PHYTOSOCIOLOGICAL NOMENCLATURE SECTION

Lazaroa ISSN: 0210-9778

http://dx.doi.org/10.5209/LAZA.54043

SyntaxonomicalupdateontherelictgrovesofScotspine( Pinus sylvestris var. iberica )andSpanishblackpine( Pinus nigra subsp. salzmannii )inthe Gredosrange(central)

JoséAntonioLópez-Sáez 1,DanielSánchez-Mata 2 &RosarioG.Gavilán 2

Received: 30September2016/Accepted:4November2016

Abstract. Pureandmixedhigh-mountainnaturalpineforestvegetationoftheGredosrange(centralSpain)wasstud - iedusing57relevésandmultivariateanalyses(hierarchicalclusteranalysisanddetrendedcorrespondenceanalysis). Classificationoftherelevésresultedin7vegetationunits,3ofwhichwererankedasassociationsandonlyoneassub - association.DCAdiagramsofrelevésandtaxaindicatedthatfloristicdifferentiationwasattributedmainlytofactors suchasaltitude.Differentialtaxaofvegetationunitswerechosenbasedontheirphicoefficientvalues.Thestatistic discriminationofsomeofthesevegetationunitshasallowedustoproposeanewassociation( Genisto cinerascentis - Pinetum salzmannii )andanewsubassociationofothercommunitydescribedinapreviouspaper( Echinosparto bar - nadesii-Pinetum ibericae pinetosum salzmannii ). Keywords: Pinus nigra subsp. salzmannii ; Pinus sylvestris var. iberica ;Gredosrange;Phytosociology;Multivariate Analyses;CommunityEcology.

[es]Estudiosintaxonómicodelosbosquesrelícticosdepinosilvestre( Pinus sylvestris var. iberica )ypinonegro( Pinus nigra subsp. salzmannii )enlasierrade Gredos(centrodeEspaña)

Resumen. SehanestudiadopinaresnaturalespurosymixtosenlaaltamontañadelaSierradeGredos(centro deEspaña)mediante57inventariosflorísticosyanálisismultivariantes(análisisdeclusterjerárquicoyanálisis decorrespondenciassintendencia).Laclasificacióndelosinventariosdeparó7unidadesdevegetación,3delascuales fueronclasificadasaniveldeasociaciónyotracomosubasociación.LosdiagramasdelDCAdeinventariosytaxones indicanqueladiferenciaciónflorísticapuedeatribuirsemayoritariamenteafactorestalescomolaaltitud.Lostaxones diagnósticosdecadaunidaddevegetaciónfueronelegidosdeacuerdoalosvaloresdesucoeficientephi.Ladiscrimina- ción estadísticadelasunidadesdevegetaciónnoshaceproponerunanuevaasociación( Genisto cinerascentis - Pinetum salzmannii )yunanuevasubasociacióndeunacomunidaddescritaenuntrabajoanterior( Echinosparto barnadesii-Pinetum ibericae pinetosum salzmannii ). Palabras clave: Pinus nigra subsp. salzmannii ; Pinus sylvestris var. iberica ;sierradeGredos;Fitosociología;análi - sismultivariantes;ecologíadelascomunidadesvegetales.

Introduction nity-levelconsequences(Soto& al. ,2010).In theIberianPeninsula, Pinus nigra subsp. Pinespeciesareimportantcomponentsof salzmannii and P. sylvestris areamongthe manynaturalecosystemsacrossthe commonestspeciesinmountainousareas, Mediterranean,especiallyinmountainregions bothintheAtlanticandtheMediterraneanbio - (Barbero& al. ,1998),andtheirhistoricaland geographicalregions(Blanco Castro& al. , geneticdynamicsmaythereforehavecommu - 1997).TheinterestintheGredosmountain

1 GrupodeInvestigaciónArqueobiología,InstitutodeHistoria,CCHS,CSIC.Albasanz26-28.E-28037Madrid.Spain.E-mail: [email protected] 2 DepartamentodeBiologíaVegetalII,FacultaddeFarmacia,UniversidadComplutense.E-28040Madrid.Spain.E-mail:dsma - [email protected],[email protected]

Lazaroa 37 2016: 153-172 153 154 López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172 populationsofbothpinespeciesisheightened throughouttheHolocene(Rubiales& al. , bytheircurrentfragmenteddistribution 2007,2012;López-Sáez& al. ,2014),suggest - (Génova& al. ,2009),sincepredictedgenetic ingthepermanenceofwidespreadhigh-moun - anddemographicconsequencesofhabitat tainpinewoodlandsinthesemountainsat fragmentationmightleadtoaprogressiveloss leastsincethemid-Holocene,aswellasits ofgeneticdiversityandtoincreasingdifferen - progressivedisappearanceinthelasttwothou - tiationamongpopulations,loweringtheirevo - sandyearsasaresultoffires,loggingandhigh lutionarypotentialtorespondtoenvironmen - pastoralpressure(López -Merino& al. ,2009; talchanges(Benito Garzón& al. ,2006, López -Sáez& al. ,2009,2016a,2016b). 2008a,2008b;Gea -Izquierdo& al. ,2015). SpanishblackpineforestsoftheGredos Blackpine( Pinus nigra s.l.)isthemost rangeareknownforalongtime(Gómez widelydistributedpinespeciesinhigh-alti - Manzaneque,1988;Regato & al. ,1992), tudeareasintheMediterraneanBasin; Pinus althoughnophytosociologicalapproachhas nigra subsp. salzmannii occursincentraland beendonesofar.OnlyGalándeMera& al. easternSpainandsouthernFrance(Catalán& (1999)haveconductedafirstapproximation al. ,1991).IntheIberianPeninsula,Spanish oftheircommunities,includingthemwithin blackpineforestsaremainlyfoundinthe the Vaccinio-Piceetea class, Pinetalia supramediterraneanbioclimaticbeltunder sylvestris orderand Cytision oromediterranei humidorsub-humidclimates(900-1500m alliance,butwithoutdescribinganyownphy - asl),althoughtherelictpopulationsofthe tosociologicalassociationoftheseforests. SpanishCentralSystemongraniticrocks Thesefactssharplycontrastwiththerestof (GredosandGuadarramaranges)arealso Spanishblackpineforestcommunitiesinthe foundundercoldperhumidclimate(Regato& IberianPeninsula,forwhichvariousassocia - al. ,1991,1992,1995;Regato & Del Río, tionsandsubassociationshavebeendescribed 2009).Scotspine( Pinus sylvestris s.l.)isa inthePyrenees,theIberianandBaeticranges Eurosiberianforesttreespeciesgrowingina (Vigo,1979;Gamisans & Gruber,1988;Valle largevarietyofecologicalconditionsfrom & al. ,1988;Regato & Escudero,1989; WesternEuropetoAsiaandthemostwidely Gamisans& al. ,1991;Regato & al. ,1995; spreadamongpinespecies(Ruby & Wright, Regato & Elena Rosselló,1995;Rivas - 1976).Itsnaturalrangeextendsnorthwards Martínez& al. ,2001,2002,2011).Itisalso fromwarmsouthernareasofSpainandthe notedthatthehighstateofdegradationof mountainsofnorthernIran(37ºN)toarticlat - Pinus nigra subsp. salzmannii forestsofthe itudesinFennoscandia(71ºN),andwestwards Gredosrangehassofarhinderedanyattempt fromlongitudes6°W(ScotlandandIberian tosyntaxonomicdescription(Regato & al. , Peninsula)to135°EinSiberia(Boratynski, 1992;Regato & DelRío,2009).Asimilarsit - 1991).Whilemostofitsnaturalrangeis uationhasoccurredwithScotspineforestsof continuous,t herearealsolargepopulationsof theGredosrange,althoughsomecommunities Scotspinedisconnectedfromthemainnatural throughouttheIberianPeninsulahavebeen rangesuchasintheIberianPeninsula,where describedinthePyrenees,thenorthernand currentlythisspeciesinhabitsdisjunctmoun - southernIberianrange,BazaandSierra tainousareas(700-2600masl)withlargedis - NevadamountainsintheBaeticrange(Rivas- tancesbetweenpopulations(Gil& al. ,1991; Martínez,1987;Loidi& al. ,1997a,1997b; Alía& al. ,2001). Rivas -Martínez& al. ,2001,2002).Although Geobotanicalinterpretationofhigh-moun - Pinus sylvestris forestsfromtheSpanish tainpineforestsoftheGredosrangehaveled CentralSystem( P. sylvestris var. iberica )have tocontroversialinterpretationsandpositions hardlybeentreatedintothephytosociological betweenthosewhoaccepttheirnaturalchar - literature,withsomeexceptionsforthe acterandthosewhorule(Costa& al. ,1990; Ayllón,GuadarramaandPelaranges(Dela Alcalde& al. ,2006).Thisdebateisstillongo - Cruz&Peinado,1996;Rivas-Martínez& al. , ingdespiteagrowingbodyofevidenceindi - 2001,2002),recently,thecombinationof catingthenativeoriginofpineforestsfrom modernpollenstudiesandfloristicrelevéshas historicalandpalaeoecologicalstudies helpedtodefinetwophytosociologicalassoci - (Génova& al. ,1992,2009).IntheGredos ationscomprisingtheScotspineforestsofthe range,macrofossilandpalynologicalrecords Gredosrange(López -Sáez& al. ,2013). showthepresenceofbothpinespecies Neverthelessthegeobotanicalrecognitionof López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172 155

Scotspineforestsasaseriesofvegetationin wholeofthismassif,whichseparatesthe theGredosmountainswashighlightedby basinsoftheDouroandTagusrivers(Figure Rivas-Martínez& al. (2011),forcommunities 1).Thisrangeisachainofsunkenandelevat - locatedonthesupramediterraneanand edblocksofgraniticandmetamorphiclitholo - supratemperate(submediterranean)bioclimatic gy,separatedbyintramontanevalleys.The belt. tectonicconstitution,atiltedhorst,withafault Inthiswork,ouraimsareto(i)determine orientatedtowardsthesouth,imposesaclear theplantcommunitiesof Pinus nigra Arnold asymmetryinboththenorthernandsouthern subsp. salzmannii (Dunal)Francoand Pinus faces.TheclimateisofaMediterraneantype, sylvestris L.var. iberica Svobodaforestvege - dryandcoldinwinter(0°C-2°C)anddryand tationintheGredosrange,and(ii)definetheir warm(20°C-22°C)insummer,withasummer ecologicalandgeographicalgradients. droughtperiodlasting3-5monthsandheavy rainfallinautumnandwinter.Precipitationis Material and Methods onaverage1400mmperyear.Thestudyarea belongstotheBejaran-Gredenseansector, Study area Carpetan-Leonesebiogeogaphicalsub - province,IberianWesternMediterranean TheGredosrangeislocatedatthewestern province,WesternMediterraneansubregion, sectoroftheSpanishCentralSystem,showing Mediterraneanregion(Rivas -Martínez& al. , thehighestaltitudes(>2000masl)ofthe 2007;Sánchez-Mata,2015).

Figure1. LocationofthestudyareaintheGredosrange(centralSpain).Forlocatingeachrelevéseethe Materialandmethodssection,Tables4to6. 156 López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172

InGredosrange, Pinus sylvestris var. iber - wereobtainedviafieldsurveysconducted ica forestsarepartof(sub)mediterraneanpine fromFebruary2010untilOctober2015,cov - forests(GonzálezMolina,2006).Thispine eringtheentireareaandknownlocationsof speciesappearsveryunevenlyonthesouthern bothspecies.Theremainingrelevés(8)were slopesofthecentralmassifofthemountain collectedfromearlierstudies(López-Sáez& andsporadicallyintheeasternone(Molina al. ,2013).Thephytosociologicalapproach Moreno,1992;Mancebo& al. ,1993),consti - (Braun-Blanquet,1979)wasusedtoobtain tuting,withsomeexception,extremelysmall relevés,includingtopographicandotherdata isolatedforestsalteredbythefireregimeand ofindividualplots,suchasaltitude,longi - pastoralpressureinthesupra-andorosub - tude,latitudeandaspect.Additionallyalder mediterraneanbioclimaticbelts(López-Sáez (Alnus glutinosa ),maritimepine( Pinus & al. ,2009,2014).However,onthenorthern pinaster s.l.)andPyreneanoak( Quercus slopeofGredosrange,itcoversanareaof pyrenaica )forestswhichlocallyinclude about2000haintheupperbasinsofthe relictSpanishblackpineswerealsosampled. AlbercheandTormesrivers,whereitforms Wechosehomogenoussamplingplotswith notablewoodlandsinHoyosdelEspino, anareaof100m 2. Navarredondaandinthe Thelocationofeachrelevéisreferredin supramediterraneanandsuprasubmediter - Tables4and5,exceptforthefollowing raneanbioclimaticbelts(Génova& al. ,2009; relevés:18,ArenasdeSanPedro,RíoPelayos, Sánchez-Mata,2015).Inadditiontothepure 530masl;19,,MonteDehesa forests,italsoformsmixedforestswith Pinus Avellaneda,805masl;23,ElArenal,Collado nigra subsp. salzmannii dependingonecolog - delaVuelta,1,175masl;39,Piedralaves, icalgradients(López-Sáez&López-García, GargantadeSerradilla,1350masl;41, 1994;Regato& al. ,1995). Piedralaves,RobledodelaAngostura,1240m Spanishblackpinepopulationsofthe asl;48,ElTiemblo,ArroyodelPuerto,880m SpanishCentralSystemareuniquetopeninsu - asl;49,ElTiemblo,CerrodelMoro,915m larlevelforbeingtheonlyonesexclusively asl;50,ElTiemblo,CasadeLasLamos,1040 developedonsiliceoussubstrates(granitic masl;and,51,ElTiemblo,ChozaLumbrero, rocks,gneissesandmicaschists)(Génova& 1180masl. Moya,2012).TheirpopulationsintheGredos rangemarkthewesternlimitofthespecies Statistical analyses throughoutitsrange(Regato& al. ,1995).In thesemountains, Pinus nigra subsp. salzman - Inordertoestablishecological,floristicand nii appearssparselyasisolatedpinecopses- phytogeographicalrelationshipsamong sometimeswithinmaritimepine( P. pinaster relevéswehavecarriedoutmultivariateanaly - s.l.)orPyreneanoak( Quercus pyrenaica ) sesbasedonclassificationandordinationof forests-onthesouthernslopesofthecentral vegetationdata. andeasternmassifs(Regato& al. ,1992), Classificationwasperformedbyhierarchi - althoughsomeisolatedpopulationsdevelop calclusteranalysis(HCA)withtheprogram onthenorthernslopesintheIruelasvalleyas IBMSPSSStatistics21usingthematrixofthe wellasaveryimportantone(40ha)inthe EuclideandistanceandWard’sminimumvari - municipalityof(Gandía& al. , ancemethod(Ward,1963),whichhasbeen 2005;Arias & al. ,2011).Somehow,itcanbe showntooptimizetheinternalhomogeneityof saidthatthispineintheGredosrangehasa clustersandtoestablishclearerdichotomies rupicolousvocation,asinmanyotherpenin - amongthem,aswellastoclassifycontinuous sularterritories(Regato& al. ,1991),occupy - variablesbetter(Wildi,1989;Escudero& al. , ingrockyoutcropsandsteepslopesoflittle 1994). edaphicdevelopment. Detrendedcorrespondenceanalysis(DCA) wasperformedusingCANOCO4.5(terBraak Data collection &Šmilauer,2002).Detrendingbysegments, square-roottransformationofthecoverabun - Atotalof57relevéscomprehendingdifferent dancesoftaxaanddownweightingofrare formationsof Pinus nigra subsp. salzmannii specieswereapplied.Althoughthelongestgra - and P. sylvestris var. iberica intheGredos dientisshorterthan3S.D.(Table1),theuni - rangewereanalysed(Figure1).Fortynine modalmethodisalsoconsideredasanappropri - López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172 157 ateanalysisofcommunitycompositionaldata, pilationsof Flora Europaea (Tutin& al. , eveninthecaseofshortenvironmentalgradi - 1964-1980)fortaxanotincludedinthefor - ents(Lepš&Šmilauer,2003).Furthermore,in mer;theexceptionsarespecificallymen - theDCA,severalvariableswereusedaspassive tionedinthefloristicappendix. explanatoryonesaimingtosupporttheinterpre - Syntaxonomicalscheme,nomenclature,and tationoftheordinationdiagram.Corre lations syntaxaauthoritiesfollowthecompilations betweenDCArelevéscoresandexplanatory andproposalsofRivas-Martínez& al. variableswereanalysedbySpearman’srank (2001,2002,2011). correlationwith STATISTICA 9.1software (http://www.statsoft.com;Table2). Results Todefinediagnosticspeciesforeachvege - tationtype,weusedthephicoefficient( Φ)of The57relevés(hereinaftersamples)were fidelity(Chytrý& al. ,2002).Asdiagnostic classifiedbymeansofHCAintosevensample species(Table3),weusedonlyspecieswith Φ groups,whichcorrespondtoecologicallyand ≥0.25andastatisticallysignificant( p<0.01) phytogeographicallyinterpretablevegetation associationwithaparticularvegetationtype units.Differentlevelsofdivisionwereaccept - accordingFisher’sexacttest,performedinthe ed.Asshownbythedendrogramanalysis JUICE7.0program(Tichý,2002).Ifataxon (Figure2),twomainclustersemerged(Aand hadafidelityvaluehigherthan25%inonly B),correspondingtothesegregationofthe onesynoptictable,thenitwasconsideredas twomainvegetationunitscomparedinthis diagnosticofthevegetationunitofthehierar - work,whereSpanishblackpineandScots chicallevelrepresentedbythecorresponding pineforestsintheGredosrangewereseparat - synoptictable,whereasifthesamevalueis ed.Whilethreeoftheseclustersrepresentthe giveninmorethanonesynoptictable,thenit Pinus sylvestris var. iberica forests,thelast wasconsideredasdifferentialofthehierarchi - fourareformedbypureormixed P. nigra callevelinwhichitpresentedthehighest subsp. salzmannii communities.ClusterA fidelityvalue. makesreferenceto Pinus sylvestris var. iberica Traditionally,thephicoefficientconsiders woodlandsandclusterBmakereferencetothe onlythepresence/absenceoftaxa,sothat remainingwoodlandscontaining P. nigra fidelityvaluescalculatedusingthiscoeffi - subsp. salzmannii ,withatruncationlevelof cientarenotinfluencedbyspeciescoveror 26.25. abundance(Tichý & al. ,2011).Itsmain advantageititsindependenceofdatasetsize Table1. ResultsofDetrendedCorrespondence (Tichý,2002).Thehighest Φ value(1)is Analysis(DCA)of57relevéshosting98species. achievedifthespeciesoccursinallrelevés ofthevegetationunitandisabsentelse - Axes Axis1 Axis2 where.Apositivevalue<1meanthatthe Eigenvalues 0.131 0.067 speciesisabsentfromsomerelevésofthe Lengthsofgradient 1.013 1.060 vegetationunitorispresentinsomerelevés Cumulativepercentage 21.3 32.1 outsidethevegetationunit.Furthermore,we varianceofspeciesdata usedanadditionalcriteriontoscreenofftaxa withasmoothdecreaseoftheirpercentage InsideclusterA,wecandetach,atasmall - constancy(Table3).A Φ valueof0is errange(17.10),thesub-clustersA1andA2; obtainedwhentherelativefrequencyofthe respectivelyseparatingorotemperatesub - species(constancy)inthevegetationunit mediterranean(oceanic)Scotspinecommuni - equalstherelativefrequencyintherestof ties(clusterA1)fromthoseofthesupratem - thedataset,thusindicatingnorelation peratesupramediterranean(oceanic)ones betweenthetargetspeciesandthetarget (clusterA2).Thefollowingthresholdofclus - vegetationunit(Tichý&Chytrý,2006). terA1(truncationlevelof11.15)clearlysepa - ratessamplescontainingSpanishblackpine Nomenclature (clusterA1b)fromthosewithout(cluster A1a).RelatingtoclusterB,themajorsegrega - Taxonomicnomenclatureandauthoritiesfol - tionattruncationlevel6.77,separatesB1and lowthepublishedvolumesof Flora iberica B2clades.Themaindisjunctionpromoted (Castroviejo& al. ,1986-2015)andthecom - hereisduetothesignificantpresenceofother 158 López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172

Figure2. Hierarchicalclusteranalysis(HCA)ofthe57relevésofhigh-mountainpineforestsofthe Gredosrange.Dottedlinesindicatethedifferentthresholdsthatsubdividethedendrogrammentionedin thetext,fromrighttoleft. López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172 159

Figure3. Detrendedcorrespondenceanalysis(DCA-biplot)ofthe57investigatedrelevés(a)andselected species(b)intheGredosrange.Vegetationunitsrepresentingtheplantcommunitiesdefinedbycluster analysis(Figure2)aremarkedbydifferentsymbolsandframed.Explanatoryvariablesdescribingthe ecological backgroundofthefirsttwoaxes(displayedbyarrows)wereplottedaposteriori.Weuse‘short writtingmode’forthefollowingtaxa: Carex elata subsp. reuteriana (C.reuteriana), Festuca elegans subsp. merinoi (F.merinoi), Festuca paniculata subsp. multispiculata (F.multispiculata), Juniperus communis subsp. alpina (J.alpina), Leucanthemopsis pallida subsp. alpina (L.alpina),and Potentilla alpina subsp. apiifolia (P.apiifolia). 160 López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172 forestspeciesinclusterB1,whilethecluster negativelycorrelatedtoDCAaxis2,although B2correspondstopureSpanishblackpine theyarenotsignificantvariablesinanyofthe forests.TheunderclustersB1atoB1c,respec - twoaxes.InFigure3bthescoresofthediag - tivelysegregateotherforestformationsthat nostictaxaofclusterB1c( Alnus glutinosa , containsporadicallysomeSpanishblack Salix atrocinerea , S. salviifolia , Carex elata pines: Quercus pyrenaica woodlands(cluster subsp. reuteriana , Ruscus aculeatus and B1a), Pinus pinaster s.l.forests(clusterB1b), Origanum vulgare )areplottedatthetopof and Alnus glutinosa ripariancommunities DCAaxis2,probablyindicatingthatthisaxis (clusterB1c). mayhavesomerelationshiptoenvironmental IntheordinationdiagramsofFigure3the orsoilmoisture. firsttwoDCAaxeswereplotted.Theeigen - valueofthefirstDCAaxiswas0.131andits Table2. Spearmanrankcorrelationscoefficients lengthwas1.013,whilethecorrespondingval - (p<0.01)betweenthesamplescoresontheDCA uesforthesecondDCAaxiswererespective - axes1and2andenvironmentalvariables. ly0.067and1.060(Table1).BothDCAaxes Variable Axis1 Axis2 accountfor32.1%ofthecumulativepercent - Altitude -0.69 0.14 agevarianceofspeciesdata(totalinertiaequal Aspect 0.04 0.18 to0.616). Latitude 0.003 -0.07 Spearmanrankcorrelationanalysis Longitude -0.007 -0.06 betweensamplescoresfromtheDCAanalysis andtheenvironmentalvariables(Table2) revealedthataltitudehadthehighestcorrela - Theclassificationofthesevenclusters,with tionwiththesamplesites(correlationwiththe statisticallydetermineddiagnosticspeciesfor firstDCAaxis: r=-0.69, p<0.01;Figure3a).In eachcluster,ispresentedinTable3. fact,intheDCAordinationdiagramofthe relevés,thereisaclearaltitudinalgradient Discussion and Conclusions alongtheDCAaxis1:orosubmediterranean Pinus sylvestris var. iberica and P. nigra Inaccordancewiththeclassificationanalysis subsp. salzmannii dominatedforestsappearat (Figure2),itseenthatthehigh-mountainfor - theleftpartoftheaxis(clustersA1aandA1b), estvegetationintheGredosrangeincludes whilesuprasubmediterraneanones(clusters sevendifferentplantcommunities.Twoof A2andB2)takeplacethroughtherightpartof themaredominatedby Pinus nigra subsp. theaxisrespectively.Spearmanrankcorrela - salzmannii (clustersA1bandB2),oneof tioncoefficientalsorevealedthataltitudehas which(clusterA1b)iscodominatedby P. asignificanteffectonvegetationalongthe sylvestris var. iberica .PureScotspineforests axis1(Figure3b): Pinus sylvestris var. iberi - compriseclustersA1aandA2,whicharesep - ca , Cytisus oromediterraneus , Echinospartum aratedintheDCAordinationdiagramalongan barnadesii , Nardus stricta and Juniperus com - altitudinalgradientintheDCAaxis1(Figure munis subsp. alpina arelocatedontheleftside 3a).Finally,clusterB1representsforestfor - oftheDCAaxis1; P. nigra subsp. salzmannii , mationsdominatedbyothertreespecies- Festuca elegans subsp. merinoi ,and Genista Quercus pyrenaica (B1a), Pinus pinaster s.l. cinerascens inthemiddleofit;and,finally, (B1b)and Alnus glutinosa (B1c)-thatcan Quercus pyrenaica , P. pinaster s.l., Daphne promptlycontainSpanishblackpines( P. gnidium and Lavandula pedunculata ,among nigra subsp. salzmannii ).Themainfactor others,ontherightpartofDCAaxis1. affectingthefloristicdifferencesbetween Thesecondaxiswasweaklycorrelated communitiesintheGredosrangeisaltitude, withaspect( r=0.18)andaltitude( r=0.14; whichissigniticantlycorrelatedwithDCA Table2),whichdoesnothaveameaningful axis1(Table2),reflecting,amongothers,a explanationonthevegetationdifferentiation. temperatureandprecipitationgradient. AlongDCAaxis2,samplesfromclusterB1c ClusterA1arepresentsxerophilous, (alderforests)werelocatedatthetopofthe orotemperatesubmediterranean(orosub - ordinationdiagram,clustersA1a,A1b,B2, mediterranean)hyperhumidrelictScotspine B1aandB1binthemiddle,andclusterA2 mesoforestsfromthemiddleoromediter - withsomesamplesfromclusterB2atthebot - raneanbeltoftheGredosrange(~1600-1800 tom(Figure3a).Latitudeandlongitudewere masl).RelevésfromclusterA1ascored López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172 161

Table3.Synoptictableof57relevésfromhigh-mountainpineforestsoftheGredosrange,basedonfidelity phicoefficient( Φ x100)andpercentageconstancyofspeciesforeachcluster(relevégroups).Fidelity wascomputedusingthephicoefficientappliedtoadatasetwiththesizeofvegetationunits(clustersA andB)equalizedto0.15%ofthesizeoftheentiredataset.Diagnosticspecies(valuesinbold)arethose with Φ ≥25;theyarerankedbyadecreasingvalueof Φ.Dashesindicatenegativefidelityorpositivebut non-significantfidelityat p< 0.01.Superscriptvaluesindicatepercentagefrequency(constancy)of species;dotsindicatespeciesabsence.Onlyspecieswithaconstancy>25%arelisted.

Cluster A1a A1b A2 B2 B1a B1b B1c Numberofrelevés 8 10 9 21 3 3 3 Diagnosticspeciesofcluster A: Pinussylvestris var. iberica 43.5 100 44.5 100 44.5 100 --- 5 --- . --- . --- . Diagnosticspeciesofcluster A1: Echinospartumbarnadesii 56 100 57.6 100 --- . --- 14 --- . --- . --- . Juniperuscommunissubsp. alpina 51.8 88 65.2 100 --- . --- 5 --- . --- . --- . Cytisusoromediterraneus 40.7 100 41.4 100 --- 78 --- 23 --- . --- . --- . Diagnosticspeciesofcluster A1a: Cytisusstriatus subsp. eriocarpus 92.5 88 --- . --- . --- . --- . --- . --- . Armeriacaespitosa 75.2 100 --- 11 --- . --- 18 --- . --- . --- . Nardusstricta 47 25 --- . --- . --- . --- . --- . --- .

Leucanthemopsispallidasubsp. 75 11 56 14 33 33 . alpina 40.6 ------Arrhenatherumelatius s.l. 36.5 100 --- 33 --- 56 --- 68 --- . --- 33 --- . Agrostiscastellana 27 100 --- 11 --- 44 24.7 91 --- 100 --- 67 --- 67 Diagnosticspeciesofcluster A1b: Corynephoruscanescens --- . 39.1 56 --- . --- 27 --- . --- . --- . Agrostisdelicatula --- . 30.5 78 --- . 26.3 64 --- 67 --- 33 --- . Diagnosticspeciesofcluster A2: Senecioadonidifolius --- . --- . 63.7 44 --- . --- . --- . --- . Hieraciumsabaudum --- . --- . 63.7 44 --- . --- . --- . --- . Spergulamorisonii --- . --- . 63.3 56 --- 5 --- . --- . --- . Lotuscorniculatus s.l. --- . --- . 57.5 89 --- 23 --- 67 --- 33 --- . Santolinaoblongifolia --- . --- 11 55.8 78 --- 23 --- . --- . --- . Anthericumliliago --- . --- . 54.7 33 --- . --- . --- . --- . Achilleamillefolium --- . --- . 53.4 100 --- 45 --- 33 --- 67 --- . Genistaflorida --- . --- . 53.4 100 --- 41 --- 100 --- 33 --- . Hypochoerisradicata --- . --- . 49.5 100 20.5 59 --- 67 --- . --- . Conopodiumpyrenaeum --- . --- . 46.5 56 --- 9 --- 33 --- 33 --- . Silenenutans --- . --- . 45.6 89 --- 50 --- 33 --- 33 --- . Violariviniana --- . --- . 44.4 67 --- 32 --- . --- . --- . Ornithogalumconcinnum --- . --- . 44.4 67 --- 27 --- 33 --- . --- . Ilexaquifolium --- . --- . 44.2 22 --- . --- . --- . --- . Stemmacanthaexaltata --- . --- . 44.2 22 --- . --- . --- . --- . Monotropahypopytis --- . --- . 44.2 22 --- . --- . --- . --- . Centaureaamblensis --- . --- . 44.2 22 --- . --- . --- . --- . Euphorbianevadensis --- . --- . 44.2 22 --- . --- . --- . --- . Potentillaalpinasubsp. apiifolia --- . --- . 44.2 22 --- . --- . --- . --- . Tanacetumcorymbosum --- . --- . 44.2 22 --- . --- . --- . --- . Seneciojacobaea --- . --- . 44.2 22 --- . --- . --- . --- . Pilosellacastellana --- . --- . 43.7 89 --- 50 --- 67 --- 33 --- . Arenariamontana --- . --- 44 43.7 89 --- 45 --- . --- . --- . Anthoxanthumodoratum --- . --- 33 41.8 89 --- 45 --- 33 --- 33 --- . Cytisusscoparius --- . --- . 37.2 100 --- 68 --- 100 --- 100 --- 67 Jasionemontana subsp. echinata --- . --- 22 32 89 25.6 73 --- 100 --- . --- . Pteridiumaquilinum --- 50 --- 11 27.4 100 --- 82 --- 100 --- 100 --- 67 DiagnosticspeciesofclusterB2: Lactucavimineas.l. --- . --- 11 --- . 56.2 50 --- . --- . --- . 162 López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172

Helianthemumapenninum s.l. --- . --- . --- . 53.3 32 --- . --- . --- . Avenulasulcata --- . --- 33 --- . 47.1 64 --- 33 --- . --- . Lavandulapedunculata --- . --- . --- 11 47.1 64 --- 67 --- 33 --- . Dianthuslusitanus --- . --- . --- . 44.3 36 --- . --- 33 --- . Genistacinerascens --- . --- 22 28.5 100 36.8 100 --- 100 --- 100 --- . Pinusnigra subsp. salzmannii --- . 27.4 100 --- . 35 100 --- 100 --- 100 --- 100 Sedumforsterianum --- . --- 11 --- . 31.5 32 --- 33 --- . --- . Armeriaarenaria --- . --- . --- . 30.2 23 --- 33 --- . --- . Koeleriacrassipes --- . --- 33 --- . 28.2 50 --- 33 --- 67 --- . Festucapaniculata subsp. --- . --- 44 --- 67 26.6 77 --- 100 --- 33 --- . multispiculata Festuca elegans subsp. merinoi --- 25 --- 33 25.1 100 26.4 95 --- 100 --- 100 --- 33 Pilosella pseudopilosella --- . --- 44 --- . 24.5 41 --- 33 --- . --- . Clinopodiumvulgare --- . --- 44 --- . 20.8 41 --- 33 --- 33 --- . Rubus ulmifolius --- . --- 11 --- 11 19.7 55 --- 100 --- 67 --- 100 Diagnostic species of cluster B1a: Quercus pyrenaica --- . --- . --- . --- 23 73.6 100 --- 33 --- . Galiumrotundifolium --- . --- . --- . --- 55 54.5 100 --- 67 --- . Rosa canina --- . --- . --- 11 --- 18 49.9 67 --- 33 --- . Diagnostic species of cluster B1b: Pinuspinaster s.l. --- . --- . --- . --- 55 --- 67 49.3 100 --- 33 Daphnegnidium --- . --- . --- . --- 18 --- 67 49.9 67 --- . Diagnostic species of cluster B1c: Alnusglutinosa --- . --- . --- . --- . --- . --- . 100 100 Salixatrocinerea --- . --- . --- 11 --- . --- . --- . 94.2 100 Salixsalviifolia --- . --- . --- 11 --- . --- . --- . 94.2 100 Carexelata subsp. reuteriana --- . --- . --- 11 --- 5 --- . --- . 89.1 100 Ruscusaculeatus --- . --- . --- 11 --- 5 --- . --- 33 84.6 100 Origanumvulgare --- . --- 11 --- . --- 9 --- . --- . 61.4 67 chieftlybetween-0.11-0.12S.D.unitsalong munityisonlyrepresentedonthesouthern DCAaxis1(Figure3a),beingforestcommu - slopebyisolatedandscatteredsmallforestsor nitiesemplacedathigherelevationsofthe copsesmostlyaffectedbyfiresandgrazing study.Diagnosticspeciesofthecommunity fromprehistory(López-Sáez& al. ,2014, are Pinus sylvestris var. iberica , Juniperus 2016a,2016b).López-Sáez& al. (2013)pre - communis subsp. alpina , Cytisus oromeiterraneus , sentedfourrelevés,southeastoriented,inthe C. striatus subsp. eriocarpus , Armeria surroundingsummitsofPuertodelPicoPass caespitosa , Nardus strict a, Leucanthemopsis (LaRubía)fromthecentralmassifofthe pallida subsp. alpina , Arrhenatherum elatius GredosRange(1710-1800masl).Inthis s.l.and Agrostis castellana (Figure3b,Tables paperfournewinventoriesfromtheeastern 3and4),aswellassomewesternendemicele - massifofthesemountainsarepresented(Table mentsuchas Echinospartum barnadesii 4).TwocomefromCerrodePinosequilloin (Sánchez-Mata,1989,1999;DelaFuente& themunicipalityofSotillodelaAdrada(1700- Ortúñez,2001).Thisclustercorrespondsto 1710masl),andtwomorefromCabezaPozo the Echinosparto barnadesii-Pinetum ibericae (Piedralaves;1340-1400masl).Intheeastern communitydescribedbyLópez-Sáez& al. massifoftheGredosrangethiscommunity (2013),representingtheScotspineforestsof appearsatloweraltitudethaninthecentral thetimberline(Sánchez-Mata,1999,2015). one(evenbelow1400masl),compensating Theseforestsareconsideredasxerophilous theirsituationwithapredominantwesternori - becausetheyarecoveredbywintersnowforat entationmoreinfluencedbythehumidwinds least4-5monthsandgrowonmoderateto fromtheAtlanticOcean(Gavilán & al. ,1998; highlystonysoilswithlowedaphicdevelop - Gavilán,2005). ment(Gavilán& al .,1998;Martínez-García& Asaresultofmultivariateanalyses(HCA Montero,2000;Gavilán,2005;López-Leiva andDCA),orosubmediterranean Pinus nigra & al. ,2009).IntheGredosrange,thiscom - subsp. salzmanni forestcommunitiesfromthe López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172 163

Table4. Echinosparto barnadesii-Pinetum ibericae Sánchez-Mata,Gavilán&López-SáezinLópez-Sáez& al. 2013 typicum (1-8); pinetosum salzmannii subass. nova (9-18) (Avenello ibericae-Pinion ibericae , Pino sylvestris-Juniperetalia sabinae , Pino-Juniperetea )

Altitude(1=10m) 177 180 175 171 140 135 170 1712 148 143 154 157 154 157 141 165 149 146 Aspect SE SE SE SE NW NW W SW SE SE SE SE SE SE SE SE SE SW Samplenumber(Fig.3) 1 2 3 4 11 12 14 15 25 26 27 29 30 33 35 36 43 46 Numberofspecies 15 15 17 20 12 15 14 14 17 15 11 11 13 16 22 22 17 28 Relevénumber 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Characteristics

Pinussylvestris var. iberica 544454441121+11212 Cytisusoromediterraneus 121122212323322223 Echinospartumbarnadesii 233433321122+23221 Juniperuscommunis subsp. 1+122112++1212121· alpina Avenellaiberica 12++11111111+·+111 Differentialsof typicum Arrhenatherumelatius s.l. 1 1 + + 1 1 1 1 · · · · · · + 1 + 1

Agrostiscastellana 1+111111 ··· ··· ··11 Armeriacaespitosa 11++1111 ··· ··· ·+· · Cytisusstriatus subsp. eriocarpus 1+++·+1+ ··· ··· ··· · Leucanthemopsispallidasubsp. · +++++· + · · · · · · ·+· · alpina Nardusstricta +· · ·· ·· + ··· ··· ··· · Differentialsof pinetosumsalzmannii Pinusnigra subsp. salzmannii ······· ·3333223232 Corynephoruscanescens ······· ·1+111++1+1 Agrostisdelicatula ······· ·212+1++1++

Shrubby,forestandperennialcompanions Festucaelegans subsp. merinoi ··++··· ·1+··1····3 Festucapaniculata subsp. ······· ·+····+1+·· multispiculata Festucagredensis 1111111+12221111·2 Genista cinerascens ······· ·++·······2 Luzulalactea 111+1+11+·· ··+11·1 Ericaarborea ·+++··+ · ··1·1·+··1 Pteridiumaquilinum ··+++·+ · ········12 Poabulbosa ·++··+· + ·····++111

Santolinaoblongifolia ······· ·+········+ Rubusulmifolius ······· · ······+··+ Thymusmastichina +· · ··+· · ··· ··· ··· · Others Cerastiumramosissimum ·+1+·1· · ······++1·

Rumexacetosella subsp. +· ·+· ·· · ··· ···+··1 angiocarpus Carduuscarpetanus +· · ·· ·· · ··+·+· ·+·1 Jasionemontana subsp. echinata ······· ·+··+······ Anthoxanthumodoratum ······· · ······++1+ Arenariamontana ······· · ·····+++1· Avenulasulcata ······· · ·····+++·· P Otherspecies:shrubby,forestandperennialscompanions: Sorbus aucuparia +in4; Cytisus scoparius 1, Crataegus monogyna 1and Pinus pinaster s.l.1in46.Others: Jasione sessiliflora and Urtica dioica +in4; Jasione montana subsp. echinata +in25; Lactuca viminea s.l.and Origanum vulgare +in26; Sedum forsterianum +in33; Galium rotundifolium , Armeria arenaria and Silene nutans +in46. Localities:Ávila,Gredosrange.1-4:CuevasdelValle,surroundingsummitsof‘PuertodelPico’and‘LaRubia’;5- 6:Piedralaves,CabezaPozo;7-8:SotillodelaAdrada,CerrodePinosequillo;9:ElArenal,ArroyodelosTorneros; holotypus subass .;10:ElArenal,ArroyodelaCabrilla;11:ElArenal,CerrodeColadillas;12-13:ElArenal,Arroyo delAndrinal;14:ElArenal,RiscodelasMorillas;15:,PortachodelAlcaraván;16:Casavieja,Gargantade laChorrera;17:Piedralaves,PeñaHorcada;18:SotillodelaAdrada,VenerodelasApreturas. 164 López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172

Gredosrange(clusterA1b)areverywelldis - demandbrightsituations(ElenaRosselló& al. , criminatedfromsamplesofclusterA1a,corre - 1985;ElenaRosselló&Sánchez Palomares, sponding,asnotedearlier,topureScotspine 1991;Regato&ElenaRosselló,1995). forestscommunities(Figures2and3).The ClusterA2includedsamplesfromxero- bulkofrelevésisplacedintheleftoftheDCA mesophilous Pinus sylvestris var. iberica for - diagram(0.15-0.33S.D.unitsalongDCAaxis estcommunities,mainlyfromthenorthern 1),althoughtotherightofsamplesfromclus - Gredosrange(samples5-10)butalsofromthe terA1aconsideringthatrelevésfromcluster southernslope(samples13,16,17),develop - A1bareatloweraltitudes(1410-1650masl; ingatanaltitudinallevelrangingfrom1300to Figure3a,Table4).SamplesfromclusterA1b 1570maslongranitesoils(Figure1).The alwaysincludeintheirrelevés Pinus sylvestris dominanceof Pinus sylvestris var. iberica in var. iberica (Table4),representingmixedfor - theirrelevésplaceswithintheclusterA,dif - mationstodaydominatedby P. nigra although feringfromsub-clusterA1byanaltitudinal inthepasttheycoulddominatebothspecies gradient(Figure2).Thisgroupincludes9 (Regato & al. ,1991,1992;López-Sáez& al. , relevéssituatedattherightoftheDCAaxis1 2014). Pinus sylvestris var. iberica , (Figure3a).ThesewereclassifiedbyLópez- Echinospartum barnadesii , Juniperus commu - Sáez& al. (2013)as Festuco merinoi-Pinetum nis subsp. alpina and Cytisus oromediterra - ibericae fromrelevésonlyonthenorthside; neus arediagnosticspeciesofbothclusters althoughinthispaperseveralonthesouthern A1aandA1b;while Corynephorus canescens , slopeoftheGredosrangearepresented(Table Agrostis delicatula and P. nigra subsp. 5).Floristically,thesesforestsrepresent Pinus salzmannii arethespecificdiagnosticspecies sylvestris var. iberica relictmesoforestsfrom ofclusterA1b(Figure3b,Table3). thesupratemperatesubmediterraneanand TheobservedvariabilitywithinclusterA1is uppersupramediterraneanhumidbeltofthe duetoanaltitudinalgradient. Pinus nigra subsp. Gredosmountains,presentbothinthenorth - salzmannii whichappearstobemoredrought- ern(NavarredondadeGredos,Hoyosdel resistantthat P. sylvestris var. iberica (Aussenac, Espino,Hoyocasero)andsouthern(La 1980),isafrequentco-dominantcanopytree Adrada,SanEstebandelValle,Guisando) onlyinlowersites(Figure1).Giventhatforest slopesinscatterednuclei.Theyarecharac - typeswithinclustersA1aandA1bsharecom - terizedbydiagnosticspeciessuchas Senecio monspecieswithsimilarecologyand,especial - adonidifolius, Hieracium sabaudum and ly,migrationhistoriesduringthelateglacialand Spergula morisonii (Figure3b,Table3), post-glacialperiods(G énova& al. ,2009;López- amongothers,aswellasdensebroomcom - Sáez& al. ,2013,2014;Abel-Schaad & al. , munitiesmainlycomposedby Cytisus scoparius 2014),allpreviouslymentionedfactsallowus and Genista florida andbywesternendemic toproposeanewpinewoodsubassociation: suchas Festuca elegans subsp. merinoi , Echinosparto barnadesii-Pinetum ibericae Centaurea amblensis , Ornithogalum concin - Sánchez-Mata,Gavilán&López-Sáezin num and Genista cinerascens (Martínez- López-Sáez&al.2013 pinetosum salzmannii García&Montero,2000;DelaFuente& subass. nova hoc loco (Table4, holotypus , Ortúñez,2001).Therelictpopulationfrom relevé9).Itrepresentsorotemperatesub - sample17islocatedat1170masl mediterranean(orosubmediterranean)humid (supramediterraneanthermotypeareas),shel - relictSpanishblackpinemesoforestsgrowing teredwithinamountainstreamandprotected throughoutthecentralandeasternGredos byalargegraniticbatholithwhichhasallowed Mountains(Figure1,Table4).Astypicalsubas - itssurvivalatlowaltitudeonthesouthern sociation( typicum ,clusterA1a),thisnew slopeoftheGredosrange. describedsubassociationisalsorepresentedon ClusterB2representspureSpanishblack thesouthernslopeoftheGredosrangebyiso - pineforestsinthelowersupramediterranean latedcopsesthathavealsosufferedrecurrent humidbioclimaticbeltoftheGredosrange actionoffires,overgrazingandindiscriminate (~900-1450masl),southeastorientedonthe felling(Regato& al. ,1992;Rubiales& al. , southernslopeandnorthwestorientedonthe 2007,2012;López-Sáez& al. ,2014).Theten northernslopeofthesemountains.Thisgroup inventoriespresentedinthispaperarealways includes21relevéssituatedattherightofthe fromsoutheastorientedpopulations(Table4), DCAaxis1(Figure3a).Diagnosticspeciesof becausetheSpanishblackpineisaspeciesthat thecommunityare Lactuca viminea s.l., López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172 165

Table5. Festuco merinoi-Pinetum ibericae Sánchez-Mata,Gavilán&López-SáezinLópez-Sáez&al. 2013( Avenello ibericae-Pinion ibericae , Pino sylvestris-Juniperetalia sabinae , Pino-Juniperetea ) Altitude(1=10m) 150 153 157 146 132 135 130 142 117 Numberofspecies 24 28 33 31 33 32 33 30 36 Relevénumber 1 2 3 4 5 6 7 8 9 Samplenumber(Figure3) 5 6 7 8 9 10 13 16 17 Aspect SE SE SE SE SE SE S SW SE Characteristics Pinussylvestris var. iberica 544455525 Genistacinerascens 122223232 Festucaelegans subsp. merinoi 222232333 Senecioadonidifolius ···+·++·+ Pilosellacastellana ++11 · +111 Hieraciumsabaudum ··+··+·++ Spergulamorisonii +++++· · · · Lotuscorniculatus s.l. · + 1 1 1 + + 1 1 Santolinaoblongifolia +++1 · +1 · 1 Anthericumliliago ····+++·· Achilleamillefolium +++11++11 Silenenutans +++1+· 111 Hypochoerisradicata +1+11++11 Genistaflorida +·++··+++ Conopodiumpyrenaeum ··+1+++·+ Violariviniana ·++··++11 Ornithogalumconcinnum ····+···+ Cytisusscoparius ++111+11+ Pteridiumaquilinum +1+11+111 Jasionemontana subsp. echinata ·1++1+111 Anthoxanthumodoratum ·+1+1+111 Arenariamontana ·+++++··· Euphorbianevadensis ····+·+·· Seneciojacobaea ····1+··· Ilexaquifolium ···+····+ Shrubby,forestandperennialcompanions Cytisusoromediterraneus 111++·++· Ericaarborea 111111222 Festucagredensis ++++1++11 Luzulalactea +1+· · +112 Poabulbosa ·1····111 Avenellaiberica 1111+1111 Agrostiscastellana ++111+1+1 Festucapaniculata subsp. multispiculata +·+· · ·111 Rosacanina ···+···+· Others Arrhenatherumelatius s.l. · + + + · · · + + Cerastiumramosissimum ·111·11+1 Rumexacetosella subsp. angiocarpus ++++· +111 Carduus carpetanus +·+· · ·+++ Leucanthemopsispallida subsp. alpina ++11 · +111 Aquilegiavulgaris ····+1··· Lychnisflos-cuculi ····+·+·· Irisxiphium ····++··· Arenariaquerioides ··+·+···· Otherspecies:shrubby,forestandperennialscompanions: Rubus corylifolius s.l.1in9; Lavandula pedunculata 1, Frangula alnus , Crataegus monogyna and Sorbus aria +in17.Others: Tanacetum corymbosum +in8; Convallaria majalis , Caltha palustris , Polygonatum odoratum and Stemmacantha exaltata +in9; Potentilla alpina subsp. apiifo - lia 1, Lilium martagon and Monotropa hypopytis +in10. Localities:Ávila,Gredosrange.1-3:NavarredondadeGredos;4:HoyosdelEspino;5-6:Hoyocasero;7:LaAdrada, PinaradelaVirgen;8:SanEstebandelValle,LaAbantera;9:Guisando,ChorrerasdeLaLancha. 166 López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172

Helianthemum apenninum s.l., Avenula sulcata , Althoughitisconsideredthatthedown - Lavandula stoechas subsp. pedunculata , weightingoffidelityvaluesinrelativelysmall Dianthus lusitanus , Sedum forsterianum , relevégroupsisdesirable(Bruelheide,2000), Armeria arenaria , Koeleria crassipes and thisfactcouldleadtoaweakerdifferentiation ofrelativelyrarevegetationunits,especially Festuca paniculata subsp. multispiculata inmicro-ormeso-scalephytosociological (Figure3b,Table3).Thiscommunityconsti - studies,asthepresentedhere,andthustothe tutessinglecohortstandstructuresothatit failureoftheirrecognition.Anadditional wasonlydominatedby Pinus nigra subsp. improvementistheuseoffidelityvaluesof salzmannii inthreelayersanditisgenerally speciesfollowingdifferenthierarchicallevels, formedbymaturestandsincentralandeastern asweproposedbelow.Forinstance, Pinus sylvestris var. iberica canbeconsidereda massifsoftheGredosrange(Regato & al. , transgressivediagnosticspecieshavingboth 1991,1992).Thesecommunitiesaremainly diagnosticfidelityvaluesinmorethanone locatedinsoutheastexposuresduetothether - group(clustersA1a,A1b,A2)andaconstancy micrequirementsofSpanishblackpine(Figure >20%inallgroupsforwhichthespeciesis 1),representingsupratemperatesubmediter - diagnostic(Table3)(Tichý&Chytrý,2006). ranean(suprasubmediterranean)humidrelict Thesamecircumstancescansignupforother forests.Infact,theGredosrangeshowshigh species: Echinospartum barnadesii , Juniperus communis subsp. alpina and Cytisus phytoclimaticsuitabilityforthisspeciesin oromediterraneus forclustersA1aandA1b; areaswithhigherrainfallinwinterandspring Agrostis delicatula forclustersA1bandB2; andcoolersummertemperatures(Romera Hypochoeris radicata , Jasione montana García& al. ,2001;GarcíaLópez& al. ,2005). subsp. echinata , Genista cinerascens and

Table6. Genisto cinerascentis-Pinetum salzmannii Sánchez-Mata,Gavilán&López-Sáez ass. nova (Avenello ibericae-Pinion ibericae , Pino sylvestris-Juniperetalia sabinae , Pino-Juniperetea )

Altitude(1=10m) 125 105 110 120 125 127 135 113 143 135 134 144 135 140 92 149 125 115 152 123 123 Aspect SE SE SE SE SE SE SE N SE SE SE SE NW SE NW NW NW NW NW NW SE Samplenumber(Figure3) 20 21 22 24 28 31 32 34 37 38 40 42 44 45 47 52 53 54 55 56 57 Numberofspecies 36 33 42 34 30 31 13 34 39 40 29 28 19 22 24 16 24 28 18 29 20 Relevénumber 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 Characteristics Pinusnigra subsp. salzmannii 433332342323343323333 Genistacinerascens 122333321223312+11121 Festucaelegans subsp. merinoi 3323332313222321+1·2+ Festucapaniculata subsp. ++11+1 · +11 1+11 1++· · ·+ multispiculata Agrostisdelicatula 1211·1111+1···· ··111· Hypochoerisradicata 1111··+1+11···· ·++·1+ Jasionemontana subsp. echinata +11+·+·1111+111··+·1+ Lactucaviminea s.l. + + + · · + + + · · + · · · · · · 1 + + + Helianthemumapenninum s.l. 3 1 1 1 · · · · · · · · · · · · 1 + · + · Avenulasulcata 1+1111· +11++· · · ·11· +· Lavandulapedunculata 211+·++·++· ···+12+·1+ Dianthuslusitanus 131+·11·+·· ···· ····+· Sedumforsterianum 111++·· ··+· ···· ···+·· Armeriaarenaria 1+1+· · · · · · · · · · · · · · · · · Koeleriacrassipes ·++·1··+··+ ···· ·1111+ Shrubby,forestandperennialcompanions Cytisusoromediterraneus ···········2···11·1·· Festucagredensis 211112+111+1111111111 Echinospartumbarnadesii ···········+······+·· Luzulalactea 1111+1·11111111··111+ Ericaarborea 21+111+112+·1++··+·1+ López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172 167

Pteridiumaquilinum ·+1222·12323212·13·22 Poabulbosa ··++11·11111111·11111 Agrostiscastellana 111111·11111111·++11+ Cytisusscoparius ··++11·2211·112··+·11 Genistaflorida ·1+++1·1+1···+······· Santolinaoblongifolia 11···1· ··+· ··+· ···· ·· Avenellaiberica 122211·1111111111111+ Pinuspinaster s.l. · · 1 · 1 1 · · 1 1 · · + · + + 1 1 · 1 · Rubusulmifolius ··++·+·++·····111+11· Crataegusmonogyna ·····+·++··+··+++·1·· Rosacanina ···············+1·11· Quercuspyrenaica ···+11········+·+···· Daphnegnidium ·····1·1·+····+······ Thymusmastichina ··++·····+···+······· Others Pilosellapseudopilosella 1+1+1· · 1+·+ · · · · · · · · +· Clinopodiumvulgare 1111·1+·+·· ····++·· ·· Arrhenatherumelatius s.l. · · · · + 1 · 1 1 1 1 + 1 + 1 · · + 1 1 + Cerastiumramosissimum ·······1111++1······· Rumexacetosella subsp. 1 +1+11·+11· ··+1++1·1· angiocarpus Carduuscarpetanus +++ ·11+111 · +· · 1++1· 1 · Leucanthemopsispallida ········+11·········· subsp. alpina Armeriacaespitosa ········++1+········· Pilosellacastellana +11+1··1·+1111· ···· ·· Achilleamillefolium 1·+1+··+1+1+1·· ···· ·· Silenenutans +·++1··++111·1· ···· ·· Lotuscorniculatus s.l. 1 1 1 · · · · · + + · · · · · · · · · · · Anthoxanthumodoratum ········+111111··+··+ Arenariamontana +++ ·+· · · ·+1+· · · · ·+· ++ Conopodiumpyrenaeum ····+······+········· Violariviniana +1+++·· ··+·+··· ···· ·· Ornithogalumconcinnum +·+···· ·++1+··· ···· ·· Galiumrotundifolium +1++···+++· ···1··+11· Merenderamontana 11··················· Corynephoruscanescens 1·+·112·1·· ···· ···· ·· Brachypodiumsylvaticum +·++1··+··· ···· ···· ·· Aspleniumtrichomanes +++ · · · · · · · · · · · · · · · · · · Origanumvulgare ·····1··+············ Urticadioica · · · · · · · · +··+···++· ···

Otherspecies:shrubby,forestandperennialscompanions: Cistus laurifolius 1and Cytisus multiflorus +in34.Others: Carex elata subsp. reuteriana and Spergula morisonii +in38; Ruscus aculeatus +in54. Localities:Ávila,Gredosrange.1:ElHornillo,LomadelaMajadilla, holotypus ass. ;2:ElHornillo,Silvona;3:Guisando,La Bardera;4:ElArenal,ArroyodelosTorneros;5:ElArenal,Majomingo;6-7:ElArenal,ArroyodelaOrtiga;8:Navalacruz;9: Casavieja,GargantadelaChorrera;10:Casavieja:FuentedelasClavellinas;11:Piedralaves,GargantadelaGraja.12: Piedralaves,LosHornillos;13:LaAdrada,GargantadelCharcodeLaHoya;14:LaAdrada,CuevadeMajaltobar;15:ElTiemblo, LasJuntas;16-18:ElTiemblo,CabezadelaParra;19-20:ElTiemblo,FuentedelasVíboras;21:ElTiemblo,LaCasquera.

Festuca elegans subsp. merinoi forclustersA2 while Genista cinerascens and Festuca ele - andB2;and,finally, Pinus nigra subsp. salz - gans subsp. merinoi wereconsideredasdiag - mannii forclustersA1bandB2(Table3). nosticspeciesoftheclusterA2( Festuco meri - Accordingtotherulesproposedinthe noi-Pinetum ibericae ).Althoughincaseslike ‘MaterialandMethods’section, Pinus nigra thatof Festuca elegans subsp. merinoi thedif - subsp. salzmannii wasconsideredasdifferen - ferentialroleofthetaxonisdifficulttodecide tialoftheclusterA1b( Echinosparto barnade - whichvegetationunitisbeingdifferentiated sii-Pinetum ibericae pinetosum salzmannii ), (Φ values:0.251clusterA2,0.264clusterB2), 168 López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172 incaseslikethatof Pinus nigra subsp. salzmannii with Pinus pinaster s.l.and Quercus pyrenaica (Φ:0.35)and Genista cinerascens (Φ:0.368) (López-Sáez& al. ,2016a).Theseforests isratherobvioustheirdiagnosticcharacter probablyhadamuchwiderdistributioninthe ofclusterB2.Thesefactsallowustopropose past,asshownbyhistorical,dendrochronolog - anewpinewoodassociationforrelevésof icalandpaleobotanicaldata(Génova& al. , clusterB2: Genisto cinerascentis-Pinetum 2008;Regato& al. ,1992,1995;López-Sáez salzmannii ass. nova hoc loco (Table6, holo - & al. ,2010,2014). typus ,relevé1). Theclassificationofthethreesub-clusters Pinus nigra subsp. salzmannii dominated ofclusterB1(Figure2),withstatistically forestsfromcalcareousanddolomiticsub - determineddiagnosticspeciesforeachsub- stratesoftheIberianPeninsulahavebeenusu - cluster,ispresentedinTable3.ClusterB1ais allyclassifiedwithintheordersof Quercetalia welldefinedby Quercus pyrenaica , Galium pubescentis and Prunetalia spinosae (Querco- rotundifolium and Rosa canina .Thiscluster Fagetea class), Quercetalia ilicis (Quercetea featuresPyreneanoakcommunities,which ilicis class)and Ononido-Rosmarinetalia locallyincluderelictSpanishblackpines. (Ononido-Rosmarinetea class)(Regato& al. , GalándeMera& al. (1999)considerthisfact 1995;Rivas-Martínez& al. ,2001,2002, asathermalrelictformofthe Festuco merinoi 2011;Regato & DelRío,2009).Additionally, -Quercetum pyrenaicae communitybetween itwasmentionedthat Pinus nigra subsp. 1100and1350masl. salzmannii forestsfromtheoromediterraneanand ClusterB1bsharessomediagnosticspecies supramediterraneanbioclimaticbeltsofsouthern withMediterraneandistribution,suchas Pinus Spain(Baza,Cazorla-SeguraandMágina pinaster s.l.and Daphne gnidium (Table3), ranges)andsomefromthesouthernIberian including P. pinaster s.l.dominatedforestsin Systemshouldbegroupedunderthe Junipero theGredosrangethatlocallyhouseSpanish sabinae-Pinetalia sylvestris order( Pino- blackpinesinthelowersupramediterranean Juniperetea class)(Valle& al. ,1988;Regato bioclimaticbelt(~900-1200masl).Probably, & al. ,1995;Rivas-Martínez& al. ,2001, aspalaeobotanicaldatahaveshown,the 2002).Giventhefloristicaffinitiesbetween Spanishblackpinehadawiderdistributionin clustersA1-A2andB2,whichsharesome thepastoccupyingloweraltitudinallevelsat commondiagnosticspeciessuchas Agrostis differenttimesoftheHolocene(Rubiales& castellana , A. delicatula , Hypochoeris radicata , al. ,2007,2012;López-Sáez& al. ,2014). Jasione montana subsp. echinata , Genista However,thegradualincreaseintemperature cinerascens and Festuca elegans subsp. merino i andcompetitionwithotherspeciesbetter (Table3),andacceptingtheconceptofmutu - adaptedtofireregimeanddisturbances,such alecological,biogeographicalandstructural as Pinus pinaster s.l.,reducedthepresenceof characteristics,webelievethatthe Genisto P. nigra subsp. salzmannii inlowerareasof cinerascentis-Pinetum salzmannii association theGredosrange(López-Sáez& al. ,2010). shouldbeputinto Junipero sabinae-Pinetalia ClusterB1crepresentsrelictpopulationsof sylvestris orderand Pino-Juniperetea class, Spanishblackpinethatfoundrefugewithin whichcomprisemainlyagroupofwestern alder( Alnus glutinosa )riparianforests~530- Mediterraneanorophilouswoodycommuni - 800masl.Itisgenerallyacceptedthatmoun - tiesdominatedbygymnosperms(Rivas- tainpines( Pinus sylvestris s.l.and P. nigra Martínez& al. ,2007,2011).Thestructureof s.l.)areTertiaryrelictswithacontinuedpres - thisvegetationcommunityintheGredosrange enceintheIberianPeninsulathroughoutthe isgenerallyconsistedbytreelayerwithmore Quaternary(Alcalde& al. ,2006;Soto& al. , orlessopencanopies,thickshrublayer,and 2010).Although Pinus nigra subsp. salzmannii ratherpoorspeciesassemblage(Table6; mainlygrowstodayinthesupramediterranean Regato& al. ,1992).Italsohasrelictcharac - bioclimaticbeltoftheIberianPeninsula, ter(Rubiales& al. ,2007,2012;López-Sáez& underhumidorsub-humidconditions,italso al. ,2014).Spanishblackpinehasdisappeared growsatlowerelevationsindryclimatewith frommanyhabitatsoftheGredosrange,main - temperatetoextremelycoldwinterconditions lyasaresultofwildfires,excessivegrazing andlowannualprecipitation(Regato& al. , anduncontrolledploughing(Génova& al. , 1995).Infact,oneofthestudiedpopulations 1992,2009;Cantero,2004;Génova&Moya, (sample18)islocatedinthemesome diterranean 2012),orinterspecificcompetition,mostly bioclimaticbeltat530masl,beingthepopu - López-Sáez J.A.; Sánchéz-Mata D.; Gavilán R.G. Lazaroa 37 2016: 153-172 169 lationatloweraltitudesinthewholeentire Acknowledgements Gredosmountains.Probably,thesefragmented populationssurvived in situ duringthelast Thisworkwasfundedbytheproject glacialmaximumratherthanhavingbeen HAR2013-43701-P(SpanishEconomyand recolonizedinthepost-glacialperiod,as CompetitivenessMinistry)“Dinámicassocio- shownbygeneticstudies(Afzal-Rafii & ecológicas,resilienciayvulnerabilidadenun Dodd,2007;Desprat& al. ,2015). paisajedemontaña:elSistemaCentral(9000 cal.BC-1850cal.AD)(DESIRÈ)”. Weare gratefultoA.LópezAndreu(RIP),P.Sáez NavasandM.TapiaArizafortheircollabora - tionandassistanceduringthefieldwork.

Syntaxonomical scheme

PiNO -J UNiPERETEA Rivas-Martínez1965 Pino sylvestris-Juniperetalia sabinae Rivas-Martínez1965 Avenello ibericae-Pinion ibericae Rivas-Martínez&J.A.MolinainRivas-Martínez, Fernández-González&Loidi1999 Echinosparto barnadesii-Pinetum ibericae Sánchez-Mata,Gavilán&López-SáezinLópez- Sáez&al.2013 typicum Sánchez-Mata,Gavilán&López-SáezinLópez-Sáez&al.2013 pinetosum salzmannii Sánchez-Mata,Gavilán&López-Sáez subass. nova Festuco merinoi-Pinetum ibericae Sánchez-Mata,Gavilán&López-SáezinLópez-Sáez&al.2013 Genisto cinerascentis-Pinetum salzmannii Sánchez-Mata,Gavilán&López-Sáez ass. nova

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Floristic Appendix

Avenella iberica (RivasMart.,Izco&Costa)RivasMart. Cytisus striatus (Hill)Rothm. subsp. eriocarpus (Boiss.&Reut.)RivasMart. Festuca elegans Boiss.subsp. merinoi (Pau)Fuente&Ortúñez Festuca gredensis Fuente&Ortúñez Jasione montana L.subsp. echinata (Boiss.&Reut.)RivasMart. Festuca paniculata (L.)Schinz&Thell.subsp. multispiculata RivasPonce&Cebolla Leucanthemopsis pallida (Mill.)Heywoodsubsp. alpina (Boiss.&Reut.)RivasMart.,Fern.Gonz.& Sánchez-Mata Pinus sylvestris L.var. iberica Svoboda