Departmental Bulletin Paper / 紀要論文 Effect of temperature and salinity on egg hatch of the rainbow trout <special>Salmo gairdneri</special> ニジマス卵の孵化に及ぼす水温と塩分の影響 Kashiwagi, Masaaki; Iwai, Toshio; Saji, Atsushi 三重大学水産学部研究報告 = Bulletin of the Faculty of Fisheries, Mie University. 1987, 14, p. 41-48. http://hdl.handle.net/10076/3541 Bull.Fac.Fish‥Mie Univ. No.14:11】19 September30.1987 First Matur‡ty andl両tia[Growth of Some Common SpecleS Of Bai’naClesinJapan. ToshiakiIwAKIand HideshiHATTORI Faculty of Fisheries,Mie University 11hebまologlCalminimum sizesand growth ratesoぎsまⅩSPeCiesofbarnacles havebeendeもerminedbyexamirlatまonorspecimenstakenfromraftsorintertまdaユ piles and by measuring the rostrかCarinaldiame廊Or animals growing on glasspanelssuspendedfromarart. ThesizeandageatrirstmaturltyWeredeterminedasfollows;及£mproぴ言古㍑ぶ 4.5mmbasaldiameも饉(17days),鼠αJわgど0ぶとα£㍑S5.2mm(17days),且reとよc比gαとαざ 7.5mm(?),且孔明痛かぬ7.3mm(22days),β.eわ比㌻花e㍑β12月mm(50days)and鼠ゐ0花ぬ− hoL)ilO.Omm(?).Fromcornparisonoftheseresultswithothcrbarnacles,itfo110WS that au ofthe barnacles sもudied here show early age and smaller size aもfirst maもurlty.Iniもは1growもhin rostro−Carinaldまrectionis slow unti16days orso arter seもtlemenも,barnacles reaching・aPprOXimatelylmmまn basaldiameter・ From6to18daystherat80fgrowth or any species accelerates and barnacles become3.5to5.Ommlarge at the end ofもhis period・Subsequent decline of the growth rateisexpectedもO be dueもo the sexualmaもurまもy・UntillO days afも群 metamorphosis barnacles tend to expandもhe basalmargln rather thanもhe OPerCularportion. Keyword:Breeding・,Growth,鮎rnacles In previous papers(IwAiく1,1975;1981),it has already been clarifiedもhat the adult ・!・.∴∴. 一丁‥・・∴∴.、∫.・・∴..り、− こ、:∴・−∴−∴・..・.、−・.・‥−∴・−・二、・‥J・・・、、●‥・・・ご・‥∴ヾ 、; −・ご・、.‥.−‥‥・.‥∴・.、.・ J∴・∴・・・∴ご・1.・.ご・、∴ 二 !.・.‥・‥.、.∴‥∴、.、 TARASOV &ZiミVINA andβα∼α花捉ぶrgと£c昆gαと㍑ぶUl、rNOM】,Showed the different reproducもive activities,COmParedwiththos80r nOrther−1SpeCies;anyindividualof theabove−mention¢d sp¢CiescouIdbreedseveraltimesdurirlgthebreedingseasom・ Several papers have been published on the breeding and the growth of other barnacles (ClヰISr)andPAでlミし1961;BAIINl三SandSTC)NFニ,1973;HINifS,1979)andrecentlytheliteratures has be8n reViewed by ClそISI)and‡ヨOURGl汀(j986).AYuNG(1976)and Hurtu三Y(1973) 12 γoshiakiIwAlくIand HideshiI−‡A′】、ri、011l attempted to rationalize th扇r daもa onlirehistories or月αgα花㍑ぶ抒ぼ0花αβandβαぬ花㍑ぶpαCよ− ficus using MACAitrE−川三Ⅰそand WILSON・s concept and reached the samc conclusion that the rapidgrowth,earlysexualmaturity,hig・hfecundityandgrcatdispersalabilitieswereadvan− tageousforadaptivestrategiesoftheso‘opportunistic’or‘fugitivQ’(トSelecもed)species. Fil、Sもmaturity andinitiaigrowもh areimportantitems of basicinformation ror under− Standingthelifehistoryand reproductivebiology of barnacles.To rnakecarefulconclusions OnthebiologlCalminimumsizeandthebreedingactivlもyinrelation to sizeor age,Observか tionsonagreatdealofmaterials coveI・ing・もhewholesizerange and froquent measurements Ofgrowlngindividuals weremade. MateriaJs and methods DuringMay−October1986specimensofβ.αJγ堺たか∼£eand且ゐ071dαたou∠atIzo−uraillthe recessofMatoyaヱさay and those of且∠〝岬rOUgぶ㍑ぶat Matiya−uIta dueeast of thecampus of MieUniv即Sityw8r8remOV¢drrom bamboopoles,Which‡ladbeensもuckintotheseabedinth¢ 1ittoralzon¢byfarmersforculturlng¢dible red andgreen algae,During A咽tlSt−September, 且eわ比r花eαぶ,β.α批coぶとαと㍑ぶand且 re乙£cαgα£混ぶWere Obtain¢d from styrofoam floats and Otherartificialstructuresunderthe rafts,Which werealso mooredin themarglmalpユots of MatoyaBayandAgoBayrorculturingoysterand peal・loysもerrespectivdy.So,thebarnacle POPulation on the bamboo poles and under the rafts w8reinfluencedbythedifferent hydro− graphicconditions. BarnaclesfromthQ¢ntiresizera咽eOfeachspecieswereselectedatrandomforprocesslng inth¢laboratory.Eachbarnacleswasmeasur¢dalongth¢apicalandbasaldiametQrOfrostro− Carinalaxis,and exa‡Ⅵined forthepresonce ofegg masses. Sixrectangularpanelsoftransparentg・1assmeasurlng18×14cmwerepreparedforporiodト Calinspectionofgrowthrate.Inorderforthelarvatoattachthemselveswell,eaChpanelwas processedbydrillinganumberofsma11pitsontoもhesurface,Thepaももern of settlemenもCan bepredeterminedbythismethods,becausethecypridssettlepreferentiallyintohemisphericai smalipits(ClえIS‡J,1960)Eachtwopanelsholdingablackpaneまbetweenthemselveswerofixed togetherinto a wooden frame and therlthree sets of panels were suspendod at O.5,2and 4m below the surrace of the sea from a raft mooredin Matoya Bay.Tbe panels w8r¢ examined every three days from Au糾Sもto SQpもQmber andもhereaぎter at somQ mOrelo咽er intervals.Ateachexamination theposition ofeach settled barnaclewastraced ontoanother Clearplate,Placedonthebackofasurfaceofthepanelwithgrowlngbarnacles,alladditionai foulinggrowthwasremovedand therlaPICaland basaldjametcrof rostro−Carinalaxis were measured underabinocularmicroscopewiもhocularmicrometer. First maもuriもy andinitまalgrowth of barnacles 13 ResuLts l)R叩rOductivecondition. Instudyi咽theareaonthePacificcoastofcentralJapan,ithasbe紺foundthatsucces− sionalbTOOdsapparentlyoccurduringApril−DecemberinB.improuisus,鼠amphitriteandB・ ebumeusandduringMay−OctoberinB.hondahoui,B.reticulaiusandB.albicostatus・They show no distincts組SOnalcyclo with synchronous release of young as observedin northern species(BA】モNES and BARNES,1954;CR‡汎1954;BARN略1959)・Å1lof the samples examined here which were collected during May−October were thereforein reproductive condition. ︵辞︶ト0⊂¢⊃bむ︼軋 l00 80 dO AO 20 0 O 1 2 3 4 5 d 7 8 9 】0 1l O 1 2 3 d 5 る 7 8 9 】O1112】314 15 Api¢a暮 diameter a10ng th◎rO$trO−Car【nalaxIs(mm) Fig.1.Percentage occurence of barnacles bearing egg masses・ In Fig.1the percentages of animals with egg massesまn the samples are shown as a flユnCtion oftheapユCaldiameもer.The size ra咽eS and frequQnCies pertainlng tO the smallest individualscarrylnぎeggmaSSeSVaried withspecies−Precisedimensjons ofthesmallest ones w8re2.15mmforβ.£汀岬rOUZぶ㍑ざ(Nニ474),2.54mmforβ.α翫coぶまα£㍑ざ(N=509),3.56mmfor且re己Z− c㍑∼α£混ざ(N=510),3.65mmfor鼠α?叩ゐかよ£g(N=842),4.85mmforβ.ゐ0花血如u∠(Nニ999)and 5.08mmforB.ebunLeuS(N=ニ660).Inan adult,eSPeCially southernspeciesofbarnaclewhichis 14 ToshiakiI\VAKiand HideshiHArユ「′ユ、ORr PrOducing broods rapidly and repeatedly durir唱the breedingseason,thefertilization occurs immediately afterma七uration ofgonads(CRIS盲〕and DAVほS,柑55).Ifitdid soinanimals Which had jus七㌻組Ched maturity for thefirst timein theirlife,thevaluesglVen above for each species can be regarded as the biologユCalminirnum size.Thereafter the proportions Ofanimalsbearingfertilizedeggsincreasedwithincreaseinsize,andonreachingcertainsize becamealmostasteadystateindistinguishablefromthoseoflargerindividuals.Thesizeat Which theぎrequencies ceasQd toincrea3e COuld be rouが11y decided as rbllows;4.Omm for β.わ研rOぴ∼ぶ混ざ,5.Ommfor且α路お0β£α己揖,5.5mmfor及re£去c昆ぬと㍑ぶ,6.Ommforβ.α〝軍兢rよ紘 6.5mmforβ.eあ比r花e㍑ざand8.Ommfor鼠如71血ゐou£.Thes¢Valu8Slikewise suggested to be Parallelwith maximum sizes of barnacles observedin naturalpopulations,aS Shown by marginalsizesrangosinFig.1. 2)Growth BeforodrawlnggrOWthcurv8StberelationsbetweenapICalandbasaldiameterwerefirst estabiished.Fig.2is based on a number of measurements of animals which were able to expandthebaserreely underexperimentalconditions.Therelationshipfoundtogivethebest COrrelation was two allometric8quations that were obtained separateiy,for thc small8r β●∫ ラ/ 0.5 1 2 3 4 5 0.5 1 2 3 4 5 0.5 1 2 3 Apicaldiameter(m†河 Fig.2.Relations between apまcaland basaldiameters measured along the rostro−Carinalaxes or shell. First maturity and initial growlh of barnacles 15 Tablel.Regressionformulashowingtheallometricgrowthofapicaldiameter(Ⅹinmm) relative to basaldiameもer(Yin mm)or barnacles. Species Smaller group Larger groups 鼠∠mprou£ぶ比ぶ Ln Y=0.692+1.362×Ln X Ln Y=0.770+0.948×Ln X 鼠 αgむ£coぶ£α£昆ぶ Ln Y=0.718十1.3ユ8×Ln X Ln Y=0.827」−0.870×Ln X 且αナナ叩仙「涼 Ln Y=0.744+1.538×Ln X Ln Y=0−698+1.298×Ln X Ln Y=0.699」−1.035×Ln X Ln Y=0.743+0.964XLn Y 風 eわ捉㍗花e比ぶ Ln Y=0.728+1.384×Ln X Ln Y=0.833+1.016×Ln X β.ゐ0花dαゐ0乙l∼ Ln Y=0.636+1.248×Ln X 鼠 re££c紀∠α餌S groupslessthan2mmandforlargergroupsexceedinglmminapICaldiameter・Theintersections ofthetwo r曙reSSionlines foreachspecies werenotlar酢1ydまfferent rrom¢aCh other,もhe co−Ordinates showing the values ofl.14tol.33mmin apicaldiameter and2・40to3・07mm inbasaldiametQr,QXCQptthat及ref加Jαと昆8hadrathersmallersizedpointand及reとょc㍑gαと昆ぶ onlygaveasl咽1elin¢aStheresultofinsufficientdat臥Abovetheseresults朋reSummarized in Tablel.Whereasin any smaller groups the relative growth coefficients,Or eXPOnentS, werelargQrthanon¢,thosevaluesinlargergroupswereequalto,OrSOmeWhatsmallerthan one.AnimalscomposlngSmallergroupsthereforehadtheirshellbaseswhichhadachievedat higherspecificgrowthrate(dy/dt/y)thanthatrate(dx/dt/Ⅹ)oftheorificeofshell,but inlargergroups both thesecompartments of shells dev8lopedisometrically(dy/dt/y= dx/dt/Ⅹ).Thisfactcan beunderstood fromtheallometricequationinitsdifferentialform一 In any sp∝ies ofbarnaclesもh¢heavy settユem孤tis observed fromJune to Octoberin Matoya Bay(ⅠⅥ∫ÅKI,H王別NO and KAWA】【IARノ\,1977).且reと∠c“gαと昆ぶand及£汀軍rOUよぶ㍑ざ mainlysettledonpanelsatd叩thof2and4mrespectivelyandotherfourspeciespreferred tosettleonuppermostone.Fig.3givesthegrowthcurvesofbarnaclesbasedonaveragesize offivetotwelveまndまvidualsforeachsp∝iesat8VQrymeaSurement,butthem組Sur¢men七SOf 鼠ゐoJldαたou£and風花乙£c比gα£昆ぶWerelimitod to a錆w number orindividuals as wellas to shortperiodsowlngtO alargenumberofmortalities・Itseemedthattherewasashortlag phaseveryearlyinanygrowthcourse,Showingthegrowthrateofca・0−1mm/dayinthefirst6 days.From6to18daysthegrowthofanyspeciesaccelerated,barnaclesreachingale咽もhof 3.5to5.Ommattheendofthisperiod.でhereaどteritsloweddown,eXCePtthat放め㍑r花eむぶCOn− tinuedtogrowin thesamemannoTaSbeforeFor30daysormore・Thegrowth rates ofB・ improuisus,B・albicosta己usandB・amPhitriteduringtheintervalfrom15to18dayswereO・28 mm/day蓼0.44mm/dayandO.55mm/dayrespcctivelyandduringthenextintervaltheratesreduced byhal壬l・CoQfficiQntSOfvariationassociatedwithaveragesizesofbasaldiameterwere13to 18%atthefirstand2】to40%attheendoぎgrowthcourseof飽Chspecies. Fig.3canbeusedinconjunctionwith bothIPig.1andダig・2givenabovetodetermine approximatesizeandageatwi−ichthebarnacleぎirstreachesmaturityandthosecorr¢SpOnding withtheintersectionoftworegrQSSionlinescorrelatingapICaldiameterswithbasaldiameもersL でoshiakiIwAK王 and HideshiHATTORI 16 8 6 ︵∈∈︶㌫芯∈万石 4 嵩め吋皿 0 3 6 9 12 15 18 21 24 27 30 Days after settlement Fig.3.GrowもhofbarnaclesallowedもOSettleong・1asspanelsinMaもOyaBayfrom midA咽uStもOmidSepもember. i:及∼mダrOぴgS比ぶ,e:及eあ㍑r几e㍑ぶ,al:風α路£co8£α払5,am:β.αmp九たr£紬, k:鼠如花血鬼0ぴ£,r:β.reと∼c捉gαと㍑∂. Table 2・Sizes and ages corresponding・tOinflection point of allometric growth of もhe shelland onset of first maturity. First Maturity 王nflecもion poinも Species Apicaldia. Basaldia. m m 2 1 2 1 3 1 且 克or乙dαゐ0乙Jよ 2 0 β.eむu/・/!ビ〟S 5883朗4007一 1 β.αmpんか£とe m m l 鼠 re乙∼e比gα£昆ぶ 2128911433⋮ 且improL壷uぶ β.α拍ね08fα£㍑8 Age Apicaldia. Basaldia. 9days 2.15mm lO−11 2.54 7 3.56 12−13 3.65 5.08 10 4.85 ユ0 Ag・e 4.46mm 5.15 17days 17 7.48 7.32 22 11.99 9.98 ca.50 Ther8SultsaregiveninTable2.It can be seen that the changes c)f positive allometry to isometryin the relative growth of barnacle shells occur first of a11before first maturity, andthatasr曙ards及∠mproび血ぶ,及α路gcoぶ£αと揖and且αmダ兢rgとetheageandsize orthe Smal】eStbarnaclesbearingeggsnearlycoincidewiththoseatwhichthegrowthfelloff(viz. 3ふ5.Omminbasaldiameter;18days aftersettlement),but thereis aslight discrepancy of Sizcs onlyin B.amphllrite. First maturlもy andinitialgrowもh or barnacles 17 Discussion A11individualbarnaclesexaminedhere werefrom the naturalpopulation undercrowded COnditions,SO they would be capable of cross−fertilizationまn naturalconditions.Barnacles Separatedfromothers byadistancegreaterthanthatoftheextendedpenisprobablydo not rertilize(CRiSP,1950).This f飽tureis quite generalin cirripeds,eXC叩tin the case of Cん£たαmαg昆ぶぶ£eggα£昆ぶ,掩rrαCαぶか℃e′花王αandβαぬ花昆叩eげorα乙昆ぶ(BA‡ヱNESandCRiS】〕,1956), Inhisstudyorbreedinginβαgα花㍑5わαJα花0£desMool11ミ(1935)suggestedもhatthesterility WaS foundin thQla柑er Or Olderindividuals,becomlng mOre frequent withincreasl昭Si狗 and was commonin members ofcrowded communities.As regards species studied here,the barnacles more than two years old wererarQly foundin their habitats wher¢Sampling was made,becauseintense settlement occurs armually over the greatly extended period andis largelyresponsiblefordeathofanimalssettledinpreceding・year.ThemajOrityofspecim¢nS brought ouもin】?ig.1arethereぎoreconsidered to be relativeけyoung and normaleondition Withouts七倍rility. Theimportant factors doterrninlng the size attained byindividuals are theintra−and inter−SPeCificcompetition,the timeofsubmergence and the water movement carrying food Withinrangeoftheanimals.Crowdingmayslowdown growth and delay the maもuration of theovaryconsiderably(CRiSP,1961)andtheraftcondiもionsarefarbetterfromthepoinもOf View ofgrowththan thosewhich areobtained onintertidalsubstrata(CRISP,1964).Using 沌eresultsrecordedherofortherateofgrowthundertheraftcondiもion,thesQWOuldth即efore glVetheminimumagesoffirstmaturityfor及αJ?Wたか∠絶息如花血ゐou£and及∠J7箪rOUよぶ㍑S, Whichweresampledfromcrowdedpopulationsonintertidalpiles, By comparlngmany reSults of this and other studies for breeding of barnacles,it can beseen that thereareconsiderabledirferencesin age and size of the first maturity of the Same kind of barnacles from different r喝10nS.New generations of及eゐ昆㌻花e昆ぶin Trinidad Mangrove Swamp carlreaCh maturitylnless than two weeks,during which the barnacles growfromabout3to5mminbasaldiamQter(BACON,1971),COmparedwiththetwomonths required at Woods Hole(GR^Vlラニ,1933)and the50days(12mm)in Matoya Bay.11iRANO and OKUSHJ(1952)found thatinlate spri咽and summer鼠α汀pゐかよ£grel飽Sedits first naupliiabout40days after settlementin Aburasubo Bay.In this area the temperature of theseaduringthesameseasonwasabouも22to27℃,SOthebroodperiodofB.amphitriteis COnSidered to be 4to 6days whichis estimated from the relationship between the brood period and temp8ratur¢preViously published for the same varieもies,及 αJれpんかょとe var. deTLticulata(PA{rH7Jand CRISP,1960)and B.amphitrite amphit7・ite(CRTSP and CosTLOW, 1963).Inthiscaseareasonableestimateofthe昭eforthefirstmaturityof及α77軍た£とr∠£eis thereforeabout34to36days,andthesize at that timequoted from HmANO and OKUSHI’s growthdatais8tolOm乱AIsoiもcanbefoundinsizeイrequencyhistogramsof及α′叩兢㌻如 inShimizuHarborgivenbyKosAKAandIsmBASm(1980)thatthesmallestanimalshavi咽 eggmassesarefrom3to3,5椚inapicaldiameter.Thesevaluoscorrespondwiththe6to7mm 18 ToshialくまIwAIくIand HideshiH▲へrl、′1、Oiモ】 basaldiameterobtainedbycalculation uslngtheequation shownin Tablel.1n Amsもerdam afewindividualsofB,improuisusobserved to contain eggmassesare5mnlin basaldiameter (Bl∧もー三汚Ml;N,1934).Itis approxjmately similar to the values obtained hereゴ予or t五e same dimension,thoughthegrowth rateis slower than thatin Ma七Oya Bay.Comparisons of the agesoffirst maturitywithin awide diversity of barnacles show that・thc values quoted for SOuthern species are generallylower than thoseま−or northern species,SuCh as月.あα∼α柁Ogdeぶ, 鼠βOrCα己昆ぶandβ.九αmer∠that do not release naupliiuntilnearly the end o壬、their first yoar(ClそISl⊃,1954;Moo兄,1935)The values of size aもrirsもmaturityパVeniflimited to SOuthern species,Vary markedly with specics from about 2mm for BalaTluS trigonus and C如九αmαg昆β/よぶぶ㍑ぶtO18mmbasaldiameter ror7甘けαCZgαぶq昆αmOSα(AYjJtNG,1976;H‡NJミS, 1979)WemaycorlCludethata1lofthebarnaclesstudiedherereachtheirmaturitiesatrelative− 1yearierag■eandsmall¢rSize, Earlygrowthjustaftersettlementwasobservedtobeslowerinthepresentstudy.Similar resultswereobtainedbyCRIS7〕(1960)forElminiusmodes乙us,Bala71uSCrenatuSandBalanus balaTとOidesandheascribedthisfacttoreductionoftherateofflowintheimmediatevicinity Ofthesolidsubstratum.Anotherpossibleexplanationofthcdclay wasthataftersettlement endogenouscontrolofthegrowthwasforawhilelimitedtolowdegreeinconsequenceofthe metamorphosis.でhe slowlng downin growth rate aft即maturi七yis obviousまy caused by meもabolicchanges associated with reproduction.Themostcriticalphasefor survivaloccurs duri咽Orjustaftermetamorphosis,Whiletheshellplatesarenotfullyformed,harde!led,and cemented to thesubstratum(ClもIS】〕,1961).Tll¢faster expansion of the basis rather than apertureofshellisth¢reforeadvantageousforsuchindividualsashaveg喝edlessthanlOdays Or SO afもer settlement andin crowding condition smallspecimen mtlSt COmPIQte aS raSt aS POSSibletofixitselffirmlytothesubstratum. AcknowLedgmenl: Theauthorswish to expresstheirgratitudeto Mr.T.Ul(A10fMieUniversity for kindly helplr将1nSamPlir噂atthefields. References AYuNG,A.M.,1976.The straもeg・y Of orientationin the barnacle Balanus trigoTluS.Mari71e β£ogogγ,36:335−342, BACON,p.R.,1971,ThemainもenanceoraresidentpopulationorβαJα花比ぶeわ㍑′【几e混ざ(Gould)in relation to salまnity rluctuatまonsin a Trinまdad Mang−rOVe Swamp.ノ.eズp.mαr.β£oZ. gco∼..6:187−198. BARNuS,H.,1959.TemperatureandthQlifecycleof鮎∼α花£ムSゐαgα氾0£deぶ(L.).工n“Marineboring・ alldroulまngor官anisms.”ed.byD.L.RAY∴Univ.Wash.,p.234−245. …………… and M.BAlもN!ミS,H,1954.The g・eneralbiolog・y Orβαgα托比S占αgα花Ogdeざ(L.)Da Cost臥 0£ゐ0ぶ,5:63−76, 】ざir・St maもurlもy arldinitialgrowth og barnacles 19 …………… and D.J.C‡∧モIS】∧),1956.Evide11CeOrSell■イertilizaもionin certain species or barnacles.ノ. Aオαr.ぁわ∼.Aぶぶ.ぴ.∬リ35:63ト639. …………… andLStone,1973.Thegeneralbiolog・yOf Verr乙ヱCαSとroemgα(0.F.MGller).fr. Reproductivecycle,POPulatまonstructure,andractorsa汀ectingreleaseornaしIPii.J eェp.mαr.β£og.∬co′り12:279−297. B】i王ニl三入用N,しⅤ∴柑34.ZurBiologievon鮎∼α花混ざよmpro〃Zぶ捉S(Darwin).Zoo∼,A花Z.,105:247−257. CRほ1〕,D.Jリ1950.Breedir唱anddistributionorCゐ£九αmα∼昆Sぶ紬ggα£比ぶ.Ⅳα£㍑re,柑6:31ト312. …………‥・ ,1954.′rhe br・eedingoi’月αZα托昆ぷpO′、Cαと比ぶ(DaCosta)inもheIristlSea.財αr,月∠og. AsぶOC.ぴ.ガリ33:473−496. …………… ,1960.Factorsinfluencinggrowth鵬ratein鮎gα/1はぶわα∼α花0∼deぶ.ノ,A扉m.だco∠リ29: 95−116. …………… ∴柑6ユ.Territc)rialbehaviourin barnaclesetも1e】11ent.ノ.g埠フ.月わgリ38:429−446 …………… ,1964.Ar盲aSSeSSmeritOrPlankton grazingby barnacles.Ill“Grazi王1ginもerresもrialarld marineenvironmer】tS.”(Brit.Eco】.Soc.Symp.4),ed.byJ〕,J.CRISP,P.251−264, …………… ,and A.DA\7=札1955.Observa貢011S玩い正和On the breedまng orだ′mgr乙∼比ぶmOdesと㍑ぶ grown on gまass slides.ノ.〟αr.わよ0∼.誼sぶOC.ぴ.∬リ34:357−380. …………… ,andIヨ▲PA′】、I三1∧.,1961.rrheinもeracもion between breedillga‡1d growth raもeirlthe barnacle ∬gmg花王㍑ぶmOdeぶf㍑βDarwin.ム∠m几Og.&OceαrとOgrり6(2):jO5」15. …………… ,andJ.D.CoSrl、LOW,1963.The toIcranceofdeveloping・Cirripede embryos to salinity and temperature.Og如s,14:22−34. …………… ,and軋BOUlモGl;;Tr,1986.Growthirlbarnacles.in“Advancesin marirle biology.”vol. 22,ed.byJ.H.S.B】∧一AX′1、lミiモetal.,Academまcpress,p,199−244, GRAVに,B.H.,1933.Rate of growれage at sexualmaもurlもy,and duration orlire of certain sessileorganisrns,atWoods fiole,Massachusetts.月∼0∼.β㍑はmαr.βgog.エαわり Ⅳ00dぶ 、抒0∼e,65:375−3B6. 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