Technical guidelines for genetic conservation and use European beech Fagus sylvatica Fagus sylvatica Fagus sylvatica Georg von Wühlisch Federal Research Institute for Rural Areas, Forestry and Fisheries, EUFORGEN Institute for Forest Genetics, Germany

These Technical Guidelines are intended to assist those who cherish the valuable European beech 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 is a good species for soil con- servation as it produces a large amount of leaf litter (ca. 900 g/m² European beech (Fagus sylvatica per year) and has extensive shal- L.) normally grows to 30–35 m tall low and intermediate roots. but in rare instances it may grow Beech is relatively resistant to over 40 m. Unlike to most diseases. It does not many other tree spe- suffer from massive predations cies, it maintains its by pests that lead to a total high growth rate dieback of stands. Late spring until late ma- frosts often damage young trees turity. Beech or flowers, which emerge simul- trees can live taneously with leaf flush. Intense for 250 years sunlight may damage the stem or more but surface. Aphids may attack the are normally bark. The fungus Nectria ditis- h a r v e s t e d sima causes bark necrosis. at 80–120 The tetrahedral shiny brown years of beechnuts are positioned in pairs age. Beech in four-sided cupules. A mast has separate year of beechnuts occurs only male and fe- every 5–8 years. The seed can male flowers be stored for about five years. on the same tree However, during storage the and it is wind pol- germination capacity of seeds linated. decreases considerably. Seed The thin, smooth, dormancy is strong but can be silver-grey bark is broken by keeping seed at 3° C highly characteristic of for at least six weeks. beech. The leaves are ellipti- Beech is highly shade tol- cal without any lobes or peaks erant. It can be regenerated and have a short stalk. Beech naturally in continuous cover EuropeanFagus beechFagus sylvaticaEuropean sylvatica beechFagus sylvaticaEuropean beech FagusFagus sylvaticaEuropean beechFagus sylvaticasylvaticaEuropean beechFagus sylvaticaEuropean Fagus beechFagus sylvaticaEuropean beechsylvaticaFagus sylvaticaEuropean beechFagus sylvatica FagEuropean beechFa

silvicultural systems. However, Distribution Importance and use where red or roe deer occur, they will browse beech seedlings and regeneration areas should Beech is widely distributed in Beech wood is homogeneous be fenced. Central and Western Europe. In with fine pores and conspicuous Beech favours damp sites the northern part of its range wood rays. The colour varies and soils that roots can eas- beech grows at low elevations from nearly white to reddish. The ily penetrate. Best growth while in the southern part it is wood has an average density of is attained in found at altitudes above 1000 m 700 kg m–3 with good stiffness moist soils asl. but little elasticity. It is resistant on calcareous Beech spread to to abrasion but is susceptible or volcanic central and north- to attack by fungi and needs to rock beds. ern Europe from be protected if used outdoors. Beech does southern France, With about 250 known uses for not grow on rocky eastern Alps–Slovenia– its wood, beech is the most sites, in very dry areas, Istria and possibly southern diversely used tree species in on sites with stagnant wa- Moravia–southern Bohemia. The Europe. ter or regularly flooded sites. populations that survived the last Beech wood is mainly used On favourable sites beech glacial period in the Mediterra- for furniture. It is also excel- is widely distributed because it nean regions (Italian and Iberian lent for flooring and staircases. out-competes other tree species peninsulas) did not spread into Beech wood is also used exten- due to its efficient use of light. central Europe. sively in the production of pulp Once beech has become the and various boards, veneer and dominant species, this creates plywood. It is used as fuelwood low light levels in the understory due to its relatively high energy where beech seedlings can sur- content. vive better than other species. Beech trees more than 100 years old frequently develop red heartwood, a discol- oration that limits the use of the wood. EuropeanFagus beechFagus sylvaticaEuropean sylvatica beechFagus sylvaticaEuropean beech FagusFagus sylvaticaEuropean beechFagus sylvaticasylvaticaEuropean beechFagus sylvaticaEuropean Fagus beechFagus sylvaticaEuropean beechsylvaticaFagus sylvaticaEuropean beechFagus sylvatica FagEuropean beechFa

Genetic knowledge is an important factor that should Genetic improvement of be considered when establishing beech has relied mainly on seed beech seed production stands stand selection. A simple system Genetic variation in beech has and gene conservation units. is usually applied, where charac- been studied in provenance trials There is continuous variation ters such as growth increment, and with genetic markers such in some characters, e.g. in the health and phenotypic appear- as isozymes and DNA markers. time of the flushing of the leaves. ance are evaluated on a large Studies using nuclear molecu- This character reflects adapta- number of seed trees (at least 80 lar markers show high varia- tion to late frost, which occurs trees more than 70 years old). In tion within populations and lit- more frequently in mild oce- each seed lot, seed should be tle differentiation among pop- anic conditions than under a harvested from at least 40 trees. ulations. However, maternally continental climate. Beech The seed stand should have a inherited DNA markers of the also shows variation in minimum size of 2.5 ha, prefer- chloroplasts (transmitted by the temperature sum re- ably larger. Individual selection seeds only) display a very quired for initiation of leaf and subsequent progeny testing high differentiation among flush, which is higher in have rarely been undertaken and populations. The analysis of oceanic beech popu- only a few seed orchards are the spatial distribution of al- lations than in conti- known to exist. leles in a naturally regener- nental populations. Mass propagation by means ated, isolated pure beech Thus, when grown at of vegetative propagation (clon- stand using molecular mark- a single site, prov- ing) is possible but is not prac- ers shows the tendency of a enances from a con- ticed due to the high costs in- strong family structure in the tinental site in the volved. Cuttings are generally distance up to 30 m. Spatial south-east of the difficult to root. Clones differ dis- genetic structures are influ- distribution range tinctly in their ability to root. Eu- enced by unpredictable fac- usually flush earlier ropean beech does not produce tors (e.g. wind direction at than those from an root-suckers like some other anthesis) and can therefore oceanically influenced beech species. In vitro culture vary from year to year. Based site in the north-west of techniques have not been devel- on these results, it is recom- the distribution range. oped to a commercially feasible mended that seed should be Provenances from the scale. collected over large areas south-east may thus suf- in order to prevent a pre- fer greater frost damage in ponderance of a few fami- spring when grown on sites in lies and a reduction of the the north-west of the distribu- adaptive potential of the tion range. next generation. Some stem form characters, Although seed dispersal of such as spiral grain, are under beech is limited. Pollen flow en- relatively strong genetic con- sures long distance gene flow. A trol. Provenances from higher study in three relatively isolated elevations may show better tree beech stands in northern Germa- form characters such as trunk ny indicated that gene flow from straightness, branching and outside beech stands is very ef- crown shape than those from ficient. External pollen gene flow lower elevations. EuropeanFagus beechFagus sylvaticaEuropean sylvatica beechFagus sylvaticaEuropean beech FagusFagus sylvaticaEuropean beechFagus sylvaticasylvaticaEuropean beechFagus sylvaticaEuropean Fagus beechFagus sylvaticaEuropean beechsylvaticaFagus sylvaticaEuropean beechFagus sylvatica FagEuropean beechFa

Threats to the present distribution range are Guidelines for genetic likely to become more favourable genetic diversity conservation and use for beech. These changes will Beech prefers sites favourable also affect genetic diversity of Genetic diversity of beech should for agriculture and subsequently beech. More knowledge about be conserved using a mix of large areas of beech forest have the genetic diversity, variability in situ and ex situ approaches. been cleared for agricultural and adaptability of beech is re- For reforestation, the minimum production. As a result, a large quired to conserve the endan- requirement should be that the proportion of beech gered populations more effec- origin of the reproductive mate- genetic diversity has tively. Such knowledge would rial is known and its adaptive probably been be valuable for any consid- characters should be appropri- destroyed. This erations to transfer re- ate for the ecological conditions land clearing productive material to at the new site. This is especially has also frag- regions which are likely important in places where beech mented the re- to become more favour- is to be re-introduced but little maining stands. able for beech. knowledge on site-adapted pop- In some regions ulations exists, e.g. where beech the reduction in is used to replace maladapted beech populations conifer stands that have been has been dramatic. For exam- planted on former beech sites. ple, in Saxony, Germany, in the Besides the present regula- centre of the distribution range, tions for documenting forest re- beech used to cover 50% of productive material in trade, a the forest area but presently it monitoring system for the use covers only 3%. Threats to the of reproductive materials should present genetic diversity can be be applied. Recommendations assumed to be small. The re- should be developed on prop- maining genetic diversity could er use of different materials in be threatened by the efforts to the face of climate change, increase the beech area if poor- together with transfer guide- ly-adapted reproductive material lines. The EU directive and is introduced. OECD scheme provide basic Some beech stands may be regulations on the transfer of at risk from climate change, par- reproductive materials. In years ticularly those in the lowlands with abundant seed production, where precipitation is expected beech seed should be harvested to decrease and summer tem- and stored in sufficient amounts peratures to increase. Thus, to capture the widest range of beech stands in the genetic diversity. southern and south- Beech can usually be con- eastern parts of the served in situ in normal stands. present distribution will In many parts of Europe seed be affected most. At the stands alone may not be enough same time conditions in for the conservation of genetic the north and north-east of resources of beech. Therefore, FagusEuropean beechFagus sylvaticaEuropean sylvatica beechFagus sylvaticaEuropean beech FagusFagus sylvaticaEuropean beechFagus sylvatica sylvaticaEuropean beechFagus sylvaticaEuropean

Distribution range of European beech

there is a need for gene re- maintain as much as possible of serve forests. These are natural the original genetic variability and stands managed to ensure suc- to allow continuing adaptation to cessful natural regeneration, e.g. local conditions. Ex situ conser- through thinning and harvesting vation stands should cover 2–5 older trees. The objective is to ha, and can be established by maintain continuous evolution of planting seedlings or by direct a tree population. Such gene sowing. reserve forests should cover at least 100 ha in order to contain sufficient genetic variability. For small, locally adapted popula- tions, however, it may be better to establish a large number of smaller reserves. The establishment of ex situ conservation plantations of beech may be necessary in order to conserve the genetic varia- tion of threatened populations that cannot be maintained at the original site. The objective is to FagusEuropean beechFagus sylvaticaEuropean sylvatica beechFagus sylvaticaEuropean beech FagusFagus sylvaticaEuropean beechFagus sylvatica sylvaticaEuropean beechFagus sylvaticaEuropean EUFORGEN

The series of these Technical Selected bibliography Guidelines and the distribution maps were produced by mem- bers of the EUFORGEN Net- Demesure, B., B. Comps and J. Petit 1996. Chloroplast DNA phylogeography works. The objective is to identify of the common beech (Fagus sylvatica L.) in Europe. Evolution 50:2515– minimum genetic conservation 2520. requirements in the long term in Madsen, S.F. editor 1995. Genetics and Silviculture of Beech. Proceedings of th Europe, in order to reduce the the 5 Beech Symposium of the IUFRO Project Group. Danish Forest and Landscape Research Institute, Hørsholm, Denmark. 272 pp. overall conservation cost and to improve the quality of standards Matič, S. editor in chief. 2003. Obična bukva (Fagus sylvatica L.) u Hrvatskoj [Common beech (Fagus sylvatica L.) in Croatia]. Akademija Šumarskih in each country. Znanosti (Academy of Forest Science), Zagreb, Croatia. 855 pp. Magri, D., G.G. Vendramin, B. Comps, I. Dupanloup, T. Geburek, D. Gömöry, M. Lata‘owa, T. Litt, L.Paule, J.M. Roure, I. Tantau, W.O. van der Knaap, R.M. Petit and J.-L. de Beaulieu 2006: A new scenario for the Quaternary history of European beech populations: palaeobotanical evidence and genetic consequences. New Phytologist 171:199–221 Citation: von Wuehlisch G. 2008. Muhs, H.-J. and G. von Wuehlisch editors. 1993. The Scientific Basis for the EUFORGEN Technical Guidelines Evaluation of the Genetic Resources of Beech. Proceedings of a scientific for genetic conservation and use workshop under the Community research programme on agriculture and agro-industry, including fisheries (“AIR”), held in Ahrensburg, Germany, for European beech (Fagus syl- 01–02 July, 1993. Working Document of the EC, DG VI, Brussels, Belgium. vatica). Bioversity International, 267 pp. Rome, Italy. 6 pages. Peters, R. 1997. Beech Forests. Kluwer, Dordrecht, The Netherlands. 169 pp. Teissier du Cros E. editor. 1981. Le Hêtre. INRA, Paris, France. 613 pp. Drawings: Fagus sylvatica, Gio- vanna Bernetti. © 2003 Biover- Vornam, B., N. Recarli and O. Gailing 2004. Spatial distribution of genetic variation in a natural beech stand (Fagus sylvatica L.) based on microsatellite sity International. markers. Conservation Genetics 5:561–570. Wang, K.S. 2004. Gene flow in European beech (Fagus sylvatica L.). Genetica ISBN 978-92-9043-787-1 122:105–113. The distribution map, showing the natural distribution area of Fagus sylvatica was compiled by members of the EUFORGEN Networks based on an earlier map published in: Pott R. (2000) Palaeoclimate and vegetation - long-term vegetation dynamics in central Europe with particular reference to beech. Phytocoenologia 30(3-4): 285-333

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