Download presentation
1 -

VerticalmovementandbehavioroftheoceansunMolamolainthenorthwestAtlantic


MolamolaNorthwestAtlanticOceansunPop-upsatellitetagVerticalmovementPop-upsatellitearchivaltagsPSATswereattachedto31oceansunMolamolaintheNorthwestAtlanticbetween2005and2008inordertoexaminetheirvertical

fanny's Recent Documents

GOALPreventionCounselingandScreeningUnderstandroleofpediatricianpreven
GOALPreventionCounselingandScreeningUnderstandroleofpediatricianpreven

DiscusstreatmentwithhypothyroidismDiscussionshouldincludeearlylimitationsnewbornscreeningswellDiscusscauseshyperthyroidismIdentifyindicatorsendocrinedisorderMedicalKnowledgeIdentifyimagingstudiesavail

published 1K
Kerala Ayurveda Paramparya  vs State Of Kerala  on 13 April 2018Author
Kerala Ayurveda Paramparya vs State Of Kerala on 13 April 2018Author

Points for consideration-Indian Kanoon - http//indiankanoonorg/doc/2987190/2Indian Kanoon - http//indiankanoonorg/doc/2987190/3DiscussionThe Indian Medical Central Council Act 1970b shall practice Ind

published 0K
abc  abc Associative Property
abc abc Associative Property

Addition Property If a b ac bc If you multiplied the same nonzero to both sides you have used the qrrs qsr Distributive Property a0 0 If you multiplied by 0 and got 0 Multiplication Property W-1

published 0K
Building Connectedness in the Classroom and
Building Connectedness in the Classroom and

on Campus A Look at McGill Universitys Architecture and InfrastructureBy Gal KramerSocial Connectedness Fellow 2017Samuel Centre for Social ConnectednesswwwsocialconnectednessorgAugust 20171TABLE OF C

published 0K
Course Title
Course Title

Course NoCreditsCourse NoCourse Title/FieldCredits10-801-195Written Communication ORENGL 101English Composition I310-801-136English Composition 110-801-196Oral/Interpersonal Communications OR10-801-19

published 0K
CONGRUENCESINVOLVINGCOMBINATIONSOFTHEBERNOULLIANDFIBONACCINUMBERSBYRIC
CONGRUENCESINVOLVINGCOMBINATIONSOFTHEBERNOULLIANDFIBONACCINUMBERSBYRIC

MATHEMATICSRPKELISKY2t/2ihtFxt22ezt/sinh22and2ext/2coshe32HenceifxisanumberandnisapositiveintegerFnxx1n-X-1nandInXX1nX-1n2nIfn0thenFox0andLox2foreverynumberxThesequenceofFibonacciisdefinedbythediffere

published 0K
PRACTICE ADVISORY
PRACTICE ADVISORY

1February 20 2017EXPEDITED REMOVAL WHAT HAS CHANGED SINCEEXECUTIVE ORDER NO 13767 BORDER SECURITY AND IMMIGRATION ENFORCEMENT IMPROVEMISSUED ON JANUARY25 2017Expedited Removal Prior to Executive Order

published 0K
KULVWRSKHU
KULVWRSKHU

/DVW1DPHLUVW1DPH7HDFKHUQGULRORULDQD/LQQDQHVKUDIDDQ/LQQDQHWPDQ6RSKLH0DWWHUQDUDMDVDQQDKUDSHUKDOHQHHNDUDSHUROH0DWWHUQXVKVDDF/LQQDQHXWWOLQDUDSHUDVKLRQ0LFKDHOUDSHUDVWHOODQRVXQWHUUDSHUDVWLOORDULQD0DWWHUQKDQ

published 0K
Download Section

Download - The PPT/PDF document "" is the property of its rightful owner. Permission is granted to download and print the materials on this web site for personal, non-commercial use only, and to display it on your personal computer provided you do not modify the materials and that you retain all copyright notices contained in the materials. By downloading content from our website, you accept the terms of this agreement.






Document on Subject : "VerticalmovementandbehavioroftheoceansunMolamolainthenorthwestAtlantic"— Transcript:

1 Verticalmovementandbehavioroftheoceansun
VerticalmovementandbehavioroftheoceansunMolamola,inthenorthwestAtlanticIngaF.Potter,W.HunttingHowellLargePelagicsResearchCenter,DepartmentofBiologicalSciences,SpauldingLifeSciencesBldg.,UniversityofNewHampshire,Durham,NH03824,UarticleinfoArticlehistory:Received30May2010Receivedinrevisedform11October2010Accepted13October2010Availableonlinexxxx MolamolaNorthwestAtlanticOceansunPop-upsatellitetagVerticalmovementPop-upsatellitearchivaltags(PSATs)wereattachedto31oceansunMolamola.intheNorthwestAtlanticbetween2005and2008,inordertoexaminetheirverticalmovementandbehavior.Tagsremainedattached JournalofExperimentalMarineBiologyandEcologyxxx(2010)xxx Correspondingauthor.POBox4524,Portsmouth,NH03802,USA.Tel.:+16033805707;fax:+16038623784.E-mailaddress:(I.F.Potter). JEMBE-49259;NoofPages9 0022-0981/$… ContentslistsavailableatScienceDirectJournalofExperimentalMarineBiologyandEcologyjournalhomepage:www.elsevier.com/locate/jembe Pleasecitethisarticleas:Potter,I.F.,Howell,W.H.,VerticalmovementandbehavioroftheoceansunMolamola,inthenorthwestAtlantic,J.Exp.Mar.Biol.Ecol.(2010),doi:10.1016/j.jembe.2010.10.014 ThepresentstudyisthersttoexaminetheverticalmovementandbehaviorofM.molainthenorthwesternAtlantic.Thestudy'sobjectivesweretoattachpop-upsatellitearchivaltags(PSATs)tooceansunshduringthesummerandautumnoffthecoastofNewEnglandinordertoexaminebehaviorandidentifymovementpatternsofM.molaintheregion.2.Materialsandmethods2.1.TaggingBetween2005and2008,31pop-upsatellitearchivaltags(PSATs:modelPTT-100,MicrowaveTelemetry,Columbia,MD,USA)weredeployedonoceansunM.mola,intheNorthwestAtlantic.Twenty-nineweredeployedduringthesummerandautumnOctober)of2005,2006,or2007.FishweretaggedintheGulfofMaine(n=7)andinshelfwaterstothesouthandeast(within80km)ofNantucketIsland(n=22).Fiveshweretaggedin2005asapilotstudy.Fourteenshweretaggedin2006,and10shweretaggedin2007.TwoadditionalshweretaggedinMarch2008inshelfwatersoffthecoastofGeorgia,USA.PSATswereprepro-grammedtoreleaseeitherveoreightmonthsafterdeployment.Fishweretaggedfromcommercialtunavessels(F/V,F/VTenacious;n=10)andachartershingvessel(F/V;n=19)inthewatersoffthenortheasternUnitedStates,andfromaresearchvesselinGeorgia(R/V,GeorgiaDNR).Fishbaskingatthesurfacewereapproachedbythevessel,andtagswereattachedviaaedwoodenharpoontaggingpole,2.5minlength.Monomenttethers(25cmlong,250-lbtest)wereusedtoattachthePSATstoblacknylonumbrelladarts(dartsdesignedandproducedbyMichaelDomeier,MarineConservationScienceInstitute).Tagswereplacedintothemusculatureapproximately15cmbelowthebaseofthedorsaln,justanteriortotheclavus.Becausetheshwerenotbroughtonboardorretainedduringthetaggingprocedure,measure-mentsofsizeandweightwerenottaken.Basedonobservation,M.molataggedintheregionwereestimatedtoweighapproximately130to230kgwithatotallengthof1.3to1.6m(weightsestimatedusingequationfromWatanabeandSato(2008ThePSATsrecordedambientlightlevels,temperature(resolution0.17°C),anddepth(resolution5.38m).Temperatureanddepthweresampledinmostcasesat15-minintervalsandstoredassummary2.2.HorizontalmovementDailygeolocationpositionestimatesoftheshwereestablishedfromrecordedhigh-resolutionlightdatastoredinthePSATusingaproprietaryalgorithm(MicrowaveTelemetry,Inc.),seasurfacetemperature,andbathymetryusingthemethodsofGaluardietal..ThehorizontalmovementofthesetaggedM.molainthenorthwesternAtlantichasbeenreportedinPotteretal.(inpress)2.3.VerticalmovementDepthsandtemperatureswererecordedbythePSATat15-minintervals.Todetermineifadielpatterninverticalbehaviorexisted,nightdifferencing(meandailydepthmeannightlydepth)wasutilized,andat-testwasappliedtotestfordifferences.Theperiodsofdayandnightweredenedforeachindividualandday,dependingonthelocationoftheshoneachparticulardayandtheresultingdaylength.Depthandtemperaturedataretrievedbythetagswereusedtoconstructdepth/temperatureprolesthatprovidedinformationonthesurroundingwaterma

2 sscharacteristics.Thedepthandtemperature
sscharacteristics.ThedepthandtemperatureprolesbasedonPSATdatawerecomparedtodatafromstationarybuoysintheGulfofMaine(partoftheGoMOOSsystem)toconrmaccuracy,andincombinationwiththecorrectedgeolocationdata,wereusedtoidentifythelocationoftaggedtemperatureplotswerecreatedfromtherawPSATdatainRusingmethodsofGaluardi(unpublished).Forsomeanalyses,theweredividedintotwogroupsnorthandsouthof35°Nlatitude,tonorthern/mid-Atlanticwhentheymovedsouthofabout35°Nlatitude,theirverticalbehaviorchanged.Thoseshdenedasnorthern/mid-Atlanticshnevermovedsouthof35°N(n=12).ThosedeweretaggedinNewEnglandandmovedsouthof35°Natsomepointduringthecourseoftheirtaggingperiod(n=11).Verticalbehaviorofshwasexaminedrelativetowatertemperaturebycomparingdepthdatafromindividualswheninwarmwater24°C)tothatofthesameindividualsinoffshore,coolerwater24°C).ThebaskingbehaviorofM.molawasexaminedbyanalyzing,foreachsh,theamountoftimeperdayspentincoldwater(10°C)andtheamountoftimeduringthatsamedaythatshspentatthesurface(06m).3.ResultsOfthe31PSATsdeployed,25successfullyreporteddatabacktoFig.1showsamapofthedeploymentandendpointlocationsforall25tags.Asummaryofthedatesandlocationsofthestartandendofeachtagdeployment,daysatliberty,distancetraveled,andpercentdatareportedisprovidedinTable1.Tagattachmentperiodsrangedfrom5to242days.Eighteentagswereattachedfor50daysormore,10ofthosewereattachedfor150daysormore,and4ofthemwereattachedforatleast240days.Meanattachmentperiodofallthetagsreportingdatawas107.2±80.6days(SD),andmeandatareportedwas69.7±33.1%(SD)(n=25).Twotagswithdataretrievaloflessthan20%wereremovedfromtheanalysis.Fishspent31.7%oftheirtimeinthetop10mofthewatercolumn,and59.8%oftheirtimeinthetop50m(n=23)(Fig.2A).Averylargeproportionoftotaltime(84.1%)wasspentinthetop200m,andthemaximumdepthrecordedbyanyshwas844m.TaggedM.molaspent24.0%oftheirtimebetween18and20°Cand68.0%oftheirtimebetween14and22°C(Fig.2B).Temperaturesexperiencedbytaggedshrangedfrom6to30°C.Examinationofthebehaviorofsouthernsh(n=11)comparedtothebehaviorofnorthernsh(n=12)showedsouthernshspentmoretimeatdepthandlesstimeatthesurfacetimethannorthernsh(Fig.3A).Southernshspent24.1%oftimeatdepthsgreaterthan200m,whereasshthatstayednorthspent10.6%oftimeatthesedepths,andtwiceasmuchtimeassouthernshinthetop10mofthewatercolumn.Southernshexperiencedwarmerwatertemperaturesthanthosethatremainednorth(Fig.3B),with17.4%oftimespentattemperatures22°Ccomparedto3.3%oftimespentinthesetemperaturesbynorthernsh.Southernshencounteredwarmseasurfacetemperaturesforalongerperiodoftimeinshelfwaters(14°Cto19°CduringthemonthsofNovemberthroughMay)andwereincloserproximitytothewarmwatersoftheGulfStream(26°C)thanshthatremainedinnorthernshelfandoffshorewaters(8to20°CduringthemonthsofAugustthroughMay).Therewasadielpatterninverticalbehavior.TaggedM.molamoretimeatdepthduringthedayandinhabitedshallowerwatersatnight.Meandaytimedepthofallshwas107.8(±21.2)mandmeannightlydepthwas72.0(±12.4)m(n=23).Meandaynightdifferenceofallshwas51.8(±12.0)m(t-test,p=0.004).Thoughshthattraveledsouthexperiencedagreaterrangeofdepth,thedaynightdifferencewasthesameasforshthatremainednorthTable2).Oceansunshchangedtheirdivebehavioroverthecourseofthetaggingperiod,fromshallowdiveswithahighproportionofsurfacetime(06m),todeeperdivesandalowerproportionofsurfacetime.Meandailydepthofeachindividual(meandepthofduringdaylighthours)increased,asdidthemeandaynightdifference(Fig.4I.F.Potter,W.H.Howell/JournalofExperimentalMarineBiologyandEcologyxxx(2010)xxx Pleasecitethisarticleas:Potter,I.F.,Howell,W.H.,VerticalmovementandbehavioroftheoceansunMolamola,inthenorthwestAtlantic,J.Exp.Mar.Biol.Ecol.(2010),doi:10.1016/j.jembe.2010.10.014 VerticalbehaviorofM.molachangedastheshtraveledsouth.WheninNewEnglandshelfwaters,taggedshexhibitedshallow,frequentdivingtodepthsof50100mwithahighpercentageofsurfacetime.Asidefromtheinitialfewdaysaftertaggingshinshallowinsho

3 rewaters,bottomdepthdidnotlimitdivedepth
rewaters,bottomdepthdidnotlimitdivedepth.Asmovedofftheshelftothesouthandeasttheychangedboththedepthandfrequencyofdiving.PlotsofthedepthdataofindividualshwithconcurrenttemperaturedatashowM.molamakingregulardescentsbelowthethermoclineintocoolerwaters.Asshmovedintoawatermasswithadeepermixedlayertheyadjustedtheirverticalbehaviortodivebelowit(Fig.5B).ThiswasespeciallyapparentwhenenteredtheGulfStream,astheyspentlittletimeatthesurfaceanddovetodepthsof400600m.Insomecases,shspentaperiodofdaysatdepthwithnosurfaceinterval.Themeanhourlydepthoffourshinwarm,GulfStreamwater(seasurfacetemperature24°C)wascantlydeeper(150400m)thanthemeanhourlydepthoftheshwhenincooler,off-shelfwater(1535m)(repeatedmeasuresANOVA,pFig.6Therewasnoobservedrelationshipbetweentheamountoftimeperdayshspentincoldwater(10°C)andtheamountoftimeshspentatthesurface(06m)inthenexttimeinterval(Fig.7).Atthebeginningofthetaggingperiod,taggedoceansunshspentlittletonotimeincoldwater,andspentahighproportionoftheirtimeatthesurface.Asthetaggingperiodprogressed,shspentmoretimeperdayincoldwater,andlittletonotimeatthesurface.Therewasalsonoobservedrelationshipbetweentheamountoftimeperdayshspentincoldwater(10°C)andtheamountoftimeshspentinwarmwater20°C).Becausedepthdatawerelimitedto15-minuteintervals,individualascentanddescentrateswerenotabletobedetermined,howeverapreviousstudyofjuvenile(small)oceansunshusingacoustictagsfoundthetypicalrateofdescenttobe8.4±5.6m/minCartamilandLowe,2004).Basedonthisinformation,itisunlikelythattaggedoceansunshweresurfacingfromdepths200minbetweenthe15-minuteintervalswhendepthwasrecordedbythetag.Twoofthe25sunshtaggedenteredtheGulfofMexico.OnetaggedinSeptember2005inNewEnglandreachedtheGulfofMexicoinDecemberbeforethetagdetached.ThesecondshwastaggedinGeorgiashelfwatersinMarch2008,traveledsouthtotheGulfofMexicoinJune,andremainedthereuntilearlyJuly.Thoughdatareturnfromthesetagswaslimited(22%and33%),analysisofdatafromthesecondshshowsthat,likeotherindividualstaggedinthestudy,theshshowedadielpatterninitsdivingbehavior,withameandaynightdifferenceof23.2m.OnceintheGulfofMexico,theshaltereditsdivingbehaviortogreaterdepthsandshortersurfacetimeswhencomparedtoitsbehaviorinSoutheastUSshelfwatersFig.8).PriortoenteringtheGulfofMexico,inthesoutheastUS,theshspent69%ofitstimeinthetop25mofthewatercolumn,and90%ofitstimeinthetop200m.OnceintheGulfofMexico,theshspentlessthan3%oftimeinthetop10mofthewatercolumn,90%ofitstimebetween25and200m,andhadoccasionaldivesto400m.While Fig.1.Mapofdeploymentlocations(graycircles)andendpoints(blacktriangles)of25PSATsattachedtoMolamolabetween2005and2008.TagswereattachedinNewEnglandwatersbetweenthemonthsofAugustandOctoberin20052007(n=23)andinGeorgiashelfwatersinMarch2008(n=2).Tagswereattachedfrom5to242days.I.F.Potter,W.H.Howell/JournalofExperimentalMarineBiologyandEcologyxxx(2010)xxx Pleasecitethisarticleas:Potter,I.F.,Howell,W.H.,VerticalmovementandbehavioroftheoceansunMolamola,inthenorthwestAtlantic,J.Exp.Mar.Biol.Ecol.(2010),doi:10.1016/j.jembe.2010.10.014 intheGulfofMexico,theshexperiencedtemperaturesrangingfrom8.3°Cto28°C,withanaveragewatertemperatureof18.23(±4.96)°C.4.DiscussionPSAT-taggedsunshexhibitedawiderangeofverticalbehavior,whichvariedoverbothspatialandtemporalscales.WefoundthatoceansunshinnortheasternUSwatersinthesummerandearlyautumnmonthsgenerallyinhabitedrelativelyshallowdepths(often10m),andwarmtemperatures(1720°C).Thewatercolumninthisregionatthistimeistypicallystratied,withastrongthermoclineandshallowmixedlayer(LalliandParsons,2006;MannandLazier,2006spentahighproportionoftheirtimeabovethethermoclinenear Table1Dateofdeployment,locationofdeploymentandpop-off,numberofdaysattached,straightlinedistance(SLD)traveled,andretentionrateof31PSATsattachedtobetween2005and2008.Deployment(°N)(°W)(°N)(°W)145389/3/200541.3769.3123.2185.73130252022%877610/2/200542.0470.0240.1968.9324222100%673

4 228/17/200643.1569.9831.3579.29815315655
228/17/200643.1569.9831.3579.298153156555%673258/18/200640.9670.5132.2378.98153120646%673218/23/200640.8270.6626.9479.94143168564%673319/1/200641.2969.9639.4274.211341582%673329/8/200642.2470.0327.1878.3374183096%673309/17/200641.2669.2838.8671.1924231240%675659/17/200641.2969.2433.8976.30242102218%673279/19/200641.2069.5932.3878.71153125031%673339/19/200641.1969.5930.4681.4124015904%673239/23/200641.1769.5832.5678.37153123551%675649/25/200641.3169.9939.9270.1714148100%673289/25/200641.3069.9728.2879.4470169078%770739/7/200741.3970.0541.4970.12530100%770799/7/200741.2969.9537.3073.179852784%770709/14/200741.2969.9533.5277.7568110592%770759/20/200741.2769.9839.6374.1031419100%770779/24/200741.1370.0341.2370.621150100%770699/24/200741.2170.1438.6467.7615333873%7707110/3/200741.2670.2240.3773.339283100%7707210/6/200742.6370.3736.5575.645079396%6732610/6/200742.3870.2736.7471.6415264572%770743/11/200830.7980.7631.7480.175711064%770783/11/200830.9080.6632.5178.3824228033% Fig.2.B.Histogramsshowingproportionoftimeatdepth(A),andproportionoftimeattemperature(B),forallsh(n=23).FishweretaggedinNewEnglandwatersbetweenthemonthsofAugustandOctoberin20052007andinGeorgiashelfwatersinMarch2008.Tagswereattachedfrom7to242days. Fig.3.B.Histogramsshowingproportionoftimeatdepth(A),andtimeattemperature(B),foroceansunshthatremainedabove35°Nandshthattraveledbelow35°N.FishweretaggedinNewEnglandwatersbetweenthemonthsofAugustandOctoberin20052007.Fishthatremainedabove35°Nhadattachmentperiodsfrom7to242days(mean=67days;n=12).Fishthattraveledbelow35°Nhadattachmentperiodsfrom68to153days(mean=127days;n=11). Table2Meandaytimedepth,meannightlydepth,andmeandaynightdifference(meandailymeannightlydepth),for23taggedoceansunsh,dividedbylocation:northern(above35°N)andsouthern(below35°N).p-ValuesindicateresultsofapairedNTotalofdaysdepth(m)depth(m)difference(m)1280267.4(±23.6)26.9(±7.01)45.3(±18.2)0.055111396150(±32.2)101.8(±15.4)45.2(±18.8)0.047sh232198107.8(±21.2)72.0(±12.4)51.8(±12.0)0.004I.F.Potter,W.H.Howell/JournalofExperimentalMarineBiologyandEcologyxxx(2010)xxx Pleasecitethisarticleas:Potter,I.F.,Howell,W.H.,VerticalmovementandbehavioroftheoceansunMolamola,inthenorthwestAtlantic,J.Exp.Mar.Biol.Ecol.(2010),doi:10.1016/j.jembe.2010.10.014 thesurface,butdovefrequentlytodepths200m.Interestingly,theobservedverticalbehaviorof(shallow,shortdives)issimilartotheverticalfeedingbehavioroftheleatherbackturtle,whichisalsofoundinthisareaduringthesametimeofyear(Jamesetal.,2005;Eckert,2006).Likeleatherbacks,oceansunsharebelievedtofeedprimarilyongelatinouszooplankton,andthedistributionofM.molahasbeenassociatedwithareaswherejellyshareabundantDesjardin,2005;Houghtonetal.,2006).SimilaritiesinverticalbehavioranddietsofthetwospeciessuggestthatM.molaisforaginginnortheasternUSshelfandslopewaters,takingadvantageofthehighconcentrationsofgelatinouspreyfoundintheregion(ShoopandKenney,1992;Madinetal.,2006).ArecentstudyonforagingpatternsofleatherbacksrelativetopreydistributionnotedsimilardivingbehaviorinturtlesinthenorthwesternAtlanticduringthesummermonths.Meandivedepthsofturtlesintheregionwereshallowerthanwheninsouthernwaters,whichtheauthorsassociatedwithhighforagingsuccess.Thispatternmatchesthatofzooplanktondistributionintheregion,suggestingaco-occurringabundanceingelatinousprey(Fosetteetal.,2010Oceansunshthatmovedsouthintothewatersofthemid-AtlanticandsoutheasternUnitedStatesinthewinterandearlyspringspentlesstimeatthesurfaceandmoretimeatdepths200m.Thisshiftinverticalbehaviorwasassociatedwithamovetowatersthataretypicallyweaklystratiedandhaveadeepmixedlayer(etal.,1999).Itisprobablethattheoceansunshadjustedtheirverticalrangetofollowthedeeperthermalgradient.BecauseM.molaareoftenlocatedwith,andpresumablyfeedongelatinouszooplank-ton(Desjardin,2005;Houghtonetal.,2006),andbecausegelatinouszooplanktonexhibitdielverticalmigrations(DV

5 M)thatrangefromafew,tohundredsofmeters(G
M)thatrangefromafew,tohundredsofmeters(Grahametal.,2001;Madinetal.,2006;HosiaandBamstedt,2008),itisalsoprobablethattheverticalmovementsofaretiedtolocatingtheirverticallymigratinggelatinousprey.AstudyonoceansunshinthePacic,aswellasstudiesofotherlargepelagicspeciessuchasbluentunaandbaskingsharksinboththePacicandNorthAtlantic,hasnotedsimilardifferencesinverticaldistributionassociatedwithwatercolumnstructureonaseasonalscale(Lutcavageetal.,1997;Kitagawaetal,2000;Simsetal.2003;Thys,2003;Wilsonetal.,2005).Likeoceanshinthepresentstudy,leatherbackturtlesinwatersoffthesoutheasternUnitedStatesdivedeeply,forlongperiodsoftime,toforageinthedeepscatteredlayerorDSL(Eckert,2006).ArecentstudycomparingtheverticalnicheoverlapofbothspeciesinwatersoffSouthAfricasuggestedthatlikeleatherbacks,oceansunshexhibitplasticityintheirverticalbehaviorinresponsetopatchilydistributedgelatinouspreywithdifferinglevelsofdielverticalmigration.Bothspeciesexperienceashiftfrompredationonseasonallyabundantsurfacemedusaduringthesummertoverticallymigratinggelatinousspeciesfoundatdeeperdepthsduringthewintermonths(Haysetal.,2009;Popeetal.,2010TherewasadielpatterninverticalbehaviorofM.molavisiteddeeperdepthsduringthedaylighthoursandremainedinshallowerwatersatnight.Thedielpatternhasbeenobservedinotherstudiesofthespecies(CartamilandLowe,2004;Thysetal.,2007;Simsetal.,2009),inotherlargepelagicshes(baskingsharks,bluesharks,swordsh,andtunas)(CareyandScharold,1990;Blocketal.,2001;Simsetal.2003),andinleatherbackseaturtles(Eckertetal.,).Inthepresentstudy,oceansunshmaintainedthesedistinctdifferencesinverticalbehaviorevenastheiroveralldepthproshiftedtodeeperwatersovertime,whichreectsthepreysearchingandforagingbehaviorofthespeciesindifferentwatermasses.TheshiftinverticalbehaviorfromshallowtodeeperdepthswasespeciallyapparentwhentaggedsunshenteredtheGulfStream.shspentlittletimeatthesurfaceanddovetodepthsof400800m,andinsomecases,spentaperiodofdaysatdepthwithnosurfaceinterval.TheGulfStreameldextendsdeepintothewatercolumnwherenutrientconcentrationsaremaximized(andLozier,2008),andamigratoryDSLhasbeendetectedintheGulfStreamwithdiurnalmigrationtodepthsof450m(Coleetal.,1971ThedeepdivingexhibitedbyoceansunshintheGulfStreamislikelyareectionofpreysearchingbehavior.Inadditiontothewarmseasurfacetemperaturesandadeeplymixedwatercolumn,swiftnorthwardmovingsurfacecurrentsmayhavemotivatedsouthwardshtoremainatdepthandavoidthesurfacecurrents.Inadditiontoashifttoincreaseddepth,oceansunshalsoalteredthefrequencyofverticalbehavior,shiftingfromhigh-frequencydivingandsurfacingtolowfrequencydivingoftenwithseveralconsecutivedaysatdepthwithnosurfaceintervals.Thehigh-frequencydivingwithextensivemovementinthewatercolumnlikelyreectspreysearchingbehavior,andhasbeenobservedinanotherstudyofM.molaintheeasternAtlantic(Simsetal.,2009).AstudyexaminingtheverticalmovementofM.mola,amongothermarinepredators,foundevidencethatthespeciesadaptsitsvertical Fig.4.C.Meandailydepth(A),meannightlydepth(B),andmeandaydifference(C),ofPSATID67328.Theshwastaggedon9/25/06offNantucketIsland.Thetagwasattachedfor70daysandpoppedoffwhentheshwasinoffshorewatersoffFloridaon12/7/06.I.F.Potter,W.H.Howell/JournalofExperimentalMarineBiologyandEcologyxxx(2010)xxx Pleasecitethisarticleas:Potter,I.F.,Howell,W.H.,VerticalmovementandbehavioroftheoceansunMolamola,inthenorthwestAtlantic,J.Exp.Mar.Biol.Ecol.(2010),doi:10.1016/j.jembe.2010.10.014 Fig.5.B.Depthandcorrespondingtemperaturedatafrom:A)sh67321;andB)sh67332.Fish67321wastaggedon8/23/06inwatersoffNewEngland.Thetagwasattachedfor143daysandpoppedoffwhentheshwasaroundtheBahamason1/23/07.Fish67332wastaggedon9/8/2006intheGulfofMaine.Thetagwasattachedfor74daysandcameoffwhentheshwasaroundtheBahamason11/25/06.I.F.Potter,W.H.Howell/JournalofExperimentalMarineBiologyandEcologyxxx(2010)xxx Pleasecitethisarticleas:Potter,I.F.,Howell,W.H.,Verticalmoveme

6 ntandbehavioroftheoceansunMolamola,inthe
ntandbehavioroftheoceansunMolamola,inthenorthwestAtlantic,J.Exp.Mar.Biol.Ecol.(2010),doi:10.1016/j.jembe.2010.10.014 behavioroverabroadrangeofdepthstooptimizeitssearchforpatchilydistributedpreyindifferentenvironments(Humphriesetal.,).TheretinaltopographyofimmatureM.molawasexaminedinarecentstudy,andresultssuggestedthatoceansunshdetecttheirpreyduringtheirdescentthroughthewatercolumn,aswellasontheoor(Kinoetal.,2009).Thefrequentdescentsthroughthewatercolumnobservedinoceansunshinthepresentstudyarepresumablypreysearchingbehavior.IthasbeensuggestedthatM.molabasksatthesurfaceasamechanismofthermalrechargingafterdeepdivesincoldwaterThys,2003;CartamilandLowe,2004).Inthepresentstudytherewasnostatisticalrelationshipbetweentheamountoftimeperdayspentincoldwater(10°C)andtheamountoftimeshspentatthesurface(06m).TaggedoceansunshspentalargeproportionoftimeatthesurfaceinNortheastUSwatersinthelatesummerandearlyautumnwherethewaterwashighlystratiedwithashallowmixedlayerandthewarmesttemperaturesatthesurface.BaskingatthesurfaceintheshelfwatersoffthenortheasternUnitedStateshasalsobeenobservedinswordXiphiasgladius,andleatherbackseaturtles,andisbelievedtobepartofarecoveryprocesstoaidindigestionandwarmbodytemperatures(CareyandRobinson,1981;Eckertetal.,1989).IfM.molaisalsofeedingonthelargequantitiesofgelatinouszooplanktonintheregion,itmayequallybenetfromrecoverytimeatthesurface.Assunshmovedsouthandencounteredwarmer,mixedwatermasses(particularlytheGulfStream)theydovedeeperand,ifseekingarecoveryperiodaftertimeatdepth,mayhavegainedthermalbenetsfromthewarmtemperaturesfoundatmid-waterdepths.ThisbehaviorhasbeenobservedinswordshintheSoutheastPacicafterfeedingonorganismsintheDSL(Abascaletal.,).Baskingbehaviorinoceansunshhasalsobeensuggestedtofacilitaterecoveryafterdivingintotheoxygenminimumlayer,ashasbeenobservedinotherlargepelagicshes(CareyandRobinson,1981;CartamilandLowe,2004;Thysetal.2007).GiventhatthenorthwesternAtlanticiswelloxygenatedatdepth,baskingbehaviormaybeobservedwithmorefrequencyinpopulationsinhabitingoceanswhereoxygenminimumzonesoccur.ThelargesizeofM.molainthepresentstudymayalsoplayaroleinbaskingbehaviorifrelatedtothermaltolerance,aslargershmaybebufferedagainsttemperaturedifferencesmoresothansmallersh.Theimportanceoflargebodymassinmaintainingelevatedbodytemperaturesrelativetotheenvironmenthasbeensuggestedforseaturtlesandntuna(Satoetal.,1994;Kuboetal.,2008BaskingbehaviormayalsobeneM.molaasamechanismofparasiteremoval.M.molaisknowntoharborupto40differentgeneraofparasites,andbaskingatthesurfaceislikelytoattractcleaningfrombelowaswellasseabirdsfromabove,bothofwhichhavebeenreportedlyassociatedwiththespecies(CartamilandLowe,2004;Popeetal.,2010shinthestudyenteredtheGulfofMexicoduringdifferenttimesofyear:winter(DecemberJanuary)andsummer(JuneM.molaisayear-roundresidentofthenorthernGulfofMexicowiththegreatestdensitiesreportedinthewinterandspring(Desjardin,2005;Fullingetal.,2007).TheshthatenteredtheGulfofMexicoduringsummerencounteredseasurfacetemperaturesapproaching30°C,warmerthanpreviousstudiessuggestisthethermalpreferenceofthespecies(Thysetal.2007;Fullingetal.,2007;Simsetal.,2009WhileintheGulfofMexico,shspentlittletimeatthesurface,moretimeatdepth,andremainedincloseproximitytotheshelfandLoop 0612182306121823 Fig.6.B.Meanhourlydepths(±SD)offourtaggedoceansunshduringaone-weekperiodwhentheywereinwarmoffshorewater(24°C)(A),andaone-weekperiodincooleroffshorewater(24°C)(B).TimeisinGMT. Fig.7.Dailysurfacetime(proportionoftimeat06mper24h)vs.timeincoldwater(proportionoftimeintemperatures10°Cper24h)forfouroceansunsh(IDs:67321,67332,67328,and67322). Fig.8.HistogramshowingproportionoftimeatdepthforoceansunshID77078whenitwasintheGulfofMexico(n=28days)andinsoutheastUSwaters(n=92days).Theshwastaggedfrom3/11/2008to11/11/2008.I.F.Potter,W.H.Howell/JournalofExperimentalMarineBiologyandEcologyxxx(2010)xxx Pleasecit

7 ethisarticleas:Potter,I.F.,Howell,W.H.,V
ethisarticleas:Potter,I.F.,Howell,W.H.,VerticalmovementandbehavioroftheoceansunMolamola,inthenorthwestAtlantic,J.Exp.Mar.Biol.Ecol.(2010),doi:10.1016/j.jembe.2010.10.014 Current.Similarbehaviorwasnotedinasharptailmola,,taggedwithaPSATinthenorthernGulfofMexico(etal.,2002).ThedeepthermoclineintheGulfofMexicowasalikelyfactorintheshiftinverticalbehaviorofM.molatogreaterdepthswhenitenteredtheregion.Ofthe31PSATsdeployedonM.molabetween2005and2008,11remainedattacheduntiltheprogrammedpop-update.Sixneverreporteddata,and14detachedprematurely.Onereasonforprematuredetachmentwastheactivationofthetag'sfail-safemechanismthatcauseditsreleaseduetowhatwasperceivedbythetagasconstantpressure(i.e.,thechangeinthesh'sdepthwaslessthantheresolutionofthetag(5.38m)overafour-dayperiod)whichoccurredinsixtags,andinallofthesecasestheshshowedsteadyhorizontalmovementinshelfwaters.Thisoftenoccurredshwereinshallowwaters(10m)aroundNantucketIslandwheretheypresumablywerefeedingonlargepatchesofgelatinousprey(salpsandjellysh)observedinthearea.Lackofanyhorizontaldisplacementfromoneprematurelyreleasedtagindicatesthedied,presumablyasaresultofbeingtagged.Otherpossiblecausesofprematuredetachmentincludetagshedding/tissuerejectionduetoimproperplacementofthetag,corrosionorfoulingofthetag,andmechanicalfailure.Arecentpaperaddressingtheissueofsignallossinsatellitetrackingstudiesidentiedantennaebreakage,exhaustionofbatteries,tagmalfunctionandanimalmortalityamongthecausesofsignalloss,andsuggestedmorediagnosticdatabeincorporatedintotagtransmissionsinordertobetterunderstandthefactorscontributingtothisproblem(Haysetal.,2007ThepresentstudyisthemostextensivetodateonthemovementandbehaviorofM.mola.ResultsestablishabaselineofmovementandbehaviordatafromoceansunshintheNorthwestAtlantic,andprovideearlyinsightsintotheunderstandingofitshabitatprefer-ences,migrationroutes,behavior,andenvironmentalassociations.DataondistributionandseasonalmovementsofM.molamayproveusefulindicatorsofnutrient-richareaswithhighproductivity,whereothermarineorganismsofimportantcommercialvalueorendan-geredstatusarefound.FuturestudiesareneededtogainamorecompleteunderstandingofthemovementandbehaviorpatternsofthespeciesintheNorthwestAtlanticandotherpartsoftheworld.Inadditiontostudiesonmovementandbehavior,studiesonsuchbasicbiologicalinformationasage,growth,andreproductionarenecessarytothoroughlyexaminethebiologyofthespeciesanditsroleintheNorthAtlanticecosystem.AcknowledgementsWearegratefultoJoshEldridgeoftheF/V,DaveLinneyoftheF/V,EricHesseoftheF/V,andWaltGoletfortheirassistancewiththisproject.WealsothankMollyLutcavage,BenGaluardi,FrancoisRoyer,KaraDodge,WinWatson,TierneyThys,RobertKenney,GregSkomal,andMarkBaumgartnerfortheircontributions.ThisresearchwassupportedbyagrantfromtheLargePelagicsResearchCenterattheUniversityofNewHampshire(toWHH),andafellowshipawardedfromtheUNHMarineProgram(toIFP).PilotresearchfortheprojectwasfundedinpartbyNewHampshireSeaGrantDevelopmentFunds.Funds.ReferencesAbascal,F.,Mejuto,M.,Quintans,M.,Espino,D.,2009.PopuptaggingofswordXiphiasgladius)intheSEPacic.AbstractinIn:Nielson,J.L.,Arrizabalaga,H.,Fragoso,N.,Hobday,A.,Lutcavage,M.,Sibert,J.(Eds.),TaggingandTrackingofMarineAnimalswithElectronicDevices:InSeries:Review:MethodsandTechnologiesinFishBiologyandFisheries,Volume9.400pp.Bass,A.L.,Dewar,H.,Thys,T.,Streelman,J.T.,Karl,S.A.,2005.Evolutionarydivergenceamonglineagesoftheoceansunshfamily,Molidae(Tetraodontiformes).MarineBiology148(2),405Block,B.A.,Dewar,H.,Blackwell,S.,Williams,T.,Prince,E.,Farwell,C.,Boustany,A.,Teo,S.,Seitz,A.,Walli,A.,Fudge,D.,2001.Migratorymovements,depthpreferences,andthermalbiologyofAtlanticbluentuna.Science293(5533),1310Carey,F.G.,Robinson,B.H.,1981.Dailypatternsintheactivitiesofsword,observedbyacoustictelemetry.FisheryBulletin79,277Carey,F.,Scharold,J.,1990.Movementsofbluesharks(Prionaceglauca)indepthandcourse

8 .MarineBiology106(3),329Cartamil,D.P.,Lo
.MarineBiology106(3),329Cartamil,D.P.,Lowe,C.G.,2004.DielmovementpatternsofoceansunMolamolasouthernCalifornia.MarineEcologyProgressSeries266,245Cole,H.P.,Bryan,G.M.,Gordon,A.L.,1971.Thedeepscatteringlayer:patternsacrosstheGulfStreamandtheSargassoSea.In:Farquhar,G.B.(Ed.),ProceedingsofanInternationalSymposiumonBiologicalSoundScatteringintheOcean.MauryCenterforOceanScience,Washington,D.C,pp.281Desjardin,N.A.2005.Spatial,temporal,anddietaryoverlapofleatherbackseaturtlesDermochelyscoriacea)andoceansunshes(FamilyMolidae).Master'sthesis,FloridaAtlanticUniversity.Eckert,S.,2006.HighuseoceanicareasforAtlanticleatherbackseaturtles()asidentiedusingsatellitetelemeteredlocationanddiveinformation.MarineBiology149,1257Eckert,S.,Eckert,K.,Ponganis,P.,Kooyman,L.,1989.Divingandforagingbehaviorofleatherbackseaturtles(Dermochelyscoriacea).CanadianJournalofZoology67(11),2834Fosette,S.,Hobson,V.,Girard,C.,Calmettes,B.,Gaspar,P.,Georges,J.,Hays,G.,2010.Spatio-temporalforagingpatternsofagiantzooplanktivore,theleatherbackturtle.JournalofMarineSystems81,225Fraser-Bruner,A.,1951.Theoceansunshes(FamilyMolidae).BulletinoftheBritishMuseumofNaturalHistory1,89Fulling,G.,Fertl,D.,Knight,K.,Hoggard,W.,2007.DistributionofMolidaeinthenorthernGulfofMexico.GulfandCaribbean19(2),53Galuardi,B.,Royer,F.,Golet,W.,Neilsen,J.,Lutcavage,M.,2009.ComplexmigrationtracksfromAtlanticbluentunataggedoffNovaScotia,Canada.AbstractinIn:Nielson,J.L.,Arrizabalaga,H.,Fragoso,N.,Hobday,A.,Lutcavage,M.,Sibert,J.(Eds.),TaggingandTrackingofMarineAnimalswithElectronicDevices:InSeries:Review:MethodsandTechnologiesinFishBiologyandFisheries,Volume9.400pp.Graham,W.,Pages,F.,Hamner,W.,2001.Aphysicalcontextforgelatinouszooplanktonaggregations:areview.Hydrobiologia451(13),199Gregory,W.K.,Raven,H.C.,1934.Notesontheanatomyandrelationshipsoftheoceansh(Molamola).Copeia4,145Hays,G.,Bradshaw,C.,James,M.,Lovell,P.,Sims,D.,2007.WhydoArgossatellitetagsdeployedonmarineanimalsstoptransmitting?JournalofExperimentalMarineBiologyandEcology349,52Hays,G.,Farquhar,M.,Luschi,P.,Teo,S.,Thys,T.,2009.Verticalnicheoverlapbytwooceangiantswithsimilardiets:oceansunshandleatherbackturtles.JournalofExperimentalMarineBiologyandEcology370(12),134Hosia,A.,Bamstedt,U.,2008.SeasonalabundanceandverticaldistributionofsiphonophoresinwesternNorwegianfjords.JournalofPlanktonResearch30(8),Houghton,J.,Doyle,T.,Wilson,M.,Davenport,J.,Hays,G.,2006.Jellyshaggregationsandleatherbackturtleforagingpatternsinatemperatecoastalenvironment.Ecology87(8),1967Humphries,N.,Queiroz,N.,Dyer,J.,Pade,N.,Musyl,M.,Schaefer,K.,Fuller,D.,Brunnschweiler,J.,Doyle,T.,Houghton,J.,Hays,G.,Jones,C.,Noble,L.,Wearmouth,V.,Southall,E.,Sims,D.,2010.EnvironmentalcontextexplainsLévyandBrownianmovementpatternsofmarinepredators.Nature.James,M.,Eckert,S.,Myers,R.,2005.Migratoryandreproductivemovementsofmaleleatherbackturtles(Dermochelyscoriacea).MarineBiology147,845Kenney,R.D.,1996.Preliminaryassessmentofcompetitionforpreybetweenleatherbackseaturtlesandoceansunshinnortheastshelfwaters.In:Keinath,J.A.,Bernard,D.E.,Musick,J.A.,Bell,B.A.(Eds.),ProceedingsoftheFifteenthAnnualSymposiumonSeaTurtleBiologyandConservation.NOAATechnicalMemorandumNMFS-SEFSC-387,NationalMarineFisheriesService,Miami,FL,pp.144147.Kino,M.,Miayzaki,T.,Iwami,T.,Kohbara,J.,2009.RetinaltopographyofganglioncellsinimmatureoceansunMolamola.EnvironmentalBiologyofFishes85,Kitagawa,T.,Nakata,H.,Kimura,S.,Itoh,T.,Tsuji,S.,Nitta,A.,2000.EffectofambienttemperatureontheverticaldistributionandmovementofPacicbluentunaThunnusthynnusorientalis.MarineEcologyProgressSeries206,251Kubo,T.,Sakamoto,W.,Murata,O.,Kumai,H.,2008.Whole-bodyheattransfercientandbodytemperaturechangeofjuvenilePacicbluentunaaccordingtogrowth.FisheriesScience74(5),995Lalli,C.,Parsons,T.,2006.BiologicalOceanography:AnIntroduction.ElsevierButter-worth-Heinemann.314pp.January6Lovgren,S.

9 ,2004.Emergingexplorerhookedonmysterious
,2004.Emergingexplorerhookedonmysteriousleviathan.NationalGeographicNewshttp://news.nationalgeographic.com/news/2004/01/0126_040126_tierneythysmola.html2004NationalGeographicSocietyWebsite.Lutcavage,M.,Kraus,S.,Hoggard,W.,1997.Aerialsurveyofgiantbluentuna,,intheGreatBahamaBank,StraitsofFlorida,1995.FisheryBulletin95,Macias,D.,delaSerna,J.M.,2002.BycatchcompositioninSpanishMediterraneanlonglineshery.198pp.ProceedingsoftheFourthMeetingoftheEuropeanElasmobranchAssociation.Societyfrancaised'Icthyologie,Paris,France.Madin,L.,Kremer,P.,Wiebe,P.,Purcell,J.,Horgan,E.,Nemazie,D.,2006.PeriodicswarmsofthesalpSalpaasperaintheslopewaterofftheNEUnitedStates.DeepSeaResearch53(5),804Mann,K.H.,Lazier,J.,2006.DynamicsofMarineEcosystems:BiologicalInteractionsintheOceans.BlackwellPublishing,Massachusetts.496pp.I.F.Potter,W.H.Howell/JournalofExperimentalMarineBiologyandEcologyxxx(2010)xxx Pleasecitethisarticleas:Potter,I.F.,Howell,W.H.,VerticalmovementandbehavioroftheoceansunMolamola,inthenorthwestAtlantic,J.Exp.Mar.Biol.Ecol.(2010),doi:10.1016/j.jembe.2010.10.014 Norman,J.R.,Fraser,F.C.,1949.FieldBookofGiantFishes.GPPutnamandSons,NewPalter,J.,Lozier,M.S.,2008.OnthesourceofGulfStreamnutrients.JournalofGeophysicalResearch(COceans)113(C6).Pope,E.,Hays,G.,Thys,T.,Doyle,T.,Sims,D.,Queiroz,N.,Hobson,V.,Kubicek,L.,Houghton,J.,2010.ThebiologyandecologyoftheoceansunMolamolareviewofcurrentknowledgeandfutureresearchperspectives.ReviewsinFishBiologyandFisheries.doi:10.1007/s11160-009-9155-9Potter,I.,Howell,H.,Galuardi,B.,inpress.Horizontalmovementofoceansun,inthenorthwestAtlantic.MarineBiology.Sato,K.,Sakamoto,W.,Matsuzawa,Y.,Tanaka,H.,Naito,Y.,1994.Correlationbetweenstomachtemperaturesandambientwatertemperaturesinfree-ranginglogger-headturtles,Carettacaretta.MarineBiology118,343Seitz,A.C.,Weng,K.C.,Boustany,A.M.,Block,B.A.,2002.BehaviourofasharptailmolaintheGulfofMexico.JournalofFishBiology60,1597Shoop,C.R.,Kenney,R.D.,1992.SeasonaldistributionandabundancesofloggerheadandleatherbackseaturtlesinwatersoffthenortheasternUnitedStates.HerpetologicalMonographs6,43Silvani,L.,Gazo,M.,Aguilar,A.,1999.SpanishdriftnetshingandincidentalcatchesinthewesternMediterranean.BiologicalConservation90,79Sims,D.,Southall,E.,Richardson,A.,Reid,P.,Metcalfe,J.,2003.Seasonalmovementsandbehaviorofbaskingsharksfromarchivaltagging:noevidenceofwinterhibernation.MarineEcologyProgressSeries248,187Sims,D.,Queiroz,N.,Doyle,T.,Houghton,J.,Hayes,G.,2009.Satellitetrackingoftheworld'slargestbonysh,theoceansunsh(Molamola)intheNorthEastAtlantic.JournalofExperimentalMarineBiologyandEcology370(12),127Thys,T.,2003.TrackingoceansunMolamolawithpop-upsatellitearchivaltagsinCaliforniawatersOceansunsh.org.Oceansunsh.orgWebsitehttp://www.ocean-Thys,T.,2005.Creator/Editor.Oceansunsh.org.WorldWideWebelectronichttp://www.oceansun2005version(2002Thys,T.,Weng,K.C.,Dewar,H.,Farwell,C.,O'Sullivan,J.,Walli,A.,Teo,S.,Tobayama,T.,Soichi,M.,Kondo,Y.,Okada,Y.,Nakatsubo,T.,Block,B.A.,2007.Trackingtheworld'slargestjellypredator,theMolamola,intheEasternandWesternPacic,CLIOTOPSymposium,BajaCalifornia.Watanabe,Y.,Sato,K.,2008.Functionaldorsoventralsymmetryinrelationtolift-basedswimmingintheoceansunMolamola.PLoSONE3,e3446.Werner,F.,Blanton,B.,Quinlan,J.,LuettichJr.,R.,1999.PhysicaloceanographyoftheNorthCarolinacontinentalshelfduringthefallandwinterseasons:implicationsforthetransportoflarvalFish.Oceanography8(2),7Wilson,S.G.,Lutcavage,M.E.,Brill,R.W.,Genovse,M.P.,Cooper,A.B.,Everly,A.W.,2005.MovementsofblueThunnusthynnus)inthenorthwesternAtlanticOceanrecordedbypop-upsatellitearchivaltags.MarineBiology146,409I.F.Potter,W.H.Howell/JournalofExperimentalMarineBiologyandEcologyxxx(2010)xxx Pleasecitethisarticleas:Potter,I.F.,Howell,W.H.,VerticalmovementandbehavioroftheoceansunMolamola,inthenorthwestAtlantic,J.Exp.Mar.Biol.Ecol.(2010),doi:10.1016/j.jembe.2010.10.0