BONUS BAMBI project has received funding from BONUS (Art 185), funded jointly by the EU and the national funding agencies Swedish research agency Formas, Estonian Research Council ETAg, German Forschungszentrum Jülich Beteiligungsgesellschaft, and the Academy of Finland.

BONUSBAMBIPROJECT(1.1.2014to31.12.2017)

ThefinalpublishablesummaryreportFebruary28th2018

Illustration:JLokrantz/Azote

Projectwebsite:www.bambi.gu.se

BONUSBAMBI

FinalpublishablesummaryreportFebruary28th2018

2

ProjectoutlineofgoalsandresultsenvisagedatthebeginningoftheprojectcycleArapidlywarmingBalticSeathatwillalsobeimpactedbylowersalinitiesthantodaywillputaheavypressureonallspecieswithmarineorigincurrentlylivingintheBalticSea.Alreadytoday,theBalticSeaisextremelyspeciespoorwithroughlyonly10percentofthenumberofspeciesfoundintheNorthSea.

TheoverallgoalofBONUSBAMBIwastogeneratenewknowledgeongeneticbiodiversityofBalticSeapopulations,inordertounderstandhowtheongoingchangesintemperatureandsalinitywillaffectkeymarinespeciesoftheBalticSea.Theprojectalsoaimedtoassessthedegreeofmanagementthatiscurrentlydirectedtowardsgeneticdiversity.Inaddition,thegoalwastocommunicatenewinsightsongeneticbiodiversitytoBalticSeamanagersandpolicy-makers.

WorkcarriedoutintheprojectAmainissuehasbeentoidentifythepopulationstructuresoffourkey-species;bladderwrackFucusvesiculosus,narrowwrackFucusradicans,BalticisopodIdoteabalthicaandthree-spinedsticklebackGasterosteus aculeatus.Forthreeofthespeciesthisinvolvedthedevelopmentofnewpowerfulgeneticmarkers,socalledRADmarkers.Thesemarkersareverymany(>10,000)andrandomlydistributedthroughoutthegenome.Withthehighnumberofmarkers,manyofthemwillsitclosetogenesunderselection,whileotherswillsignalpopulationgeneticstructureandgeneflow.RADanalysisrequireshigh-qualityDNA,andforthetwoFucusspeciesthisturnedouttobeachallenge.Finally,however,DNAfrommorethan40populationsofFucusvesiculosus/Fradicansand37populationsofIdoteabalthicawereextractedandsequencedfor>10,000polymorphicsites(SNPs).

BONUS BAMBI target species. The seaweeds (Fucus vesiculosus and F. radicans) are grazed upon by the isopod (Idotea balthica), that is eaten by the fish (Gasterosteus aculeatus), the fish using the seaweed to its nests. This model ecosystem constitutes a dominant part of coastal hard bottom ecosystems of the Baltic Sea.Photos:BONUSBAMBI.

BONUSBAMBI

FinalpublishablesummaryreportFebruary28th2018

3

Weusedsticklebacktoassesstowhatextentenvironmentalchangemaydirectlyinducegeneticdifferencesintermsofchangesingeneexpression,andifthesechangesarebeingtransferredtothenextgeneration.Theprocessforthisisnamed"epigenetics"andthereareseveraldifferentmechanismsinvolved.OneismethylationoftheDNAofaparentandthismethylationwillchangetheregulationofgenesandbeinheritedintothenextgeneration.Sixpopulationsofsticklebackhavebeencarefullyanalysedforfixedgenomevariation,differencesingeneexpression,andanyeffectsofepigenetictype.

NewmodelsforconnectivityanddistributionWehavepartlydevelopedandusednewmodellingapproachestomapconnectivityamongpopulations.Connectivitymodellingcombinesstatisticaloceanographicdatadescribingdirectionsanddispersalratesofcurrents,withbiologicaldataoflarvalduration,larvalbehaviour(depthdistribution)andhabitatchoice.Thesemodelscanbeusedtopredicthowfarandinwhatdirectionalarvaislikelytobetransportedafterreleasefromacertainspot.Inourmodelling,wetypicallyusedmultiplereleasespotsandinthisway,itwaspossibletopredictconnectivitypatternsforthewholeBalticSeaenvironmentinthesameanalysis,foraspecieswithcertaincharacteristics.

WehavealsomappedthedistributionofFucusandIdoteafromusingacombinationofnewandexistingground-truthingdataandclimateenvelopemodellingforthewholeBalticSeaarea.Thisisthefirsttimethecompletedistributionofthesekeyspeciesiscompiled.ThecurrentdistributionactasareferencetomodelhowthedistributionswillchangeuponwarmingandfresheningoftheBalticSeaovercomingdecades.Inthemodellingtoolswedeveloped,wealsoincludedexperimentaldatawheretolerancetohighertemperatureandlowersalinitywereexperimentallyassessed.

Mapswereproducedthatsummarisedtheresultsoftheconnectivityanddispersalmodelling,andthesemapsweremadeavailabletotheendusersthroughtheBaltGenewebpage.

ExperimentaltestsoftolerancelevelsTodeterminetoleranceofFucusandIdoteatocombinedstressfromincreasedtemperatureanddecreasedsalinityweusedexperiments.Intheseexperiments,wetestedpopulationsfromtheentranceoftheBalticSea,fromthemid-partandfromtheinnermost(marginal)partsofthespecies'distributions.Theseexperimentswereusedbothtodeterminetheaveragetoleranceofindividualstostresslevelscorrespondingtofuturecombinationsoftemperatureandsalinity,andtofindoutifthetolerancevaryamongindividualsduetodifferentgenotypes.Ifdifferentgenotypeshavedifferenttolerances,thereispotentialforevolutionofpopulationsthataremoretoleranttofutureclimatethanisthepresentpopulations.

Tolerancewasassessedforfitnessparameterssuchassurvivalandgrowthbutalsotogametereleaseandfertilizationsuccess.Thegeneticbackgroundoftolerancewasassessedbysequencingofthetranscriptometoseewhichgeneswereexpressedandnotundervariousenvironmentalconditions.

StudiesofknowledgetransferandeffectsofmanagementmeasuresToprotectgeneticvariation(geneticbiodiversity)isamajortargetundertheRioConventionandtheAichiBiodiversitytargets.Tounderstandtherolegeneticbiodiversityactuallyplayinmanagement,weusedinterviewsofauthoritypersonneltofindouthowinformationongeneticvariationiscurrentlyusedinthedesignandmanagementofMPAswithintheBalticSea.Inaddition,wealsoassessedcurrentlyappliedpolicyandmanagementagainstthebackgroundofexistinginternationalagreements,inparticulartheStrategicPlanoftheConventionofBiologicalDiversity.

Weanalysedtheeffectivenessofknowledgetransfertostakeholderscomparinglecturesandgroupdeliberations,andevaluatedbothshort-termandlong-termeffectsofthetransferonknowledgeandbeliefs.

Finally,weevaluatedtheeffectsofmanagementmeasures(positionsandsizesofMPAs)bybothadirectcomparisonofthegeneticvariationmaintainedinsideandoutsideMPAs,andbycomparingthelocationofexistingMPAswiththeoptimalpositionofBalticSeaMPAsasgeneratedfromamodelofconnectivity.

WedisseminatedresultsfromBONUSBambiandinitiatedanend-userdialogthroughthewebpage“BalticSeaGeneticsforManagers”(https://bambi.gu.se/baltgene)andthrougha2-dayworkshop.

BONUSBAMBI

FinalpublishablesummaryreportFebruary28th2018

4

MainresultsachievedduringtheprojectInBONUS-BAMBIweshowthatforthespecieswehavestudied,populationsofmarineoriginhavechangedtheirgeneticcompositioninanumberofdifferenttraitstobeabletosurviveandreproduceinthelow-salineBalticSea.WhatwealsoshowisthatthischangehasresultedinahighlycomplexgeneticstructurewithpopulationslocallyadaptedtodifferentpartsoftheBalticsalinitygradient.Inaddition,somespecies,haveacquiredcompletelynewtraits,suchas,cloninginBalticFucus,whichradicallychangethewaysinwhichgeneticvariationismaintainedandevolving.

InFucus,geneticdifferencesamongsitesintheBalticProperandGulfofBothnia,GulfofFinlandandGulfofRigaareevenmorepronouncedthandifferencesaccumulatedovertheBalticSeaentrance.Allthesedifferencesareduetoisolationandlackofgeneflow,butalsotolocaladaptation.

InbothIdoteaandsticklebackwehaveidentifiedgenesunderselection,andlocaladaptationtothelowsalinityoftheBalticSea.WeexpectsimilarresultsinFucus(analysisstillongoing).InFucusweseeanextremelycomplexpicturewiththeseparationofF.radicansfromF.vesiculosusinsomeareas,butnotinothers

NewtoolsforselectingMPAsandestimatefuturedistributionsUsingbiophysicalmodellingtoestimatetheconnectivitybetweenbasinsandregionswithintheBalticSeawehaveidentifiedseveralpartialdispersalbarrierswithintheHELCOMarea,whichmaysuggestmanagementunitsandlocallyadaptedsubpopulations.Biophysicalmodellingexplained,forexample,alargepartofthepopulationstructureofIdoteabalthicarevealedbyourdetailedgeneticsurvey.ThisshowsthatbiophysicalmodellingisatoolthatmaybeusedforafirstapproximationtoidentifyareasforMPAs.

ResultsofdispersalmodellingofthefutureFucusvesiculosuspopulationspointedoutthatlocallyadaptedgenotypesareatahighriskofextinctionbecauseFucusareunabletoshifttheirrangesfastenoughduetorestricteddispersalandlonggenerationtime.TheworstscenarioisthatF.vesiculosusmaydisappearfromlargepartsoftheBalticSeabeforetheendofthiscentury,withlargeeffectsonthebiodiversityandecosystemfunctioning.Toreducethisrisk,conservationactionslikeassistedcolonizationandassistedevolutionmaybenecessaryinthefuture.

VariationenablesfutureadaptationsWedeterminedtoleranceofFucusvesiculosus,F.radicansandIdoteabalthicatofuturesalinity-temperatureregimes,forpopulationsfromtheentrance,central,andmarginalBalticSearegions.FutureconditionshamperedgrowthandsurvivalofthecentralandmarginalBalticSeapopulationsofF.vesiculosuswhereastheentrancepopulationsfaredwell.Wealsofoundthatgametereleaseandfertilizationsuccessarehighlysensitivetosalinityconditionsandmaybethemostcriticallife-historystageunderclimatechange.

However,wefoundgeneticvariationintoleranceandthisvariationprovidestherawmaterialforadaptationtoachangingenvironment.Thisfindinghighlightstheneedformonitoringgeneticvariationinkeyecosystemspecies.InFucuswealsofound72genesthatweredifferentiallyexpressedbetweenthecurrentandfuturepredictedsalinityconditionsimplyingstronggeneticbackgroundofsalinitytolerance.

UnknownconsequencesofecosystemchangeTheendemicF.radicanstoleratedbetterthanF.vesiculosusthefuturecombinationofhyposalinityandwarmingandphenotypicplasticityoccurredevenwithinclonallineages.Fromtheseresults,itappearsthatF.radicansisakey-speciesforthefutureBalticSeaenvironmentandmayeventuallyreplaceF.vesiculosusthroughouttheBalticSea.However,ourexperimentsalsoshowthatresistancetograzingfromIdoteabalthicadecreasesinfutureconditions,andthismaybeatoogreatchallengeforbothspeciesofFucus.Incontrast,wecanbeconfidentthatsticklebackswillbeabletoadaptandsurvivefuturesalinitychangesintheBalticSea.ResultsfromourexperimentsshowthatBalticsticklebackhasabettertolerancetowardsdecreasedthanincreasedsalinity.ThesechangeswillstronglyaffectthefutureBalticSeaecosystem,butinwhatwayremainsunclear.

BONUSBAMBI

FinalpublishablesummaryreportFebruary28th2018

5

AneedfordialogueOurpoliticalsciencestudiesclearlyshowthatthereisagapbetweenwhatthescientificcommunityknowsaboutgeneticbiodiversityandhowitishandledinconservationmanagement.ThisgapisperhapsmoreworryingfortheBalticSeabiodiversitythanforbiodiversityelsewhere,becausegeneticdiversityisofparticularimportanceinthisyoungandspecies-poorecosystem.Weshowthatcontinuedinformationanddialoguebetweenresearchersandauthoritiesarenecessarytoestablishandimplementmanagementmeasuresprotectingcriticaland/orendangeredpartsofthegeneticdiversity.Moreover,wecanshowthatthecurrentdistributionofMPAsdoesnotgiveenoughsupporttoimportantgeneticvariationortositesimportanttomaintainahighconnectivityamongBalticSeapopulations.

ThecontinuityplanoftheprojectManyoftheresearchresultsgeneratedintheBONUS-BAMBIprojectarecurrentlybeingqualitycheckedandmadepubliclyavailablethroughpublicationsinopenaccessscientificjournals.Alreadymorethan25scientificpapershavebeenpublishedthathadfullorpartialsupportfromtheBONUS-BAMBIproject,andanother20publicationsareinpipelineasmanuscript.DuringtheBONUS-BAMBIprojectamassiveamountofsequencingdatahasbeengeneratedthatwillbecontinuedtobeanalysedforthecoming1-2yearsandgeneratepublicationsinthecomingyears.

MainhighlightsfromourresultsarecurrentlybeingmadeavailableontheBONUSBAMBIwebsiteformanagers,https://bambi.gu.se/baltgene.AnimportantconclusionfrominterviewsanddiscussionswithmanagersaroundtheBalticSea,istheneedforbettercommunicationsbetweenscienceandmanagement.

Asadirectconsequenceoftheproject,apilotprojectfundedbytheSwedishAgencyforMarineandWaterManagement(SWAM)hasinitiatedandisledbyJohannessonandLaikre.ThisprojectwillbenefitconsiderablyfromtheknowledgegeneratedinBONUS-BAMBI.TheprojectwillsuggestamonitoringprogramforgeneticdiversityinSwedishwaters.Moreover,theknowledgegeneratedinBONUS-BAMBIisfedintoalargerstudyofgeneticvariationinnorthernandwesternEurope,intheBiodiversaprogramMARFOR.

End-user conference BaltGene 2017 gathered almost 60 managers, policy-makers and scientists from around the Baltic Sea. For two days, knowledge and experiences was shared on why and how genetic biodiversity should be included in Baltic Sea management. Photos:BONUSBAMBI.

Overall,theBONUS-BAMBIprojecthasgeneratednewandcriticaldataongeneticvariationofBalticSeapopulations.Contrastingearlierbeliefs,localadaptationofBalticSeapopulationsistheruleratherthantheexception,andBONUS-BAMBIhasdemonstratedthisinanumberofnon-commercialspecies.Withthisknowledgeathand,itisworryingthat–asdocumentedinBONUS-BAMBI–thereissolittlefocusongeneticbiodiversityinmanagementofnon-commercialspeciesimportanttothefunctioningofthemarineecosystems.