iJ) and lhcir North Ameican fishcrics. Ca
n a d i a nB u l l e r i n o f F i \ h e r i e sa n d A q u a t i c S c i e n c e s
2 2 7 :I E , l .
lan der Veer,H. \.. R. Berghahn,lllld A. Rijnsdorp. 199,1.
Inpact ofju\,enile gfowth on recruitnent in flrtfish.
Netherlandi.lourno
a fl S e al i e s e a r c h3 2 : l 5 l 1 7 3 .
Weatherley.A. H. and H. S. Cill. 1987.The Biologr- of Fish
Growth. Academic Pfes\. London. England.
Wildeftuer T.K.. C. E. \Lrlters. and R. C. B,rkliala. 1992.
Yello\\'fin sole. Plturotp.ttt dvrf. of the eastefn
Bering Searbiologicil characlerislics.history oi ex
pbitation, and managemenr.
NlarineFisheriesRevie\\'
5 ' 1 ( ' l ) : 11 8
Wit{ing, D. A. and K. \{ Able. 1993.Eftects of bod} size on
probabiliq of predationfor ju\'enile summerandwinter
tlounder bascd on laboratort expcrinrcnts. Fishcr!
B u l l e f i ( U . S . )9 1 : 5 7 7 5 8 1 .
Zijlsra. J. J.. R. I)appcr.J. lJ Witte. 1982.Scttlcnrcnl.growth
a n d m o r t a l i t y o f p o s t - l a r v a lp l a i c e ( 1 ' l e u t o n . c l t : \
p/.J/er'.ld)in thc scslcm \laddcn Sca. Ncthcflands
Journalof SeaResearch15: 150 172.
Received9 April 2001
Atcepted.fttrpublit:ation 2EAugust 2001
JuvenileFlatfish Growth and Behavior
,15