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EUROPEAN FOREST GENETIC RESOURCES PROGRAMME (EUFORGEN) 

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23-25 October 1997 

Bordeaux, France 

J .. Turok, A .. Kremer and S .. de Vries, compilers

ii EI.JE0SGEN: S0€IAI.ili BS0ADI.iliEAXlES . 

The International Plant Genetic Resources Institute (IPGRl) is an autonomous international scientific 

organization, supported by the Consultative Group on International Agricultural Research (CGIAR). 

IPGRl's mandate is to advance the conservation and use of plant genetic resources for the benefit of 

present and future generations. IPGRl's headquarters is based in Rome, Italy, with offices in another 14 

countries worldwide. It operates through three programmes: (1) the Plant Genetic Resources Programme, 

(2) the CGIAR Genetic Resources Support Programme, and (3) the International Network for the 

Improvement of Banana and Plantain (INIBAP). The international status of IPGRl is conferred under an 

Establishment Agreement which, by January 1998, had been signed and ratified by the Governments of 

Algeria, Australia, Belgium, Benin, Bolivia, Brazil, Burkina Faso, Cameroon, Chile, China, Congo, Costa 

Rica, Cote d'Ivoire, Cyprus, Czech Republic, Denmark, Ecuador, Egypt, Greece, Guinea, Hungary, India, 

Indonesia, Iran, Israel, Italy, Jordan, Kenya, Malaysia, Mauritania, Morocco, Pakistan, Panama, Peru, 

Poland, Portugal, Romania, Russia, Senegal, Slovakia, Sudan, Switzerland, Syria, Tunisia, Turkey, 

Uganda and Ukraine. 

Financial support for the Research Agenda of IPGRl is provided by the Governments of Australia, 

Austria, Belgium, Brazil, Bulgaria, Canada, China, Croatia, Cyprus, Czech Republic, Denmark, Estonia, 

F.R. Yugoslavia (Serbia and Montenegro), Finland, France, Germany, Greece, Hungary, Iceland, India, 

Ireland, Israel, Italy, Japan, Republic of Korea, Latvia, Lithuania, Luxembourg, Malta, Mexico, Monaco, 

the Netherlands, Norway, Pakistan, the Philippines, Poland, Portugal, Romania, Slovakia, Slovenia, South 

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Bank, Common Fund for Commodities, Technical Centre for Agricultural and Rural Cooperation (CTA), 

European Union, Food and Agriculture Organization of the United Nations (FAO), International 

Development Research Centre (IDRC), International Fund for Agricultural Development (IFAD), 

International Association for the promotion of cooperation with scientists from the New Independent 

States of the former Soviet Union (INTAS), Interamerican Development Bank, United Nations 

Development Programme (UNDP), United Nations Environment Programme (UNEP) and the World 

Bank. 

The European Forest Genetic Resources Programme (EUFORGEN) is a collaborative programme 

among European countries aimed at ensuring the effective conservation and the sustainable utilization 

of forest genetic resources in Europe. It was established to implement Resolution 2 of the Strasbourg 

Ministerial Conference on the Protection of Forests in Europe. EUFORGEN is financed by 

participating countries and is coordinated by IPGRl, in collaboration with the Forestry Department of 

FAO. It facilitates the dissemination of information and various collaborative initiatives. The 

Programme operates through networks in which forest geneticists and other forestry specialists work 

together to analyze needs, exchange experiences and develop conservation objectives and methods for 

selected species. The networks also contribute to the development of appropriate conservation 

strategies for the ecosystems to which these species belong. Network members and other scientists 

and forest managers from participating countries carry out an agreed workplan with their own 

resources as inputs in kind to the Programme. EUFORGEN is overseen by a Steering Committee 

composed of National Coordinators nominated by the participating countries. 

The geographical designations employed and the presentation of material in this publication do not 

imply the expression of any opinion whatsoever on the part of IPGRl or the CGIAR concerning the legal 

status of any country, territory, city or area or its authorities, or concerning the delimitation of its frontiers 

or boundaries. Similarly, the views expressed are those of the authors and do not necessarily reflect the 

views of these participating organizations. 

Citation: Turok, J., A. Kremer and S. de Vries, compilers. 1998. First EUFORGEN Meeting on Social 

Broadleaves. Bordeaux, France, 23-25 October 1997. International Plant Genetic Resources Institute, Rome. 

ISBN 92-9043-377-9 

IPGRl, Via delle Sette Chiese 142, 00145 Rome, Italy 

© International Plant Genetic Resources Institute, 1998

CONTENTS 

iii 

Contents 

Introduction 

Workplan 

1 

3 

Country Reports 

Beech and oak genetic resources in Romania 

loan Blada 

5 

Present status of the conservation and use of broadleaved forest 

genetic resources in Moldova 

Gheorghe Postolache 11 

Conservation of genetic resources of Social Broadleaves in Ukraine 

19or M. Patlaj, Svitlana A. Los, Roman M. Jatzyk and lhor M. Shvadchak 13 

Conservation of genetic resources of white oaks and beech in Hungary 

Attila Borovics, Zoltan Somogyi and Csaba Matyas 20 

Conservation of beech and oak genetic resources in Slovakia 

Ladislav Paule 29 

Social Broadleaves in the Czech Republic 

Vladimir Hynek 34 

Conservation of genetic resources of oaks and beech in Austria 

Thomas Geburek 41 

Conservation of Social Broadleaves genetic resources in Switzerland 

Patrick Bonfils 45 

Fagus sylvatica, Quercus petraea and Quercus robur genetic resources in Italy 

Paolo Menozzi 55 

Beech and oak genetic resources in Slovenia 

19or Smolej, Robert Brus, Marjanca Pavle, Saso Zitnik, Zoran Grecs, 

Nevenka Bogataj, Franc Ferlin and Hojka Kraigher 64 

Oak genetic resources in Malta 

Darrin T. Stevens 75 

Management and conservation of beech (Fagus sylvatica L.) and oak 

(Quercus petraea (Matt.) LiebL Q. robur L.) genetic resources in France 

Eric Teissier du Cros, lsabelle Bilger, Alexis Ducousso and Antoine Kremer 77 

Oak and beech genetic resources in Luxembourg 

M. Wagner, F. WoIter and J.F. Hausman 86 

Genetic conservation strategy for Social Broadleaves in Belgium 

Dominique Jacques and Bart de Cuyper 88 

Activities concerning Social Broadleaves genetic resources 

in the Netherlands 

Sven M.G. de Vries 97 

Beech and oak species in Germany: occurrence and gene 

conservation measures 

B. Richard Stephan 102 

Conservation strategy for beech and oaks in Denmark 

Jan S. Jensen 112 

Genetic resources and conservation of Quercus robur L. in Lithuania 

Virgilijus Baliuckas and Julius Danusevicius 117 

5

iv 

EUF0RGEN: S0GIALE BR0ADLEEAVES 

Oak and beech resources in Latvia 

Arnis Cailis and Edgars Smaukstelis 121 

Genetic resources of oaks and their conservation in Finland 

Anu Mattila 127 

Social Broadleaves in Sweden 

Lennart Ackzell 131 

Overview Presentations 133 

Structure of gene diversity, geneflow and gene conservation in 

Quercus petraea 

Antoine Kremer, Remy J. Petit and Alexis Ducousso 133 

Oak decline in European forests 

Tomasz Oszako 145 

Genetic diversity of beech populations in Europe 

Ladislav Paule and Dusan Comory 152 

A Network of International Beech Provenance Trials 

Ceorg von Wuehlisch, Mirko Liesebach, Hans-J. Muhs and Richard Stephan 164 

Programme 173 

List of Participants 174

INwR0Duewl0N 1 

Intr0dueti0n 

The first EUFORGEN meeting on Social Broadleaves was held from 23 to 25 October 1997 at 

the Station de Recherches Forestieres (INRA) in Bordeaux-Cestas, France. Participants 

representing 23 countries attended the meeting (see List of Participants). They presented 

country reports, identified common needs, agreed on a joint workplan and established a 

Network concerned with the conservation and sustainable use of genetic resources in Social 

Broadleaves. 

For the immediate future, the scope of the Network includes European white oaks 

(Quercus robur, Q. petraea, Q. pubescens and related species) and beech (Fagus sylvatica, 

F. orientalis). 

The establishment of this Network - a fifth one within the framework of EUFORGEN - 

had been previously requested by the participating countries through their National 

Coordinators. It was felt that the existing Networks (Noble Hardwoods, Picea abies, Populus 

nigra and Quercus suber) do not cover adequately the European temperate oaks and beech, 

considering their importance, and in view of the specific gene conservation strategies they 

require. The coverage of species by the Network and its title will be addressed at the next 

Steering Committee meeting of National Coordinators (to be held in Vienna, Austria, 26-29 

November 1998). 

The common needs concern in particular: 

• to improve information flow among countries 

• to harmonize research priorities and disseminate available research results 

• to address legislation-related issues 

• to develop joint, long-term, practically oriented strategies and standardize or develop 

methodologies 

• to raise awareness of decision-makers, the general public and forest owners about the 

necessity of conserving genetic resources of Social Broadleaves. 

The participants of the meeting developed a common workplan with shared 

responsibilities. It aims at strengthening collaboration among European countries by 

providing practical outputs such as technical guidelines for the sampling, design and 

management of gene conservation units, databases, information resources and public 

awareness tools (see Workplan). 

To obtain a comprehensive overview of the current developments, four participants were 

invited to deliver introductory presentations during the first part of the meeting. They 

reviewed the results of research on the genetic diversity of oaks and beech throughout 

Europe, summarized the concerns associated with oak decline and introduced the 

international network of beech provenance trials established recently. Overview presentations 

as well as country reports presented at the meeting are published in this volume. 

The country reports typically include, but are not strictly limited to, information on the 

occurrence and origin of the European temperate oaks and beech in each country, their 

economic importance, silvicultural approaches used, health state of the forest stands and 

threats to their genetic diversity, genetic conservation activities, relevant nature protection 

policies and activities, tree-improvement, use of reproductive material, institutions involved, 

research capacities, needs and priorities for international collaboration. The order of the 

country reports is based on ecogeographic regions where these species can be characterized 

in a similar way as relevant to gene conservation. It does not imply any order of importance 

or priorities. 

The meeting was held immediately after the French National Committee for the 

Conservation of Forest Gene Resources meeting. This enabled participants of both meetings 

to interact closely.

2 EUFORGEN: SOCIAL BROADLEAVES 0 

The participants of the meeting elected Dr A. Kremer (France) as Chair of the Network. 

Drs T. Geburek (Austria) and L. Paule (Slovakia) were elected to act as Vice-chairs. 

Both research and practice of gene conservation in Social Broadleaves are fast evolving 

and have attracted significant attention in European countries during recent years. Regular 

Network meetings are foreseen to further the exchange of information and experience, to 

review the progress made in the implementation of the workplan and to coordinate action at 

the all-European level. The second meeting will be hosted by Switzerland in early 1999.

, ' WGRKRL.1AN 3 

WGrkplan 

Country reports 

These will be completed according to the discussion at the meeting. They should be no more 

than 10 pages of text (standard format: Palatino font size 12, single line spacing). Figures 

and statistics (e.g. area occupied by the species, overview of conservation activities) should 

be given in additional tables, The reports should follow the structure outlined previously 

(see circular letter dated 1 October 1997). All participants will send their reports (electronic 

version and in printed form) to Jozef Turok for compilation and inclusion in the Report of 

the meeting before 1 December 1997. J, Turok will contact countries not represented in the 

meeting and ask for their input as well according to the agreed structure and deadline. The 

Report will be available in April 1998. It was agreed that the introductory presentations by 

A. Kremer, T. Oszako, L. Paule and R. Stephp.n also be included in the Report (to be 

submitted no later than 1 February 1998), 

Synthesis of legislation and other regulations related to 

genetic resources of Social Broadleaves 

This will be prepared by Sven de Vries on the basis of information available in the country 

reports. It will be presented and discussed during the next Network meeting (circulated 

one month before the meeting). Lennart Ackzell will send a copy of the latest version of the 

OEeD and EU regulations to all participants. 

Overview of ongoing research projects 

An overview of the objectives, available results and collaborating partners of ongoing EUfunded 

projects, related IUFRO Working Parties and other relevant international projects 

will be compiled by Antoine Kremer (oaks) and Richard Stephan (beech) before 1 February 

1998. A compilation of ongoing national projects and programmes related to genetic 

resources will be extracted from the country reports by J. Turok. Participants are, therefore, 

encouraged to collect and include information representing the whole research spectrum of 

their respective country. Both outputs will be included in the Report of the meeting. 

Development of joint, long-term, practically oriented gene conservation strategies 

This was considered to be the fundamental task of the new Network. A working group 

composed of Thomas Geburek (responsible for the task), Patrick Bonfils, Richard Stephan 

and Alexis Ducousso will send a questionnaire asking for information about methodologies 

currently used for the in situ and ex situ conservation in European countries. The questionnaire 

will be circulated by 1 February 1998 and replies are to be sent back to T. Geburek 

before 1 July 1998. This basic information (empirical knowledge and experience available in 

countries) will lead to preparing a background document to be presented at the next 

Network meeting. The activity will help to identify topics where additional research is 

needed (spatial structure of diversity in gene conservation units, influence of silvicultural 

practices, etc.). The ultimate aim is to provide technical recommendations (guidelines) for 

the sampling, design and management of gene conservation units of oaks and beech. The 

draft background document will be circulated by T. Geburek one month before the next 

meeting. 

Terminology 

The terminology of the Network will be harmonized using the agreed Norway spruce 

glossary. All participants should send comments and additional terms they wish to include 

to J. Turok by 10 November 1997. The adjusted list will be circulated to all participants

4 ,EU~()RGEN: S()GI~I!! BB()~BI!!E~MES 

before 20 November 1997, in order to facilitate the use of common terms in the country 

reports (to be submitted by 1 December 1997). 

Descriptors and databases 

Jan S. Jensen prepared a list of common minimum descriptors for Noble Hardwoods. This 

was recognized as a good starting point for Social Broadleaves as well. J. Turok will circulate 

this list to the participants before 20 November 1997. Comments should be sent directly to 

J.S. Jensen by 1 March 1998. The list will be discussed at the next meeting (circulated one 

month before the meeting). The objective is to provide a minimum common format for 

databases on gene conservation units at a national level. An international database may be 

set up later. 

Public awareness 

Awareness about the importance of the genetic resources of beech and oaks as cultural 

heritage will be promoted by the Network. loan Blada in collaboration with Jozef Turok will 

formulate a one-page draft about the importance of conserving genetic resources of oaks and 

beech and describing the role of the Network. J. Turok will send him the leaflet produced by 

Noble Hardwoods Network by 10 November 1997. The draft will be circulated by loan 

Blada before 15 December 1997 and comments sent back to him by all participants before 1 

January 1998. It will then be included in the Report of the meeting. 

A collection of slides linked to the gene conservation of Social Broadleaves will be 

established. Dominique Jacques will prepare and circulate a letter with proposed topics to 

be covered by the collection. Ladislav Paule and S. de Vries will assist with organizing this 

Network's collection. It will be presented at the next meeting and completed afterwards. 

Conclusion 

The participants of the meeting elected A. Kremer as Chair of the Network. T. Geburek and 

L. Paule were elected to act as Vice-chairs. The host of the meeting, M. Arbez, INRA Station 

de Recherches Forestieres in Cestas, was commended for the excellent organization of the 

meeting. Following several offers, it was agreed to hold the next meeting in Switzerland, in 

February-March 1999. The final dates will be announced.

· €eUNmR¥ REeeRf'nS 5 

Country Reports 

Beech and oak genetic resources in Romania 

loan Blada 

Forest Research and Management Institute, Bucharest, Romania 

Occurrence and origin 

According to the Romanian Forest Inventory (Anonymous 1984) the total forest area was 

6341472 ha, including 6 223 416 ha of forests and 118056 ha not covered by forested stands. 

The major broadleaved species are beech and oaks. 

Fagus sylvatica is the most widespread broadleaved species, naturally distributed from 

low hills to mountains, i.e. between about 400 and 1400 m (Negulescu and Savulescu 1957). 

It covers 1915657 ha, or 30.8% of the forest area (Table 1). It grows mostly in pure stands of 

large extent, but is also mixed with other native species such as Abies alba, Picea abies, 

Quercus petraea, Ulmus glabra, Acer pseudoplatanus, etc. (Blada 1995). The species is 

characteristic with very good natural regeneration and therefore possesses a high genetic 

variability. It must be emphasized that Romania still has many virgin populations that 

should be protected for future generations. 

Fagus orientalis has a sporadic distribution at low elevations, mostly in the southern part 

of the country but also in southern Transylvania (Stanescu et al. 1997). 

Oaks (Quercus spp.) are widely distributed throughout the country, covering 1 339 065 ha, 

i.e. 18.5%. 

The genus Quercus is represented by seven native species: Quercus petraea, Q. robur, 

Q. pedunculiflora, Q. jrainetto, Q. cerris, Q. pubescens and Q. virgiliana (Georgescu and Moraru 

1948; Stanescu et al. 1997). 

Quercus petraea is the most widespread oak in Romania, covering 670 319 ha or 10% of the 

total forest area (Table 1). It is a native species distributed throughout the country, from low 

hills up to the lower part of the mountains. Its upper limit is 800 m in the eastern 

Carpathians, 1000 m in the southern Carpathians and 900 m in the western Carpathians 

(Negulescu and Savulescu 1957). It grows in pure and mixed stands together with Fagus 

sylvatica, Quercus robur, Carpinus betulus and several Noble Hardwoods. 

Quercus robur is naturally distributed in lowlands but also in low hills, covering 139 856 

ha. Quercus pedunculiflora, Q. jrainetto, Q. cerris, Q. pubescens and Q. virgiliana cover 332325 

ha or 5.3% of the total forest area. 

mable 1. Beech and oaks in Romania 

Species 

motal area 

ha 

% 

Total forests 

Fagus sylvatica 


Quercus robur 

Other Quercus spp.t 

motal beech 

motaloaks 

6223416 

1915657 

670319 

139856 

332325 

6223416 

1 339065 

100.0 

30.8 

10.8 

2.3 

5.3 

30.8 

18.5 

t Q. pedunculiflora, Q. frainetto, Q. cerris, Q. pubescens, Q. virgiliana. 

Economic importance 

Beech and oaks are very important in several industries. They also play a major role in 

reforestation. Details are given in Table 2.

6 EUEORGEN: SOGIA.l.ill BROA.Dl.illEA.MES 

Table 2. Economic importance of beech and oaks in Romania 

Use Beech Oaks 

Silviculture for reforestation 

Furniture industry (with or without veneer) 

Paper industry 

Leather industry (tanning) 

Chemical industry (acetic acid, methylic alcohol, tar) 

House, boat, vehicle manufacture 

Flooring and interior finishes 

Handicrafts and woodwork goods 

Fuel 

Landscaping 

Cooperage (barrel/butt), railway sleepers manufacture 

Woodwheel manufacture 

Plywood 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

Silvicultural approaches used 

• Exclusively natural regeneration for beech. 

• Natural regeneration combined with artificial regeneration (planting but no coppice) for 

oaks. 

Health state of the stands and threats to their genetic diversity 

According to Patrascoiu et al. (1985), the present health state of beech and oaks is as follows. 

Beech has a good to very good health state and there is no threat to diversity. 

The situation is different for oaks: 

• not very good health state for Q. petraea, poor for Q. robur and Q. pedunculiflora and 

very poor for Q. cerris, Q. jrainetto, Q. pubescens and Q. virgiliana 

• negative factors: drought, pollution, defoliating insects, vascular diseases, seed 

predators, unsuitable silvicultural approaches (coppice) used in the past 

• negative effects: slow growth, lack of fructification, decline in health state, death of 

some populations (Blada, unpublished) 

• genetic diversity threatened at population level for all oaks but mainly Q. cerris, 

Q. jrainetto, Q. pubescens and Q. virgiliana. 

Research activities and tree improvement 

The following research activities are under way: 

• Research on natural regeneration in both beech and oaks to maintain the genetic 

diversity of ancestral populations. 

• Research on genetic variation at population level. Thirteen provenance trials were 

laid out about 20 years ago, five in Quercus petraea and eight in Q. robur. Two 

international beech trials were planted in 1996 (Table 3). 

The aim of these studies was to detect the best provenance for reforestation and to 

determine what transfer between regions is possible without loss of adaptability, resistance, 

yield and quality. 

Needs for better research in the future: 

• More comprehensive provenance trials, for all regions of provenance, in both beech 

and oaks. 

• Besides field testing, new and modern techniques should be applied (isoenzyme and 

DNA analyses) to describe genetic diversity; at present the necessary equipment is not 

available. 

Tree-improvement activities consist of the phenotypical selection of plus trees and 

establishment of seed orchards.

COUNmR'V RERORmS 7; 

Species 


Quercus rabur 

Fagus sylvatica t 

Total 

mable 3. Provenance trials 

mrials 

5 

8 

2 

t International trials. ' Approximately. 

Number of: 

Provenances 

33 

27 

44 

Area (ha) 

4.5 

7.7 

4.0' 

16.2 

Conservation of genetic resources 

According to the Romanian Forestry Law (Anonymous 1996) all forests are to be managed 

according to the following principles: 

• sustainable, close-to-nature and multifunctional forest management, for dynamic 

gene conservation 

• active protection and conservation of the biological diversity of forests 

• support of the biological and economic stability and continuity of forests, by 

promoting natural regeneration and improving the planting stock 

• natural regeneration supported in all forests, where possible. If seedlings are used, 

they should derive from adequate seed sources, and only suitable species/ 

provenances can be used. 

Upon this basis the Forest Research and Management Institute in Bucharest initiated a 

national programme devoted to the conservation of genetic diversity in forests. The main 

objective of the programme was the conservation and utilization of the genetic adaptability 

of forest tree populations. According to the OECD regulations, Romania was divided into 

regions of provenance, and consequently, the transfer of the reproductive material between 

these regions must be done according to OECD rules (Enescu et al. 1988). 

In situ conservation 

The goal of in situ conservation in beech and oaks is to maintain the evolutionary genetic 

adaptability of populations over generations. For beech, most of the original natural 

populations are still present in Romania, but it is not the case in oaks where many 

populations were lost for various causes. 

According to Koski (1997) a successful in situ gene conservation must fulfil certain 

fundamental prerequisites: 

• a network of gene conservation stands is to be created with sufficient coverage of the 

spatial genetic variation of the species 

• the number of individual genotypes per population must be large enough to include 

most of the genepool existing in the respective population 

• the system of regeneration must maintain the population, and the regeneration stock 

should predominantly originate from matings within the respective population. 

All the following activities of gene conservation in beech, and partially in oaks, took into 

consideration the above prerequisites. 

Selected seed stands 

To be considered for conservation, a forest stand had to be representative of the natural 

populations, and regenerated naturally. These requirements were fully met for beech and in 

most cases for oaks. 

According to the national register (Enescu 1986), to date 1085 in situ conservation (i.e. 

seed) stands are declared as forest genetic resources, comprising a total area of 34179 ha 

(Table 4). Taking into consideration beech and oaks separately, it can be noted that:

8 EUEaRGEN: SaCI~l1; BRa~[)l1;E~MES ~ 

• out of the 1085 seed stands, 871 were oaks and and 214 beech 

• out of a total area of 34 179 ha, 26057.8 ha (76%) represent oak while 8 121.2 ha (24%) 

represent beech 

• out of a total area of 26057.8 ha of oak, 25077.7 ha (96%) represent natural 

populations and only 980.1 ha (4%) represent artificial ones 

• no artificial beech population was selected. 

It must be stressed that a new inventory of the seed stands is under way and will be 

finished in 1999. 

Table 4. Selected seed stands 

Total 

stands Total area {ha} 

Species {no.} In situ Ex situ Total {ha} 

Fagus sylvatica 214 8 121.2 0 8 121.2 

Quercus petraea 498 17056.8 352.6 17409.4 

Quercus robur 211 4658.8 535.7 5194.5 

Quercus cerris 69 705.1 6.7 711.8 

Quercus frainetto 46 1 851.9 0 1 851.9 

Quercus pedunculiflora 41 785.3 85.1 870.4 

Quercus pubescens 6 19.8 0 19.8 

Total Fagus 214 8 121.2 0 8 121.2 

Total Quercus 871 25077.7 980.1 26057.8 

Total 1085 33198.9 980.1 34179.0 

National parks and biosphere reserves 

In addition to seed stands, beech and oak genetic resources are also conserved in 13 national 

parks and biosphere reserves covering 397761 ha or 2.3% of the total forest area (Table 5). 

These forests are allowed to develop with almost no human interference, leaving the trees to 

reach their natural age and leaving dead wood in the forest. The forests serve as nature 

reserves and research populations. The structure and development of such forests are being 

analyzed and results from these studies used to establish guidelines for close-to-nature 

silviculture. It must be stressed that because of their large areas, national parks and biosphere 

reserves include almost all native species; therefore they represent significant reserves. 

Table 5. National parks and biosphere reserves 

Local name of the park 

Area {ha) 

Retezat (Biosphere Reserve) 

54400 

Rodna (Biosphere Reserve) 

56700 

Domogled - Valea Cernei (National Park) 

60100 

Cheile Nerei - Beusnita (National Park) 

45561 

Apuseni (National Park) 

37900 

Bucegi (National Park) 

35700 

Semenic - Cheile Carasului (National Park) 30400 

Ceahlau (National Park) 

17200 

Cozia (National Park) 

17100 

Calimani (National Park) 

15300 

Piatra Craiului (National Park) 

14800 

Cheile Bicazului (National Park) 

11 600 

Gradistea de Munte (National Park) 

1 000 

Total 

397761

~ G()UN1FRM REP()R1FS 9 

Conservation units 

The areas covered by gene reserves are supposed to be large enough to conserve the genetic 

structures and allow evolution of the population to continue. Since pollen contamination 

from outside sources is undesirable, the core area should be surroundpd by a wide buffer 

zone. 

An inventory of this category started in 1993 and will be finished in 1998. By the end of 

1996 the following results were available (Lalu and Nicolescu 1996): 

• 347 units were selected in 42 counties, each unit consisting of a core area and a 

surrounding buffer zone 

• the total core area, at the country level, was 11 304 ha, with an average of 33 ha per 

core area 

• genetic reserves represented 0.17% of the total national forest area 

• the total buffer zone was 25 805 ha. 

The major criteria for selecting these units were origin, timber production and quality, 

health state and adaptability to environment. Such gene reserves contain the principal forest 

tree species including beech, oaks and Noble Hardwoods. 

Ex situ conservation 

According to Enescu et al. (1989), 1004 ha of clonal and seedling orchards of selected plus 

trees were established in Romania. 

Out of the 1004 ha, 87 ha belong to oak species. No seed orchards were established for 

beech, because in this country it is regenerated only naturally. 

Provenance trials represent another method of ex situ conservation. As mentioned earlier, 

16.2 ha were established (12.2 ha of oaks and 4 ha of beech) (Table 3). 

Beech and oak trials are not sufficient, and more trials should be established. 

Use of reproductive material 

Reproductive material is used according to the Romanian Forestry Law 'Codul Silvic' and 

the OECD rules. According to these regulations, the reproductive material must be collected 

from selected seed stands and seed orchards, taking into consideration that: 

• seed collecting must be done only in years of good fructification, where more than 

50% of the trees have a good crop 

• the number of individuals chosen for seed collecting should be more than 100 

• seed collecting must be overseen by a specialist 

• all significant details about the population from which the seed was collected must be 

recorded. 

Institutions involved in genetic resources activities 

The following institutions are currently involved in the conservation and utilization of forest 

genetic resources: 

• the Ministry of Waters, Forests and Environment Protection, through its Department 

of Forests 

• ROMSILV A - The Romanian National Company of the Forests 

• Forest Research and Management Institute 

• The Romanian Academy of Sciences.

10 EUE0RGEN: S0CIAJ..: BR0ADJ..:EAVES 

Country priorities and capacities 

e Restoration of genetic diversity in oaks, mainly in the southern part of the country, 

where many populations disappeared or are in decline. 

e Stimulation of fructification in oaks, in both seed stands and seed orchards - the 

constraint is lack of 'know-how'. 

e Long-term ex situ conservation or storage of oak acorns - the constraint is no facilities. 

e In situ and ex situ conservation of the most representative oak populations. 

e Creation of artificial mixed stands of oaks with other species, preferably Noble 

Hardwoods; also, re-introduction of shrubs should be supported. 

e Description of genetic diversity within populations of the main species by using modern 

techniques (genetic markers). 

Current international collaboration 

The following collaborations are under way: 

1. Project 'Genetics and improvement of forest trees', cooperation between INRA (France) 

and the Forest Research and Management Institute, Bucharest. 

2. Project INCO-COPERNICUS 'Geographic map of oak gene diversity at an European 

scale for conservation and utilization of genetic resources'. 

3. Project 'Genetic resources of broadleaved forest tree species in southern Europe', 

cooperation between IPGRI, Luxembourg, Bulgaria, Moldova and Romania. 

Needs for international collaboration 

Without international support, the Forest Research and Management Institute from 

Bucharest cannot solve the following: 

1. Conservation/storage of oak acorns for at least 2 years; lack of know-how and facilities. 

2. Genetic inventories based on genetic markers (isoenzymes, DNA), because there are no 

specialists and facilities and because the financial situation of our Institute is very difficult. 

3. Construction of geographic maps based on the genetic differences (chloroplast DNA 

analyses). 

Bibliography 

Anonymous. 1984. Romanian Forest Inventory (unpublished). 

Anonymous. 1996. Legea Codului Silvic No. 26/1996. Monitorul Oficial. 

Blada, 1. 1995. Conservation of Romanian forest genetic resources. In Romania, Country 

Report, International Conference and Programme for Plant Genetic Resources (ICPPGR). 

Enescu, V. 1986. Seed stands catalogue. ICAS, Bucuresti. 

Enescu, V. et al. 1988. Zonele de recoltare a semintelor forestiere in Romania. ICAS, Seria a H 

a, Bucuresti. 

Enescu, V. et al. 1989. Ameliorarea prin selectie si incrucisare a arborilor plus si crearea 

plantajelor pentru producera semintelor genetic ameliorate de. foioase si rasinoase. 

Unpublished manuscript, ICAS. 

Georgescu, c.c. and 1. Moraru. 1948. Monografia stejarilor din Romania. Ed. Universul. 

Koski, V. 1997. In situ conservation of genetic resources. Pp. 5-13 in Technical guidelines for 

genetic conservation of Norway spruce (Picea abies (L) Karst.) (V. Koski, T. Skmppa, 

L. Paule, H. Wolf and J. Turok). IPGRI, Rome. 

Lalu, 1. and L. Nicolescu. 1996. Catalogul National al resurselor genetice forestiere, partea 1. 

Unpublished manuscript, ICAS. 

Negulescu, E. and A. Savulescu. 1957. Dendrologie, E.A.S., Bucuresti. 

Patrascoiu, N., O. Badea, and N. Geambasu. 1985. Studiu privind dinamica starii de sanatate 

a padurilor pe baza informatiilor obtinute din monitoringul forestier. Unpublished 

manuscript, ICAS. 

Stanescu, V., N. Softelea and O. Popescu. 1997. Flora forestiera lemnoasa a Romaniei, Ed. 

Ceres, Bucuresti.

· GOI.JNmR¥ REBORmS 11 

Present status of the conservation and use of broad leaved forest genetic 

resources in Moldova 

Gheorghe Postolache 

Institute of Botany, Chi;>inau, Moldova 

The total forest fund occupies 11.7% of Moldova's territory (394400 ha), including 325 400 ha 

covered by forests (9.6% of the territory). The total growing stock is 35.14 million m 3 

constituting only 8.1 m 3 of timber stock and 0.075 ha of forest land per capita. 

The state forest organizations manage 345600 ha or 87.5% of the total area, of which 

295300 ha (or 85.6%) are covered with forests. The rest of the area, which is made up of 

48800 ha or 12.4% of the territory, is managed by municipalities and agricultural farms. 

The principal species in the natural forest are oaks, which occupy 140500 ha. Three 

species of oaks occur: Quercus petraea (Matt.) Liebl., Q. robur L. and Q. pubescens Willd. 

Forest types are distributed according to the zonal and vertical division of the territory. 

Forest associations of Q. petraea occupy watersheds and slopes of different expositions in the 

central part of Moldova (180-400 m). Quercus robur occupies sites of low relief. In the 

northern part of the country natural forests predominate. Forest associations of Q. pubescens 

are widespread in the south. European beech (Fagus sylvatica) grows only in the central part 

of Moldova. 

Thus, in spite of the small territory occupied by forests, the forest vegetation is rather 

diversified. Twelve 'zonal' and six 'azonal' types of forests were recognized (Gheideman et 

al. 1964; Postolache 1995). 

According to the forest planning inventory (1985) the area of forest stands, which do not 

correspond to the ecological site conditions, constitutes around 40% of all the forests. 

Ninety-five percent of this area is covered by species such as Robinia pseudo-acacia (52%), ash 

(15%) and hornbeam (8%). 

A tendency to invasion by the introduced maple tree (Acer negundo) was noticed in the 

meadow forests in the valleys of the Nistru and Prut rivers. This maple species invades 

mainly willows and poplar stands. Reconstruction and careful silvicultural management are 

now carried out with the aim of re-establishing the original stands (including native oaks). 

The health condition of many forest stands worsened considerably recently. The average 

annual surface of forests damaged by defoliating insects constitutes 50 000-70 000 ha (16-22% 

of the area occupied by forests); this area has to be subjected to some active measures of 

control every year. 

Droughts, frosts and reduced biological resistance of some forest plots lead to the further 

spreading of pathogens and overall decline. 

The natural regeneration of most oak stands is satisfactory (Ivanov 1962), and most stands 

were regenerated in this way. Natural regeneration is very difficult in the damaged or 

declining stands. 

In the application of natural regeneration, priority is given to stands which consist of 

native species, thus contributing to gene conservation. Artificial regeneration methods were 

applied after clear-cutting, successive logging, etc. The application of different methods of 

regeneration aims at achieving a structure similar to that of natural forests. 

Forest plantations have been created during the last 50 years on a total area of 171 000 ha. 

Forest stands (113 000 ha) and forest belts (21 000 ha) have been planted. 

The concept of creating stands similar to the natural ecosystems through the use of the 

native species was established. The origin of forest reproductive material for the creation of 

more productive and resistant forest plantations in extreme ecological conditions was 

recognized and controlled. That is why conservation of the forest genetic resources is one of 

the principal objectives of our silviculture.

~2 EUF(,;)RGEN: S(,;)CIAI,;j BR(,;)ADI,;jEAXlES 

Other measures of forest gene conservation were initiated: 

• designation and conservation of most valuable forest stands 

• identification and conservation of some rare species 

• conservation of trees with exceptional qualities 

• creation of genetic collections of valuable species, subspecies, clones, etc. 

• conservation of seeds, pollen, meristem, etc. of valuable genotypes. 

Three forest reserves, five nature reserves, 10 protected natural landscapes and 22 'natural 

monuments' were designated by the Government of Moldova with the aim of conserving 

valuable forest stands, rare species and their populations. 

Most valuable stands of beech and oak (Q. petraea, Q. robur) are conserved within three 

reserves: Codrii, Plaiul Fagului and Pad urea Domneasca. 

Local valuable populations which were not contained within the reserves were 

distinguished as an independent protected category of stands. These are represented by 

valuable populations from the standpoint of timber production, genetic properties, etc. 

The reserves with large surfaces give the possibility to protect and conserve species in 

different types of forests. 

The establishment of forest plantations on the basis of ecological genetics is a central issue 

in creating new forests. Great attention is being paid to this issue. 

The old trees are a particular category of conservation of forest resources. Owing to 

favourable ecological conditions and their valuable genotype, old trees are distinguished by 

vitality and resistance to extreme conditions. They are characterized by a larger size, height 

and diameter. The total of 372 old trees identified in the Republic of Moldova included 265 

Quercus robur, 10 Q. petraea and 10 Fagus sylvatica. 

The creation of clonal archives and collections of some valuable tree species is another 

form of conservation of the forest genetic resources. Clonal archives of some valuable forms 

of oak trees were created in the forest enterprises of Bender and Stra~enni. 

Until now, the species forming natural populations - beech, Fagus sylvatica (Istrati 1975); 

oak, Quercus robur (Cuza 1994; Gumeniucm et al. 1994) and cherry tree, Prunus avium 

(Gumeniuc 1987) - have been studied. Following from this research, recommendations were 

formulated for division of the forest into seed districts for oak, and for the creation of a forest 

'seed base'. 

References 

Cuza, P. 1994. Variabilitatea frunzelor stejarului pedunculat (Quercus robur L.) din Republica 

Moldova. Revista padurilor 2:2-8. 

Gheideman, T., B. Ostapenco, L. Nicolaeva, M. Ulanovski and N. Dmitrieva. 1964. Tipi lesa i 

lesnye assotiatii Moldavskoi S.s.R. Khisinev. 

Gumeniuc, 1. 1987. Morfologhiceskaia i biochimiceskaia izmencivost Cerasus avium L. 

Moench. v Moldavii. Rastitelinye resursi 23(2):239-245. 

Gumeniuc, 1., P. Cuza and C. Istrati. 1994. Polimorfizmul ~i diferentierea populatiilor de 

stejar pedunculat (Quercus robur L.) din nordul Republicii Moldova dupa spectrele 

izoenzimatice. Revista padurilor 1:16-18. 

Istrati, A. 1975. Izmencivosti listiev buka v Moldavii. Izv. AN MSSR. Ser. bioI. i chim. nauc. 

6:3-11. 

Postolache, Gh. 1995. Vegetatia Republicii Moldova. Chi~inau, $tiinta. 

State Forestry Association Moldsilva. 1997. Report on the forest fund's status of the Republic 

of Moldova. Chi~inau.

, GOUNlllRM REPORlllS 13 

Conservation of genetic resources of Social Broadleaves in Ukraine 

Igor M. Patlaj 1, Svitlana A. Los 1, Roman M. J atzyk 2 and Ihor M. Shvadchak 3 

1 Ukrainian Research Institute of Forestry & Forest Melioration, Kharkiv, Ukraine 

2 Research Institute of Mountain Forestry, Ivano-Frankivsk, Ukraine 

3 State University of Forestry and Wood Technology, Lviv, Ukraine 

Occurrence and origin of oaks and beech 

Ukraine is not very rich in forests. Only 14.2% of its territory is covered by forests, with a 

total forest area of 8.6 million ha. Because of the diversity of climatic conditions, the 

distribution of the forests over the country's territory is quite irregular. Usually, five natural 

zones are differentiated: mixed forests, forest-steppe, steppe, and the mountain regions of 

the Carpathians and the Crimea. The major part of the forests is concentrated in the 

Carpathians and the mixed forests zone where they occupy 37.5 and 29.8% of the territory, 

respectively. In the relatively small territory of the Crimea, 28.7% is forests. Only small 

forest areas can be found in the forest-steppe and steppe zones. In these zones the land 

covered by forest amounts only to 12 and 4%, respectively. 

The species composition of Ukrainian forests is diverse. The forests with predominance 

of Social Broadleaves occupy just above 2 million ha, of which 1.69 million ha are forests 

with a predominance of oaks. 

The range of Quercus robur covers almost all the plain territory of Ukraine. The forests 

with predominance of this species occupy 27% of the total forest area (1.57 million ha). In 

the Carpathian region Q. robur grows on foothills and reaches the altitude of 500 m asI. 

The forests with predominance of Quercus petraea are concentrated in the southwestern 

regions and in the Crimea. They reach the altitude of 700 m in the Carpathians, some stands 

reaching 1000 ffi. Quercus petraea grows on poorer soils, where Q. robur and Fagus sylvatica 

cannot compete. Quercus petraea stands occupy 20 000 ha in the Carpathians. 

The forests with predominance of F. sylvatica occupy 560 000 ha. They are situated in the 

mountain regions of the Carpathians and the Crimea. Most beech forests (524000 ha) are 

concentrated in the western regions of Ukraine. About 80% of the beech forests grow in the 

mountain regions of the Carpathians (420000 ha, 38.3% of all forests of this region). Beech is 

the principal forest tree there. It grows from 150 to 1300 m asl. 

It should be pointed out that the taxonomic position of the Crime an beech has not been 

explained until now. Poplavskaja (1928, 1936) described this beech as a distinct species - 

Fagus taurica PopI. Some time later the beech populations from the lower zone of the 

Crimean mountains were more frequently described as Fagus orientalis Lipsky and from the 

upper one as F. taurica (Molotkov 1966; Milescu et al. 1967). Sometimes the Crimean beech 

stands are described as a mixture of individuals of F. sylvatica and F.orientalis with the 

occurrence of several hybrid forms (Wulff and Tsyrina 1925) or as stands formed 

predominantly by F.orientalis with an admixture of F. sylvatica (Privalova 1958). Several 

authors consider it to be a transitional form between F. sylvatica and F. orientalis, which is 

morphologically closer to F. orientalis (Czeczott 1932, 1933), or as a subspecies identical to the 

Balkan beech Fagus sylvatica subsp. moesiaca (Maly) Cerny. (Didukh 1992, 1997). It should be 

noted that all the authors based their classifications exclusively upon morphological 

characters.

;14 EI..JE0RGEN: S0GIAIiZ BR0A[)IiZEAMES . 

Current economic importance for the forestry sector 

The wood of Q. robur, Q. petraea and F. sylvatica is widely used in furniture, aircraft, 

shipbuilding, the chemical industry, buildings, etc. because of its high mechanical and 

aesthetic characteristics. 

Presently the wood of oak and beech is widely exported, particularly to countries of 

western Europe (some 25% of the total export of forestry production). 


The forests of Ukraine are subdivided into two groups. The forests of the first group (I) 

represent 50.1 % of the forests and they perform protective functions. These are forests of 

green zones around the cities, water-protective forests, soil-protective forests, roadside belts 

and reserves. The management carried out in these forests aims at their conservation and 

enhancing their protective role. The commercial forests, which are the main source of wood, 

comprise the second group (H). 

Oaks and beech are the principal tree species. They predominate in natural stands in the 

forests of groups I and H. These species are widely used for the establishment of artificial 

stands in suitable soil and climate conditions. 

Health state of the forest stands and threats to genetic diversity 

An important part of oak stands has low resistance to biotic and abiotic factors at present. 

Climatic factors, particularly droughts, play a major role in the weakening of 45.9% of oak 

stands affected by decline. Unsuitable forest management measures were the cause of 

decline in 6.5% of oak stands. The majority of these stands originated from vegetative 

reproduction. 

Diseases cause weakening of 15.6% of declined oak stands, and 17% of declined stands 

were affected by defoliating insects, which are the main factor of damage in Ukrainian 

forests. 

The considerable extent of forest harvesting in mountainous conditions reflected on the 

state of Social Broadleaves forests of the Carpathian region. Many high-quality beech stands 

were replaced by pure spruce stands regularly subjected to wind and snow breakage, 

diseases and insect pests. This resulted in disequilibrium of the forest ecosystems. 

Research activities and capacities related to genetic resources/diversity 

The Laboratory of Forest Tree Breeding, Research Stations and Carpathians branch of the 

Ukrainian Research Institute of Forestry and Forest Melioration have worked for a long time 

on the problem of conservation and rational use of the forest genetic resources. In the 1960s 

some research on the selection of plus trees was carried out under the guidance of Prof. 

Piatnitsky. Subsequently the investigations were oriented toward the creation of a constant 

seed-production base for the principal forest species under the guidance of Pr of. Molotkov. 

All this research covered all regions of Ukraine and included programmes for gene 

conservation in situ and ex situ: 

• surveys of natural forest and selection of gene reserves, plus trees and seed stands, 

including research on population structures, genetic diversity and the dynamics of 

taxation indexes 

• creation of clonal archives of plus trees at experimental stations 

• creation of seedling and clonal seed orchards 

• creation and study of progeny tests of the best-performing trees and stands. 

Social Broadleaves hold a prominent place in this research.

~ G0UNmB¥ BER0BmS 15 

Current genetic conservation activities in situ and ex situ 

Since the early 1980s, gene reserves have been designated as follows: 6789.1 ha for Q. robur, 

265.2 ha for Q. petraea and 2820.4 ha for F. sylvatica. The distribution of gene reserves in 

Ukraine is given in Table 1. The largest reserve areas are concentrated in forest regions. 

Presently the inventory of gene reserves is being undertaken. This will provide the 

opportunity to estimate the current situation of genetic resources. 

For several years, studies on European beech diversity in the Carpathian region and 

adjacent territories have been carried out by the Ukrainian State University of Forestry and 

Wood Technology (Lviv), together with Technical University in Zvolen, Slovakia. Genetic 

diversity and differentiation of beech populations (20 European beech and 7 Crimean beech) 

have been studied by electrophoresis. 

Genetic differentiation was estimated on the basis of genetic distances. The results 

confirm a low degree of differentiation within F. sylvatica compared with F. orientalis, the 

values of subpopulation differentiation of European beech populations being only 

approximately 30% of the differentiation among F. oriental is populations. Within European 

beech populations from western Ukraine a slight differentiation between populations from 

the southwestern and northeastern macroslopes of the Carpathians, and plain populations 

from the northeastern limit of beech distribution range was observed. Crimean beech is also 

much more differentiated than Carpathian beech. Crimean beech occupies an eccentric 

position between both beech species; however, it is much more similar to F. orientalis than to 

F. sylvatica (Paule et al. 1993; Vysny et al. 1995; Gomory et al. 1998a, 1998b). 

The intensity of reproduction in beech root meristem cells was examined in cytological 

studies by the University together with the Institute of Forestry and Forest Melioration. The 

results led to the conclusion that some differences exist in the mitotic activity of root 

meristem cells of beech seedlings from different populations. Since the mitotic kinetics of 

the meristematic cells of all plants depend not only on ext~rnal factors but also on internal 

factors, it is quite possible that the results reflect the genetic diversity between populations 

(Kyrychenko et al. 1995). 

Clonal propagation of European beech and sessile oak is studied at the Laboratory of 

Tissue Culture. For this purpose explants were taken from different vegetative and 

generative plant parts of beech (buds of different-aged trees, hypocotyls, cotyledons) 

(Bazyuk and Fedyaj 1995). 

Relevant nature protection policies and activities 

Among the legislative acts related to conservation of genetic resources, the Law on Natural 

Reserves of Ukraine (of 5 May 1993) is in force. According to this law any activity which 

might affect negatively the state of protected areas is forbidden. The Committee of 

Environmental Protection is the administrative unit specialized in this field. The protection 

is implemented by the institutions on the territory in which they are situated. Protection of 

forest areas is carried out by regional forest offices. 

Table 1. Gene reserves of Social Broadleaves in Ukraine 

Quercus robur Quercus eetraea Faflus syJvatica 

Natural zone Area {ha} Number Area {ha} Number Area (ha) Number 

Mixed forest 2526.3 80 52.4 1 2.0 1 

Forest-steppe 3935.8 160 29.0 2 1789.8 60 

Steppe 269.0 1 128.0 7 0.0 0 

Carpathians 58.0 1 22.1 2 1028.6 7 

Crimea 0.0 0 33.7 4 0.0 0 

Total 6789.1 257 265.2 16 2820.4 68

16 EUE0RGEN: S0GIAU BR0ADIlEAVES 

Tree-improvement activities 

Research activities on the conservation of genetic resources are usually associated with tree 

breeding. Conservation of genetic resources aims at the reproduction of genetic diversity in 

the most valuable trees and stands on a broad scale for the creation of productive and stable 

forests. 

The phenotypically best-performing and most productive stands were classified as seed 

stands. Plus stands of Social Broadleaves occupy 1758.6 ha. Among them Q. robur is 

represented by 1588.6 ha, Q. petraea by 30.2 ha and F. sylvatica by 139.8 ha (Table 2). By now 

1564 plus trees of Social Broadleaves have been selected in the stands of Ukraine. Among 

them Q. robur is represented by 1212 trees, Q. petraea by 165 trees and F. sylvatica by 181 

trees. Forest management in these stands should be carried out in such a way to ensure the 

conservation of the genetic information. 

Clonal archives were created for the conservation and study of the plus trees. There are 

23.1 ha of clonal archives of Q. robur and 0.6 ha of Q. petraea in Ukraine. The seedling and 

clonal seed orchards were created on large areas to provide forestry with seeds. Another 

objective of the creation of seed orchards is gene conservation. The distribution of the seed 

orchards over the territory of Ukraine is given in Table 2. The total area of clonal seed 

orchards of Q. robur amounts to 492.8 ha, the seedling seed orchards to 60.2 ha. Seed 

orchards of Q. petraea and F. sylvatica are small, and need to be extended. 

Research on the inheritance of useful traits of Social Broadleaves is carried out through 

progeny tests. The areas and numbers of the progenies in the progeny tests are shown in 

Table 2. Progeny tests of Q. robur occupy an area of 32.05 ha where 852 progenies are tested. 

Progeny tests of Q.petraea are limited (6.4 ha, 134 progenies). The trees and stands whose 

progeny had the best growth performance and high adaptability were selected. 

From the beginning of the century to the present, 12 plots of provenance tests of Q. robur 

were established in Ukraine on a surface of 69.3 ha (323 provenances). The bulk of the area 

is concentrated in the forest-steppe region (Table 2). As for F.sylvatica there is only one 

provenance trial zone (1.2 ha) in the Carpathians, where 45 provenances of this species are 

represented. The delimitation of seed zones is developed on the basis of the provenance 

tests and altitudinal-ecological studies. In 1995, on the territory of an experimental training 

forest of the Ukrainian State University of Forestry and Wood Technology, a provenance test 

of European beech were established on an area of 2.4 ha, where 70 provenances of the 

species were represented from western, central, southern and eastern Europe. These studies 

are carried out within the framework of the European Network on the Evaluation of Genetic 

Resources of Beech, and are coordinated by the Institute for Forest Genetics and Forest Tree 

Breeding in Grosshansdorf (Germany) (see von Wuehlisch et al., this volume). 

The Laboratory of Forest Tree Breeding carries out similar studies. We study the growth, 

morphology, phenology and reproduction of Q. robur in the clonal seed orchards and in 

progeny tests. A compilation of morphological descriptors for Q. robur is under way. 

Differences between clones are studied by using biochemical analyses of the phenolic 

compounds. A test for selection of biotypes in different seed zones and technologies for 

collecting, storage and grafting have been developed. Seed orchards have also been 

established. 


The reproductive material from valuable stands is used for natural regeneration and for 

establishment of artificial stands. Artificial plantations are created for the conservation of 

the most valuable stands and trees. The main part of seeds in these cases is produced in 

seed orchards (see section on Tree improvement). The clonal seed orchards of oak begin to 

produce seed when 15-30 years old, and beech at 20-50 years old. The average seed crop of 

the Q. robur seed orchard is almost 300 kg/ha. In the majority of cases the seedlings are 

grown in nurseries for 2 years and then are planted out. Vegetative material is used for 

creation of the clonal seed orchards.

Table 2. Tree im~rovement activities of Social Broadleaves in Ukraine 

~ ~ 

---- 

Progeny tests 

Provenance tests 

Species and Seed stands Plus trees Clonal Clonal seed Seedling seed No. of No. of 

natural zone {ha} (no.} archives {ha} orchard {ha} orchard (ha} Area {ha} ~rogenies Area {ha} ~rogenies 

Quercus robur 

Mixed Forest 330.3 294 11.0 76.9 15.0 1.5 52 0.0 0 

Forest -Steppe 1224.7 459 18.2 382.0 35.2 27.1 634 47.4 215 

Steppe 30.3 323 5.6 28.6 10.0 3.5 119 9.5 60 

Carpathians 3.3 138 0.0 5.3 0.0 0.0 0 12.4 48 

Total 1588.6 1214 34.8 492.8 60.2 32.1 805 69.3 323 


Mixed Forest 27.0 4 0.0 0.0 0.0 0.0 0 0.0 0 

Forest-Steppe 0.0 35 0,0 0.0 0.0 0.0 0 0.0 0 

Steppe 2.6 1 0.6 0.0 0.0 0.0 0 0.0 0 

Carpathians 0.6 59 0.0 1.2 0.0 1.0 14 0.0 0 

Crimea 0.0 99 0.0 0.0 0.0 5.4 120 0.0 0 

Total 30.2 198 0.6 1.2 0.0 6.4 134 0.0 0 

Fagus sy/vatica 

Mixed Forest 0.0 0 0.0 0.0 0.0 0.0 0 0.0 0 

Forest-Steppe 87.6 134 0.0 2.0 16.0 0.0 0 0.0 0 

Carpathians 52.2 41 0.0 1.2 0.0 0.0 0 1.2 45 

Crimea 0.0 12 0.0 0.0 0.0 0.0 0 0.0 0 

Total 139.8 187 0.0 3.2 16.0 0.0 0 1.2 45

18 EUF;eBGEN: SeGIAUl BBeAI1)UlEA~ES 

Institutions involved in genetic resources activities in Ukraine 

The Ukrainian Institute of Forestry and Forest Melioration (Kharkiv) is the leading research 

institution working on the conservation of forest genetic resources in Ukraine. This Institute 

has a network of Research Stations covering the whole country. The regional Ukrainian 

Research Institute of Mountain Forestry (URIMF, Ivano-Frankivsk) was created on the basis 

of the Institute's Carpathian branch in 1993. In the Carpathians, related investigations are 

also carried out by the Ukrainian State University of Forestry and Wood Technology (Lviv). 

Summary of country capacities and priorities 

The forests of Ukraine have a valuable genepool of Social Broadleaves which needs to be 

studied, conserved and reproduced. The best-performing natural stands were selected and a 

network of clonal archives, seed orchards, provenance and progeny tests established in 

previous years. On the other hand, the activity of the mentioned institutions in the field of 

conservation of genetic resources is considerably limited by the lack of funds. 

Since in the past more emphasis was placed on the conservation and study of plus trees, 

now it is necessary to focus on study and conservation of the most valuable populations. The 

majority of gene reserves was selected in the early 1980s; thus it needs repeated inventory 

with biochemical, cytological and molecular genetic methods. Unfortunately, the lack of 

financial support resulted in a decrease in research activities and in the impossibility of 

applying modern methods requiring expensive equipment. 


Undoubtedly the mentioned institutions need international collaboration related to the 

conservation of genetic resources. In addition to exchange of information and participation 

in meetings, the scientific potential of Ukraine should be drawn upon and utilized within 

the framework of European programmes and projects. The following measures are needed 

for the consolidation of international cooperation: 

• exchange of databases 

• exchange of information on research activities and legislation 

• exchange of reproductive material 

• establishment of an international network of provenance tests and altitudinalecological 

studies 

• establishment of an international network for the monitoring of genetic resources 

• development of fellowship programmes in the field of molecular genetics. 

Bibliography 

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Bazyuk, O.F. and L.W. Fedyaj. 1995. P. 1 in In vitro propagation of Fagus sylvatica. Abstracts 

of 6th IUFRO Beech Symposium, Lviv, October 1995. 

Ghensiruk, S.A. 1992. The forests of Ukraine [in Ukrainian]. Naukova dumka, Kyiv. 

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· G0l:JNmRM REB0RmS ~ 9 

Meshkova, V.L. and I.M. Ustskiy. 1997. Forest decline in Ukraine. Proceedings of XI World 

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Patlaj, LM. 1986. Conservation and utilization of gene pool of Ukrainian forests [in Russian]. 

CBNTIleskhoz, Moscow. 

Paule, L., J. Vysny, I. Shvadchak, J. Sabor and D. Gomory. 1993. Genetic resources of 

European beech (Fagus sylvatica L.) in the Slovak, Polish and Ukrainian Carpathians. Pp. 

79-88 in The Scientific Basis for the Evaluation of Forest Genetic Resources of Beech. 

Proceedings of an EC Workshop, Ahrensburg (H.-J. Muhs and G. von Wuehlisch, eds.). 

Working Document of the EC, DG VI, Brussels. 

Poplavskaja, G. 1928. Die Buche in der Krim und ihre VariabilWit. Osterr. Botan. Z. 77:23-42. 

Poplavskaja, G.L 1936. K izucheniju sistematiki krymskogo buka [About the experimental 

study of the systematics of Crime an beech]. Trudy Leningradskogo Obshchestva 

Estestvoispytatelej 65:353-371. 

Privalova, L.A. 1958. Rastitel'nyj pokrov nagorij Babugana i Chatyr-Daga. Obshcheje 

zaklyuchenije po vsemu Krymskomu nagoriyu [Plant cover of the Babugan and Chatyr- 

Dag mountains. General conclusions for the Crimea mountains]. Trudy Gos. Nikitskogo 

botan. sada, Tom 28, Yalta. 

Vysny, J., I. Shvadchak, B. Comps, D. Gomory and L. Paule. 1995. Genetic diversity and 

differentiation of beech populations (Fagus sylvatica L.) in Western Ukraine: The 

Ukrainian Carpathians and Adjacent Territories. Russ. J. Genetics 31(11):1309-1319. 

Wulff, E.V. and T.S. Tsyrina. 1925. Materialy dlya izucheniya krimskogo buka [Materials for 

the study of Crime an beech]. Zapiski Krymskogo Obshchestva Estestvoispytatelej i 

Lubitelej Prirody (Simferopol) 8:75-82.

20 EUE0RGEN: S0€IA~ BR0AD~EAMES 

Conservation of genetic resources of white oaks and beech in Hungary 

Attila Borovics 1 , Zoltan Somogyl and Csaba Mdtyas 3 

1 Forest Research Institute, Department of Tree Breeding, Sarvar, Hungary 

2 Forest Research Institute, Department of Silviculture, Budapest, Hungary 

3 University Sopron, Department of Environmental Sciences, Sopron, Hungary 

Introduction 

Hungary is situated in the central, lowest part of the Carpathian basin. Out of the total area 

of 93 000 km 2 , only slightly more than 2% of the country's area is above 400 m altitude. Hills 

of low elevation, and medium-high mountains between 200 and 400 m occur on 14% of the 

territory. The tw:o lowlands bear a certain similarity of appearance with the loess-covered 

Sarmatian plains of eastern Europe. The topographical regions and natural zones of the 

country are the Great Plain (S and E), the Small Plain (NW), the Sub alpine region (W), the 

Transdanubian Hills (SW), the Transdanubian Central Mountains (mid-W) and the Northern 

Central Mountains (NE). 

The country lies in the temperate zone, at the meeting point of three large climatic 

regions. The continental climate of the eastern European plains is characterized by extremes 

such as hot summers, cold winters and medium precipitation; the well-balanced, cool 

Atlantic climate has cool summers and mild winters; and finally the Mediterranean climate 

is characterized by hot, dry summers and mild, rainy winters. The Hungarian climate is 

therefore very diverse. 

The natural plant cover of Hungary is extremely variable and rich in species. 

Origin and occurrence of oaks and beech in Hungary 

Introduction to Hungarian oak taxa 

Although oaks are still the most widespread tree species in Hungary, their taxonomy and 

genetics are not yet sufficiently known. Oaks can be found under a great variety of site 

conditions and tend to form different races. Transitional forms between species are very 

common too. This is partly because, except for Quercus cerris, oaks hybridize among 

themselves. In addition, variability of important traits, such as early flowering and bud 

burst, is similarly observable. 

It is generally recognized that the Hungarian oak taxa belong to the white oaks in a 

broader sense (subgenus Quercus s. lat.), in particular to two groups, usually designated as 

section Cerris (here Q. cerris only) and section Quercus s. str. (here Q. robur, Q. petraea, 

Q. pubescens and rare species of Q. frainetto). Within Q. petraea and Q. pubescens further 

taxonomic subdivision has been advocated by numerous botanists, but is not yet accepted by 

forestry practice. 

Accordingly, Q. petraea s. lat. is divided into Q. dalechampii Ten., Q. polycarpa Schur and 

Q. petraea s. str.; and Q. pubescens s. str. is differentiated from Q. virgiliana Ten. 

Pedunculate oak 

The species has been exposed to very intense human effects (seed transfer, artificial 

regeneration, selective logging). In central Europe, and first of all in Hungary, pedunculate 

oak of Slavonian origin has been extensively planted since the end of the last century. 

Although the Slavonian populations can be easily distinguished by their exceptionally 

straight stem, relatively regular crown and upward pointing branches, recent genetic 

investigations do not justify the separation of this provenance from other populations in 

central Europe.

Sessile oak 

Three small species belong to the classical sessile oak species. 

III Quercus petraea s.str. is an Atlantic species occurring jointly with hornbeam and 

thriving on cooler mountain slopes with seeping water supply. 

III Quercus dalechampii is the species of the southern slopes, often associated with Q. cerris. 

This species reaches the northern limit of its distribution in Hungary. 

III Quercus polycarpa is also a southern-type species. Its ecological features resemble those 

of pubescent oak, and it often grows mixed and intercrosses with other oaks. 

Pubescent oak 

Pubescent oaks inhabit the extremely dry, calcareous mountain slopes. These sites have been 

mostly planted with Austrian pine in recent decades. Quercus pubescens and the less known 

Quercus virgiliana are, however, very valuable because of their tolerance to arid conditions, 

and should therefore be conserved and promoted. 

Beech 

European beech (Fagus sylvatica L.) is one of the most important and widespread native 

forest tree species in Hungary. It is distributed in the hills and mountains of Transdanubia 

and in the Northern Central Mountains. Beech is mainly regenerated naturally. 

Within its area of distribution, beech grows under diverse climatic, geological and soil 

conditions. The Hungarian beech populations are isolated from each other. They show 

different growth and morphological traits (for example, stem form, bark colour, crown 

structure, timing of bud burst of these populations are easily distinguishable). However, the 

hereditary character of these traits has not been proved yet. 

Economic importance and silviculture of oaks and beech 

The total forest area in Hungary is 1586000 ha, which represents approximately 17% of the 

total land area. Roughly 35% of the forest area is composed of stands of oak species 

(Q. robur, Q. petraea, Q. pubescens, Q. cerris and a negligible proportion of Q.frainetto as well 

as introduced oak species), while 7% of the Hungarian forests are composed of beech stands 

(Table 1). It is worth mentioning that, whereas almost all pedunculate oak forests are planted 

artificially, most sessile oak stands and about 80% of beech stands are regenerated naturally. 

Since Turkey oak is considered less valuable, it is planned to replace 40% of its stands with 

sessile and pedunculate oak species on the appropriate sites. 

Health state of the forest stands and threats to their genetic diversity 

Sessile oak 

Oak decline has caused considerable losses in the Hungarian forest recently. The health state 

of sessile oak has been assessed on 65 plots in Hungary for 10 years. Data for all observed 

trees from 1989 to 1994 are given in Table 2. It can be noted that 12.2% of the observed trees 

have died to date, and that mostly forests in the sub alpine region are damaged. Oak decline 

is also shown to cause growth losses (Somogyi and Standovar 1995). Preliminary results 

suggest that possible causes of sessile oak decline are the following: 

III permanent drought (lack of precipitation, as well as dry air) 

III outbreaks of defoliating insects 

III outbreaks of oak death-watch beetle (Scolytus intricatus) 

III multiplication of xylophagous insects 

III multiplication of bud- and shoot-destroying insects 

III increase of the pathological effect of Armillaria spp. 

III faulty silvicultural practices, e.g. lack of thinning 

III air pollution.

22 EUE0RGEN: S0elAI.:. BR0A[)I.:.EAMES . 

Table 1. Economic importance of oaks and beech (Source: Eorest Research Institute, 1991) 

Natural Growing Annual 

regeneration stock allowable cut 

Species Area (ha) % (%) (1000 m 3 ) % (1000 m 3 ) % 

Q. robur 144326 9.2 10 28993 10.2 635 7.0 

Q. petraea high 91 222 5.8 16304 5.7 371 4.1 

forest 

Q. petraea coppice 94658 6.1 24571 8.6 537 6.0 

Q. petraea total t 185880 11.9 60 40875 14.4 908 10.1 

Q. pubescenst 15657 1.0 100 1 878 0.7 54 0.6 

Q. cerris high 107000 6.9 21 621 7.6 606 6.7 

forest 

Q. cerris coppice 68585 4.3 15182 5.3 442 4.9 

Q. cerris total 175585 11.2 10 36803 12.9 1048 11.6 

F. sylvatica 102457 6.6 80 38952 13.7 799 8.9 

Total forest area 1586760 100 15 284556 100 8997 100 

t Data are for the sessile oak aggregate. 

t Including Q. virgiliana. 

Table 2. Health state of sessile oak (Source: Varga et al. 1995} 

Height class 1 +2+3 Height class 4 

Total 

Total 

No. trees Dead trees trees Dead trees trees Dead trees 

Region observed No. % (No.} No. % (No.} No. % 

Northern Central 9505 961 10.1 8124 537 6.6 1385 424 30.6 

Mountains 

Transdanubian Central 2183 289 13.2 1917 183 9.5 266 106 39.8 

Mountains 

Transdanubian Hills 2487 354 14.3 2148 189 8.8 339 165 48.7 

West-Transdanubia 1689 336 19.9 1307 114 8.7 382 221 57.9 

Total 15868 1940 12.2 13496 1023 7.6 2372 916 38.9 

Height classes: 1 - dominant; 2 - codominant; 3 - intermediate; 4 - overtopped. 


Relevant data for pedunculate oak for the years 1990-94 are summarized in Table 3. 

Table 3. Health state of pedunculate oak (Source: Varga et al. 1995} 

Height class 1 +2+3 Height class 4 

Total 

Total 

No. trees Dead trees trees Dead trees trees Dead trees 

Region observed No. % (No.} No. % (No.} No. % 

Great Plain 1 1586 303 19.1 1434 165 11.5 152 138 90.8 

Great Plain 2 863 125 14.5 786 88 11.2 74 37 50.0 

Transdanubian 1888 159 8.4 1830 128 7.0 58 31 53.4 

Hills 

Transdanubian 651 128 19.6 581 82 14.1 70 46 65.7 

Central Mtns. 

Mezof61d 1555 312 20.1 1262 163 12.9 293 147 50.2 

West- 929 34 3.7 923 31 3.4 6 3 50.0 

Transdanubia 

Total 7472 1061 14.2 6816 657 9.6 653 402 61.6 

Height classes: 1 - dominant; 2 - codominant; 3 - intermediate; 4 - overtopped.

eGUNliR¥ REeGRliS 23 

The causes of pedunculate oak decline are partly similar and partly different from those 

of sessile oak. They are as follows: 

III outbreaks of defoliating insects 

III too much gley and slack water, as well as chalk formation in compacted soils 

III mildew 

III disequilibrium in the water balance within the tree 

III appearance of parasites (Armillaria spp.) 

III outbreaks of scale insects and xylophagous insects. 

Pedunculate oak decline is a periodical phenomenon. Intensive drying of trees occurred 

when the outbreaks of Lymantria dispar followed periods with a rainfall higher than average. 

This happened in 1907-08, 1914-17, 1924, 1962-65 and 1972-74. 

Beech 

The health state of beech stands has declined in recent years, but problems are less serious 

than for oaks (Table 4). The most common cause associated with damaged beech trees is 

drought. Air pollution or acid rains are not considered an important damaging agent in the 

Carpathian basin. 

liable 4. Percentage of health~ stems in beech 

Difference 

Difference 

Region 1992 1993 {1993 - 1992} 1994 (1994 -1993) 

Northern Central Mtns. 1 67.9 59.6 -8.3 55.7 -3.9 

Northern Central Mtns. 2 45.4 32.0 -13.4 35.5 +1.5 

Transdanubian Central 73.2 69.1 -1.4 71.4 +2.3 

Mtns. 

Transdanubian Hills 1 74.4 67.1 -7.3 63.1 -4.0 

Transdanubian Hills 2 73.3 69.1 -4.2 71.6 +2.5 

West-Transdanubia 87.7 82.7 -5.1 86 +3.3 

Source: T6th et al. 1995. 

Research activities related to genetic resources/diversity 

We have hardly any information about the genetic variation of the Hungarian oak and beech 

populations. In collaboration with other countries, we could only conduct a few 

investigations including: 

III Genetic differentiation by RAPD markers of oak species in Hungary (see Bordacs and 

Burg 1997). 

The genetic differentiation of four oak taxa in Hungary was investigated by RAPD 

analysis. In total, 99 trees were sampled to compare levels of genetic diversity and to 

identify the taxa. Among 26 single decamer primers screened, the four resulting profiles 

showed significant differences among the four taxa. 

III Genetic variation in beech populations along the Alpine chain and the Hungarian Basin 

(Comps et al. 1998): 

Seventy-eight European beech population from the Alp Chain and the Hungarian Basin 

were analyzed using eleven alloenzymatic loci. Four pools of populations could be 

discriminated. 

Experiments are under way in the following fields: 

III inter- and intraspecific variation of Q. robur and Q. petraea, and within the Q. petraea 

aggregate for morphological traits 

III controlled crossing among native oak species 

III provenance tests.

24 El.JFORGEN: SOCIAE BROADEEAVES 

Some of the respective results are as follows: 

The status of oak species within the oak aggregate is still very doubtful. To try to separate 

these species taxonomic ally, numerical methods were applied. The frequency distribution of 

discriminant scores, calculated from leaf morphological characteristics of pedunculate and 

sessile oak s. [at., shows that the two oaks can be separated according to these characteristics. 

Similarly, multigroup discriminant analysis of the three minor species of sessile oak resulted 

in three distinguishable groups based on the same morphological characteristics (Fig. 1). 

4r-----~~--~~--------------~------~----~ 

3 

.::.=-.... : "-"···::-··········1;.·······1······················· 

1;.: 1;. : : 

Vl 

x 

ro 

ro 

·u 

c 

ro 

c 

E 0 

.L: 

u 

Vl 

:.a -1 

N 

-2 

-3 

_4L-~--~--~--~--~---L----~~------~----~ 

-6 -4 -2 o 2 4 6 

1 discriminant axis 

Fig. 1. Multigroup discriminant analysis of three sessile oak taxa: 0 = Q. petraea; .11= Q. da/echampii; 

+ = Q. po/ycarpa. 

In the second research field mentioned above, we carried out controlled crossing trials 

with native oaks. In general, intraspecific crossings were more successful than interspecific 

ones, with Q. frainetto showing the highest success rate (Fig. 2). 

It is worth noting that, when grown, combinations of Q. petraea female parents with 

Q. robur male parents were equally successful as the reciprocal combinations. The opinion 

that reciprocal combinations give different success rates may arise from the fact that most 

combinations are made under field conditions where the microclimate within the isolation 

bags may differ considerably. It must be mentioned that the timing of male and female 

flowering can also be synchronized easily in a greenhouse. Another noteworthy result of 

our research so far is that the combination of Q. robur x Q. pubescens showed transient 

morphological characters (for example in density of pilosity, form of lobe, etc.), and that 

pollinated flowers of the Q. robur x Q. cerris combination rapidly aborted around a month 

after pollination (Borovics, unpublished).

GOUNTR¥ REPORTS 25 

N umber of flowers/acorns 

900,---------------------------------------------------, 

Pollinated flower 804 

800+------------------=~----------------------~~--~ 

700+------------------ 

600+-----------------------------------------~ 

500+-----------------------------------------~ 

400+-----------------------------------------~ 

300 +----------1 

200 

100 

0+-'----"" 

rob pet pub fra 

Pollen donor species 

Selfpollin. 

Total 

Fig. 2. Total number of pollinated flowers and viable acorns in a controlled crossing with O. robur 

female individuals in the Bejcgyertyanos seed orchard in 1995. 

Current genetic conservation activities 

Seed stands represent a general basis for genetic conservation (Table 5). No ex situ 

conservation measures for oaks and beech have been taken yet. 

Table 5. Area of seed stands in Hungary 

Species 

Area (ha) 

Ouercus robur 1590.4 

Q. petraea 583.8 

Q. frainetto 12.1 

Q. cerris 308.6 

Fagus sylvatica 308.6 

Source: National Institute for Agricultural Quality Control, 1996. 

The forest reserves system has recently been established' in Hungary. The proportion of 

beech stands in these reserves is higher than that in the country, while oak is underrepresented. 

This is clearly shown in Figure 3, where the area of forests is compared in the 

reserves and the country mean according to the forest climate zones defined in Hungary. 

Country-wide 

Forest steppe 

zone 

23% 

Closed oak 

28% 

Beech 

10% 

Mixed 

hornbeamoak 

39% 

Forest steppe 

zone 

Forest reserves 

20% Mixed 

hombeam-oak 

16% 

Beech 

40% 

Fig. 3. Area of forest reserves. Riparian and other wetland forests are grouped in the f,orest steppe 

climate zone, whereas all types of plantations (hybrid poplar, pine, etc.) are excluded from these 

statistics.

26 EI..JF0RGEN: S0GI~12: BR0~DI2:E~VES • 


Neither oaks (except for pubescent oak) nor beech are endangered species, from a nature 

conservation viewpoint. However, there are many endangered plant or animal species in 

several stands of these indigenous trees; therefore, some of these stands are protected. In the 

case of pubescent oaks, a research programme will be launched next year, which will be 

supported by the Ministry of Environment, and which will focus on the conservation of 

genetic variability. 

Tree-improvement activities and use of reproductive material 

Tree improvement 

The Carpathian basin was once inhabited by a rich and well-adapted oak flora. Human 

activity (clearing of forests, selective logging, coppicing) greatly affected these forests and 

their ecological conditions were severely damaged by the deterioration of the water regime 

of the sites. The excessive planting practice, disregarding the ecological demands of the 

species and the importance of provenance compatibility, resulted in a mixture of valuable 

stands of local provenance and stands of unknown or unsuitable origin, these often 

occurring side by side. Conservation and natural regeneration of ecotypes that are well 

adapted to a particular site must be given high priority in the future. This underlines the 

importance of seed stands. 

There is no tree improvement programme for beech in Hungary, because it is mainly 

regenerated naturally. It is the silvicultural practice that may improve relevant features of 

the popula tions. 

The establishment of Hungarian seed stands of oak and beech was initiated by Matyas in 

the 1950s. These stands were continuously observed and classified from the points of view 

of quality of morphological characteristics, species identity and state of health. Systematic 

provenance testing with oaks started in 1985 at the Forest Research Institute. There are 

provenance tests with stands established over 5 years and approximately 60 populations (in 

a random block design with four repetitions). 

Oak plus tree selection in Hungary was initiated by Harkai in the late 1960s. The plus 

trees were generally selected in seed stands. These plus trees were grafted and planted in 

seed orchards established in several locations in recent years: three for Quercus robur and 

two for Q. petraea. One of the biggest, situated in Bogdasa (southwest Hungary), was 

established jointly by the Forest Research Institute and the Mecsek State Forest Company in 

1989. In this orchard, trees of Q. robur subsp. slavonica were planted. The total area of the 

orchard is approximately 16 ha, and the 2186 grafts are found in six simple random blocks. 

The grafts are clones of 40 plus trees selected in local seed stands along the Drava river. 

Clones were tested for flowering capacity and fertility by the National Institute for 

Agricultural Quality Control (Bordacs 1997). 

Reproductive material 

Seed production of oaks and beech is rather irregular. Therefore, there have always been 

problems with the availability of their reproductive material. Oaks and beech in Hungary 

produce a sufficient quantity of acorns only once every 6-8 years. 

There was an exceptionally large acorn production in 1995 and therefore, a considerable 

surplus of seedlings is available at the moment. In future years, however, a shortage of 

seeds that may amount to 10-90% of demand is expected. Unfortunately, only a negligible 

part of the reproductive material is produced from seeds collected in stands (Fig. 4).

. 0eUNillRM REReRillS 2T/; 

80000~------------~------------r===~mI---------' 

70000 

60000 

50000 

40000 

30000 

20000 

10000 

o 

seedlings 

more 

year 

seedlings 

seedlings 

demand 

~ 

~ 

Vi 

'-< 

~ 

,,' 

28 EUFGRGEN: SGGIAI;.; BRGABI;.;EAllES 

Needs for international cooperation 

• Investigations on the genetic structure of the species (DNA, isoenzyme analysis). 

• Establishment and evaluation of provenance tests. 

• Investigations on the mating system. 

• Standardization of numerical taxonomic investigation procedures: establishment of 

taxonomic keys. 

References 

Bordacs, S. 1997. Pedunculate oak (Quercus robur L.) seed orchard and clone tests in 

Hungary. Conference report of Diversity and Adaptation in Oak Species, 12-17 October 

1997, State College, Pennsylvania, USA. 

Bordacs, S. and K. Burg. 1997. Genetic differentation by RAPD-markers of oak species in 

Hungary. Conference report of Diversity and Adaptation in Oak Species, 12-17 October 

1997, State College, Pennsylvania, USA. 

Borovics, A. 1997. A kocsanytalan tolgyek levelmorfol6giai vizsgalata. Erdeszeti Lapok 86- 

87:125-142 [in Hungarian]. 

Borovics, A. 1997. Tolgyek nemesitesi alapanyagainak vizsgalata. Novenygenetikus 

Szakmernoki dolgozat. GATE, Genetika Tanszek [in Hungarian]. 

Comps, B., Cs. MMyas, J. Letouzey and T. Geburek. 1998. Genetic variation in beech 

populations (Fagus sylvatica L.) along the Alp Chain and in the Hungarian Basin. For. 

Genet. 5(1):1-9. 

Forest Research Institute. 1991. A faallomanyok elofakeszlete, novedeke es a fakitermelesi 

lehetosegek. Kutatasi Jelentes, ERTI Kozponti Konyvtar [in Hungarian]. 

National Institute for Agricultural Quality Control. 1996. Orszagos erdeszeti csmeteleltar 

1996/97 szezon. OMMI, Budapest [in Hungarian]. 

Somogyi, Z. and T. Standovar. 1995. Kocsanytalan tolgyesek egeszsegi allapota, novekedese 

es a termohelyi viszonyok kozotti osszefiiggesek vizsgalata egy erdoreszletben. Pp. 68-76 

in Changing health condition of forests, Magyar Tudomanyos Akademia Kiadvanya [in 

Hungarian]. 

T6th, J., H. Pagony and P. Szontagh. 1995. A magyarorszagi biikkosok egeszsegi allapota. 

Pp. 77-81 in Changing health condition of forests, Magyar Tudomanyos Akademia 

Kiadvanya [in Hungarian]. 

Varga, F., J. T6th and H. Pagony. 1995. A tolgypusztulas Magyarorszagon. In Changing 

Health Condition of Forests. Magyar Tudomanyos Akademia Kiadvanya [in Hungarian].

€GUNmS¥ SEPGSmS 29 

Conservation of beech and oak genetic resources in Slovakia 

Ladislav Paule 

Faculty of Forestry, Technical University, Zvolen, Slovakia 

Introduction 

Forest land in Slovakia represents about 1.9 million ha, of which 90% are in the Carpathians. 

Approximately 40% of the forest land is covered by beech and oak stands. Basic information 

on the gene conservation units for both species is given and gene conservation programmes 

are outlined for both species. 

Occurrence, origin and distribution of beech and oak species 

The forest area in Slovakia covers 1904339 ha in total (Table 1). The proportion of beech is 

29.6% and of oak species (Quercus robur and Q. petraea) 11.26% . The proportion of Turkey 

oak (Quercus cerris) is 2.45%. The present proportion of beech and oaks is lower than the 

original one (HanCinsky 1972). During the last century, beech was replaced in some places 

by Norway spruce and oak stands by Scots pine. 

Natural distribution of the beech forests in Slovakia is in altitudes between 330 and 

1200 m. It forms pure forest stands in the beech optimum and mixed stands with oak, or 

mixed stands with conifers (silver fir and Norway spruce). The natural distribution of beech 

covers all Slovakia except the Western and Eastern Lowlands and the Southern Slovakian 

Karst region. It is also missing in the highest mountains. 

Natural distribution of Q. petraea is in the hills of western, central and eastern Slovakia. It 

forms forest stands up to 700 m, although there are several occurrences even in higher 

altitudes, e.g. Sitno up to 900 m. Natural distribution of Q. robur is on loamy soils and in wet 

lowlands. It is a tree species of lowlands and lower hills up to 450 m. In several parts of 

Slovakia the natural ranges of both species overlap and they form mixed populations. 

One of the classifications of forest vegetation cover was elaborated by Prof. A. Zlatnik 

who, on the basis of intensive investigation of Carpathian forests in Slovakia and in 

Transcarpathian Ukraine, described the natural forest cover in eight vegetation zones which 

could be defined as the climax geobiocenoses determined by geography and macro- and 

mesoclimate in given altitudes. The vertical forest vegetation zones are usually named 

according to prevailing tree species and they correspond with the altitudinal zones usually 

applied in geobotany (planar, colline, submontane, montane, boreal, subalpine, alpine and 

nival): 

• Oak forest vegetation zone 

• Oak-beech forest vegetation zone 

• Beech-oak forest vegetation zone 

• Beech forest vegeta tion zone 

• Beech-fir forest vegetation zone 

• Beech-fir-spruce forest vegetation zone 

• Spruce forest vegetation zone 

• Mountain pine forest vegetation zone. 

The principal characteristics of forest vegetation zones were given in a previous paper 

(Paule 1995).

30 EUFGRGEN: SG01AI... BRGADI...EAVES . 

Table 1. 

Slovakia 

Forest land t 

Actual area (ha) 

% 

Forest area, economic importance and breeding activities in beech and oaks in 

Beech 

563453 

29.60 

107678 

29.41 

Growing stock (1000 m 3 ) 

% 

Annual allowable cut (1000 m 3 ) 

Final logging 1366 

% 

Thinnings 

% 

Total 

Seed stands ~ 

A-category 

2471 

B-category 

14642 

Plus trees 

38 

Seed orchards 

Seed plantations 

Gene reserves 

t As of 1 January 1994. 

36.69 

363 

33.40 

1729 

Sessile + 

pedunculate 

oak 

214506 

11.26 

38587 

10.54 

191 

5.15 

92 

8.42 

283 

745 

3971 

262 

7.00 

149 110 

1200 1119 

t As of 31 December 1997. 

Turkey oak 

46732 

2.45 

8050 

2.20 

59 

1.60 

23 

2.05 

82 

Broadleaves 

1083440 

56.89 

182877 

49.97 

7300 (mixed) 

Oak species in Slovakia 

Except for the two principal white oak species - Q. petraea and Q. robur - the forestry practice 

also recognizes Turkey oak (Q. cerris) and Quercus pubescens. These two species reach their 

northern limit of distribution in Slovakia. The fifth, introduced oak species is Q. rubra which 

grows in several 60- to 70-year-old plantations and numerous younger ones. 

Besides the four indigenous species, taxonomists (Magic 1974, 1975) differentiated 

another five oaks (may be of hybrid origin) with a lower but indigenous occurrence in 

Slovak forests: Quercus dalechampii, Q. polycarpa (both from the section Roburoides), 

Q. pedunculiflora (section Robur) and Q. virgiliana and Q. frainetto (section Dascia). They can 

be found in the inner Carpathians as a continuation of their more abundant occurrence in 

the Hungarian lowlands and hills. 

The significance of these minor oak species for the forestry practice is questionable, since 

it uses only the two principal oak species (Q. petraea and Q. robur) and does not include the 

others in forest management plans and forestry statistics. Their occurrence is, however, 

known and might be interesting from the point of view of gene conservation, independently 

of their taxonomic status. 

The northernmost occurrence of Q. pubescens has been proved in several nature reserves 

within the natural distribution range in Slovakia. Quercus cerris forms lower-quality forest 

stands usually mixed with hornbeam or other oak species. 

Threats to beech and oak genetic resources 

Several factors negatively affect genetic resources of oaks in Slovakia, e.g. grazing by game, 

improper silvicultural practices, lack of natural regeneration, and recently the oak dieback. 

Originally this was considered as a bark beetle outbreak (Scolytus intricatus) and subsequently 

tracheomycosis, but later the scientists proved the complex disease was caused by abiotic 

(drought) and biotic factors combined with human activities (summer logging of oak). 

Beech has mostly been regenerated naturally. Most changes in genetic composition, even 

in the case of natural regeneration, are due to the improper forestry and silvicultural 

practices which led to failure of natural regeneration. Part of the beech stands was 

converted in the past to Norway spruce stands or mixed beech-spruce stands, frequently on 

inadequate sites.

" COUNTRY REPORTS 31 

Conservation aims and current state of conservation activities 

On the territory of Slovakia the first regulation aimed at seed procurement from approved 

stands was adopted in 1938. In keeping with this legal regulation, the seeds of conifers 

(Norway spruce, silver fir, Scots pine and European larch) could be collected only in 

approved stands (A and B category) and seed transfer was allowed only within 'silvicultural' 

seed zones. The silvicultural zones were defined according to their geographical distribution 

and the length of vegetation period. 

Subsequently these regulations were updated in 1965 and again in 1985 and 1988. The 

last one also includes seed procurement from the approved seed stands and the seed transfer 

within seed zones for pedunculate and sessile oak and beech. 

Oaks and beech were absent from the first regulations because these tree species were 

mainly regenerated naturally. In oaks the artificial regeneration by seeding and planting 

was more common than in beech. At present the proportion of beech reforestation in 

Slovakia is rather high. Beech is frequently a component of combined regenerations to 

establish mixed forest stands. 

For beech there are four seed zones within the natural range (Sub-Tatra, Eastern Slovakia, 

Central Slovakia and Little Carpathians) and two zones outside the natural range (Tatra and 

Southern Slovakia), for sessile oak there are three zones within the natural range (Eastern 

Slovakia, Central Slovakia and Southern Slovakia) and three seed zones outside the natural 

range (Northern Slovakia, Western Slovakia and Southeastern Slovakia). For pedunculate 

oak there are three zones within its natural range (Western Slovakia, Southeasern Slovakia 

and Southern Slovakia) and three zones outside the natural range (Northern Slovakia, 

Eastern Slovakia and Central Slovakia). Vertical transfer of seeds is allowed within ±200 m 

for beech and ±150 m for oaks from the altitude of approved stands where the seed was 

collected. 

At present, 111 gene reserves are defined for all tree species in Slovakia, five (1200 ha) 

and six of them (1119 ha) are established only for beech and oaks, respectively, seven of 

them (3637 ha) for mixed stands composed of oaks and beech, and seven gene reserves 

(3663 ha) for mixed stands composed of tree species other than beech and oaks. They occur 

mainly in central Slovakia and some in western and eastern Slovakia. 

Facilities for long-term conservation of acorns and beech nuts have been built up in 

Liptovsky Hradok. In these facilities the storage of acorns and beech nuts can be maintained 

up to 5 years to cover the time gaps between the seed years. 

Four provenance trials have been established with beech. Those established in 1968-1972 

and 1982 comprise the Slovak and Czech provenances (20 and 27, respectively) (Cervenka 

and Paule 1982; Paule 1982). The latter two are a part of international provenance 

experiments coordinated by the Federal Forestry Research Center in Grosshansdorf, 

Germany, containing 100 and 31 provenances, respectively. Trials were established in 1996 

and 1998. 

At present there are 74 forest reserves, of which 5% are in the oak forest vegetation zone, 

2.42 and 3.78% are in the beech-oak and oak-beech forest vegetation zones, respectively, and 

9.19% in the beech forest vegetation zone. For comparison, the proportion of nature reserves 

in the first four forest vegetation zones (20.39%) is about one-third of the total proportion 

(69.41%) in the forests in Slovakia. It means that there is a lower proportion of virgin and 

natural forests with oaks and beech than would be expected according to the general 

occurrence (V oloscuk 1993). 

Nevertheless, there are also some natural oak forests with Q. robur (Palarikovo) or 

Q. petraea (Kasivarova, Bujanov, Sitno) as well as numerous beech virgin forests in eastern 

Slovakia (e.g. Vihorlat, Stuzica, Rozok,) whose size ranges from 16 ha (Kasivarova) to 600 ha 

(Stuzica) (Korpel' 1993). The forest reserves are in most cases a part of gene reserves, 

although the legal status of both categories is defined by two different Acts (Nature 

Protection Act and Forestry Act and accompanying regulations). The most common 

occurrence of beech forests in Slovakia (and also the forest reserves) is in eastern Slovakia

32 EUFORGEN: SO€IAl.ill BROAIl>l.illEA\lES 

where conifers are rmssmg in the natural composition (Carpathian disjunction). The 

Carpathian disjunction is on the southern side of the Carpathians about 200 km wide (from 

Khust in Transcarpathian to Kosice in Eastern Slovakia) due to climate. 

Relevant research activities and needs 

Genetic diversity of indigenous populations was investigated not only from the territory of 

Slovakia but also on the adjacent regions of the Western Carpathians (Czech Republic, 

Poland, Ukraine and Romania) (Gomory et al. 1992, 1995, 1998; Paule et al. 1995; see Paule 

and Gomory, this volume). 

There is, however, a lack of information on genetic diversity and differentiation of oak 

populations from Slovakia, although some material has been used for broader genetic 

investigations on DNA polymorphisms by Kremer et al. Studies of genetic diversity and 

differentiation of oak populations are planned for the coming years. Further research is 

needed on the taxonomy and genetic structures of forest stands with the occurrence of minor 

species as well as on the genetic composition of the forest stands with the occurrence of the 

principal oak species, Q. petraea and Q. robur. 

The geographic variation has been investigated in beech in provenance trials and 

significant differences were found between eastern Slovakian provenances and the 

remainder of Slovakia, although in western and central Slovakia some good-performing 

provenances were also found along with slow-growing ones. Beech provenances have also 

performed well in the international provenance trials (Cervenka and Paule 1982; Paule 1982). 

There is only a single open-pollinated progeny test of beech, established in Slovakia in 

1961 (Cervenka and Paule 1979). 

Significant differences among beech populations concerning the crown shape and quality 

and stem quality (including forking) among individual regions were observed (Vesely 1977). 

Oak provenance trials are few, established by the Forest Research Institute in Zvolen but 

these were not inventoried systematically (Korenek 1980). We have little information on the 

geographic variation of oak populations. 

The occurrence of minor oak species was also studied and a comparative trial established. 

Unfortunately this trial, owing to a restricted number of parent trees of each species, did not 

give sufficient information on the intra specific variation of oak species involved. There is 

also a lack of information concerning the spatial distribution of individual oak species. 

Further investigations of population structure based on morphological traits as well as on 

genetic markers and provenance trials are needed for these species. 

Beech, in contrast with other tree species of economic importance, is in a favourable 

situation. Owing to its ecological features it has seldom been regenerated artificially, and 

the common silvicultural practice has always relied on natural regeneration. In eastern 

Europe there are, however, also traces of improper forest management of beech stands. 

Coppice stands, mainly in mixtures with oak, are characteristic for the contact zone with 

agricultural land in lower altitudes. There were many cases when deforestation and 

replacement of beech stands by more productive coniferous ones took place. Artificially 

regenerated beech stands were established during the last two centuries. 

Natural regeneration will remain the best way for gene conservation of indigenous beech 

populations. The significance of gene conservation is emphasized by the prognosis of 

climate changes. If, in the case of climate change, the necessity to replace local populations 

by more southern ones occurs, genetic resources should be available. 

Science is already partially prepared for this event. The provenance trial with European 

beech has been established which includes 155 provenances in two series, established in 20 

sites (incomplete provenance numbers); the previous series contained over 150 provenances 

planted on 10 sites (see von Wuehlish et al., this volume). The main aim of these provenance 

experiments is to test the adaptation potential of individual provenances to changed 

environmental conditions.

, GOI.JNTB¥ RERORmS 33 

Acknowledgements 

Thanks are due to Drs D. Gomory, and D. Magic for their valuable advices and to Drs 

J. Hoffmann and R. Longauer for providing the latest statistical information on seed stands 

and gene reserves in Slovakia. 

References 

Cervenka, E. and L. Paule. 1979. Wachstum der Nachkommenschaften von freiabgebhihten 

Auslesebiiumen der Buche. Acta Facultatis Forestalis, Zvolen 21:47-66. 

Cerve~a, E. and L. Paule. 1982. Vyskovy a hrubkovy rast slovenskych proveniencii buka 

(Fagus sylvatica L.) [Height and diameter growth of Slovak provenances of beech (Fagus 

sylvatica L.)]. Lesnicky casopis 28(6):409-422. 

Blattny, T. and T. St'astny. 1956. Prirodzene rozSfrenie lesnych drevin na Slovensku [Natural 

distribution of forest trees in Slovakia]. SVPL, Bratislava. 

Domin, K. 1932. The beech forests of Czechoslovakia. Pp. 63-165 in Die Buchenwiilder Europas 

(E. Riibel, ed.). Vlg. Hans Huber, Bern-Berlin. 

Fekete, L. and T. Blattny. 1913/1914. Die Verbreitung der forstlich wichtigen Biiume und 

Striiucher im Ungarischen Staate. Commissionsverlag, Schemnitz. 

Gomory, D., B. Comps, L. Paule, B. Thiebaut and J. Vysny. 1997. Genetic variation of beech 

(Fagus sylvatica L.). Pp. 61-68 in Medzinarodnavedecki konferencia Les - Drevo - Zivotne 

prostredie '97. Sekcia 1: Ekol6gia lesa a jeho integrovana ochrana (E. Krizova and J. 

Kodrik, eds.). Technicka univerzita, Zvolen. 

Gomory, D., V. Hynek and L. Paule. 1998. Delineation of seed zones for European beech (Fagus 

sylvatica L.) in the Czech Republic based on isozyme gene markers. Ann. Sci. For. (in press). 

Gomory, D., J. Vysny, L. Paule and B. Comps. 1992. Genetic structure of European beech 

(Fagus sylvatica L.) populations in Czecho-Slovakia. Pp. 27-32 in Fytotechnika a 

hospodarska uprava lesov v sucasnych podmienkach. Technicka univerzita, Zvolen. 

HanCinsky, L. 1972. Lesne typy Slovenska [Forest types of Slovakia]. Prfroda, Bratislava. 

Korenek, J. 1980. [Results of oak provenance trials]. In Proveniencni vyzkum lesnich drevin. 

vULHM, Jiloviste-Strnady. 

Korpel', S. 1995. Die Urwiilder der Westkarpaten. Gustav Fischer Verlag, Stuttgart. 

Magic, D. 1974. [Additional species of oak in the forests of Slovakia]. Les 30(6):244-252. 

Magic, D. 1975. [Taxonomic notes from research on oaks in the Western Carpathians]. 

Biologia (Bratislava) 30(1):65-74. 

Ostrolucka, M.G. and M. BolvanskY. 1992. Umela hybridizacia druhov rodu Quercus 

[Artificial hybridization of Quercus species]. Lesnicky casopis 38(3):239-251. 

Paule, L. and D. Gomory. 1993. Significance of forest reserves for studies of the population 

and evolution genetics. Pp. 153-157 in European Forest Reserves (M.E.A. Broekmeyer, W. 

Vos and H. Koop, eds.). Pudoc Scientific Publishers, Wageningen. 

Paule, L. 1982. Untersuchungen zum Wachstum der slowakischen Provenienzen der 

Rotbuche (Fagus sylvatica L.). Silvae Genet. 31(4):131-136. 

Paule, L. 1994. Biodiversity of the Western Carpathians' forest ecosystems. Pp. 31-38 in Conservation 

of Forests in Central Europe (J. Paulenka and L. Paule, eds.). Arbora Publishers, Zvolen. 

Paule, L. 1995. Conservation of Norway spruce genetic resources in Slovakia. Pp. 51-58 in 

EUFORGEN. Picea abies Network. Report of the first meeting, 16-18 March 1995, Stara 

Lesna, Slovakia (J. Turok, V. Koski, L. Paule and E. Frison, compilers). IPGRI Rome, Italy. 

Paule, L. 1995. Gene conservation in European beech (Fagus sylvatica L.). For. Genet. 2(3):161-170. 

Paule, L., D. Gomory and J. Vysny. 1995. Genetic diversity and differentiation of beech 

populations in Eastern Europe. Pp. 159-167 in Genetics and Silviculture of Beech. Proc. of 

the 5th IUFRO beech symposium 1994, Denmark (S. Madsen, ed.). Forskningsserien no. 

11-1995. Danish Forest and Landscape Institute, H0rsholm, Denmark. 

Voloscuk,1. 1993. The virgin forests and reserves in Slovakia. Pp. 69-74 in European Forest 

Reserves (M.E.A. Broekmeyer, W. Vos and H. Koop, eds.). Pudoc Scientific Publishers, 

Wageningen.

34 EUFeRGEN: SeGIA~ BReAD~EAVES 

Social Broadleaves in the Czech Republic 

Vladimir Hynek 

Forestry and Game Management Research Institute, Praha Zbraslav, Czech Republic 

Introduction 

Forest land covers about 2 630 000 ha in the Czech Republic. The composition of forest tree 

species was considerably changed in the past two and a half centuries of intensive forest 

management. Plantation of coniferous tree species was recommended since the 18th century. 

The natural species composition is as follows: beech (about 40%), oaks (about 18%), fir 

(16%), spruce (15%) and pine (3%). The present species composition is different. The 

proportions of spruce (55%) and pine (18%) are higher. Oaks (6%), beech (5.6%) and fir 

«1%) are underrepresented with regard to the original situation. Most of the beech stands 

were harvested and used in glass manufactories and for charcoal production. Mixed beech 

stands were replaced by Norway spruce monocultures and oak stands by pure plantations 

of pine. 

The territory of the Czech Republic is divided into 41 Natural Forest Regions (NFRs, see 

Fig. 1 and Table I), delimited by geographic, geomorphological and climatic conditions. 

Ecological conditions affect the representation and rise of regional populations which are 

adapted to local conditions. 

Within these NFRs, populations are merged into seed zones. The tree seed zones are 

proposed for the following reasons: 

• current regional populations should not be mixed with regard to the use of forest 

reproductive material 

• there is insufficient knowledge on their variability (in contrast to coniferous species - 

spruce, pine and larch - which are better known), and their contamination by other, 

not well-adapted populations or species should be avoided. 

Unfortunately, it seems that some representatives of the forestry practice prefer not to 

have seed zones, with the possibility to handle reproductive material arbitrarily. 

Beech grows naturally in all 41 NFRs, oaks grow naturally in 37 NFRs. 

Occurrence and origin of oaks and beech in the Czech Republic 

In the Czech Republic there are eight forest vegetation belts (FVB) (Table 2). Pedunculate 

oak occurs naturally from FVB 1 to 3, and up to FVB 5 if artificially planted. Sessile oak 

occurs naturally from FVB 1 to 4. Beech occurs naturally from FVB 2 to 7. 

Pedunculate oak stands survived, especially on humid sites. Sessile oak survived on 

extremely shallow soil as coppice forest. Beech stands survived in extremely steep areas, 

where it was impossible to carry out artificial regeneration with spruce. In these localities 

beech regenerates in a natural way. 

Pedunculate oak was artificially distributed, for example, around dams and ponds 

(especially in southern Bohemia and southern Moravia). Many stands with pedunculate oak 

are situated in floodplain forest after stream regulation. Important were imports of 

pedunculate oak from Croatia, specially from Slavonia. Populations of oaks from Slavonia 

are more productive and have better wood quality than local oaks. 

Sessile oak was not regenerated artificially as well as the pedunculate oak. Generally in 

the past foresters did not respect site requirements of these two species. In oak forest stands, 

both species often mixed. 

Artificial regeneration of beech started at the beginning of the 20th century. In the last 40 

years, beech reproductive material from Slovakia and western Ukraine has been used more 

than beech of local origin.

,. COUNmR¥ REBORmS 35 

Fig. 1. Natural forest regions of the Czech Republic. 


Oak wood is used in the furniture industry, and beech wood is used for example for railway 

sleepers and in the building industry. Spruce and pine are still more important 

economically. It depends on the level of wood industry. According to the new Forest Law 

from 1995, a minimum of 30% of 'melioration' species has to be used, which increases the 

resistance of newly established stands. Oaks and beech are often used as the 'melioration' 

species. 

Silvicultural approaches 

Mixed forest stands which contain native broadleaved species such as oaks and beech are 

more stable and resistant to air pollution. Air pollution in the Czech Republic is one of the 

highest in all Europec;tn countries. Most of the forest stands ar'e artificial with a majority of 

monoculture of spruce and pine. An increase in the percentage of broadleaved tree species 

increases the stability and resistance to other abiotic and biotic factors. 

Therefore, the current forest law prescribes the raising of the proportion of broadleaved 

species. 


Oaks 

At the moment oaks are the second most endangered genus in the Czech Republic after 

elms. This situation is a result of the tracheomycosis disease and repeated damages by an 

oak leaf roller moth (Tortrix viridana L.). In forest stands without chemical protection no 

acorn has been found for several years. The seed crop is not a problem nationally but 

regionally. To conform with the new Forest Law (30% of 'melioration' species), practical 

forestry needs to import reproductive material from foreign countries. This practice 

decreases the native oaks genepool in the Czech Republic and also contaminates with 

untested genetic material. From all the imported oaks material, Slavonic oak (pedunculate 

oak) has good practical results, but the imports from Croatia are practically nonexistent.

Table 1. Forest veqetation zones in natural forest belts reqions of the Czech Republic 

FVZ 2a/b I 3 4 5 6 7 I 8a/b I 9 I 10 I 11 I 12 I 13 I 15 I 16 I 17 I 18a/b I 19 I 20 I 21 I 22 I 23 I 24 I 25 I 26 I 27 I 28 I 29 I 30 I 31 I 32 I 33 I 34 I 35 I 36 I 37 I 38 I 39 I 40 I 41 

9 I 0.9 0.2 0.9 0.2 r 10.3 1.3 

8 I 9.9 I I 0.5 0.2 I 112~5 12.9.125.5 I + I I 2 12.0 0.1 

7 128.91 + 11.1 ... .7 0.1 3.7 I + 126:9 1.0 2.9 10.9119,31 + 10.1 116& 2.1;81 0.7 + + 1.9 

6 127~21 1.0 1.21 .. 0 0.1 I 0.2 Il1t6 '30,81 6.2 1.55.8 23.6 22 .. 9118:4141 :3.141.31 4.1 1.1}:2148.91 0.3 13SC8!.1U I 9.6 I 0.7 I 1.0 8.4 I 0.4 

5 L27.9114.0 15$:8128.41 6.5 2.4 137.3 1.5 164.0 j 65:71 3.5 4.3 163.7 24.5J52.8126.81 3.6 1.43.8173~O 1 32.5 !3!1:8 1.27.4 !54;8137:S I. 14:5131.8 0.1 8.4 1.1 1.87.91·70.0 

4 0.1 8.2 0.6 1~.71 5.0 I 9.2 117.6 3.1 121.21 0.3 1.13~8 38.8.1 7.0 I 0.2 0.7 I 0.2 I I 0.9 I 0.8 I I 2.1 I I 7.7 I 9.9 127.2133:3 29;4 7.3 .20:7115.91 0.2 8.7 

3 4.6 I 6.6 1l6,7147;8149.91~3.5J 35.814~.~1 0.6 1.12.8 17,81 4.0 2.2 I 9.9 12:4.01 7.6 42.11 3.9 0.3 148:71 0.6 123.7139JI140.3127;7 i.9M 127:1 0.3 155.$ 16!t715.9.~Jt2.71 1.5 119:4 

2 0.1 0.4 I 3.6 127.6 j 20.41 1.5 I 47.0123.5 1.2 5.6 1 0.3 22 + 0.2 + 6.1 10~1 1.2 3.2 114.8 1 4.7 7.5 128.5.126.51 7.8 133.6130.3119;71 3.2 1.5 

0.1 I 0.2 I 1.4 111.71 1.8 I 0.3 I I 2.8 3.6 0.4 0.1 1.6 0.1 173.81 I 0.1 0.6 0.3 + 3.0 0.3 0.1 2.3 0.6 1.9 I 14::H1:tSI9t.91 3.3 0.7 7.1 + 

o 0.3 3.0 I 0.2 1.3 116.1 I 0.1 11} I 0.4 + 0.2 0.4 131.90.1 0.3 1.8 0.3 2.7 5.0 + + 0.1 + 0.2 0.9 0.7 + + +

COUNiliR¥ REBORiliS 

3'7: 

iliable 2. Characteristics of the forest vegetation belts in the Czech Re~ublic 

% of total Altitude Mean annual Annual Vegetation period 

Vegetation belt forest area {m as\} tem~. re} ~reci~. {mm} {da~s} 

O. Pine 3.73 

1. Oak S.31 S.O 165 

2. Beech-oak 14.S9 350-400 7.5-S.5 600-650 160-165 

3. Oak-beech 1S.41 400-550 6.5-7.5 650-700 150-160 

4. Beech 5.69 550-600 6.0-6.5 700-S00 140-150 

5. Fir-beech 30.04 600-700 5.5-6.0 SOO-900 130-140 

6. Spruce-beech 11.95 700-900 4.5-5.5 900-1050 115-130 

7. Beech-spruce 5.00 900-1050 4.0-4.5 1050-1200 100-115 

S. Spruce 1.69 1050-1350 2.5-4.0 1200-1500 60-100 

9. Mountain (2ine 0.29 >1350 1500

38 EU~eRGEN: SeGI~~ BRe~D~E~VES 

Fig. 2. Map of tree seed zones proposed for Fagus sy/vatica L. 

Fig. 3. Map of tree seed zones proposed for Quercus petraea. 

Fig. 4. Map of tree seed zones proposed for Quercus robur.

- G0llJNmBM BEB0SmS 39 


In situ 

Passive gene conservation of oaks and beech populations in situ takes the form of state 

reserves. The gene reserves (bases) are considered as an active way of gene conservation 

and reproduction. Gene reserves are groups of stands with a minimum surface area of 

100 ha of forest land. Regeneration is usually natural. If natural regeneration is not 

successful, it is possible to use artificial reproductive material but only with origins from 

these gene reserves. Suggestions for management methods in gene reserves are elaborated 

by the Institute. 

Ex situ 

The most important activities on ex situ gene conservation are grafting and establishment of 

clonal archives and seed orchards. This method is not used often for oak propagation, 

because of incompatibility problems. Only one seed orchard has been established and no 

others are planned. This method of conservation is more important for beech. We 

established clonal archives of beech from the seven forest vegetation belts only in the most 

polluted areas (Ore Mountains). Another method of ex situ conservation is the establishment 

of special plantations for obtaining secondary cuttings. Genebanks are used within the 

existing seed banks and tissue culture banks. 


In 1992, a Law on Nature Protection was issued, and later in 1995 a new Forest Law (No. 

289). Both aim to conserve the remaining natural forest tree genepool. 

nee-improvement activities 

Tree-improvement activities are carried out on the basis of provenance tests. Practical 

implications of the provenance tests include rules for effective transfer of reproductive 

material, delimitation of seed zones, preparation and implementation of breeding 

programmes including different methods of vegetative reproduction (grafting, cuttings and 

in vitro methods). 


In the Czech Republic it is allowed to use reproductive material of spruce, pine and larch 

only from approved sources (approved seed stands, seed orchards, clonal archives). For all 

forest trees (including oaks and beech) a rule regarding the limitation of vertical transport of 

reproductive material is in force. This limit is plus or minus one forest vegetation belt (FVB). 

Beech origins from the seven FVBs are suggested as special climatic ecotypes. 

The proposed seed zones for oaks and beech (also for beech originating from the seven 

FVBs) have not been approved yet. 

Import of reproductive material of all forest tree species (including oaks and beech) from 

foreign countries is regulated by the Ministry of Agriculture. The Ministry also defined the 

areas where this reproductive material can be used. 


Most of the work is done by the Forestry and Game Management Research Institute. The 

central seed bank is managed by the state forest company and situated in the town TyniSten, 

Orlici. Both Forestry faculties (University Bmo and University Prague), the Academy of 

Science and the other state and private institutes and organizations are concerned with the 

problems of oaks and beech genetic resources to a limited extent. The Ministry of 

Environment and its organizations are interested in the conservation of oaks and beech in 

state nature reserves and national parks.

40 EUFORGEN: SOCIAL. BROADL.EAVES 


In the Forestry and Game Management Research Institute there are only four scientists 

involved in research on oaks and beech (breeding, vegetative propagation, breeding 

programmes concerned with conservation and reproduction of the genepool). 

The main priority is to conserve the remaining natural and good-quality populations of 

pedunculate and sessile oaks and beech by means of natural regeneration. 


The proportion of oaks should increase from 6 to 9% and beech from 5.5 to 12-18%. 

Considering these goals, the local sources of reproductive material will not be sufficient. 

Testing of new provenances, establishment of new series of provenance tests for both oak 

species are very important to increase the variability of newly established forest stands. In 

the case of beech a sufficient number of provenance tests in the framework of the 

international series (von Wuehlisch et al., this volume) will be established. 

To collect enough information on the genetic variability of populations of forest trees, 

international projects concerned with DNA and isoenzyme studies should be carried out. A 

laboratory in the Forestry and Game Management Research Institute has just been 

established, and accordingly started work. 

Selection on resistance to tracheomycosis and Tortrix viridana L. would also be interesting.

e0UNTR¥ REF!0RTS 41 

Conservation of genetic resources of oaks and beech in Austria 

Thomas Geburek 

Institute of Forest Genetics, Federal Forestry Research Centre, Vienna, Austria 

Introduction 

Austria is densely forested, with 46.2% of its surface covered by forests. Owing to the 

predominantly mountainous terrain, the proportion of conifers is high (77.3%), whereas 

broadleaved tree species and shrubs cover 22.7% of the total land (3 877000 ha) (BMLF 

1995). Since more than two-thirds of the forest area is located in the Alps, besides their 

economic importance forests play a very important role in the protection from erosion, 

torrents and avalanches. These reasons led the Federal Forestry Research Centre (Vienna) to 

start a national programme on the conservation of genetic diversity in forests. Launched in 

1986 and implemented through cooperative actions of the Institute of Silviculture and the 

Institute of Forest Genetics, the programme aims at the conservation of the genetic 

adaptability of forest tree populations. 

Distribution of beech and oaks 

Beech is the second most widespread forest tree species in Austria (9.8%), while oaks 

represent only 2.2%. A meticulous description of the natural range of Fagus sylvatica was 

published in the late 1920s by Tschermak (1929). The following remarks are based on his 

work and on the Austrian Forest Inventory (Schadauer 1994). 

With the exception of major parts of the Central Alps, beech is found in the other areas of 

Austria, although at highly variable densities. In the Weinviertel, northern parts of the 

Waldviertel in Lower Austria, and in the Miihlviertel, beech is scarce. Further west and 

south of the river Danube, beech is more frequent in the Hausruck and the 

Kobernausserwald. In the Northern Limestone Alps, it is distributed from the 

Bregenzerwald close to the Bodensee to the northeastern edge of the Wienerwald. 

In the Allgauer Alps and in the northern parts of the Tyrolean Alps, beech is commonly 

found in mixed stands. In the province of Salzburg, mixed beech forests are typical on slaty 

sites and at high altitudes of the Northern Limestone Alps. Widespread pure stands are 

common in the foothills of the Salzkammergut. In the Northern Limestone Alps of Upper 

and Lower Austria, extensive pure beech stands are also frequently found, especially on 

sediments of the Cretaceous and Tertiary period. Pure Fagus sylvatica stands are also typical 

of the Wienerwald, whereas foothills of the eastern Alps (Northern Limestone and Central 

Alps) are only partly covered by beech in mixed forests. Disjunct beech stands are further 

found in the hills of the Geschriebestein close to the Neusiedler Lake. 

Beech is also found in the Southern Limestone Alps, namely in the Karawanken, the 

Carnic Alps, and the Gailtaler Alps. However, pure stands are more common in the 

Karawanken than in the rest of the Southern Limestone Alps. As a scattered forest tree 

species it is indigenous at the southern foothills of the Central Alps, in the Lavantal at the 

eastern edge of Carinthia, and on elevations of the Klagenfurter Becken. In the Central Alps 

beech is nearly completely absent (see also Tschermak 1958). There is a good coincidence of 

the natural range of beech with the occurrence of limestone (Geburek and Thurner 1993). 

The altitudinal range varies from 170 to 1700 m. A mixed beech stand is located at 170 m 

close to the river Danube, at Greifenstein in Lower Austria. Pure stands are found in this 

area above 225 m. At the Gapfahler Falben in the province of Vorarlberg, shrubby forms 

were found close to 1700 m. The upper limit within the Northern Limestone Alps rises from 

east to west. Within the Southern Limestone Alps, the upper tree line of beech is generally 

higher than in the northern Alpine range.

42 EUE0RGEN: S0GI~t:; BR0~[)t:;E~"ES 

Quercus petraea and Q. robur are found on (moderately) fresh mollisol and interceptisol 

types. Both species most commonly occur in mixed forest stands in the Weinviertel, 

Marchfeld, Wiener Becken, Burgenland and Oststeirisches Hiigelland. In the northeastern 

part of its range the climate is Pannonian-subcontinentally influenced with high summer 

temperatures and low annual precipitation ranging from 450 mm in the north to 700 mm in 

the south. In the southeastern part precipitation is much higher. At lower elevations up to 

350 m Quercetum petraea-cerris, pure Quercus pubescens stands on sunny, dry and calcareous 

sites or the Quercus petraea-Carpinus betulus forest type are typical, while at higher elevations 

up to 500 m'Quercus petraea-robur is associated with beech and hornbeam. Scattered stands 

are also found in the foothills of the Northern and Southern Limestone Alps (e.g. 

Klagenfurter Becken). In this region Quercus is found in the Quercus robur-Carpinus betulus 

forest type at low elevations or in association with beech at the submontane level (Kilian et 

al. 1994). 

Economic importance of beech and oaks 

Beech is the most important hardwood economically and yields approximately 1 600 000 

m 3 /year. Less than 2% of the wood is of veneer quality. On average 75 ECU 1m 3 can be 

expected for beech timber, while only 30 ECU I m 3 are paid on average for poor-quality wood 

by the pulp and paper industry. In the past, beech forests have been mainly harvested for 

fuel wood. Low profits and low yield of beech in comparison with conifers have caused 

conversion of beech forest into coniferous stands. More recently, the demand for fuel wood 

and beech lumber has increased and consciousness of a close-ta-nature forest management is 

more pronounced. This has commonly resulted in new planting of beech forests. Forest 

enterprises based on beech logging face a fairly stable economic situation, while returns of 

softwood have been diminishing in the last decade. Oak species account for approximately 

450000 m 3 (2.3% of the total). On average 100 ECU/m 3 can be expected for oak timber, 

whereas prices for veneer quality are much higher (1000-5000 ECU Im} 

Silviculture 

Oaks and beech are mainly managed in high forests. Beech is predominantly regenerated 

naturally in different shelterwood systems. Oaks are also managed in high forest, but the 

coppice with standards system still plays an important role locally. Oaks are regenerated 

artificially more often than beech. 

Reproductive material 

In beech, 760000 plants are produced annually and 470000 plants are imported. This 

accounts for 1.3% of the 57 million plants produced and 23% of the total import (some 

2 million plants). In oaks, 1.1 million plants are produced per year and 350000 plants, i.e. 

18%, are imported. Thus, domestic plant production is low, while Austrian import of plants 

of both genera is fairly high (roughly 40% of the total plant imports). According to the 

Austrian Forest Reproductive Material Act (Anonymous 1996), beech and oak seeds have to 

be harvested in selected stands. In line with this regulation, some 840 ha of beech stands, 

231 ha of sessile oak and 238 ha of pedunculate oak have been selected and registered. 

Health and threats 

Early loss of leaves and the resulting crown sparseness indicate a state of individual 

damage. Usually the health status of beech is much better than that of oaks in Austria. Less 

than 1% of the beech trees and more than 4% of oaks monitored exceed 60% defoliation. 

Beech accounts for approximately 4% of the trees in the defoliation class 25-60%, oaks for 

more than 16%. More than 50% of beech trees and 35% of oaks are (nearly) unaffected. 

Damage by browsing of game and livestock is negligible for both genera.

. CQUNlTS¥ REPQRlTS 43 

Gene conservation 

Genetic resources of both genera are mainly conserved by in situ measures, to secure the 

unknown genetic variability, guarantee evolutionary development, and thus conserve 

adaptability to changing environmental conditions. Since Austrian virgin forests are low in 

numbers and small in size, conservation is practised in managed forests which are 

integrated into the economic cycle (Muller 1996). In situ management plans are developed 

that comprise gene reserves, each covering a minimum area of 30 ha. This size is assumed to 

be large enough to conserve the genetic structure and to allow an evolutionary change of the 

resource population. As pollen contamination from outside sources is unwanted, the core 

area ought to be framed by a 300-500 m wide buffering zone. A suitable forest stand is 

declared an in situ gene conservation stand by a voluntary agreement of the owner. 

Unfortunately the Austrian forest ownership structure is unfavourable to declaration of 

genetic resources. Of the total forest area, 36.3% is cultivated by small forest enterprises 

(productive area under 50 ha). There is no means to declare forest genetic resources by law. 

Since beech and oaks are often found in mixed forest types, preference is laid on so-called 

large-scale gene reserves that exceed 30 ha in size to ensure a sufficient effective population 

size (see Table 1). 

The pragmatic approach chosen is based on the assumption that different site conditions 

determine the forest communities and shape genetic structures of the forest tree species. The 

following main criteria are considered for declaration of a gene conservation stand: 

• representative of a natural forest community 

• conform to nature with regard to the plant community 

• autochthonous or at least well adapted (high vitality, no damage by biotic and/or 

abiotic factors, economically important phenotypical traits (e.g. straight stem forms) 

not regarded) 

• naturally regenerated, at least potentially. 

Silvicultural measures are taken in the gene conservation stands. However, forest 

managers must aim to achieve a sustainable balance by permanent stocking, all-aged stand 

structure, heterogeneous stand development, long natural regeneration with overlapping 

reproductive periods, and self-differentiation processes during all growth phases (for more 

details see Muller 1996). 

Breeding 

The Institute of Forest Genetics of the Federal Forestry Research Centre currently contributes 

to the International Beech Provenance Trial (see von Wuehlisch et al., this volume), but there 

are no breeding activities in a strict sense for either Quercus or Fagus. 

Table 1. Large-scale gene reserves (>30 ha) with beech and oaks in Austria 

Beech 

Oak 

Province 

Burgenland 

Carinthia 

Lower Austria 

Salzburg 

Styria 

Tyrol 

Vorarlberg 

lTotal 

No. 

2 

15 

7 

1 

5 

6 

1 

37 

Average size (ha) 

32.6 

66.3 

48.5 

35.0 

43.9 

170.0 

52.6 

No. 

2 

Average size (ha) 

103.0 

59.8

44 EUEORGEN: SOCIAL BROADLEAMES 

Research and capacities 

Three institutions conduct genetic studies on beech and oak in Austria: 

• the Austrian Research Centre, Seibersdorf 

• the Centre of Applied Genetics of the University of Agriculture, Vienna 

• the Federal Forestry Research Centre. 

At present these institutions perform provenance, isoenzyme and DNA marker studies 

(e.g. Steinkellner et al. 1997; Comps et al. 1998). 

Institutions 

The Federal Forestry Research Centre with its Institute of Forest Genetics and Institute of Silviculture 

acts to put forest gene conservation into action. However, the cooperative programme 

to conserve forest genetic resources relies on the support of forest owners and managers. 

Needs and perspectives 

Gaps in the knowledge related to the conservation of genetic resources of beech and oaks are 

as follows: 

• within- and among-population variation for morphological and genetic traits 

• links between adaptive traits and genetic markers 

• cryopreservation 

• Multiple Population Breeding System (MPBS) vs. unmanaged in situ populations 

• required number and size of in situ populations 

• silviculturally managed vs. unmanaged in situ populations. 

Answers to the above-mentioned questions would strongly contribute to the national 

conservation strategy. 

References 

Anonymous. 1996. Bundesgesetzblatt fur die Republik Osterreich, Jahrgang 1996:419. 

Bundesgestz uber forstliches Vermehrungsut (Forstliches Vermehrungs-gutgesetz), 

Bundesgesetz, mit dem das Forstgesetz 1975 geandert wird, und Bundesgesetz, mit dem 

das Dungemittelgesetz 1994 geandert wird [Austrian Forest Reproductive Material Act] 

BMLF (Bundesministerium fur Land- und Forstwirtschaft). 1995. Osterreichischer 

Waldbericht. 

Comps, B., Cs. MMyas, J. Letouzey and T. Geburek. 1998. Genetic variation in beech 


Genet. 5(1):1-9. 

Geburek, Th. and G. Thurner. 1993. Current status of common beech (Fagus sylvatica L.) in 

Austria. Pp. 27-38 in The Scientific Basis for the Evaluation of Forest Genetic Resources of 

Beech. Proceedings of an EC Workshop, Ahrensburg (H.-J. Muhs and G. von Wuehlisch, 

eds.). Working Document of the EC, DG VI, Brussels. 

Kilian, W., F. Muller and F. Starlinger. 1994. Die forstlichen Wuchsgebiete Osterreichs. 

FBV A-Berichte 82. 

Muller, F. 1996. Ausscheidung und waldbauliche Behandlung von Genreservaten in 

Osterreich. Pp. 330-340 in Biodiversitat und nachhaltige Forstwirtschaft (G. Muller-Starck, 

ed.). Ecomed, Landsberg. 

Schadauer, K. 1994. Baumartenatlas fUr Osterreich. Die Verbreitung der Baumarten nach 

Daten der Osterreichischen waldinventur. FBVA-Berichte 76. 

Steinkellner, H., S. Fluch, E. Turetschek, C. Lexer, R. Streiff, A. Kremer, K. Burg and J. Glossl. 

1997. Identification and characterization of microsatellite loci from Quercus petraea. Plant 

Mol. BioI. 33:1093-1096. 

Tschermak, L. 1929. Die Verbreitung der Rotbuche in Osterreich. Einiges uber die fur die 

Verbreitung der Rotbuche maBgebenden Standorsfaktoren. Schweiz. Z. Forstw. 

Tschermak, L. 1958. Das Fehlen der Buche (Fagus sylvatica L.) in den Innenalpen. CbI. ges. 

Forstwesen 75:208-223.

Conservation of Social Broadleaves genetic resources in Switzerland 

Patrick Bonfils 

Swiss Federal Institute for Forest, Snow and Landscape Research, Birmensdorf, Switzerland 

Occurrence and origin of beech and oaks in Switzerland 

Switzerland is commonly divided into five different geographic regions: the Jura Mountains, 

the Central Plateau, the Lower Alps, the Alps and the Southern Alps (Fig. 1). These regions 

are characterized by specific ecological conditions. Overall, a great variety of different site 

types can be found, ranging from basic to acidsoils and from atlantic to continental climates. 

Fig. 1. The five geographic regions of Switzerland: Jura Mountains, Central Plateau, Lower Alps, Alps 

and Southern Alps. 

Beech 

Fagus sylvatica is mainly found in the Jura Mountains and the Central Plateau. It is not 

present in the dry and continental climates of the Alpine region (Fig. 2). In the subatlantic 

climate regions of Switzerland, beech forests form a broad vegetation belt between the 

subalpine vegetation zone dominated by Norway spruce and the lowland vegetation zone 

which is characterized by mixed deciduous forests. Beech occurs in almost all forest 

communities except those from subalpine mountain forests (Leibundgut 1984). Beech 

appears commonly as a species of mixed forests and only in 20% of the stands is its 

abundance higher than 66%. It is often associated with Norway spruce and silver fir, but 

also with sycamore maple, ash, oak and Scots pine (Brandli 1996). Fresh deep fertile soils, 

high humidity and high precipitation (>750 mm/year) are beneficial to beech (ETH-Zurich 

1993). Beech requires a cooler climate than oak but a slightly warmer one than silver fir 

(Leibundgut 1984). 

In the past, forest stands were considerably affected by human activities. Destruction of 

beech forests started in Roman times and continued throughout the Middle Ages. A 

significant proportion of beech stands disappeared as a result of clearing. Large areas were

46 EUFORGEN: SOCIAL BROADLEAVES . 

converted to agricultural land, especially in the Central Plateau. Today this region is the 

most populated part of Switzerland. Remaining beech forests were often replaced by 

Norway spruce, which is not native to the Central Plateau. 

Oaks 

Four different oak species occur in Switzerland: pedunculate oak (Q. robur L.), sessile oak 

(Q. petraea (Matt.) Liebl.), pubescent oak (Q. pubescens Willd.) and Turkey oak (Q. cerris L.). This 

report focuses on the two most abundant oak species, Q. petraea and Q. robur. 


The main range of pedunculate oak is in the Central Plateau and the eastern Jura Mountains. 

It is abundant in the western part of the country. It is rarely found or is even absent in areas 

of the more continental alpine regions. Pedunculate oak is most frequently found in forests 

below 400 m whereas solitary trees are encountered up to 1400 m in the upper montane 

vegetation zone (Brandli 1996). Although it has no specific demands on soil, it is often 

found on heavy clay and loam soils with high nutrient content and high water supply. 

Pedunculate oak is moderately heat-demanding during summer and sensitive to winter cold 

(less than sessile oak). 

Sessile oak 

Sessile oak is common and widespread in the Central Plateau, the Jura Mountains and the 

Southern Alps. It is most frequently found in the western part of the Central Plateau. Threequarters 

of the sessile oaks grow in the lowland and submontane vegetation zones. Its main 

range is 400 to 600 m. Solitary trees are found above 1400 m (Brandli 1996). The demands 

for site conditions are different from those of pedunculate oak. Sessile oak is less nutrientdemanding 

and grows also on drier sites (Schweingruber 1990). On heavy soils it is replaced 

by pedunculate oak. Its demand for summer heat as well as its sensitivity to winter cold are 

higher than that of pedunculate oak (ETH-Ziirich 1993). 

Sessile oak is estimated to be twice as abundant as pedunculate oak. Pedunculate oak 

occurs mainly in hardwood riparian forests and mixed ash woodlands, whereas sessile oak 

dominates on very dry sites in acidophilous mixed oak forests. In a great number of forest 

communities (especially beech forests) both oak species are present in varying proportions. 

The most common associated species is beech. Norway spruce, silver fir, ash and sycamore 

maple are also often found in mixture with oak, owing to the impact of forest management. 

Pure oak forests (approximately 12000 ha) are found only on 12% of the oak area, either on 

very dry sites or as artificial plantations (Brandli 1996). Natural oak forests are not found on 

fertile sites (Leibundgut 1984). 

In the past, sessile and pedunculate oaks used to be more common in the Central Plateau 

and the Jura Mountains. In the former coppice with standards system, oaks were not only 

valued for their wood production but also for the acorns which were used for pig feed. The 

importance of oak species decreased with the introduction of potato starting in 1740. Oaks 

became less attractive than other species (e.g. Norway spruce) owing to the transformation 

of the coppice with standards into high forests. Since 1850 the construction and 

maintenance of the railway also devoured huge amounts of oak wood (Brandli 1996).

COUNTRY REPORTS 47 

M. 

1500 

1000 

500 

J 

CP P-A A SA 

1500 

1000 

500 

J 

CP P-A A SA 

100 

1500 

~ 

~ 90 

0 

Qj 1000 

i ~ 

jIIIIIIIIIIj 

.,.... 

i 

50 

rJ'J 

i 

., 

rJ'J 

500 : 

Qj 

00 

J 

CP P-A A SA 

Fig. 2. Altitudinal and geographic distribution of beech, pedunculate oak and sessile oak.

48 EU60RGEN: S0GIJ.\U BR0J.\[)UEJ.\XlES v 

Growing stock 

Beech is the most important broadleaved tree species in terms of growing stock 

(approximately 59 million m 3 ) and after Norway spruce (179 million m 3 ) the second most 

important species in total (Table 1). Both oak species together represent approximately 2% 

of the total growing stock in Switzerland (EAFV 1988). 

Table 1. Percentage of the total growing stock of beech and oak in different regions of 

Switzerland (EAFV 1988} 

Central Uower- Southern 

S~ecies Jura Plateau AI~s AI~s AI~s Switzerland 

Fagus sy/vatica 30.0 20.4 13.3 6.6 13.1 16.2 

Picea abies 31.2 42.9 57.3 62.6 35.2 49.1 

Quercus petraea 2.0 2.3 0.1 0.2 1.7· 1.1 

Quercus rabur 1.2 2.3 0.2 0.1 0.5 0.9 

Others 35.6 32.1 29.1 30.5 49.5 32.7 

Total in 1000 m 3 63574 92785 88139 97481 23148 365128 

Ice age refugia 

Fossil pollen maps of Europe indicate that oak survived the last maximum glaciation in 

refugia in southeastern Europe (Greece/Turkey), southern Europe (Italy) and southwestern 

Europe (Iberian Peninsula) (Huntley and Birks 1983). Analyses of genetic variation in 

chloroplast DNA (cpDNA) suggest that oaks recolonized Switzerland from at least two 

refugia (Dumolin-Lapegue et al. 1997; Ferris et al. 1997). 

Beech populations north of the Alps are likely to originate from southeastern Europe 

(Balkan). Beech from an additional refugium in southern Italy may have expanded to south 

of the Alps (Burga and Perret 1998). 


The economic value of oaks and beech remains limited because of their low abundance. The 

demand of Swiss non-coniferous wood for paper and cellulose industry has been stagnant 

for 20 years (Anonymous 1997). The use of steel, concrete and synthetic products is often 

preferred in construction and interior design industries. For railway construction, wooden 

sleepers have also been replaced by concrete and steel. 

Beech and oaks, however, can be locally important especially for the production of 

veneer. During the last eight years, the average selling price for beech stem wood increased 

by approximately 6% (Anonymous 1997). 


Regeneration 

Swiss forestry has a long tradition of close-to-nature silviculture and thus of enhancing 

natural regeneration. Almost 90% of the beech forests are regenerated naturally (Table 2) 

without major problems. Beech can be regenerated in small patches because of its shade 

tolerance. In association with Norway spruce and silver fir, beech is an important 

component in selection system forests. In other silvicultural systems it is usually 

regenerated with the shelterwood or strip-shelterwood method. 

Artificial regeneration is much more important for oaks than for beech. Especially 

pedunculate oak is often planted as shown in Table 2. Both oak species are rarely sown. 

Natural regeneration is carried out in the shelterwood system and has to be fenced to avoid 

damage caused by game. The area of regeneration is usually at least 0.3 ha to minimize the

00l:JNlliB¥ BER0BlliS 49 

lliable 2. Mode of regeneration used in Switzerland for beech and oak (Brandli 1996) 

Regeneration (%) 

Natural Natural and artificial Artificial No information 

Beech 

85.5 

6.1 1.5 

6.9 

Pedunculate oak 54.8 

18.4 17.3 

9.5 

Sessile oak 70.7 

4.9 4.1 

20.3 

number of bent growing trees due to phototropic reaction (especially for Q. robur). Forest 

tending at the'thicket and the early pole stages aims to homogenize quality and density of 

the trees. The growth in pure stands is considered to be an essential prerequisite for the 

production of good-quality wood. The cultivation of oaks is quite difficult and costintensive. 

Foresters are currently evaluating alternative silvicultural systems (e.g, planting 

in final distance - wide spacing) in order to lower costs. 

Exploitation 

The rotation period for beech is approximately 120 years, whereas oak is harvested after 120- 

200 years for sawn timber and after 240 years for veneer. 

Health condition of the forest stands 

Forest condition 

In Switzerland a systematic monitoring of forest condition has been carried out since 1984 by 

evaluating the defoliation of crowns. The long-term trend suggests that the proportion of 

trees with unexplained defoliation higher than 25% is increasing (Fig. 3). Oak species seem 

to be more affected than beech (BUW ALjWSL 1992) . 

..c: .... 

.§: ~ 

Ul 0 

a>LO 

a> C\I 

.!:: /\ 

-o C 0 

a>+= 

Cl ,!!:! 

S '0 1 

c 

a> a> 

0"0 

"- 

a> 

0. 

l!) 

0) 

en 

~ 

50 EUFORGEN: SOCIAL BROADLEAVES 

Tree diseases 

The Swiss Federal Institute for Forest, Snow and Landscape Research (WSL) publishes a 

periodical bulletin of relevant forest pathology problems based on a survey of all forest 

districts (Meier et al. 1996). In 1995 the following observations were made. 

Oaks 

Three different kinds of damage were observed in oaks. 

First, in the development stages from saplings up to pole stage, signs of dieback of the 

crown were observed. The cause for this phenomenon was infestation by different cortical 

fungi. These damages were without exception caused by secondary pests, which affected 

the bark of weakened oaks. 

Second, clear yellowing of leaves was observed. All age groups were affected. Often all 

the leaves on a branch or a bough were affected whereas the neighbouring leaf material of 

the same tree showed no signs of damage. In extreme cases the entire tree was affected. 

Third, old oaks often showed a high percentage of dry branches with crown thinning. In 

advanced stages the root-infecting fungus Armillaria mellea was found to speed up the 

process of dying. 

Although the health condition of older oaks in Switzerland has worsened, it does not 

seem that the observed symptoms and the damages point to an oak decline. The causes for 

the worsening health state of oak are probably the result of unfavourable weather (droughts, 

frost), pollution and to some extent also silvicultural activities (e.g. stress induction due to 

the transformation from coppice with standards into high forest) (Hammerli and Stadler 

1988). The ageing of certain oak stands could also be important. 

Beech 

In 1995 some cases of beech bark disease were registered. Various causes may explain this 

complex disease: disturbance of the hydrological regime of the soil, infestation with the 

canker fungus Nectria coccinea and attacks of the scale insect Cryptococcus fagi. Apart from 

beech bark disease, beech canker (Nectria ditissima) is locally important. Especially saplings 

can be affected (Meier et al. 1996). However, in Switzerland beech is not considered as 

seriously threatened by pathogens. 


Various research activities are focused on the native oak species. Two projects are carried out 

at WSL. Within the project 'Conservation of Genetic Resources in Forests' the genetic diversity 

within and among oak populations is characterized. The second project 'Synthetic Maps of 

Gene Diversity and Provenance Performance for Utilisation of Oak Resources in Europe' is 

carried out within the frame of the EU-FAIR programme. A third project is carried out at the 

Swiss Federal Institute of Technology. This project aims at the characterization of genetic 

diversity of Q. pubescens within Switzerland. In the project 'Conservation of Genetic Resources 

in Forests' the variation of cpDNA was assessed in more than 600 individuals from 135 

locations covering the natural range of the three native oak species (Quercus robur, Q. petraea, 

Q. pubescens) in Switzerland. Results from these analyses are expected to provide information 

on postglacial migration routes and transfer of seeds by humans. Assessment of genetic 

diversity within and among populations of oak species is also planned at nuclear marker gene 

loci, i.e. isoenzymes and microsatellites. Populations of all three native oak species will be 

included in the genetic inventory. Selection of populations for investigation will take into 

account the insights into the evolutionary history of oaks gained from chloroplast DNA 

studies. Geneflow and mating system will be investigated in some populations by observing 

the temporal dynamics of genetic structures in mature forests and progenies. Results will be 

used to identify valuable genetic resources of oaks for the designation of in situ gene reserves 

and to establish guidelines for the management of these reserves. 

At present, no research activities are performed for beech.

, COUNTRY REPORTS 51 

Current genetic conservation activities in situ 

Gene reserves 

In 1988 the Conference of Cantonal Forest Services approved a programme for the 

conservation of genetic resources in gene reserves. As a result, the Federal Office of 

Environment, Forests and Landscape (BUWAL) financed various projects for the 

implementation of genetic conservation activities. Within the framework of the current 

project 'Conservation of Genetic Resources in Forests' (1996-99) several projects for gene 

reserves for Norway spruce and silver fir were initiated and are presently under negotiation 

with the local forest services and the forest owners. Projects for oak gene reserves will be 

started within the next 2 years. The objective is to enter into long-term contracts (50 years) 

for the special management of gene reserves. One gene reserve for oaks was established in 

1993 in the Galm forest, canton Fribourg. 

Gene reserves will be under strict forest management regulations (Bonfils 1995). 

Introduction of foreign genetic material is forbidden and natural regeneration has to be used 

as far as possible to ensure transmission of all genetic information to the subsequent tree 

generation. The gene reserves are divided into four zones (zones 0-3) and for each zone 

management regulations are defined. In zone 0 selective thinnings are forbidden. This zone 

covers a relatively small area (about 2 ha) but ensures a selection process close to nature. 

Zone 1 represents the main part of the gene reserve. This zone surrounds zone 0 and is 20- 

100 ha in size. Within this zone traditional close-to-nature silviculture is performed to 

enhance natural regeneration. If natural regeneration, is not possible, artificial regeneration 

has to rely on reproductive material from the local source. Zone 2 is realized if trees of 

foreign origin grow within the gene reserve. These trees have to be eliminated at the latest 

by the end of the production period. Zone 3 is a buffer zone that prevents or reduces 

geneflow from trees of surrounding stands to the gene reserve. No particular silvicultural 

regulations are defined for this zone. Gene reserves will be incorporated in the general 

management plan of the forests. The Federal government jointly with the cantons will 

compensate the forest owner's expenditures that occur as a result of the special management 

of the gene reserves. 

Seed stands 

In Switzerland, the cantons are responsible for the supply of seed material. According to the 

Swiss forest law they determine the forest stands in which reproductive material may be 

collected. The Swiss Forest Agency is responsible for a national register of the seed stands 

and its registration according to OECD regulations. Currently the two categories 'Sourceidentified' 

and 'Selected' are available in Switzerland. The number and categories of seed 

stands for beech and oak species are shown in Table 3. 

The minimum size of seed stands for beech and oak is as follows: 0.25 ha for seed stands 

providing source-identified reproductive material; 1 ha for stands providing selected 

reproductive material. The registered stands vary from 0.25 to 700 ha (Fiirst, pers. comm.). 

Table 3. Number of seed stands classified in the national register for beech and oak 

Seed stands 'Source-identified' 'Selected' Total 

Beech 81 25 106 

Pedunculate oak 46 18 64 

Sessile oak 38 6 44

52 EUE0BGEN: S0el~1.;l BB0~DI.;lE~MES 


Close-to-nature silviculture 

According to the 1991 Federal Forest Law the cantonal forest service has to perform close-tonature 

silviculture. This is achieved by promoting natural regeneration and using natural 

growth patterns to obtain uneven-aged and well-structured stands. The resulting diversity 

in species, stand structures and age classes is important for sustainable benefits from forests 

and, furthermore, very valuable for nature protection. The use of natural regeneration is one 

of the key points of the concept of close-to-nature silviculture. In the context of in situ gene 

conservation this appears to be a prerequisite for successful management of gene reserves. 


According to the Federal Forest Law (1991) forest tending and exploitation are not 

compulsory. for the total forest area. It is possible to renounce, partially or entirely, to 

silviculture activities. Forest reserves can be declared for the protection of a diversity of 

species. Currently the cantonal forest services are working on recommendations for such 

forest reserves with special management or even strict ban of management. However, strict 

forest reserves have existed in Switzerland since the beginning of this century. There are 

numerous reserves ranging in size in which human activities are forbidden. They make up 

approximately 0.5% (6000 ha) of the total forest area (FOEFL 1995). 

Forest reserves can be integrated into gene reserves. When possible, it will be taken into 

consideration that in addition to these banned' areas there will also be areas in which an 

active genetic conservation policy is applied. 


Currently there are no ongoing tree improvement activities in Switzerland. The demand for 

reproductive material has clearly decreased during the past 20 years (Fig 4). From the mid- 

1980s to the beginning of the 1990s the total number of plants of broadleaved trees slightly 

increased. From 1991 on, the use of reproductive material decreased, showing the same 

trend as conifers had for 20 years. The annual demand for plants is currently estimated to be 

200000 for oaks and 380 000 for beech (Fiirst, pers. comm.) . 

10000 

s 

o 

,.... 8000 

s::: 

~ 

(/)' 

c 

-a. 6000 

-o 

Qi 

.c 

E 

::::I 

Z 

4000 

2000 

.... "".,,""_ ..... , 

: -+-coniferous : 

!~~r~Cl9_~e~,:,e9 ~p~ci~~; 

0 

G0l:JNmRM RER0RmS 53 


I» Swiss Federal Institute for Forest, Snow and Landscape Research (WSL), Birmensdorf 

Contacts: 

P. Bonfils (extension service) all: WSL 

R. Finkeldey (research) Gruppe Forstgenetik 

G. MMyas (research) CH-8903 Birmensdorf 

C. Sperisen (research) 

I» Swiss Federal Institute of Technology (ETH), Zurich 

Contacts: 

P. Rotach (Professur fur Waldbau) 

B. Muller (Professur fur Forstschutz und Dendrologie) 

ETH -Zentrum 

ETH-Zurich, CH-8092 Zurich 

I» Swiss Forest Agency, Berne 

Contacts: 

M. Bolliger (coordination EUFORGEN-Programme) 

E. Fu,rst (seed stands / reproductive material) 

BUWAL 

Eidg. Forstdirektion 

3003 Bern 

I» Forest services of the cantons 

Contacts and addresses can be provided by the author on request. 

Country capacities and priorities 

Capacities for conservation and research activities in Switzerland are limited. Ongoing 

projects therefore are focused only on a few species. Oak is included in these programmes 

whereas beech is low priority. 


The coordination of research projects and the collaboration of different groups could 

contribute to an overall reduction in costs and to an increase of the output on European 

level. Important factors for successful conservation of genetic resources is information flow 

among countries and availability of research results. Practical guidelines for the 

conservation of genetic resources should be discussed. In addition, research activities 

should be designed according to the needs of gene conservation. 


I thank for helpful discussions and support: U.-B. Brandli, M. Dobbertin, R. Engesser, R. 

Finkeldey, E. Furst, F. Meier, W. Ortloff, C. Sperisen. The Swiss programme 'Conservation 

of Genetic Resources in Forests' is financed by the Federal Office of Environment, Forests 

and Landscape (BUWAL). 

References 

Anonymous. 1997. Wald- und Holzwirtschaft der Schweiz. Jahrbuch 1997. Bundesamt fUr 

Statistik / BUW AL. 

Bonfils, P. 1995. Erhaltung genetischer Resourcen im Wald. Genreservat Galm. Schweiz. Z. 

Forstwes.146:295-303. 

Brandli, U.-B. 1996. Die haufigsten Waldbaume der Schweiz. Ergebnisse aus dem 

Landesforstinventar 1983-85: Verbreitung, Standort und Haufigkeit von 30 Baumarten. 

Ber. Eidgenoss. Forsch. anst. Wald Schnee Landsch. 342:1278.

54 EUFORGEN: SOCIAl... BROADI...EAVES 

Burga, c.A. and R Perret. 1998. Vegetation und Klima der Schweiz seit dem jungeren 

Eiszeitalter. Ott Verlag, Thun. 

BUWAL/Eidgenossische Forstdirektion; Eidgenossische Forschungsanstalt fur Wald, 

Schnee und Landschaft (eds.) 1992. Sanasilva-Waldschadenbericht 1992. BUWAL/ 

Eidgenossische Forstdirektion; Eidgenossische Forschungsanstalt fur Wald, Schnee und 

Landschaft (WSL). Bern. 

Dumolin-Lapegue, S., B. Demesure, S. Fineschi, V. Le-Corre and RJ. Petit. 1997. Phylogeographic 

structure of white oaks throughout the European continent. Genetics 146(4):1475- 

1487. 

EAFV (Eidg. Anstalt fUr das forstliche Versuchswesen) and BFL (Bundesamt fur Forstwesen 

und Landschaftsschutz) (eds.) 1988. Schweizerisches Landesforstinventar. Ergebnisse der 

Erstaufnahme 1982-1986. Ber. Eidgenoss. Forsch.anst. Wald Schnee Landsch. 305:1375. 

ETH-Zurich. 1993. Mitteleuropaische Waldbaumarten. Artbeschreibung und Okologie unter 

besonderer Berucksichtigung der Schweiz. Hrsg. Professur fur Waldbau und Professur 

fUr Forstschutz & Dendrologie. 

Ferris, c., A.I. Davy and G.M. Hewitt. 1997. A strategy for identifying introduced 

provenances and translocations. Forestry 70(3):211-222. 

FOEFL. 1995. Forests and Wood in Switzerland/Wood and Forest in Switzerland. Federal 

Office of Environment, Forests and Landscape (FOEFL), Swiss Forest Agency. 

Hammerli, F. and B. Stadler. 1988. Eichenschaden. Eine Ubersicht zur Situation in Europa 

und in der Schweiz. Eidg. Anstalt fur das forstliche Versuchswesen (EAFV), 

Phytosanitarischer Beobachtungs- und Meldedienst (PBMD). Bulletin No. 3. 

Huntley, B. and H.J.B. Birks. 1983. Pp. 285-305 in An Atlas of Past and Present Pollen Maps 

for Europe: 0-13 000 years ago. Cambridge University Press, Cambridge. 

Leibundgut. 1984. Unsere Waldbaume. Huber, Frauenfeld und Stuttgart. 

Meier, F., R Engesser, B. Forster, E. Jansen and E.O. Odermatt. 1996. PBMD-Bulletin, 

Forstschutz-Uberbllick 1995. Eidgenoss. Forsch.anst. Wald Schnee Landschaft (WSL). 

Schweingruber, F. 1990. Baum und Holz in der Dendrochronologie. 2. Auflage Birmensdorf. 

Eidgenoss. Forsch.anst. Wald Schnee Landschaft (WSL). 

WSL. 1995. Pressemitteilung zum Waldzustand 1994 (20.2.1995). Eidgenoss. Forsch.anst. 

Wald Schnee Landschaft (WSL).


Fagus sylvatica, Quercus petraea and Quercus robur genetic resources 

in Italy 

Paolo Menozzi 

Dipartimento di Scienze Ambientali, Universita di Parma, Parma, Italy 

Beech 

Biogeographical range and origin of beech 

In Italy, European beech (Fagus sylvatica) is a dominant species between 800 and 1500 m asl 

in part of the Alps and between 1000 m and the upper tree line in the Apennines (see Table 2 

in Bucci 1997). It should be stressed that Italian forests in general are mountain forests, with 

only 17% of the total forest area located below 500 m. 

Beech is the dominant species in 16% of Italian forests. Numerous populations exist 

outside the continuous biogeographical range, likely remnants of a wider coverage of the 

Italian peninsula that probably reached a maximum 5000-6000 years ago (Chiarugi 1939). In 

central Italy a number of fragmented populations (27) affected by various levels of isolation 

(marginal, remote-summit, remote-abyssal) have been, studied to investigate the genetic 

effects of fragmentation (Leonardi et al., unpublished). In southern Italy the largest beech 

forest separated from the continuum of the biogeographical range is the beech stand growing 

at relatively low altitude (500 m) in the Gargano peninsula, which extends to the Adriatic sea. 

There is clear palynological evidence of a southern origin of Italian beech. A compilation 

of dates of the first appearance of beech after the last glaciation shows a clear geographic 

gradient from Sicily to the Alps (Leonardi and Menozzi 1995; Watson 1996). Not enough 

evidence exists to demonstrate the importance of refugia in the Illyrian Alps as a 

recolonization source of Italian beech. 

The demonstrated presence of glacial refugia gives Italian beech populations a particular 

value for reconstructing the natural history of the species, and as a potential reservoir of the 

genetic variability associated with recolonization sources. 

Current economic importance 

Italy is the third largest importer of sawn lumber in the world, after the USA and the UK. 

This is not surprising given the large Italian furniture industry (almost US$40 billion in 

1997). Against this background the economic relevance of beech production is dwarfed: 

between 1975 and 1985, 200000 m 3 of sawn lumber (about 5% of the total production from 

all species) were produced annually, mainly from the Calabria and Campania regions 

(Bernetti 1995). Until about 30 years ago firewood used to be a much larger part of beech 

forest production (almost three times in the 1950s). In the last decades the increase in labour 

cost and the convenience of using other less bulky fuels has reduced the production of 

firewood to an amount equivalent to timber production. The conditions leading to an 

increased conversion of coppice to high forest seem to be present. 

In this situation there is clearly a great potential for expanding the economic importance 

of beech wood. 


In Italy, out of a total forest surface of 6.2 million ha, beech covers about 1 million ha: 

240000 ha of high forest, 330000 ha of coppice (or coppice being converted to high forest), 

the rest in mixed types. High forest is mostly found in southern Italy (80%). 

Silviculture followed a close-to-nature approach for a long time, producing a large 

(although difficult to quantify) proportion of beech forests with heterogeneous, uneven-aged

56 El!JE0BGEN: S0GIJ,\E BB0J,\DEEJ,\MES 

structure with natural regeneration and overlapping production periods. These are some of 

the requirements necessary for establishing seed sources. 

In general, beech forest ownership is mostly in private hands with only about one-third 

belonging to public institutions, although a large variation exists among regions. Forest 

management was transferred in the 1970s to regional authorities. The lack of coordination 

among regions has generated different policies and regulations in different regions and a 

different level of enforcement of such policies in different areas. 

Health state of the forest stands 

A critical reappraisal of previous work claiming widespread beech decline in Italy was 

published recently (Bussotti 1994). Forest damage seems within physiological limits for 

stands growing in favourable locations; ecologically marginal stands tend to show signs of 

decline from the combined effect of environmental stress and the associated secondary pestrelated 

damage. 

Research activities on genetic diversity 

Recently, the genetics of Italian beech has been the object of a series of studies reported in 

the international literature. 

The genetic variability of 21 Italian populations was studied assaying 10 (9 polymorphic) 

enzyme loci by starch gel electrophoresis (Leonardi and Menozzi 1995). The estimates 

obtained by different parameters (e.g. 16.8% average expected heterozygosity) were in line 

with the range of values (17.1-31.7) reported for European beech populations for the same 

parameter (Comps et al. 1990; Merzeau et al. 1994; Paule 1995). A closer match (24.5% for 

Italian populations compared with 23.1 % for European ones) was obtained by recomputing 

the variability estimated using only the loci in common among the different studies. A 

value of 23.2% is reported for 11 stands from Piedmont region, northwestern Italy (Belletti 

and Lanteri 1996). A slightly higher variability estimate (30%) was obtained for a population 

from the Northern Apennines using I-SSR and RAPD markers (Troggio et al. 1996). This is 

not surprising given the well-known higher power of these DNA-based markers in detecting 

variability (Miiller-Starck and Ziehe 1991). 

The excess of homozygotes usually reported for European populations (Comps et al. 1990) 

was also found in most Italian populations (Leonardi and Menozzi 1995; Rossi et al. 1996). 

No definitive explanation was given for this observation. The discussion of this 

phenomenon touches upon the partitioning of variability among and within populations. 

As with other forest tree species (all characterized by wind pollination, high outcrossing 

rates, large geographical range and long life cycles), beech populations carry most of the 

variability at the within-population level. A correspondingly low estimate of the amongpopulation 

component was found in the survey of 21 Italian populations and in the 11 

stands from Piedmont (F s1 

=0.046 and F s1 

=0.043). A very close estimate had been reported for 

a large survey of European populations (F s1 

=0.054) in Comps et al. (1990). 

Inbreeding rates were estimated for two Italian populations between 2 and 4% values, in 

line with what is reported in the literature (Rossi et al. 1996). The paternity analysis of a set 

of open-pollinated sibships carried out using RAPD markers in a Northern Apennines 

population allowed a preliminary estimate of effective pollen migration (Troggio et al. 1996). 

The mean average distance between sibships (mother tree) and potential father trees (31 m) 

was larger than the mean distance from incompatible ones (29 mt Distance effects have to 

be investigated at greater spatial scales. 

A large study of the spatial distribution of genetic variability within populations 

confirmed limited structuring at the microgeographicallevel (Leonardi and Menozzi 1996a). 

An autocorrelation study at 11 enzyme loci in 14 populations over the Italian biogeographical 

range of the species found significant spatial structuring for only 11.5% of all 

genotypes. No correlation between the amount of spatial structuring and environmental 

(latitude, longitude, altitude), structural (mean and standard deviation of tree size) and

genetic characteristics (mean expected heterozygosity, mean F ) i 

of the population was 

found. No significant differences seem to exist among loci if low heterozygosity loci are 

excluded from the analysis. 

In spite of the low genetic differentiation among populations and the low level of spatial 

structuring at the microgeographical scale, a clear pattern of variation was observed at the 

regional level: using multivariate statistics (principal components) southern and northern 

populations clustered in separate groups. Moreover the values of the synthetic multivariate 

statistics from Sicily to the Alps showed a gradient strikingly similar to the sequence of dates 

(obtained from palynological records) tracking the postglacial recolonization of the Italian 

peninsula (Leonardi and Menozzi 1995). The principal component gradient was found 

significant by a test based on resampling techniques (bootstrap) (Leonardi and Menozzi 

1996b). 

At a different scale, using different markers, a high variability was found in southern 

populations in a survey of European populations to investigate variability of chloroplast 

DNA by PCR-RFLP markers: the two populations from southern Italy included in the study 

(Pollino (Calabria) and Sicily) were, along with other southern stands from Crimea and to a 

lesser extent from the Pyrenees, among the most variable in Europe (Demesure et al. 1996). 

No correlations between allele frequencies and soil type or altitude were found (Leonardi 

and Menozzi 1995). In the Piedmontese populations a slightly higher genetic variation was 

observed in north-facing (cooler and wetter microclimates) populations than in more 

stressed southern exposures (Belletti and Lanteri 1996). Provenance studies revealed 

differences in bud flushing (Borghetti and Giannini 1982), chilling needs (Bagni et al. 1980), 

xylem embolism, growth parameters and phenology (Borghetti et al. 1993) and drought 

resistance (Tognetti et al. 1995). 

A complete diallelic cross (4x4) produced a progeny of more than 800 viable plants, ideal 

material for developing a deeper understanding of the genetics of the species using modern 

molecular tools (Ceroni et al. 1996). 

Genetic conservation in situ and ex situ 

In Italy genetic conservation for most species of forest trees is usually limited to seed stands. 

An official Register of seed stands for all principal forest tree species containing about 150 

seed stands selected according to OECD IEEC regulations has been set up at the national 

level. 

For beech a biogenetic reserve of the European Council was established for one stand 

(Cansiglio forest, Veneto region) in recognition of the outstanding features of this locality. 

Biogenetic reserves have been established for a long time for other high-quality stands 

(Vallombrosa in Tuscany and Sassofratino on a watershed between Tuscany and Romagna). 

Recently (July 1997) the status of Biogenetic Reserve has been granted to all the seed stands 

of the official Register. Beech is represented in the Register by a total of seven stands 

belonging to six different regions (Table 1). The seed stands are representative of the whole 

biogeographical range of beech in Italy. 

There is very limited replanting in Italian forests in general (see below) and it is basically 

nonexistent for beech. Seed collecting can be seen as a form of ex situ conservation. The 

state seed company in Peri (Verona) maintains a sizable seed bank from the seed stands and 

from other locations. In total, 3590 kg of beech seeds are available. 

Essentially no other ex situ conservation initiative has been taken in Italy. Although the 

availability of genetic material in controlled ex situ situations would be useful for other 

purposes (see below) the complex of nature conservation initiatives seems to protect the 

genetic resources of beech to such an extent to make ex situ conservation programmes 

unnecessary.


Table 1. Fagus sylvatica seed stands in Italy 

Locality Region Area (ha) 

Molveno Trento 16 

Cansiglio Veneto 243 

Alta Bormida Liguria 78 

Abetone Tuscany 590 

Amiata Tuscany 4 

Campo Ceraso Abruzzo 270 

Cinquemiglia Calabria 174 

Source: Ministry for Agricultural Policies. Libra nazionale dei 

Boschi da Seme 1976. 

It is interesting to stress that the population genetic investigations carried out on the 

spatial structuring of genetic variability in Italian stands can help in designing rationally the 

form of genetic reserves for the species. The low spatial structuring of genetic variability, 

with patches of significant clustering of less than 100 m and paternity not attributable to the 

nearest trees, calls for a complex design of reserves that guarantee the conservation of the 

genetic variability present in the population. 


A great conservation effort is being programmed in Italy. The total area of the national 

territory designated to be under some degree of protection in the near future is quite large 

for a highly densely populated country like Italy (3 041 000 ha, 10.09% of the whole country). 

At the moment 508 areas (18 national parks, 147 state natural reserves, 71 regional parks and 

171 regional natural reserves, 101 protected biotopes, 7 marine reserves) cover a total of 

2000232 ha (7.4% of the country). This is a great increase from just 10 years ago (in 1987 

only 3.3% of the national territory was under some form of protection). 

Many of these areas located in all parts of the country, mostly in mountain areas, contain 

beech forests (for details, connect to the National Forest Service Web site at 

http://www.corpoforestale.itlhome.htm). Although in general not designed with the 

specific purpose of conserving genetic resources, the system of Italian protected areas with 

its latitudinal span from Sicily to the Alps guarantees a vast source of beech genetic 

resources. 


No large-scale tree improvement is carried out in Italy for beech. Provenance trials have 

been established for ecological research purposes. For instance the Institute for Forest Tree 

Breeding (IMGPF-CNR, Florence) maintains the following provenances in three localities in 

Tuscany: 

• in Vallombrosa, provenances from Liguria (2), Abruzzi (4), Calabria (4) 

• in Rincine, provenances from Emilia (I), Tuscany (I), Abruzzo (2), Puglia (I), 

Basilicata (9), Calabria (3), Campania (2) 

• in Pisanino, provenances from Emilia (I), Tuscany (I), Abruzzo (2), Puglia (I), 

Basilicata (8), Calabria (2), Campania (2). 


According to data of the Statistical Institute, artificial reforestation covered on average 

approximately 10000 ha/year during the decade 1979-88 (60% broadleaves and 40% 

coniferous trees). From 1989, the afforestation activities have been supported by the EU 

(EEC 1094/80, 'set aside') related to 2500 ha in 1989, 7000 ha in 1990 and 10500 ha in 1991. 

Given the silvicultural practices used in Italy for beech, it not surprising that no sizable 

planting is carried out.



As previously illustrated, with the exception of the state seed company, there is no 

institution specifically dedicated to genetic resources activities: many share responsibility for 

their management and protection, from the Ministry of Agricultural Policies (M IPA) with its 

National Forest Service, to the regional Forest Services, the Parks and protected areas at 

various levels. Research activities on genetic resources are carried out in Universities (about 

20 of them offer a programme in Forestry) and in other research institutions: CNR (National 

Research Council), MIP A, and other public and private concerns. 


Italy has great potential for contributing to the conservation of beech genetic resources. The 

ecological variety of the biogeographical range that hosted glacial refugia makes Italian 

forests very rich in genetic resources. The numerous institutions involved in the 

management of forest resources, the political initiatives for nature conservation and the 

potentially sound research capacities all contribute to a favourable scenario for successful 

action in genetic resources conservation. A coordination capable of focusing all this 

potential on clear goals seems necessary to achieve factual results. 

International collaboration and participation in multinational programmes could play a 

very important role in overcoming the multiplicity of objectives of the different actors, and 

foster the shaping of an effective national policy in genetic conservation of beech natural 

resources. 

Oaks 

Biogeographical range and origin ofQuercus robur andQuercus petraea 

In Italy, 12 species of deciduous oaks exist, but only five have a non-negligible presence 

(Bernetti 1995). The taxonomy of oak species is an open question, and is attracting a lot of 

attention in Italy. In this paper, information on the distribution of the species most 

frequently found in Italy is given only to evaluate the relevance of the two species of interest 

for the EUFORGEN Social Broadleaves Network: Q. robur and Q. petraea. 

Out of a total of almost 1 million ha of forest prevalently made up of deciduous oaks, 

Quercus pubescens is the most frequent, followed by Q. cerris (Table 2). Quercus frainetto is 

found in the peninsular part of Italy from south of Tuscany to Calabria. 

Species (Italian name) 

Quercus pubescens (roverella) 

Quercus cerris (cerro) 

Quercus robur (pedunculata) (farnia) 

Quercus petraea (sessiliflora) (rovere) 

Quercus frainetto (farnetto) 

Table 2. Deciduous oaks in Italy 

Area (ha) 

150000 ha (high forest) 540000 ha (coppice) 

70000 ha (high forest) 200000 ha (coppice) 

a few thousand ha 

sporadic 

in mixed fomations with Q. cerris, area difficult to quantify 

The two species of interest for the Network (Q. robur and Q. petraea) are scarcely 

represented. The biogeographical range for Q. petraea is reported to extend as far south as 

the Abruzzi region (east of Rome), although stands of the species have been reported from 

the whole peninsula and Sicily. 

There is clear palynological evidence of a southern origin of Italian oaks. Italian 

populations of Q. robur and Q. petraea are at the southern margin of the European 

biogeographic range and their study is of great interest for the understanding of the 

recolonization process and to investigate the effects of fragmentation on the genetic structure 

of the species. 

The natural plain forests hardly exist in Italy, having been replaced long ago by crops of 

larger agricultural value. The ecological preferences of Q. robur for this kind of environment

60 EUEGBGEN: SGGIALE BBGADLEEAVES 

severely limit its presence in the country. Although exhaustive quantitative data on its 

distribution are not yet available, one could say with caution that the species 1s in a status 

warranting some protective measures. We will see that this statement will have to be 

validated in light of the new evidence that is being collected. 

Quercus petraea, relatively more tolerant of environmental conditions less desirable for 

agricultural use, is found in a more important extent. 


Timber production from most oak species is mainly used as firewood (1.8 million m 3 in 1989) 

while production of sawn timber is limited (70 000 m 3 in the same year) (Bernetti 1995). 

Quercus robur and Q. petraea, given their scarce presence in Italian oak forests, are at 

present of marginal economic interest. Given the high quality of their wood, the increasing 

availability of land made available by European Union agricultural policies, and the 

importance of the Italian wood manufacturing industry (see part on beech), a great potential 

can be seen for expansion of the two species. 

Silvicultural approaches and health state of the forest stands 

Quercus petraea is managed like the much more frequent Q. pubescens with which it can easily 

produce hybrids. Quercus petraea has been found to be more abundant than expected in 

central Italy, while little information exists on its presence in southern Italy (Buresti et al. 

1995). The largest population of Q. petraea in central Italy (10 ha) has been coppiced in the 

past (Cutini and Mercurio 1995). 

The sparse distribution of Q. robur makes it hard to formulate meaningful generalizations. 

The small Q. robur Policoro population (Basilicata region, Fineschi, pers. comm.; Dumolin- 

Lapegue et al. 1997), part of a small protected area, is made up by a group of old trees in poor 

health with almost no natural regeneration. 

Research activities on genetic diversity 

Given the limited distribution of the two species in Italy, the amount of research carried out 

on them is almost surprising. The potential practical use of a genetic characterization of the 

existing populations makes such activities of special interest. Five Q. robur and five 

Q. petraea populations from Piedmont have been investigated using 11 enzyme loci (Belletti 

and Leonardi 1997). Variability is mostly found between populations (G st 

=0.023 for Q. robur 

and G st 

=0.056 for Q. petraea). Sicilian populations of Q. petraea and Q. pubescens have been 

investigated using chloroplast DNA markers (Fines chi et al. 1997). Populations of Q. petraea 

from several Italian locations are included in a Europe-wide investigation carried out using 

chloroplast DNA markers (Dumolin-Lapegue et al. 1997). 

Research on the genetic variability of oaks using microsatellites is being carried out at 

Florence University (PhD thesis of P. Bruschi, under the supervision of Pro£. Grossing). 

Genetic conservation in situ andex situ, tree improvement activities and use of 

reproductive material 

The national Register lists only three stands for Quercus: one of Q. robur (70 ha in Piedmont), 

two of Q. cerris in the Lazio and Molise regions. The official availability of seeds is not 

negligible and of diversified origin (Tables 3 and 4). 

According to data of the Italian Statistical Institute, artificially reforested surfaces were on 

average approximately 10 000 ha/year during the decade 1979-88 (see information given for 

beech).

Table 3. Seed availability (kg) for Quercus robur 

Seed provenance (province)t 

Total weight (kg) BL SO MN BO VE UD RO BG VR CN PR VI TN 

8459 41 342 1105 633 447 181 325 4538 270 88 72 189 228 

t BL=Belluno, SO=Sondrio, MN=Mantova, BO=Bologna, VE=Venezia, UD=Udine, RO=Rovigo, BG=Bergamo, 

VR=Verona, CN=Cuneo, PR=Parma, VI=Vicenza, TN= Trento. 

Source: Ministry for Agricultural Policy. Uficio Amministrazione Produzione Sementi Forestali di Peri (Verona). 

Table 4. Seed availability for Quercus petraea 

Seed provenance (province)t 

Total weight (kg)· PR TS BO PD VI TN+VR VC TO CN 

5136 1657 407 5 199 176 238 570 219 1665 

BO=Bologna, VR=Verona, CN=Cuneo, PD=Padova, PR=Parma, VI=Vicenza, TN= Trento, TS= Trieste, 

TO=Torino. 

Source: Ministry for Agricultural Policies. Uficio Amministrazione Produzione Sementi Forestali di Peri (Verona). 

In response to the demand for reproductive material for afforestation of agricultural land 

freed by the 'set aside' European Union policy, a great deal of interest has been dedicated to 

the evaluation of possible sources of genetic material. These efforts are still in their initial 

phase. For instance, reproductive material was sampled from about 70 natural stands for 

both species in northern and central Italy. An ex situ collection of nine provenances was 

established by the Forest Research Institute of Arezzo in collaboration with regional 

authorities. A 4-ha plot has been established for the production of reproductive material. 

These initial efforts reflect the awareness of the importance of inventorying, evaluating 

and developing the genetic resources of the country, which are potentially very rich for the 

above-mentioned reasons. 


The general information given for beech on this topic also applies to oaks. In addition, given 

their economic importance and sparse distribution, there is an obvious need for a complete 

mapping of the extant populations of Q. robur and Q. petraea. As is the case for beech, the 

populations most valuable for genetic conservation are probably already under some kind of 

protection. But it is likely that given the small size of the remaining populations, many of 

those still have to be identified. Their evaluation through appropriate procedures is 

necessary so that they can be granted the protection they deserve. 


The same institutions involved in genetic resources activities of beech are also involved in 

the handling of genetic resources of Q. robur and Q. petraea. 


Italy has a great potential for contributing to the conservation of Q. robur and Q. petraea 

genetic resources. Owing to the ecological variety of the biogeographical range that has 

hosted glacial refugia and to the fragmentation and reduction in size of the populations of 

both species, the study of their genetic resources present in Italy is very interesting. Their 

potential economic importance should be able to stimulate the numerous institutions 

involved in forest resources management to coordinate their action to achieve factual results. 

International collaboration and participation in multinational programmes is even more 

necessary for Q. robur and Q. petraea than for beech, to overcome the organizational 

difficulties and to acquire the standard of scientific knowledge necessary for effective 

conservation of their genetic resources.

62 EUF'ORGEN: SOCIAl.. BROADI..EAVES 

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petraea) nell'Italia centrale. Linea Ecologica 27(4):18-24. 

Demesure, B., B. Comps and RJ. Petit. 1996. Chloroplast DNAphylogeography of the 

common beech (Fagus sylvatica L.) in Europe. Evolution 50:2515-1520. 

Dumolin-Lapegue, S., B. Demesure, S. Fineschi, V. Le Corre and RI. Petit. 1997. 

Phylogeographic structure of white oaks throughout the European continent. Genetics 

146:1475-1487. 

Fineschi, S., A. Diarra, A. Balijja, G. Scuderi and D. Taurchini. 1997. Polimorfismo del DNA 

cloroplastico in popolazioni di querce (sezione robur) in Sicilia: una risorsa genetica da 

salvaguardare. Primo Congresso SISEF. Summary. 

Leonardi, S. and P. Menozzi. 1995. Genetic variability of Fagus sylvatica L. in Italy: the role of 

postglacial recolonization. Heredity 75:35-44. 

Leonardi, S. and P. Menozzi. 1996a. Spatial structure of genetic variability in natural stands 

of Fagus sylvatica L. (beech) in Italy. Heredity 77:359-368. 

Leonardi, S. and P. Menozzi. 1996b. Methodological aspects in spatial analysis of genetic 

structure in Italian populations of beech (Fagus sylvatica L.). Statistica Applicata 8:281-297. 

Merzeau, D., D. Comps, B. Thiebaut, J. Cuguen and J. Letouzey I. 1994. Genetic structure of 

natural stands of Fagus sylvatica L. Heredity 72:269-277. 

Miiller-Starck, G. and M. Ziehe. 1991. Genetic variation in populations of Fagus sylvatica L., 

Quercus robur L. and Quercus petraea Liebl. in Germany. Pp. 125-140 in Genetic Variation 

in European Populations of Forest Trees (G. Miiller-Starck and M. Ziehe, eds.). J.D. 

Sauerlander's Verlag, Frankfurt am Main.


Paule, L. 1995. Gene conservation in European beech (Fagus sylvatica L.). For. Genet. 2:161- 

170. 

Rossi, P., c.c. Vendramin and R. Giannini. 1996. Estimation of mating system parameters in 

22 Italian populations of Fagus sylvatica L. Can. J. For. Res. 26:106-111. 

Tognetti, R., J.D. Johnson and M. Michelozzi. 1995. The response of European Beech (Fagus 

sylvatica L.) seedlings from two Italian populations to drought and recovery. Trees 9:348- 

354. 

Troggio, M., E. DiMasso, S. Leonardi, M. Ceroni, G. Bucci, P. Piovani and P. Menozzi. 1996. 

Inheritance of RAPD and I-SSR markers and population parameters estimation in 

European beech (Fagus sylvatica L.). For. Genet. 3:173-181. 

Watson, C.S. 1996. The vegetational history of the northern Apennines, Italy: Information 

from three new sequences and a review of regional vegetational change. J. Biogeography 

23(6):805-841.

64 EUE(i)RGEN: S(i)el~U BB(i)~DUE~MES 

Beech and oak genetic resources in Slovenia 

Igor Smole/, Robert Brus 2 , Marjanca Pavli, Saso Zitnikl, Zoran Grecs 3 , Nevenka Bogata/, 

Franc Ferlin 1 and Hojka Kraigherl 

1 Slovenian Forestry Institute, Ljubljana, Slovenia 

2 Forestry Department, Biotechnical Faculty, University of Ljubljana, Ljubljana, Slovenia 

3 Slovenian Forest Service, Ljubljana, Slovenia 

Introduction 

Forests cover 53% of Slovenia. Because of the mountainous and karstic character of the 

country, which makes access to many forests difficult, the degree of human intervention has 

been lower than in most central European countries. In addition, owing to highly diverse 

ecological conditions, forest sites and their tree species are characterized by a high diversity. 

Because of the traditional close-to-nature forestry management for sustainable and 

multifunctional use, the species composition in 87% of the Slovenian forests is equal to or 

very similar to the potential one. Only in 9% of all forests has the species composition been 

changed significantly, and in 4% the species composition is completely different from the 

natural one (Grecs and Kraigher 1997). The potential forest types are presented in Table 1. 

However, the proportion of the most important tree species in the growing stock of 

Slovenian forests is different (Table 2). 

Occurrence, origin and current economic importance of beech and oaks 

for the forestry sector 

European beech (Fagus- sylvatica L.) is an indigenous tree species in Slovenia. It represents 

29% of the total growing stock but in the potential natural vegetation its share would be as 

high as 58%. A great majority of the beech forests are natural, for in the past practically no 

beech was planted in Slovenia. During the glacial period the majority of beech populations 

survived in the refugia of southern Italy, southern France and on the Balkans peninsula. In 

the postglacial period, approximately 8000 BP, beech started to spread into the territory of 

Slovenia again. The so-called beech stage, following the Quercus-Corylus stage, starts in the 

period before Boreal and is subdivided into a Fagus-Abies substage, when fir and alder 

prevailed, a pure Subboreal Fagus stage, when beech increased due to a warmer and drier 

climate, and another Fagus-Abies stage in which the living space of beech was reduced and 

which lasts up to the present (Sercelj 1972; Horvat-Marolt 1992). 

Table 1. Forest sites in Slovenia 

(The Forest Development Programme of Slovenia, 1996) 

Forest sites 

Beech forests on carbonate ground rock 

Acidophilous beech forests 

Forests of beech and silver fir 

Forests of beech and oak 

Forests of oak and hornbeam 

Thermophilic broadleaved forests 

Silver fir forests 

Alpine forests 

Pine forests 

Oak forests 

Norway spruce forests 

Forests of willow and alder 

Total 

Area (ha) 

286074 

179451 

163581 

115166 

87373 

57936 

49228 

41 525 

39394 

33769 

15471 

7508 

1 076474 

% 

27 

17 

15 

11 

8 

5 

4 

4 

4 

3 

1 

1 

100

~ G0UNmB¥ BER0BmS 65 

Table 2. Proportion of the most important tree species in the growing stock of Slovenian 

forests (The Forest Development Programme of Slovenia, 1996) 

Valuable Other 

Vegetation 

Potential (%) 

Current (%) 

European 

beech 

58 

29 

Norway 

spruce 

8 

35 

Silver 

fir 

10 

11 

Oaks 

8 

8 

broad leaved 

species 

6 

3 

broad leaved 

species 

8 

7 

Pine 

2 

7 

Lowland' forests of oaks are the most changed forest ecosystems in Slovenia. As a result 

of agricultural activities and urbanization, they have been transformed into cultivated 

steppes and urban areas. Only a few remnants subsist and even these are affected by 

various disturbances like changes in water table, input of fertilizers, air pollution, etc. 

However, the species structure in these remnants is largely natural. There are seven 

indigenous oak species (MartinCie and Susnik 1984; Azarov 1991; Batie et al. 1994; Batie et al. 

1995) and two introduced oak species in Slovenia (Table 3). 

Most of the pedunculate oak (Quercus robur L.) forests have been cleared in the past for 

agricultural use. The biggest residues of once widespread forests are in lowland, 

occasionally flooded areas of Krakovski gozd. This Krakovo Forest is also known because of 

the largest virgin oak forest reserve, formed predominantly by pedunculate oak. Other 

complex localities are sited north from Breice and along the rivers Mura and Ledava. 

Residual forest fragments are known from the valleys of Drava, Paka and Mislinja rivers, on 

the Ljubljana's moor and even on some carbonate soils in the Midlands (Notranjska). 

Sessile oak (Quercus petraea (Matt.) Liebl.) is the most widespread oak species in Slovenia. 

It forms approximately 87% of all oak species in the growing stock, grows equally well on 

carbonate and silicate ground rock materia1, mainly up to 700 m asl, but also above 1000 m. 

The overview on the areas and growing stock in the forest management units (Slovenian 

Forest Service 1990) in Slovenia are presented in Table 4. 


Silvicultural measures in Slovenia depend on the Forest Development Programme of Slovenia 

and the Forest Management Plans. On the basis of these, detailed silvicultural plans are 

prepared for each forest management unit (on average 50 ha of forest), belonging to districts 

(area from 3000 to 6000 ha), these being included in forestry regions (14 in Slovenia). 

According to the Forestry Act (1993) the detailed silvicultural plans define: 

• necessary cultivation work for regeneration and tending of the forests, including the 

first selective thinning (before the first commercial thinning) 

• necessary protection work 

• guidelines and time limits for carrying out tending and protective work 

• quantity and structure of trees for the maximum allowable cut 

• guidelines and conditions for felling and skidding timber. 

On the basis of silvicultural plans and other operational projects or plans within the 

framework of the investment programme for forests, drawn up by the Slovenian Forest Service 

for the current year, the state subsidizes the following silvicultural and protective measures: 

• measures for preventing or mitigating the disturbances in the functioning of the forest 

and forest work in protective forests and torrent watersheds 

• measures for silvicultural protection and for the maintenance of wildlife habitats 

• production of seeds and seedlings in a nursery and investments in forest tree nurseries 

• restoration of forests if the party responsible for the damage is unknown 

• reforestation after fires and natural disturbances 

• thinning of pole stands and conversion into private forests.

66 EUFORGEN: SOCIAl... BROADl...EAVES 

Table 3. Beech and oak (Azarov 1991) species in Slovenia, their status (Wraber and Skoberne 

1989) and the annual felling in 1996 (Zavod za gozdove Slovenije 1996) 

Latin name Common name Status t Annual felling in 1996 (m 3 ) 

Fagus sylvatica L. European beech common 540 906 

Quercus robur L. pedunculate oak 7% of all oaks 10 913 

Quercus petraea (Matt.) Liebl. sessile oak 82% of all oaks 76 128 

Quercus ilex L. holm oak EN, L 0 

Quercus cerris L. Turkey oak 8% of all oaks 6 260 

Quercus pubescens Wild. pubescent oak 2% of all oaks 328 

Quercus crenata Lam. false-cork oak EN, L 0 

Quercus virgiliana Ten. Croatian oak EN, L 0 

Quercus palustris Muenchh. swamp oak I 134 

Quercus rubra L. red oak I 101 

t I = introduced species; EN = endangered species in terms of IUCN categories; L = the species being 

in its geographical borderline and occurring in limited numbers in Slovenia. 

Table 4. Areas and growing stock in the forest management units for beech and commercially 

im~ortant oak s~ecies in Slovenia 

Area of 

"10 tree sp. 

units Stock of tree s~. {m 3 l Total stock {m 3 l in total 

Tree species (ha) Units Units/ha Units Units/ha stock No. units 

European beech 903501 61 323849 67.87 190874477 211.26 32.13 64619 

Sessile oak 528266 12297359 23.28 94169135 178.26 13.06 39434 

Pedunculate oak 30379 1 010362 33.26 5209483 171.48 19.39 2708 

Turkey oak 84446 1 257606 14.89 10265231 121.56 12.25 4572 

Pubescent oak 28770 338424 11.76 1 657657 57.62 20.42 1394 

Swamp oak 724 2550 3.52 122363 169.01 2.08 38 

Red oak 2261 14953 6.61 371 877 164.47 4.02 161 

Source: The Slovenian Forest Service, 1990. 


Forest decline monitoring activities started in Slovenia in 1985. A 4x4 km network was 

established, while a 16x16 km grid (part of the 4x4 km network) was used for a few more 

detailed studies (foliar nutrient analysis, etc.). The health state of beech and oaks is speciesspecific: 

beech shows a slight trend of decline (where it is necessary to mention extreme 

droughts as well as air-pollution effects), while oaks have shown high decline trends since 

1990. The probable causes for oak decline are several: changed water table, air pollution, 

root pathogens, defoliating insects, etc. The results from forest decline monitoring for the 

last 10 years are shown in Figure 1. 

In the three most air-polluted areas in Slovenia the genetic structure of beech populations 

has been studied and compared with the unpolluted populations. Significant differences of 

allelic frequencies between the groups of polluted and unpolluted populations on two 

isoenzyme loci were determined. A substantial change of allelic frequencies is indicated by 

the fact that genetic distances are highest when we compare polluted and unpolluted 

populations or polluted populations only. There are some indications that selection against 

some alleles is present, but this was not unambiguously confirmed. Only some small 

differences in genetic diversity between polluted and unpolluted groups were revealed 

(Brus 1996b). 

Other disturbances which were very important in the last 15 years are extreme winters, 

extreme droughts, snow and ice-breaks in the last 2 years. The whole range of disturbances 

has resulted in high sanitary felling during this period (as presented in Tables 5 and 6). 

In previous studies, oak decline has been studied on nine forest research plots (Golob 1991). 

These were followed by two small oak projects: on taxonomy, cytogenetics and isoenzyme 

studies of oaks (Batic 1996; Rogl et al. 1996), which included also several Diploma and MSc

. COUNTRY REPORTS 67" 

theses (Mavsar 1996; Breznikar 1997), the genetics of oaks in Slovenia (Benedik 1997), and on 

oak decline in Slovenia (Batic 1997). 

Silvicultural studies and provenance trials with beech progressed well between 1951 and 

1971, when the main Slovenian geneticist, M. Brinar, was active at the Institute. He partially 

described two beech races and mapped several interesting beech morphotypes. His work 

was largely forgotten until recently, when he offered collaboration with some young 

researchers. 

At the Forestry Department of the Biotechnical Faculty studies on population genetics of 

beech in Slovenia have just started. The main goal is to compare the genetic characteristics 

of the Slovenian populations with the populations from other parts of Europe, study the 

existence of different races or types of beech in Slovenia and define the influence of 

ecological parameters on the genetic structure of populations. Reconstruction of potential 

postglacial migrations of beech will also be attempted, since the territory of Slovenia is 

supposed to be an important crossing of a number of tree species' routes. 

Research activities and capacities related to genetic resources 

Within the National Research Programme 'The Forest', lasting from 1995 to the end of 1999, 

one single project, financed at 50% by the Ministry for Agriculture, Forestry and Food and 

50% by the Ministry for Science and Technology, with less than 1 FTE (full time equivalent) 

staff, was accepted for forest genetics, physiology, seeds and nursery studies. It is run by 

both research organizations, working in forestry in Slovenia, the Slovenian Forestry Institute 

and the Forestry Department of the Biotechnical Faculty. This project is enhanced by two 

young researchers' projects: on the role of phytic acid in physiology of acorns during 

storage, and on the genetic variability of Norway spruce (Table 7). 

40 

-';:/2.. 

0 

.......- 30 

Cl) 

(]) 

(]) 

'"- 

....- 

-0 

(]) 

0) 20 

CI:l 

-0- European beech 

Fagus sy/vatica 

"0· Sessile oak 


E o. 

CI:l . 

-0 

p' 

D'" .0, 

..- '0 

0 10 

....- 

C 

(]) 

U 

'"- 

(]) 

0... 0 

1985 1987 1989 1990 1991 1993 1994 1995 1996 1997 

Year 

t1 

0 

Fig. 1. Monitoring of beech and sessile oak decline in Slovenia: on 4x4 km network (on approximately 

700 plots) for 1985, 1987, 1991 and 1995; on 16x16 km network (43 plots for all tree species) for 1989, 

1990,1993,1994,1996,1997.

Table 5. Structure of felling of Slovenian beech and oaks in 1996 in % of total 635000 m 3 (for 

felling per tree species see Table 3), which represents 27% of total yearly felling of all tree 

species or 76% of all broad leaved tree species 

Selective thinning Artificial regeneration Sanitary felling Other felling 

Tree sp. (%) (%) 

(%) 

(%) 

Beech 

80 0.1 

Sessile oak 

57 0.1 

Pedunculate oak 37 10 

Red oak 

35 0 

Swamp oak 84 o 

Downy oak 50 

Turkey oak 74 

Source: Slovenian Forest Service, 1996. 

o 

2 

13 

31 

43 

10 

15 

49 

16 

7 

12 

10 

15 

1 

1 

8 

Table 6. Silvicultural measures in Slovenian forests in 1996 

Individual 

protective Tending of Tending of 

measures young growth thickets 

Programme 96 (ha) 2224 3575 6334 

Realisation (ha) 1709 1329 2375 

Real.lProg. 96 {%} 77 37 37 

Source: Slovenian Forest Service, 1996. 

Selective Total 

thinning tending 

4077 16210 

2091 7504 

51 46 

Table 7. Current research activities t related to forest genetic resources 

No. of SFI SF For. Other 

Programme 

projects (FTE) (FTE) (FTE) 

National Programme - Forest 1 0.5 0.3 0 

National research programme 1 0.8 0 0.2 

Bilateral projects (F, eRO) 

1 visits 0 0 

International projects 

o 0 0 0 

Young researchers (MSc, 

PhD) 

2 2 0 0 

Total 

(FTE) 

0.8 

1 

visits 

o 

2 

Duration 

(years) 

1995-99 

1997-99 

1998-99 

o 

1996-99 

Students (BSc, MSc) 2 0 0 0 O 

Public Forest Service 1 2 0 0 2 permanent 

t 

FTE = Full Time Equivalent (1700 working hours per year per person); SFI = Slovenian Forestry 

Institute, BF For. = Forestry Department at the Biotechnical Faculty; the duties of the Public Forest 

Service are written in the Forestry Act and the Forest Development Programme of Slovenia and are 

performed mainly by the Slovenian Forest Service, while some aspects, such as The Seed Objects and 

The Forest Gene Bank, are done or led by law by the Slovenian Forestry Institute. 


According to the Slovenian Forestry Act of 1993 all forests are managed in a close-to-nature 

way, classified as Category VI of the IUCN management categories; 'protected area managed 

mainly for the sustainable use of natural ecosystems' (Wraber and Skoberne 1989). 

Sustainable, close-to-nature and multifunctional forest management implies: 

• small-scale flexible forest management, easily adapted to site conditions and natural 

development of forests 

• active protection of natural populations of forest trees 

• protection and conservation of biological diversity in forests 

• support of the bio-ecological and economic stability of forests by improving the 

growing stock 

• tending of all developmental stages and all forest forms for support of vital and highquality 

forest trees, which could optimally fulfil all functions of forests

• natural regeneration is supported in all forests; if seedlings are used, they should be 

derived from adequate seed sources/provenances, and only adequate species can be 

used. 

Additionally, about 135000 ha of forests are protected under different IUCN categories 

(Table 8). The network of virgin forest reserves was established in the 1970s on all possible 

forest sites (Mlinsek et al. 1980). They take areas from 1.6 to 500 ha. Beech grows as the 

dominant tree species in 62% of them, in 30% it grows together with oaks, while oaks are 

dominant species in 4% of all reserves. Beech and oaks occur in 96% of forest reserve areas, 

while beech forms 50% and oaks 3% of the growing stock in these reserves. 

Most forest stands are regenerated naturally, only one-tenth are regenerated from nursery 

seedling material, while seeds are mostly collected from acknowledged seed stands. 

Therefore, no special attention is given to ex situ conservation of forest genetic resources in 

Slovenia. 

The use of reproductive material from selected seed stands started in Slovenia in the 

1950s (Wraber 1951; Brinar 1961). Evaluation and selection of seed stands, the register of 

seed stands and the Slovenian forest gene conservation projects are run through the Public 

Forest Service, defined in the Act on Forestry (1993). The first revision of seed stands was 

done by 1987, the second has just been finished (Pavle 1987, 1997). The main change over 

the last 10 years is in the planned search for seed stands of broadleaved tree species, 

whereby the recorded number of these seed stands has risen from 68 in 1987 (Pavle 1987) to 

about 168 this year (Pavle 1997). The numbers and distribution of seed stands of oaks and 

beech in Slovenia are presented in Figures 3 and 4, their seed units in Tables 9 and 10. 

Tree-improvement activities and the use of reproductive material 

There are no tree breeding programmes for oaks or beech in Slovenia. However, for 

conservation of biodiversity and support of natural or site-adapted populations of forest 

trees, only seedlings derived from seeds collected in seed stands of the same seed unit from 

certain regions of provenance can be used for regeneration. Seed units are defined. 

considering the potential forest type, bedrock and elevation (Pavle 1997). They are groups of 

similar potential forest types, among which exchange of seeds and seedlings is allowed. 

They belong to four altitudinallevels (0-400 m, 401-700 m, 701-1000 m, above 1000 m) and 

two groups based on bedrock material (carbonate, non-carbonate). The seed units are 

separately formed for each tree species. For beech and oaks there are eight seed units in 

Slovenia (on carbonate and silicate ground rock material and in four altitudinal zones). Each 

potential forest type can be included into one or more seed units, considering its altitude. 

Plans for 1997 included natural regenetation of 2150 ha of forests and planting or sowing 

of 750 ha of forests. For this, about 3 million seedlings, 60% of which broadleaved, were 

needed.

70 EUFORGEN: SOCIAl.. BROADl..EAVES 

lJ 

lJ 

lJ .. 

••• 

lJ 

.. .. 

• 

... 

lJ 

.. 

.. 

lJ 

)( 

lJ 

• 

.. 

A[] 

• 

Fig. 3. Seed stands of different oak species in Slovenia: • Q. robur (9 seed stands); II1II Q. petraea 

(10 seed stands); D Q. rubra (9 seed stands); x Q. cerris (2 seed stands); f::, Q. palustris (1 seed 

stand). 

• • • 

• • • 

• • 

• • • 

• • • • 

• • • 

• 

• • • 

• •• 

• • 

• • • • • • • • 

• 

• 

• 

• • 

• • • 

• • 

• 

• • 

• 

• 

• • 

•• 

• 

~ 

Fig. 4. Seed stands of beech in Slovenia (65 seed stands). 

Table 8. Protected forest areas 

Protected forests IUCN category Area of forests (ha) 

Triglav National Park 11 / V 36 240 

36 regional parks V, one III 30045 

173 forest reserves I 10 421 

Protection forests I / V 55 400 

Seed stands IV / VI 2313 

Sources: modifed from Kraigher et al. 1996; Pavle 1997. 

Protected since 

1924,1981 

most from 1984 or later 

some from 1887, 1973 

most from 1852 

1955, updated in 1997

o COUNTRY REPORTS 71 

Table 9. Beech seed stands in different seed units in Slovenia 

Seed unit Altitude {m asJ} Area {ha} No. of seed stands 

B-1k

!7i2 EUE(,:)BGEN: S(,:)GI~1!l; BB(,:)~DI!l;E~MES 


In Sloverua, 53% of the country is covered by forests, which have been grown in a close-tonature 

and sustainable way for almost 150 years. The highly diverse ecological conditions 

have supported the high biodiversity at the ecosystem, species and genetic levels, which 

have all been well conserved. European beech is the most naturally widespread tree species 

in Slovenia. It presents 29% of the current growing stock. Of the seven indigenous oak 

species, three are at their geographical borderlines and occur in limited numbers. The other 

four and the two introduced oak species (of minor importance) form 8% of the current 

growing stock in Slovenia, of which 82% represents the sessile oak. The silvicultural 

approaches in Slovenia depend on the Forest Development Programme and the Forest 

Management Plans, used as a basis for preparation of detailed silvicultural plans according 

to the Forestry Act. The majority of felling is by selective thinning, while in the last few 

years heavy snow and ice damage have increased sanitary felling. The health state of beech 

shows a slight decline in the last 10 years, while the health state of oaks has worsened 

remarkably since 1990. However, a single study of air pollution impacts on genetic diversity 

of beech populations did not show significant differences. Research activities comprise a 

small project on oak decline, an MSc project on water stress, another one on acorn 

physiology during storage and a PhD project on population genetics of beech. 

Within the framework of 173 forest reserves, beech grows as the dominant species in 62%, 

in 30% it grows together with oaks, while oaks grow as dominant species in 4% of them. 

The use of forest reproductive material deriving from selected seed stands has been applied 

since 1951. At the moment there are 65 beech seed stands in eight seed units, defined after 

the phytocenological associations, the groundrock material and altitudinal levels; and 31 

seed stands for five oak species. However, seed collecting and distribution need a better 

control system. This should be done through acceptance of a new Act on forest reproductive 

material, which is being prepared, following as much as possible the directives of the ED. 

International collaboration is needed for the above-mentioned research and legislationrelated 

activities. 


Collaboration in research projects: 

.. Genetic studies of beech: international collaboration is essential. The sampling has been 

planned to include beech populations from Slovenia, Italy, Croatia, Bosnia and 

Herzegovina and a few other countries. The laboratory part of the studies with 

isoenzyme markers has been carried out at the Forestry Faculty of the Technical 

University in Zvolen, Slovakia. 

11 The studies of the role of phytic acid in long-term storage and physiology of acorns are 

done in collaboration with the Department of Plant Sciences, University of Cambridge, 

UK. Also a bilateral project has just started to enable collaboration with the Forestry 

Research Center at INRA, Nancy, France. 

Collaboration in preparation of the new legislation on forest reproductive material: 

11 In order to prepare the new act on forest reproductive material as close to the EU 

directives as possible, an unofficial collaboration has started through the bilateral DAAD 

visits scheme with the Institute for Forest Genetics and Tree Breeding, Grosshansdorf, 

Germany. More visits, also to different institutions and possibly to the EU officials from 

this field, would be needed. 

.. Inclusion of the Slovenian strategies for sustainable use and support of the high 

biodiversity at all levels in the Slovenian forests into the European strategies might help 

in further harmonizing our legislation.


We would like to thank Robert Ogrizek from the Slovenian Forest Service for handling some 

of the data from the 1996 Report, and Matja Cater and Gregor Bozie for comments. This 

report is part of the Public Forest Service of the Slovenian Forestry Instit~te, belonging to the 

Slovenian Forest Genebank project. 

Bibliography 

Anonymous. 1973. Act on Seeds and Seedlings - Ur.L.5RS 42/73. 

Anonymous. 1976. Regulation on Protection of Rare or Endangered Plant Species. - Ur.L.SRS 

15/76. 

Anonymous. 1990. Popis gozdov. Slovenian Forest Service Databases, Ljubljana. 

Anonymous. 1993. Act on the Protection of the Environment - Ur.L.32/93. 

Anonymous. 1993. Forestry Act - Ur.L.RS 30/93. 

Anonymous. 1994. Decree on Financing and Co-Financing Investments in Forests - URl.RS 

58/94. 

Anonymous. 1996. Annual Report. Slovenian Forest Service, Ljubljana. 

Anonymous. 1996. The Forest Development Programme - Ur.L.RS 14/96. 

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Ljubljana. 

Batie, F. 1997. Oak deCline in Slovenia. End report, GIS, Ljubljana. 

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severovzhodni Sloveniji. MSc Thesis, Univ. Ljubljana. BF Forestry Dept., Ljubljana. 

Brinar, M. 1961. Naeela in metode za izbiro semenskih sestojev. GozdV 19:1-20. 

Brus, R 1996a. Hrast oplutnik (Quercus crenata Lam.) tudi na Krasu. GozdV 54(10):511-515. 

Brus, R 1996b. Vpliv onesna evanja ozraeja na genetsko strukturo bukovih populacij v 

Sloveniji. Zbornik gozdarstva in lesarstva 49:67-103. 

Golob, S. (ed.) 1991. Forschung der Waldokosysteme und der Forstlichen Umwelt, Bericht 

iiber die Forschungszusammenarbeit Slowenien-Osterreich, GIS, Ljubljana. 

Grecs, Z. and H. Kraigher. 1997. Interakcije v mikorizosferi in komplementarnost naravne 

obnove in obnove s sadnjo ali setvijo. Pp. 297-308 in Znanje za gozd. Spominski zbornik 

ob 50-letnici GIS. GIS, Ljubljana. 

Horvat-Marolt, S. 1992. A historical analysis of beechwoods. Pp. 3-16 in Actas del Congreso 

Internacional del Haya, Pamplona, Vol. 1. Investigacion Agraria (R Elena Rosello, ed.). 

Itnik, S., G. Bozie, M. Pavle and H. Kraigher. 1997. Gospodarjenje in zakonodaja na podroeju 

gozdnih genskih virov v Sloveniji in Srednji Evropi. Pp. 309-320 in Znanje za gozd. 

Spominski zbornik ob 50-letnici GIS. GIS, Ljubljana.

7'4 EUFORGEN: SOCIAL. BROADL.EAVES 

Kraigher, Z. 1996. Kakovostne kategorije gozdnega reprodukcijskega materiala, semenske 

planta e in ukrepi za izboljsanje obroda. Zbornik gozdarstva in lesarstva (Tematska st. 2: 

Kvaliteta v gozdarstvu) 51. Pp. 199-215. 

Kraigher, H., G. Bozic, R. Brus, A. Golob, M. Pavle and M. Veselic. 1996. Forest genetic 

resources. in The Republic of Slovenia, Country report, International Conference and 

Programme for Plant Genetic Resources (ICPPGR). Ministry of Agriculture, Forestry and 

Food (M. Cerne and H. Kraigher, eds.). 27 p. 

MartinCic, A. and F. Susnik. 1984. Mala flora Slovenije. DZS, Ljubljana. 

Mavsar, R. 1996. Morphological variability of common oak (Quercus robur L.) populations in 

Slovenia. BSc Thesis, Univ. Ljubljana. BF, Forestry Dept., Ljubljana. 

Mayer, E. 1958. Pregled spontane dendroflore Slovenije. GozdV 16:161-191. 

Mlinsek, D. et al. 1980. Gozdni rezervati v Sloveniji. Pp. 1-414 in J. Bozic, ed. GIS, Ljubljana. 

Pavle, M. 1987. Semen ski sestoji v Sloveniji. Register, GIS, Ljubljana. 

Pavle, M. 1997. Semen ski sestoji v Sloveniji. Register, GIS, Ljubljana. 

Rogl, S., B. Javornik, T. Sinkovic and F. Batic. 1996. Characterisation of oak (Quercus L.) seed 

proteins by electrophoresis. Phyton (Horn, Austria) 36:159-162. 

Sercelj, A. 1972. Verschiebung und Inversion der postglacialen Waldphasen am sudostlichen 

Rand der Alpen. Ber. Deutsch. Bot. Ges. :123-128. 

Smolej,1. 1991. Bericht uber die Entnahme von Blattproben der Autochtonen Eichenarten in 

Slowenien fur Taxonomische Determinierung. Pp. 146-152 in Forschung der 

Waldokosysteme und der Forstlichen Umwelt. Bericht uber die Forschungszusammenarbeit 

Slowenien-Osterreich. GIS, Ljubljana (S. Golob, ed.). 

Smolej, 1. and F. Batic. 1992. Pomen morfoloskih znakov pri dolocanju hrastovih vrst. 

Zbornik gozdarstva in lesarstva 39:133-172. 

Trpin, D. and B. Vres. 1995. Seznam praprotnic in semenk Slovenije. CD, ZRC-SAZU, 

Ljubljana. 

Wraber, M. 1951. Nova pota gozdne semenarske sluzbe. GozdV 9:3-14. 

Wraber, T. and P. Skoberne. 1989. Rdeci seznam ogro enih praprotnic in semenk SR 

Slovenije. Varstvo narave 14/15, Ljubljana.


Oak genetic resources in Malta 

Darrin T. Stevens 

Environment Protection Department, Floriana, Malta 

I ntrod uction 

Oaks in Maltese are called 'ballut', whilst the acorns are referred to as 'gandar'. Only one 

species is known to be native, namely Quercus ilex subsp. ilex, the holm or evergreen oak, 

which is locally very rare and also listed in the Red Data Book for the Maltese Islands 

(Lanfranco 1989). 

Quercus robur subsp. robur, the pedunculate oak (or English oak), occurs as a rare planted 

street tree but also as a very rare naturalized alien species (Borg 1927; Haslam et al. 1977). It 

was introduced by the British, as the Maltese Islands formed a colony. Thus, Q. robur is 

referred to in Maltese as Balluta Ngliza, literally English Oak, but also as Sigra tar-Ruvlu, 

Oak-Wood Tree. 

Other species occur - Quercus cerris and Quercus infectoria subsp. veneris - but these are 

very rare planted ornamentals, occurring in few public gardens or public areas. 

With respect to habitat, the few Q. robur occur planted in public gardens and forest 

remnants, or as street trees. Quercus robur is naturalized in the semi-natural woodland of 

Buskett. 

Uses 

At present, no local uses of Q. robur are known. Acorns used to be fed to goats, pigs and 

other domesticated animals, but these were most probably acorns of Q. ilex (Lanfranco 1993). 

Conservation and silviculture 

Few ex situ conservation measures for Q. robur are undertaken in the Maltese Islands. 

However, acorns of this oak are collected at the beginning of autumn, and sown in normal 

calcareous soil immediately after collecting. They are usually planted out 2-3 years after 

germination, or at a size of about 50 cm. 

With respect to in situ conservation measures, all oaks at an age of more than 200 years 

are fully legally protected via the Antiquities Act of 1925. Apart from this, legal protection 

of both Q. robur and Q. ilex has been proposed in the Tree and Forest Protection Regulations 

1998; analogously, all trees occurring in the holm oak forest remnants (four in all) and 

Buskett have been proposed as Woodland Nature Reserves in the same regulations. 

Departments and agencies responsible for genetic resources 

The following departments are involved. Work relevant to genetic resources conservation 

and utilization is included. 

Environment Protection Department 

• Environmental legislation and its enforcement. 

• Production of environmental education material. 

• Formulation of management plans for protected areas. 

• Monitoring of local flora and fauna, including oaks. 

• Producing strategic countryside policies and guidelines. 

• Providing expert advice on conservation issues.

iZ6 

EWE0R6EN: S0el~1t1i BR0~[)It1iE~MES 

Environmental Management Unit (EMU) 

Within the Planning Directorate in the Planning Authority (PA) 

.. Countryside and nature conservation. 

.. Scheduling of areas or sites of ecological and/ or scientific importance. 

.. Issuing of Conservation Orders and Tree Preservation Orders. 

.. Providing expert advice on conservation issues. 

.. Processing applications related to development in rural areas. 

.. Coordinating the environment impact assessment process associated with development 

applications. 

Departments of Agriculture and Fisheries 

.. Soil conservation, afforestation and landscaping. 

.. Issuing of phytosanitary, veterinary and other certificates in connection with importation 

and exportation of flora and fauna. 

Department of Works 

.. Cleaning of valleys, building sites and other sites of heavy rubble. 

Police 

.. Enforcement of legislation relating to the environment and to trade. 

Customs Department 

.. Control of importation of flora and fauna and enforcement of trade regulations. 

Local Councils 

.. Each Local Council has its own jurisdiction including some environmental aspects. 

At the moment, no legislation prohibits introduction of foreign genetic stock; Q. robur is, 

however, rarely imported for afforestation and ornamental purposes, since it is rarely used 

as such. The few pedunculate oaks which are employed are usually grown from the local 

existing stock. 

Priorities 

In general, priority for in situ and ex situ conservation projects is given to indigenous 

(autochthonous) species, and in this sense Q. robur is not native. However, as stated 

previously, legal protection of this species has been proposed, also because some old trees 

(probably not older than 100 years, but no dating has been carried out) occur in the Maltese 

Islands. 

Bibliography 

Borg, J. 1927. Descriptive Flora of the Maltese Islands. Government Printing Office, Malta. 

Haslam, S.M., P.D. Sell and P.A. Wolseley. 1977. A Flora of the Maltese Islands. Malta 

University Press. 

Lanfranco, E. 1989. The Flora. Pp. 5-70 in Red Data Book for the Maltese Islands (P.J. 

Schembri and J. Sultana, eds.). Department of Information, Malta. 

Lanfranco, G.G. 1993. Hxejjex medicinali u ohrajn fil-Gzejjer Maltin. Media Centre 

Publications, Malta. 

Schembri, P.J. and J. Sultana. 1989. Red Data Book for the Maltese Islands. Department of 

Information, Malta.

- e()UNl11RM REB()Rl11S "ltl 

Management and conservation of beech (Fagus sylvatica L.) and oak 

(Quercus petraea (MaU.) Liebl., Q. robur L.) genetic resources in France 

Eric Teissier du erOSl, Isabelle Bilgel, Alexis Ducousso 3 and Antoine Kremer 3 

1 Unite de Recherches Forestieres Mediterranneennes, INRA, Avignon, France 

2 CEMAGREF, Domaine des Barres, Nogent sur Vernisson, France 

3 Laboratoire de Genetique et d'Amelioration des Arbres Forestiers, INRA, Gazinet-Cestas 

Cedex, France 

Occurrence and origin 

France has seven native oak species. The main species are pedunculate oak (2.1 million ha) 

and sessile oak (1.51 million ha), which represent about 30% of the total forest area. 

Temperate oak forests have a very large ecological niche from podzolic to calcareous soils, 

and occur from sea level to 1750 m. They are very common in France except in the 

Mediterranean region and Corsica. 

In French forests, beech is a native species in the major part of its range. It is the second 

most important broadleaved species after oaks. It covers (in pure and mixed stands) over 1.7 

million ha (12% of the total forest area). It is a species of low to high elevation (0-1600 m) as 

long as the water supply is at least 800 mm rainfall per year. 


The average annual timber production is 2.25 m 3 /ha for pedunculate oak and 2.4 m 3 /ha for 

sessile oak. The total felling of oak wood is 3215000 m 3 /year. Prices vary according to 

quality and diameter from 70 to 2600 FRF /m 3 (10-400 ECU). 

Annual production of beech varies from 2 to 7 m 3 /ha according to site fertility. A recent 

forecast shows that beech should remain one of the most attractive broadleaves for timber 

production with current prices easily reaching 800 FRF per standard quality cubic meter 

wood under bark (120 ECU). 

Stand management 

Oaks 

About 50% of the regeneration is natural and 50% artificial. The system of registered seed 

stands and regions of provenance is applied only for sessile and pedunculate oaks. French 

foresters plant about 20 million oak seedlings per year. Even with a conversion to high 

forest started in 1830, this treatment represents only 22% (Table 1). The main types are 

coppice with standards and coppice. 

Table 1. Types of white oak forest in France 

Types Area (ha) Percentage 

High forest 921 236 22 

Coppice with standard 2 672 175 64 

Coppice 423 170 11 

Miscellaneous 140 270 3 

Total 4 156851 100 

Beech 

Most of the regeneration is natural with a variety of silvicultural treatments including high 

forest (majority), uneven-aged forest, and coppice with standards under conversion to high 

forest. These treatments also vary according to regions, topography, elevation, history and 

of course, of the stands: timber production, soil conservation, protection of stands at the 

upper tree line, amenity, etc. But part of the regeneration has been, and still is, done by 

artificial regeneration. Up to 3000 ha/year were planted in the mid-1970s, when beech was


highly ranked among species used for reforestation for its relative ecological plasticity and 

when foresters had to face severe failures in natural regeneration. 

Threats to oaks and beech genetic diversity 

Oaks 

The genetic resources of oaks are not endangered in France but marginal situations exist 

such as in the Mediterranean region and Corsica, or in extreme ecological situations (sand 

dune, peat bogs). In these cases, the populations of sessile and pedunculate oaks are very 

scattered and very small. They very often have only a few tens of individuals. The other 

threatening factor is the introduction of foreign origins. 

Beech 

In spite of a decline in certain regions (Teissier du Cros et al. 1993) beech resources are not 

endangered in France. Unfortunately several mistakes were made during the 1970s when 

beech was planted with little or no consideration of the origin/provenance of the material. 

Three measures resulted to counteract this 'wild' reforestation: (1) selected seed stands, (2) 

provenance testing, and (3) in situ conservation. 

Use of forest reproductive material 

Following OECD and EU rules, CEMAGREF has established a network of selected seed 

stands for most forest tree species used for reforestation. Their main purpose is wood 

production. For pedunculate oak, 143 stands have been selected in 10 regions of provenance 

and for sessile oak 132 stands in 15 regions of provenance (Table 2). They represent 

3512.84 ha for pedunculate oak and 10799 ha for sessile oak. 

For beech, 183 stands have been selected and grouped in 20 regions of provenance (Fig. 1 

and Table 3). They represent 10 000 ha. All the seed used for reforestation is collected there. 

CEMAGREF with the help of INRA has also released recommendations for the possible 

transfer of beech reproductive material between regions. A golden rule was stressed: when 

possible, give "absolute priority to the local material" (CEMAGREF 1991). 


Provenance and progeny testing 

Oaks 

A provenance experiment has been established in France which comprises four plantations. 

Their main characteristics and locations are given in Table 4. The tested populations come 

from the whole natural range of Quercus petraea. Sessile oak is represented by 108 

populations and pedunculate oak by 17 populations (Table 5). 

Beech 

In 1974, when provenance testing was started in France, the aim was to identify adequate 

reproductive material and to determine what transfer between regions was possible without 

loss of adaptability, resistance, yield and quality (Table 6). Several results concerning 

adaptation, growth and form (morphology) have been published (Teissier du Cros 1993; 

Teissier du Cros et al. 1980; Teissier du Cros and Lepoutre 1983; Teissier du Cros and 

Thiebaut 1988). First recommendations, essentially based on bud bursting date and early 

growth, have been provided to practitioners.

, , e0UNTR¥ REP0RTS 79 

Table 2. Regions of provenance and selected seed stands of oak (1 January 1997) 

a. Sessile oak 

Regions of ~rovenance 

Registered stands 

No. Name Number Total area (ha) 

01 Secteur Ligerien 9 3089.24 

02 Charentes-Poitou 8 591.48 

03 Picardie 3 217.19 

04 Sud du Bassin Parisien 8 935.16 

05 Centre Sud 7 1112.61 

06 Allier 11 1072.09 

07 Nord-Est greseux 18 598.84 

08 Vallee de la Sa6ne 4 76.29 

09 Est du Bassin Parisien 11 375.01 

10 Morvan-Nivernais 7 477.61 

11 Nord-Est Limons et argiles 29 613.11 

12 Bretagne 2 38.08 

13 Sud du Massif Central 5 101.69 

14 Quest du Bassin Parisien 6 1489.94 

15 Gascogne 4 51.20 

Total 132 10799.54 

b. Pedunculate oak 




01 Bourgogne 23 1077.77 

02 Plateaux du Nord-Est 19 797.88 

03 Nord 3 248.92 

04 Vallee du Rhin 6 84.54 

05 Sud-Quest vallees 35 488.96 

06 Loire moyenne 8 77.40 

07 Quest 12 124.77 

08 Bassin superieur de la Sa6ne 28 572.90 

09 Sud-Quest hors vallee 2 15.76 

10 Quest Massif Central 7 23.94 

Total 143 3512.84 

Table 3. Regions of provenance and selected seed stands of beech 




01 Perche 4 348 

02 Bordure Manche 11 2192 

03 Picardie 4 1134 

04 Nord-Est (limestone) 41 3042 

05 Nord-Est (acid) 8 206 

06 Nord-Est Massif Central 2 197 

07 Sud Massif Central 18 354 

08 Bretagne 2 87 

09 Bassin Superieur de la Sa6ne 22 890 

10 Charentes-Poitou 2 147 

11 Plateaux du Jura 17 375 

12 Auvergne 12 227 

13 Pyrenees Centrales 13 512 

14 Argonne 4 63 

15 Quest Massif Central 2 15 

16 Prealpes 2 19 

17 Est Massif Central 2 5 

18 Alsace-Sudgau 8 201 

19 Pyrenees Qrientales 2 77 

20 Pyrenees Qccidentales 7 161


o selected 

seed stands 

* 

conservation stands 

representative of the main regions of provenance 

• stands in particular conditions (range limit, timber line, ... ) 

100 km I.B. f 10-97 

Fig. 1. Regions of provenance, selected seed stands of beech and the in situ conservation network. 

Table 4. Location and main characteristics of the provenance test 

Soil 

Site Location, region Type Texture Climate 

Petite Charnie Pays de Loire, West brown forest soil sand and silt or silt and clay Atlantic 

Vierzon Centre, Centre podzol sand dry Atlantic 

Vincence Bourgogne, Centre brown forest soil silt or silt and clay cold Atlantic 

Sillegn~ Lorraine, North-East brown forest soil silt or silt and c1a~ continental


Table 5. Origins of the populations tested in the provenance experiment 

Country Species Number of populations 

France Quercus petraea 66 

Quercus robur 4 

Great Britain 

Germany 

Austria 

Denmark 

Poland 

810vakia 

Hungary 

Romania 

Russia 

Turkey 

Total 


Quercus robur 


Quercus robur 




Quercus robur 


Quercus robur 


Quercus robur 





Quercus robur 

All species 

Table 6. French provenance and progeny testing network for beech 

Planting 

Planting density Number of entries 

Series Location year (trees/ha} F = French, E = "Exotic" 

1976 Ecouves (W France) 1979 5000 14 provenances F 

Chaud (Centre) 1979 5000 21 provenances F 

80mmedieue (NE) 1979 5000 11 provenances F 

Montagne Noire (8) 1978 5000 12 provenances F 

1977 Ligny en Barrois (NE) 1980 5000 6 provenances F 

1979 Ligny en Barrois (NE) 1984 4000 30 provenances FE 

Plachet (NE) 1982 5000 22 provenances FE 

Ormancey (NE) 1982 5000 38 provenances FE 

Retz (N) 1983 5000 49 provenances FE 

Lyons (NW) 1983 5000 40 provenances FE 

Guimont (Centre) 1982 5000 26 provenances FE 

International Lyons (NW) 1986 10000 24 provenances FE 



Progeny tests Eawy (NW) 1997 3900 51 open-pollinated progenies 

Ha~e {NE} 1997 2600 48 open-pollinated progenies 

3 

3 

17 

2 

2 

3 

6 

4 

1 

3 

1 

1 

2 

108 

17 

125 

Progeny tests 

A plantation programme was started this year. The project concerns two experimental tests 

(N.F. La Petite Charnie and N.F. Russy). The experimental design for the test at La Petite 

Charnie involves two species (Q. petraea and Q. robur), 30 progenies per species, 30 

individuals per progeny and each individual is propagated 12 times. For the second test, the 

experimental design is more conventional: 30 progenies of Q. petraea, 120 individuals per 

progeny.

82 EUF()RGEN: S()CIAL BR()ADLEAVES - 

Progeny testing aims at estimating the level of genetic control of traits used when 

selecting 'seed trees' for natural regeneration. A sound use of this knowledge should result 

in a 'mild', long-term genetic improvement of naturally regenerated stands. For beech, two 

open-pollinated progeny tests, including copies of the mother trees, were established in 1997 

(Table 6). They allow the estimation of several genetic parameters such as broad and narrow 

sense heritability, genetic correlation between traits and juvenile x mature correlation. 

Beech 

Conservation of genetic resources 

Current situation 

In France, beech is not endangered as such. But the integrity of certain stands has probably 

been disturbed by 'wild' planting of reproductive material either of unknown origin or from 

other parts of the range. The long-term effect of such a practice which has now fortunately 

been stopped is (1) lack of adaptation (for instance to local climatic extremes) and (2) genetic 

pollution of local sources. Beech is theiirst broadleaved species in France for which an in 

situ conservation network has been established. The underlying principles at the time of 

creation of this network were based on pragmatism: conservation stands were to be 

representative of a majority of usual beech ecological conditions but some of them were also 

to originate from marginal sites. Furthermore, since in situ conservation of forest genetic 

resources is a long-term process, public forests should be involved as much as possible. The 

current conservation network includes 27 stands (Fig. 1). Twenty of them have been chosen 

in the main regions of provenance representing the majority of beech ecotypes. Seven 

represent rare conditions or phenotypes such as: protection stands near upper tree line, 

Mediterranean conditions or the population with curly branching phenotype. Each element 

of this network is composed of a central core covering at least 5 ha (an effective genetic 

population size of at least 300 trees) and a minimum 50-m-wide buffer to protect the core 

from allogenic pollen. Of course consistent administrative sources should be given to certify 

that the core and the buffer were of local origin (Teissier du Cros and Bilger 1995). 

To convert the pragmatism initially used to establish this network into a scientifically 

based sampling, complementary studies were done by B. Comps, University of Bordeaux- 

Talence. Each element (stand) of the conservation network has been characterized by 

isoenzyme markers and compared with neighbouring stands. Results were compiled with 

those of other scientists working in other parts of the range. French beech populations could 

be divided into four main groups: (1) northern half of France, (2) centre: Massif Central 

mountains, (3) Alps, (4) Pyrenees mountains. The curly branching population cannot be 

distinguished from other populations in the vicinity (northern half of France). One southern 

population is genetically similar to the northern population. Two important additions will 

be made in the near future to the current network. Both the Alpine and the Pyre ne an parts 

of the range appear under-represented (Teissier du Cros 1997): 

• the former because the French Alps are a melting pot of two colonizing currents after 

the last glaciations: the northern current originating from the Balkans, the southern 

one from south Italy 

• the latter because the Pyrenees mountains include a much greater diversity than 

originally expected. 

Management of the conservation network 

In France, because the beech conservation network was the first to be established, beech is 

used as a pilot species for the management of in situ conservation units. But the 

management recommendations given to foresters in charge have sometimes seemed 

inadequate for local site conditions or stand evolution. For instance, natural regeneration is 

sometimes scanty because seed trees are too old, declining or too dense, and therefore do not 

produce seed. If such a lack of regeneration occurs in the core, at least two solutions should

· . COUNTRY REPORTS 83 

be proposed: (1) moving the core to more favourable plots of the same conservation unit, (2) 

collecting beechnuts in buffer plots to plant seedlings in the core. The consequence of such 

measures during the life cycle of the conservation units needs to be carefully analyzed. 

Another important point is the compatibility of gene conservation with stand 

management. The current trend is to promote diversity and therefore to reduce the 

'beech/total tree density' ratio, which incidentally is likely to favour stand health and 

natural regeneration. Conservation recommendations may have to consider this point and 

to become more flexible. 

The different institutions involved in the conservation of beech gene resources are INRA 

(research and network coordination), Bordeaux University (research), Office National des 

Fon~ts in management of conservation stands (State and Community forests). 

Oaks 

Up to now oak genetic resources are not really endangered in France, except in some situations 

such as marginal populations (coastal sand dunes, peat bogs, altitude higher than 1400 m) and 

in the southeast of France and Corsica. Several problems can threaten these resources such as 

introduction of exotic genotypes, neglected practices, conversion to high forest, etc. For these 

reasons, the committee for the conservation of forest gene resources (Commission technique 

nationale) has launched a programme of gene conservation for oaks with four objectives: 

1. Sampling the gene diversity: 20 populations are selected for this objective. The 

sampling strategy has been defined according to the results obtained with molecular 

and quantitative markers (Ducousso et al. 1995, 1996c; Petit et al. 1993; Zanetto et al. 

1994; Zanetto and Kremer 1995, 1997). The list is given in Table 7. 

2. Conservation of evolutionary mechanisms: white oaks have the highest observed 

genetic diversity; this results from evolutionary mechanisms like interspecific 

hybridization. Several mixed stands will be used for this network (Bacilieri et al. 1993, 

1995, 1996; Bodenes 1996; Ducousso et al. 1996c). 

3. Conservation of oak ecosystems: ecotypes adapted to different types of management 

for wood production (high forest, coppice with standards, coppice) and for acorn 

crops (provender for cattle). Most of these types are neglected because the foresters 

have undertaken a conversion to high forest. The objective will be included in the 

'reference forest network' managed by the French National Forest Service (Office 

National des Fon~ts). 

4. Conservation of marginal or endangered populations requires actions. The first step 

of this objective is to take a census, and then to define a policy for each situation. 

Each reserve designated according to objectives 1, 2 and 3 is composed of a central core 

covering 15 ha surrounded by a buffer zone of at least 100 ha. The central core and the 

buffer must have a local origin. This origin is known by administrative records and by using 

chloroplast DNA. All the populations belong to the French provenance experiments. 

Capacities for the conservation of forest gene resources 

A committee for the conservation of forest gene resources (Commission Nationale de 

Conservation des Ressources Genetiques Forestieres) was formalized in 1992. It has 

proposed four pilot networks involving beech and silver fir for in situ conservation, elm and 

wild cherry for ex situ conservation. These networks are almost all complete. The committee 

and the institutions involved have sponsored research to establish five other conservation 

networks: European white oaks, black poplar, Norway spruce, Maritime pine and Sorbus 

species. Methodological research is under way for ex situ static conservation 

(cryopreservation) and for the interaction between human activity on the one hand, 

including stand management, and gene resource integrity on the other. Oak populations 

will essentially be used to assess this interaction.

84 EUE'(i)RGEN: S(i)GI~1.l! BR(i)~DI.l!E~MES 

Table 7. List of the candidate populations for objective 1 (Mallance and Ducousso 1997) 

National Forest Region Location 

Bareille Midi Pyrenees Southwest 

Berce Pays de Loire West 

Bommiers Centre Centre 

Bussieres Champagne Ardennes Parisian Basin 

Compiegne Picardie North 

Fontainebleau Region Parisienne Parisian basin 

Gresigne Midi Pyrenees Southwest 

Hagueneau Alsace Northeast 

Loche Centre Centre 

Orleans Centre Centre 

Premery Bourgogne Centre 

Reno Valdieu Normandie Northwest 

Saint Aubin du Cormier Bretagne West 

Serqueux Champagne Ardennes Parisian Basin 

Sturzelbronn Lorraine Northeast 

Tronc;::ais Auvergne Centre 

Vacheres Provence Alpes Cotes d'Azur Southeast 

Valbonne Provence Alpes Cotes d'Azur Southeast 

Vouille Poitou Southwest 

Westhoffen Alsace Northeast 

Needs for international cooperation 

In its natural range beech plays an important role in forestry. It has many of the 'advantages' 

which the 'Greens' wish to promote: it is autochthonous, broadleaved, rather flexible 

(provided the right reproductive material is chosen), it produces timber whose attractiveness 

for industry is increasing. For instance, it is one of the species which could replace certain 

tropical hardwoods for veneer and high-quality sawn timber if the former were to become 

scarce on the international market. It is excellent in many situations where conifer stands, 

extensively planted since the last world war, need to be converted into to more adapted 

stands devoted to the long-term sustainable production of high-value raw material for 

industry, amenity and soil conservation. 

France has been involved since 1985 in the international diversity study on beech, which 

covers almost the entire range of the species. This study, sponsored by the European Union 

and coordinated by the Federal Forest Research Center in Germany, includes provenance 

testing and different types of genetic markers, either physiological or molecular. France has 

also cooperated with other European countries, particularly Slovakia, for the description of 

beech genetic diversity throughout its distribution range. 

Any other proposal for international research and development is welcome for a common 

scientific approach on beech and oak genetic resources conservation, and for increasing the 

capacity of institutions requesting it. 

Bibliography 

Bacilieri, R, A. Ducousso and A. Kremer. 1995. Genetic, morphologic at ecological and 

phenological differentiation between Quercus petraea (Matt.) Liebl. and Quercus robur L. in 

a mixed stand of northwest of France. Silvae Genet. 44:1-10. 

Bacilieri, R, A. Ducousso and A. Kremer. 1996. Comparison of morphological characters and 

moleculars markers for the analysis of hybridation in sessile and pedunculate oak. Ann. 

Sci. For. 53:79-9l. 

Bacilieri, R, G. Roussel and A. Ducousso. 1993. Hybridization and mating system in a mixed 

stand of sessile and pedunculate oak. Ann. Sci. For. 50(1):122-127. 

Bodenes, C. 1996. Differenciation moleculaire entre le chene sessile (Quercus petraea (Matt.) 

Liebl. et le chene pedoncule (Quercus robur L.). These de l'Universite de Bordeaux 1.

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Conseils d'utilisation. Nogent sur Vernisson, France. Dix-neuf fiches d'especes. 

Ducousso, A, R Bacilieri, B. Demesure, R Petit, A Zanetto and A Kremer. 1997. Dernieres 

donnees sur la genetique du chene sessile: la notion de region de provenances et de 

peuplements classes. Bull. Tech. O.N.F. 33:7-19. 

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europeens. Foret-Entreprise 112:49-56. 

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burst in western populations of sessile oak (Quercus petraea (Matt.) Liebl.). Ann. Sci. For. 

53:775-782. 

Ducousso, A, R. Petit and A 

Kremer. 1995. Proposition pour creer un reseau de 

conservation des ressources genetiques des chenes blancs europeens. Report for 

'Commission Nationale de Conservation des Ressources Genetiques Forestieres'. 8 p. 

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ressources genetiques des chenes blancs europeens: premieres propositions pratiques. 

Report for 'Commission Nationale de Conservation des Ressources Genetiques 

Forestieres'. 15 p. 

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polymorphisms in European oaks. Theor. Appl. Genet. 87:122-128. 

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Workshop, Ahrensburg (H.-J. Muhs and G. von Wuehlisch, eds.). Working Document of 

the EC, DG VI, Brussels. 

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forestieres. Convention DERF-INRA nO 61.21.06/96. INRA B01267. Compte rendu a 6 

mois.11p. 

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symposium 1994, Denmark (S. Madsen, ed.). Forskningsserien no. 11-1995. Danish Forest 

and Landscape Institute, Horsholm, Denmark. 

Teissier du Cros, E., J. Kleinschmit, P. Azoeuf and R Hoslin. 1980. Spiral grain in beech. 

Variability and heredity. Silvae Genet. 29(1): 5-13. 

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sylvatica L.). For. Sci. 29(2):403-411. 

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VI, Brussels. 

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tree form. Ann. Sci. For. 45(4):383-398. 

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'Commission Nationale de Conservation des Ressources Genetiques Forestieres'. 5 p. 

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(Matt.) Liebl. 1. Monolocus patterns of variation. Heredity 75:506-517. 


(Matt.) Liebl. H. Multilocus patterns of variation. Heredity 78:476-489. 

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1(2):111-123.

86 EUFORGEN: SOCIAl.. BROADI..EAVES 

Oak and beech genetic resources in Luxembourg 

M. Wagner1, F. WoIter1 and Jean-Fram;ois Hausman 2 

1 Administration des Eaux et Fon~ts, Service de l'Amenagement des bois et de l'economie 

forestiere, Luxembourg 

2 Centre de Recherche Public-Centre Universitaire, CREBS Research Unit, Luxembourg 

Occurrence and origin of oaks and beech in Luxembourg 

The total forest area in Luxembourg is about 90000 ha which represents 33% of the total 

land area. The country is divided into four main geological and climatic regions: 

• Oesling (hills, 300-500 m, acid slate soils (Devon» 

• Gutland (plains on heavy clay soils (200 m) and sandy hills (300 m» 

• Moselle (shell-limestone valley) 

• Minette (limestone hills, 300 m). 

Oak(Quercus robur, Quercus petraea) 

Origin: 

Q. robur: indigenous; and from Westdeutsches Bergland, Plateau du Nord-Est France 

Q. petraea: indigenous; Rheinisches and Saarbergland, Nord-Est France, Bassin Parisien 

Est 

Occurrence: 

All over the country 

13 000 ha coppice mainly in the northern region Oesling 

12 000 ha even-aged high forest with more than 60% from 100 to 180 years old. 

Beech (Fagus sylvatica) 

Origin: 

Indigenous; Rheinisches and Saarpfalzer Bergland, Harz, Hessen and Weser hills, Nord- 

Est calcaire France, Vosges du Nord, Belgium (Ardennes, Foret de Soignes, Arlon) 

Occurrence: 

25000 ha even-aged high forest with more than 70% from 100 to 140 years old, all over 

the country, mainly on hills (300 m) in the Gutland and Minette regions. 


The economic importance of oak and beech stands in Luxembourg relates to three sectors: 

wood production, recreation and nature protection (drinking water). With an overall yearly 

wood production of about 300000 m 3 , oak (40 000 m 3 ) and beech (150 000 m 3 ) represent more 

than 60% of the total production. A major part of this production is exported for further 

processing. 


Except for the old oak coppice stands in the northern part of the country which are now 

being transformed into high forest stands or replanted with conifers and broadleaves, almost 

all oak and beech stands are even-aged high forests. 

Most beech stands have a small amount of oak (less than 20%) admixed to stabilize the 

stands, especially on sandy hills. Natural regeneration of beech is possible, but difficult on 

sandy soils. After regeneration or plantation (5000 plants/ha) and cleaning, the stands are 

thinned regularly (5-10 years) until small-scale clear-cutting (maximum 2 ha) or progressive 

clear-cutting at about 140 years is done to obtain natural regeneration.


Many even-aged high forest oak stands have originated from coppice with standards, so 

that stem quality is often rather poor. Natural regeneration is the rule, with a mean thinning 

interval of 5-15 years and a mean rotation period of about 180 years. 

Health state of the forest stands 

Since 1986, the health state (defoliation and decoloration) of forest stands in Luxembourg 

has been monitored by a yearly inventory of plots within a 4x4 km grid on the basis of the 

European ICP forest standards. From 1986 to the present, the health state has been 

constantly decreasing, so that for the moment, not more than one-third of the trees can be 

considered as absolutely healthy. In beech, 15% of trees are without damage (class a), 52% 

are slightly damaged (class 1) and 33% are damaged (classes 2, 3 and 4). In oak high forest, 

the respective percentages are 22, 32 and 46% and in oak coppice stands 11, 36 and 52%. 

The poor health state is mainly due to a lack of rainfall for 10 years and more recently 

because of massive attacks by insects over the last 2 years. 


The CREBS Research Unit of the Centre de Recherche Public-Centre Universitaire is 

involved in two research programmes on genetic resources of forest trees. The first project 

focuses on the conservation ex situ of oaks and beech as well as of Noble Hardwoods. This 

conservation is realized using in vitro techniques. Up to now Quercus robur, Q. petraea, Fagus 

sylvatica, Fraxinus excelsior, Prunus avium and Sorbus domestica have been initiated and are 

multiplied in vitro. Mid-term conservation will be realized and the further viability, as well 

as the rooting and acclimatization performance, will be tested. The second research project 

deals with the development of techniques leading to the characterization of the genetic 

diversity of broadleaves. The techniques that will be used are mainly based on molecular 

biology. 


• 17 nature protection areas (1862 ha). 

• Protection of 16 beech and oak stands for collecting of reproductive material (180 ha). 

• Selection of areas for the network 2000 of the "Directive HABITAT" (EU). 


Tree-improvement activities are limited to phenotypic selection during seed collecting, 

propagation of forest trees and silvicultural operations (cleaning, thinning, etc.). 


Information on the private forest sector is not available. The Forestry Administration has 

four main tree nurseries producing from 500000 to 1000000 plants per year, covering 

approximately 50% of the plant demand of the forests managed by the Administration. The 

seeds are mainly imported or collected in the protected stands in Luxembourg. 

Institutions involved in genetic resources activities in Luxembourg 

• Administration des Eaux et Fon~ts (Forestry Administration), Service de l'Amenagement 

des bois et de l't:§conomie forestiere, BP 411, L-2014 Luxembourg. 

• Ministry of Environment, 18, Montee de la Petrusse, L-2918 Luxembourg. 

• Centre de Recherche Public-Centre Universitaire, CREBS Research Unit, 162a, av. de la 

Fai'encerie, L-1511 Luxembourg.

88 EUEQRGEN: SQGI~1l: BRQ~DIl:E~VES 

Genetic conservation strategy for Social Broadleaves in Belgium 

Dominique J acques 1 and Bart de Cuyper2 

1 Station de Recherches Forestieres, Gembloux, Belgium 

2 Institute for Forestry and Game Management, Hoeilaart, Belgium 

Introduction 

Belgian forests cover around 600000 ha, mainly concentrated in the southern part, Wallonia 

(80%). With 140000 ha, beech and oaks are the most important broadleaves. Natural 

regeneration is commonly used but artificial plantations have become more and more 

frequent since the 1980s. 

For a good comprehension of the Belgian conservation strategy, it is important to know 

that since 1981, the National Forest Service, followed by the Forest Research Institutes eight 

years later, were regionalized with the creation of a federal structure. This situation 

involves specific policies regarding forest management and conservation objectives followed 

by the Regions. 

Conservation strategy in Wallonia 

Present situation 

Indigenous oaks (Quercus petraea (Matt.) Liebl. and Quercus robur L.) and beech (Fagus 

sylvatica L.) represent the major broadleaved species of Wallonia. Considering the regional 

inventory, the growing stock can be estimated at 13 and 19 million m 3 respectively for beech 

and oaks, with 8 and 18% respectively of a total productive forest area of 473 750 ha 

(Table 1). Oaks and beech represent, respectively, the second and third species in 

importance in Wallonia after Norway spruce. 

Except for High Ardenne, the oaks are potentially well adapted to Wallonia. On the other 

hand, the potential range for beech covers the whole region. 

At the economic level, beech and oaks constitute the main part (>80%) of the annual 

production of hardwoods which can be estimated at about 860 000 m 3 of wood. 

Based on this evaluation, primary income from hardwoods could be roughly estimated at 

1 million BEF. 

Threats 

Since the 1980s, different signs of decline in oaks and beech have appeared (Table 2). A 

combination of factors seems to be involved. Claes (1997) cites in particular drought, fungi 

and insects (mainly caterpillar). Weissen (1996), on the other hand, stresses the influence of 

mineral elements deficiency, like magnesium shortage. 

This disquieting situation should be taken into account for the elaboration of a general 

conservation strategy. 

Another problem is the lack of natural regeneration due to two main factors. The first is 

the long period between good fructification years; this observation is particularly true for 

oaks. The second is the very high pressure of game (red deer, roe deer and wild boar), 

whose numbers have increased regularly since 1985 (Table 3). 

The third threat is using material of foreign origin for the artificial plantations which are 

not necessarily adapted to local ecological conditions. The difficulties linked to the low rate 

of natural regeneration increase this risk. 

Genetic conservation strategy 

In Wallonia, the conservation programme is integrated in the general breeding programme 

and has been progressively developed for some years. 

The classical steps of our general forest tree breeding programme are presented below.

C()UNTR¥ RER()RTS 89 

Table 1. Importance of oaks and beech in Wallonia 

Forest High forest Coppice with Coppice Total 

management Species {haf standards {haf {halt {halt 

Private owners Oaks 22250 13000 2250 37500 

Beech 10250 750 11 000 

Forest under public Oaks 27000 17750 1 250 46000 

control Beech 27750 500 28250 

Total Oaks 49250 30750 3500 83500 

Beech 38000 1 250 39250 

Surlace occupied by more than 66% in sectional area by the concerned species. 

Table 2. Decline of oaks and beech in Wallonia 

Rercentage of defoliation 

Species 1993 1994 1995 

Beech 10.8 10.7 12.2 

Sessile oak 7.3 9.2 9.1 

Pedunculate oak 8.7 9.1 12.7 

Table 3. Numbers of big game, after hunting and before births (Anon. 1996) 

Species 1975 1985 1994 1994/1975 

Red deer 5 144 4830 8095 +57% 

Roe deer 19 504 22 300 31 338 +61 % 

Wild boar 8 484 6 348 12 609 +49% 

Conservation in situ 

Seed stands 

For the last 5 years, an important effort has been made to increase the number of seed stands 

of different hardwood species. Today, the results are sufficient for beech to meet the needs 

of foresters. That is not yet the case for indigenous oaks where some additional surveys and 

selection have to be carried out (Table 4). These selections nevertheless are not directly 

linked to a general conservation purpose but are mainly done to ensure good timber 

production potential for the future. 

Individual selection 

Although a small number of plus trees were selected in the 1950s, mainly to study genetic 

variability, this work of selection was given up later in favour of a more adequate genetic 

improvement programme. In fact, a clonal seed orchard with oaks and beech seems to be 

rather ineffective. Therefore, other components such as seed stands or seedling orchards 

have to be considered. 


Besides the general genetic improvement programme, the concept of forest reserves has been 

developed since 1973. Today, eight forest reserves for a total of 244 ha have been registered. 

They generally comprise special ecological sites including beech and oaks. 

Conservation ex situ 

Provenance/progeny trials 

In the 1950s, different tests were established to study genetic variability in beech at different 

levels (individual, population, ecological type, provenance) and were analyzed by Galoux 

(1966) and Hubert (1988). These tests, mainly limited to Belgian populations, completed by

90 EUFORGEN: SOCIAL BROADLEAVES . 

Table 4. Present situation of the genetic conservation of Social Broadleaves 

Quercus Quercus Fagus 

Type of gene conservation unit petraea robur sylvatica Total 

In situ 

Seed stands 

Number 4 6 19 29 

Area (ha) 47 121 552 720 

Plus trees 21 23 70 114 

Ex situ 


Number 1 2 3 

Area (ha) 3.8 2.5 6.3 

Provenance/Progeny tests 

Number 2 4 6 

Area (ha) 1.3 2.5 3.8 

observations in natural forests, show an important variability between populations for 

different characteristics like flushing, morphology of leaves and growth. 

In addition, Belgium took part in an international provenance trial in 1988 from which 

one site was established in Paliseul and where 74 provenances are compared. 

These different trials should give us more basic information to elaborate a complete longterm 

conservation programme. 

Prospects for further conservation and priorities 

A general programme adapted to all tree species has been outlined and partly completed. 

The main aspects of this programme (Nanson 1993, 1995; de Cuyper and Jacques 1996) are 

described hereunder. 

Promoting the use of highly diversified reproductive material 

With the regional grants delivered for hardwoods to private and public owners and 

considering the problem of natural regeneration, the share of Social Broadleaves plantations 

has been increasing for several years. 

To favour the use of diversified autochthonous seed sources, the Walloon Region has 

created a new seed centre. This centre has been operational since 1996 and is in charge of 

forest seed harvesting complying with strict genetic rules like t~e respect of a minimum 

number of trees collected in each stand. 

In parallel, selection of new seed stands is pursued. 

Active conservation plantations of the registered seed stands 

Up to now, ex situ conservation was included in the general genetic improvement 

programme through conservation of particular origins in provenance or progeny trials. 

Nowadays, through collaboration between the regional forest administration, nurseries~ 

seed centre and research institute, it is possible to set up an integrated programme to 

organize conservation of the major part of our officially registered seed stands in the long 

term. This programme should be implemented in the near future. 

Trace-keeping in forest management and data storage 

In the field of forest management, general rules are applied for all the Walloon public 

forests. Numerous data are stored in databases that allow us to know the method of 

regeneration and the name of the origins used in any artificial plantation at management 

unit level. These data and other additional information could be stored afterwards in a 

specific database at the regional level.


A new database developed with Germany for Douglas Fir within an EU Research 

Contract (No. CT95-09091 - EUDIREC) could probably also be used for indigenous oaks and 

beech in the future. 

More details can be found in Nanson (1993,1995) or de Cuyper and Jacques (1996). 

Improvement of knowledge about forest genetic resources 

As for Noble Hardwoods, a better knowledge of the genetic structure of populations is 

useful to put in place an efficient genetic conservation programme. 

With the potential offered by new biochemical techniques (isoenzymes, DNA markers) 

and the growing interest of many organizations, it seems possible to propose a 

comprehensive programme to explore genetic diversity in the natural range. 

In this field of activity, provenance trials are a step of major importance to evaluate 

adaptive traits, growth and form characteristics. These tests should be promoted and 

carefully evaluated. 

Improvement of seed conservation techniques 

Acorns, and beechnuts to a certain extent, are known as seeds difficult to store for a long 

period. The rarity of crops (10-12 years between two good fructification years in Belgium for 

oaks), associated with the problem of conservation, forces nursery owners to buy seeds 

harvested in more favourable regions where frequent fructifications occur. 

The consequences of this situation could be very detrimental to forest owners by reducing 

adaptability and performances of the tree species in the future. 

To avoid these problems, long-term storage techniques should be developed to reduce the 

lack of seeds between two good crops by increasing research in this field. 

Conservation strategy in Flanders 

Present situation 

Indigenous oaks (Q. robur and Q. petraea) and beech (Fagus sylvatica) are to be regarded as 

major broadleaved species in Flanders, as they cover 8 and 3.5% respectively of the total 

forest area of 150 000 ha. 

The available information on the current importance of Social Broadleaves in Flanders is 

still incomplete (Table 5). As an example, figures concerning the occurrence of oaks and 

beech in the different types of forest (coppice, high forest, coppice with standards) are not 

yet available. 

More accurate data are expected in 1998, as an overall forest inventory will be initiated by 

the Flemish Forest Service. This inventory is to be based on some 2500 circular sampling 

plots of about 10 ares each, centred at the intersections of a 2x1.5 km grid. 

Threats 

Forest dieback 

Since 1987 an annual survey of the vitality of forests in Flanders has been carried out by the 

Institute for Forestry and Game Management (IFG). Health condition is assessed by estimating 

premature leaf loss. Trees are considered to be damaged when leaf loss exceeds 25%. 

Observations reveal a fast decline of the vitality of pedunculate oak and beech, 

culminating in an alarming situation in 1995 when respectively 42.5 and 44.4% of the trees 

were to be regarded as damaged (Muller-Edzards 1997). Fortunately, during the last 2 years, 

the health condition of both species has seemed to stabilize and, in some cases, even to 

improve. 

Regarding the cause of this dieback, no unequivocal explanation can yet be given. It is 

generally assumed that the simultaneous occurrence of stress factors (repeated years with 

extreme drought, insect damage) has led to a severe weakening of the trees which then 

become susceptible to infection by secondary parasites.

92 ELJEQRGEN: SQCIA~ BRQAD~EAMES . » 

Table 5. Area (ha) of Social Broadleaves in Flanders 

Forest ownership Oaks Beech Total 

State Forests 1529 2565 4094 

Public Forests 1634 430 2064 

Private Forests 9032 2026 11058 

Total 12195 5021 17216 

Inconsiderate forest management 

Social Broadleaves species can be endangered by 'skimming off' natural populations by 

excessive felling of trees with high economic value. This genetic erosion by eliminating the 

best genotypes heavily burdens the quality and vitality of future stand generations. 

Trace-keeping 

Any genetic resource of Social Broadleaves, identified, selected or created within the scope 

of a breeding or conservation prograrrune, should be recorded in a well-managed, accessible 

database. Permanent and frequent updating of the database are essential to prevent 

valuable genetic elements from passing into oblivion. In addition, lack of follow-up 

inevitably entails unawareness of possible critical threats (felling, game damage, pests, etc.), 

eventually leading to accidental loss. 

Conservation strategy 

Conservation of genetic resources of Social Broadleaves has mainly been a logical spin-off of 

breeding and selection prograrrunes, driven by motivations with a strong economic bias. 

Only recently, efforts are made to conserve genetic elements (i.e. local populations) with no 

direct economic importance through actual protection (forest reserves). 

Conservation in situ 

Seed stands 

Seed stands are to be defined as stands which are phenotypically superior for most forest 

characteristics (cf. OECD/EEC regulations). These stands are selected and included in an 

official Register of Basic Materials (Table 6). 

Plus trees 

Plus trees are selected as such on the basis of their outstanding phenotype (Table 6). 

In view of a continuous extension of this collection, a scouting campaign is resumed 

every year, mainly based on an inquiry addressed to all local forest services. 

Conservation ex situ of selected clones is compulsory as plus trees are not only subject to 

the normal exploitation term, but are, in some cases, threatened by premature felling, 

especially in privately owned forests. Ex situ conservation can be achieved by creation of 

seed orchards and/ or clone collections (see below). 


The Flemish Forest Decree of 1990 provides a legal framework for deSignating forest 

reserves, aiming at a final overall surface of some 2000 ha. A major criterion for selecting 

forest reserves is the presence of autochthonous tree species. 

Sessile and pedunculate oak and beech constitute major stand-building species in 17 of 

these reserves (31 in total), covering an overall surface of 677 ha, i.e. 49% of the total area yet 

designated. 

This policy ensures the conservation of Social Broadleaves as the majority of the reserves 

is assigned the status of 'integral reserve', meaning that 'doing nothing' is adopted as a 

management option, except for averting external threats.

G0UNmRY RER0RmS 93 

Conservation ex situ 

Provenance trials 

The principal and initial objective for setting up provenance trials is to identify provenances 

which, through their adaptation and performance (growth, form, pest resistance), can 

contribute in a significant way to forest practice in Flanders. This will finally result in the 

drawing up of a list of 'recommendable' provenances. In a further stage, the best individuals 

will be selected within the most promising provenances, thus offering the possibility of 

creating seedling seed orchards. 

A major spin-off of these experiments is the conservation of provenances outside their 

range, ensuring the possibility of restoring the original populations should they be lost. 

Since 1989, provenance trials of Social Broadleaves have been included in the research 

programme of the IFG (Table 7). 

Yearly efforts are made to broaden this spectrum, either by using the offer of reliable and 

reputable professional nurseries, or through collaboration with other foreign research 

institutes (e.g. exchange of breeding material). Whenever possible, the highest level of 

accuracy for identifying provenances will be striven for (e.g. identification at a stand level). 

Seed orchards 

A clonal seed orchard for sessile oak is being created at the IFG: scions were collected from 

50 selected plus trees and first graftings were carried out in 1995. Considering the largely 

insufficient seed crops in selected seed stands of Social Broadleaves, seed orchards might 

offer an answer to the high demand for autochthonous reproductive material. 

Although conservation of the component clones is inherent to the existence of a seed 

orchard, seed production still remains the prime purpose of its establishment. Thus, 

whenever seed productivity should prove to be insufficient, the normallifespan of such an 

orchard (50 to 100 years) could be severely shortened by either felling or mere neglect, as its 

further maintenance loses its economic justification. 

Progeny trials 

Comparative progeny trials have only been established for sessile oak, involving the half-sib 

offspring of each of the 50 selected plus trees. These trials were initiated in 1994 and will be 

repeated whenever a good seed crop occurs. 


Forest authorities strongly support the natural regeneration of, among others, oak and beech 

stands, in view of the resulting conservation of autochthonous populations. Hence, attempts 

are made, through adequate forest management, to induce natural regeneration in state and 

public forests. Promotion of natural regeneration in private forests is achieved by granting 

financial support whenever stands are successfully regenerated in a natural way (subvention 

policy). 

However, unfavourable conditions, such as high game pressure, invasive herbaceous 

vegetation and compact soil, often constitute barriers for natural regeneration. In these 

cases, artificial regeneration becomes imperative and requires high-quality, selected planting 

material. The availability of breeding material from authochthonous sources (i.e. seeds) is 

insufficient because: 

• the number of seed stands is rather limited (Table 6) 

• owing to their inherent phenology, good crop years in oak and beech stands are 

separated by long-term intervals 

• as selection and breeding efforts have been focused on broadleaves only very recently, 

seed orchards (in casu sessile oak) are still in their infancy and are far from being 

productive.

94 EUFORGEN: SOCIAL BROAD LEAVES 

Table 6. Selected seed stands and plus trees of Social Broadleaves in Flanders 

Seed stands 

Species Number Area Plus trees 

Pedunculate oak 11 63.6 8 

Sessile oak 1 5.0 50 

Beech 1 1452.6 17 

Total 14 1521.2 75 

Table 7. Provenance trials of Social Broadleaves set up by the IFG 

Species 

Sessile oak 

Beech 

Year 

1989 

1993 

1996 

Origin 

Seed stand 

Provenance region 

Seed stand 

Number 

19 

11 

32 

Countries 

B/F/D/DKlGB/H/N/PLITR t 

F/D/SLO 

D 

t International provenance trial with 24 participating countries, coordinated by the Danish Forest 

Research Station, Lyngby, Denmark. 

In this way, public as well as private nurseries are forced to appeal to foreign seed 

origins. As provenance trials have only been set out very recently, first results are not 

available yet. Therefore, adaptive the value of most of these provenances is still unknown. 

This entails a high risk for genetic pollution which increases the vulnerability of the forest 

ecosystem. 

Silvicultural approach 

Management of Social Broadleaves forests is 'close-to-nature'. In general it can be 

characterized as silviculture on an ecological basis, with attention to all components of the 

forest ecosystem, and pursuing stable, healthy forests with a durable economic and 

ecological value. Such management offers best guarantees for sustained forest use. To 

achieve this ecological management, a set of principles is observed: 

1. Short rotations are to be omitted. On average, the rotation period in oak and beech 

stands is set at 150 and 200 years, respectively. 

2. A complex and varied stand structure is striven for, characterized by individual or 

group-wise mixture of age and size classes. 

3. Damage caused by forest exploitation is reduced to a strict minimum by imposing 

regulations concerning machinery, log dimensions, hauling roads and exploitation 

period. 

4. As management must be based on self-regulating processes, natural regeneration is 

regarded as a general rule. 

5. Clear-cutting is altogether prohibited as it causes a severe disturbance of the forest 

microclimate over a long period. Felling is to be carried out on individuals or groups. 


The economic importance for the forestry sector of Social Broadleaves mainly evolves from 

their ability to produce high-quality timber, suitable for multiple purposes (furniture, 

veneer, construction, etc.), rather than from the timber volumes produced. 

The economic importance can be assessed by using the yearly timber sales as a simple 

criterion: the yearly revenue from oak and beech fellings in state forests represents some 0.18 

million and 0.87 million ECU respectively, representing 10.5 and 51.7% of the total income 

for all species. Data concerning the yearly timber sales in the remaining public and private 

forests are not readily accessible. Rough estimates could be made, taking into account that 

state-owned oak and beech forests represent 12 and 51%, respectively, of the total area of 

these species in Flanders.

COUNTR¥ REPORTS 95 


Forest Research Stations of Gembloux (FRS) and Geraardsbergen (IFG) are the two sole 

scientific organizations dealing regularly with forest genetic resources management in 

Belgium. Both are integrated in their own region (Wallonia and Flanders), in a general 

administration involved in the fields of the natural resources and the environment. With 

their seed centres, nurseries and forest areas, these two structures possess the useful tools to 

set up a rational and practical genetic conservation policy. 

Besides this main structure, different organizations (Annex 1) are indirectly involved in 

this programme. They belong to universities or to other scientific research organizations. 

These structures possess valuable knowledge or tools to complete and improve the basic 

work realized by the two Forest Research Stations. 

Conclusion 

Beech and oaks represent the major broadleaved species in Belgium. They are not really 

threatened in our country but some observations seem to indicate problems of decline. 

The conservation strategy mainly relies on the application of the general genetic 

improvement programme (provenance trials, seed stands) and on the general forestry policy 

(grants, scientific support) in favour of an optimal utilization of the forest reproductive 

material. 

Further activities could be focused on: 

.. promoting the use of highly diversified reproductive material of good genetic quality 

.. effective genetic conservation methods 

• better scientific knowledge on genetic diversity 

• improving seed conservation techniques. 

References 

Anonymous. 1996. La gestion durable de la fon2t en Wallonie. Document provisoire. Min. 

Reg. Wal. Belgique. 

Claes, V. 1997. Etat sanitaire des chenes en Region Wallonne: ecologie du deperissement et 

moyens de lutte. Rapport final. FUSAGX, Gembloux, Belgique. 

de Cuyper, B. A. and D. Jacques. 1996. Conservation strategy for Noble Hardwoods in 

Belgium. Pp. 111-119 in Noble Hardwoods Network. Report of the first meeting, 24-27 

March 1996, Escherode, Germany (J. Turok, G. Eriksson, J. Kleinschmit and S. Canger, 

compilers). IPGRI, Rome, Italy. 

Galoux, A. 1966. La variabilite genecologique du hetre commun (Fagus sylvatica L.) en 

Belgique. Trav. Sta. Rech. Eaux et Forets, Groenendaal, Ser. A, no. 11. 

Hubert, D. 1988. Test de provenances et de descendances du Detre en Belgique. Trav. Fin 

d'Etud., Fac. Sci. Agron. Gembloux. 

Muller-Edzards, A. et al. 1997. Ten years of monitoring forest condition in Europe. UN/ECE- 

EC Report, Pp. 145-147. 

Nanson, A. 1993. Gestion des res sources forestieres. Annales Gembloux 99:13-36. 

Nanson, A. 1995. Situation of the conservation of Norway spruce in Belgium. Pp. 44-50 in 

Picea abies Network. Report of the first meeting, 16-18 March 1995, Stara Lesna, Slovakia 

0. Turok, V. Koski, L. Paule and E. Frison, compilers). IPGRI, Rome, Italy. 

Weissen, F. 1996. Scenarios de deperissement et du retablissement chez le hetre et l'epicea. 

Risques nouveaux pour la sylviculture du douglas et des melezes. Rapport. IRSIA, 

Gembloux, Belgium.


Annex 1. Institutions involved in forest genetic conservation in Belgium. 

Walloon Region 

Ministry of Walloon Region 

Forest Research Station 

Avenue Marechal Juin, 23 

B-5030 Gembloux 

Tel: +32 81 61 11 69 

Fax: +3281 61 5727 

Agricultural Research Center 

Avenue de la Faculte, 22 


Tel: +3281 611955 

Fax: +3281 614941 

Catholic University of Louvain-la-Neuve 

Agronomic Sciences Faculty 

Place Croix du Sud, 2, boite 9 

B-1348 Louvain-la-Neuve 

Tel: +32 10473707 

Fax: +32 10 47 36 97 

Faculty of Agricultural Sciences 

Forestry Department 

Passage des Deportes, 2 


Belgium 

Tel: +3281622320 

Fax: +32 81 62 23 01 

Flanders 

Ministry of the Flemish Community 

Environment, Nature and Land 

Development Administration 

Institute for Forestry and Game 

Management 

Duboislaan, 14 

B-1560 Hoeilaart 

Tel: +32 2 65703 86 

Fax: +32 2 657 96 82 

University of Brussels 

Faculty of Sciences 

Lab. for General Botany and Nature 

Management 

Pleinlaan, 2 

B-1050 Brussels 

Tel: +322 629 34 16 

Fax: +32 2 629 34 13 

University of Ghent 

Faculty of Sciences 

Lab. for Molecular Genetics 

K.L. Ledeganckstraat, 35 

B-9000 Ghent 

Tel: +32926451 71 

Fax: +329 26453 49 

University of Ghent 

Faculty of Agronomic and Applied 

Biological Sciences 

Lab. for Forestry 

Geraardsbergsesteenweg, 267 

B-9090 Melle-Gontrode 

Tel: +3292522113 

Fax: +:32 9 252 54 66


Activities concerning Social Broadleaves genetic resources in the 

Netherlands 

Sven M.G. de Vries 

Institute for Forestry and Nature Research (IBN-DLO), Wageningen, the Netherlands 

Introduction 

The current importance of Social Broadleaves in the Netherlands is increasing rapidly these 

days, both within the forestry sector and in landscaping. 

The species covered by the name Social Broadleaves in the Netherlands include beech 

(Fagus sylvatica) and oaks (Quercus petraea and Q. robur). 

Beech and oak species are not threatened so much at the species level, since they have 

always been much used. However their genetic resources are threatened. Autochthonous 

material is scarce, often located on private estates or nature reserves and therefore less 

accessible, and the price of seeds and plants is high. As a result of an often negative 

selection it appears that the quality of trees from these rare sources does not meet the 

requirements in terms of forestry standards. 

A start was made some years ago for a national strategy of conservation of forest genetic 

resources, including inventories and background research on diversity. However, Social 

Broadleaves are not considered as a group of species, but treated like all other species, 

individually. 

Besides in situ gene conservation of ecosystems, a specific gene conservation strategy is 

being or will be applied to every individual species. 

Within the framework of the EU, the genetic diversity of presumed autochthonous oak 

populations is studied with molecular techniques. A national programme provided the 

possibilities for research and practical application of gene conservation in Q. petraea. 

General situation of indigenous tree species 

From the total of about 80 indigenous species of trees and shrubs in the Netherlands, about 

9% almost disappeared, 33% are very rare, 36% rare and only 21% more or less common, 

though possibly threatened locally (Maes 1993). 

Juniperus communis is the only species protected by law. However, a number of 

regulations and incentives favouring genetic resources do exist. Following the International 

Union for Conservation of Nature and Natural Resources (IUCN), a Red List was 

constructed for the Netherlands, containing eight species of trees and shrubs. 

Besides the protection of the species, it is very important to be able to protect 

environments and locations where they occur. 

Restoration of typical forest types and reintroduction of autochthonous forest material 

take place at forestry level and in landscaping. 

Inventories have been made since 1992, whereas seeds and plant material are collected 

and grown on a contract basis to serve special purposes and plantations. 

A proposal for a nationwide inventory was drawn up by Maes (1993) including a certain 

time schedule. Parts of the inventories in this proposal have been carried out by now, while 

other parts will have to wait for funding (Fig. 1). 

Some of the species have been introduced into clonal archives, or into seed orchards, or 

both. In this way rare genotypes can be combined in order to increase genetic variability. 

Collecting of the material, maintenance of the clonal archives and the layout of the seed 

orchards has so far been carried out by the Institute for Forestry and Nature Research (IBN- 

DLO).


Fig. 1. Proposal for a nationwide inventory of indigenous species of trees and shrubs (Maes 1993). 

Establishment and maintenance of the seed orchards has been taken care of by the State 

Forest Service (SBB). 

Future activities in relation to genetic collections, recently initiated by the Government, 

also include private companies working together with SBB and IBN-DLO. 

Value of Social Broadleaves 

Beech is frequently planted in the Dutch forestry system. Natural regeneration is rarely used 

for reforestation. For this re.ason it is difficult to trace autochthonous material. The 

probability of the presence of autochthonous material concerns only cases where planting 

stock of local origin was used. Presumed autochthonous beech occurs only in a few places 

in the country and even in these cases the origin is not absolutely certain.

· COUNTRY REPORTS 99 

Quercus robur is a very important forest tree species in the Netherlands. Among 

generatively reproduced indigenous broadleaved species, it is the most widely spread and 

planted in the country. It is present in most of the natural forest ecosystems. Quercus petraea 

is of less importance than Q. robur in the Netherlands. Its importance is mainly based on 

ecological grounds rather than on wood quality criteria. 

At the moment oak is the dominant tree species on 16% of the total forest area. 

In addition about 25% of line and roadside plantations are established with oak. The 

Forest Policy Plan indicates an even higher proportion of oak in the Netherlands in the 

future. 

Oak has been subjected to two different selection systems throughout time. In earlier 

days oak was a major species in the community forests. When local people needed wood 

they usually harvested the best material (by phenotypic criteria) and thus created a negative 

selection on the genetic material of oak. No positive selection was carried out in the coppice 

type of forest either. On the other hand, the use of oak for roadside plantations was a 

tradition. Seeds from these plantations were the source of new roadside plantations. The 

advantage of this long tradition of harvesting tree seeds in roadside plantations is the 

recurrent positive selection toward a certain desirable type of tree. The tree nursery selects 

among all its populations about 2-5% for the special purpose of establishing the next 

generation of roadside trees. These trees finally come to stand in a roadside plantation 

which after many years delivers a series of new generations in which the tree nursery again 

selects 2-5% to proceed with the next generation of roadside trees. 

Despite this, too little emphasis has been put in the past on the use of material with high 

genetic quality. Seeds of good stands were very often mixed with that of poor stands for the 

sake of quantity only. On top of that, also in the past, a lot of seeds were imported in poor 

production years from unknown sources. The chance of this material being less adapted to 

local conditions is rather high. 

Beech and oaks very often grow together in harmony in those places where they seem to 

have grown undisturbed for a long time. Some still existing large estates illustrate this 

(Maes 1993). 

A major problem these days is the sudden change in water tables. In both directions, 

either too dry or too wet, this could mean a disaster to the ancient trees. Climate fluctuations 

and the need for high water supply are often the cause of these changes in the water table. 

Identification of the genetic background of autochthonous oak populations 

For many reasons it is considered important to learn about the genetic background of oak in 

the Netherlands. 

During a survey of genetic resources of trees and shrubs in the Netherlands, 13 

autochthonous populations of oak were identified, based on historical, taxonomic and site 

criteria (Maes 1993). 

Within the framework of the EU project 'Synthetic maps of gene diversity and 

provenance performance for utilization and conservation of oak genetic resources in 

Europe', the genetic diversity in these presumed autochthonous oak populations is studied 

with molecular techniques. Laboratories and research institutes from Austria, Denmark, 

France, Germany, Italy, Netherlands, Spain, Switzerland and the United Kingdom are 

involved in this project. 

Eight mixed populations of Q. robur and Q. petraea and five pure Q. robur populations 

were sampled to determine the polymorphism of the chloroplast DNA. 

Three haplotypes (Petit et al. 1993) are dominant in the Netherlands. Two of them follow 

the pattern of the geographic distribution of haplotypes in Europe, one originates from the 

Balkans. Further studies should reveal whether this last haplotype migrated into the 

Netherlands during postglacial recolonization of Europe or was introduced artificially. 

Combining the information gathered in Europe might allow identification of the original 

origin of our selected seed stands.

102 EUFORGEN: S001AL BROADLEAVES 

Beech and oak species in Germany: occurrence and gene conservation 

measures 

B. Richard Stephan 

Bundesforschungsanstalt fur Forst- und Holzwirtschaft, Institut fur Forstgenetik und 

Forstpflanzenzuchtung, Grosshansdorf, Germany 

Introduction 

Beech (Fagus sylvatica L.) and the two oak species, sessile oak (Quercus petraea (Matt.) Liebl.) 

and pedunculate oak (Quercus robur L.), are the most common and most economically 

significant broadleaved species in Germany. According to the last forest inventory of 1990 

the total forest cover in Germany is 10.8 million ha with 14.0% beech forests and 8.6% oak 

forests (BML 1990, 1994). The two oak species are not separated in forest statistics. These 

three broadleaved species occur in nearly all parts of Germany with a few regional 

exceptions (Rohrig 1980). They grow on sites with a wide range of ecological conditions 

(Fig. 1). 

A third oak species, the pubescent oak (Q. pubescens Willd.), is a submediterranean tree 

species native only on very warm and dry sites of southwestern Germany. The North 

American red oak (Q. rubra L.) is an important introduced and fast-growing species and was 

planted in large numbers in the 19th century. The Turkey oak (Q. cerris L.) is also grown in 

some regions on drier and poor sites. These last species regenerate also naturally, but will 

not be considered in this report. 

dry 

shade-intolerant trees 

wet (Pinus) Fraxinus excelsior Ulmus species 

Betula pubescens 

Alnus glutinosa 

acid ----------------------~. base-rich 

Fig. 1. Water and nutrient site requirements of main tree species in submontane regions (Ellenberg 

1982, modified) [shaded = beech area; dark shaded = oak area].

COUNTRY RERORTS 103 

The species 

Beech occurs under atlantic to subatlantic climatic conditions. The species grows mainly in 

western and central Europe in pure and mixed stands, has a very strong competitive 

capacity, is shade-tolerant and very adaptive. Beech is distributed naturally from the coastal 

areas to an elevation of about 1500 m (1700 m) in the northern parts of the Alps. It prefers 

sites with a good water and nutrient supply and avoids very moist or dry and poor sites. 

Beech occurs typically on sites with limestone as basic rock. It can be assumed that many 

stands in Germany are autochthonous. 

Sessile oak and pedunculate oak occur sympatrically in Germany from atlantic to 

continental climatic regions. As both oak species occur under various climatic conditions, 

differentiated populations have developed. The populations show a high intraspecific 

genetic variation and differ by phenology, growth performance, form and other traits 

(Kleinschmit 1993). The differences in flushing are particularly important in connection 

with the susceptibility to late frost (Stephan et al. 1995). 

Mating between the two species is obviously frequent, although the proportion of 

tentative hybrids in natural stands is not known so far (Rushton 1993). The species concept 

of the two oaks is still in discussion. Sessile oak and pedunculate oak are considered 

taxonomic ally and ecologically as two separate species (e.g. Aas 1996). The distinction 

between sessile oak and pedunculate oak is difficult as many intermediate forms occur, and 

hybrids cannot be identified morphologically on an individual level. Comparing various 

morphological and genetic traits, Kleinschmit et al. (1995) did not detect any single character 

which had disjunctive expression. Many other studies have shown that it was in no case 

possible to differentiate the two species by one single trait or marker. In some cases the 

combination of various characters opened the possibility to distinguish between Q. petraea 

and Q. robur, but the genetic differentiation was very low (Muller-Starck et al. 1993). The 

combination of morphological, phenological and growth traits of the same seedling over 

several years also resulted in some significant correlations between traits for the 

differentiation between the two species (Liesebach and Stephan, unpublished). 

Hybridization between Q. petraea and Q. robur is possible by controlled crosses, but with 

different fertility rates depending on whether Q. petraea or Q. robur were taken as pollen 

parent (Steinhoff 1993). Some authors (Kleinschmit et al. 1995) concluded that sessile oak 

and pedunculate oak may represent different ecotypes of the same species. 

The sessile oak type inhabits warmer sites and lighter soils. The species is prevalent in 

the warmer hilly regions of western and southwestern Germany and endures drier site 

conditions. Regions with famous sessile oak forests are in southwestern Germany, the 

Spessart and the Pfi:ilzer Wald. In the Bavarian Alps sessile oak can be found up to an 

elevation of about 900 m. 

The pedunculate oak type grows optimally on rich loamy soils with a good water supply 

and occurs mainly in the lowlands from sea level up to an elevation of 950 m in the Bavarian 

Alps. 

Beech, sessile oak and pedunculate oak are included in the Act on Forest Seed and 

Planting Stock of Germany (Anonymous 1979). In the Regulations on the Regions of 

Provenance for Forest Reproductive Material 26 regions are defined for beech, 13 for sessile 

oak, and 9 for pedunculate oak, respectively (Anonymous 1994). 

Current economic importance and use of reproductive material 

Timber from beech and oak has a high economic importance. The amount of fellings during 

the past years was about 7 million m 3 wood under bark per year for beech (= 20% of the total 

felling), and about 1.3 million m 3 wood under bark per year for oak (= 4%) (Hein and Bitter 

1997). In both groups of these two tree genera about 50% was used as stem wood and 50% 

as industrial wood. Prices for stem wood remained more or less stable for beech, but 

decreased for oak. Prices for industrial wood decreased generally in the last 10 years.

104 EUEORGEN: SOGIA~ BROA[i)~EAMES 

Nevertheless, high prices are paid when the timber has a good veneer quality, as the 

following example shows: for an oak stem with a length of 6.5 m and a mean diameter of 

0.85 m, 51 765 DM were paid for the whole stem (14 500 DM/m 3 solid volume). 

Besides timber, seed trade of approved beech and oak stands is also of great economic 

significance. Seed trade is an important factor at both national and international levels. The 

number of approved stands and seed orchards of the EEC and OECD category 'Selected', 

and of the approved stands of the category 'Tested' is shown in Table 1. More than 81000 ha 

approved stands - seed orchards included - are registered for beech, about 32000 ha for 

. sessile oak, and about 9100 ha for pedunculate oak. About 166000 kg seeds of beech, about 

400000 kg acorns of sessile oak, and about 256000 kg acorns of pedunculate oak were 

collected annually on average over the last 14 years. 

Table 1. Summary list of approved basic material 

No. of 

stands or 

Reduced area (ha) of stands or seed orchards 

seed Autoch- Non-autoch- 

Tree species orchards thonous thonous Unknown Total 

Approved stands of the category 'Selected' 

European beech 14199 70434.5 691.7 9969.3 81095.5 

Sessile oak 8336 26079.2 286.1 5278.1 31643.4 

Pedunculate oak 2145 3858.5 535.7 4684.0 9078.2 

Approved seed orchards of the category 'Selected' 

European beech 1 1.5 1.5 

Sessile oak 

Pedunculate oak 2.0 2.0 

Approved stands of the category 'Tested' 

European beech 28 244.1 13.3 257.4 

Sessile oak 39 202.4 5.2 2.3 209.9 

Pedunculate oak 8 2.5 11.7 28.3 42.5 


In general, beech is regenerated naturally. Oak can be sown directly or planted using wild 

saplings from the forest or nursery-grown seedlings. If the species are regenerated 

artificially, the use of local and well-adapted provenances is recommended. The aim of any 

silvicultural and forest management activity is to guarantee the sustainable use of the 

resources, the maintenance of forests as the basis for the multiple forest functions, such as 

production of wood as renewable raw material, regulation of watershed, various social 

functions, and last but not least sustainable conservation of biodiversity. Beech and oak 

forest communities in particular are rich forest ecosystems, given their various combinations 

with other species (Fig. 1). 


Although forest decline has been observed for a long time, the situation developed 

dramatically in the early 1980s, when heavy and increasing damage occurred in silver fir 

(Abies alba Miller), followed by severe damage in Norway spruce (Picea abies (L.) Karst.). For 

the last 10 years, broadleaved trees, including beech and oak, also have shown increasing 

and heavy disease symptoms. Forest decline was registered officially in 1984 and the 

monitoring results published in annual reports, the latest in 1997 (BML 1997). 

Damage is recorded on a five-step scale with increasing grades from 0 to 4. The 

proportion of damaged forest trees of all ages was 20% (grades 2 to 4) in 1997. Beech was 

damaged at about 29%, oak at 46%. The annual proportion of damaged beech and oak trees 

from 1984 to 1997 is shown in Figures 2 and 3. Older stands in particular show a high 

percentage of damaged trees. For trees more than 60 years old, the situation was serious in 

1997: 38% of beech trees and 55% of oaks showed remarkable damage (grades 2 to 4).

GeUNmR¥ REBeRmS 105 

During recent years a Europe-wide decline of oaks was recorded, dramatic in some 

regions. A complex of several biotic and abiotic factors seems to be responsible for the 

symptoms (Wulf and Kehr 1996). 

Forest decline is mainly caused by anthropogenic environmental loads (emissions) and 

has obviously strong effects on the genetic diversity of certain tree populations (compare, for 

example, Miiller-Starck 1993). 

Besides damage by emissions, beech and oak are sensitive to late frost in spring, beech 

also to drought. During recent years several epidemics by insects (e.g. Agrilus sp., Lymantria 

dispar, Tortrix viridana and others) attacked in'particular the oak species in various parts of 

Germany (Wulf and Kehr 1996). Excessively increased populations of game can also be a 

burden for young forest stands. 

01 = Northwest German States 

02 = South German States 

113 = East German States 

1114 = Germany (in total) 

Fig. 2. Development of forest decline in beech in Germany from 1984 to 1997. The summary of 

damage classes 2 to 4 in % is shown for all age classes (source: BML 1997). 

% 

01 = Northwest German States 

02 = South German States 

1!1113 = East German States 

114 = Germany (in Total) 

1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 

Fig. 3. Development of forest decline in oak in Germany from 1984 to 1997. The summary of damage 

classes 2 to 4 in % is shown for all age classes (source: BML 1997).

106 EUF0RGEN: S0CIAIl BR0ADIlEAVES 


After the establishment of the Federal and State Working Group 'Conservation of Forest 

Genetic Resources in the Federal Republic of Germany' in 1984, a concept was developed for 

conservation measures and research. Among many other tree and shrub species, the 

conservation activities also include beech and the two main oak species (BLAG 1989, 1997). 

Existing measures for direct and indirect gene conservation are carried out by federal and 

state institutions (Uinder) of Germany. There is little private activity in this respect. Special 

legal regulations concerning the conservation of forest genetic resources are missing. But the 

following legal regulations contribute indirectly to the protection and conservation of forest 

genetic resources: 

• forest laws of the Federal and State Governments 

• legal regulations concerning forest reproductive material 

• laws and regulations on nature protection. 

The following in situ and ex situ conservation measures can be applied: conservation of 

stands by reduced management (fellings), support of natural regeneration, sowing and 

planting in situ and ex situ, establishment of seedling and clonal seed orchards, 

establishment of clone collections or clonal archives, conservation of seed, pollen, plants, 

parts of plants including tissue in genebanks, and conservation by macro- and 

microvegetative propagation. All these measures are applied in beech and oaks, often 

concurrently, minimizing the risk. The various measures have advantages and 

disadvantages (BLAG 1997). 

Additional activities include conservation of material within the framework of breeding 

programmes, as well as the conservation of provenances, families and clones in field trials. 

The material designated as 'worthy for conservation' includes, for example: approved 

basic material for forest reproductive material, selected or comparable populations covered 

by the Act on Forest Seed and Planting Stock (Anonymous 1979), populations under specific 

ecological conditions, marginal populations and material severely endangered by current 

damage or by its rarity. 

The state (1996) of gene conservation activities in beech, sessile oak and pedunculate oak 

is shown in Table 2 with respect to in situ and ex situ measures. Emphasis was laid on in situ 

conservation of stands including natural regeneration (Fig. 4). As an ex situ measure the 

establishment of seed orchards is of great significance (Fig. 5). 

In connection with the above-mentioned activities research is also needed on variability, 

genetic structures of populations, effects of silvicultural and conservation methods on 

genetic diversity, and methods for long-term storage of seeds. Despite the importance of 

beech and oak in German forestry, there is little knowledge about the genetic structure of 

populations throughout their range of distribution (Milller-Starck and Ziehe 1991). There 

are many results of isoenzyme studies in beech. A practical guide concerning electrophoretic 

methods of separation and for the evaluation of zymograms is in preparation. Although 

regional differences can be found, it was shown that generally the variation within 

populations is larger than between populations. Concerning oak, DNA studies have shown 

very recently the existence of genetic types with differences in the chloroplast genome 

(chloroplast DNA), which are distributed geographically (Kremer et al. 1991; Petit et al. 1993; 

Dumolin-Lapegue et al. 1997). These results can probably be used for studies about the 

postglacial migration, about the delimitation of regions of provenance, about the 

autochthonous origin of populations, and other questions. Studies on the genetic diversity 

in specific amplified chloroplast regions of the two main oak species in Germany are under 

way (Konig et al. 1998). 


Valuable beech and oak forest stands with a rich vegetation are often protected as nature 

areas. Very old and attractive single trees are also under special protection.

CC>UN!I"RY REPC>R!I"S 

10'7' 

Very recently a plan is under discussion to establish at least two large areas with natural 

beech forests as national parks: Hainich (Thuringia) and Kellerwald (Hesse). 

Public awareness about the importance of beech and oak is high in Germany. Since 1989, 

each year a tree species is elected 'Tree of the Year'. Oak was the first species elected, 

followed by beech in 1990. Special leaflets are prepared and articles are published in 

newspapers to provide the public with information on the 'Tree of the Year'. 


Selection of single plus trees is conducted in all three tree species for the establishment of 

seed orchards or clonal archives (Table 2). 

In comparison with the significance of beech and oak, few older provenance trials exist. A 

review of the international trial series with more than 300 beech provenances since 1983 was 

presented at this EUFORGEN meeting (von Wuehlisch et al., this volume). 

There are also oak provenance trials (trees about 40 years old) initiated by Krahl-Urban, 

which give interesting results on growth performance, stem form and flushing (Kleinschmit 

and Svolba 1995). More recently, an international series with more than 30 sessile oak 

provenances was organized by Madsen (1990) and established in various countries, 

including Germany. First results give information about growth, flushing, bud-setting, frost 

tolerance and several other traits (Stephan et al. 1995). 

The intensive breeding work with oak concluded at the Forestry Research Station of 

Lower Saxony, Escherode, must also be mentioned (Steinhoff 1993; Schiite 1995). Controlled 

crossings between and within the two oak species have been carried out successfully. The 

results give an insight about the possibility and importance of hybridization between the 

species. 


The Federal Research Institute, 10 Institutes of the states (Liinder), and Institutes of four 

Universities are involved in the in situ, ex situ activities or in research. The federal and the 

state institutions constitute the working group 'Conservation of Forest Genetic Resources in 

the Federal Republic of Germany', exchange information intensively and coordinate the 

activities of the member institutions by considering regional peculiarities. 

Summary of country capacity and priorities 

Studies on the genetic structures in beech and oak, funded by several national and 

international projects (EU, Deutsche Forschungsgemeinschaft - German Research 

Association, etc.) have a high priority. German institutions are partners or coordinators of 

the following national and international projects on beech and oak: 

• Concerted Action: 'European Network for the Evaluation of the Genetic Resources of 

Beech for Appropriate Use in Sustainable Forestry Management' (AIR3-CT94-2091) 

• Shared-Cost Action: 'Common Beech for Forestation and Diversification: Development 

of Forestation Techniques and Assessment of the Genetic Variation in Reproductive 

Material' (FAIR3-PL96-1464) 

• Shared-Cost Action: 'Synthetic Maps of Gene Diyersity and Provenance Performance 

for Utilization of Oak Resources in Europe' (FAIR1-CT95-0297) 

• Testing the Frost Tolerance of Acorns by Differential Temperature Analysis [Priifung 

der Frostharte von Eicheln mittels Differenztemperaturanalyse] (BML 115-0762-A-3- 

5/363). 

Still very little is known about optimal conditions for long-term storage of acorns of the 

recalcitrant tree species beech and oak. Projects were started recently. 

The mentioned international provenance trials in beech and oak will improve our 

knowledge about the genetic variation of beech, sessile oak and pedunculate oak and will 

support gene conservation measures. Therefore, it is necessary to start or to continue 

international collaboration with all countries in which beech and oaks occur. A first step 

was made by the establishment of this EUFORGEN Network on Social Broadleaves.

Table 2. In situ and ex situ conservation measures for beech and oak in Germany (at 31.12.1995) 

In situ 

Ex situ 

Stands Stands Seed orchards 

Area No. single Area No. single Area No. of 

Species No. (ha) trees NC). (ha) __!!"~~s; No. ... (~ families 

Fagus sylvatica 255 2201 145 93 152 10 13 64 

Quercus petraea 75 345 444 81 96 7 8 

Quercus robur 217 480 246 55 77 11 20 

No. of 

clones 

216 

222 

158 

Clonal archives 

No. of 

No. clones 

1 6 

Seed storage 

Stands/seed orchards Single trees Pollen storage Tissue storage 

Species No. Quantity (kg) No. Quantity (kg) No. Quantity (cm3) No. of samples 

Fagus sylvatica 256 1961 72 16 19 

Quercus petraea 2 23 16 4 42 685 35 

Quercus robur 1 < 1 35 535 51 

Source: BLAG 1996.

COUNTR¥ REPORTS 109 

ha 

2500 

2000 

1500 

1000 

2201 

48 67 

Fig. 4. Stands for in situ conservation (state: 1995) (sources: BLAG 1996; Tabel 1997). 

number 

200 

180 

160 

140 

120 

100 

80 

60 

40 

20 

Fig. 5. Seed orchards for ex situ gene conservation (state: 1995) (sources: BLAG 1996; Tabel 1997).


References 

Aas, G. 1996. Morphologische and okologische Variation mitteleuropaischer Quercus-Arten: 

Ein Beitrag zum Verstandnis der Biodiversitat. Habilitation Thesis, ETH Zurich. 

Anonymous. 1979. Gesetz uber forstliches Saat- und Pflanzgut [Act on Forest Seed and 

Planting Stock]. BGBI I, p. 1242. 

Anonymous. 1994. Verordnung uber Herkunftsgebiete fUr forstliches Vermehrungsgut 

(Forstsaat-Herkunftsgebietsverordnung) vom 7. Oktober 1994 [Regulation on the Regions 

of Provenance for Forest Reproductive Material - Forest Seed Provenance Region 

Regulation of 7. October 1994]. BGBI I, p. 3578. 

BLAG (Bund-Lander-Arbeitsgruppe "Erhaltung forstlicher Genressources"). 1989. Konzept 

zur Erhaltung forstlicher Genressourcen in der Bundesrepublik Deutschland. Forst und 

Holz 44:379-404. 

BLAG. 1996. Tatigkeitsbericht der Bund-Lander-Arbeitsgruppe "Erhaltung forstlicher 

Genressourcen". Berichtszeitraum 1994-1995. 

BLAG. 1997. Concept for the conservation of forest genetic resources in the Federal Republic 

of Germany. Silvae Genet. 46:24-34. 

BML (Bundesministerium fUr Ernahrung, Landwirtschaft und Forsten). 1990. Bundeswaldinventur. 

Band 1. 

BML. 1997. Waldzustandsbericht der Bundesregierung 1997. Ergebnisse der Waldschadenserhebung. 

BML. 1994. Der Wald in den neuen Bundeslandern. 

Dumolin-Lapegue, S., B. Demesure, S. Fineschi, V. Le Corre and RJ. Petit. 1997. Phylogeographic 

structure of white oaks throughout the European continent. Genetics 146:1475-1487. 

Ellenberg, H. 1982. Vegetation Mitteleuropas mit den Alpen in okologischer Sicht. 3. Aufl. 

Verlag E. Ulmer, Stuttgart. 

Hein, W. and W.-G. Bitter. 1997. ZMP-Bilanz Forst und Holz 1997. Zentrale Markt- und 

Preisberichtstelle, Bonn. 

Kleinschmit, J. 1993. Intraspecific variation of growth and adaptive traits in European oak 

species. Ann. Sci. For. 50, Suppl. 1:166-185. 

Kleinschmit, J. and J. Svolba. 1995. Intraspezifische Variation von Wachstum und 

Stammform bei Quercus robur und Quercus petraea. Mitteilungen aus der Forstlichen 

Versuchsanstalt Rheinland-Pfalz 34:75-99. 

Kleinschmit, J., R Bacilieri, A Kremer and A Roloff. 1995. Comparison of morphological 

and genetic traits of pedunculate oak (Q. robur L.) and sessile oak (Q. petraea [Matt.] 

Liebl.). Silvae Genet. 44:256-269. 

Konig, A, K. Groppe and B. Ziegenhagen. 1998. First results of chloroplast-DNA investigations 

in German populations of Quercus petraea and Q. robur. Presented at the meeting, "Diversity 

and Adaptation in Oak Species", State College, Pennsylvania, USA, 12-17 October 1997. 

Kremer, A, R Petit, A Zanetto, V. Fougere, A Ducousso, D. Wagner and C. Chauvin. 1991. 

Nuclear and organelle gene diversity in Quercus robur and Q. petraea. Pp. 141-166 in 

Genetic Variation in European Populations of Forest Trees (G. Muller-Starck and M. 

Ziehe, eds.). J.D. Sauerlander's Verlag, Frankfurt am Main. 

Madsen, S. F. 1990. International provenance trial with Quercus petraea. The 1989 series of 

provenance experiments with Quercus petraea (Matt.) Liebl. Danish Forest Experiment 

Station, December 1990. 

Muller-Starck, G. 1993. Auswirkungen von Umweltbelastungen auf genetische Strukturen 

von Waldbestanden am Beispiel der Buche (Fagus sylvatica L.). Schriften der Forstlichen 

Fakultat der Universitat Gottingen und der Niedersachsischen Forstlichen 

Versuchsanstalt 112:1-163. 

Muller-Starck, G. and M. Ziehe. 1991. Genetic variation in populations of Fagus sylvatica L., 

Quercus robur L. and Q. petraea Liebl. in Germany. Pp. 125-141 in Genetic Variation in 

European Populations of Forest Trees (G. Muller-Starck and M. Ziehe, eds.). J.D. 

Sauerlander's Verlag, Frankfurt am Main.


Miiller-Starck, G., S. Herzog and H.H. Hattemer. 1993. Intra and inter population genetic 

variation in juvenile populations of Quercus robur L. and Quercus petraea Liebl. Ann. Sci. 

For. 50, Suppl. 1:233-244. 

Petit, R, A. Kremer and D.B. Wagner. 1993. Geographic structure of chloroplast DNA 


Rohrig, E. 1980. Waldbau auf okologischer Grundlage. Band 1. 5. Aufl. Verlag Paul Parey, 

Hamburg und Berlin. 

Rushton, B.s. 1993. Natural hybridization within the genus Quercus L. Ann. Sci. For. 50, 

Suppl. 1:73-90. 

Schiite, G. 1995. Kontrollierte Kreuzungen und Entwicklung der Hybriden von Stiel- und 

Traubeneiche (Quercus robur L. und Quercus petraea [Matt.] Liebl.). Mitteilungen aus der 

Forstlichen Versuchsanstal t Rheinland -Pfalz 34:38-49. 

Steinhoff, S. 1993. Results of species hybridization with Quercus robur L. and Quercus petraea 

(Matt.) Liebl. Ann. Sci. For. 50, Suppl. 1:137-143. 

Stephan, B.R, A. Konig, K Liepe and H. Venne. 1995. Klimakammer- und Freilandversuche 

zur Frostharte und Phanologie von Herkiinften der Traubeneiche (Quercus petraea 

[Mattuschka] Liebl.). Mitteilungen aus der Forstlichen Versuchsanstalt Rheinland-Pfalz 

34:50-74. 

Tabel, U. 1997. Erhaltung forstlicher Genressourcen in der Bundesrepubik Deutschland. 

Allgemeine Forst Zeitschrift/Der Wald 52:259-261. 

Wulf, A. and R Kehr (eds.). 1996. Eichensterben in Deutschland. Oak decline in Germany. 

Mitteilungen der Biolog. Bundesanstalt f. Land- und Forstwirtsch., Berlin, Heft 318.

112 EUFOBGEN: SOCIAL BBOADLEAVES 

Conservation strategy for beech and oaks in Denmark 

Jan S. Jensen 

Danish Forest and Landscape Research Institute, H0rsholm, Denmark 

Introduction 

The Strategy for the Conservation of Genetic Resources of Trees and Shrubs in Denmark was 

prepared by the National Forest and Nature Agency in 1991-93. The general principles and 

approaches to genetic conservation of forest trees in Denmark are discussed in detail by 

Graudal et al. (1995). 

The objective is to secure the ability of the species to adapt to environmental changes, and 

to maintain the basis for future tree-improvement work. For both beech (Fagus sylvatica) and 

oaks (Quercus robur and Quercus petraea) this strategy was initiated in 1992. Several ex situ 

areas have been established, but many areas for in situ conservation have not been 

designated yet. 

Occurrence and origin of beech and oaks 

Denmark has mainly been formed by the last Glacial (Weichselian), and the development of 

the forest has taken place during the last 10 000 years. 

Oaks have been present since 7000 BC. Pedunculate oak (Quercus robur) is distributed all 

over the country, found on a large variety of ecological conditions. Sessile oak (Q. petraea) 

grows mainly in the central parts of Jutland, on sandy hilly locations. Sessile oak is found 

sympatric to pedunculate oak. 

After invading Denmark in 1000 BC, beech reached its maximum distribution only 1000 

years ago, replacing lime as the dominating forest species. Approximately 5000 years ago, 

substantial human impact on the landscape began, due to agriculture and grazing, and the 

forest reached a minimum of 3% of the total land area toward the end of the 18th century. 

Since then the forest area has increased to 12%, mainly through the establishment of 

coniferous plantations. Beech is dominant on better soils and is very competitive with both 

sessile and pedunculate oaks. 

Beech occupies 72 000 ha and is slowly increasing in area after a long period of decrease. 

The area covered by oaks has increased from 13 000 ha in 1907 to 30000 ha in 1996. Oaks 

and beech cover about 25% of the forest area. 

Forest management 

The expansion of beech since the Glacial is a result of both anthropogenic and natural 

processes. The reduction in forest area until 1800 resulted in serious fragmentation of beech 

and oak forests. 

Human impact has influenced qualitative properties of forest trees, especially beech and 

oaks. Substantial genetic variation in tree form can be shown for oaks. 

Slash-and-burn management reduced forest area, and instead created heathland and 

brushwoods in western Denmark. Large amounts of wood were used for fuel. Evidently 

large-dimension timber has been removed for construction purposes; straight trees have 

been selected primarily. Heavy grazing by domestic animals also had an effect on the forest 

stands. Acorns were used for fodder in historical times. 

Large numbers of pigs roaming around in the forest probably favoured beech 

regeneration as compared with other species. All these processes have probably been acting 

in a complex way at different rates and strengths at various locations in the country. 

Since 1800 most of the forests including coppice, shelterwoods and brushwoods has been 

converted into high forests, favouring straight and fast-growing trees. Management of 

single species has been preferred and mixed forests have almost disappeared. Beech has 

been planted widely in areas where it was absent before.

CGUNmRM REPGRmS 113 

In contrast to other species, the Danish beech forests still retain their genetic continuity as 

most are regenerated naturally. 

Several oak stands in Jutland were converted from coppice forest into high forest. Several 

methods of coppice management may have been practised in the past; this may influence 

genetic structure in different ways. 

Both oaks and beech are important species for forestry. On better soil types, their mean 

economic production can reach 500-1000 DM per year based on the 130-year rotation period. 

The timber is used for veneer and board production. Both species possess high recreational 

value and historical and cultural importance. 


Most areas with beech and oaks are owned by the public, but a number of large estates 

includes large stands with beech and oaks. Danish forestry is dominated by management of 

small compartments between 2 and 6 ha. Oaks and beech are mainly found in pure stands 

and along with drainage of wet biotopes, the genetic structures may be affected. Coppice 

forestry is seldom practised nowadays, only by a very few small private forest owners. 

Beech 

For the last 200 years, beech management has mainly been high forest with a rotation age 

110-140 years. Intensive thinning has been practised. On a few occasions, 100-150 years ago, 

large compartments of 10-30 ha beech stands were established. 

Oaks 

Oak forests on better location have been managed as high forest with strong thinning and 

short rotation - 130 years. On poor sites, moderate thinning is often practised and rotation 

period may be up to 150-160 years. 

Health state 

The health state of beech varies from year to year, depending on climate, flowering, etc. 

Beech is considered to have endangered genetic resources owing to fragmentation or heavy 

imports of forest reproductive material. 

The health state of oak also varies. Moderate oak dieback appeared from 1989 to 1994, 

but now it has almost stopped. Two extreme years with bad flushing occurred (in 1993-94) 

in several locations. 

Research activities 

Isoenzyme studies have been carried out for both oaks and beech. The highly outcrossing 

species show low variation between population and moderate to high variation within 

populations (Larsen 1996; Siegismund and Jensen, pers. comm.). The results are comparable 

to international studies. They do not indicate genetic erosion, which could be expected 

following fragmentation. However, what is mostly needed is a more intensive assessment of 

variation, mainly of beech. 

Studies of provenance variation were initiated a hundred years ago, and were recently 

reported (Jensen 1993). We found high variation between populations and some clinal 

differences between Danish and foreign provenances. Within the Danish provenances, there 

seems to be a clinal pattern in flushing, as western provenances are flushing later than 

eastern provenances. This indicates the presence of different ecological zones in Denmark. 

In 1989-90 a large international collection of sessile oak provenances was initiated by the 

Forest and Landscape Research Institute. For the time being, a study of Scandinavian 

pedunculate oak is under way including studies on adaptive traits (frost and drought stress) 

and genetic markers. 

The Forest and Landscape Research Institute and the Agricultural University are involved 

in common EU projects on beech and oaks along with several partners from the EU

114 EUFORGEN: SOCIAL BROA[)LEAVES 

countries. This includes management of a common database for oak provenance trials, a 

synthetic map of chloroplast DNA variation, and mapping of within-population variation 

with microsatellite DNA markers. 


Two different strategies have been adopted concerning gene conservation of trees and 

shrubs in Denmark: 

'A Strategy for the Conservation of Genetic Resources of Trees and Shrubs in Denmark' 

by the National Forest and Nature Agency was prepared in 1991-93. The strategy provides 

an overview of gene conservation needs and required gene conservation measures, and a 

plan of implementation for gene conservation in Denmark. The strategy is closely linked to 

the tree improvement and seed procurement programmes in Denmark. 

The Strategy for Natural Forest and Other Forest Types of High Conservation Value in 

Denmark', adopted in 1992. The strategy focuses on ecosystem conservation, and does not 

address the conservation of forest genetic resources. 

Both strategies can be considered as national responses to recent international agreements 

concerning the conservation and sustainable use of biological diversity (Agenda 21, Helsinki 

and Strasbourg Resolutions). 

The Danish Forestry Act protects separately oak brushwoods and coppice forest. In 1987, 

314 ha of oak brushwoods were protected. 


Ex situ 

Activities have been initiated for oaks and beech. Some of the seed sources are regarded as 

very valuable, especially first and second generation stands. Even if landraces have not 

evidently developed, these seed sources should be conserved for future use. On a few 

locations, a limited number of combined seed production and conservation stands have been 

or will be established. Several stands of beech covering 40-50 ha will be established. 

Further ex situ tasks for oaks are also carried out through the tree improvement 

programme and the provenance trials. 

In situ 

Until recently only a limited number of forests have been protected formally. A number of 

virgin stands have been described, covering a total of 217 ha. Nature protection areas 

include forest of autochthonous origin. A number of private and public stands have been 

protected administratively. In 1992, The Danish Strategy for Natural Forest was adopted 

(see above). The Strategy should secure quantitative and qualitative distribution of nature 

protection areas for beech and oaks. The principle of natural forest is mainly based on a 

definition of genetic origin. As beech can be easily regenerated, this will give no problems. 

Oaks may be more difficult to assess. In total the Strategy concerns about 35 000 ha, most of 

it beech. 

Within the strategy for forest genetic resources, the in situ stands of beech will be selected 

within the natural forest areas. A number of 5-7 isolated stands of beech covering the 

geographic range will be identified. Each stand comprises at least 500 individuals. 

For pedunculate oak, a number of 8-10 stands will be designated, and for sessile oak 5-6 

stands. Sessile oak occurs naturally in 4-7 ecogeographic zones, in two of these zones only 

on one site each. The conservation status of many of the sessile oak brushwoods, so-called 

'purs' is good, but in situ conservation should mainly protect one or more of these areas 

against pollination from external sources.

, COUNTRY REPORTS 115 

Tree improvement activities 

A tree improvement programme for oaks was initiated in 1993 by the Arboretum. As 

natural regeneration is not practised, there is a large demand for acorns. The breeding 

programme should enhance the production of valuable seeds, and for oaks of Danish origin, 

stem quality should be improved. 

About 180 plus trees of eastern Danish origin were selected and acorns collected in 1995. 

Two seedling seed orchards will be established in 1998-99 (about 10 ha). This programme 

will also include clonal seed orchards. 

Another two seedling seed orchards of oak based on plus trees of Dutch origin will be 

established in 1998-99 (about 8 ha). 

For sessile oak used in harsh locations in western Jutland, 126 plus trees have been 

selected. From these trees, two clonal seed orchards are planned to be established. 

A breeding programme for beech has not been considered. 


Beech is mainly regenerated naturally. For planting, a substantial amount of seed material 

was imported from abroad during the last 200 years. Between 1880 and 1935, the major 

imports were from the Carpathians, the Netherlands, Belgium, Sweden and Germany. 

Today reproductive material comes from approved selected stands (36 in 1997) or foreign 

imports. From 1960 to 1980, 86% of the total beech seeds used were imported (Larsen 1983). 

In recent years, the amount of seed from Danish stands has increased. From 1990 to 1995 

Danish nurseries received 18 tonnes of beech seeds per year (Madsen and Sogaard 1996). 

Seed-processing has improved significantly and an increased number of approved stands 

has been selected. 

Large amounts of acorns have been imported for plantations of oaks since 1780. 

Especially Germany and the Netherlands, but also Norway, Sweden and France have 

contributed with acorns. Stem form and growth have been better in the imported material; 

provenances from the Netherlands have shown significantly better stem straightness than 

Danish provenances (Jensen 1993). Furthermore, imported seed material is relatively 

cheaper than Danish. Within the period 1960-90 only 27% of acorns for forestry and 

landscape use were harvested in Danish stands (Madsen 1991). Most of the acorns were 

imported from approved Dutch stands, mostly comprising roadside trees. From 1990 to 

1995, 39 t of sessile oak acorns and 97 t of pedunculate oak acorns were used annually in 

Danish nurseries. Eighty-three stands have been approved for seed production (1996). Only 

a few of these are presumably of Danish origin. 

Suml1)ary of country capacities and priorities 

In Denmark, beech is considered a well-protected species. In spite of high forest 

degradation (fragmentation), low interpopulation diversity.and high outcrossing have been 

revealed. Existing genetic resources are given high priority and are well protected. Active 

gene conservation efforts and tree improvement are not immediately needed. 

Both pedunculate oak and sessile oak are high-priority species. Generally, oak resources 

are considered threatened. This is due to massive import of foreign seed sources and no 

tradition of natural regeneration. An active programme involving tree improvement and 

gene conservation has been initiated. 


International collaboration in the fields of tree breeding and gene conservation is valuable 

for further research and management of genetic resources. 

Development of descriptive methods (morphology, phenology and biochemical markers) 

and more provenance experiments are necessary in general. A more precise description of 

seed sources, especially commercial, is also needed.


References 

Graudal, L, E.D. Kjeer and S. Canger. 1995. A systematic approach to the conservation of 

genetic resources of trees and shrubs in Denmark. For. Ecol. Manage. 73:117-134. 

Huntley, B. and Birks, H.J.B. 1983. Pp. 352-369. in An Atlas of Past and Present Pollen Maps 

for Europe, 0-13 000 years ago. Cambridge University Press, Cambridge, UK. 

Jensen, J.5. 1993. Provenances of Pedunculate Oak (Quercus robur) and Sessile Oak (Quercus 

petraea) in Denmark. PhD thesis. Research Series, The Royal Vet. And Agric. University. 

The Danish Forest and Landscape Research Institute, No. 2. 

J0hnk, N. and H.R. Siegismund. 1997. Population structure and Post-glacial migration routes 

of Quercus robur and Quercus petraea in Denmark, based on Chloroplast DNA Analysis. 

Scand. J. For. Res. 12:130-137. 

Larsen, A.B. 1996. Genetic structure of populations of beech (Fagus sylvatica L.) in Denmark. 

Scand. J. For. Res. 11:220-232. 

Larsen, J.B. 1983. Danske Skovtrceer, raceforhold, fmforsyning og proveniensvalg. Dansk 

Skovforenings Tidsskrift. 68. 

Madsen, S.P. 1991. Fm og planter til det danske marked. 1985-1990. Skoven 3:129-133. 

Madsen, S.P. and J. S0gaard. 1996. Fm og planter til det danske marked 1990/1995. 2 L0vtrce. 

Skoven 10:458-463.

Genetic resources and conservation of Quercus roburL. in Lithuania 

Virgilijus Baliuckas and Julius Danusevicius 

Lithuanian Forest Research Institute, Girionys, Kaunas, Lithuania 

Introduction 

Lithuania covers 65300 km 2 • The landscape is basically flat with minor hills. Sod podzolic 

soils are prevalent. The average altitude is 99 m. The climate is transitional between 

maritime and continental. The average annual temperature is +6°C and the average annual 

precipitation is 650 mm. The vegetation period lasts for 175 days. The southern part of 

Lithuania belongs to the temperate vegetation zone, the northern part to the Hemiboreal 

zone. Forest area represents 30.1 % (1 860300 ha). 

Occurrence and origin of oaks and beech 

In Lithuania deciduous hardwoods cover 4.7% of the total forest area, and pedunculate oak 

(Quercus robur L.) 1.7%. It forms mixed stands, often together with Norway spruce and 

softwood deciduous trees, and is only exceptionally found in pure stands. Oak forests are 

mostly concentrated on fertile sites of the coastal lowlands and in the central part of the 

country. 

Lithuania is crossed by the northern boundary of the distribution area of Carpinus betulus 

L. Behind this, Quercus petraea grows on an area of 70 ha in the Trako forest in the 

southeastern part of the country. This natural stand is almost 100 years old and 90% of the 

oaks growing there are Q. petraea (Tuminauskas 1957). 

Beech stands are presumably not native and grow in a small part of southwestern 

Lithuania near the Baltic Sea. 

The optimum climate for pedunculate oak and the maximum broadleaved forests cover 

reached was in 4000-5000 BP (Kabailiene 1990). Climatic changes alone were not sufficient 

to account for the decrease of broadleaved forests. It is likely that human activity further 

reduced their distribution. Biological and ecological features of Q. robur suggest that, even 

in the period of maximum distribution of the species, the share of oak forests was less than 

30% of total forest area. According to Lukinas (1967), in the 16th century Lithuania had 15- 

20% of oak stands, while in 1895 only 2-3% remained. Therefore, for economic reasons and 

biological characteristics it is not necessary to extend the area of pedunculate oak in 

Lithuanian forests above 5-7% (Karazija 1997). 

Current economic importance in the forestry sector 

Forests are one of the most valuable natural resources in Lithuania. Mature pedunculate oak 

stands produce on average 246 m 3 /ha of timber. Some stands have a very high productivity: 

the Juravos oak stand, 660 m 3 /ha at the age of 110 years; the 80-year-old Dusnioniu oak 

stand in Alytus, 343 m 3 /ha; the 120-year-old Kulupenu oak stand in Kretinga, 354 m 3 /ha, 

etc. 

The rotation period for oak ranges from 120 to 140 years. The average age of oak stands is 

80 years although the greatest part of the area is constituted of middle-aged stands (about 

58%). In 49% of the stands in which pedunculate oak occurs, it represents less than 5% of the 

species present in the stand; not more than 5% stands have more than 50% oak trees. Stands 

with oak admixture cover about 15% of the forest area. Every year about 15 000 m 3 (or about 

1.1% of the total amount of timber harvested) of mature and overmature deciduous 

hardwoods, mainly pedunculate oak and European ash, are logged. Felling of oak stands is 

increasing and will rise to 0.7% of the total growing stock in 1998. The needs of the national 

market in oak wood products are not fulfilled completely. Some oak stands are still owned 

by private owners (one-sixth) and the process of privatization is not finished yet.

118 EUF'ORGEN: SOCIAL BROADLEAVES 


The regeneration of oak stands is prescribed in the Reforestation Regulations approved by 

the Ministry of Agriculture and Forestry. The planting of other species in the areas of oak 

felling is prohibited as well as clear-cutting. For these reasons acorns have to be collected in 

the seed reserves. Seed transfer is controlled by the responsible state institutions. 

Health state of the forest stands and threats to genetic diversity 

Defoliation of oak stands affects 24% of the trees, though the investigations on oak wood 

rings made at the Lithuanian Forest Research Institute did not reveal any significant 

increment changes. Greater defoliation is observed in older stands and it represents about 

17% of the total area occupied by pedunculate oak stands. Young stands (about 16% of total) 

are to a large extent damaged by deer. Wood rot fungi (Phelinus robustus and Laetiporus 

sulphureus) heavily infect older trees on a large scale. In some cases very old trees have 

survived. One of the oldest pedunculate oaks in Europe (approximately 3500 years old) 

grows in the eastern part of Lithuania. Over 100 Q. robur trees are included in the list of 

nature conservation units. In spite of the small percentage of pedunculate oak stands in 

Lithuania, it is the 'national tree' and is mentioned very often in the national folklore. 


Research on pedunculate oak stands in Lithuania was mostly related to typology and ecology 

studies (Lukinas 1956). For conservation of genetic diversity and tree breeding purposes an 

open-pollinated progeny test (>100 families from 10 populations) was established in 1996. The 

network of pedunculate oak experimental plantations will be established in 1999. 

Protected stands represent 1.7% of the total pedunculate oak area (Table 1, Fig. 1). It is 

the highest proportion of all forest tree species. 

Scientific research on pedunculate oak genetic resources is funded directly by the State 

and by forest enterprises through the Department of Forestry ,of the Ministry of Agriculture 

and Forestry. Forest enterprises realize the need for implementing the results of research in 

the forestry practice. A State programme for oak restoration in Lithuania is in preparation at 

the Forest Research Institute. Part of this programme deals with gene conservation. The 

programme plans to increase the area of oak stands by almost three times, up to 90 000 ha, 

by using reproductive material from the best selected stands and bred material. 

In 1993-95 the Lithuanian Forest Research Institute carried out a complex study on the 

state of oak stands. The monograph on regeneration and conservation of oak stands in 

Lithuania is in press. The long-term research project on conservation of plant genetic 

resources will be initiated next year. This project will include genetic studies of oak and 

development of a programme for dynamic gene conservation in accordance with the MPBS 

(multiple population breeding system) concept. 

Table 1. Breeding and conservation units of Quercus robur L in Lithuania 

Forest No. of Seed 

ecoclimatic plus Gene reserves Seed reserves orchards Total oak stand 

region trees No. ha No. ha No. ha Area {ha} 

I 3 1 54.6 1 8.6 5239.6 

IIA 20 13 203.6 2 11.2 5429.9 

liB 3 17.7 3 17.5 2942.7 

III 7 9 43.2 3 3.6 6485.8 

IVA 35 6 93.4 2 20.7 1.2 10617.2 

IVB 1057.8 

Total 65 32 412.5 11 61.6 1 1.2 31773.0


.c. " 

1 

~/ number of objects 

ctr gene reserves 

&. seed reserves 

2 number of plus trees 

* Quercus petraea stand 

2A 

the limits of forest ecoclimatic regions 

Fig. 1. Stands of Quercus robur and mixed stands with more than 10% proportion of Q. robur in stand 

composition in Lithuania. 


The Forest Research Institute is involved in the joint project 'Resources of Domesticated 

Plants' which includes research on forest genetic resources and has been carried out since 

1994. The 'Lithuanian Open State Science and Study Fund' supports this programme related 

to the utilization of forest genetic resources. Neither national nor international oak genetic 

resources committee has been established. The Department of Forestry of the Ministry of 

Agriculture and Forestry has appointed a coordinator for international relations in the area 

of forest genetic resources. There are no specialized laws for the conservation of forest 

genetic resources; however, they are protected at Ministry level. The Law on Protected 

Territories, the Forest Law, the Law on Wild Plants still do not recognize the importance of 

gene diversity and conservation. The catalogue of Lithuanian forest genetic resources is 

prepared at the Lithuanian Forest Institute. These are registered in the National Register of 

Forest Genetic Resources at the Lithuanian Forest Tree Breeding and Seed Management 

Center. All data are compiled and continuously updated in databases. A genebank was 

recently established at the Agricultural Institute in Dotnuva. 

The state programme 'Lithuanian Forestry and Timber Industry Development' was 

approved in 1996 for the period until 2003. The 'Conception and Programme for Protection 

of Biodiversity in the Forests' is under preparation by the Lithuanian Forest Research 

Institute. The 'Conceptional Program for Forest Regeneration', approved in 1994, will last 

until 2010. It includes the perspective development of a basis for genetic diversity 

conservation. 

The Convention of Biologcal Diversity from Rio de Janeiro was ratified in 1995.

~20 EUFORGEN: SOGIAIl BROADEEAVES 

Forest gene conservation units are partly covered by the following categories in the Forest 

Law: strict reserves, reserves, protected landscape units. The definition of reserves includes 

location for botanical reserves in which forest resources are also mentioned. However, in 

general, these types of reserves are aimed at conserving the plants or mushrooms and their 

biotopes, but not the genetic diversity. The State has exclusive property rights on the 

selected genetic resources. 

The Forest Research Institute approves the gene reserves and the Ministry of Agriculture 

and Forestry funds the work undertaken for the selection of genetically valuable units. 

Institutions involved in genetic resources activities in Lithuania 

Three institutions collaborate closely in national forest genetic resources activities: 

• The Department of Forest Genetics and Reforestation of the Lithuanian Forest Research 

Institute carries out genetic studies, develops programmes and recommendations for 

selection, conservation and utilization of the forest genetic resources, forest tree 

breeding, creates PC databases on forest genetic resources. The department carries out 

the selection and designation of gene conservation units, updates and maintains the 

databases. 

• The Lithuanian Forest Tree Breeding and Seed Management Center is responsible for 

the Lithuanian forest genetic resources. The Center carries out the inventory and 

designation of in situ genetic resources, provides periodic inventory of the gene 

conservation units, controls their utilization, manages the documentation, compiles the 

data in databases. The Center also carries out forest tree breeding, guides the 

establishment of second-stage seed orchards, grows seedlings and graftings for seed 

orchards, supplies the forest enterprises with improved seed and planting material, and 

is responsible for the trade of seeds inside and outside Lithuania. 

• The Forest Seed Control Station controls the utilization of forest genetic resources, the 

origin of seed and planting material, seed transfer, seed collecting, processing and 

storage, seed quality and trade. The system for the documentation of forest reproductive 

material is based on OECD standards. 

International collaboration 

The Lithuanian Oak Society was established in 1992 in collaboration with Swedish 

colleagues. The agreement on oak diversity conservation signed by the Polish Forest 

Research Institute and the Lithuanian Forest Research Institute in 1996 will help in the 

exchange of material and information, and also deals with pedunculate oak genetic 

conservation. The most common needs are: 

• collaboration in joint research projects with technical support or assistance from other 

countries 

• exchange of information with relevant institutions of European countries and 

international organizations 

• development of appropriate policy related to forest genetic resources and use of 

effective models for its implementation. 

References 

Kabailiene, M. 1990. Lietuvos Holocenas [Lithuanian Holocen] [in Lithuanian]. Mokslas, 

Vilnius. 

Karazija, S. 1997. Azuolynu atkurimo perspektyva [Perspectives of oak regeneration] [in 

Lithuanian]. Musu girios 11. 

Lukinas, M. 1956. Teritorinis azuolynu fondas ir jo iskyrimo tipologiniai pagrindai 

[Territorial oak's fund and the principles of its ascription on typological basis] [in 

Lithuanian]. Rink. LTSR misku ukio isvystymo klausimai, Vilnius. 

Tuminauskas, S. 1957. Bekotis azuolas (Quercus petraea Liebl.) pietu Lietuvoje [Sessile oak 

(Quercus petraea Liebl.) in South Lithuania] [in Lithuanian]. Musu girios 5.

· GOUNmR¥ REPORmS ~2~ 

Oak and beech resources in Latvia 

Arnis Gailisl and Edgars Smaukstelis 2 

1 Latvian Forest Research Institute Silava, Latvia 

2 Forest Research Station Kalsnava, Latvia 

Occurrence and Qrigin of oak and beech in Latvia 

The share of boreal coniferous species (Scots pine and Norway spruce) in Latvia is 60.5% of 

the total forest area. The remaining 39.5% is covered by broadleaved forests. 

Native silver birch, downy birch, common alder, grey alder, aspen, rowan, willow species 

and bird cherry are typical boreal species and are situated in the optimum of their natural 

distribution range. 

Other indigenous broadleaved species - pedunculate oak, ash, small-leaved lime, 

Norway maple, wych elm, European white elm, European hornbeam and white poplar - are 

or could be referred to as temperate forest species. Latvia is at the northern limit of the 

species' natural range; therefore the occurrence of these species is scattered and fragmented. 

Pedunculate oak(Quercus robur L.) 

Forests with oak as the dominant species cover about 10000 ha or 0.35% of Latvia's 2.882 

million ha forest area. According to inventory data, the area of forest stands with oak as 

admixed species is about 157 700 ha or 5% of the total forest area. The growing stock of oak 

is estimated at 7.36 million m 3 (1.5% of the total 500 million m 3 growing stock). 

In spite of its insignificant share in the country's economy, oak in Latvia is regarded as a 

national symbol. For this reason, single trees are maintained as landscape elements and left 

untouched in clear-cutting areas. 

Historically, oak was much more widely distributed over the Baltic region. The main 

decline of oak forests w'as related to the development of agriculture, when the most fertile 

forest areas were felled, and with the shipbuilding in the Russian Empire at the beginning of 

the 18th century. 

Over the past 50 years forestry in Latvia was mainly oriented toward native coniferous 

species. Broadleaves (including oak) were neglected. As a result the proportion of young 

oak stands is less than 5% (Fig. 1). 

1-20 41-60 81-100 121-140 161-180 201-220 

Age 

Fig. 1. Age structure of oak forests in Latvia.

122 EUFORGEN: SOCIA~ BROAD~EAVES 

The uneven distribution of oak forests does not seem to be linked with definite climatic 

conditions or agroclimatic zones (see Fig. 2). Few forest districts where oak stands are more 

than 1 % can be found. 

CJ-O,19 00,2-0,39 _ 0,4-0,59 _ 0,6-0,99 _ 1- 

Fig. 2. Distribution of oak forests in Latvia (percentage of total oak area). 

Sessile oak(Quercus petraea (Matt.) Liebl.) 

Quercus petraea is an exotic species introduced from unknown ongms, represented in 

separate forest parks mostly in the western part of Latvia. The latest introductions, in the 

past 15 years (Arboretum of the Forest Research Station 'Kalsnava,), originate from 

Lithuania, Kaliningrad Oblast (Russian Federation) and the Ukraine. 

Beech (Fagus sylvatica L) 

Introduction of Fagus sylvatica started in the middle of the 19th century, from unknown 

origin(s). The first beech stands were established in 1885 in western Latvia. A quite severe 

climate and temperature extremes (down to -45°C) prevented beech introduction in eastern 

Latvia. At present beech plantations are located only in the west. 

The total area covered by pure or mixed beech stands is 210 ha (the age distribution is 

presented in Fig. 3). Recent studies showed that cultivation of beech was successful. It is 

possible to obtain high-quality timber. The following example can be mentioned: within 

one small stand at the age of 101 years, the average tree height was 36 m, DBH 36 cm, stem 

volume 1.63 m 3 and the mean growing stock 867 m 3 /ha. 

The older stands produce quite viable seed (24-81%) in different seed years. It seems that 

young stands mostly appear from natural regeneration around the older ones. 

Besides beech plantations in the forest, single trees were planted in parks, represented by 

different ornamental forms. The most common are Fagus sylvatica 'Pendula', 'Laciniata', 

'Purpurea' and 'Tricolor'. 

According to the data of the National Botanical Garden, separate beech trees and their 

ornamental forms are registered in more than 100 parks. 

Other species include Fagus orientalis 1. (a few trees in the National Botanical Garden) and 

F. grandifolia (represented in Latvia only by one tree in the Skriveri Arboretum).


Fig. 3. Age structure of beech forests in Latvia. 


Oak 

As already mentioned, the current economic role of oak is insignificant because of its scarce 

distribution and national traditions to maintain single trees after felling. 

At the same time it should be mentioned that the importance of oak could significantly 

increase in the private forestry sector and be used for forthcoming afforestations. Therefore 

it was expected that the share of oak stands in the future would increase up to 200000- 

250000 ha (i.e. 7-9% of the total forest area) 

Beech 

Beech would be successfully used as a commercial species in the western part of Latvia by 

using the locally adapted seed material, after obtaining knowledge about the genetic 

structure of first beech introductions, and studying the possibilities of new introductions. 


Oak 

According to the latest observations, the old stands (120-250 years) are mainly regenerated 

naturally. Starting from the end of the 19th century artificial plantations were established, 

frequently with reproductive material of unknown origin. At present we can find very few 

successful plantations with good productivity at the age of 100 years and more. 

Normally oak on fertile soil forms mixed stands with aspen, Norway spruce, ash, lime 

tree, grey and common alders and maple, on less fertile soil with birch and Scots pine. 

Over the past 50 years forestry in Latvia was mainly oriented toward native coniferous 

species. Broadleaves (including oak) were neglected. The area of annual oak reforestation is 

shown in Figure 4. The survival rate in these reforested areas is about 10-15%. 

As a result today we have little knowledge about appropriate oak regeneration, 

afforestation and silvicultural methods.

~24 EWE0RGEN: S0GIAt.: BR0ADt.:EAVES 

1948 1958 1968 1978 1988 

Fig. 4. Area of oak regeneration, 1948-88. 

Beech 

No appropriate programme has been developed yet. It was suggested that after the 

provenance trials (including the relevant provenances from Europe) have been evaluated, it 

would be reasonable to include beech in common silvicultural practice, mainly in the 

western part of Latvia. 

Health state of the oak stands and threats to their genetic diversity 

The mentioned diseases do not cause significant threats to genetic diversity of oak forests in 

Latvia. Decline of pedunculate oak stands has become a serious problem since the early 

1980s. The typical symptoms are as follows: yellowing of leaves, thinning of crown, dying 

of branches, numerous water sprouts, necrotic patches in bark and phloem, discolouration of 

sapwood, loosening of bark, root deformation, dying of the thin roots. The frequency of 

these symptoms and fungi present in the necrotic tissues has not been investigated in Latvia. 

Investigations on oak decline in Europe have led to the hypothesis that a combination of 

unfavourable weather conditions (drought, very wet spring and dry summer, low 

temperature affecting overground parts of trees), unfavourable soil conditions (especially 

too much or too little water) and repeated defoliation by insects has weakened oaks to such a 

degree that they can easily be invaded by fungi. It may be supposed that the fungi are not 

the primary cause of the symptoms of oak decline. Microsphaera alphitoides Griff et Maubl. 

(powdery mildew), Armillaria spp. (root pathogen) and fungi of the genus Ceratocystis are 

most frequently reported in connection with oak decline, although no causal relationship 

has yet been established between the presence of Ceratocystis spp. or Ophiostoma spp. and the 

intensity of decline. 

Wild game cause damage in young oak plantations or naturally regenerated stands. 

The different degree of frost damage in oak stands could be related to origin. At the same 

time it should be mentioned that up to now we have no information about the possible 

sources of introduced oak, nor on the degree of pollution of the local genepool with 

allochthonous origins.


Research activities 

Three research projects on broadleaves were started in recent years. Their main objectives 

are: 

• Inventory of all Latvian forests (including oak) with the aim to: 

select seed stands 

establish gene reserve forests 

study the genetic structure of species. 

• Collection of seed material from selected stands. 

• Growing of planting material for provenance (population) trials and progeny tests. 

A first provenance trial was established in 1996, where eight local oak stands are 

represented at two locations. 

Further provenance trials will be established in spring 1998 at three locations with 18 

local stands represented. 

The genetic structure should be studied with molecular markers to determine the 

effective size of in situ conservation stands, genetic reserves and ex situ collections. 

Current oak genetic conservation activities 

In situ conservation 

• Since 1986: 

Cesvaine district forest - 60.3 ha 

Gauja National Park - 65 ha 

• Newly approved (1997): 

Aizpute district forest (Apriki) - 322 ha 

• Combined with tree improvement programmes: 

Seed stands - 15 stands, 72 ha 

Ex situ conservation 

• Seed orchards: 1 ha (1972) 

• Clonal archive of Apriki genetic reserve. 

Relevant nature conservation activities 

In addition to genetic conservation and according to traditions since 1924, about 800 old 

'noble oaks' with DBH> 1.3 m were registered and maintained throughout the country. The 

age of these trees is about 300-1500 years. This activity, performed by the National Botanical 

Garden, is to be continued. 

Genetic conservation is provided by protected forests: 

• in nature reserves, 

• in specially protected forest compartments, 

• in 600 protected old parks or forest parks (oaks were mainly introduced in these 

locations) - total area about 2200 ha. 


Oak 

Tree-improvement activities are aimed at: 

• the establishment of seed sources/ seed orchards for each agroclimatic region 

• ensuring improved stem quality and adaptability to changing environment 

• the elaboration of appropriate afforestation and silvicultural methods.


For these reasons, 

• a seed orchard was established in eastern Latvia (area = 1 ha) 

• progeny' tests are to be established in spring 1998 (36 open-pollinated families) 

• 50 clones were grafted for a new seed orchard in western Latvia 

• a new plant growing technology was developed (containers), allowing the accelerated 

establishment of trials 

• selected seed stands include 15 stands, 72 ha. 

Beech 

A first trial was established in 1997 using 15 local seed sources (both forest stands and old 

parks). 


The use of broadleaves is expected to increase in the future for: 

• afforestation needs (according to information from different sources, there are at 

present in Latvia about 300000-500000 ha of abandoned agricultural land) 

• increasing the species diversity in the commercial forests. 

The technology for container plants was developed during the past 2 years. Container 

plants will be used for the trials mentioned above. 


• Forest Research Institute 'Silava', in close collaboration with: 

• Forest Research Station 'Kalsnava' 

• East Tree breeding and seed procurement centre (within 'Kalsnava') 

• West Tree breeding and seed procurement centre in Kuldiga 

• the National Botanical Garden. 


• European oak provenance trials 

• Joint genetic studies of oak and beech 

• Technical guidelines for in situ and ex situ conservation of Social Broadleaves 

• Integrated Social Broadleaves databases (genetic resources) 

• Improved silvicultural methods (exchange of literature and experience).


Genetic resources of oaks and their conservation in Finland 

AnuMattila 

Foundation for Forest Tree Breeding, Helsinki, Finland 

Occurrence and origin of oaks in Finland 

Forests cover two-thirds, or over 200000 km 2 , of the total area of Finland. There are over 20 

indigenous tree species growing in Finland, but only Scots pine (Pin us sylvestris), Norway 

spruce (Picea abies) and silver birch (Betula pendula) are commercially important at the national 

level. These three species contribute to 96% of the total growing stock of 1886.6 million m 3 • In 

addition a few species, downy birch (B. pubescens), Carelian curly birch (B. pendula var. carelica), 

Siberian larch (Larix sibrica) and aspen (Populus tremula, P. tremula x P. tremuloides), have some 

commercial significance in forestry. The most abundant broadleaved tree species (B. pendula, 

B. pubescens, P. tremula and Alnus spp.) dominate 8.2% of the forest area, and only 0.1% is 

dominated by other broadleaved trees. Pedunculate oak (Quercus robur) is the main deciduous 

hardwood species of interest to forestry in Finland. It is also the only native oak. 

The natural range of pedunculate oak is narrow and restricted to the southwest corner of 

the country, but in favourable environments it can successfully grow at least 100 km 

northward. The present oak populations mainly occur as small patches rather than as large 

and continuous stands. Beech (Fagus sylvatica) does not occur naturally in Finland, and the 

cultivation attempts have not been successful in the mainland, but it might be grown on the 

island of Ahvenanmaa. 

With the exception of a few arctic plants, species have migrated to Finland during the 10000 

years after the latest glaciation. Hardwoods such as oak migrated about 9000 years ago mainly 

from the east, but also across the Gulf of Finland, and later on from the west. During the warm 

and humid Atlantic period 7500-4500 years ago their distribution was largest and extended 

about 400 km further north. The following slow cooling of the climate favoured Norway 

spruce, which spread vigorously from the east, whereas hardwoods withdrew to the south. 

More recently the demand for agricultural land has resulted in the fragmentation of these 

previously continuous forests. Oak was also used extensively for shipbuilding, and in the 18- 

19th centuries oak trees had a special status, belonging to the Crown. This resulted in the 

destruction of oak in certain areas as Finnish landowners drowned their oak logs in the sea 

rather than hand them over to the Crown. During the last few decades forest management 

practices were unfavourable to oak as well, as many suitable sites were turned into pure spruce 

plantations. 

Importance for the forestry sector 

Broadleaves in general are of minor importance for forestry in Finland. Statistics are available 

for birch, oc~asionally also for aspen and alder, whereas all other broadleaves are grouped 

together. In 1994 the share of broadleaved trees other than birch in the growing stock was 

3.1 %. Broadleaves are annually planted on about 500 ha while the yearly regeneration areas 

amount to 180 000 ha. Annually 150-180 million seedlings are delivered from about 30 central 

nurseries (88%) and 70 smaller family-owned nurseries (12%). The quality of nursery seedlings 

in Finland is supervised by the Ministry of Agriculture and Forestry in accordance with the 

Forest Reproductive Material Act of 1979 and the related Decision of 1992 (No. 1533/92). 

Recent changes in Finnish forestry policies have increased the interest for growing 

hardwoods; especially alder (Alnus glutinosa) and pedunculate oak have been suggested as 

alternatives to conifers and birch in afforestations. The interest arises rather from the will to 

protect the cultural landscape than from forestry. These species have for centuries been an 

essential part of the landscape in southwestern Finland, but in some cases all afforestation 

plans have caused local conflicts: open fields are considered at least as important for the 

maintenance of the cultural landscape.

128 EUEORGEN: SOCIAL BROAOLEAXlES 

Valkonen et al. (1995) made an inventory of cultivated stands in Finland. The present 

natural and artificial oak stands are largely of too inferior quality to be of interest for the 

industry, e.g. owing to stem defects. Moreover, the present resources without protection 

mesures correspond to only a few year's use. The higher costs (e.g. growing tubes and/or 

fencing to protect from vole, hare and moose) and labour-intensive management required for 

producing high-quality timber makes the profitability of oak growing difficult to predict. 

The wood of oak and beech is valued in traditional carpentry, and oak is also used for 

several special purposes. Louna and Valkonen (1995) made an inventory of the role of Finnish 

raw material in the industrial use of broadleaves. The value of the import of Noble 

Hardwoods 1 timber in 1993 was 150 million FIM, of which the value of oak and beech import 

were 56 and 51 million FIM, respectively. Oak is mainly imported from other European 

. countries (Q. robur, Q. petraea), Canada, the USA (Q. alba, Q. rubra) and China (Q. mongolica). 

There are no commercial beech plantations in Finland, and all beech wood is imported. The 

Finnish resources of oak wood are of very limited local importance: less than 100 in 3 of Finnish 

oak wood is yearly on the market, whereas in 1993 the total oak wood use was 26 400 m 3 • 

Oak, ash and maple have a well-established, rather stable market share in traditional 

carpentry, and beech is largely used as an alternative for oak. In 1993, 11.7% of all parquets 

were made of oak, covering 75% of the total oak wood consumption. Beech was used for 9.4% 

of all parquet production. Oak is also used for several small-scale purposes, e.g. for 

constructing boats and parts of musical instruments. 

Selected seed sources 

Three oak seed collection stands have been selected, and the total number of seed collection trees 

is about 250. New selection is under way as additional natural or artificial stands with good 

quality for seed sources are needed. So far the only oak seed orchard was established by the 

Forest and Park Service in 1978 with 279 grafts from 24 clones. In 1995 the first seed crop was 

collected, but the material was not used for forest regeneration because of the low number of 

flowering clones. 

Conservation 

The important role of Noble Hardwoods including oak and beech for the conservation of 

biodiversity in general has been widely acknowledged in Finland, and various programmes 

emphasize the importance of diversity at the genetic level. Pedunculate oak is not 

threatened as a species in Finland, but the popula:tions are susceptible to increasing 

fragmentation by road construction and enlargements of settlements. Most of the natural 

oak stands are protected. Genetic conservation, however, may have special needs that are 

not met by strict habitat conservation. Many oak stands would benefit from light 

management to promote natural regeneration. On the other hand, many of the stands are 

fairly small and isolated, and thus inbreeding and genetic erosion may play a role in them. 

Conservation practices should take into account the underlying genetic constitution of the 

stands and more knowledge, especially on the structure of small and isolated populations, is 

needed. Results of the population genetic research suggest that the differentiation among 

populations at neutral markers is more pronounced in oak than in the more common tree 

species (Mattila et al. 1994; Mattila and Vakkari 1996). 

The general in situ gene reserve forest programme was started in 1992, and four Noble 

Hardwoods stands are included in the total of 33 stands (by the Finnish Forest Research 

Institute (FFRI), see Rusanen 1996). So far there are no gene reserve forests for pedunculate 

oak. For the main tree species the minimum area of a gene reserve forest is 100 ha, and 

silvicultural measures such as thinning and harvesting are allowed. However, for the 

conservation of Noble Hardwoods the gene reserve forest system is only seldom applicable. 

Most of the valuable stands are included in various forest protection programmes, which 

I Noble Hardwoods as defined in Finnish forestry include oak and beech (see Rusanen 1996).

COUNTRY'REPORTS 129 

prevents the active measures used in gene reserve forests and required for gene conservation 

(such as collecting of seeds). Also the minimum target area of the main tree species can 

seldom, if ever, be met in Noble Hardwoods stands. There is only one large oak stand in 

Finland (Ruissalo, Turku), where oak dominates on 90 ha, of which 50 ha is pure oak; all 

other oak stands are significantly smaller. 

The main approach to the conservation of the genetic resources of oak, and other Noble 

Hardwoods, will be in ex situ collections. The operational ex situ plan for oak (by FFRI) has 

been developed. The first seed collections were made in 1995, land areas have been chosen 

and their preparation is in progress. In the future these collections may also serve in the 

production of forest reproductive material. 

Breeding and research 

There is no intensive breeding programme for pedunculate oak in Finland. Three progeny 

tests were established in 1985-93 (in total 4150 individuals, by FFRI), and in 1995 a joint 

research project on oak cultivation practices (by S. Valkonen, FFRI: sowing/planting, 

individually /in groups, growing tubes/no tubes) and on the distribution of quantitative and 

adaptive genetic variation in Finnish oak stands was started. Five progeny trials will be 

established in 1998 by the Foundation for Forest Tree Breeding (FFTB). 

A common research project on the genetic structures of Noble Hardwoods was started in 

1994 by the FFTB and the FFRI. The results will give some insight into the genetic processes 

in fragmented populations, and thus help to make operative plans for genetic conservation, 

but they will also be helpful in giving recommendations for the selection of seed sources for 

forest regeneration. Estimates of the level and distribution of neutral genetic variation are 

assessed by isoenzymes in pedunculate oak, Norway maple (Aeer platanoides) and European 

white elm (Ulmus laevis). The first results support the hypothesis that the population 

structure of rare species with a scattered distribution is different from that of the main tree 

species (Mattila et al. 1994; Mattila and Vakkari 1996). A new joint project, financed by the 

Academy of Finland and based on the isoenzyme work, was started in 1997 to acquire 

knowledge on the genetic processes (family structures, pollen migration, etc.) in oak and 

maple populations by microsatellite markers. 

To elucidate the colonization history of oak in Finland R. Vi:iinbli:i (Univ. of Helsinki) used 

chloroplast DNA (cpDNA) markers (in collaboration with C. Ferris et al., Leicester Univ., 

UK). 

The rate and potential for pollen migration between oak stands was evaluated in 1995 by 

direct measurements on the concentration of airborne oak pollen within and outside the 

largest oak stand in Finland. There was a significant decrease in the concentration of oak 

pollen in the air at a short distance from the pollen source (Lahtinen et al. 1996). 

Differences between species in the onset of the hardening process and in maximum frost 

tolerance as measured by, e.g. differential thermal analysis, differential scanning calorimetry 

and magnetic resonance imaging, will be examined by T. Repo (Univ. of Joensuu). The 

project started in 1997; the species involved are pedunculate oak, Norway maple, wych elm 

(Ulmus glabra) and birch. 

Public awareness 

Forests are an important element in the daily life of virtually every Finn. In Finland onethird 

of the forests is owned by the state, less than 10% by companies and about 55% by 

private persons. The number of private forest holdings is 439000, with an average size of 

28 ha of forest land. Forests are open for everybody's unrestricted recreation use based on 

public rights. 

The predicted climate change has also increased the interest in Noble Hardwoods 

(including oak and beech) as the warming of the climate might make their cultivation north 

. of their present natural distribution possible. The forests are also subject of public debate, 

with topics ranging from conservation to economic aspects of forests and timber production.


As nature conservation has become popular and acceptable within the last decade, it has 

inevitably resulted in changes within the forestry sector. In the 1990s, forestry has 

increasingly emphasized the importance of forest ecology and the promotion of multiple use 

of forests, and both nature conservation and forest legislation have been revised (Nature 

Conservation Act 1096/96, Forestry Act 1093/96). 

References 

Lahtinen, M.-L., P. Pulkkinen and M.-L. Helander. 1996. Potential gene flow by pollen between 

English oak (Quercus robur) stands in Finland. Pp. 46-50 in Conservation of forest genetic 

resources. Proe. of Nordic Group for Forest Genetics and Tree Breeding Meeting (M. Kurm 

and U. Tamm, eds.), 3-7 June 1996, Tartu, Estonia. 

Louna, T. and S. Valkonen. 1995. Kotimaisen raaka-aineen asema lehtipuiden teollisessa 

kayt6ssa. Metsantutkimuslaitoksen tiedonantoja 553. 

Mattila, A and P. Vakkari. 1996. Genetic variation of Quercus robur and Ulmus laevis in Finland. 

Pp. 63-68 in Conservation of forest genetic resources. Proe. of Nordic Group for Forest 

Genetics and Tree Breeding Meeting (M. Kurm and U. Tamm, eds.), 3-7 June 1996, Tartu, 

Estonia. 

Mattila, A, A Pakkanen, P. Vakkari and J. Raisio. 1994. Genetic variation in English oak 

(Quercus robur) in Finland. Silva Fennica 28:251-256. 

Ranta, H. 1996. Tammen ja eraiden muiden lehtipuiden tuholaiset Suomessa ja Euroopassa; 

lajisto, merkitys ja ilmastonmuutoksen seuraukset [Pests of oak and certain broadleaved 

trees in Finland and Europe; species, significance and implications of climate change.] 

Metsanjalostussaati6n tiedonantoja 12. 

Rusanen, M. 1996. Noble hardwoods genetic resources and their conservation in Finland. Pp. 

155-157 in Noble Hardwoods Network. Report of the first meeting, 24-27 March 1996, 

Escherode, Germany (J. Turok, G. Eriksson, J. Kleinschmit and S. Canger, compilers). IPGRI, 

Rome, Italy. 

Salonen, K. 1997. Research on genetic diversity of Finnish nature (English summary). Finnish 

Environment Institute. Suomen ymparist6keskuksen moniste 85. 

Statistical Yearbook of Forestry. 1996. The Finnish Forest Research Institute, Helsinki. 

Valkonen, S., S. Rantala and A Sipila. 1995. Jalojen lehtipuiden ja tervalepan viljely ja 

kasvattaminen. Sammandrag: Odling och uppdragande av adla l6vtrad och klibbal. 

Metsantutkimuslaitoksen tiedonantoja 575.


Social Broadleaves in Sweden 

Lennart Ackzell 

National Board of Forestry, J6nk6ping, Sweden 

None of the Social Broadleaves occurring in Sweden (Quercus robur, Quercus petraea and 

Fagus sylvatica) reaches the boreal zone, approximately delimited by latitude 60 0 N. These 

species altogether make up less than 2% of the growing stock of forests in Sweden, 

predominantly composed of Scots pine and Norway spruce. 

The regeneration methods used are planting in the case of oak and natural regeneration 

for beech. The planting stock for oak is to a large extent imported because relatively small 

quantities are required by users. Plantations of oak ha·ve to be protected from browsing by 

wild game. The health status of these species is considered good despite some indications of 

recent decline due to air pollution and climatic factors. These are currently under study. 

Organizations involved in research and gene conservation activities on Social 

Broadleaves are the Swedish University of Agricultural Sciences (SLU), the Forestry 

Research Institute and the National Board of Forestry. Research activities on silviculture 

take place at SLU's southern research locality of Alnarp. Genetics is studied at SLU in 

Uppsala. The Swedish Forestry Research Institute (SkogForsk) maintains provenance trials 

for beech and breeding activities. 

The Swedish Forest Gene Conservation Programme conducted by the National Board of 

Forestry comprises conservation activities for oak. Six gene conservation archives were 

established in 1995 (Fig. 1). They consist of a total of 20000 seedlings from 513 families of 

autochthonous origin. It would be desirable to perform cpDNA (chloroplast DNA) analyses 

on this material. 

Fig. 1. Six gene conservation archives.

132 EUFOBGEN: SOCIAl.. BROADl..EAVES 

The genetic resources used for forest regeneration represent 115 selected seed stands of 

Quercus robur and 17 selected seed stands of Quercus petraea. Two seed orchards of oak are 

being tested. Nature protection activities concentrate on 5500 ha of hardwood forest (mainly 

oak and beech) in the three southern provinces of Sweden. 

Sweden is very interested in the international collaboration concerning genetic resources 

of Social Broadleaves. As it is on the northern fringe of the species' distribution areas, the 

country can contribute to comprehensive studies and conservation efforts extending over the 

entire geographic range.

OVERVIEW PRESENTATIONS 133 

Overview Presentations 

Structure of gene diversity, geneflow and gene conservation in 


Antoine Kremer, Rimy J. Petit and Alexis Ducousso 

INRA, Laboratoire de Genetique et d'Amelioration des Arbres Forestiers, Gazinet Cestas 

Cedex, France 

Abstract 

A review of the genetic diversity of Quercus petraea (sessile oak) for gene markers and 

adaptive traits is presented. Remarkable range-wide trends of variation exist in sessile oak 

for all traits investigated and for the corresponding measures of diversity. We illustrate 

these trends for isoenzyme and quantitative trait data (bud burst and height growth 

assessed in a provenance test). A comparison with cpDNA (chloroplast DNA) information, 

which had been shown to closely reflect postglacial recolonization routes, allows testing to 

determine whether these general genetic trends were established during or after 

recolonization, or more recently, as a consequence of isolation-by-distance or adaptation to 

local environments. Results obtained so far indicate that contrasting trends can be observed. 

For example within-population diversity for bud burst follows a longitudinal pattern of 

variation, whereas population differences for the same trait follow a latitudinal pattern of 

variation. The situation is even more complex when different traits are compared. A general 

conclusion is that, although the geographic structure of diversity is quite strong, there is no 

constant trend across traits that would allow sampling of specific populations for in situ 

conservation. The review extends to investigations on inter- and intraspecific geneflow in 

Q. petraea. It is shown that Q. petraea is almost exclusively an outbreeding species, 

hybridizing in natural populations with Q. robur, and most likely hybridizing with other 

related species. Furthermore experimental results obtained with paternity analysis suggest 

pollen flow at rather long distances. As a result geneflow through pollen is considered a 

major mechanism for conserving diversity. Because of the contradictory conclusions that 

may be drawn depending on the traits considered, the geographic structure of diversity or 

differentiation would not be recommended as relevant criteria for in situ conservation 

decisions. Rather, methods conserving mechanisms contributing to the maintenance or 

increase of diversity as inter- and intraspecific geneflow are considered of primary 

importance for in situ conservation decisions. 

Introduction 

Gene conservation decisions rely on scientific and socioeconomic criteria. For the scientific 

community, the challenge is to provide policy-makers with relevant information on the 

potential of a species to adjust to future predictable or unpredictable ecological conditions. 

This task requires an inventory of the existing genetic diversity and insights in its future 

dynamics. In the case of Quercus petraea (sessile oak), a major component of European 

forests, our aim in this contribution is to update the current knowledge of the distribution 

and organization of genetic diversity at different spatial scales, but also to pinpoint the 

mechanisms that generate or reduce diversity. In the case of oaks, not only intraspecific 

geneflow but also interspecific geneflow should be considered as a prevalent force for 

shaping diversity. 

During the past 10 years extensive literature was published on the distribution of genetic 

diversity in the sessile oak. A common characteristic of these studies was their large


sampling of populations covering most of the natural range. Results from studies conducted 

at range-wide scale clearly indicated clinal trends of variation. Zanetto and Kremer (1995) 

showed that most of the allozymes exhibited longitudinal patterns of variation. Similarly 

Dumolin-Lapegue et al. (1997) confirmed earlier observations of Petit et al. (1993) that 

cpDNA polymorphism was strongly differentiated among populations along an east-west 

gradient. Finally, results from provenance tests showed that bud burst was most likely to 

follow a latitudinal trend of variation (Ducousso et al. 1996). As data on genetic diversity 

increased, interests in the Quaternary history of oaks was raised as well. Compilations of 

fossil deposits on a European scale in the form of isopollen maps drawn at different time 

slices (Huntley and Birks 1983) had revealed that deciduous oaks recolonized Europe from 

three separate refugia and had occupied most of the extant range about 7000 years ago. The 

recolonization routes have further been reconstructed with the help of cpDNA 

polymorphism (Dumolin-Lapegue et al. 1997). As a result, today's oak stands may still be 

considered as 'young' on an evolutionary time scale. This raises the question of whether the 

origin of the populations, in terms of the glacial refugium they are stemming from, still has 

an impact on today's distribution of diversity. The alternative hypothesis is that genetic 

differences between the colonizing populations were erased by extensive pollen flow 

favoured by the continuous distribution of the sessile oak in Europe. Finally historical 

impacts and ongoing geneflow may have been rapidly counterbalanced by local selection 

pressures, especially for phenotypic traits. 

In this contribution we attempt to re-analyze some of the data previously published with 

the aim to depict major trends of variation for different criteria relevant to the structure of 

genetic diversity: the level of within-population diversity and the differentiation among 

populations. We will particularly compare results of genetic markers, assumed to be 

neutral, with those of phenotypic traits assumed to be adaptive, by computing parameters 

making comparisons possible. Because oaks are species for which the recent Quaternary 

history is now well understood, we will address the issue of the potential historical impacts 

on the structure of diversity. Finally inter- and intraspecific geneflow will receive particular 

attention as key mechanisms for future evolution of gene diversity. 

Geographic structure of within-population diversity 

The data set for the geographic structure of diversity and variability 

One hundred populations sampled throughout the natural range of the species were 

collected. Harvests were made as bulk collection on a surface varying between 10 and 20 ha 

(Zanetto and Kremer 1995; Le Corre et al. 1998a). The seeds were then "distributed for 

different analyses: 

• Assessment of allozyme diversity on 13 isoenzyme loci (Zanetto and Kremer 1995), 

with a sample size of 120 seeds per population. 

• Installation of a provenance test comprising a subset of 69 provenances among the 100 

collected. The provenance test is located in the Northwest of France (Petite Charnie 

forest) according to a complete block design comprising 10 repetitions with a plot size 

of 24 trees (Ducousso et al. 1996). Total height was measured at age 7 (4 years after 

plantation) and bud burst was assessed at age 6 according to a" grading system 

varying from 0 (dormant bud) to 5 (elongating flush). 

• Assessment of cpDNA diversity on the basis of 5 sampled trees per provenance. 

Protocols for the cpDNA survey are given in Dumolin-Lapegue et al. (1997).


Geographic trends of within-population diversity 

For allozymes, within-population diversity was estimated by calculating the mean number 

of alleles per locus (A) and Nei's mean gene diversity (H) (Nei 1987), which is equivalent to 

the expected heterozygosity per population. For phenotypic traits, levels of withinpopulation 

genetic variability were estimated by computing the coefficient variation of 

additive genetic values (CVA' Kremer 1981; Houle 1989). The CV A values were obtained by 

assuming that heritability (h 2 ) was of similar magnitude in all populations: 

where CV A , V plt 

and X are the coefficients of genetic variation, the phenotypic variance and 

the mean value of the phenotypic character in a given population, respectively. 

Heritability values of the two phenotypic traits were considered to be 0.10 for height 

growth and 0.30 for bud burst. These values were chosen from the review on heritability 

values in forest trees (Cornelius 1994). Because actual values may be different for the two 

characters than those chosen here, levels of CV A 

will not be compared among the two 

characters, but only between populations for a given trait. 

Sessile oak is extremely diverse and variable at a within-population level, as illustrated by 

several inventories conducted with either gene markers or phenotypic traits. For isoenzymes 

in this data set, the mean number of alleles per locus averages 2.73, and expected 

heterozygosities 0.245, which makes Q. petraea among the most variable tree species. 

Additive coefficient of variation amounted to 0.39 (for constant heritabilities of 0.30) and to 

0.11 (for constant heritabilities of 0.10). However, in the same time, Q. petraea also exhibits 

an important inbreeding depression (Kleinschmit and Kleinschmit 1996); part of the 

diversity may therefore be due to the genetic load that exists in oaks. Experimental evidence 

of genetic load is suggested by segregation distortion that occurs quite often in control 

crosses of oaks (Miiller-Starck et al. 1996; Zanetto et al. 1996). 

To depict the geographic distribution of diversity, correlations between the withinpopulation 

levels of diversity for phenotypic traits and isoenzymes and three geographical 

variables (latitude, longitude and altitude) were calculated (Table 1). Six of the 12 

correlations are significant, and each of the four measures of diversity is correlated with 

either latitude or longitude, demonstrating that diversity is always spatially organized in the 

sessile oak at the studied scale. The following associations are significant: 

• bud burst is more variable within population in the western part of the range and at 

high elevations 

• height growth is more variable within population in the south 

• the number of alleles per population increases toward the east (Zanetto and Kremer 

1995) 

• the expected heterozygosity increases toward the west and decreases with altitude 

(Zanetto and Kremer 1995). 

Flowering and fruiting are known to be extremely variable across stands or years. 

Climatic constraints increase in the eastern part of Europe, where late frosts occur more 

often and impede regular flowering. One would therefore expect that geneflow among 

populations might be higher in western than in eastern populations. As a consequence, 

within-population diversity or variation is likely to be maintained at higher levels in the 

west than in the east. This could explain why bud burst, for example, exhibits higher CV A 

values. Furthermore bud burst is also highly differentiated among populations (see Table 3).

136 EUFORGEN: SOCIAL BROADLEAVES ~ 

Table 1. Correlation between four genetic diversity measures and geographic data 

CV A 

CV A 

bud burst height Number of alleles Expected heterozygosity 

N 64 65 100 100 

Latitude 0.16 NS -0.33** 0.18 NS 0.12NS 

Longitude -0.35** -0.21 NS 0.43** -0.53** 

Altitude 0.38** O.OONS 0.06 NS -0.31 ** 

* = Significant at P=0.05; ** = significant at P=0.01; NS = nonsignificant at P=0.05. 

N = number of populations. 

Geneflow between highly differentiated populations contributes to maintaining high 

levels of within-population variation. However geneflow should increase diversity for all 

traits, including isoenzymes. Our results indicate that allozyme diversity is higher in the 

east. We would advocate the higher diversity existing in the eastern refugia to interpret the 

contrasting conclusions between the geneflow hypothesis and the observed data. Another 

possibility is that the highest within-population diversity of bud burst in the west may be 

related to a less predictable climate in the spring in this part of the range, and especially the 

occurrence of late spring frosts that can damage growth of early flushing phenotypes 

(Ducousso et al. 1996). On the contrary, in the east, springs are usually shorter, and 

diversifying selection may not be so high. The significant relationship of bud burst 

variability and altitude still need to be explained. Defoliating insects may somehow play a 

role but this remains to be investigated. 

That height growth is more variable in the south may be related with the fact that 

selection for this trait is relaxed there: this would be the case if not all small individuals were 

eliminated there. Scattered cases of hybridization with the short-sized pubescent oak 

(Quercus pubescens), which can be found in sympatry with sessile oak in the south, could also 

account for this pattern. 

The causes for the strong longitudinal pattern of measures of diversity based on 

isoenzymes data have been discussed by Zanetto and Kremer (1995). The opposite trend for 

the two diversity measures (allelic richness and heterozygosity) is remarkable. Given the 

rare occurrence of sessile oak in the Iberian peninsula, it is possible that this species was 

absent or rare there during the last ice age, hence the lower allelic richness in the west. 

Furthermore, there are many closely related species or subspecies of the sessile oak in the 

Balkans, but not in western Europe, which may also have contributed to the higher allelic 

richness in the east. For heterozygosity estimates, it seems difficult to stick to neutral 

explanations for the trend observed. Interestingly, it appears that the regions displaying the 

highest heterozygosity correspond to the best growing areas for sessile oak in Europe. It is 

possible that rare alleles are counterselected in the climacic range of the sessile oak, which is 

centred to the west of Europe, which would account for the difference of allelic richness but 

not of heterozygosity. Finally, it should be recalled that for cpDNA, more haplotypes had 

been detected in the southern part of the range: only a subset of these maternal variants have 

indeed migrated up to the northern part of the range of the species (Dumolin-Lapegue et al. 

1997). 

Geographic structure of population differentiation 

Levels of population differentiation 

New methodological tools are now available to compare the subdivision of genetic diversity 

for molecular markers and quantitative traits. For allozymes, population differentiation was 

calculated as G st 

(Nei 1987). It has been shown that when the number of populations is high, 

then G st 

becomes very close to Weir and Cockerham's (1984) e (Chakraborty and Leimar

G"ER"IEW PRESENlTAlTlGNS ~ 137 

1987). Population differentiation for phenotypic traits was derived from ANOVA following 

the method described in Kremer et al. (1997) as 

F,., 

v;, 

where Vp and V ph are the between-popul


The following associations are significant: 

• provenances from southern populations flush earlier (Ducousso et al. 1996; Deans and 

Harvey 1996; Stephan et al. 1996) 

• provenances from higher altitudes flush earlier (Deans and Harvey 1996; Ducousso et 

al. 1996) 

• western provenances grow more than eastern populations 

• there is a clear east-west trend of variation for allozyme frequencies (Zanetto and 

Kremer 1995). 

For illustration purposes, the maps of the mean score for bud burst, the mean height 

growth, and the allelic frequencies of allele 7 at the isoenzyme locus Aap are provided in 

Figures I, 2 and 3. 

·1 

~~ . 

&J7 

1f~ 7 

~ :'~~ 

. 6 S --\ 

Uf 

kYS 

~~.:Jj 

~ 

') 0 

~ ·0' 

Budburst l '-->-A •• ~ • o~ 

~o Cl· 0 • ~ 

• Early ~ * .~o 

o Late ~ c:fl~ 

* provenance:Jest 

~---- .• 

~ . 

o 

• 

Fig. 1. Mean bud burst score. Values above the overall mean (early flushing individuals) are in black, 

values below the mean (late-flushing) in white. The larger the circle, the more different is the value from 

the mean. 

A discussion concerning the origin of this trend may be found in Ducousso et al. (1996). 

They likely reflect adaptation to cold and warm conditions and to predators. Southern 

provenances are flushing earlier than northern as found in three separate studies (Deans and 

Harvey 1996; Ducousso et al. 1996; Stephan et al. 1996). However, it is remarkable that this 

extremely clear-cut latitudinal trend is the opposite to that documented for northern red oak 

and for most tree species, including conifers, where late-flushing individuals are typically 

found in the south.


Fig. 2. Mean height growth. Values above the overall mean are in black, values below the mean in 

white. The larger the circle, the more different is the value from the mean. 

Fig. 3. Allelic frequencies of allele 7 of locus Aap. Values above the mean allele frequency are in 

black, values below the mean in white. The larger the circle, the more different is the value from the 

mean. Mean regional frequencies are provided for arbitrary regions corresponding to the circled areas.

140 EUF()RGEN: S()CIAL BR()ADLEAVES 

The highest growth rates are found at intermediate to high latitudes and seem to 

correspond to the areas with the highest heterozygosity. A detailed analysis is needed to 

understand this pattern, as it may be highly dependent on the location of the plantation site, 

contrary to the previous character (bud burst), which is much more stable. However, it is 

possible that the more pronounced height growth for the populations from the central part 

of the distribution range results from the combined effect of competition and silviculture. 

Under optimal conditions for growth, competition for light cmd mineral resources between 

trees is enhanced. Competition would in turn promote selection for higher growth potential. 

This trend may be reinforced by the long history of silviculture in the central part of the 

natural range. Silvicultural treatments of even-aged oak stands favour thinning practices 

that eliminate poorly growing trees. Natural selection and human intervention may have 

acted together to increase growth in these central populations. 

A longitudinal differentiation is apparent for allele 7 of locus Aap, present in higher 

frequencies in the west than in the east. This appears to be the general tendency for most 

studied alleles of all isoenzyme loci, as detailed in Zanetto and Kremer (1995). Le Corre et al. 

(1998a) by usin& geostatistical methods confirmed the preferential direction from southeast 

to northwest along which allele frequencies varied continuously. This pattern most likely 

reflects the genetic divergence among populations in different glacial refugia scattered from 

east to west, and the subsequent postglacial migration routes. The remaining oak forests 

were probably differentiated genetically between the glacial refugia since they were 

separated for more than 80 000 years. 

Causes of population differentiation 

Two hypotheses concerning the possible ongm of the present distribution of nuclear 

diversity are tested: are geographic trends of variations footprints of the origin of 

populations (historical hypothesis), or are they the result of geneflow among established 

populations, leading to isolation-by-distance (geographical hypothesis) (Le Corre et al. 

1998b). The two hypotheses are tested by comparing a set of pairwise distances between 

populations. Because cpDNA is only transmitted by seed (Dumolin et al. 1995) the spatial 

distribution of cpDNA polymorphisms indicates the past colonization routes. Therefore if 

the first hypothesis is favoured, a significant correlation is expected between nuclear genetic 

distances and cpDNA distances. Conversely if the isolation-by-distance hypothesis holds, 

the nuclear genetic distances should be correlated with the geographic distance among 

populations. 

Pairwise genetic differences for traits controlled by nuclear genes (isoenzymes and 

phenotypic traits) were compared with differences in respect of their origin (glacial refugia) 

or their present geographic location. Comparisons were made using a stepwise procedure. 

First three different distance matrices between populations were constructed. The first one 

(CD) corresponds to the genetic differences among populations (Nei's (1987) genetic 

distance for isoenzymes, or absolute difference between population mean values for 

phenotypic traits). The second distance, called the 'historical' distance (HD), is based on 

cpDNA information. It is calculated as the number of restriction fragment polymorphisms 

that distinguish two populations (Nei 1987; Le Corre et al. 1998b). The third distance is the 

geographic distance between two populations (CeoD), calculated as the Euclidean distance 

between the Mercator-projected coordinates. In a second step, the product moment 

coefficient of correlations between distances (CD and CeoD, CD and HD) was calculated 

between populations. And finally the correlation coefficients were compared with a Mantel 

test (Mantel 1967). The Mantel test consists of constructing a null hypothesis (Ho: the two 

distances are not correlated) through a Monte Carlo procedure. Cells of one distance matrix 

(for example CD) are permuted, whereas the second matrix (for example CeoD) is 

maintained as such. For each permutation a product moment correlation coefficient is 

calculated. The procedure is then repeated 1000 times, and the actual value of the

OVERVIEW RRESENTATIONS 141 

correlation is compared to the distribution corresponding to the null hypothesis. Because 

cpDNA polymorphisms follow a strong geographic distribution (Petit et al. 1993), partial 

correlation coefficients were calculated to remove the effects due to the correlation between 

the two explicative variables (GeoD and HD) as suggested by Smouse et al. (1986). 

Table 4. Mantel's tests. Second column: partial correlation between genetic distance (based on 

isoenzymes) and historical distance (geographical distance constant). Third column: partial 

correlation between genetic distance (based on isoenzymes) and geographical distance 

(historical distance constant) 

Traits R (GD,HD),GeoD R (GD,GeoD),HD 

Isoenzymes +0.02S NS +0.408** 

Bud burst +0.140* +0.288** 

Height growth +0.060 NS +0.088 NS 

* = Significant at P=O.OS; ** = significant at P=0.01; NS = nonsignificant at P=O.OS. 

Geographical distances between two populations are always more strongly correlated 

with the genetic distances, compared with historical distances, for all three traits 

investigated (it is highly significant for Nei's distance based on isoenzymes and for 

differences in bud burst scores). However, historical distances (based on the information 

contained in the maternal genomes of the trees) do seem to account for some of the variation 

in phenology, but not in isoenzymes. This is a somewhat surprising result that needs to be 

confirmed. 

Geneflow 

The data set for geneflow 

Mating system and geneflow were studied in a natural stand comprising 296 adult Q. petraea 

and Q. robur. The study stand is a square area of 5.76 ha (240x240 m) and is part of a 

continuous forest called La Petite Charnie in the northwest of France, near the city of Le 

Mans. Trees were assigned to either Q. petraea or Q. robur according to a multivariate 

analysis of leaf morphological traits. Outcrossing and hybridization rates were calculated 

with allozyme markers with the help of a "two species mixed mating model" (Bacilieri et al. 

1996). A second estimation of the same parameters in the same study site was made on a 

different pollination year by paternity analysis using microsatellites (Streiff et al. 1998b). 

Paternity analysis furthermore allowed reconstruction of all mating events in the stand, and 

as a result tracing of pollen dispersion. 

Interspecific geneflow 

As for other oaks, sessile oak is part of a complex of species exchanging genes (Kremer and 

Petit 1993). Interfertility between Q. petraea and Q. robur was shown by controlled 

hybridization experiments. Hybridization occurs also in natural populations as suggested 

by the extremely low species differentiation; reported interspecific G st 

values amount to 

about 3% (Table 2). All alleles, including rare alleles were shared by the two species. We 

estimated hybridization rates in a stand where the two species were mixed, with two 

different markers, isoenzymes and microsatellites. As shown in Table 5, there are gene 

exchanges between the two species in both directions; the preferential direction of 

pollination from petraea to robur that was recorded in one single year of observation (Bacilieri 

et al. 1996) has yet to be confirmed in other studies. Because species differentiation is 

extremely low throughout the natural distribution of the two species (Bodenes et al. 1997), 

hybridization is not likely to be restricted to specific geographic areas, but should be 

considered as a general mechanism. Furthermore, Q. petraea can also fertilize Q. pubescens or 

other species from central Europe (Q. dalechampii, Q. polycarpa and Q. virgiliana).


Table 5. Inter- (tb) and intra- (tw) specific outcrossing rates and selfing (s) rates in a mixed 

Q. petraea and Q. robur oak stand 

Molecular Q. robur Q. petraea 

markers S tw tb S tw ts Reference 

Isoenzymes 0.05 0.63 0.32 0 1.20 -0.20t. Bacilieri et al. 1996 

Microsatellites 0.03 0.94 0.03 0.01 0.89 0.11 Streiff et al. 1998b 

f Negative values of hybridization rates indicate that Q. petraea was most likely pollinated by extreme 

Q. petraea-like trees. 

The potential of Q. petraea to hybridize with other species should be seen as a mechanism 

to maintain and enrich its own diversity and those of its related species, and as a mechanism 

of Q. petraea to colonize new sites. The importance of hybridization as an evolutionary 

mechanism is witnessed by the complete sharing of cytoplasmic genomes among Q. petraea 

and Q. robur when they cohabit in the same forest. This indicates that the two species had 

continuous gene exchanges, since one of them colonized that forest; and that the second 

species, by continuously pollinating the recipient species through backcrosses, colonized the 

site as well (Petit et al. 1997). In other words, Q. petraea was 'regenerated' in each fon:!st by 

hybridiza tion. 

Intraspecific geneflow 

Paternity analysis on the experimental study site of 5.76 ha resulted in the assignment of a 

male parent for only 310 acorns out of a total of 984 analyzed (Streiff et al. 1998a). More than 

68% of the pollen that pollinated mature trees of the study stand originated from outside the 

stand! The construction of the pollen dispersion curve resulted in extremely flat tails, 

meaning that the pollen cloud is composed of pollen grains of widely spaced trees. 

Calculation of pollination distances in Q. petraea resulted in a mean value of 287 m with a 

standard deviation of 139 m (Streiff et al. 1998a). These figures should be considered as only 

rough estimates, since male parents from outside the stand were not identified, and 

pollination distances were inferred from mathematical models. However, reported studies 

in other oak species using the same technology also mentioned the long-distance pollen flow 

(Dow and Ashley 1996). Lahtinen et al. (1996), by monitoring pollen dispersal of Q. robur 

with traps on the edges of the natural distribution of the species, showed that pollen density 

was still present at 7 km from the source. Even at low rates, long-distance pollen flow is a 

strong homogenizing force, as shown by the extremely low population differentiation of 

Q. petraea for nuclear markers (about 3%, Table 2). Quercus petraea is widespread and its 

distribution is almost continuous except in alpine and Mediterranean regions, suggesting 

that genes could rapidly diffuse throughout the natural distribution. 

Conclusions 

Remarkable range-wide trends of variation exist in sessile oak for all traits investigated and 

for the corresponding measures of diversity. Actually, the trend for a trait and for its 

associated measure of diversity can differ, as in the case of bud burst, where the 

intrapopulation means vary with the latitude, whereas the corresponding coefficients of 

variation vary with the longitude. In the case of isoenzymes, most allele frequencies vary 

according to longitude, but depending on the measure of variation chosen (allelic richness or 

expected heterozygosity), the more variable region is either the east or the west. A striking 

result of this review is also the contrasting subdivision of diversity according to the traits; 

more than one-third of genetic diversity exists among populations for bud burst, whereas 

only 3% account for population differentiation for allozymes. Furthermore, depending on 

the goal and purpose of each investigator, at a given time, some traits may be more relevant 

for conservation than others. Consequently, given our results, this means that no region of 

the natural range should deserve more attention than others, as long as there is no serious


threat for the maintenance of the species. Because of the contradictory conclusions that may 

be drawn depending on the traits considered, the geographic structure of diversity or 

differentiation would not be recommended as relevant criteria for in situ conservation issues. 

However an alternative attitude would be to favour the maintenance of the mechanisms 

generating diversity, rather than the maintenance of diversity per se. As shown by our 

results, recent evolutionary factors as geneflow and diversifying selection in various areas of 

the natural distribution had a profound impact on today'S structure of gene diversity, 

erasing progressively the initial structure established at the end of last glaciation. As for 

practical issues for in situ conservation, management measures conserving inter- and 

intraspecific geneflow among oak stands coexisting in the form of a metapopulation would 

be an appropriate solution. Owing to the high levels of diversity that exist in Q. petraea 

stands, and the extensive geneflow through pollen dispersal, this species has a great 

evolutionary potential and is probably able to react extremely rapidly to ecological changes. 

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relation to budburst and growth cessation. Pp. 165-184 in Inter- and Intra-specific 

Variation in European Oaks: Evolutionary. Implications and Practical Consequences (A 

Kremer and H. Muhs, eds.). European Union, Brussels. 

Streiff, R, A Ducousso, Ch. Lexer, H. Steinkellner, J. Glossl and A Kremer. 1998a. Pollen 

dispersal inferred from paternity analysis in a mixed oak stand of Quercus robur L. and 

Quercus petraea (Matt) Liebl. Heredity (in press). 

Streiff, R, M. san Cristobal and A Kremer. 1998b. Paternity assignment, paternal identity 

and male mating success in a mixed stand of Quercus robur L. and Quercus petraea (Matyt.) 

Liebl. Genetics (in press). 

Weir, B.s. and c.c. Cockerham. 1984. Estimating F-statistics for the analysis of population 

structure. Evolution 38: 1358-1370. 


(Matt.) Liebl. I. Monolocus patterns of variation. Heredity 75:506-517. 

Zanetto, A, A Kremer, G. Miiller-Starck and H.H. Hattemer. 1996. Inheritance of isozymes 

in Pedunculate oak (Quercus robur L.). J. Heredity 87:364-370. 

Zanetto, A, G. Roussel and A Kremer. 1994. Geographic variation of inter-specific 

differentiation between Q. petraea (Matt.) Liebl. For. Genet. 1(2):111-123.


Oak decline in European forests 

Tomasz Oszako 

Forest Research Institute (IBL),Warsaw, Poland 

c/o European Forest Institute, Joensuu, Finland 

The spread of oak decline, a concern in many parts of Europe 

Oak forests are important for the forest economies of many European countries. Because of 

the importance of oak, the increasing number of cases of oak decline in many European 

countries is causing much concern. Results of the 1995 survey on Forest Condition in 

Europe showed abundant occurrence of severely damaged oak trees in Central Europe (EC- 

UN/ECE 1996). The most severe deteriorations were observed in Germany, the Czech 

Republic, Poland and the Slovak Republic. Are the oak forests going to follow the 

coniferous forests in suffering from air pollution or from the new vascular mycosis disease 

which is spreading over the European continent Oak decline is widespread in many 

districts of Italy (e.g. Lazio, Marche, Toscana, Puglia, Basilicata and Calabria) and has been 

spreading during the last 10 years. A survey carried out in 1988 on 1000-m 2 plots showed 

that around 85% of Q. cerris trees were declining or already dead in Puglia, 76% in Lazio and 

51 % in Basilicata. Even more alarming were the figures for Q. pubescens in Puglia, where 

approximately 95% of trees were declining or dead (OEPP/EPPO 1990). 

In the Netherlands, according to the national forest health survey which has taken place 

annually since 1984, the health of oaks is deteriorating. In 1988 only 21% of trees were 

estimated as healthy. Mortality in individual stands varied from 20 to 50% (OEPP/EPPO 1990). 

In Poland in 1984-85 oak decline spread very quickly all over the country, affecting 

145000 ha (2% of the total forest area) (Oszako 1994). The progression of the disease was so 

swift that for example in one forest district (Krotoszyn) during the period 1982-89, foresters 

had to remove 30000 m 3 of timber from the forest (40% in 1984 and 40% in 1985). The 

amount of salvage felling expressed in m 3 was 690 times higher in 1984 (20 194 m 3 ) than in 

1980 (29 m 3 ). Such a large supply of wood interfered not only with the harvesting plan of a 

particular forest division but also with the local wood market. 

Even in countries which do not report oak decline as a major problem, a number of types 

of dieback can be recognized. In one of the few studies undertaken in Great Britain, about 

40% of the 60-year-old trees in an affected area of about 8 ha had died. Additionally, figures 

obtained during a sample survey of hedgerow oak in 1990 indicated that some 18% of oaks 

were suffering from dieback affecting more than 10% of the crown (OEPP/EPPO 1990). 

There is a long list of descriptive names given to instances of sudden oak decline: cohort 

senescence, T' disease, bark canker, epidemic wilt, Eichensterben, new-type damage, 

damage of various types, oak mortality, vascular disease, spiral disease, lethal yellowing and 

oak wilting are the most common (Ragazzi et al. 1995). 

Why is it important to study oak decline 

Although it can be asked "Why engage in research on oak decline while major and 

important tree species are endangered by general forest decline", the following arguments 

support the necessity for engaging in further oak research: 

• Oaks are ecologically and economically indispensable. Many forest sites on 

pseudogleyic soils would be easily destabilized without oaks. Oaks are essential for a 

wide range of forest products and are a valuable asset for forest enterprises. 

• The health of oaks in Europe has steadily declined in the last decade in contrast to the 

recovery of other tree species (e.g. fir, spruce). Oak may be a tree of minor importance 

on a national scale, but on a local or provincial scale it is quite important.


e 

e 

Oak and mixed oak hardwood stands improve the effect of forests on climate and 

hydrology in areas where the 'agricultural steppe' is dominant. Forests in these areas 

provide valuable habitats for many plant and animal species. A change in tree species 

would endanger many of them. 

With global warming the potential range of oak silviculture will be expanded. At the 

same time, the steadily deteriorating conditions for coniferous species make oak 

forests more attractive or even indispensable in some European regions. 

Oak decline is not a new phenomenon in Europe but when it occurs it causes important 

economic perturbations. Therefore it is important to understand this decline syndrome - to 

predict its appearance and mitigate its effect on the forest economy. 

A review of the literature on oak decline in Europe shows a chronological and 

geographical progression (Table 1). The first report originated more than 250 years ago and 

other cases have been reported regularly since the 18th century. However, in the last 15 

years the incidence of oak decline has shown a dramatic increase all over Europe, and even 

by 1989 was reported in every European country. Such a situation has prompted many 

investigations in both Europe and the United States. Up to now, no single universal causal 

factor (abiotic or biotic) has been found which could explain the widespread abnormal death 

of oaks. In the beginning of the 1980s some researchers from Romania, Poland and the 

former Czechoslovakia, after preliminary studies, reported the fungus Cera tocys tis fagacearum 

as a cause of oak disease. Actually, the symptoms were very similar to these observed in 

North America. We now know that this was a mistake and that this serious pathogenic 

fungus is not yet present in Europe. However, the possibility of its appearance is a real 

threat because of the steadily growing wood timber trade. For this reason quarantine 

regulations do not allow the importation of oak wood timber with bark or without chemical 

or drying treatments. As a consequence of the panic over a possible outbreak within the 

European oak forests of the well-known and dangerous USA vessel disease called 'oak 

wilting', many trees showing first decline symptoms were immediately removed from the 

stands. During necessary sanitary harvesting operations many live trees in the stands were 

injured. These wounds of plant tissues were open gates for fungi infections, mostly Phellinus 

robustus, which colonized internal parts of the trees, causing internal rot of stems. This 

wood is now only used for firewood. In some stands, up to 40% of trees are infected by this 

fungus and have lost their primary high quality value. The removal of many trees changed 

the light condition in the oak ecosystem. More light reached the litter level in the oak stands 

and enhanced weed grass development, causing quick site degradation and silvicultural 

problems (e.g. with regrowth of natural regeneration). Light stim\llated the development of 

water sprouts which decreased stem wood quality. Oak decline has caused not only direct 

financial losses because declining trees had to be removed before their maturation or 

harvesting age, but also indirectly affected wood quality and growing conditions in the 

ecosystem. The total economic cost should also include the negative effect on the site (soil). 

According to the literature, oak decline is a result of a general weakening of the trees and 

is manifested by a dieback of the crown. Trees either die within a few years or suffer for 

many years. Some of them do recover, producing re sprouts along the trunk and branches. 

Usually, oak decline is recorded when numerous trees die or when symptoms are exhibited 

over quite a large area. The symptoms can vary among sites according to environmental 

(ecological and silvicultural) differences and the oak species affected. Symptoms common to 

all oak species throughout the European range are crown thinning, growth of epicormic 

shoots, and bleeding. 

Some researchers view oak decline as a pernicious new event that will lead to the 

disappearance of oak forests. Others regard it as a chronic occurrence common to all plants 

and forest ecosystems. There are at least two hypotheses to explain the nature of this 

complex disease (Ragazzi et al. 1995):


• The first emphasizes the synergistic action of different factors which previously operated 

separately, and which have caused an increase in oak decline during the last decade. 

• The second suggests that it is a quite new phenomenon which first appeared about 15 

years ago. Earlier cases were sporadic and localized events which have no real 

bearing on current oak decline. 

Researchers are agreed that numerous biotic and biotic causes are involved in the 

development of the disease. Their importance depends on varying local circumstances and 

their interrelations. Research carried out in many European laboratories detected many 

different organisms, e.g. fungi, nematodes, bacteria, MLOs and viruses: 

• A high number of fungi isolated from declining oak tissues have been reported by 

numerous scientists. They mainly belong to the Deuteromycotina and Basidiomycotina 

classes, the most frequently recorded being Armillaria spp., Fusicoccum quercus, 

Hypoxylon mediteraneum, Phomopsis quercella and Phytophtora cinnamoni. Most of those 

species are already known to be pathogenic and to occur on declining oaks. 

• Sapwood nematodes (Bursaphelenchus spp.) were found to be common in declining and 

dying oaks. A correlation was found between the quantity of nematodes and vitality 

(expressed by conditiometer measurements). 

• Only a few bacteria have been found in Romania in association with oak decline: Erwinia 

quercicola Georges et Bad. and Erwinia valachica Georges et Bad. (Petrescu 1974). 

• Mycoplasma Like Organisms (MLOs) have been detected on declining oaks in 

Romania (Ploaie et al. 1987). 

• The only virus identified so far in declining oaks is the tobacco mosaic virus (TMV) on 

Quercus robur, Q. petraea and Q. cerris in Germany (Nienhaus 1975) and Hungary 

(Horvat et al. 1975). 

There are two common models of oak decline accepted by most researchers: the classical 

chain disease model of Keller (Dominik 1971) in which sets of factors act in succession, and 

the spiral model developed by Manion (1981) which consists of three main groups of factors 

acting simultaneously in the same space and time. Initially, factors acting over the long term 

(e.g. climate, site conditions, air pollution, genotype) predispose trees to the inciting factors 

which cause direct damage to the trees (e.g. insect defoliation, frost damage, etc.). It is a 

combination of these factors which ultimately causes the death of the trees. 

References 

Dominik, J. 1971. Ochrona lasu PWN, Warszawa. 

Horvat, H.J., 1. Eke, T. Gal and M. Dezesery. 1975. Demonstration of virus-like particles in 

sweet chestnut and oak with leaf deformations in Hungary. Z. Pflanzenkr. 

Pflanzenschutz. 82:498-502. 

Manion, P.D. 1991. Tree Disease Concepts. 2nd edn. Prentice Hall, Englewood Cliffs, New 

Jersey. 

Nienhaus, F. 1975. Viren und virusverdachtige Erkrankungen in Eichen (Quercus robur und 

Quercus sessiflora). Z. Planzenkr. Pflanzenschutz. 82:739-749. 

OEPP IEPPO. 1990. OEPP IEPPO Bulletin 20:405-422. 

Oszako, T. 1994. Choroby drzewostanow lisciastych, przyczyny, przebieg I ograniczanie 

szkod, IBL, Warsaw. 

Petrescu, M. 1974. Le deperissement du chene en Roumanie. Eur. J. For. Pathol. 4:222-227. 

Ploaie, P.c., M. Ionica and A. Alexe. 1987. Oak decline: a disease caused by mycoplasma-like 

microorganism Buletinul Proctecia Plantelor 1:13-21. . 

Ragazzi, A., S. Vagniluca and S. Moricca. 1995. European expansion of oak decline: involved 

microorganisms and methodological approaches. Phytopathol. Medit. 34:207-226.

Table 1. Causal factors involved in incidences of oak decline in some European countries 

Defoliation by 

Year Oak species Drought insects Secondary pests 

Fungi 

Silviculture 

Other 

Author 

Austria 

1982- 

83 

only in some cases 

in 1986 

Scolytus intricatus, Jassidae, 

Hemiptera, Cynipidae, 

Buprestidae 

Armillaria spp. 

Ohiostoma piceae, 

0. cf. valachicum, 

0. prolifera, 0. cf. 

roboris 

Nematodes 

Bursaphelen-chus 

spp. 

0fTl:-i 

-lOO 

o 0 CD 

3 :::J () 

o. ~_:::J" 

N 0" 

CD III 

AC 

~ 

Slovakia 

1979 Q. dalechampii, 

Q. polycarpa, 

Q. petraea x . 

robur, Q. petraea, 

Q. robuT, 

Q. slavonica, 

Q. virgiJiana, 

Q. cerris, 

Q. pedunculifora 

Q. frainetto, 

Q. pubescens 

Denmark 

1910 Q. robur, 

Q. petraea 

predisposing factor 

is soil moisture 

dropping below 

wilting point in 

summers 1976-77 

Operophtera brumata, 

Tortrix viridana 

Scolytus intricatus as a vector 

of fungal diseases 

0. quercus, 

0. valachicum, 

0. roboris, 

0. kubanicum, 

0. piceae, Diaporthe 

fasciculata 

Microsphaera 

alphitoides, 

Armillaria meJlea s./., 

Gloeosporium 

quercinum, Stereum 

rugosum, Nectria 

dittisima, Colpoma 

qercinum, 

Myxosporium 

lanceola, Pezicula 

cinnamomea 

summer fellings, 

increase of 

volume and size 

of mechanical 

damage and 

attractiveness 

for beetles 

high temperatures, 

temperature 

increase since 

1970 about 1- 

1.5°C, 

atmospheric 

pollution, fluorine, 

aluminium 

damage to bark by 

woodpeckers 

:0 

r 

CD 

o 

:::J 

er 

< 

o· 

;t> 

-< 

Q. 

CD 

» 

:::J 

Q. 

CD 

Cri 

CD 

:::J

Year 

Oak species 

Drought 


insects 

Secondary pests 

Fungi 

Silviculture 

Other 

Author 

France 

1921 Q. robur 

1942 

1976 

1986 

does not always 

lead to decline, limits 

radial growth 

Coflybia fusipes, 

Armi/laria meflea, 

Microsphaera 

alphitoides 

o 

o 

CD 

~ 

~ 

Germany 

1 989 Q. robur 

summer 1983 

Lepidoptera 

Cytospora sp., 

Pezicula 

cinnamomea, 

Armilfaria sp., 

0. piceae, O. roboris 

reduced groundwater 

level, winter 

extreme 

temperature 

1984/85 

;t> 

~ 

c 

:::;; 

Hungary 

1978 Q.peifaea 

Italy 

1988 Q. cerris, 

Q. pubescens 

outbreaks of leaf- 

eating insects 

last decade, soil 

water content below 

critical level in July 

last 10 years small 

amount of 

precipitation in 

August 

Scolytidae, Cerambycidae, 

Buprestidae, etc. Scolytus 

intricatus as vector of fungi 

Armiflaria melea 

(sensu la to), 

Ophiostoma spp., 

Botryosphaeria 

stevensii (an. 

Diplodia mutila), 

Cytospora spp., (C. 

leucosperma), 

Phomopsis sp., 

0. piceae, 

0. pilifera, 

0. moniliformis 

0. coerulescens, 

0. piceae, Diplodia 

mutila, Graphium 

penici/lodes, 

Hypoxylon 

mediterraneum, 

Phoma cava, 

Phomopsis quercina, 

pyrenochaeta 

quercina, 

Acremonium spp., 

Sporotrix spp., 

Armi/laria spp. 

inadequate 

forest 

management 

practices, lack of 

thinning and 

sanitary felling, 

too large game 

population 

atmospheric 

pollution 

lack of snow, 

shallow root 

system of cop-pice 

stands and 

increased depth of 

underground 

water, air pollution, 

acid precipitation, 

acidification of soil, 

aluminium toxicity 

c.... 

~ 

_:::J" 

,- 

< 

.l! 

:::J 

Pl 

;t> 

< 

Pl 

:::J 

:::J 

_2: 

z 

r 

c 

~:


Year Oak species Drought insects Secondary pests Fungi Silviculture Other Author 

Netherlands 

1984 Q. robur, 1982, wet spring and caterpillars of Tortrix Pezicula Winter 1984/85 

:t> 

1986 Q. petraea dry summer 1983, viridana and cinnamomea, mild until late 

most serious on wet Operophtera brumata Armilfaria spp., December and 

0 

0 

sites with a 1983 Microsphaera sudden severe 

rJ) 

cri 

groundwater level alphitoides, frost C- 

O> 

which fluctuates Leptographium spp., 0> 

:::J 

0. piceae, 

Acremonium s2. 

Poland 

1982 Q. robur 1982-84 Tortrix viridana, Saperda scalaris, Scolytus Armi/laria spp., Unsuitable soil -l 

1984 Operophtera brumata, intricatus, Plagionotus Microsphaera conditions 

A 

0. fagata, Hiberia arcuatus, Ragium mordax, alphitoides, (especially too 0 

:;: 

defoliaria, Lymantria Leiopus nebulosus, Agrilus Phelfinus robustus, much or too little 0> 

iii 

dispar, L. monacha, sulcico/lis, A. angustatus, Colpoma quercinum, water) 

p. 

Malacosoma neustria, A. biguttatus, Phymatodes Coryneum 

' 

Biston histortata, testaceus, Plagionotus umbonatum, "U 

Tischeria dodonea, detritus, Xylotrechus anti/ope, Dothiorelfa advena, N 

'< 

Phalera bucephala, Dryocoetes villas us, Xyloterus Cytospora 

0- 

~ 

Euproctis domesticus x saxeseni, intermedia, 

chrysorrhoea, Aftica A. monograph us, Xipphydria Ophiostoma spp., :-l 

quercetorum longicoflis, Hylecoetus Ceratocystis 0 

rJ) 

dermestoides 

fimbria ta, 

Fusiccocum 

quercus, Pezicula :- 

cinnamomea, 

(J) 

Phomopsis 

Pi 

N 

quercefla, 

'< 

Amphiporte 

'" 

leiphaemia, 

COlJ:.neums2· 

United Kingdom 

1989 0. robur caterpillars of Tortrix scale insect Kermes quercus Armilfaria spp. I ntensification of 

:z 

viridana in several Sphaeropsis spp. farming (hedgerow 

(j) 

consecutive years oak) cr 

0- 

rJ) 

N 

0> 

~ '" 

Yugoslavia 

1985 Q. pubescens, no correlation with Coraebus bifasciatus Armilfaria spp. air pollution, 

0 

1987 Q. robur, soil moisture and strong winds in c- 

o. petraea, state of health 1985, shallow c 

0 

Q. cerris, 0. rubra soils, interrupted 

forest structure


Year Oak species Drought insects Secondary pests Fungi Silviculture Other Author 

Russia 

Central Part 

1941- Q. robur dry summer and low Microsphaera Hard winter of 

43 snow wi nter of 1938- alphitoides 1939-40 and 

1939 1941-42 

1971- Q. robur drought of 1972 Microsphaera 

73 alphitoides 

1991- Q. robur Periodical defoliation Secondary defoliation by PheIJinus robustus, improper forest Periodic change of 

z 

94 by Limantria dispar L., Ocneria dispar L. and others Polyporus management, wet and dry 

:< 

Operphthera brumata sulphureus, destruction of periods (climatic z 

L., etc Daedalea quercina, complex oak factor - ecological Pl 

-0 

ArmiIJaria spp., stand structure, stress) Pl 

;;;: 

Ceratocystis spp. pasture, 0 

< 

damages by elk 

Middle Povolzhje region 

1941 Q. robur Hard winter frost 

1968- Q. robur Little snow and 

69 hard winter frost 

1978- Q. robur Microsphaera Hard winter frost 

79 alphitoides, 

PhelJinus robustus, 

Polyporus 

sulphureus, 

Daedalea quercina 

1991- Q. robur Tortrix viridana, and Secondary defoliation by Microsphaera inadequate 

94 crataegana, Ocneria dispar L. and others alphitoides, forest 

(Q 

Q 

Operophtera brumata PheJ/inus robustus, management ~ 

and others Polyporus practices, -< 

Pl 

sulphureus, 

destruction of 

Daedalea quercina complex forest 

'" 0 

< 

stand structure, 

iD 

< 

decreasing 

stand density, 

pasture

152 EUFORGEN: SOGIAI.;. BROADI.;.EAVES 

Genetic diversity of beech populations in Europe 

Ladislav Paule and Dusan Gomory 

Faculty of Forestry, Technical University, Zvolen, Slovakia 

Introduction 

European beech (Fagus sylvatica L.) is considered at present the most widespread economically 

important broadleaved tree species in Europe. The extent of beech forests (Fagus sylvatica and 

Fagus orientalis) in Europe and Asia Minor is estimated to be between 17 and 20 million ha (e.g. 

Milescu et al. 1967 estimate 16.8 million ha) and represents approximately 10% of European 

forests. The proportion of beech forests in individual regions represents frequently up to 30% 

of the total forest area (e.g. Croatia, Slovakia, Romania, etc.). 

Natural range and systematics 

Both F. sylvatica and F. orientalis belong to the forest tree species with the widest natural 

distribution range in the western part of Eurasia. Fagus sylvatica is distributed in western, 

central and southern Europe with individual occurrences in southern England and southern 

Scandinavia. Fagus oriental is is distributed in Asia Minor, in the Caucasus, in the Amanus 

mountains (Syria), and in the Elburz mountains (Iran). A contact zone between the natural 

ranges of both species is located in northern Greece and Bulgaria. Isolated occurrences of 

F. oriental is outside the natural range were recorded in eastern Serbia (GliSi 1973), in 

Macedonia (Cernavski ex Milescu et al. 1967), in Banate and Moldova (Milescu et al. 1967), 

and in Dobrudja and Central Bulgaria (Czeczott 1932). 

Problematic taxonomic identity of beech exists in the Crimea. Poplavskaja (1927) 

described beech in the Crimean peninsula as an independent species - F. taurica. Beech 

occurs in the Crimea in two altitudinally separated zones. The lower zone was more 

frequently described as F. oriental is and the upper one as F. taurica (Molotkov 1966; Milescu 

et al. 1967), but Wulff (1932) described it as F. sylvatica. It is necessary to point out that the 

name F. taurica is used in different ways. While Poplavskaja described it as the intermediate 

form between F. sylvatica and F. orientalis, Milescu et al. (1967) considered it another species 

with its occurrence not limited to the Crimea. 

A further dubious taxonomic unit is F. moesiaca with the most frequent occurrence in the 

Balkans. Misic (1957) considered it a separate species of Tertiary origin. It is most 

frequently considered a subspecies of F. sylvatica. 

It is obvious that taxonomic status of beech in a rather large zone is unclear. The original 

description of F. moesiaca and F. taurica was based mainly on the morphological traits of 

leaves. Seldom do these presumed species occur in comparative provenance trials together 

with F. sylvatica. 

The aim of recent investigations carried out in Slovakia, France and other countries was to 

describe both species - F. sylvatica and F. orientalis - using genetic markers, to define the 

zone of introgressive hybridization and the limits or the direction of geneflow between 

them, and to characterize the structure of diversity within the genus Fagus in Europe and 

Western Asia. 

Fagus sylvatica is a tree species of oceanic and suboceanic climate. Its eastern distribution 

limit is parallel to the limit of continentality (Stanescu 1979) and is defined by the air 

humidity and late frost (Pukacki 1990). Although it is resistant to fairly low temperatures, 

which does not exclude it from the higher altitudes, it is sensitive to late spring frost - a 

limiting factor at lower altitudes with a great accumulation of cold air (Becker 1981). For this 

reason beech does not occur in frost valleys or in the regions with a more pronounced 

continental climate.

BXlERXlIEW BRESENrIlArIllBNS 153 

Phylogeny 

The oldest fossils that can be attributed to the genus Fagus originate from the late 

Mesozoicum (Milescu et al. 1967). The remnants of different forms of beech are much more 

abundant from the Tertiary. They were even found in regions where beech does not occur at 

present, e.g. Greenland, Iceland and Svalbard (WaIter ex Milescu et al. 1967). Among the 

Tertiary fossils, at least four different species were described: F. attenuata, F. pliocenica, 

F. sylvatica and F. feroniae. Ettingshausen (ex Milescu et al. 1967) considers all these species 

the different forms of one collective species F. feroniae. The forms similar to F. ferruginea 

(grandifolia) were also found in Europe. From both beech species found in Europe and 

western Asia, F. orientalis is generally considered a more ancient, ancestral form, of which 

F. sylvatica has evolved. 

During the Pleistocene, beech underwent several retreats within the glacial periods and 

expansions during the warm interglacials. Unfortunately, it is not possible to reconstruct the 

early history completely on the basis of fossil remnants. A more or less reliable description, 

based on pollen diagrams, is possible only for the last Wiirm/Weichsel glacial period and 

the recolonization of Europe in the Holocene (Huntley and Birks 1983). The refugia of beech 

were probably situated in central Pyrenees, southern France, Sicily, Apennines and Balkan 

Peninsula. The expansion started ca. 12 500 years ago. Further development of pollen 

values indicates that the main source of expansion was the refugium in the southern Balkans 

(Macedonia, western Bulgaria, southeastern Serbia). Beech from this area recolonized the 

whole present range, intermixing with the populations from local, smaller refugia. There 

seems to be one exception - beech in the Apennines and in Sicily, which preserved its 

continuity and was not affected by immigration from outside. 

This view, although generally accepted, seems to contradict in some aspects the 

differentiation patterns identified utilizing isoenzyme genetic markers. 

Genetic inventories 

History 

The application of genetic markers to investigate genetic diversity of beech populations 

started later than in conifers. Kim (1980) identified the first enzyme gene locus by studying 

zymograms of parent trees and their offspring. He used in his investigations only two loci- 

Acp and Lap. Paule (1992) and Hattemer et al. (1993) published reviews of the isoenzyme 

systems, their biochemical analyses and controlling gene loci which were applied in genetic 

inventories of beech. They have listed a total of 17 isoenzyme systems controlling 27 gene 

loci (Table 1). 

Miiller-Starck and Starke (1993), Thiebaut et al. (1989) and Merzeau et al. (1989) studied 

inheritance patterns of the enzyme gene loci in progenies from controlled crossings and 

single trees. 

SDS-PAGE was applied to find species-specific differences in the seed protein content of 

F. sylvatica and F. orientalis populations from Bulgaria. The comparative analysis of their 

patterns showed a greater similarity than dissimilarity between both species (Busov 1993). 

Finally, the polymorphism in the chloroplast genome has been detected by resolutive 

restriction site studies of PeR-amplified fragments. Eleven haplotypes, which could be 

phylogenetically ordered, were detected in a large survey (399 individuals in 85 populations) 

encompassing most of the natural range of the species (Demesure et al. 1996). 

Institutions involved 

Beech is now considered to be a tree species with the most extensive genetic inventory of the 

broadleaves. The first investigations used a quite small number of isoenzyme gene loci (3-6) 

(Comps et al. 1987, 1990). In later studies the number of polymorphic isoenzyme loci 

increased to 10-16 (Miiller-Starck and Ziehe 1991; Gomory et al. 1992; Hattemer et al. 1993; 

Turok 1993, 1996; Leonardi and Menozzi 1995; Trober 1995; Larsen 1996).


Table 1. 

Euro~ean beech 

Review of the isoenzyme systems used for population genetic investigations in 

Authors who used the isoenzyme systems and loci 

Locus 1 2 3 4 5 6 7 8 9 10 11 12 13 14 

6Pgd-1 x x x x x x x x x x 

6Pgd-2 x x x x x x x 

6Pgd-3 x x x x x 

Aap-1 

x 

Aap-2 

x 

Aco-1 x x x x x 

Aco-2 x x x x x x 

Acp-1 x x 

Acp-2 

Adh-1 x x 

Adh-2 x x 

Adh-3 

x 

Amy-1 

x 

Dia-1 x x x x x 

Fum-1 

x 

Gdh-1 x x x 

Got-1 x x x x x x 

Got-2 x x x x x x x 

Idh-1 x x x x x x x x x x 

Lap-1 x x x x x 

Lap-2 

Mdh-1 x x x x 

Mdh-2 x x x x x x x x x x x x 

Mdh-3 x x x x x x x x x x x x 

Mnr-1 x x x x x x 

Ndh-1 

x 

Pgi-1 x x 

Pgi-2 x x x x x x x x x x x x 

Pgm-1 x x x x x x x x x x 

Px-1 x x x 

PX-2 x x x x x x 

Skdh-1 x x x x x x x x x 

Skdh-2 

x 

Sod-1 x 

1 Comps et al. 1991 9 MOlier-Starck and Ziehe 1991 

2 Gomory et al. 1998 10 Trober 1995 

3 Konnert 1995 11 Turok 1996 

4 Larsen 1995 12 Kitamura et al. 1992 (F. crenata and 

5 Leonardi and Menozzi 1995 F. japonica) 

6 Lochelt and Franke 1995 13 Houston and Houston 1994 (F. grandifolia) 

7 MOlier-Starck and Starke 1993 14 Premoli 1996 (Nothofagus) 

8 MOlier-Starck 1996


Genetic inventories of beech populations have been carried out at several institutions in 

Europe. To these belong mainly the French group from Bordeaux which focused attention 

on the western European populations and compared them with some central European and 

Balkan ones (Comps and Thiebaut and other coworkers). The second group is from Zvolen, 

Slovakia, which focused on the eastern European populations and the transition zone of 

F. sylvatica and F.orientalis in the Balkans, Asia Minor and Caucasus. Numerous other 

groups analyzed regional sets of populations in Italy (Leonardi, Belletti, etc.), Germany - 

Bavaria (Konnert), Saxony (Trober), North Rhine-Westphalia and Rhineland-Palatinate 

(Turok, Ziehe, etc.), Denmark (Larsen), or combined their studies with the air pollution 

effect (Muller-Starck in Germany, Brus in Slovenia, Vysny et al. in Bohemia), silvicultural 

practices (Starke et al.). A rough estimate of the number of populations analyzed within the 

European genetic inventories would be around 800. 

Genetic diversity 

The levels of genetic diversity and multiplicity are strongly affected by the choice of marker 

loci. As mentioned above, the number of loci, used in different studies on beech, is quite 

restricted, in general not exceeding 16. Loci exhibiting at least a minor polymorphism are 

used. These loci, which are generally considered monomorphic, are not utilized, they are 

not even recorded although they appear on the gels stained (an example is the fastest 

migrating monomorphic zone, appearing on gels stained for malate dehydrogenase in 

F. sylvatica. It was proven that it is controlled by one locus, which is polymorphic in 

F. orientalis). Therefore, caution is necessary when comparing results from different studies. 

An incomplete overview of the published levels of genetic multiplicity, diversity and 

differentiation measures is given in Table 2. 

Despite this, there are no substantial differences when comparing genetic multiplicity in 

different parts of the distribution range. The mean number of allel~s per locus is in general 

about 2.5. Although there is no clear trend, it seems that mean number of alleles tends to 

increase in the eastern and southern regions. Low estimates for the Italiah Piedmont and/ or 

for Denmark could be influenced by a lower number of loci used (8 and 7, respectively) than 

in the other studies. 

Among the genetic diversity measures, the values of the expected heterozygosity are 

frequently available. However, no geographic trends can be identified, probably because of 

the choice of different sets of marker loci. Expected heterozygosity values (which measure 

gene diversity) range between 0.2 and 0.4, effective number of alleles between 1.25 and 1.6. 

Contrasting estimates of diversity for Denmark (0.177, Larsen 1996) and Sweden (0.290, 

Comps et al. 1993) underline the presumed effect of the choice of loci. 

Genetic differentiation 

As indicated by Gst-values presented in Table 2, isoenzyme markers do not allow discovery 

of much differentiation. The major part of the genetic variation (mostly over 95% of the total 

variation) is observed within populations. Despite this, several studies succeeded in 

identifying geographic trends of single allelic frequencies, or geographic patterns of the 

genetic variation based on multilocus approach. 

One of the first large-scale studies on beech, focusing on the Atlantic region of western 

Europe (Comps et al. 1987), proved the existence of a latitudinal trend for Px-l/105 

(peroxidase) allele (frequency decreasing toward the north). For other loci, certain trends 

were also observed: the frequency of the Px-2/13, Got-l/105 and Pgi-1/87 alleles is highest in 

the Pyrenees and decreases toward the north as well as the south. The frequency of the Got- 

1/105 is significantly correlated with altitude. Similar trends were reported by Felber and 

Thiebaut (1984), Cuguen et al. (1985) and Barriere et al. (1985). They found the heterozygosity 

as well as frequencies of alleles Px-l/l05, Px-2/13 to be highest in extreme climatic conditions 

within Europe.

156 EUFORGEN: SOCIAl.. BROADl..EAXlES 

Table 2. Selected values of measures of the genetic multiplicity, diversity and differentiation of 

beech forests in Euroee 

Country/Region N. H. ne G st 

Reference 

Spain 0.301 0.046 Comps et al. 1993 

France 0.306 0.040 Comps et al. 1993 

Germany 2.3-2.9 0.188-0.229 1.32-1.44 0.045 t MOller-Starck and Ziehe 

1991 

Bavaria 2.1-2.9 0.247 1 .299-1 .404 0.019 Konnert 1995 

North Rhine- 2.1-2.8 0.23-0.38 1.30-1.59 Turok 1996 

Westphalia 

Baden-WOrttemberg 2.2-2.6 0.226-0.289 Lochelt and Franke 1995 

Czecho-Slovakia 2.1-2.6 0.281-0.346 Gomory et al. 1992 

Italy - Piedmont 2.0-2.3 0.177-0.278 0.043 Belletti and Lanteri 1996 

Italy 2.0-2.9 0.046 t Leonardi and Menozzi 

1995 

Balkans 2.4-3.2 0.230-0.260 1.295-1.346 0.014-0.040t Gomory et al. (1998) 

0.228-0.282 Hazier et al. 1997 

Croatia 2.2-2.5 0.297-0.315 0.053 Comps et al. 1991 

Ukraine 2.4-2.6 0.186-0.247 Vysny et al. 1995 

Denmark 2.0-2.4 0.164-0.187 0.006 t Larsen 1996 

Sweden 0.290 0.039 Comps et al. 1993 

t - 

F st 

Na - mean number of alleles per locus, He - expected heterozygosity, ne - effective number of alleles, 

G st 

- interpopulation component of the genetic variation. 

Leonardi and Menozzi (1995) found a very dear differentiation between Italian stands 

from the southern and northern parts of the country. They attributed these differences to the 

glacial and postglacial history. 

In a recent study, Comps et al. (1998) investigated 78 populations on a west-east transect 

from the Alpine Chain to the Hungarian Basin. A principal component analysis of 

multilocus allelic frequencies allowed assignment of the populations to four groups: French 

Alps, Swiss Alps, Northeastern Alps (Germany, western Austria), and eastern Austria 

together with Hungarian Basin. 

In Ukrainian Carpathians and the adjacent lowlands, Vysny et al. (1995) found differences 

between populations from the southwestern slope, northeastern slope and the lowlands, but 

the grouping was not unequivocal. However, principal coordinate analysis of the genetic 

distance matrix proved the existence of a dear altitudinal dine. 

Comps et al. (1991) analyzed 35 populations from Croatia. Discriminant analysis based 

on allelic frequencies allowed distinction of two strongly differentiated groups of 

populations originating from higher altitudes (mountain regions) and lowlands (maritime 

regions). At the same time, the populations belonging to the phytosociological association 

Selerio-Fagetum exhibited significant differences, compared with the other associations. 

The populations from the transition zone between F. sylvatica and F. orientalis represent a 

specific problem and were subjected to several studies. Eastern beech itself seems to be 

much more differentiated than European beech, but there are features of allelic profiles 

which are common for all regions with the occurrence of F. orientalis: the loci Got-2 and Mdh- 

3 are almost or completely monomorphic, while in F. sylvatica the representation of the most 

frequent allele does not exceed 0.8. The opposite proportion is at Mdh-l and other loci 

where a higher degree of polymorphism is shown in the populations of F. orientalis (Paule et 

al., unpublished).


Balkan beech (designated by local botanists as Fagus moesiaca Czeczott) differs from the 

typical F. sylvatica by morphological traits of leaves, flowers and fruits, a high sprouting 

capacity and a considerably higher frequency of seed years, as well as ecological 

requirements (MiSic 1957). HazIer et al. (1997) compared beech populations from the 

northwestern Balkans (Croatia), generally considered pure European beech, with 

populations from Macedonia and Bulgaria. The southeastern populations differed by the 

occurrence of specific rare alleles (Mnr-l/131, Pgi-l/76, Pgi-l/113, Pgm-l/93, Pgm-l/l09) and 

could be clearly distinguished from the northwestern ones by means of discriminant 

analysis of multilocus allelic profiles. 

The gap of information from the central part of the former Yugoslavia was fulfilled in the 

study of Gomory et al. (1998), where populations from Serbia and Bosnia-Herzegovina were 

included. Although they found a clinal character of the overall genetic variation as 

expressed by multilocus genetic distances, the stands classified as Fagus moesiaca could be 

distinguished from the remaining ones. They exhibited a similarity of allelic frequencies and 

of the presence of rare alleles not only with the neighbouring F. orientalis beech populations 

from eastern Balkans and Asia Minor, but, strikingly, also with beech in Calabria. 

Overviews of the investigations performed in different regions and covering the whole 

range in detail have not been published yet. Preliminary results for the eastern part of range 

were published by Paule et al. (1995) and Gomory et al. (1995). The results indicate that 

differentiation is higher within F. oriental is populations than within F. sylvatica, although the 

populations of F.orientalis originate from a much smaller territory. The F. sylvatica 

populations originating sampled the Carpathians exhibit a rather compact group, thus 

having very similar genetic structures. There seem to be two links between the recent beech 

species. One link is represented by Crimean beech (sometimes described as F. taurica Popl.). 

There is a rather smooth transition of allelic frequencies from F. orientalis in the Caucasus 

through Crimea, Moldova to the Romanian Carpathians (Gomory et al. 1998). The second 

link is represented by Balkan beech, as described above. Although both Balkan and Crimean 

beech seem to be transitional taxa, they differ clearly by their allelic structure. It is quite 

difficult to decide, solely on the basis of the knowledge about the allelic frequencies, which 

of these links (if any) represents a true phylogenetic transition from F. orientalis to F. sylvatica 

and which is only a product of later introgression and geneflow processes. The capacities of 

isoenzyme markers are rather limited and seem to be reached by the described studies. 

Further range-wide studies employing DNA markers, similar to that of Demesure et al. 

(1996), would therefore be very useful and could bring solutions to questions which could 

not be answered using other tools. 

Mating system and intrapopulation spatial structure 

A very detailed study of beech mating systems was performed by Merzeau (1991), the 

results were later summarized in Merzeau et al. (1992, 1994). She analyzed six populations in 

total with different characteristics (isolated treed, forest edge, forest massive, populations in 

different altitudes) and investigated also the effects of tree height and crown density on 

outcrossing rates. In general, she found very high outcrossing rates, approaching 1.0. 

Significant heterogeneity of outcrossing pollen pool frequencies was found not only among 

stands, but especially among individual trees within populations in five out of six analyzed 

stands. She concluded that although beech is a highly outcrossing species, beech 

populations cannot be considered panmictic. The intrapopulation spatial structure was 

investigated using spatial autocorrelation analysis. In most cases, the spatial distribution of 

genotypes was random, only in one forest she found a weak patchy structure. 

In two natural beech stands from Italy, Rossi et al. (1996) obtained very similar results. 

Only in dormant seeds from one stand, the outcrossing rate was significantly lower than 1.0. 

As in the previous study, pollen pool allele frequencies were significantly heterogeneous 

among parent trees.


A wide study of the spatial structure was performed in 14 Italian beech populations 

(Leonardi and Menozzi 1995). They revealed a patchy structure in several populations. 

Although they did not use equal distance classes, they found in general a high proportion of 

positive autocorrelations between like genotypes and negative autocorrelations between 

unlike genotypes in first distance class (upper limit of 32 to 77 m), with decreasing 

proportions of significant values in further distance classes. 

Starke (1996) estimated outcrossing rates in stands managed by different silvicultural 

systems. She found substantially lower outcrossing rate estimates (0.862 to 0.929). This may 

be due to a different estimation method, since she did not use the mixed mating model as in 

previous studies, but estimated outcrossing rate on the basis of xenoheterozygosity. 

Selection processes 

The discussion in the scientific community, of whether isoenzymes are selectively neutral or 

not, does not seem likely to end in the foreseeable future. However, there are several studies 

available which indicate the effect of selection due to different external factors on the allelic 

and genotypic structure of populations of forest trees, including beech. 

One of the most intensively investigated factors was air pollution. Miiller-Starck (1989) 

investigated the differences between sets of individuals apparently tolerant and sensitive to 

air pollution. He found substantially higher heterozygosity values in the set of tolerant trees 

than in the sensitive ones. The difference amounted to 25% of the actual heterozygosity and 

almost 10% of the conditional heterozygosity. Similarly, the genetic multiplicity as 

measured by the number of alleles was slightly higher in the 'tolerant' set. About 5% of 

alleles were found only in the sensitive trees. 

In the investigations of changes of the genetic structure under complex environmental 

stress, Miiller-Starck and Ziehe (1991) observed a reduced heterozygosity in 2-year-old 

seedlings, compared with the seed stage. On the other hand, the number of alleles 

decreased, which the authors ascribe to a reduced sample size. They also showed a 

directional selection at Lap-A locus. 

Brus (1996) found significant differences in allelic frequencies at Lap-A and Idh-A in a 

comparison of polluted and unpolluted beech stands in Slovenia. In addition, a complex 

multilocus response was identified: in general, the polluted stands were considerably more 

differentiated and exhibited unpredictable deviations of the allelic structures. 

Studies on the effects of other stress factors on the genetic structures of beech are scarce. 

Recently, a paper on the genetic differences in susceptibility to infestation of young beech 

trees to Cryptococcus fagisuga was published by Gora et al. 1995. They found significant differences 

of infestation rates associated with particular genotypes at Idh-A, Per-B and Mdh-B loci. 

The effect of human activities on the genetic structures is a specific problem. Kim (1980) 

compared the viability selection processes in natural conditions and in the greenhouse. He 

found a selective advantage of homozygotes at Lap-A locus in the greenhouse, whereas in 

the forest, the heterozygotes containing the a specific allele are favoured. The changes of the 

genotypic structure were observed at Acp-A locus as well. At Skdh-A locus, rare genotypes 

decreased in frequency or even vanished in the studied seedlings after 2 years in field 

conditions (Hattemer et al. 1993). 

A more profound study was published by Starke et al. (1996). They investigated the 

effects of different soil-preparation procedures on the genetic structures. Strong changes of 

genotype frequency distributions were observed at most loci. It was proved that the 

modification of environmental conditions changes selection regimes and leads to changes of 

genetic structures.

0VERVIEW PRESENTATI0NS 159 

DNA analyses 

The lack of sufficiently variable genetic markers for studying haploid tissue of broadleaved 

forest trees gives cause to develop a method for analyzing DNA from single pollen grains. 

Using the PCR technology the DNA from single pollen grains of beech (F. sylvatica) could be 

amplified. The pattern generated from a 10-base random oligonucleotide of haploid and 

diploid tissue was compared. The described method will offer a new approach for paternity 

analysis in angiosperms (Vornam 1996). 

Heinze and Geburek (1995) used PCR analysis of the total DNA from leaves to find a 

marker linked to the locus controlling leaf colour in copper beech. Among five markers 

exhibiting Mendelian segregation they found one weakly linked with the purple leaf colour. 

The Mendelian inheritance of RAPD and I-SSR markers has been assessed using a 

progeny from controlled crossings of F. sylvatica. Out of a total of 165 amplification 

products, 30 Mendelian markers with dominant mode of gene action were found. A subset 

of 11 markers has been used to estimate population parameters in a sample of 46 trees from a 

natural stand in the Northern Apennines (Italy). As expected, higher expected 

heterozygosity was observed than reported in the literature for allozyme studies of this 

species. The same subset of markers was used for assessing paternity of 81 seedlings 

belonging to 9 open-pollinated sibships of 9 individuals each. In spite of the low power of 

the analysis (only 2 unambiguous paternity assignments out of 46 potential fathers) its 

results seem in agreement with the lack of spatial clustering already known for the species 

(Troggio et al. 1996). 

Demesure et al. (1996) used the polymorphisms in the chloroplast genome of F. sylvatica, 

which have been detected by resolutive restriction site studies of PCR-amplified fragments. 

Eleven haplotypes, which could be phylogenetically ordered, were detected in a large 

survey (399 individuals in 85 populations) encompassing most of the natural range of the 

species. The high level of genetic differentiation (G st 

= 0.831), together with the highly 

structured geographic variation, contrast with the low level of nuclear genetic differentiation 

measured in previous studies with isoenzymes and indicate a low level of geneflow by 

seeds. The northernmost populations are genetically uniform, suggesting a bottleneck at the 

time of postglacial recolonization, a scenario which fits with palaeobotanical reconstructions. 

The description of the matrilineal genetic structure of this important tree species enables the 

detection of a clear case of introduction of an exotic population due to long-distance seed 

transfer. 

Practical application 

The investigations of the genetic diversity and differentiation of beech all over Europe have 

been a good basis to understand the inner structures of beech populations and the processes 

that maintain them. Naturally, parallel provenance experiments are the best way to answer 

the question of whether an intentional move of reproductive material could. be 

recommended for the establishment of beech forest stands under conditions of the 

environmental stress, or on abandoned agricultural lands outside or even inside the beech 

natural distribution range. 

The artificial regeneration will become more common for regenerating beech stands in the 

future. There are vast areas in Europe that were converted from mixed or broadleaved 

stands to coniferous monocultures. It is a common case that these stands are frequently not 

on the appropriate sites and their ecological stability is at risk. Mainly in the air-polluted 

areas the consequences are already visible. It is expected to convert these stands again into 

mixed or broadleaved stands with higher resistance potential. 

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Molotkov, P.L 1966. Bukovyje lesa i khozyajstvo v nikh [Beech forests and their 

management]. Lesnaya promyshlenhost, Moscow. 

Miiller-Starck, G. and R. Starke. 1993. Inheritance of isozymes in European beech (Fagus 

sylvatica L.). J. Heredity 84:291-296. 

Miiller-Starck, G. and M. Ziehe. 1991. Genetic variation in populations of Fagus sylvatica L. 

Quercus robur L., and Q. petraea LiebI. in Germany. Pp. 125-141 in Genetic Variation in 

European Populations of Forest Trees (G. Miiller-Starck and M. Ziehe, eds.). J.D. 

Sauerlander's Verlag, Frankfurt am Main. 

Miiller-Starck, G. 1989. Genetic implications of environmental stress in adult forest stands of 

Fagus sylvatica L. Pp. 127-142 in Genetic Effects of Air Pollutants in Forest Tree 

Populations (F. Scholz, H.R. Gregorius and D. Rudin, eds.). Springer Verlag, Berlin, 

Heidelberg. 

Paule, L. 1992. Geographic variation and genetic diversity of the European beech (Fagus 

sylvatica L.) in Europe. In Actas del Congreso Internacional del Haya (R. Elena Rosello, 

ed.). Investigacion Agraria. Sistemas y Recursos Forestales 1:C197-206. 

Paule, L., D. Gomory and J. Vysny. 1995. Genetic diversity and differentiation of beech 

populations in Eastern Europe. Pp. 159-167 in Genetics and Silviculture of Beech. Proc. of 



Poplavskaja, G.L 1927. Materialy po izucheniyu izmenchivosti krymskogo buka. Zhurnal 

russkogo botanicheskogo obshchestva (10):59-82.. 

Premoli, A. 1996. Allozyme polymorphisms, outcrossing rates, and hybridization of South 

American Nothofagus. Genetica 97:55-64. 

Pukacki, P. 1990. Odpornosc na niskie temperatury. Pp. 185-192 in Buk zwyczajny (Fagus 

sylvatica L.) (S. Biatobok, ed.). Panstwowe wydawnictwo naukowe, Poznan. 

Rossi, P., G.G. Vendramin and R. Giannini. 1996. Estimation of mating system parameters in 

two Italian natural populations of Fagus sylvatica. Can. J. For. Res. 26:1187-1192. 

Stanescu, V. 1979. Dendrologie. Universitaeta din Brasov. 

Starke, R. 1996. Die Reproduktion der Buche (Fagus sylvatica L.) unter verschiedenen 

waldbaulichen Gegebenheiten. Pp. 135-159 in Biodiversitat und nachhaltige 

Forstwirtschaft (G. Miiller-Starck, ed.). Ecomed, Landsberg. 

Starke, R., M. Ziehe and G. Miiller-Starck. 1996. Viability seleciton in juvenile populations of 

European beech (Fagus sylvatica L.). For. Genet. 3(4):217-226. 

Thiebaut, B., R. Lumaret and P. Vernet. 1982. The bud enzymes of beech (Fagus sylvatica L.) 

genetic distinction and analysis of polymorphism in several French populations. Silvae 

Genet. 31(2-3):51-60. 

Trober, U. 1995. The genetic variation in Saxon beech populations (Fagus sylvatica L.) - 

preliminary results. Pp. 168-179 in Genetics and silviculture of beech. Proc. of the 5th 

IUFRO beech symposium 1994, Denmark (S. Madsen, ed.). Forskningsserien no. 11-1995. 

Danish Forest and Landscape Institute, H0rsholm, Denmark. 

Troggio, M., E. DiMasso, S. Leonardi, M. Ceroni, G. Bucci, P. Piovani and P. Menozzi. 1997. 

Inheritance of RAPD and I-SSR markers and population parameters estimation in 

European beech (Fagus sylvatica L.). For. Genet. 3(4):173-18l. 

Turok, J. 1993. Levels of genetic variation in 20 beech (Fagus sylvatica L.) populations from 

Western Germany. Pp. 181-195 in The Scientific Basis for the Evaluation of Forest Genetic




Turok, J. 1996. Genetische Untersuchungen bei der Buche. Genetische Anpassungsprozesse 

und die Erhaltung von Genressourcen in Buchenwaldern (Fagus sylvatica L.). 

Schriftenreihe der Landesanstalt fur Okologie, Bodenordnung und Forsten. Landesanstalt 

fur Agrarordnung Nordrhein-Westfalen 8. 

Vornam, B. 1996. DNA amplification from single pollen grains of beech (Fagus sylvatica L.). 

For. Genet. 3(4):213-216. 

Vysny, J., 1. Shvadchak, B. Comps, D. Gomory and L. Paule. 1995. Geneticheskoe 

raznoobrazie i differentsiyatsiya populyatsij buka (Fagus sylvatica L.) na Ukraine. 

Ukrainskie Karpaty i prilegayushchie territorii [Genetic diversity and differentiation of 

beech populations (Fagus sylvatica L.) in Western Ukraine]. Genetica (Moscow) 

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Wulff, E.V. 1932. The beech in Crimea, its systematic position and origin. Pp. 223-261 in Die 

Buchenwalder Europas (E. Rubel, ed.). Hans Huber, Bern - Berlin.

Table 1. Number of ~rovenances in the Beech Trial Series established in 1986, 1987, 1988 and 1995 and the Institution in charge of the trial 

Person 1986 1987 1988 1995 

Country res~onsible Institution Series 1 Series 2 Series 3 Series 4 

Austria U. Schultze Federal Forest Research Centre, Vienna 49 

Belgium D. Jacques Station de Recherches Forestieres, Gembloux 73 

Bulgaria V. Karamfilov Forest Committee, Sofia 49 

Croatia J. Gracan For~st Research Institute, Jastrebarsko 51 

Czech Republic V. Hynek Forest Research Institute, Jiloviste - Strnady 49 

Denmark S. F. Madsen Ministry of Environment and Energy, H0rsholm 48 49 

France E. Teissier du Cros INRA, Avignon 17 58 50 

Germany BY W. Ruetz Centre for Tree Improvement, Teisendorf, Bavaria 57 

Germany NW M. Rogge Forest Gene Bank, Arnsberg, Northrhine-Westfalia 116 

Germany SN H. Wolf Saxonian State Centre for Forestry, Graupa, Saxony 104 

Germany ST F. Schuffenhauer District Government, Dessau, Sachsen-Anhalt 49 

Germany SH H.-J. Muhs Institute for Forest Genetics, Grosshansdorf 48 31/68 t 147 

Germany NW H.-J. Muhs Institute for Forest Genetics, Grosshansdorf 54 28 

Germany HE H.-J. Muhs Institute for Forest Genetics, Grosshansdorf 48 58 

Germany BW H.-J. Muhs Institute for Forest Genetics, Grosshansdorf 29 33 

Luxembourg F. Theisen Administration for Water and Forests, Luxembourg 49 

Ireland D. Thompson COILL TE Research Laboratory, Newtownmountkennedy 49 

Italy R. Giannini Faculty of Forestry, University, Florence 49 

The Netherlands S. de Vries Institute for Forestry and Nature, Wageningen 32 70 

Poland Z. Rzeznik Forest Dept., Akademia Rolnicza, Poznan 71 

Poland M. Sulkowska Forest. Research Institute, Warsaw 49 

Romania V. Enescu Academy of Agriculture and Forestry, Bucharest 27 

Romania V. Enescu Academy of Agriculture and Forestry, Bucharest 44 

Sweden M. Werner Forest Research Institute, Ekebo 36 

Slovakia L. Paule University of Forestry and Wood Technology, Zvolen 100 

Spain F. Puertas Tricas State Forest Service, Navarra 100 

United Kingdom N. Cundall Northern Research Station, Roslin, Midlothian 53 

Ukraine I. Shvadchak Institute of Forestry and Wood Technology, Lvov 70 

Total number of trials per series 3 5 7 23 

t There are two parallel trials at this location.

OVERVIEW PRESENTATIONS 16'7 

Seed collecting and establishment of trials 

It was aimed to have trials in all regions covered naturally by beech. However, owing to 

differing seed sets and availability because of political restrictions (e.g. former East 

Germany, Bosnia-Herzegovina) the number of trials and the regions represented vary 

considerably between the trial series. Figure 1 shows the location of the 38 field trials in 19 

countries throughout the range of distribution of beech. A trial is located in most of the 

regions, except in Bosnia-Herzegovina/Serbia (Dinaric mountains), and Switzerland/France 

(western Alps/Jura) trials, which should have been included to give an even better 

coverage. A number of trials are near the border of the range of distribution and the trial in 

Ireland is outside the natural range of beech. 

l:!,. 1986 

I> 1987 

\l 1988 

III 

1995 < 100 pr. 

11 1995> 100 pr. 

III 

11 11 

Fig. 1. Location of the trials. The 38 trials were established in 1986, 1987, 1988 and 1995 in 19 

countries. The larger squares indicate trials with 100 or more provenances; the other trials contain 

between 17 and 73; most trials have 49 provenances.


Figure 2 shows the origins of the seed samples in the different trials. The different parts 

of the area of distribution of beech are sampled with different intensity. Czech Republic, 

Germany, Romania and Slovakia are very well represented. Bulgaria, France, Poland, Spain, 

Slovenia and Ukraine are quite well represented, whereas Austria, Croatia and Italy, 

especially southern Italy, are poorly represented. The area between Bosnia-Herzegovina, 

Serbia, Macedonia, Albania and Greece is not represented. This is regrettable because in this 

area, owing to the high variety in site conditions and the long period during which beech 

has been able to adapt to these sites, a large genetic variation and special ecotypes can be 

expected. 

Fig. 2. Location of the 335 provenances included in the 38 trials established in 1986, 1987 and 1988 

(triangles) and 1995 (dots). 

Seed sample quality 

The seed samples received at the Institute differed strongly in many respects: cleanliness, 

means and duration of transport, collecting method (by hand, by nets), commercial, state or 

scientific collection, pretreatments, etc. Seed quality differed accordingly. So~e samples 

were too dry «10% water content), others were mouldy because there was too little air 

exchange in the shipping bag, some seed samples showed a lot of damaged seeds. 

Generally, seed samples from distant places which had a longer journey were in worse 

condition than samples from nearby places. 

Seed sample treatment 

The seed samples were first cleaned and their moisture content determined. Depending on 

the freshness of the seeds they were left for a period of after-ripening of up to 4 weeks. The 

seed samples from 1990 and 1991 were stored by reducing the moisture content to about 10% 

and freezing at -S°e. Before seeding, the seeds were carefully remoistened to 30% water 

content and stratified at 3°e. When dormancy was broken, the seeds were frozen with water 

at -2°C until seeding. The seeds collected in 1993 were stratified to break dormancy and, as 

all other seed samples, frozen with water at -2°C until seeding time.


The time required to break dormancy differed strongly between the provenances. This 

was due to the preconditions (e.g. after-ripening) and possibly also to genetic differences. 

These effects could not be studied because the preconditions (harvesting, transport, 

treatments) were too different. 

Seeding and rearing of plants 

Seeding was done at the nursery of the Institute at Grosshansdorf and, because there was 

not enough space, also at commercial nurseries. Germination frequency and rate differed 

strongly between provenances. There seemed to be a relationship between the time required 

to break dormancy and germination: provenances which needed a long time to break 

dormancy seemed to have low germination frequency and were slow to germinate. Also, 

these provenances were prone to fungi attack in the seed bed. Shortly after germination and 

development of cotyledons the plantlets were struck severely by various damping-off fungi 

(e.g. Phytophthora sp.) from which they usually did not recover and dropped out quickly. 

The different sensitivity of seed samples toward the fungi attack is probably not due to 

provenance differences. The duration of storage also had some influence, although there 

were seed samples collected in 1990 and 1991 which yielded a high number of plants. The 

sensitivity to fungi attack may also be the result of improper seed handling before storage 

which led to a general loss of vitality of the seeds. This can be presumed because dampingoff 

fungi are mostly not specialized but rather occur ubiquitovsly. A general loss of vitality 

might also explain the longer periods required to break dormancy, and the slow, as well as 

low, total germination. 

After the critical postgermination stage had passed, no further attack by fungi was 

observed. Following this, there was some irregular attack by Phyllaphus fagi L. The plants of 

the 1995 series, sown in spring of 1993, were undercut during the first growing season for 

easier lifting and lifted in autumn of 1994 at 2 years of age. The plants of the other series were 

lined out after 1 year, transplanted for a further 2 years and were planted as 1+2 seedlings. 

Design of the field trials 

The layout of the trials is a randomized incomplete (series 1-3) or complete (series 4) block 

design with three replications. Planting was done in rows with a space of 2xO.5 m (series 1- 

3) or 2x1 m (series 4). Each plot was laid out with 100 plants (series 1-3) or 50 plants (series 

4), resulting in a plot size of 10x10 m. Thus, a trial with 49 provenances occupies an area of 

about 1.5 ha. Plots are considered large enough to maintain the trials for 60 years. There are 

no bordering rows between the plots, usually only two rows were planted around the trials. 

Table 1 gives the number of provenances included in each trial. In most trials there are 

about 49 or more provenances included, in five trials 100 or more provenances are 

represented. In the series planted in 1995, of the total of 147 provenances only 19 are 

represented in all of the trials and 36 provenances are included in more than 75% of the 

trials. The remainder of the provenances are represented on a differing number of field 

trials depending on the number of plants available. A number of joint partners added up to 

24 mostly local provenances to their trial. 

Traits to be recorded 

The large number of trials and joint partners involved in this experiment requires precise 

definition of the characters to be recorded. Survival, plant height, trunk diameter (when 

trees have reached measurable sizes), flushing time, growth cessation, lammas shoots, stem 

form and any damages that might be observed will be recorded primarily. Additional traits 

are other form characters (branching, spiral grain, etc.) and genetic markers (isoenzymes, 

molecular). An important feature is that the position of the single trees measured is 

recorded to enable calculation of the effects of neighbouring trees as well as stand density 

effects and meaningful age-age correlations.

17'0 EUFGRGEN: SGGIAI... BRGADI...EAVES 

Participating institutions 

Table 1 lists the persons and Institutions that have established and are maintaining one or 

more trials. We are very thankful for their endeavour and contribution. We are also very 

thankful to all those, mostly foresters, who provided the numerous seed samples for this 

triaL Neither without the field trials nor without the seed samples would the establishment 

of the trial network have been possible on a Europe-wide scale. 

Database 

A data bank is located at Grosshansdorf, where the geographical and site data of the 

provenances (stand age, soil, climate, etc.), the location data of the field trials (position, soil, 

climate, etc.), management of the trials, and the data collected in the field trials for the 

different traits will be kept centrally and made available to the joint partners. Processing 

will also be performed at the data bank. 

Survival 

Survival varied between trials, depending on the local conditions during planting and the 

weather in the year of establishment. Drought and strong late frosts after planting caused 

higher mortality in some trials (Bjelovar, Croatia; Nedlitz, Sachsen-Anhalt; Bayreuth, 

Bavaria). At other locations (Attendorn, Northrhine-Westfalia), extreme high soil acidity 

(pH 3.2-3.7), and damage by voles (Sweden) caused high mortality rates. At a polder site in 

the Netherlands there was high mortality due to flooding. Of the total of 38 trials, three had 

to be given up and two trials are strongly reduced in their usability because the survival rate 

is less than 50%. Thus, the remaining 34 trials are in a state to give reliable data potentially 

for a long period, which is important in a species with long rotation periods like beech. 

Differences in the survival rates between the provenances were observed. However, no 

general trends could be established so far. 

Flushing 

The times of bud burst and leaves flushing have been recorded on a number of the field 

trials in different years (Muhs 1985, von Wuehlisch et al. 1993, 1995a; Madsen 1995). These 

results show large differences in leaves flushing time in spring. Generally, provenances from 

the eastern and southeastern part of the distribution area (e.g. Slovakia, Romania, Bulgaria), 

as well as provenances from high elevations, require a smaller heat sum for flushing and 

thus flush early. Provenances from the western part of the distribution area (Spain, France, 

The Netherlands, Belgium, England) and from low elevations where late frosts occur, 

require a higher heat sum and flush late. Depending on the temperature development 

during spring, the time difference between the earliest and last individuals to flush in a set 

of provenances in a trial can be 4 to 6 weeks. Between the different years and over a number 

of sites, a certain stability in the ranking of flushing time was found in the provenances 

analyzed. More studies are necessary to also study year-to-year and site-to-site interactions 

in the flushing reaction of the provenances. It can be concluded that this character is 

adaptive and reflects the adaptation to a certain site in respect to late frost occurrence. This 

makes flushing a very interesting character for the estimation of adaptation and adaptability 

of beech populations. 

Temperature sum requirement for bud burst 

In a further study on bud burst and leaf flushing the temperature sums at which flushing 

occurs were estimated (von Wuehlisch et al. 1995b). When summing up all degree-hours 

above 5°C from beginning of January, it was found that the investigated total of 159 

provenances required an average of 9750 degree-hours for bud burst. The first tree to flush 

required 7600, the last one 14750 degree""hours before bud bursting, which is about twice as 

much. Large differences were also found between the provenances. The first to flush

OllERlllEW PRESENTA.TIONS 1'71 

required an average of 8500, the last one an average of 11 000 degree-hours. These 

differences underline the adaptive character of this trait. It may be assumed that when a 

provenance is frequently struck by late frosts because it flushes too early, it is probably also 

not adapted to that site in other characters. 

Height growth 

Plant heights have been recorded in the year after planting and in the following years at 

regular intervals. However, at this early stage, the height increment does not reflect the 

potential growth and the effect of the local environment on the genotype. The height 

increment reflects mainly how the plants took root and how they overcame the planting 

shock. Therefore it is too early, especially for the trial planted 1995, to present height data. 

For the trials established in 1986, 1987 and 1988 some results have been published (von 

Wuehlisch et al. 1992, 1995c; Madsen 1995). No general trends could be established. In all 

regions studied, fast- as well as slow-growing provenances could be recorded. In some trials 

distinct differences between provenances were shown, in others not. In some trials with 

contrasting sites, e.g. poor, extremely acidified soil, as opposed to neutral soil, rich in 

nutrients, distinct genotype x environment interactions were found. However, among other 

trials no genotype x environment interactions could be proven. To conclude, in a species 

known for its compatibility and ability to change rank even above 42 years of age 

(Kleinschmit and Svolba 1995), the trials are still far too young to reflect the growth potential 

of the provenances. 

Significance of the trial network for the genetic conservation of beech 

Primarily, the collection of provenances and the network of trials throughout the area of 

beech occurrence opens the possibility of an evaluation of beech populations on a specieswide 

scale. This is of great value and provides the means to observe and document the 

growth and other adaptive or economically important traits of a set of provenances 

originating from most regions, on sites located in most of the regions of beech occurrence. 

The trial data will show how well populations have adapted to certain site-inherent 

environmental features, e.g. late frost occurrence, acidic or calcareous soil, etc., how nonadapted 

populations react to such situations, and how successfully they might cope with 

them. This is of great significance for formulating evaluation criteria to be able to assess the 

value of a given population in respect of the conservation of the genetic resources of beech. 


The establishment of the beech trials has been funded by the Federal Ministry of Food, 

Agriculture and Forestry, Bonn, and since January 1995 by the Concerted Action of the 

Commission of the European Communities, AAIR3 Programme, Grant No. CT94-2091, 

which is gratefully acknowledged. We also thank many colleagues who helped in the 

collecting of numerous seed samples throughout Europe and the participating institutions 

for establishing the field trials. Thanks are also due to Gertrud Willich, Anja Forster, Marion 

Korsch and many others for valuable advice and technical assistance. 

References 

Boratynska, K. and A. Boratynski. 1990. Systematyka i geograficzne rozmieszczenie. Pp. 27-73 in 

Buk Zwyczajny. Panstwowe Wydnawnictwo Naukowe, Poznan-Warszawa. 

Kleinschmit, J. and J. Svolba. 1995. Results of the Krahl-Urban beech (Fagus sylvatica L.) 

provenance experiments 1951, 1954, and 1959 in northern Germany. Pp. 15-34 in Genetics 

and Silviculture of Beech. Proc. of the 5th IUFRO beech symposium 1994, Denmark (S. 

Madsen, ed.). Forskningsserien no. 11-1995. Danish Forest and Landscape Institute, 

H0rsholm, Denmark.


Madsen, S.F. 1995. International beech provenance experiment 1983-85. Analysis of the 

Danish member of the 1983 series. Pp. 83-89 in Genetics and silviculture of beech. Proe. of 



Muhs, H.-J. 1985. International provenance trial of beech (Fagus sylvatica L.) from 1983/85. 

Mitteilungen der Bundesforschungsanstalt fur Forst- und Holzwirtschaft 150:99-104. 

Muhs, H.-J. 1988. Die Anlage des Internationalen Buchenherkunftsversuchs 1983-1985. Pp. 

77-83 in 3. IUFRO Buchensymposium (S. Korpel' and L. Paule, eds.).Hochschule fUr 

Forstwirtschaft und Holztech-nologie, Zvolen. 

Muhs, H.-J. and G. von Wuehlisch. 1992. Research on the improvement of beech in the last 

decade. Pp. 311-318 in Proc. of Int. Congress on Beech, Pamplona 1992, Investigacion 

Agraria, Sistemas y Recursos Forestales, Vol. I (R. Elena Rossello, ed.). 

Muhs, H.-J. and G. von Wuehlisch. 1993. Research on the evaluation of forest genetic resources 

of beech - a proposal for a long-range experiment. Pp. 257-261 in The Scientific Basis for the 


Ahrensburg 1993 (H.-J. Muhs and G. von Wuehlisch, eds.). Working Document of the EC, 

DG VI, Brussels. 

Teissier du Cros, E. and 1. Bilger. 1995. Conservation of beech genetic resources in France. 

Pp. 196-209 in Genetics and Silviculture of Beech. Proe. of the 5th IUFRO beech 


and Landscape Institute, H0rsholm, Denmark. 

Turok, J. and H.H. Hattemer. 1995. Gene resources in beech: which population should be 

chosen Pp. 210-225 in Genetics and Silviculture of Beech. Proc. of the 5th IUFRO beech 


and Landscape Institute, H0rsholm, Denmark. 

Wuehlisch, G. von and H.-J. Muhs. 1992. International provenance trial of Fagus sylvatica L.: First 

results at 2 sites up to 7 years of age. Pp. 311-318 in Proe. of Int. Congress on beech, Pamplona 

1992, Investigacion Agraria, Sistemas y Recursos Forestales, Vol. I (R. Elena Rossello, ed.). 

Wuehlisch, G. von, D. Jacques and H.-J. Muhs. 1993. Phenological differences between beech 

provenances. Pp. 229-232 in The Scientific Basis for the Evaluation of Forest Genetic 



Wuehlisch, G. von, H. Duval, D. Jacques and H.-J. Muhs. 1995a. Stability of differences in 

flushing between provenances in different years and at different sites. Pp. 83-89 in 

Genetics and silviculture of beech. Proe. of the 5th IUFRO beech symposium 1994, 

Denmark (S. Madsen, ed.). Forskningsserien no. 11-1995. Danish Forest and Landscape 

Institute, H0rsholm, Denmark. 

Wuehlisch, G. von, D. Krusche and H.-J. Muhs. 1995b. Variation in temperature sum 

requirement for flushing of beech provenances. Silvae Genet. 44:343-346. 

Wuehlisch, G. von., H.-J. Muhs and D. Krusche. 1995e. Early provenance variation at sites of 

the international beech provenance trial 1983/85. Pp. 45-50 in Genetics and Silviculture of 

Beech. Proe. of the 5th IUFRO beech symposium 1994, Denmark (S. Madsen, ed.). 

Forskningsserien no. 11-1995. Danish Forest and Landscape Institute, H0rsholm, Denmark.

PROGRAMME 173 

Programme 

22 and 23 October 

Arrival of participants at Bordeaux-Merignac Airport 

23 October 

OSh30 

09hOO-llhOO 

IlhOO-12hOO 

12hOO 

13hOO-14h30 

15hOO-15h15 

15h15-15h30 

15h30-16h15 

16h15-16h30 

16h30-17h15 

17h15-1ShOO 

lShOO-lSh45 

19hOO 

20hOO 

24 October 

OSh30-10hOO 

10hOO-l0h30 

10h30-12h30 

12h30-14hOO 

14hOO-16h30 

16h30-17hOO 

17hOO-lSh45 

19hOO 

20hOO 

25 October 

OSh30-10hOO 

1 OhOO-l Oh30 

10h30-12h30 

12h30-13h30 

13h30-17hOO 

17hOO-lSh30 

19h30 

Departure for a visit of the molecular marker labs at INRA Forest Research 

Station 

Visit of the molecular marker labs 

Registration 

Welcome aperitive with members of the 'Commission Nationale de 

Conservation des Ressources Genetiques Forestieres' 

Lunch at INRA restaurant in Pierroton 

Welcome address (M. Arbez, INRA) 

Introduction and format of the meeting (J. Turok, IPGRI) 

Genetics of European oaks (A. Kremer) 

Coffee break 

Oak decline in Europe (T. Oszako) 

Genetic variation of beech in Europe (L. Paule) 

International provenance experiment on beech (R. Stephan) 

Transfer by bus to Bordeaux city 

Dinner in Bordeaux City 

Regional working groups (country reports): session I 

Coffee break 

Regional working groups (country reports): session II 

Lunch 

Regional working groups (country reports): session III 

Coffee break 

Summary: Common needs, priorities and existing capacities on the 

conservation and sustainable use of genetic resources of oak and beech in 

Europe 

Transfer by bus to Bordeaux 

Dinner in Bordeaux City 

Network session I: Coordination of ongoing activities at a European level 

Coffee break 

Network session II: Development of shared Network tasks according to the 

needs and capacities 

Lunch 

Excursion to local oak forests (riverside and coastal sand dunes) 

Establishment of a workplan, conclusions and approval of the Report of the 

meeting 

Farewell dinner at restaurant Les Pavois in Arcachon Bay 

26 October 

Departure of participants by shuttle bus service to the Airport

174 EUFORGEN: SOCIAl... BROADl...EAVES 

Email: menozzi@dsa.unipr.it

PARTICIPANTS 175 

Mr Edgars Smaukstelis 

Forest Research Station Kalsnava 

Jaunkalsnava 

4860 Madonas distr. 

Latvia 

Tel: +371-48 37591 

Fax: +371-48 23 891 

Mr Virgilijus Baliuckas 

Dept. of Forest Genetics and Reforestation 

Lithuanian Forest Research Institute 

4312 Girionys, Kaunas 

Lithuania 

Tel: +370-75472 45 

Fax:+370-754 7446 

Email: a 

Mr Jean- Fran


Mr Patrick Bonfils 

Swiss Federal Institute for Forest, Snow and 

Landscape, WSL 

Ziircherstr. 111 

8903 Birmensdorf 

Switzerland 

Tel: +41-17392363 

Fax: +41-1 739 22 15 

Email: 

Ms Svitlana A. Los 

Ukrainian Institute of Forestry and Forest 

Melioration 

Pushkinska str. 86 

310024 Kharkiv 

Ukraine 

Tel: +380-572 431549 

Fax: +380-572 43 25 20 

Email 

Mr Jozef Turok 

EUFORGEN Coordinator 

IPGRI 

Via delle Sette Chiese 142 

00145 Rome, Italy 

Tel: +39-0651892250 

Fax: +39-06 575 03 09 

Email:

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