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Plant Propagation Protocol for Pedicularis bracteosa ESRM 412 – Native Plant Production Source: Burke Herbarium 2017 Photo credit: G. D. Carr TAXONOMY Plant Family Scientific Name Orobanchaceae (formerly placed in Scrophulariaceae) Common Name Broomrape family (formerly in figwort family) Plant Species Scientific Name Pedicularis bracteosa Benth. Varieties and Sub-species Pedicularis bracteosa var. atrosanguinea Pedicularis bracteosa var. bracteosa Pedicularis bracteosa var. flavida Pedicularis bracteosa var. latifolia Pedicularis bracteosa var. pachyrhiza Pedicularis bracteosa var. siifolia (Burke Herbarium 2017) Cultivar None listed Common Synonym(s) None listed Common Name(s) Bracted lousewort, wood betony, fernleaf Species Code (as per USDA Plants database) PEBR GENERAL INFORMATION Geographical range Widely distributed throughout the mountainous areas of Washington; British Columbia south to California, east to Montana, Colorado and

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PlantPropagationProtocolforPedicularisbracteosaESRM412–NativePlantProduction

Source:BurkeHerbarium2017Photocredit:G.D.Carr

TAXONOMY

PlantFamily ScientificName Orobanchaceae(formerlyplacedinScrophulariaceae) CommonName Broomrapefamily(formerlyinfigwortfamily)PlantSpecies ScientificName PedicularisbracteosaBenth.

VarietiesandSub-species

Pedicularisbracteosavar.atrosanguineaPedicularisbracteosavar.bracteosaPedicularisbracteosavar.flavidaPedicularisbracteosavar.latifoliaPedicularisbracteosavar.pachyrhizaPedicularisbracteosavar.siifolia(BurkeHerbarium2017)

Cultivar NonelistedCommonSynonym(s) NonelistedCommonName(s) Bractedlousewort,woodbetony,fernleafSpeciesCode(asperUSDAPlantsdatabase)

PEBR

GENERALINFORMATIONGeographicalrange WidelydistributedthroughoutthemountainousareasofWashington;

BritishColumbiasouthtoCalifornia,easttoMontana,Coloradoand

NewMexico(BurkeHerbarium2017).

Source:USDAPlantDatabase2017

Source:BurkeHerbarium2017

Ecologicaldistribution

Moist,openforestthickets,meadowsandclearingsinthemountains(Pojaretal1994).Shade-intolerant,subalpine,WesternNorthAmericanforbdistributedmoreintheCordilleranthanthePacificregion.Speciesoccursincontinentalborealandcooltemperateclimatesonverymoisttowetnitrogen-mediumsoils.Commoninsubalpinemeadowsandopen-canopy,highelevationforestsonwater-receivingsitesinthecoastalinteriorecotone(E-floraBC2017).

Climateandelevationrange

Occursinmontaneboreal&cooltemperateclimatesatanaverageelevationof1694meters,withaminimumelevationof8metersandamaximumof2520meters.(E-floraBC2017)

Localhabitatandabundance

CommonandabundantatsubalpineandalpineelevationsfromsouthernmainlandBritishColumbiasouthtoCalifornia(Pojaretal1994).Growsonturfyslopes,nearalpinebrooksorseepages,oramongwillowsandscrubconifersnearorabovetreeline(Porsild1979).

Plantstrategytype/successionalstage

Noinformationfound.MuchworkhasbeendonewithpollinationstrategiesforthePedicularisgenus(see“Workcitedbutnotused”sectionforpapersonpollinationbiology),butlimitedinformationwasavailableaboutgeneralplantstrategies.

Plantcharacteristics General:Perennialherbfromamixofcoarsefibrousandtuberous-thickenedroots,smoothexceptlong-hairyintheinflorescence;stemssingleorclustered,erect,30-100cmtall,un-branched.Leaves:Basalleavessimilartothestemleavesbutlonger-stalked,ormuchsmallerorlacking;stemleavesalternate,short-stalkedtoun-stalked,7-16cmlong,deeplypinnatelycleftintonarrowlyoblongtolanceolatesegments,themainsegments1-7cmlong,jaggedlyincisedandalsofinelysaw-toothedormerelydouble-toothed,theleavesreducedandlessdividedupward.Flowers:Inflorescenceadense,elongate,terminalspikeofnumerousflowers,aboveseveralhairy,leaf-likebutundividedbracts;corollasyellowortingedwithredorpurpletowhollypurple,13-21mmlong,2-lipped,theupperliphood-like,beaklessorveryshort-beaked,aboutaslongasthetube,thelowerlip3-lobed;calyces7-10mmlong,hairy,5-lobed,thelobesglandular-hairytonearlysmooth,theupperlobeshortest;stamens4.Fruits:Capsules,asymmetrical,flattened,curved,smooth,10-12mmlong;seedsseveral,2-5mmlong,net-veined.Source:E-flora2017.

Source:E-floraBC2017

PROPAGATIONDETAILS:SEEDSFivedifferentscientificstudiesforalpineseedpropagationwerefoundthatincluded

thegenusPedicularisinthestudygroup,buttheonlyonewhospecificallygrewP.bracteosaaspartofthestudywasKayein1997.

Ecotype 1. Alsosetal2012:ArctictundrainIsfjordenareaofSvalbard,Norway.

2. Bliss1958:ArctictundranearUmiat,Alaska.

3. Cooperetal2004:ArctictundraonColesdalen,NWcoast,andAdventdaleninSpitsbergen,Svalbard,Norway.

4. Kaye1997:AlpinetundrainOlympicNationalPark,ClallamCo.,Washington:onthesouthfaceofBlueMountainat1700-1800melevation;alongthetrailtotheElwhaRiverfromHurricaneHillat1460-1500mandat1675-1700mneartheFirstDivide

areaofMountAngeles.

5. Lietal2007:AlpineandsubalpinemeadowsfromthenorthwesternYunnanProvinceinChina.

PropagationGoal GerminatedseedsPropagationMethod SeedProductType GerminatedseedsStockType Nativeseedfrom:

1. Alsosetal2012:ArctictundrainIsfjordenareaofSvalbard,Norway.

2. Bliss1958:ArctictundranearUmiat,Alaska.

3. Cooperetal2004:ArctictundraonColesdalen,NWcoast,andAdventdaleninSpitsbergen,Svalbard,Norway.

4. Kaye1997:AlpinetundrainOlympicNationalPark,ClallamCo.,Washington.

5. Lietal2007:AlpineandsubalpinemeadowsfromthenorthwesternYunnanProvinceinChina.

TimetoGrow 1. Alsosetal2012:Noinformationlistedforhowlongseedsweregerminated.

2. Bliss1958:Seedswereconsideredtohavegerminatedwhentheradicleappeared(p.181).(theauthordidnotlistatimeframeforgermination).

3. Cooperetal2004:Soilsamplesfromthepolarheath,birdcliff,

proglacialhabitatandthermophilicheathweregrownover12weeks(11forthermophilicheath)simulatingthemaximumSvalbardsummerperiod.SamplesfromintactanddisturbedDryasheathweregrownover14weeks(p.117).

4. Kaye1997:Themaximummonitoringperiodwas145days,butwaslessthan80daysformostspecies(Kaye1997,p.116).

5. Lietal2007:100%oftheseedsgerminatedafter10daysof

incubation(p.1260).TargetSpecifications Germinatedseeds.PropaguleCollectionInstructions

1. Alsosetal2012:SeedswerecollectedbetweenAugust27thandSeptember19th,2008intheIsfjordenareaofSvalbard.M.rubellaseedswerecollectedonSeptember10th,2008in

Sassendalenina“disturbedmosstundra”habitat(Appendix1).Whenpossible,seedswereshakenoutoftheplantstoensureonlymatureseedswerecollected(p.820).

2. Bliss1958:SeedsofthealpinespecieswereobtainedfromplantsgrowinginthealpinetundraoftheSnowyRangeoftheMedicineBowMountains,11,000feetabovesea-level,approximately40mileswestofLaramie,Wyoming,in1954and1955(p.181).

3. Cooperetal2004:SeedscollectedinearlyJulytolateAugustin

theyear2000(p.116).Thetop2cmoforganicsoilwascollectedtogetherwithbryophytesandlitter.Soilsampleswerecollectedwithin10cmoffocusspeciestomaximizechanceofcapturingdispersedseed(p.117).

4. Kaye1997:Seeds(orsingleseededfruitssuchascaryopsesand

achenes)wereharvestedbyhandinAugustorSeptemberof1987fromsinglepopulationswithinOlympicNationalPark,ClallamCo.,Washington(p.116).

5. Lietal2007:SeedsofPedicularisspeciesoftenmaturefrom

AugusttoSeptemberinnorthwesternYunnanProvince.Inthisexperiment,seedsofeightPedicularisspecieswerecollectedfromAugusttoNovember2004(p.1259).

PropaguleProcessing/

PropaguleCharacteristics

1. Alsosetal2012:Iftheplantswerewet,seedcapsuleswerecollectedandleftinpaperbagstodry.Seedsthatwereobviouslynotripewerenotcollected(p.820).

2. Bliss1958: Seedswereplacedinglassvialsorpaperbags,airdriedandshippedtoDukeUniversity(p.181).

3. Cooperetal2004:Samplesofseedcollectedinthefieldwere

cooled2-6°Cduringtransport(p.117).

4. Kaye1997:Seedswerehand-cleaned,sorted,andstoredinpaperpacketsat7°C(p.116).

5. Lietal2007:Seedswereallowedtodryatroomtemperature

forabouthalfamonthandwerestoredinpaperbagsat4°Cinarefrigeratoruntilused(p.1259).

Pre-Planting 1. Alsosetal2012:Iftheplantswerewet,seedcapsuleswere

PropaguleTreatments

collectedandleftinpaperbagsat5-8°Cin35%relativehumidity(RH)todry.Seedsthatwereobviouslynotripewerenotcollected.Seedsofspecieswhichdidnotseem100%ripewereleftinpaperbagsat5-8°CuntilSeptember19thtopermitripening,afterwhichtheywereassumedtoberipe(p.820).

2. Bliss1958: Seedswereplacedinglassvialsorpaperbags,airdriedandshippedtoDukeUniversity,wheretheywerestoredat5°Ffor6to7months.Afterremovalfromstorage,theseedswerecleaned,sortedandtheunfilledandshrunkenonesremoved(p.181).

3. Cooperetal2004:Samplesofseedcollectedinthefieldwere

cooled2-6°Cduringtransportandstoredinpaperbagsat0.5°Cfor5to7weeks(p.117).

4. Kaye1997:Seedswerehand-cleaned,sorted,andstoredin

paperpacketsat7°C.Severaltreatmentstobreakdormancywereapplied:after-ripeningindrypaperpacketsat7°Cfor9months;darkness;scarification(abrasion)oftheseedcoat;and/ormoistcoldstratificationof10montholdseedfor6,12,and18weeksat5°C.Onlyasubsetoftreatmentswasappliedtoeachspecies(seeTable1onp.117).ForP.bracteosa,onlydarknessandstratificationweretested,butnogerminationnumberswerereported.Darknesswasappliedbywrappingtwolayersofaluminumfoilaroundtheplasticboxcontainingthepetriplates.Thedark-boxwasplacedforaminimumof18weeksonthesameshelfinthesamegerminatorastheotherseeds(p.116).

5. Lietal2007:Seedswereallowedtodryatroomtemperature

forabouthalfamonthandwerestoredinpaperbagsat4°Cinarefrigeratoruntilused.GerminationexperimentswereconductedinDecember2004(p.1259).Basedonstatisticalanalysis,scarifyingseedcoatwithsandpaperresultedinsignificantlyimprovedgerminationinalltestedPedicularisspecies(exceptP.lutescens).Scarificationwasthemosteffectivegerminationpromotingmethod,showingasignificantlypositiveeffectinsixofeightspecieswhencomparedwithothertreatments(p.1260).

GrowingAreaPreparation/AnnualPractices

1. Alsosetal2012:3to50seedsofeachspecieswereplacedona9cmdiameterpetridishwith10%agarsolution(p.821).

forPerennialCrops 2. Bliss1958:Seedswereplacedinsterilizedpetridishesbetweenlayersofmoistfilterpaperandincubatedat72°F(p.181).

3. Cooperetal2004:Soilsamplesfromthepolarheath,birdcliff,

proglacialhabitatandthermophilicheathwerespreadthinlyonfilterpaperinplasticpetridishes.SamplesfromintactanddisturbedDryasheathwereplacedoncommercialsterilizedsoilinaluminumfoilboxes.Unidentifiedseedlingsweretransplantedtoamixtureofpeatandperliteandgrownat15°Cuntilidentificationwaspossible(p.117).

4. Kaye1997:Dependingonseedavailabilityandtreatment,45–

300seedswereputonmoistcellulosepadsin10cmpetriplates,stackedina40x25x15cmclearplasticbox,andplacedina20°Cconstant-temperaturegerminatorequippedwithcool-whitefluorescentlightsonan8hourphotoperiod(p.116).

5. Lietal2007:Allgerminationtestswerecarriedoutonthree

layersoffilterpapersaturatedwithdistilledwaterintransparentglasspetridishes(p.1259).

EstablishmentPhaseDetails

1. Alsosetal2012:Thelighttemperaturewas4,000K(Osram35W,840HE)andthebrightnesswas3,300lm(manufacturer’sinformation).Theprotonfluxwasapproximately40umolpersquaremeterpersecondmeasuredwithaquantumfluxsensorattheleveloftheseeds.Ifthegerminationpercentageobtainedwaslow,buttheseedsstillseemedviable,anewgerminationtestwasattemptedfollowinganadditionalperiodofstratification(p.821).

2. Bliss1958:SeedswereplacedinsterilizedPetridishesbetweenlayersofmoistfilterpaperandincubatedat72°F,onesampleinthelightandtheotherinthedark.Distilledwaterwasaddedwhennecessarytokeepthefilterpapermoist.Thedisheswereremovedtwiceaweekforseedcounting,atwhichtimethecontinuousdarkperiodwasbrokenforapproximately1hour.Thenumberofseedstestedvariedaccordingtothequantityofseedavailable.Seedswereconsideredtohavegerminatedwhentheradicleappeared(p.181).

3. Cooperetal2004:Soilsamplesfromthepolarheath,birdcliff,

proglacialhabitatandthermophilicheathwerespreadthinlyonfilterpaperinplasticpetridishesandgerminatedat18°Cinagreenhouse,usinga24-hrphoto-period(150umol)over12weeks(11forthermophilicheath)simulatingthemaximum

Svalbardsummerperiod.SamplesfromintactanddisturbedDryasheathwereplacedoncommercialsterilizedsoilinaluminumfoilboxesandgerminatedat22°Cinagreenhouseover14weeks.Thesesampleswerestirredinweeks3and11,andgibberillicacid(1ppm)wasaddedinweek12.Allsamplesweremoistenedeverysecondday,andseedlingscountedweekly.Unidentifiedseedlingsweretransplantedtoamixtureofpeatandperliteandgrownat15°Cuntilidentificationwaspossible(p.117).

4. Kaye1997:Petriplateswereleftinthegerminatoruntilallseeds

hadgerminatedornonewgerminationoccurredforatleastfourdays.Themaximummonitoringperiodwas145days,butwaslessthan80daysformostspecies(p.116).

5. Lietal2007:Allgerminationtestswerecarriedoutonthree

layersoffilterpapersaturatedwithdistilledwaterintransparentglasspetridishes.Toguaranteeconstantmoistureinthedishes,theywereconnectedtoanextradishfilledwithdistilledwaterbyafilterpaperband.Twolightregimes(16hlight/8hdarkandcompletedarkness)andthreeartificialtreatmentswereusedinourexperiments.Forthetreatmentindarkness,thedisheswereputintoapaperboxwrappedwithaluminumfoil.Forartificialtreatments,seedswereeitherscarifiedbyrubbingagainst120-gritsandpaperuntiltheseedcoatwaspenetratedorwassoakedingibberellicacid(GA3;100

and500mgL-1,respectively)for2h.Alltestswerecarriedoutinagrowthchamber(Yi-HengTechnologyCo.,Ltd.,modelMGC-300A,seriesBluePard;Shang-hai,China)withacycleof16hlight(44.4mmolm–2s–1,400–700nm)at25°Cand8hdarkat18°C.Thistemperatureandlightregimewaschosenbasedonthetemperatureandlightfoundtoproduceoptimalgerminationinotherstudiesofhigh-elevationspecies.Eachtreatmentconsistedofthreereplicationsof20seeds(p.1259-1260).

LengthofEstablishmentPhase

1. Alsosetal2012:Noinformationlistedforhowlongseedsweregerminated.

2. Bliss1958:Seedswereconsideredtohavegerminatedwhentheradicleappeared(p.181).(theauthordoesnotlistatimeframeforgermination).

3. Cooperetal2004:Soilsamplesfromthepolarheath,birdcliff,

proglacialhabitatandthermophilicheathweregrownover12weeks(11forthermophilicheath)simulatingthemaximumSvalbardsummerperiod.SamplesfromintactanddisturbedDryasheathweregrownover14weeks(p.117).

4. Kaye1997:Themaximummonitoringperiodwas145days,butwaslessthan80daysformostspecies(Kaye1997,p.116).

5. Lietal2007:100%oftheseedsgerminatedafter10daysof

incubation(p.1260).ActiveGrowthPhase Noinformationfound.LengthofActiveGrowthPhase

Noinformationfound.

HardeningPhase LengthofHardeningPhase

Noinformationfound.

Harvesting,StorageandShipping

Noinformationfound.

LengthofStorage Noinformationfound.GuidelinesforOutplanting/PerformanceonTypicalSites

Noinformationfound.

OtherComments Somegreenhousetrialsfailedtomeetthegerminationrequirementsofsomespeciesthatsuccessfullygerminatedinthefield,suchasPedicularishirsute,possiblyduetoitshemiparasiticassociationwithS.polaris(Cooperetal2004).

INFORMATIONSOURCESReferences SeebelowOtherSourcesConsulted

Seebelow

ProtocolAuthor SageStowellDateProtocolCreatedorUpdated

May24th,2017

Workscited:JournalarticlesAlsos,I.,Müller,G.,&Eidesen,E.(2013).Germinatingseedsorbulbilsin87of113testedArcticspeciesindicatepotentialforexsituseedbankstorage.PolarBiology,36(6),819-83.

Bliss,L.(1958).SeedGerminationinArcticandAlpineSpecies.Arctic,11(3),180-188.Cooper,E.,Alsos,I.,Hagen,D.,Smith,F.,Coulson,S.,&Hodkinson,I.(2004).PlantrecruitmentintheHighArctic:SeedbankandseedlingemergenceonSvalbard.JournalofVegetationScience,15(1),115-124.Li,A.R.Guan,K.Y.Probert,R.J.(2007).EffectsofLight,Scarification,andGibberellicAcidonSeedGerminationofEightPedicularisSpeciesfromYunnan,China.HortScience,42(5):1259–1262.

BooksKaye,T.,&Oregon.StateDepartmentofAgriculture.(1997).SeedDormancyinHighElevationPlants:ImplicationsforEcologyandRestorationchapterinConservationandmanagementofnativeplantsandfungi:ProceedingsofanOregonConferenceontheConservationandManagementofNativeVascularPlants,Bryophytes,andFungi.(pp.115-120).Portland,OR:NativePlantSocietyofOregon.

Porsild,A.E.(1979).RockyMountainWildflowers.Ottowa,Canada;NationalMuseumsofCanada.

Pojar,J.,MacKinnon,A.,&Alaback,PaulB.(1994).PlantsofthePacificNorthwestcoast:Washington,Oregon,BritishColumbia&Alaska.Redmond,WA;Vancouver,Canada:LonePinePub.WebsitesBurkeHerbarium:Giblin,D.andKnoke,D.(n.d)Pedicularisbracteosa.Available:http://biology.burke.washington.edu/herbarium/imagecollection.php?Genus=Pedicularis&Species=bracteosa[AccessedonMay15th,2017] E-FloraBC:ElectronicAtlasofthePlantsofBritishColumbia.(n.d.)Pedicularisbracteosa.Availableat:http://linnet.geog.ubc.ca/Atlas/Atlas.aspx?sciname=Pedicularis%20bracteosa[AccessedonMay15th,2017]USDAPlantsDatabase.(n.d.)Pedicularisbracteosa.Availableat:https://plants.usda.gov/core/profile?symbol=PEBR[AccessedonMay15th,2017]Workscitedbutnotused:JournalArticlesAmen,R.(1966).TheExtentandRoleofSeedDormancyinAlpinePlants.TheQuarterlyReviewofBiology,41(3),271-281.

Bell,K.,&Bliss,L.(1980).PlantReproductioninaHighArcticEnvironment.ArcticandAlpineResearch,12(1),1-10.Bliss,L.,&Gold,W.(1999).Vascularplantreproduction,establishment,andgrowthandtheeffectsofcryptogamiccrustswithinapolardesertecosystem,DevonIsland,NWT,Canada.CanadianJournalOfBotany-RevueCanadienneDeBotanique,77(5),623-636.Cannone,N.,Lewkowicz,A.,&Guglielmin,M.(2010).Vegetationcolonizationofpermafrost-relatedlandslides,EllesmereIsland,CanadianHighArctic.JournalofGeophysicalResearch:Biogeosciences,115(G4).Cox,C.(1933).AlpinePlantSuccessiononJamesPeak,Colorado.EcologicalMonographs,3(3),299-372.Eriksen,B.,Molau,U.,&Svensson,M.(1993).ReproductivestrategiesintwoarcticPedicularisspecies(Scrophulariaceae).Ecography,16(2),154-166.Forbis,T.,Bowman,William,&Seastedt,Timothy.(2002).SeedlingEstablishmentonAlpineTundra:CommunityPatterns,PopulationDemography,andDisturbanceEffects,ProQuestDissertationsandTheses.Hagen,D.(2002).PropagationofnativeArcticandalpinespecieswitharestorationpotential.PolarResearch,21(1),37-47.Kikvidze,Z.(1993).Plantspeciesassociationsinalpine-subnivalvegetationpatchesintheCentralCaucasus.JournalofVegetationScience,4(3),297-302.Lindgren,Å,Eriksson,O.,&Moen,J.(2007).TheimpactofdisturbanceandseedavailabilityongerminationinalpinevegetationintheScandinavianmountains.Arctic,AntarcticAndAlpineResearch,39(3),449-454.Macior,L.(1970).ThePollinationEcologyofPedicularisinColorado.AmericanJournalofBotany,57(6),716-728.Marcante,S.,Winkler,E.,&Erschbamer,B.(2009).Populationdynamicsalongaprimarysuccessiongradient:Doalpinespeciesfitintodemographicsuccessiontheory?AnnalsofBotany,103(7),1129-1143.Molau,U.(1993).RelationshipsbetweenFloweringPhenologyandLifeHistoryStrategiesinTundraPlants.ArcticandAlpineResearch,25(4),391-402.Sayers,R.,&Ward,R.(1966).GerminationResponsesinAlpineSpecies.BotanicalGazette,127(1),11-16.

Schmidt,S.,Reed,S.,Nemergut,D.,Grandy,A.,Cleveland,C.,Weintraub,M.,Martin,A.(2008).TheEarliestStagesofEcosystemSuccessioninHigh-Elevation(5000MetresaboveSeaLevel),RecentlyDeglaciatedSoils.Proceedings:BiologicalSciences,275(1653),2793-2802.Tscherko,Hammesfahr,Zeltner,Kandeler,&Böcker.(2005).Plantsuccessionandrhizospheremicrobialcommunitiesinarecentlydeglaciatedalpineterrain.BasicandAppliedEcology,6(4),367-383.Williams,J.,&Batzli,G.(1982).PollinationandDispersionofFiveSpeciesofLousewort(Pedicularis)nearAtkasook,Alaska,U.S.A.ArcticandAlpineResearch,14(1),59-73.BooksDenverBotanicGardens,&AmericanRockGardenSociety.1986.RockyMountainAlpines:ChoicerockgardenplantsoftheRockyMountainsinthewildandinthegarden(1sted.).Portland,OR:TimberPress.Hulme,J.K.(1982).PropagationofAlpinePlants.London,England:AlpineGardenSociety.Nicholls,G.(2002).AlpineplantsofNorthAmerica:AnencyclopediaofmountainflowersfromtheRockiestoAlaska.Portland,OR:TimberPress.WebsitesBaskin,JerryM.;Baskin,CarolC..2002.PropagationprotocolforproductionofContainer(plug)PedicularisbracteosaBenth.Plants.UniversityofKentuckyLexington,Kentucky.In:NativePlantNetwork.URL:http://NativePlantNetwork.org(accessed2017/05/23).USDepartmentofAgriculture,ForestService,NationalCenterforReforestation,Nurseries,andGeneticResources.