Technical guidelines for genetic conservation and use Mediterranean Abies spp Abies spp Abies spp Abies spp P.G. Alizoti1 , B. Fady2, M.A. Prada3, G.G. Vendramin4 1 Aristotle University of Thessaloniki, Greece 2 INRA, URFM, Ecology of Mediterranean , Avignon, France 3 GV, CIEF- Bank, Valencia, Spain 4 IGV-CNR, Florence, These Technical Guidelines are intended to assist those who cherish the valuable Mediterranean 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.

Biology and ecology A. nordmanniana and A. numid- ica). are linear, flattened with acute (A. cephalonica, A. Mediterranean firs are inter-fer- borisii-regis, A. equi-trojani and tile species that belong to two A. pinsapo) or round (A. cilicica, sections: i) Section Abies A. nebrodensis, A. nordmanniana ( and A. numidica) apex. Mattf., A. borisii-regis The firs are obligate seeders, Mattf., A. cephalonica monoecious and wind-pollinated Loud., A. equi-trojani and their are wind-dis- Asch., A. nebrodensis persed. The cones ripen in one Lojac and A. nor- season, while abundant seed dmanniana Spach.), production occurs only every 3-5 and ii) Section Pi- years. ceaster (Abies cili- They form pure or mixed for- cica de Lannoy, A. ests that span a considerable marocana Trabut, altitudinal range (600-2000 m) A. numidica Car- and grow in humid bioclimates, rière and A. pinsapo except for A. cephalonica, A. Boissier). cilicica and A. nebrodensis that They are ev- may also grow in sub-humid re- ergreen gions. A. borisii-regis, A. bornm- growing up to 50 uelleriana, A. cephalonica and A. m in height, with cilicica can tolerate wider ranges straight trunk and of mean annual temperature (7.5 pyramidal crown to 16 ºC) than the rest of the spe- that later turns to cies. Bud bursting occurs from flat-topped. The 3-5 early April (A. cephalonica and A. vegetative buds, at the cilicica), mid April (A. marocana, apex of the shoots, are A. numidica and A. pinsapo) to conical to ovoid and res- early May (A. bornmuelleriana inous (except in A. cilicica, and A. nordmanniana). MediterraneanAbies firsAbies spp Mediterranean spp firsAbies sppAbies Mediterranean firsAbies spp Mediterraneanspp firs AbiesAbies spp Mediterranean firsAbies spp Mediterranean firsAbiesAbies spp Mediterranean firs Abiesspp spp Mediterranean Abies firsAbies spp Mediterranean firssppAbies spp Mediter

Distribution Importance and use Genetic knowledge

Mediterranean firs have discon- Fir is generally consid- Disjunct geographic distribu- nected and limited distributions ered of lesser technical value tions and different population that are relict ranges of mostly than pine wood, but is used for sizes explain the generally high endemic species. The distribu- carpentry purposes due to its level of genetic diversity found tion area of most fir species softness and workability. It is among and within Mediterranean is concentrated in the eastern also used for general construc- fir populations. No variation ex- Mediterranean and the tion, paper, glued and composite ists at chromosome level (2n=24) re- wood products, veneer, ply- but broad genetic variation has gion. wood, panels and poles, been recorded at morphological as well as fuel wood. The (e.g. needle traits), anatomical bark, buds and cones may (e.g. position of resin canals), contain a large amount biochemical (e.g. terpenes, of fine, highly res- isozymes) and mo- inous turpentine. A lecular (e.g. micros- fine oil of turpentine atellites) levels. can be distilled from Paleo-ecologi- the crude material, while cal records and ge- the residue forms a coarse netic studies based on resin named colophony or markers with low evolu- rosin. Fresh oleoresin is mainly tionary rates (mitochondrial used for pharmaceutical pur- DNA and isozymes) suggest poses. that Mediterranean firs might Because of their fragrance, have stemmed from a common colour, good form and excep- Tertiary ancestor and were later tionally long retention af- separated into different genepo- ter being cut, most of the firs ols: eastern (A. cephalonica, A. are used as ornamental bornmuelleriana, A. equi-trojani and are grown in plantations for and A. nordmanniana, except Christmas trees (e.g. A. borisii A. cilicica that differed from the regis, A. cephalonica and A. nor- other species), southern (A. nu- dmanniana). midica) and western (A. pinsapo Mediterra- and A. marocana). More rapidly nean firs are evolving markers (chloroplast often found microsatellites) indicate that A. in protected cilicica is related to the eastern areas either Mediterranean group and that A. because of their marocana and A. pinsapo con- level of endemism and stitute different species. Simi- limited distribution or be- larly, A. nebrodensis appears as cause of their vital role as key- an original genetic group. stone species in Mediterranean Within-population genetic mountain ecosystems. variability increases from west to MediterraneanAbies firsAbies spp Mediterranean spp firsAbies sppAbies Mediterranean firsAbies spp Mediterraneanspp firs AbiesAbies spp Mediterranean firsAbies spp Mediterranean firsAbiesAbies spp Mediterranean firs Abiesspp spp Mediterranean Abies firsAbies spp Mediterranean firssppAbies spp Mediter

east as a result of harsher past Threats to Polyporus fulvus and Trametes climate in the western than the radiciperda. Other damaging genetic diversity eastern Mediterranean. A. ne- fungi include Aecidium elatinum, brodensis constitutes an excep- Overharvesting, grazing, pro- Armillaria gallica, Dioryctria abi- tion with high genetic diversity, longed drought and wild fires are etara, Fusarium sp., Heteroba- although the single remaining major threats to Mediterranean sidion annosum, Phytophthora natural population is extremely firs. The of cactorum (for young seedlings) small (29 individuals). the firs ranges from criti- and Trichosphaeria parasitica. Because of their coancestry, cally endangered to least Climate change is predicted Mediterranean firs can hybridize concerned according to the to intensify summer droughts in naturally and artificially. A. bo- IUCN Red List of Threatened the Mediterranean. It represents risii-regis is considered a natural species. A. nebrodensis is a serious threat for fir forests and hybrid between A. cephalonica recognized as critically en- is also likely to increase the fre- and A. alba. Successful artificial dangered and A. numidica is quency and extent of wild fires, crossing also suggests weak as threatened-vulnerable. A. with devastating effects on sur- reproductive barriers among cephalonica, A. marocana and vival, reproduction and regen- Mediterranean firs, as opposed A. pinsapo are considered as eration of the firs. Furthermore, to the relatively strict barri- near threatened species. A. ci- climate change may decrease ers that exist between North licica, A. nordmanniana and A. winter hardening and cause ear- American and Mediterranean borisii regis are recognized as lier bud burst, thus increasing firs. This supports the argu- least concerned species, while the risk of frost damage. This ment that geographic isolation A. bornmuelleriana and A. equi can jeopardize the development has been the main driver of trojani are not mentioned in of seedlings and young trees es- speciation in the Mediterra- the IUCN Red List. pecially in open areas, and cause nean basin. Even though no seri- fir forests to disappear from their Species and provenance ous insect-related threats low altitude sites. Temperature tests have revealed strong vari- seem to exist, Mediterranean rise during summer com- ation in adaptive traits such firs are usually damaged by bined with water stress as survival, growth, bud-break, Cacoecia muriana, Chroris- can result in extensive drought resistance, morphologi- toneura murinana, Crypha- crown defoliation and cal and anatomical traits. For- lus piceae, Dioryctria aulloi, mortality. The fir est reproductive material origi- Ips vorontzowi, Mindarus forests growing at nating from the Peloponnesos abietinus, Phaenops cya- low elevation, su- in Greece (A. cephalonica) per- nea, Pissodes picea, perficial soils or on forms well under many types of Pityokteines curvidens, southern slopes Mediterranean climate. Pityokteines spinidens are those currently and Xyloterus lineatus facing the greatest (especially after dry peri- threats. ods). Seed insects of the genus Megastigmus are known to significantly reduce seed crops. The most dangerous fungi for the firs include Armil- laria mellea, Fomes annosus, MediterraneanAbies firsAbies spp Mediterranean spp firsAbies sppAbies Mediterranean firsAbies spp Mediterraneanspp firs AbiesAbies spp Mediterranean firsAbies spp Mediterranean firsAbiesAbies spp Mediterranean firs Abiesspp spp Mediterranean Abies firsAbies spp Mediterranean firssppAbies spp Mediter

Guidelines for genetic 29 remaining individuals of the is conserved in a nature reserve species) in Arezzo, in botanical in Morocco and seven ex situ conservation and use gardens (40 000 in the Bo- stands have also been estab- Due to the threats, endemism tanical Garden of Palermo), ar- lished for the species. and geographically scattered boreta and in private properties Climate change will have an distribution, the conservation of in the Madonie Mountains close impact on the current in situ Mediterranean firs and their ge- to the natural habitat. A. borisii- conservation efforts but it is dif- netic resources is a major chal- regis and A. cephalonica are ficult to predict its effect on seed lenge. protected in situ in various pro- production, natural regeneration The genetic resources of the firs tected areas in Greece. Genetic and recruitment of the firs as well are currently conserved in vari- material, representing almost the as on the risks from insects and ous protected areas that have whole natural distribution of pathogens. The dynamic rarely been established for this the fir species, is included in gene conservation units purpose. Due to their evolution- provenance trials established should be monitored in ary history and specific adap- in Greece and France. A. ci- order to ensure that the tation, the fir forests harbour licica is protected in national populations are not seri- unique genetic resources that parks, nature reserves and ously affected and that are important beyond the Medi- seed stands in ten areas in they retain their evolution- terranean. Thus, the establish- and in Lebanon while ary potential and regener- ment of conservation units for in Syria it is considered as ate naturally. Management the firs that meet pan-European an endangered species. A. of the units should aim minimum requirements for dy- equi-trojani is conserved mainly at assisting natural namic gene conservation is of in situ in the Kazdagi Goknari regeneration and when this crucial importance. nature reserve in Turkey. A. nor- is not possible, the area should At present, several of the spe- dmanniana is also covered by be artificially regenerated with cies and their genetic resources protected areas in Turkey and local genetic material. Manage- are protected either in situ (na- several provenances are growing ment of natural forests should tional parks, nature reserve- ex situ in test sites, plantations also safeguard genetic resources sand gene conservation units) and arboreta in Denmark and by allowing natural selection to or ex situ (conservation seed France. The A. pinsapo forests occur on regeneration in a vari- orchards and stands). The are included in three pro- ety of situations. Ex situ conser- critically endangered A. tected areas in vation efforts should focus on nebrodensis Spain. A. small populations that have an is conserved numidica endangered status, insufficient in situ in the is pro- seed production or unsuccessful Madonie Re- tected in pollination in their natural envi- gional Park in the Dje- ronment. This approach is useful Sicily, but the bel Barbor especially in case of rare spe- reinforcement nature re- cies or species with limited or of the species has serve located in scattered distribution as ex situ been problematic mainly the Petite Kabylia stands with a sufficient number due to soil degradation in its Mountain range of and of genotypes form new inter- natural habitat. A. nebrodensis the same provenance is report- breeding populations that will is also conserved ex situ in a edly also conserved in ex situ produce seeds with a potentially seed orchard (with grafts of the stands. At present A. marocana high genetic diversity. 10

10 Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies spp Mediterranean firsAbies 8spp Mediterranean firsAbies spp Mediterranean 10 6 7 Abies spp Abies spp 7 Abies spp 6 9

10 5 Distribution2 range of Mediterranean 8 6 4 firs 10 9 5 6 7 10 7 1 3 6 9

10 5 6 2 8 10 4 9 1 Abies8 marocana 5 6 7 7 1 6 3 7 7 2 6 9 6 3 Abies9 numidica

5 6 4 Abies nebrodensis 2 5 2 6 4 9 1 Abies4 marocana 5 5 Abies cephalonic9a 5 1 1 33 2 Abies pinsapo 6 Abies borisii-regis 3 7 Abies equi-trojani 4 Abies nebrodensis 8 Abies bornmuelleriana 1 1 Abies Abies marocana marocana 5 9 Abies cilicica 2 2 Abies Abies pinsap pinsapoo 6 Abies borisii-regis 10 3 3 Abies Abies numidic numidicaa 7 Abies equi-trojani 4 Abies nebrodensis 4 Abies nebrodensis 8 Abies bornmuelleriana 5 Abies cephalonica 5 Abies cephalonica 9 Abies cilicica Mediterranean6 Abies borisii- firsre giofferss an 6 Abies borisii-regis 10 Abies nordmanniana opportunity7 Abies to equi-trojanitackle the pre- 7 Abies equi-trojani dicted forest8 Abies decline bornmuellerian in southerna Europe8 9Abies asAbies a bo cilicicaresultrnmuellerian of climatea change.9 10Abies A. Abies nordmanniana cilicica nordmanniana has al- ready 10been Abies used nordmanniana for reforestation in . Other Mediterranean firs (particularly A. cephalonica, A. bornmuelleriana and A. cili- cica) are far less water demand- ing and could represent an al- ternative for silver fir (A. alba) in Europe. Fir provenance tests in the Mediterranean include mate- rial that has demonstrated good growth, adaptation to drought and late bud burst in spring. Such provenances of Mediterra- nean firs could be of interest for the European forestry. MediterraneanAbies firsAbies spp Mediterraneanspp firsAbies Abiesspp Mediterranean firsAbies sppspp Mediterranean firs AbiesAbies spp Mediterranean firs Abiesspp spp Mediterranean

These Technical Guidelines were Selected bibliography produced by members of the EUFORGEN Conifers Network. The objective of the Network Aussenac, G. 2002. Ecology and ecophysiology of circum-Mediterranean firs is to identify minimum genetic in the context of climate change. Ann. For. Sci. 59:823-832. conservation requirements in the Liepelt, S., Mayland-Quellhorst, E., Lahme M and Ziegenhagen, B. 2010. Con- long term in Europe, in order to trasting geographical patterns of ancient and modern genetic lineages in Mediterranean Abies species. Syst. Evol. 284: 141-151 reduce the overall conservation cost and to improve the quality of Liu, T.S. 1971. A monograph of the genus Abies. Dept. of Forestry, National Taiwan University, Taipei, Taiwan, China. standards in each country. Parducci, L., A.E. Szmidt. 1999. PCR-RFLP analysis of cpDNA in the genus Abies. Theor. App. Genet. 98:802-808. Parducci, L., A.E. Szmidt, A. Madaghiele, M. Andizei and G.G. Vendramin. 2001. Genetic variation at chloroplast microsatellites (cpSSRs) in Abies nebrodensis (Lojac.) Mattei and three neighboring Abies species. Theor. Citation: Alizoti, P.G., B. Fady, App. Genet. 102:733-740. M.A. Prada and G.G. Vendramin. Scaltsoyiannes, A., M. Tsaktsira and A. Drouzas. 1999. Allozyme differentiation 2011. EUFORGEN Technical in the Mediterranean firs (Abies, ). A first comparative study with phylogenetic implications. Pl. Syst. Evol. 216:289-307. Guidelines for genetic conserva- tion and use of Mediterranean Terrab, A., S. Talavera, M. Arista, O. Paun, T.F. Stuessy and K. Tremetsberger. 2007. Genetic diversity of chloroplast microsatellites (cpSSRs) and geo- firs (Abies spp.) Bioversity Inter- graphic structure in endangered West Mediterranean firs (Abies spp., national, Rome, Italy. 6 p.. Pinaceae). Taxon 56(2):409-416. Ziegenhagen, B., B. Fady, V. Kuhlenkamp and S. Liepelt. 2005. Differentiat- Drawings: Abies pinsapo, Clau- ing groups of Abies species with a simple molecular marker. Silv. Gen. 54(3):123-126. dio Giordano © Bioversity, 2009.

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