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Mediterranean firs are inter-fer- tile species that belong to two sections: i) Section Abies (Abies bornmuelleriana Mattf., A. borisii-regis Mattf., A. cephalonica Loud., A. equi-trojani Asch., A. nebrodensis Lojac and A. nor- dmanniana Spach.), and ii) Section Pi- ceaster (Abies cili- cica de Lannoy, A. marocana Trabut, A. numidica Car- rière and A. pinsapo Boissier). They are ev- ergreen conifers growing up to 50 m in height, with straight trunk and pyramidal crown that later turns to flat-topped. The 3-5 vegetative buds, at the apex of the shoots, are conical to ovoid and res- inous (except in A. cilicica, A. nordmanniana and A. numid- ica). Leaves are linear, flattened with acute (A. cephalonica, A. borisii-regis, A. equi-trojani and A. pinsapo) or round (A. cilicica, A. nebrodensis, A. nordmanniana and A. numidica) apex. The firs are obligate seeders, monoecious and wind-pollinated and their seeds are wind-dis- persed. The cones ripen in one season, while abundant seed production occurs only every 3-5 years. They form pure or mixed for- ests that span a considerable altitudinal range (600-2000 m) and grow in humid bioclimates, except for A. cephalonica, A. cilicica and A. nebrodensis that may also grow in sub-humid re- gions. A. borisii-regis, A. bornm- uelleriana, A. cephalonica and A. cilicica can tolerate wider ranges of mean annual temperature (7.5 to 16 ºC) than the rest of the spe- cies. Bud bursting occurs from early April (A. cephalonica and A. cilicica), mid April (A. marocana, A. numidica and A. pinsapo) to early May (A. bornmuelleriana and A. nordmanniana). These Technical Guidelines are intended to assist those who cherish the valuable Mediterranean fir genepool and its inheritance, through conserving valuable seed sources or use in practical forestry. The focus is on conserving the genetic diversity of the species at the European scale. The recommendations provided in this module should be regarded as a commonly agreed basis to be complemented and further developed in local, national or regional conditions. The Guidelines are based on the available knowledge of the species and on widely accepted methods for the conservation of forest genetic resources. Technical guidelines for genetic conservation and use Abies spp Mediterranean firs P.G. Alizoti 1 , B. Fady 2 , M.A. Prada 3 , G.G. Vendramin 4 1 Aristotle University of Thessaloniki, Greece 2 INRA, URFM, Ecology of Mediterranean Forests, Avignon, France 3 GV, CIEF-Forest Seed Bank, Valencia, Spain 4 IGV-CNR, Florence, Italy Biology and ecology

Mediterranen Firs Abies Spp

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Mediterranen Firs Abies Spp

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Mediterraneanfirsareinter-fer-tilespeciesthatbelongtotwo sections:i)SectionAbies (Abiesbornmuelleriana Mattf.,A.borisii-regis Mattf.,A.cephalonica Loud.,A.equi-trojani Asch., A. nebrodensis LojacandA.nor-dmannianaSpach.), andii)SectionPi-ceaster(Abiescili-cicadeLannoy,A. marocanaTrabut, A.numidicaCar-rire and A. pinsapo Boissier).Theyareev-ergreenconifers growingupto50 minheight,with straighttrunkand pyramidalcrown thatlaterturnsto flat-topped.The3-5 vegetative buds, at the apexoftheshoots,are conicaltoovoidandres-inous (except in A. cilicica, A.nordmannianaandA.numid-ica).Leavesarelinear,flattened withacute(A.cephalonica,A. borisii-regis,A.equi-trojaniand A.pinsapo)orround(A.cilicica, A. nebrodensis, A. nordmanniana and A. numidica) apex. The firs are obligate seeders, monoecious and wind-pollinated andtheirseedsarewind-dis-persed.Theconesripeninone season,whileabundantseed production occurs only every 3-5 years. They form pure or mixed for-eststhatspanaconsiderable altitudinalrange(600-2000m) andgrowinhumidbioclimates, exceptforA.cephalonica,A. cilicicaandA.nebrodensisthat mayalsogrowinsub-humidre-gions. A. borisii-regis, A. bornm-uelleriana, A. cephalonica and A. cilicica can tolerate wider ranges of mean annual temperature (7.5 to 16 C) than the rest of the spe-cies.Budburstingoccursfrom early April (A. cephalonica and A. cilicica),midApril(A.marocana, A.numidicaandA.pinsapo)to earlyMay(A.bornmuelleriana and A. nordmanniana).These Technical Guidelines are intended to assist those who cherish the valuable Mediterranean firgenepoolanditsinheritance,throughconservingvaluableseedsourcesoruseinpractical forestry. The focus is on conserving the genetic diversity of the species at the European scale. Therecommendationsprovidedinthismoduleshouldberegardedasacommonlyagreed basistobecomplementedandfurtherdevelopedinlocal,nationalorregionalconditions. TheGuidelinesarebasedontheavailableknowledgeofthespeciesandonwidelyaccepted methods for the conservation of forest genetic resources.Technical guidelines for genetic conservation and useAbies spp Abies spp Abies sppAbies sppMediterranean firsP.G. Alizoti1 , B. Fady2, M.A. Prada3, G.G. Vendramin41 Aristotle University of Thessaloniki, Greece2 INRA, URFM, Ecology of Mediterranean Forests, Avignon, France3 GV, CIEF-Forest Seed Bank, Valencia, Spain4 IGV-CNR, Florence, Italy Biology and ecologyMediterraneanfirshavediscon-nectedandlimiteddistributions thatarerelictrangesofmostly endemicspecies.Thedistribu-tionareaofmostfirspecies isconcentratedintheeastern Mediterraneanandthe BlackSeare-gion.Firwoodisgenerallyconsid-eredoflessertechnicalvalue thanpinewood,butisusedfor carpentrypurposesduetoits softnessandworkability.Itis alsousedforgeneralconstruc-tion, paper, glued and composite woodproducts,veneer,ply-wood,panelsandpoles, aswellasfuelwood.The bark, buds and cones may contain a large amount offine,highlyres-inousturpentine.A fineoilofturpentine canbedistilledfrom the crude material, while the residue forms a coarse resinnamedcolophonyor rosin. Fresh oleoresin is mainly usedforpharmaceuticalpur-poses. Becauseoftheirfragrance, colour,goodformandexcep-tionallylongleafretentionaf-terbeingcut,mostofthefirs areusedasornamentaltrees andaregrowninplantationsfor Christmastrees(e.g.A.borisii regis, A. cephalonica and A. nor-dmanniana). Mediterra-neanfirsare oftenfound inprotected areaseither becauseoftheir level of endemism and limiteddistributionorbe-causeoftheirvitalroleaskey-stonespeciesinMediterranean mountain ecosystems. Disjunctgeographicdistribu-tionsanddifferentpopulation sizesexplainthegenerallyhigh levelofgeneticdiversityfound among and within Mediterranean firpopulations.Novariationex-ists at chromosome level (2n=24) butbroadgeneticvariationhas beenrecordedatmorphological (e.g.needletraits),anatomical (e.g.positionofresincanals), biochemical(e.g.terpenes, isozymes)andmo-lecular(e.g.micros-atellites) levels.Paleo-ecologi-calrecordsandge-netic studies based on markerswithlowevolu-tionary rates (mitochondrial DNAandisozymes)suggest thatMediterraneanfirsmight havestemmedfromacommon Tertiaryancestorandwerelater separated into different genepo-ols:eastern(A.cephalonica,A. bornmuelleriana,A.equi-trojani andA.nordmanniana,except A.cilicicathatdifferedfromthe otherspecies),southern(A.nu-midica) and western (A. pinsapo andA.marocana).Morerapidly evolvingmarkers(chloroplast microsatellites)indicatethatA. cilicicaisrelatedtotheeastern Mediterranean group and that A. marocanaandA.pinsapocon-stitutedifferentspecies.Simi-larly,A.nebrodensisappearsas an original genetic group. Within-populationgenetic variability increases from west to Abies spp Abies spp Abies spp Abies spp Abies sppDistribution Importance and use Genetic knowledgeMediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Meditereastasaresultofharsherpast climateinthewesternthanthe easternMediterranean.A.ne-brodensisconstitutesanexcep-tionwithhighgeneticdiversity, althoughthesingleremaining naturalpopulationisextremely small (29 individuals). Because of their coancestry, Mediterraneanfirscanhybridize naturallyandartificially.A.bo-risii-regis is considered a natural hybridbetweenA.cephalonica and A. alba. Successful artificial crossingalsosuggestsweak reproductivebarriersamong Mediterranean firs, as opposed totherelativelystrictbarri-ersthatexistbetweenNorth AmericanandMediterranean firs.Thissupportstheargu-mentthatgeographicisolation hasbeenthemaindriverof speciationintheMediterra-nean basin. Speciesandprovenance testshaverevealedstrongvari-ationinadaptivetraitssuch assurvival,growth,bud-break, droughtresistance,morphologi-calandanatomicaltraits.For-estreproductivematerialorigi-natingfromthePeloponnesos inGreece(A.cephalonica)per-formswellundermanytypesof Mediterranean climate.Overharvesting,grazing,pro-longed drought and wild fires are majorthreatstoMediterranean firs.Theconservationstatusof thefirsrangesfromcriti-callyendangeredtoleast concerned according to the IUCN Red List of Threatened species.A.nebrodensisis recognizedascriticallyen-dangeredandA.numidicais asthreatened-vulnerable.A. cephalonica, A. marocana and A.pinsapoareconsideredas near threatened species. A. ci-licica,A.nordmannianaandA. borisiiregisarerecognizedas leastconcernedspecies,while A. bornmuelleriana and A. equi trojaniarenotmentionedin the IUCN Red List. Eventhoughnoseri-ousinsect-relatedthreats seemtoexist,Mediterranean firsareusuallydamagedby Cacoeciamuriana,Chroris-toneuramurinana,Crypha-luspiceae,Dioryctriaaulloi, Ipsvorontzowi,Mindarus abietinus,Phaenopscya-nea,Pissodespicea, Pityokteinescurvidens, Pityokteinesspinidens andXyloteruslineatus (especially after dry peri-ods). Seed insects of the genusMegastigmusare knowntosignificantly reduceseedcrops.The mostdangerousfungi forthefirsincludeArmil-lariamellea,Fomesannosus, PolyporusfulvusandTrametes radiciperda.Otherdamaging fungi include Aecidium elatinum, Armillariagallica,Dioryctriaabi-etara,Fusariumsp.,Heteroba-sidionannosum,Phytophthora cactorum(foryoungseedlings) and Trichosphaeria parasitica. Climatechangeispredicted tointensifysummerdroughtsin theMediterranean.Itrepresents a serious threat for fir forests and isalsolikelytoincreasethefre-quencyandextentofwildfires, withdevastatingeffectsonsur-vival,reproductionandregen-erationofthefirs.Furthermore, climatechangemaydecrease winter hardening and cause ear-lierbudburst,thusincreasing theriskoffrostdamage.This canjeopardizethedevelopment of seedlings and young trees es-pecially in open areas, and cause fir forests to disappear from their lowaltitudesites.Temperature riseduringsummercom-binedwithwaterstress canresultinextensive crowndefoliationand treemortality.Thefir forestsgrowingat lowelevation,su-perficialsoilsoron southernslopes arethosecurrently facingthegreatest threats. Abies spp Abies spp Abies spp Abies spp Abies sppThreats togenetic diversityMediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp MediterDuetothethreats,endemism andgeographicallyscattered distribution,theconservationof Mediterraneanfirsandtheirge-neticresourcesisamajorchal-lenge. The genetic resources of the firs arecurrentlyconservedinvari-ousprotectedareasthathave rarelybeenestablishedforthis purpose.Duetotheirevolution-aryhistoryandspecificadap-tation,thefirforestsharbour uniquegeneticresourcesthat areimportantbeyondtheMedi-terranean.Thus,theestablish-mentofconservationunitsfor thefirsthatmeetpan-European minimumrequirementsfordy-namicgeneconservationisof crucial importance.At present, several of the spe-ciesandtheirgeneticresources areprotectedeitherinsitu(na-tionalparks,naturereserve-sandgeneconservationunits) orexsitu(conservationseed orchardsandstands).The critically endangered A. nebrodensi s isconserved insituinthe MadonieRe-gionalParkin Sicily,butthe reinforcement ofthespecieshas beenproblematicmainly duetosoildegradationinits naturalhabitat.A.nebrodensis isalsoconservedexsituina seedorchard(withgraftsofthe 29remainingindividualsofthe species)inArezzo,inbotanical gardens (40 000 plants in the Bo-tanicalGardenofPalermo),ar-boretaandinprivateproperties intheMadonieMountainsclose tothenaturalhabitat.A.borisii-regisandA.cephalonicaare protectedinsituinvariouspro-tected areas in Greece.Genetic material, representing almost the wholenaturaldistributionof thefirspecies,isincludedin provenancetrialsestablished inGreeceandFrance.A.ci-licicaisprotectedinnational parks,naturereservesand seedstandsintenareasin Turkey and in Lebanon while inSyriaitisconsideredas anendangeredspecies.A. equi-trojaniisconserved insituintheKazdagiGoknari naturereserveinTurkey.A.nor-dmannianaisalsocoveredby protectedareasinTurkeyand several provenances are growing exsituintestsites,plantations andarboretainDenmarkand France.TheA.pinsapoforests areincludedinthreepro-tectedareasin Spain.A. numidica ispro-tectedin theDje-belBarbor naturere-serve located in thePetiteKabylia MountainrangeofAlgeriaand thesameprovenanceisreport-edlyalsoconservedinexsitu stands.At present A. marocana isconservedinanaturereserve inMoroccoandsevenexsitu standshavealsobeenestab-lished for the species. Climatechangewillhavean impactonthecurrentinsitu conservationeffortsbutitisdif-ficult to predict its effect on seed production,naturalregeneration and recruitment of the firs as well as on the risks from insects and pathogens.Thedynamic geneconservationunits shouldbemonitoredin ordertoensurethatthe populationsarenotseri-ouslyaffectedandthat they retain their evolution-arypotentialandregener-ate naturally.Management oftheunitsshouldaim mainlyatassistingnatural regenerationandwhenthis isnotpossible,theareashould beartificiallyregeneratedwith localgeneticmaterial.Manage-mentofnaturalforestsshould also safeguard genetic resources byallowingnaturalselectionto occuronregenerationinavari-ety of situations. Ex situ conser-vationeffortsshouldfocuson smallpopulationsthathavean endangeredstatus,insufficient seed production or unsuccessful pollinationintheirnaturalenvi-ronment. This approach is useful especiallyincaseofrarespe-ciesorspecieswithlimitedor scattereddistributionasexsitu standswithasufficientnumber ofgenotypesformnewinter-breedingpopulationsthatwill produce seeds with a potentially high genetic diversity.Abies spp Abies spp Abies spp Abies spp Abies sppGuidelines for geneticconservation and useMediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp MediterMediterraneanfirsoffersan opportunitytotacklethepre-dicted forest decline in southern Europeasaresultofclimate change. A. nordmanniana has al-ready been used for reforestation inEurope.OtherMediterranean firs(particularlyA.cephalonica, A.bornmuellerianaandA.cili-cica) are far less water demand-ingandcouldrepresentanal-ternativeforsilverfir(A.alba)in Europe.Firprovenancetestsin the Mediterranean include mate-rialthathasdemonstratedgood growth,adaptationtodrought andlatebudburstinspring. SuchprovenancesofMediterra-neanfirscouldbeofinterestfor the European forestry.1Abies marocana 2Abies pinsapo3Abies numidica4Abies nebrodensis5 Abies cephalonica6Abies borisii-regis7Abies equi-trojani8 Abies bornmuelleriana9 Abies cilicica10 Abies nordmanniana123451067899610567Abies spp Abies spp Abies sppDistribution range of Mediterranean firsMediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean1Abies marocana 2Abies pinsapo3Abies numidica4Abies nebrodensis5 Abies cephalonica6Abies borisii-regis7Abies equi-trojani8 Abies bornmuelleriana9 Abies cilicica10 Abies nordmanniana1234510678996105671Abies marocana 2Abies pinsapo3Abies numidica4Abies nebrodensis5 Abies cephalonica6Abies borisii-regis7Abies equi-trojani8 Abies bornmuelleriana9 Abies cilicica10 Abies nordmanniana1234510678996105671Abies marocana 2Abies pinsapo3Abies numidica4Abies nebrodensis5 Abies cephalonica6Abies borisii-regis7Abies equi-trojani8 Abies bornmuelleriana9 Abies cilicica10 Abies nordmanniana123451067899610567Abies spp Abies spp Abies sppThese Technical Guidelines were producedbymembersofthe EUFORGENConifersNetwork. TheobjectiveoftheNetwork istoidentifyminimumgenetic conservation requirements in the longterminEurope,inorderto reducetheoverallconservation cost and to improve the quality of standards in each country.Citation:Alizoti,P.G.,B.Fady, M.A. Prada and G.G. Vendramin. 2011.EUFORGENTechnical Guidelinesforgeneticconserva-tionanduseofMediterranean firs(Abiesspp.)BioversityInter-national, Rome, Italy. 6 p..Drawings:Abiespinsapo,Clau-dio Giordano Bioversity, 2009.ISBN 978-92-9043-827-4Aussenac, G. 2002. Ecology and ecophysiology of circum-Mediterranean firs in the context of climate change. Ann. For. Sci. 59:823-832. Liepelt, S., Mayland-Quellhorst, E., Lahme M and Ziegenhagen, B. 2010. Con-trasting geographical patterns of ancient and modern genetic lineages in Mediterranean Abies species. Plant Syst. Evol. 284: 141-151Liu,T.S.1971.AmonographofthegenusAbies.Dept.ofForestry,National Taiwan University, Taipei, Taiwan, China.Parducci,L.,A.E.Szmidt.1999.PCR-RFLPanalysisofcpDNAinthegenus Abies. Theor. App. Genet. 98:802-808.Parducci,L.,A.E.Szmidt,A.Madaghiele,M.AndizeiandG.G.Vendramin. 2001.Geneticvariationatchloroplastmicrosatellites(cpSSRs)inAbies nebrodensis(Lojac.)MatteiandthreeneighboringAbiesspecies.Theor. App. Genet. 102:733-740. Scaltsoyiannes, A., M. Tsaktsira and A. Drouzas. 1999. Allozyme differentiation in the Mediterranean firs (Abies, Pinaceae). A first comparative study with phylogenetic implications. Pl. Syst. Evol. 216:289-307.Terrab, A., S. Talavera, M. Arista, O. Paun, T.F. Stuessy and K. Tremetsberger. 2007.Geneticdiversityofchloroplastmicrosatellites(cpSSRs)andgeo-graphicstructureinendangeredWestMediterraneanfirs(Abiesspp., Pinaceae). Taxon 56(2):409-416.Ziegenhagen,B.,B.Fady,V.KuhlenkampandS.Liepelt.2005.Differentiat-inggroupsofAbiesspecieswithasimplemolecularmarker.Silv.Gen. 54(3):123-126.Selected bibliographyMore informationwww.euforgen.orgMediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp MediterraneanEUFORGEN Secretariatc/o Bioversity InternationalVia dei Tre Denari, 472/a00057 Maccarese (Fiumicino)Rome, ItalyTel. (+39)0661181Fax: (+39)[email protected]