taxonomy plant familycourses.washington.edu/esrm412/protocols/pebr.pdf · together with bryophytes...

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.