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sporophytes Figure5 de nedasthematurestageofthebull and giantkelpdiploidlifecycles wereindiscriminatelycollected formorphometricmeasurement andcanopybiomass determinationa crossmultiplelocationsincentraland nort hernCaliforni a Table2 in2018and2019 including veofthesixdivesitessurveyedin2019 NoyoHarbor PointPiños OtterCove HopkinsMarineStation andS an CarlosBeach Spec i cally wemeasuredmorphologyatfour sitesf orgi antkelp totalof11sporophytes in2019andsixsites for bullkelp totalof86sporophytes in2018and2019 Table2 Diversremovedsporophytesfromthesubstrate manuallybycuttingtheprimarystipejustabovetheholdfast Figure5 Supplemen taryFigureS1A broughtthemtothe surfaceandthentosho rewheremorphometricmeasurements wereconductedonaclean surface SupplementaryFigures S1B Œ F Whentissuehydrationcouldnotbemaintainedusing freshseawater a portablepop u ptentwas usedtoshadesamples SupplementaryFigures1C D Bullkelpmorphometricmeasurementsweremadeforstipe lengthandwidth bulbdiameter sub bulbdiameter 15cmbelow thebaseofthebulb longestbladelength width longestblade weight numberofblades canopyweight top1mofstipe includingallthebladebiomass andtotalsporophyteweight Figure5 CentralandnorthernCaliforniadatafrom2018to 2019 n 86 werecombinedwithdatacollectedonthewestern FIGURE4 MicaSenseRedEdge Mnear infrared re ectance NIR foreachdivesurveysiteoverlayedwiththelocationsforthemidpoint ofeachtransect interval withthecolorof thedotcorrespondingtothemeasured insitu canopybiomass A NoyoHarbor NH B PortugueseBeach PB C PointPiños PP D OtterCove OC E SanCarlosBeach SCB F Hopkins MarineStation HMS Frontier sinEnvironmentalScience www frontiersin orgJuly2022 Volume10 Article690963 6 McPhersonandKudela Patch Speci cCharacteristicsChallengeBiomassEstimation coastof Prince of WalesIsland Alaskain 2018 Pearson and Eckert 2019 n 55 Bulbdiameterwasrandomlymeasuredwithinthe3 600m 2 divesurveyplot Thesite speci c mean bulbdiameterwasusedto estimatethemeancanopybiomass persporophyteateachbull kelpsite The totalwetbiomassforeachtransectinterval 10m 2 was calculatedbytakingtheproductofthestipecounts within eachtransectintervaland thesite speci cmeancanopybiomass persporophytederivedfromallometry Giantkelpsporophytesweredividedintotwosections the sub surfacecanopyandsurfacecanopy Figure5 Thesurface canopywas determinedbymeasuringthedepthoftheholdfast priortocollection Withineachsection morphometric measurementswere madefortotaltissueweight numberof blades andnumberoffronds CentralCaliforniagiantkelp datawerecombinedwithSBCLTERmeasurementsof canopy biomassandfrondcountsfrom2002to2003 Reedand Rassweiler 2018 n 36 Ateachgiantkelpdivesite the canopybiomasspertransect intervalwasestimatedusingthe allometricrelationshipdevelopedbetweenthetotalnumber of stipes fronds sporophyte andthecanopybiomass

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text:
Wecounted withineach transectinterval tocalculatethe corresponding canopybiomass AnANCOVAwasrunusingthePython 3 1 pingouin statisticsmodule Vallat 2018 todetermineif FIGURE 5 Relevantmorphometriccharacteristics ofbullkelp Nereocystisleutkeana andgiantkelp Macrocystispyrifera adultsporophytes imagecredi t Niky Taylor UCSC TABLE2 Detailedinformationofsite speci csporophy te collectionincludingsitename region kelpgenera latitude longitude and collectiondate s SiteNameSiteLocationKelpGeneraLatitude LongitudeCollection Date s ofSporophytes CasperCoveMendocino Countybullkelp39 362 123 820Sept 17 2018 Œ 10 AlbionCoveMendocino Countybullkelp39 228 123 772Sept 18 2018 Œ 15 BodegaMarineLabSonomaCountybullkelp38 311 123 071Sept 25 2018 Œ 17 Noyo HarborMendocino Countybullkelp39 429 123 812Sept 19 2018 Œ 12 7 Aug 2019 Œ 22 PointPiñosMontereyPeninsulabullkelp36 641 121 931Aug 28 2019 Œ 10 HopkinsMarineStationMontereyPeninsulagiantkelp36 622 121 9028 July2019 Œ 2 10July2019 Œ 1 17July2019 Œ 1 SanCarlosBeachMontereyPeninsulagiantkelp36 613 121 89522July2019 Œ 1 31July2019 Œ 1 OceanCoveMontereyPeninsulagiantkelp36 630 121 91926July2019 Œ 1 Aug 8 2019 Œ 1 Aug 26 2019 Œ 1 Steamer LaneSantaCruz giantkelp36 952 121 023Aug 16 2019 Œ 2 Frontier sinEnvironmentalScience www frontiersin orgJuly2022 Volume10 Article690963 7 McPhersonandKudela Patch Speci cCharacteristicsChallengeBiomassEstimation region andseason hadanin uenceontheslopeandintercepts oftherelationshipbetweenthedependentvariable bulbdiameter or frondcount andcanopybiomass RetrievingcanopybiomassfromNDREB Œ Aspatialanalysiswas conductedtocompare insitu derivedcanopy biomassestomean NDREBkelpclassi cationatfourdif ferentradiiaroundthetransect interval 0 5 1 5 2 5 5m Wecho setheseradiibecausetheyrelate torelevantspacebornsensor sgroundsa mplingdista nces PlanetScope 3m PlanetRapidEye 5m ESASentinel 2 10m spectralcharacteristicsaredescribedin Supplementary TableS1 Allpixelsthatfellwithintheradiuswereaveraged r esultingina

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text:
singlemeanNDREBvaluepertransectintervallocation Afullsite levelcomparisonofmeanbiomassandmeanNDREBwasalso conductedwithrelevancetoderivingcanopybiomassfromthe Landsatsuite 30m a ndSBCLTERretrievalsofcanopybiomass withLa ndsa t Cavanaughetal 2011 Belletal 2015 Semi variogramanalysis Œ Ageostatisticalapproachwasused todescribekelppatch spatialautocorrelationandpatch characteristicsbycalculatingthesemi variance of eachsurvey site sNDREB Figure3D and in situ biomassmeasurements Figure3F Prior toconductingthevariogramanalysis insitu biomassdatawereinterpolatedinto a2 Dgrid withtheMATLAB function griddata usingatriangulation basedlinear interpolationmethod WeusedthePython3 1Variogramand DirectionalVariogramclasseswithin theSciKit GStat skgstat moduletodeterminethesemi variance whichcanbe describedashalfofthemeasuredvariancebetweenpairsof valuesseparatedby an increasinglagdistancebetweenpixels h h 1 2 N h p N h i 1 NDREB x i NDREB x i h 2 2 whereNDREB or insitu biomass are the observationsat locations x i and x i h and N h isthenumberof pointpairs atthatlag Semi varianceparameterswereestimatedby tting sphericalmodelstotheempiricalsemivariograms C p 1 5 p h r 0 5 p h 3 3 where isthenugget C isthesill and istherange The describesthetotal unresolvedvariability ornoise while C describesthetotal resolvedvariability The describesthe distanceatwhichthesemi variance reachesamaximumand therefore thedistanceofspatialautocorrelation Spherical modelsareidealwhentheincreaseinsemi varianceissteep orbeing estimatedwithinasmallregion such asthe60 mlength scale ofthisstudy However because thiswasadescriptive analysisforpatch characteristics wedo notreportspeci c C or informationfromthedifferentsites RESULTS Allometricrelationshipsforca nopybiomassw ere developedfor bullkelpandgiantkelpbasedonmorphometricmeasurements andtheirrelationshiptocanopybiomass Figure5 Longest bladelengthwasthestrongestpredictorofbullkelpcanopy biomass r 2 0 78 Table3 Sub bulbdiameter bulbdiameter andnumberofbladeswer eals osigni cant predictorsof canopy biomass r 2 0 72 0 64 and0 48 respectively Table3 Our results were consistentwith Stekolletal 2006 forbullkelpin

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text:
Al aska Bul b diameterwasselected to useforderivingcanopy biomassinthisstudybecausebulbdiametercanbequickly measured bydiversorfromaboatatthesurfaceofthekelp canopy Thestrongest predictorof giant kelpcanopybiomass wasthenumbe r ofcanopyblades r 2 0 81 Table3 Additionally mean canopyfrondlength andnumberof frondsatthebaseofthesporophytewerestatistically signi cantpredictorsofcanopy biomass r 2 0 57and0 48 respectively p 0 05 Oursamplesizeofsporophyteswa s relativelysmall collectedinJulyandAugust2019 n 11 However combiningourdatase twithSBCLTERmeasurements ofstipe frond countandcanopy biomasscollectedmonthly in 2002and2003increasedthesamplesizeton 40 increasedthe F statistic23 1 p 0 05 but reducedtheoverallcoef cientof determinationto 0 32 Table3 All insitu divermeasurementsofbull kelpbulbdiameter displayedaGaussianprobabilitydistribution Figures6A Œ C Therangeofbulb diameteratPointPiños Figure6C 7 06 0 96 n 148 wassigni cantlydifferentthanNoyo Harbor Figure6A 6 50 0 76 n 80 t statistic 4 5 p 0 05 and PortugueseBeach Figure6B 6 44 0 64 n 79 t statistic 5 12 p 0 05 Additionally themorphologyofMontereyPeninsulabull kelpbulbdiameterwassigni cantlydif ferentfromthemorphology ofbullkelpinAlaskaandMendocino Table4 Asaresult there weresigni cantdifferencesbetweenthebullkelpallometric TABLE3 Linearregressionstatistics for thediffer entmorphometricmeasurements for estimatesofwetweight BullKelpSlope m Intercept b r 2 F Statisticn Longestbladelength cm 24 1 2 4153 4 14 40 78103 232 Sub bulbdiameter cm 0 2 0 032 6 0 160 7276 232 Bulbdiameter cm 0 2 0 035 8 0 200 6453 132 Numberof blades3 0 0 640 7 40 70 4827 732 GiantKelpSlope m Intercept b R 2 F Statisticn Numberof blades

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canopy 0 03 0 010 2 4 10 8138 811 Meanfrondlength canopy 0 07 0 02 7 9 9 60 5710 610 Numberof fronds base Œ thisstudy0 9 0 3 0 8 8 60 488 211 Numberof fronds base Œ SBCLTER0 4 0 14 2 3 60 3223 140 Frontier sinEnvironmentalScience www frontiersin orgJuly2022 Volume10 Article690963 8 McPhersonandKudela Patch Speci cCharacteristicsChallengeBiomassEstimation relationshipbyregion Figure6D Table5 However giantkelpwas notin uencedbyregionorseason Table5 Thepersporophyte canopybiomassmeasurementsweregenerallyhigherforgiantkelp thanbullkelpandmaximawere 15and60kgfor bullkelpand giant kelp respectively Figure 6 Thoughmeasuredcano pybi omassvaluesrangedsigni cant ly across siteandgenera NDREBval uesweregenerallywithinrange ofeachother Figure7 toprow bullkelpsites bottom row giant kelpsites Apredictiverelationshipbetweencanopybiomass and NDREBwasobservedattwoofthest u dysites PortugueseBeach bull kelp Figure7B andSan CarlosBeach giantkelp Figure7E forall samplingradii 0 5 1 5 2 5m Table6 Regressionstatisti csforthese twositeswascharacterizedbyhigh F statisticand r 2 valuesrelativeto theotherfoursites However ac rossallsites therewasno in uenceof averagingradiion thepredictivepoweroftherelationship between FIGURE6 Theprobabilitydistributionofbullkelpbulbdiameterfor A NoyoHarbor B PortugueseBeach C PointPiños D Scatterplot ofwetbullkelp canopybiomassagainstbulbdiameter ttedwithOLSlinearregressionsforAlaska blue line Mendocino tealline Monterey greenline andallregionscombined b lackline E Scatterplotofwet giantkelpcanopybiomassagainstfrondnumber ttedwithOLSlinearregressionsforalldatafromSantaBarbara CA blueline summerdatafromSantaBarbara b luedashedline Monterey CA greenline andallregionscombined blackline Regressiondetailslistedin Supplementary TableS1 Greyshadedareasrepresentregression95 CI TABLE4 ResultsfrompairwiseTukey sHSDpost hoctestforregionspeci c bull kelpbulbdiameters denotessigni cance LocationALocationBMean A Mean B Std Error p value AlaskaMendocino2 6 3 90 470 012

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AlaskaMonterey2 69 00 91 0 001 MendocinoMonterey 3 9 9 00 90 0 001 Frontier sinEnvironmentalScience www frontiersin orgJuly2022 Volume10 Article690963 9 McPhersonandKudela Patch Speci cCharacteristicsChallengeBiomassEstimation canopybiomassand NDREB Port ugueseBeach Figure7C and PointPiñosexhibite dthelargestr angeinmeasuredc anopybiomass 0to 60 kg wwm 2 Figure7 thatwereassociatedwithhighdiver recordedstipedensities Figures3E F Conversely giantkelp stationshadasmallerrangedinmeasuredc anopybiomassasa result of lowerdivermeas uredstipedensities 0 Œ 15 kg wwm 2 Figures7D Œ F Nosigni cantrelationshipwasobservedwhensitelevelmeansof NDREBand insitu canopybiomasswerecompared Figure8 PortugueseBeachandPointPiñosshowedhighmeancanopy biomassrelativetothemeanstationN DREBre ectinghigher diver measuredstipedensities for thesetwobullkelpsites Figures3E F Therema iningfourstationswe recharacteristic ofrelativelylowmeancanopybiomass andarangeofmeanNDREB valuesfrom0 18to0 39re ectingsparserd ivermeasuredstipe densitiesatthose fourstations Thedescriptivevariogramanalysisconducted forNDREBand divermeasuredcanopybiomassindicateddifferencesinpatchlevel spatialpatternsbetweenPortugueseBeach SanCarlosBeachandt he othersurveysites Figure9 Althoughwewerenotabletoconsistently developpredictionsofcanopybiomassfromNDREB thepatternsof semi varianceacrossthesite sspatialextentsweresimilarbetweenthe diver surveys andNDREB indicatingthatsimilar spatialpatterns were beingobserved with bothmethods ForPortugueseBeachand San CarlosBeach thetwositeswhererobustrelationshipsofcanopy biomasswereachieved adominant u shaped patternwasvisiblei n thesemi variograms wherevariab ilityrapidlyrisestotheedgeofthe patch the peakofthe u shape and fallsoutsideofthepatch beyond approximately30mfromthecenterofthesurveysite DISCUSSION In thenortheastPaci cregion bullkelpandgiantkelpform thebaseofanecologicallyandeconomicallyimportant tempe ratenearshorecoastalecosystemwithextensive oatingsurfacecanopies Remotesensingisadvantageous fordetectingtheir oatingcanopyandderivinglarge scale estimatesofsurfacecanopybiomass whichcomprisesmore than90 ofbullandgiantkelpstandingstock Rassweiler etal 2008 Developmentandvalidationofremotelyderived kelpcanopybiomassislackingbecausetheassociated methodologyis timeintensiveandcostly butimportant becausebiomassisnecessaryfordeterminingratesof net primaryproductivityandquantifyingcarboncyclingin temperatenearshoreregions Inthis study wecompiled bullandgiantkelpmorphometricmeasurementsfrom differentregionsin theNE Paci c toassessthe validity of applyingasinglebiomassconversionrelationshipacross disparateregions Wealsosur veyedsixspatiallydistinct sites 3bullkelp and 3giantkelp with varyingstipeand canopycharacteristicsformeasurementsof in situ canopy biomassviadiverandUAVsurveyswiththeaimtoretrieve canopybiomassestim atesfromremotesensingimagery The resultsexhibitedthat1 kelpmorphologyd ifferedslightly acrosstheregionsweexaminedbutdidn tstrongly

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in uence insitu estimatesofcanopybiomass and2 reliable consistentremotesensingretrievalofcanopy biomasswasdif cultusingthesu rveyap proachdevelo ped hereanddependentonspeci c kelppatchcharacteristics acrosssites Kelpallometryforcanopybiomassmeasurements Œ Der iving andutilizingallometricrelationshipsforcanopybiomassisa keycomponentofdevelopingmethodsforcanopybiomass retrievalfromspaceborneandaerialplatforms Stekolletal 20 06 Cavanaugheta l 2010 Inthis study wecomparedthe regionaldifferencesinbullkelpbulbdiameterandgiantkelp stipe countaspredictorsofcanopy biom assandfoundthat acrossallregions thesemetricswererobustpredictorsof canopybiomass Theeffectofregionontheslopesand intercept of thelinearrelationshipsweresmall forboth genera Figure6 Table4 and Table5 andindicatesthatit maybegenerallyacceptabletoa pplyarelationshipd erived fromoneregiontoothersacrosstheNEPaci c Itispossible thatthemorphologicaldifferencesweobservedwerearesultof local scalehydrodynamicsin uencingtissuemorphology ratherthanregional scaleprocesses KoehlandAlberte 19 88 Koehl 2022 However thesamplesizesforsomeofourregionswere relativelylow e g giantkelpfromtheMontereyregion Samplesizesofstudiescollectingsporophytemorphologyis in uencedbythepersporophyteeffortitrequirestocollect thisinformation Asinglegiantkelpsporophytecantakeupto 15person hourstoconductacompletemorphometricsurvey Bullkelpislesslaborintensive ontheorderoftwoperson hours per sporophyte BothgenerarequireSCUBA from shore orboat tocollectanddeliversporophytesthatcan weighupto 60 kgto thelab forprocessing Whilethelogistical challengesmaybetheprimaryreasonforthelackofdata the collec tionofthisdatahavenotbeenprioritizedbyfundingand managementagenciesin thepast Weadvocateforallocating resourcesinto developingpredictiverelationshipsforcanopy TABLE5 ANCOVAresultsfortheeffectofregion andseasonsforgiant kelp ontheslopeandinterceptsoftherelationshipbetweenthedependentvariable bu lbdiameter or frondcount andcanopybiomass BullkelpregionsareMonterey Mendocino andAlaska Giantkelpregions season sareSantaBarbara all Sant a Barbara summer andMontery denotessigni cance GeneraGroupDegreesof Freedom Df F p valueEffectSize Nereocystis Region211 6 2 3 10 5 0 15 Nereocystis BulbDiameter cm 184 7 5 3 10 16 0 38 Macrocystis Region season22 68 5 10 2 0 074 Macrocystis Sporophytefrond144 6 6 9 10 9 0 41 Frontier sinEnvironmentalScience www frontiersin orgJuly2022 Volume10 Article690963 10 McPhersonandKudela Patch Speci cCharacteristicsChallengeBiomassEstimation biomasstofurtherimproveaccuracyof kelpcanopyretrieval fromspacebornesensorsvia long termmonitoringand targetedstudieswithUAVs The in uenceofkelppatchcharacteristicsoncanopybiomass retrievals Œ Descriptiveresults fromthesemi variogramanalysis

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text:
indicatedthatkelpbed characteristicsplayarolein accurately determiningcanopybiomassretrievals Ifakelppatchwaslarger thanthespatialareaofthedivesurvey i e abreakdownin autocorrelationdidnotoccurwithinthesurveyarea wewere limitedin our abilityto developarobustworkingrelationship betweencanopybiomassandNDREB Ifthekelppatchwas smallerthan thespatialareaofthedivesurvey i e pixel autocorrelationbrokedownwithinthedivesurveyarea we wereabletoidentifyarelationshipbetweencanopybiomassand kelpfraction Thispatterninthesemi varianceresultsdescribed siteswithadistinctkelppatchsurroundedby waterandbiomass atthesesitesalsohad arobustcorrelationwithNDREB Outof thesixsites PortugueseBeach Figure4B Figure7 andSan CarlosBeach Figure4E Figure7 wereclearexamplesofthis pattern exhibitingdense relativelyhomogenousstipe frond countsinsidethekelpbedand arelativelyhomogenousabsence ofstipes fronds outsideofthe kelpbed Thisindicatesa potential mismatchbetweenthescaleofmeasurementsby diversurveyandremotesensingbecause 1 surveyinglarge patchesisquanti edbestbysatelliteremotesensingbutisnot feasiblefor in situ surveys and2 veryhighspatial resolution TABLE6 OriginalLeast Squared OLS regressionstatisticsfrom Figure7 withslopeand interceptvalues 1standarddeviation StationNDREBSampli ng Radii m nSlope m Intercept b r 2 F Statist ic Noyo Harbor NH 0 528 1 1 3 233 3 0 960 0040 12 1 5 282 2 3 063 1 0 970 0200 52 2 5 282 3 3 153 0 0 980 0250 68 PortugueseBeach PB 0 55637 3 7 335 7 2 070 32425 9 1 5 5642 1 8 785 2 2 210 29923 1 2 5 5645 6 10 04 7 2 350 26420 8 PointPiños PP 0 556 1 5 7 8221 7 2 11 0 0010 04 1 556 2 4 10 821 6 2 21 0 0010 05 2 556 6 2 13 521 2 2 37 0 0040 21 OtterCove OC 0

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text:
5 562 7 1 331 7 0 580 0724 20 1 5 563 5 1 421 5 0 580 1036 22 2 556 3 7 1 551 49 0 59 0 0975 79 SanCarlosBeach SCB 0 556 6 5 1 481 2 0 450 26419 3 1 5 566 5 1 491 2 0 480 26319 3 2 5 566 8 1 511 1 0 480 27320 3 HopkinsMarineStation HMS 0 5 562 7 1 901 4 0 780 0362 00 1 5 562 6 1 981 4 0 800 0321 78 2 5 562 4 2 051 5 0 820 0261 42 FIGURE7 Transectintervalcanopybiomassinkgwet weight kg ww againstaverageNDREBvaluesfroma1 5mradiusaround each transectinterval A Noyo Harbor B PortugueseBeach C PointPiños D OtterCove E SanCarlosBeach F HopkinsMarineStationwithOLSregression blacklines and95 con dence inte rvals greyshading Signi canceoftheOLSregressionisindicatedwithan asterisk See Table6 foreach samplingradii and site s regressionstatistics Frontier sinEnvironmentalScience www frontiersin orgJuly2022 Volume10 Article690963 11 McPhersonandKudela Patch Speci cCharacteristicsChallengeBiomassEstimation measurementsofcanopycanbe observedwithUAVs butdiver surveymethodsarelimitedinthecoarsenessoftheir observations Thein uenceofkelpgeneraoncanopybiomassretrievals Œ Bull andgiantkelphavedistinctlydifferentmorphologiesandsurface canopycharacteristicswhichin uencedaccurateretrievalsof canopybiomassestimatesfrom UAVbasedobservations Bull kelphasasinglegas lledstipeandpneumatocyst Figure5 withlongblades length 1m thatcannumberuptothe hundredsconcentratedatthetopofthesporophyte Unlikegiant kelpblades thebladesofabullkelpdonot oatonthewater surfacewithoutsuf cienttidalandcurrentforcing Becausetides and currentwerenotproducingsigni cantdragonthebull kelp canopyatoursurveysites asvisibleintheimagery Figure4 muchofthebiomassinthecanopy Figure5 bullkelpblades was belowthewatersurfaceatourstudysites Therefore thespectral signatureofthekelpcanopyobservedwas

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text:
emergentstipesand bulbs This mayhaveresultedinaheterogeneoussurface comprisedof kelptissueandwaterandrelativelylow NDREB incomparisonto insitu biomass measurements Diverestimatesofcanopybiomassesrangedsigni cantly dependingonthedensityofbullkelpsporophytesresultingin areanormalizedcanopybiomassesatPortugueseBeachandPoint Piñosthat wereapproximately 7times greater thanthegiantkelp sites Figure7 Althoughbulbdiametercanpredict anindividual sporophyte scanopybiomass withrelativeaccuracy usingstipe countsdidnottranslatewelltoareanormalized canopybiomass atintervallocations eitherbecausethesporophytesfannedout anddistributedatthesurfaceor becausebladeswerebelowthe watersurfaceand werenotdetectedbythesensorontheUAV Furtherconsiderationshould be giventothedeepestsurveysite Point Piños 30m whereitislikelythataportionofthe sporophytescountedby diversdidn treachthesurface resulting inanoverestimationof insitu canopybiomass Giantkelpbladesgrowalongtheentirefrondfromthebase tothegrowingtip meristem Figure5 Moreofthecanopy biomassis oatingonthewatersurfaceorjustbelowbecause thebaseofeachgiantkelpblade length 30cm containsa single pneumatoc yst Asingle giantkelpsporophyte c an containupto 100fronds clusteredtogether Figure5 SupplementaryFigureS1A Therefore dense oating canopiesof giant kelpfrondscanform oftenwithmany FIGURE9 S emi variogram foreachstationNDREB grey anddivermeasured canopybiomass black NDREBareoverlayed withasphericalmodel t A NoyoHarbor B PortugueseBeach C PointPiños D OtterCove E SanCarlosBeach F HopkinsMarineStation FIGURE8 Meanstationcanopybiomass plottedagainmeanstation NDREBwith 1SEbarsin grey Differentcirclesrepresentconditionsofstipe densityatthedifferentdivesites Frontier sinEnvironmentalScience www frontiersin orgJuly2022 Volume10 Article690963 12 McPhersonandKudela Patch Speci cCharacteristicsChallengeBiomassEstimation frondslayingontopof eachotherat thesurface However since the distributionofindividualsporophytesatthe substrateis oftensparseandn on homogenouslydistributed acrossak elppatch disparitiescandevelopbetweensparse divercountsoffrondsand theexpressionofdensesurface canopy andanunder representationof insitu canopy biomass Itislikelythatstipec ount thoughrelativelyeasy fordiverstoconductisanunreliableapproachforremote sensingvalidationstudiesofgiantkelp Th ein uence ofsurveydesignoncanopy biomassretrievals The insitu diversurveyapproachdevelopedinthisstudywassimilarto methodologyforLandsatderivedgiantkelpcanopybiomass Rassweileretal 2008 Cavanaughetal 2011 andaerial surveyderivedAlaskabull kelpbiomass Stekolletal 2006 in thatdisparate kelpbedsweremeasuredtoquantifycanopybiomass andthenassociatedwithspectral characteristicsofkelpcanopy However thegoal ofoursurvey designwastorapidlyassessas manylargesitesas possibleby onlycollectingstipe frond counts withineach 5 msectionofthe transect The radialsurveypattern allowedustohaveeighttransectsof reasonablelength 35m withineachsurveysite Byassessing alargearea wecouldincrease thesamplesize andtherefore thenumberof matchuplocations

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text:
between insitu canopybiomassandUAVimagery However itis possiblethatpersistenterrorsandoffsetswereintroducedtothe transectinterval sgeospatiallocationsviathisapproach Despite SBCLTER s robustpredictionofSantaBarbaragiantkelpcanopy biomassestimates from30mLandsat Cavanaugh etal 2011 Bell etal 2020b wehadlittlesuccessdeveloping asimilarrelationship usingmeansitelevel 60 60m NDREBandcanopybiomass likelyduetolimitationsinthenumberofsitesthatcouldbe sampledwithinthegiventimeframeof thestudy Our ndingsshedlightontheadvantagesofsite speci clong term monitoringforretrievingcanopybiom assfrom remotesensing platformsatthekelpbedscale 10 Œ 30m allows for the developmentoftemporally andspatiallyrobustrelationships However thisapproachis n otalwaysfeasibleforsubtidal mo nito ringpro grams which initself arelacking Werecognize thereshouldbealternativemethodstoretrievingcanopybiomass UAVsprovide exibilityforsurveydesignandapproachtoadjust measurementtemporalfrequency thespatialextentofthesurvey a singlebedtomultiple and postproc essing pixelspatialbinning pixel sizesfrom10cm Ifgiven alongenoughtimeseriesorenoughdata points UAV scan collecttherequired information butas demonstratedhere itrequire smultiplemeasurements i e a timeseries tobuildreliablerelationships Suggestedsurveydesign forfutureUAVmethods Œ Basedonthe resultspresentedhereandlessonslearned we providea recommended surveydesignaimedatretrievingcanopy biomassestimatesfromremote sensingmethods Primarily we believemeasurementsshouldfocusdirectlyonthesurfacecanopy ratherthansubtidalstipecountsorbiomass Figu re10 Thisallows fordirect comparisonof insitu surfacebiomass measurementsto pixel basedkelpclassi cationmetrics Speci cally werecommend conducting insitu canopymeasurementsacrossavarietyofdensities eitherwithinasinglebedormultipledifferentbeds withina boundingboxmarkedbybuoys markersinthecenterandcorners Figure10 whiteandorangedots Buoysshouldremaind eployed for theextentofthe UAVsurveyfor groundtruthingandgeospatial positioningoft he insitu site Quadrat samplingpointsforsurface measurements withintheboundingboxcanberandomizedby heading distancetoremovebiasassociatedwithusing underwater compassheadingsforcalculationofGPSlocations AssuchGPS measurementsatthecenterofeachquadratlocationcanbetaken Withineach1 m 2 quadrat measurementsofthebulbdiameter bull kelp sub bulbdiameter bullkelp meancanopystipelength giant kelp andstipecount bullandgiantkelp canbeusedtoestimate canopybiomass Table3 Taking subtidalstipe countsisa good comparisontoothersurveysbeingconductedbyrelevant monitoringorganizations i e ReefCheckCalifornia CDFW HumbleStateUniversity etc but asindicatedbytheresults presented herecan beunreliablefordeveloping nespatialscale estimatesof biomass WerecommendconductingUAVsurveyson thesamedayjustbeforeor after insitu canopy measurementsorata similartidalheight FIGURE10 Suggestedsurveydesignfor future insitu validation studiesof canopy biomassusingaUAVplatform Frontier sinEnvironmentalScience

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www frontiersin orgJuly2022 Volume10 Article690963 13 McPhersonandKudela Patch Speci cCharacteristicsChallengeBiomassEstimation CONCLUSION Estimationofkelpcanopyareahasrapidlyadvanced leading toregional scale e g Belletal 2020b toglobal scale e g Mora Soto etal 2020 estimates andassessmentsofkelp canopyareainveryremotelocations Friedlanderetal 2020 Houskeeperetal 2022 Despitetheseadvances itisdesirable insomecasestomovebeyondcanopyareatoassessbiomass andthenprimaryproductivity asthatprovidesinformation onkelpbedhealth carbonsequestration andviability Validationstudiesofkelpcanopybiomassretrievalsvia remotesensingarelackingbecausethedatais dif cultto collect but thisdataisimportant becauseaccuratelarge scale estimates ofproductivityandcarboncyclingcannotbemade withoutthem Weattemptedtoclosethedatagapby developingarapidandthoroughapproachtogatheringthe insitu andremotesensingdatathat arerequiredtoretrieve biomassfromremotes ensingplatforms Although speci c kelppatchcon gurationandsurveydesignlimitedthe numberofsiteswherewesuccessfully retrievedkelp canopybiomass weconsiderour ndingsvaluabletothe kelpremote sensingcommunity towhomthereis verylittleof this dataavailable Understandingthechallenges in determining remotely derivedkelpcanopy biomassis importantandourgeneralconclusionscanbeusedto extrapolatet ootherregionsandpotent iallyot herkelp genera Toaidinfutureefforts weoutlineanimproved surveydes ignforfuture insitu validationstudiesusing UAVs Finally westronglyrecommend funding implementationoflong termmonitoringprograms suchas thoseused bytheSBCLTER acrosstheregion forboth giant andbullkelp Critically thisrequiresbothstandardizationof methodologyand consistentfundingbymanagementagencies forsuchefforts Thealternativeistorelysolelyoncanopyarea whichcanbeapoorproxyforbiomassandproductivity limitingourunderstandingof howkelpbedsrespondtoshort andlong termenvironmentalconditions DATA AVAILABILITYSTATEMENT Therawdatasupporting theconclusionof thisarticlewillbe madeavailablebytheauthors withoutunduereservation AUTHORCONTRIBUTIONS Allauthorscontributedtothismanuscriptincluding conceptualization MLMandRMK methodology MLMand RMK writing Š originaldraftpreparation MLM writing Š reviewandediting MLMandRMK FUNDING Financial sup portwasprovid edb ytheDr Earl H MyersandEthel M MyersOceanographicandMarineBiologyTrust andtheUCSC DepartmentofOceanSciences Vesselsupportandlodgingwas providedbyLauraRogers Bennett sMarineInvertebrateFisheries andC onservationLaboratory Funding forthe UAVwassupported bytheIdaBenson LynnOceanHealthendowment TheMicaSense RedEdge MandPix4Dsoftwarewassupport edby thePackard ScienceandTechnologyendowmenttotheInstituteofMarine Sciences ACKNOWLEDGMENTS Manythanks totheallthepeoplewhomadethisworkpossible includingHeidiHirsh StanfordUniversity scienti cdivers LoganGrady JeannettePeters andSara Hamilton eld volunteers alphabetically DennisFinger AimeeFoy Megan Gallagher KyleHughes KevinHum JosieIselin Quinn Letitia John Mizell RichMuller Brynn O

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