e u r o p e a n f o r e s t g e n e t i c r e s o u r c e s p r o g r a m m e Use and transfer of forest EUFORGEN reproductive material in Europe in the context of climate change

Monika Konnert, Bruno Fady, Dusˇ an Gömöry, Stuart A’Hara, Frank Wolter, Fulvio Ducci, Jarkko Koskela, Michele Bozzano, Tiit Maaten and Jan Kowalczyk U s e a n d T r a n s f e r o f f r M

Bioversity International is a global research-for-development organization. We have a vision – that ag- ricultural biodiversity nourishes people and sustains the planet. We deliver scientific evidence, manage- ment practices and policy options to use and safeguard agricultural and tree biodiversity to attain sus- tainable global food and nutrition security. We work with partners in low-income countries in different regions where agricultural and tree biodiversity can contribute to improved nutrition, resilience, produc- tivity and climate change adaptation. Bioversity International is a member of the CGIAR Consortium – a global research partnership for a food-secure future.

European Forest Genetic Resources Programme (EUFORGEN) is an instrument of international co- operation promoting the conservation and appropriate use of forest genetic resources in Europe. It was established in 1994 to implement Strasbourg Resolution 2 adopted by the first Ministerial Conference of the FOREST EUROPE process, held in France in 1990. EUFORGEN also contributes to implementa- tion of other FOREST EUROPE commitments on forest genetic resources and relevant decisions of the Convention on Biological Diversity (CBD). Furthermore, EUFORGEN contributes to the implementation of regional-level strategic priorities of the Global Plan of Action for the Conservation, Sustainable Use and Development of Forest Genetic Resources (GPA-FGR), adopted by the FAO Conference in 2013. The Programme brings together experts from its member countries to exchange information and experienc- es, analyse relevant policies and practices, and develop science-based strategies, tools and methods for better management of forest genetic resources. Furthermore, EUFORGEN provides inputs, as needed, to European and global assessments and serves as a platform for developing and implementing European projects. EUFORGEN is funded by the member countries and its activities are mainly carried out through working groups and workshops. The EUFORGEN Steering Committee is composed of National Coordi- nators nominated by the member countries. The EUFORGEN Secretariat is hosted by Bioversity Interna- tional. Further information on EUFORGEN can be found at www.euforgen.org.

The geographical designations employed and the presentation of material in this publication do not im- ply the expression of any opinion whatsoever on the part of Bioversity 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 organizations.

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Citation: Citation: Konnert, M., Fady, B., Gömöry, D., A’Hara, S., Wolter, F., Ducci, F., Koskela, J., Bozzano, M., Maaten, T. and Kowalczyk, J. European Forest Genetic Resources Programme (EUFORGEN). 2015. Use and transfer of forest reproductive material in Europe in the context of climate change. European Forest Genetic Resources Programme (EUFORGEN), Bioversity International, Rome, Italy. xvi and 75 p.

Cover photos/illustrations: Ewa Hermanowicz (on the right) and Fulvio Ducci (on the left) Layout: Ewa Hermanowicz

ISBN 978-92-9255-031-8

Bioversity International Via dei Tre Denari, 472/a 00057 Maccarese Rome, Italy © Bioversity International 2015 ii A ut h o r s

Authors

Monika Konnert Fulvio Ducci Bayerisches Amt fuer forstliche Saat- und Consiglio per la ricerca in agricoltura e Pflanzenzucht (ASP), Teisendorf l’analisi dell’economia agraria, Centro di Germany Ricerca per la Selvicoltura (CREA-SEL), Arezzo Bruno Fady Italy Institut national de la recherche agronomique (INRA) - Ecologie des Forêts Jarkko Koskela Méditerranéennes (URFM), Avignon Bioversity International, Rome France Italy

Dušan Gömöry Michele Bozzano Technická univerzita vo Zvolene, Lesnícka Bioversity International, Rome fakulta, Zvolen Italy Slovakia Tiit Maaten Stuart A’Hara Eesti Maaülikool, (EMÜ) Forest Research, Northern Research Station, Metsandus- ja maaehitusinstituut Roslin Tartu United Kingdom Estonia

Frank Wolter Jan Kowalczyk Administration de la nature et des forêts Instytut Badawczy Leśnictwa (IBL) Diekirch Sękocin Stary Luxembourg Poland

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Preface

Forest regeneration, whether natural or artificial, is based on the utilization of forest genetic resources (i.e. genetic material of forest trees that is of actual or potential use for humans). Natural regeneration relies on genetic material that is already available on a particular site, while artificial regeneration, carried out through seeding or planting, typically involves transferring forest reproductive material (FRM) (i.e. the parts of a tree that can be used for reproduction) from other locations to the site. In Europe, much of the material used for artificial regeneration is produced and transferred within a single country. However, FRM, usually in the form of seeds or cuttings, is increasingly traded across national borders, especially within the European Union. As forest managers and owners seek to minimize the costs of establishing new forests, mistakes are made in matching the material purchased with ecological site conditions, and in ensuring high genetic and physiological quality of the material.

This phenomenon is nothing new as FRM has been traded in Europe for centuries. However, climate change is likely to increase the future demand for imported FRM as forest managers and owners try to identify tree species and provenances that will be able to grow in their land under new climatic conditions. According to the Intergovernmental Panel on Climate Change (IPCC), the global average surface temperature (combined land and ocean temperatures) has increased by nearly 1°C during the period 1901–20121. The recent IPCC report concludes that it is very likely that temperature will continue to increase throughout the 21st century in different parts of the world, including Europe. In the same report, the IPCC projects that summer temperature (June-August) will increase by 3–4°C in most part of Europe by 2081–2100, and even 4–5°C in some places in the Mediterranean region. This will alter the environmental conditions to which European forest trees are adapted and may even create novel climatic conditions. As a result, some parts of the current distribution ranges of forest trees are expected to become unsuitable while new areas are likely become suitable for many species in higher latitudes or altitudes. The accelerating speed of climate change has raised serious concerns on how tree species can cope with the projected changes. Furthermore, the warming climate will probably facilitate the spread of pests and diseases, creating an additional threat to forest trees and their populations.

1 IPCC. 2013. Climate Change 2013: The Physical Science Basis. Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change [Stocker, T.F., D. Qin, G.-K. Plattner, M. Tignor, S.K. Allen, J. Boschung, A. Nauels, Y. Xia, V. Bex and P.M. Midgley (eds.)]. Cambridge University Press, Cambridge, United Kingdom and New York, NY, USA. 1535 p. doi:10.1017/CBO9781107415324.

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Most European countries have recommendations or guidelines for selecting species and provenances that can be used in a given site or zone. However, these recommendations are mostly based on present or past climatic conditions. They therefore provide limited advice for selecting FRM so that the next tree generations will thrive until the end of their rotation periods under rather different climatic conditions than today. More and more FRM is traded across borders but the national guidelines rarely offer help in deciding where the imported material should be used.

Solutions for these problems and challenges have been explored through European collaboration on forest genetic resources. Various European projects have tested the feasibility of harmonizing national provenance regions across the continent and have re-evaluated existing provenance trials to predict how climate change will affect forest growth and how individual species and provenances will perform under changing climatic conditions. Unfortunately, much of the new information produced by these efforts is being discussed and debated only by scientists, while forest owners, managers and policy-makers often remain unaware of the potential that the use of forest genetic resources offers for facilitating the adaptation of forests to climate change.

During the past decade, the European Forest Genetic Resources Programme (EUFORGEN) has increased its efforts to promote the conservation and sustainable use of genetic resources that can be adapted under climate change and to facilitate dialogue among scientists, forest owners, managers and policy-makers on this issue. EUFORGEN was established in 1994 to coordinate pan-European collaboration on forest genetic resources as part of the Ministerial Conferences on the Protection of Forests in Europe (now called the FOREST EUROPE process). During Phase IV (2010-2014), EUFORGEN had three objectives: (1) promote appropriate use of forest genetic resources as part of sustainable forest management under climate change; (2) develop and promote pan-European genetic conservation strategies and improve guidelines for management of genetic conservation units and protected areas; and (3) collate, maintain and disseminate reliable information on forest genetic resources in Europe.

This report presents the findings and recommendations of the EUFORGEN working group on FRM. The report presents possible approaches for the use and transfer of FRM under climate change and identifies critical factors; it does not attempt to provide field-level guidelines for using and transferring the material. The working group held two meetings, the first one hosted by Bioversity International in Maccarese, Italy, 28–30 March 2012, and the second one by the Bavarian Office

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for Forest Seeding and Planting (ASP) in Freising, Germany, 4–6 July 2012. The EUFORGEN Steering Committee discussed an earlier draft of this report during its 8th meeting, held in Paris, France, 27–29 November 2012. The updated draft report was then presented to a larger group of experts for further discussion at the EUFORGEN workshop on FRM that was organized in Kostryca, Poland, 1–3 October 2013. After the workshop, the working group incorporated the comments received to the revised draft report and presented it to the Steering Committee at its 9th meeting, held in Tallinn, Estonia, 3–5 December 2013, for approval. The Steering Committee endorsed the draft report, provided some additional minor comments, and requested the working group to finalize the report for publication. The working group members gratefully acknowledge inputs and comments received from all national experts who contributed to the preparation of this report.

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Contents

Authors iii Preface v Acronyms used in the text xi Executive summary xiii Introduction 1 Climate change and the use of forest reproductive material 5 The impact of natural processes and biological factors on genetic variation 5 Identification of critical factors for persistence under climate change 7 What is ‘local’ and is the local forest reproductive material always the best? 10 The challenges of using the right forest reproductive material under climate change 11 Legal and policy frameworks 15 Council Directive 1999/105/EC 15 OECD forest seed and plant scheme 16 National law on forest reproductive material – similarities and differences 16 Recent developments in policy and legal frameworks 19 New traceability and certification systems for forest reproductive material by means of genetic markers 19 The Nagoya Protocol on Access and Benefit-Sharing 20 Current guidelines and recommendations related to forest reproductive material 23 Recommendations adopted by FOREST EUROPE 23 Regional activities – the Silva Mediterranea example 25 National provenance recommendations and support tools 26 Scientific and practical considerations on the choice of provenances under climate change 29 Alternative approaches for seed zone delineation and seed transfer recommendations under climate change 30 Scientific basis for forest reproductive material transfer guidelines 35 Methodological aspects of forest reproductive material transfer based on provenance research 35 Case studies and transfer recommendations in selected species 38 Challenges and opportunities 51 Potential of tree breeding to meet conservation needs and climate change challenges 51 Marker-assisted breeding and genomics approaches 55 Molecular markers for genetic characterization of forest reproductive material 57 Advances in phenotyping and phenomics 59 Recommendations 61 References 65 Annex 1 75

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x A c r o n y m s

Acronyms used in the text

ABS Access and Benefit Sharing COST European Cooperation in Science and Technology EST expressed sequence tag EUFORGEN European Forest Genetic Resources Programme FAO Food and Agriculture Organization of the United Nations FGR forest genetic resources FRM forest reproductive material G×E genotype-by-environment GEBV genomic estimated breeding values GIS geographical information system GWAS genome-wide association studies INC Intergovernmental Negotiating Committee IPCC Intergovernmental Panel on Climate Change IUFRO International Union of Forest Research Organizations masl metres above sea level MAT mutually agreed terms MPBS Multiple Population Breeding System NGS next-generation sequencing OECD Organisation for Economic Co-operation and Development PIC Prior Informed Consent RAD restriction-site-associated DNA [marker] QTL Quantitative Trait Locus SNP Single Nucleotide Polymorphism

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Executive summary

Forest regeneration may be natural or artificial. Both depend on forest genetic re- sources, but while natural regeneration relies on what is already available on a given site, artificial regeneration often makes use of the deliberate transfer from elsewhere of forest reproductive material (FRM). This material may be in the form of seeds, seedlings or cuttings.

The selection of suitable FRM has assumed a new importance both because trees are long-lived species and because rapid climate change will have an impact on the environmental conditions of the trees as they grow and mature. The long-standing importance of FRM to forestry and the cross-border trade in FRM have resulted in several European countries exercising some control over sources of FRM and their selection. Climate change is one reason why countries need to re-evaluate and mod- ify their policy framework and guidelines on the use of FRM. An important addi- tional practical challenge is that to an unprecedented degree forest managers must now consider the climate that a new generation of trees might experience in future, in order to select material that will thrive now, under the present climate, and also be able to withstand predicted climate. Furthermore, many forest owners think of FRM as a cost to be minimized rather than as an investment for which they should be seeking better returns.

Against this background, a working group under EUFORGEN was tasked to report on the use and transfer of FRM to respond to the challenges of climate change. The working group examined scientific research on provenance and adaptation, includ- ing several case studies of transfer, the existing regulatory framework and recent policy developments, guidelines on FRM transfer and their scientific basis, and fu- ture challenges and opportunities.

The group concluded that:

• Transfer of FRM is a valuable option to adapt forests to climate change, although there may be limits to the transfer of FRM. • Local provenances may not always be the best source of FRM. • Before considering changing tree species, forest managers should consider de- ploying other, well-tested provenances of the existing tree species. At the Euro- pean level, recommendations on FRM transfer must be revised and harmonised

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and at the same time, all stages of production and marketing should be more stringently controlled.. • Tree breeding also offers opportunities to assist forests and forest management to adapt to climate change. • Improved documentation is crucial to ensuring that today’s use of FRM can inform tomorrow’s choices, just as past efforts have helped to guide today’s recommendations. • Basic research on adaptation of forest tree populations, along with provenance research, should continue and be strengthened, and the results disseminated in forms that forest owners and managers and policy makers can use.

Provenance, adaptation and suitability The term “provenance” refers to the source of FRM, and material of known prov- enance has been a feature of forest regeneration for many decades. The working group considered historical and current uses of FRM of known provenance, includ- ing the use of provenance trials to supply additional information. Provenance trials, where material of different provenances are planted in a single place (common gar- den trials) or at different locations spanning a range of environmental conditions, have been instrumental in revealing genetic variation among provenances, as well as differences among provenances in the plasticity of phenotypic responses to fluc- tuating environmental conditions. A key conclusion is that local provenances are often outperformed by other provenances for many traits.

In long-lived species like trees, populations have been experienced natural selection, possibly over multiple generations, on a given site. However, the original popula- tion on which natural selection acted consisted only of those seedlings present at the start of selection. They may not have the genetic diversity or phenotypic plasticity to guarantee good performance under changed conditions. A different population from further away, which may have experienced selection under conditions more like those forecast for the site to be reforested, might represent a more suitable seed source. Modern molecular tools and other analytical approaches offer new scientific methods for evaluating the suitability of different provenances for different sites, in addition to underpinning a deeper understanding of adaptation. Research can also deepen understanding of the strong GxE interaction (interaction between genotype performance and environmental conditions) that operates on many forest tree spe- cies. The best provenance on one site is unlikely to be the best on a different site.

The scientific breeding of tree species, like all forms of selection, inevitably results in the loss of genetic diversity. Nevertheless, intensive tree-breeding efforts are useful

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in improving FRM, provided attention is given to traits that will be most important in the future under different climate regimes. Drought resistance and phenotypic stability (i.e. the ability to maintain superior productivity across a range of sites) could be useful targets for breeding, which will be helped by molecular techniques such as marker-assisted breeding, high-throughput phenotyping and others.

Law and policy Council Directive 1999/105/EC provides the legal basis for regulating the produc- tion and marketing of FRM within the European Community, and all Member States have enacted national legislation to meet the goals of the directive. The working group examined the details of the directive and of national legislation, along with other sets of standards, and points out that none of them currently takes account of climate change. The report identifies opportunities to change some aspects of reg- ulation, for example making use of modern genetic tools for provenance control. A fruitful avenue may be to extend control from the production and marketing of FRM to its use, possibly in the form of decision-support tools to help forest managers to choose the best source of FRM.

Going forward, the European Commission is currently reviewing the control of plant reproductive material. This review will result in a Regulation, rather than a Directive, harmonising pan-European approaches. The implications of the Nagoya Protocol for the exchange of forest genetic resources for research and development purposes remains unclear.. The Nagoya Protocol can potentially hamper research and development efforts by, for example, increasing transaction costs.

A suitably adapted policy framework could do much to improve the outlook for the transfer of FRM in response to climate change. To ensure that such policy framework can be put in place, however, additional research will be needed, along with the dis- semination of research results in usable form.

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Introduction

The use and transfer of forest reproduc- Throughout the 19th century and early tive material (FRM) (i.e. seeds, cuttings 20th century, the international transfer or other propagating parts of a tree) of seeds continued in large quantities have a long history in Europe. Millen- in many parts of Europe. The demand nia ago, ancient Greeks and Romans for seed was high as many countries cultivated chestnut (Castanea sativa) started to regenerate artificially their and stone pine (Pinus pinea) mainly for overexploited forests. Despite the fact food production, and also traded their that the importance of using appropriate seeds across the Mediterranean region seed sources was demonstrated by (Conedera et al., 2004; Vendramin et practical experiences and the early al., 2008). In the 18th century, seeds of scientific studies, many reforestation several forest trees, such as Scots pine efforts seem to have paid little attention (Pinus sylvestris), Norway spruce (Picea to this aspect. Historical documents abies), European larch (Larix decidua) and indicate that the reforestation efforts oaks (Quercus spp.), were widely traded were not always successful and that and transferred for forestry purposes several countries tried to limit the use of across European countries (Tulstrup, foreign or unknown seed sources. 1959). The field testing of tree species and their different seed sources, includ- At the beginning of the 20th century, ing several tree species introduced from more systematic assessment efforts North America, started more than two were initiated for several tree species hundred years ago (see König, 2005, in Europe (see König, 2005). The first for a comprehensive review on prove- international provenance trial for Scots nance research in Europe). These early pine was established in 1907 under the testing efforts revealed that the origin auspices of the International Union of of the seed had a major influence on the Forest Research Organizations (IUFRO). success of tree planting efforts. This is This was followed by the establishment considered the starting point for mod- of a second series of IUFRO trials ern provenance research, which has for Scots pine in 1938–39, and then increased our understanding on the ef- a third series in 1982. As part of the fects of climate, site conditions and ge- IUFRO collaboration, an international netic resources on tree growth (König, provenance trial was also established 2005). for Norway spruce in 1938. Later on,

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many countries established additional to different environmental conditions provenance trials for these and several (e.g. Rehfeldt, Wykoff and Ying, 2001; other native tree species, such as Rehfeldt et al., 2002) and the degree of silver fir (Abies alba), Mediterranean this plasticity varies among populations firs (Abies spp.), European larch (Larix and species (Aitken et al., 2008; Skrøppa decidua), sessile oak (Quercus petraea) and Kohlmann, 1997). Climate change and pedunculate oak (Q. robur), as well is likely to favour genotypes with high as introduced tree species (e.g. grand levels of plasticity, whereas low plasticity fir (Abies grandis), Sitka spruce (Picea may lead to extinction (Rehfeldt, Wykoff sitchensis) and Douglas fir (Pseudotsuga and Ying, 2001). However, the buffering menziesii)). Subsequently, several capacity offered by phenotypic European countries launched breeding plasticity is limited (e.g. Mátyás, 2007). programmes for many of these tree Furthermore, the natural migration species in the 1950s and 1960s. speed of forest trees is considerably slower than the pace of climatic changes In the 1990s, when climate change (Jump, Mátyás and Penuelas, 2009) so started raising concerns, the provenance their migration potential is also limited. trials were re-discovered as they offer readily available empirical data for In the past, environmental conditions studying the impact of climate change largely guided the selection of FRM to on tree growth in field conditions (e.g. be used in a given site. Now climate Mátyás, 1996). Although European change makes this more difficult as forests are relatively well managed, forest owners and managers will also climate change is considered to be an have to consider what sort of climate the additional threat, especially for many new tree generation might experience scattered tree species and marginal during the next 50–100 years and try to populations of even common and widely select material which would not only occurring species. Tree populations have thrive under the present climate but three options to avoid extinction under also withstand the predicted changes changing climate: (1) local persistence in climatic conditions. Unfortunately, using their inherent flexibility (or many forest owners and managers “phenotypic plasticity”) to a wide do not pay enough attention to this. range of environmental conditions; Furthermore, they rarely consider the (2) local persistence via reproduction selection and purchase of FRM as an and increased genetic adaptation to new investment, but rather as a cost that conditions; and (3) survival through should be minimized. migration (Aitken et al., 2008). The provenance trials have shown that trees The use and transfer of FRM in the exhibit different phenotypic responses context of climate change was one

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of the topics which was discussed provenance requirements as part of their during a FOREST EUROPE workshop grant schemes supporting tree planting organized by EUFORGEN and IUFRO efforts. in 2006 (see Koskela, Buck and Teissier du Cros, 2007). The workshop noted The Forest Management Network noted that the impacts of climate change on that the production part (seed sources forests will vary in different parts of and seed supply) works rather well in Europe, bringing both opportunities European countries, while there are and threats. It stressed the fact that some problems with the information. forest genetic resources play a key role Scientists and forest professionals have in maintaining the resilience of forests a wealth of information, which is not to the threats and in taking advantages always easily accessible or available of the opportunities. Furthermore, the for forest owners and those who are workshop recommended development contracted to plant seedlings. When the of pan-European guidelines for the information is available, it is sometimes transfer of FRM on the basis of the latest neglected by forest owners and tree scientific knowledge, to maintain the planters. The Network concluded that productivity of European forests and the use part is the most critical issue, as to facilitate the adaptation of forests to lack of knowledge, market forces and climate change. trade mechanisms often work against the use of high-quality FRM (in terms of Between 2005 and 2009, the EUFORGEN both genetic and physiological quality). Forest Management Network also The Network also found that, when focused on issues and problems related low-quality material was used or when to the use of FRM. Experts participating the material was not well adapted to site in this network conducted two surveys conditions, it usually took 5 to 10 years in 2006–2007: one on relevant policies for problems (e.g. frost damage; low and practices related to genetic resources vigour; susceptibility to pests, diseases, and forest management, and another one wind or snow; etc.) to become visible. on tools and mechanisms to promote the However, in some cases, these problems use of high-quality FRM. The surveys surfaced only after more than 30 years. found that many countries promote tree planting efforts through legal and policy In 2010, the EUFORGEN Steering instruments. However, several of these Committee, consisting of representatives countries promote largely the use of from all member countries, decided to local provenances, and some countries establish a working group to review have even banned the use of non-local the latest information available on provenances or introduced tree species. the use and transfer of FRM in the In addition, some countries have specific context of climate change, and to

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make recommendations for further 6. Analyse if any relevant information action at the pan-European level. More should be added to the accompanying specifically, the Steering Committee documents as specified in the EC Di- requested the working group to: rective and other relevant schemes covering the movement of FRM. 1. Review existing work from 7. Compile a list of existing models and EUFORGEN Networks and relevant tools that can be used for future for- European projects. est management planning and trans- 2. Synthesize existing (national) fer of FRM. guidelines. 8. List the issues related to the climate 3. Select (widely used) model species. change context. 4. Identify critical factors related to cli- 9. Prepare a draft report (including mate change and future needs associ- recommendations). ated with transfer of FRM. This report presents in detail the 5. Summarize lessons from provenanc- findings and recommendations of the es trials for seed transfer. working group.

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Climate change and the use of forest reproductive material

Genetic diversity is the basis of individuals (or groups of individuals) evolution. Selection, gene flow and as distinct from variation caused by drift together form the engine of environmental influences. The unique evolution, the processes by which genetic composition of an individual genetic diversity can be modified when (its genotype) will determine its environmental conditions change. performance (its phenotype) at a Without genetic diversity, adaptation particular site. Successful deployment to novel environmental conditions is of FRM can be achieved, now and in the nearly impossible. Because of climate future, by assessing genetic variation change, local conditions will be and understanding how it is structured severely modified throughout species at the individual, population and species distribution ranges by the end of the levels. 21st century. Choosing the right FRM is a challenge for forestry when average Genetic variation can be categorized and critical environmental conditions into two types: neutral and adaptive. are changing fast. Neutral variation results from DNA sequence differences among individuals that do not directly affect their ability The impact of natural processes and to survive and reproduce. Adaptive biological factors on genetic variation variation results from DNA sequence The suitability of FRM to a particular differences which do affect the overall site is judged by its ability to survive, fitness of an individual and thus can be grow and reproduce at that location. selected for or against in a given habitat. This ability derives from how particular Several key evolutionary processes genes and gene combinations are affect genetic variation: mutation, drift, expressed at the location, i.e. what gene flow and natural selection (Box 1). phenotypic traits will arise under critical biotic and ecological factors at that site A part of the observed heritable and whether these traits can be modified phenotypic variation does not result when environmental conditions change from DNA sequence changes but (i.e. phenotypic plasticity). Genetic rather from changes in gene expression diversity is the term used to describe triggered by particular environmental differences in DNA sequence between conditions during key life stages. This

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Box 1. The major processes that affect genetic diversity within and among tree populations Mutation is a rarely occurring event that acts on the DNA to create new genetic variation within a species. It is a random event that can alter both neutral and adaptive genetic variation. Mutations at adaptive genes can either confer a selective advantage or, more frequently, disadvantage, for the individual, or have no effect at all.

Gene flow and migration Gene flow – the migration of genetic variants via natural dispersal of seed, pollen or vegetative propagules – acts to homogenize both neutral and adaptive variation between tree populations. This connectivity may increase the capacity of a given forest to respond to changing selection pressures by providing a regular input of genetic material from trees adapted to different ecological conditions. However, gene flow can also be maladaptive, particularly towards range margins or at ecologically marginal sites, because the material that arrives at a new location may come from a habitat with widely differing ecological conditions.

Natural selection is any factor that prevents or reduces reproduction of certain individuals in a population, and represents a selection pressure. Since mortality is often highest when trees are at the seedling or sapling stage, this is the point at which natural selection could have its greatest influence, at least for certain traits. Natural selection acts on phenotypic traits, thus on the adaptive variation, but it can also be indirectly observed in neutral variation if selection has caused demographic bottlenecks.

Genetic drift is the random reduction of genetic variation in a population of finite size. Genetic drift affects mostly neutral genetic diversity. It increases when population size decreases and its effects are largest in small isolated populations, typically at ecological and range margins.

potentially reversible variation is apply to other fitness-related traits referred to as “epigenetic”. Epigenetic as well. Both genetic and epigenetic changes are generally induced by variation influence how trees survive, environmental stimuli; in this way, a grow and reproduce in a given tree may “remember” the environment location. The extent of this variation it is exposed to and transmit this can be measured using common- information to its progeny. There is garden trials and molecular markers. evidence for such memory effects and Common-garden trials, also known their epigenetic nature for phenology as transplant experiments, i.e. field in conifers (Johnsen et al., 2005; Skrøppa tests that compare genetic material of et al., 2009), but such phenomena different geographical (provenance) may be much more widespread and or genealogical (progeny) origins in

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a common environment, have a long the circum-annual ontogenetic rhythm history in forestry. They have been is internally regulated and proceeds used for assessing adaptive variation almost regardless of external signals, and selecting material for breeding. Up climatic conditions play an essential until recently, molecular techniques role in the temporal regulation of typically provided information on growth and reproduction. Timing of neutral variation (drift and gene biological processes in plants results flow) and mating patterns in natural from evolutionary trade-offs between populations and seed orchards. Recent conflicting needs. In the case of advances in molecular techniques are vegetative phenology (budburst,­ shoot beginning to allow links to be made growth cessation, frost hardening, etc.), between adaptive traits and molecular for example, the trade-off is mainly variation, and to decipher the genetic between the length of the growing and molecular basis of adaptation in season and the risk of frost damage forest trees. (Gömöry and Paule, 2011; Leinonen and Hänninen, 2002). Under rapidly changing climate conditions, these Identification of critical factors for persistence strategies and trade-offs are severely under climate change challenged (Pigliucci and Marlow, 2001; Life history traits, such as duration of Roff, 2000). juvenile and adult phases, timing and intensity of growth, onset of aging, Climate change may push species beyond or the temporal sequence of events their ecological niche limits and thus and processes such as reproduction, drive tree populations to local extinction. nutrient storage, seed dispersal or Alternatively, extreme weather events some physiological or morphological or shifts in annual weather pattern characteristics (i.e. bark thickness or may disturb climatic signals received sprouting ability) are traits of species, by plants and consequently disrupt populations or individuals that are adaptive strategies and trade-offs. shaped by the interplay of biotic and Extinction of local populations is the abiotic factors. They are associated ultimate consequence of these processes, with variation in fertility and survival and even when tree populations throughout life and, thus, have manage to persist (for example through evolutionary implications. Life history processes such as phenotypic plasticity traits are subjected to complex trade- and genetic adaptation), productivity offs that have been shaped over long losses are a serious concern for practical evolutionary time scales and constitute forestry. Furthermore, if climate change evolutionary strategies under mostly adversely influences reproduction stable environments. Although a part of processes, it may also endanger

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the persistence of tree populations. Newton et al., 1991). Ultimately, drought Mitigation of, and adaptation to, climate stress can lead to adult tree dieback change through a proper choice of FRM and mortality. Xerothermic conditions requires that there is genetic variation (when high temperatures combine with in resistance or tolerance to climate- intense drought) may be accentuated in associated damaging factors. Several the Mediterranean, where summers are abiotic factors and life history traits are already dry and year-to-year variability critical for European FRM under climate of rainfall is high (Scarascia-Mugnozza change. et al., 2000). The Mediterranean region of southern Europe is expected to lose Among the critical abiotic factors, almost one-quarter of its plant species drought stress is generally considered by 2100 (EEA, 2007; Thomas et al., 2004; to be the main problem. Most Bakkenes, Eickhout and Alkemade, climate scenarios predict increasing 2006). temperatures and prolonged drought periods, resulting in increased Increased drought and xerothermic continentality in much of Europe conditions throughout Europe will (IPCC, 2007, 2014). High temperatures make wild fires a more widespread in combination with water stress threat and selective force than it is disturb physiological processes such currently. Forest trees exhibit two as nutrient uptake or photosynthesis basic responses to endure fire: they (Saxe et al., 2001). Physiological effects can either recover through a fire- of a single severe drought may last not triggered germination of a seed-bank just one growing season but can persist (e.g. low elevation Mediterranean over several years (Jonard et al., 2012). pines like Pinus halepensis/brutia) or In addition to decreasing tree growth they can survive and regenerate the (Cailleret and Davi, 2011), reproduction destroyed aboveground biomass may also be threatened by recurrent (Mediterranean evergreen broadleaved droughts. Summer droughts have species). Increased frequency of forest been demonstrated to decrease seed fires due to climate change can have crops and seedling recruitment in a detrimental effect on species that many tree species (e.g. Perez-Ramos do not have mechanisms to endure et al., 2010; Silva et al., 2012). Plants forest fires, such as mountain conifers react to drought stress by a variety of like Abies spp., Picea abies, Pinus nigra, heritable physiological and structural Pinus sylvestris or Pinus leucodermis, as mechanisms, such as changing they can severely reduce population morphology or anatomy of assimilatory sizes and consequently increase genetic organs or root systems (Ahuja et al., drift. Even fire-adapted low-elevation 2010; Kozlowski and Pallardy, 2002; Mediterranean forest trees can be at risk

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if the frequency of forest fires increases induce shallower levels of hardiness beyond the age of reproductive maturity. (Saxe et al., 2001) causing frost damage in the case of sudden frost events. Changed winter precipitation patterns The phase during which maximum represent another threat. Heritable cold hardiness is ultimately achieved features of tree architecture, such is induced by freezing temperatures as crown shape or branching form, (Weiser, 1970) and is associated with frequently result from evolutionary structural changes in cells. As growth adaptation to snow pressure and processes require a normal physiological occurrence of hoarfrost and ice (Geburek, state of cells, the whole process is Robitschek and Milasowszky, 2008). A reversed in the spring. Warm minimum shift of wet and heavy snow towards temperatures decrease hardiness after a higher altitudes may bring excessive few days, but low minima can slow or damage and thus important economic even reverse de-hardening, resulting losses. in fluctuating levels of frost hardiness (Leinonen, Repo and Hänninen, Elevated-temperature events during 1997). However, temperature-driven winter may induce winter desiccation in ontogenetic development towards bud­ conifers, when frozen soil does not allow burst is irreversible, and once bud­burst take up of water lost by transpiration. begins, the shoots cannot harden again This has reportedly caused damage in in response to low temperatures (Repo, the form of needle loss in Douglas fir Makela and Hänninen, 1990; Leinonen, (Larsen, 1981) and Norway spruce at Repo and Hänninen, 1997). Changed high latitudes (Kullman, 1996). As winter temperature patterns due to climate temperatures are expected to increase warming can thus lead to lower winter more than summer temperatures under hardiness and premature de-hardening, climate change, winter desiccation with the risk of frost damage to shoots associated with xylem cavitation in spring. and needle loss may thus decrease productivity in coniferous forests. Formation and growth of new shoots and vegetative organs and cessation Changed temperature patterns (mainly of growth are other cyclic sequences of the timing of seasonal temperature phenomena associated with the annual changes) in autumn and winter may cycle. In addition to photoperiod, they influence the frost hardening and de- are partly related to temperature during hardening processes. Frost hardening is a a year. Shoots and vegetative buds enter mechanism that prevents damage to cell a state of dormancy in the autumn. membranes due to ice crystal formation. Growth cessation is triggered primarily Warmer autumns and winters may by night length, but low minimum daily

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temperatures may induce earlier growth Reproduction, i.e. flower and seed cessation (Hänninen and Tanino, 2011). production, undergoes similar cyclic However, even when only photoperiod development triggered by climatic plays a role, entering dormancy in a time signals, physiological trade-offs and when temperature still allows further possibly predator escape strategies, growth would mean that trees do not a phenomenon known as masting fully exploit the growing season. (Herrera et al., 1998). Saxe et al. (2001) argued that trees can afford the risk of Dormancy means that cell division losing or damaging reproductive organs and growth have ceased and bud as these can be easily compensated for development is completed. Thereafter, by a heavy seed crop during a more buds require chill temperatures (along favourable year. This is true if such with night length) to become responsive losses or damages remain a rare event. to warm temperatures. Chilling Frost events can significantly reduce requirements differ among species and seed crops, but so can unfavourable genotypes within species: trees with climatic conditions during seed low chilling requirements flush earlier. maturation (for example high summer The process of cell growth and division temperatures), which are predicted to resulting in bud­burst is induced by increase under most climate change temperature sum accumulation. The scenarios. Moreover, a changing climate lack of chilling caused by climate can de-synchronize pollen production warming can cause abnormal bud­burst and female flower receptivity, leading in some species (Morin et al., 2009). to low effective density and poor seed Moreover, elevated air temperatures quantity and quality (e.g. increased during dormancy induction in late inbreeding in self-compatible species) summer increase depth of dormancy, (Alizoti, Kilimis and Gallios, 2010), meaning that more chilling is required particularly at range edges (Restoux et to break the dormancy and more al., 2008). With increasingly temporally degree-days need to be accumulated to distant suitable seed crops and proper induce budburst­ (Granhus, Floistad and recruitment conditions, the persistence Sogaard, 2009). Finally, leaf unfolding of local populations can be endangered patterns during bud break vary strongly in the long term. depending on life history traits such as evergreen vs. deciduous and shade tolerant vs. shade intolerant (Davi et al., What is ‘local’ and is the local forest 2011). The final effect of climate change reproductive material always the best? on the course of budburst­ phenology Environmental conditions vary is thus hard to predict (Hänninen and between sites and therefore the degree Tanino, 2011). of selection pressure also varies in

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different populations. This process more strongly in the insect pollinated leads to differentiation in adaptive Fraxinus ornus (FRAXIGEN, 2005). traits of tree populations between sites. Natural selection operates to remove The genetic diversity of a given FRM the least suited genotypes at a site and is the product of the original genetic this is used as a basis for the “native material that colonized the site on species and local provenances should which natural selection, drift and gene be preferred where appropriate” flow have acted. The relative influence principle (MCPFE, 1993). In sites with of evolutionary and demographic a long period of occupancy (multiple factors can be investigated using a generations), trees that are present combination of molecular markers currently are considered to have the and provenance trials to provide most optimal genotypes after having information on the geographical scale undergone several cycles of natural of local adaptation. This information is selection. However, Gould (1997) crucial for selecting and using FRM in argued that natural selection can only different sites and habitats. work on the material that is available. Therefore, the adult genotypes at a given site are only the best ones among The challenges of using the right forest those genotypes that were present at reproductive material under climate the site at the seedling stage, rather change than being the best possible option The current pattern of genetic variation for that site. In addition, it is difficult in forest trees and their adaptation to to define the exact size of a local climatic conditions has occurred over population in forest trees (such as for evolutionary time scales lasting from those trees that exchange genes during thousands to millions of years. The rate reproduction), particularly when of adaptive responses to environmental there are no strong environmental changes at population level depends gradients and in species where pollen on several factors, such as the level of is dispersed over long distances. genetic variation among inter-breeding Topography, population density, individuals, the size of the population, flowering intensity and phenology the heritability of fitness-related traits, and life history traits all influence the gene flow between populations, and size of the local population. In wind the intensity, direction and duration pollinated European ash species, for of the selection pressure. Many forest example, adaptation operates at very tree species are known, or believed, to broad scales of up to several hundreds have high genetic variation in important of kilometres, whereas geographical adaptive traits (Petit and Hampe, 2006). distances influence differentiationThey also have high fecundity, which

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creates a large gene pool for natural FRM will occur, with uncertain results selection to work on. However, the in terms of adaptation. The impact of unprecedented pace of anthropogenic this introduced diversity will depend climate change (Loarie et al., 2009), upon the scale of planting, relative to coupled with the comparatively long that of the local population, and the generation time of many forest trees, degree of genetic difference between the means that there may be insufficient populations. time for natural selection to give rise to populations adapted to new climatic Genetic adaptation will depend on the environments (Jump et al., 2006). Only amount of genetic variation present in when a tree population is initially natural populations, as it is unlikely that large, environmental changes are not new variation will be created via new too severe and evolutionary potential mutations. Heritability measures the is high, can it effectively survive by part of the variability of a phenotypic genetic adaptation (Gomulkiewicz and trait that is heritable in a population. Holt, 1995). The higher the heritability of a trait is, the more efficient selection is for a new, Concerns that there is insufficient more adapted, phenotype is. Heritability time left to ensure that adaptation of cold hardiness or flushing date is occurs before tree populations face low to moderate (Bower and Aitken, extinction have provided the impetus 2006; Rweyongeza, Yeh and Dhir, 2010). for intervention in the form of assisted Phenological traits frequently exhibit migration (Aubin et al., 2011; Hoegh- clinal variation along longitudinal, Guldberg et al., 2008), i.e. in the context latitudinal or altitudinal gradients, of this report, man-made transfer of as demonstrated by common-garden FRM to a non-autochthonous place. experiments with different provenances The rationale behind assisted migration (Dæhlen, Johnsen and Kohmann, 1995; is that the predicted future habitat at von Wuehlisch, Krusche and Muhs, the location to where the material is 1995). This gives good prospects for FRM translocated matches the current habitat recommendations related to phenology. of stands from whence the material is However, the responses to increasing taken. However, current and future temperature in the timing of various habitat similarity is judged based on processes associated with the annual climate, which is far from enough for cycle have been demonstrated to differ defining a suitable habitat. Also, the among species, i.e. the same change translocated FRM being significantly in temperature may accelerate a bud­ different from the local material, burst in one group of species, but delay complex genetic interactions between it in another (Hänninen and Tanino, the autochthonous and translocated 2011). The question is whether such

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differences in reaction also exist among (Saxe et al., 2001). It remains genotypes or provenances within a questionable, however, whether any species. Research has not sufficiently reliable predictions can be made about addressed this issue. Moreover, a part phenological behaviour of transferred of the heritable variation is epigenetic. material under climate change. In conifers, day length and temperature during zygotic embryogenesis and Several options are offered by FRM for seed maturation (associated with both mitigating climate change and adapting latitudinal and altitudinal differences) forests to climate change. However, there affect spring flushing, autumn growth is no general consensus on the best way cessation, height growth and frost to use FRM, either for favouring local hardiness (Johnsen et al., 2005; Skrøppa, adaptation or for assisted migrations. 1994). Ultimately, such epigenetic Options on how to use FRM will depend memory leads to rapid changes in on the objectives of management phenological behaviour of the offspring (production, protection, amenities), of transplanted provenances after one on the evolutionary past of the forest generation (Skrøppa et al., 2009). At (whether it is genetically diverse or not), present, no reliable mechanistic models on its position in the distribution range of phenological processes are available of the species (rear edge, core, or leading as there is much uncertainty about the edge) and within the ecological niche of background physiological processes species (marginal or central). Research, and external factors triggering them. particularly provenance research and The existing models accurately simulate molecular genetics and ecology, will past long-term phenology records, be foremost in helping design proper even if different models are based on strategies, and legal requirements and substantially different assumptions regulations for implementing them.

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14 L e g a l P o l i c i e s a n d F r a m e w o r k s

Legal and policy frameworks

Council Directive 1999/105/EC The objectives of this Directive are to: The legal framework in the European Union for the production and marketing • ensure free movement of FRM with- of forest reproductive material (FRM) in the EU; within the Union is based on Council • protect against the introduction Directive 1999/105/EC1 which applies and spread of organisms harmful to to all EU Member States. EU Directives plants in the EU; lay down certain end results that must • provide high quality and genetically be achieved in every Member State. suited FRM for the various site con- National authorities then have to revise ditions; and their national laws to meet these goals, • conserve forest biodiversity, includ- but are free to decide how to do this. The ing genetic diversity. requirements of Directive 1999/105/EC are organized in relation to four different The scope of the Directive is limited categories of FRM (‘source-identified’, to 47 species and artificial hybrids ‘selected’, ‘qualified’ and ‘tested’). important for forestry purposes (listed in Annex 1 of the Directive). The Directive The Directive sets up rules for the provides a common set of minimum approval of basic material (determined requirements, but allows the EU Member by each member state) for the production States to impose additional and more and marketing of FRM. It also prescribes stringent requirements for the approval a registration, labelling and control of basic material and production of system to allow the identification of FRM in their own territory, using their FRM from the collection phase to the national legislation. The exception is delivery to the end user. marketing – no other restrictions are

1 Available at http://ec.europa.eu/food/plant/propagation/forestry/forestry_leg_en.htm

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acceptable other than those provided National law on forest reproductive in the Directive. This provides special material – similarities and differences flexibility for Members States to regulate Council Directive 1999/105/EC has the production and marketing of FRM been implemented into national law by of the category ‘source-identified’ all 28 Member States. Supplementary concerning the approval of basic third states, such as Serbia, have adopted material and the marketing to the end- similar rules. Turkey has also adopted user. The Directive does not include the text of the Directive and applies requirements concerning the end use it strictly. All definitions concerning of FRM, but Member States are free to the type of material and categories determine requirements for this in their were introduced without changes into national legislation. national laws and regulations and are fully accepted by national authorities, forest owners and producers of FRM. OECD forest seed and plant scheme The term ‘production’ is adopted in The Council Directive on marketing a uniform way when seed processing of FRM is harmonized with the OECD and handling is considered. The Scheme for the Certification of Forest multiplication of parts of plants or in vitro Reproductive Material Moving in propagation is considered as production International Trade2. This voluntary of FRM only in some countries. scheme is open to all Members of the OECD as well as to any Member of In all Member States, regions of the United Nations. There are four provenance (also called seed zones) categories of reproductive material are delineated, based on a number under the scheme (‘source-identified’, of criteria, including ecological units ‘selected’, ‘qualified’ and ‘tested’). or vegetation zones, phenotypic or genetic similarities, or a combination. According to the decision of the The number and size of regions of European Union, forest seed and provenance vary greatly between planting stock of all four categories countries due to different approaches produced in six non-EU countries used for delineation, the complexity under the OECD scheme and officially of site conditions, the composition of certified by the national authorities shall forest ecosystems in a country, and the be considered equivalent to seed and economic importance of the species. planting stock complying with Council Aspects related to climate change are Directive 1999/105/EC. not yet taken into consideration.

2 Available at www.oecd.org/fr/tad/code/46092739.pdf

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Generally, the national legislations field inspections with different temporal follow the requirements for approval regularity and control of books and of seed stands from the Directive. In accompanying papers (master certificate nearly all Member States, an official and delivery papers). To facilitate inspection service checks the basic the control for cross-border transfer, material before approval. In states with a Commission recommendation was federal structures, such as Germany, adopted in February 2012 on guidelines Italy and Spain, this is done not at for the presentation of information the national but at the regional level. for the identification of lots of FRM Review of the approved basic material is and the information to be provided recommended at between 10 years (most on the suppliers label or document. countries) and 15 years (e.g. Italy). In This document contains harmonized some national laws (e.g. Germany), no identification codes for different revision period is indicated, but revision requirements, as listed in Article 13 of is being done continuously. Italy applies Council Directive 1999/105/EC, and special legislation concerning forest tree serves as numerical translations of the clones due to the economic value of text in the supplier’s documents or poplar clones in the country. labels3.

Generally, an FRM supplier needs an In many countries there are restrictions official licence for trading material for related to marketing of FRM, based on forestry purposes. In many countries the protection of valuable local genetic a register of suppliers is maintained. resources. The restriction to marketing The licensing and registration is done of FRM is based on results of ecological by official bodies, but professional and and genetic tests for certain species. In ability tests are not always requested. the case of ‘source identified’ it is based on quality criteria for approval of basic The master certificate is issued in all material of this category, which in these countries by a public authority, generally countries is subject to ‘more stringent the local forest service or the local office requirements’ regarding criteria for of the national body. Seed testing is also approval, or restriction of this category done by authorized laboratories under for certain species as a whole. public authority.

The official control is done with different intensities across countries. It includes

3 Available at http://ec.europa.eu/food/plant/propagation/forestry/forestry_leg_en.htm

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Table 1. Additional regulations, schemes and recommendations made by countries on forest reproductive material (FRM) (details are based on replies of member states to a questionnaire of the Working Group) Detailed specification Countries 1. Legal framework 1.1 Controls or limitations regarding the production & commercialization of FRM (basic material, collection, breeding, controls Limitations concerning list of species BE, FR, SI, IE Additional list of species HU, ES Limitations for category source identified BE, FR, DE, HU, LU, SI, CZ, IE, RS, IT,SK Ministerial authorization for import of non-EU material BG, FR; PL, HR, RS, SI (after professional opinion by SFI) Voluntary certification scheme including reference samples BG, FR, DE, UK, SI, NL (for additional non-directive species) Re-inspection of seed stands HU, PL 1.2 Requirements for maintaining minimal genetic diversity Number of individuals to collect from seed stands, varying by species and BE, CY, BG, HR, DK, DE, GR, IT, LU, category NO, RO, SI, SK, UK, CZ, PL Number of clones in seed orchards HR, EE, FI, FR, DE, HU, NO, SK, SE, SI, CZ, PL, IT, SI Minimum area to collect from DE, GR, HU, LT, SI, UK, CZ, PL, IT (for selected species) Distance between trees to collect from BG, IT, PL, CZ, SI Age of seed stands; number of mature trees to collect from DE, HU, CZ, PL, SK Designation of trees to collect from BE, SI Minimum number of genetically different trees NL Special rules for source identification UK, NL (specifically for autochthonous seed sources), IT (collection and use within the same region of provenance), SI (additional criteria for approval) 1.3 National or regional regulations for the use of FRM in the forest Rules, specific exceptions, limitations, administrative authorization for BG, CY, HR, EE, FR, LT, NO, SK, SI, transfer of FRM into and between eco-geographical regions or regions of SE, UK, RS, PL provenance Species adapted to site conditions BE, BG, FI, FR, GR, LT, SI Species restrictions in protected areas (Natura2000, National parks, etc.) BE, HU, SK, IE, HR, CZ, ES Provenances adapted to site conditions for important species BE, EE, FI, FR, GR, LT, NO, SI Limitations on exotic species BE, BG, CY, EE, FI, GR, HU, NO, PL, SK, SE, SI, ES (national legislation with list of invasive exotic species) Use of local FRM required GR, CZ (only in genetic conservation units), SI (recommended, but with a ranking of appropriateness) Limitation on clonal FRM SE, SI 1.4 National or regional regulations regarding subsidies linked to FRM Forest subsidies funding only if recommended FRM used CY, FR, DE, GR, IT, PL, SK, SI, ES, UK, IE, CZ, RS Forest regeneration funding linked to special conditions and/or use rules of FI, LT, LU, SI, RS, ES FRM, like natural hazards, use of local FRM, adaptation to site conditions Funding for production of FRM SI, FI, RS Funding for promotion of hardwoods SE, IE, RS, ES Funding according to positive list of FRM DK, ES 2. Guidelines and recommendations 2.1 Transfer of species or provenances (documents, web, etc.) BG, FI, FR, HU, LT, SK, SI, ES, SE, UK, CZ, IT, NO 2.2 Use of species by ecological regions (documents, web, etc.) BE, BG, DK, FR, HU, LT, LU, RO, SK, SI, ES, SE, UK, CZ, IT 2.3 Use of provenances by ecological regions (docs., web, etc.) BE, BG, DK, FR, DE, HU, LT, NO, ES, SE, UK, IE, CZ, IT, SI 2.4 Other guidelines and recommendations Number of trees to be collected FR, NO, DE, ES, SI Production of FRM: diversity, no. of trees, no. of ramets per clone ES, FI, DE, SI Diversity recommendations for state forests PL Use of FRM for close to nature forestry DK Quality requirements for marketing of FRM ES

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Recent developments in policy and legal frameworks

New traceability and certification systems Reproductive Material in Southern for forest reproductive material by means Germany (ZüF; http://www.zuef.de/) of genetic markers and the FfV Certification, supported by For official controls, proper documen- ISOGEN and the Association of Forest tation through master certificates and Seeds (ISOGEN; http://www.isogen. delivery papers is an important ele- de/) (Konnert and Hussendörfer, 2002; ment. The often long-distance transport Konnert and Behm, 2006; Konnert during production, from seed harvest and Hosius, 2008). Both systems aim to planting, facilitates mislabelling or to verify and trace the origin of FRM erroneous declaration of the origin of from harvesting and seed processing FRM. Therefore, methods to specifical- to the raising of seedlings in nurseries. ly test the origin of FRM, irrespective They are essentially based on reference of at which step the samples are taken samples and on the comparison of their (harvesting sites, seed lots, or reproduc- genetic composition using a species- tive material grown in nurseries), are specific array of different molecular clearly needed. DNA markers open up markers. The reference samples are possibilities for control techniques that collected at different stages of FRM are yet more precise and more efficient. production (harvest, cleaning of seed Such systems are used to verify the ori- and/or mixing of the seed lots, and plant gin of FRM, to check the number of trees delivery) and they are centrally stored. from which the seed was collected, to To increase the efficiency and to reduce identify tree species, hybrids or clones, costs, only a randomly selected subset and to check the parent-offspring rela- of samples is selected and analysed. The tionships. Control tools based on genet- molecular marker allele frequencies are ic tests are generally not foreseen in the used to assess the genetic similarity or legislation, but they can be implement- dissimilarity of the reference samples. ed under private law. Both systems are open for producers of FRM in all European countries. In Germany, two almost identical certification systems are currently in In Austria and Slovenia, reference place, i.e. the Certification Scheme samples are taken from every tree for Tracing the Origin of Forest harvested for seed and stored for

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prospective controls. In Slovenia, the from their Utilization (Access and Benefit regulation on confirmations and master Sharing – ABS). The aim of the protocol certificates for FRM and its modifications is – in very simple terms – to facilitate (UL RS 11/2003, 19/2004, 55/2012) access to genetic resources and to ensure demands that the responsible person of the fair distribution of benefits arising the producer collects a sample of living from their use by establishing a clear and tissue (e.g. a living branch, three wood transparent legally binding framework. cores including living cambium, cones, The “utilization of genetic resources” is conelets, beechnuts, etc.) from each tree defined in the Nagoya Protocol as “to used for seed production, delivers it to the conduct research and development on the authorized professional field officer, who genetic and/or biochemical composition then sends all material together with a of genetic resources, including through copy of the confirmation document to the the application of biotechnology” (CBD, Slovenian Forestry Institute (SFI). Once 2011). Therefore, the protocol does not the material is received, DNA is extracted apply to the use of genetic resources for and stored at -85°C until processing. production purposes, such as obtaining seeds for growing and planting seedlings A new approach based on DNA as part of normal forestry operations. fingerprinting can efficiently verify the The Nagoya Protocol requires that users, origin of seed sources without the above- including scientists, companies and mentioned use of multiple reference individuals utilizing genetic resources, samples (Degen, Holtken and Rogge, and traditional knowledge associated 2010). The authors used only a sample with these resources, from other countries of adult trees within oak seed stands and for research and development purposes, the control was directly made for each have to obtain in advance: suspicious plant or a group of suspicious plants by the use of multi-locus genotype • Permission from the country of or- assignment. However, this new method igin of the genetic resource and a is not yet in general practice. written endorsement, termed “prior informed consent” (PIC). • After obtaining the PIC, the user The Nagoya Protocol on Access and of the genetic resource has to ne- Benefit-Sharing gotiate conjointly on a bilateral In October 2010, the 10th Conference of level with the country of origin Parties to the Convention on Biological the terms and conditions of the ex- Diversity (CBD) adopted an international change (including a set of benefits agreement called the Nagoya Protocol on on “mutually agreed terms” – MAT). Access to Genetic Resources and the Fair and Equitable Sharing of Benefits Arising

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The protocol is intended to increase There is a consensus that the exchange legal certainty and transparency for of forest genetic resource (FGR) for the exchange of genetic resources research and development needs to be and related information. Rights and facilitated and intensified under climate obligations deriving from any existing change. However, at the moment it is not international agreement shall not be entirely clear how the implementation affected by the implementation of the of the Nagoya Protocol and the EU protocol. In October 2012, the European regulation ((EU) No 511/2014) will affect Commission presented a proposal for the exchange of forest genetic resources an EU ABS Regulation to implement the for research and development, including elements on compliance of the Nagoya between EU member states in order to Protocol for the European Union. maintain the free circulation of FRM on Subsequently, the European Parliament the EU internal market. Unfortunately, it and the Council adopted Regulation is expected to increase transaction costs ((EU) No 511/2014) on ABS on 16 April and create administrative hurdles before 2014. It entered into force on 9 June 2014 countries have managed to establish and applies from the date the Nagoya fully functional and transparent national Protocol itself entered into force (12 ABS regulatory systems (e.g. Koskela et October 2014). al., 2014).

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22 C u r r e n t G u i d e l i n e s a n d R e c o mm e n d a t i o n s

Current guidelines and recommendations related to forest reproductive material

Recommendations adopted by FOREST According to Helsinki Resolution EUROPE H1 (1993), native species and local The FOREST EUROPE process (formerly provenances should be preferred the Ministerial Conference on the where appropriate. The use of species, Protection of Forests in Europe – MCPFE) provenances, varieties or ecotypes has adopted several resolutions on forest outside their natural range should be genetic resources, including FRM. discouraged where their introduction would endanger important or valuable Strasbourg Resolution S2 (1990) set indigenous ecosystems, flora and fauna. up principles for the conservation Introduced species may be used when of forest genetic resources in Europe their potential negative impacts have and the cooperation between the been assessed and evaluated over countries in this field. It requested sufficient time, and where they provide the establishment of a functional but more benefits than do indigenous species voluntary instrument of international in terms of wood production and other cooperation to promote and coordinate: functions. Whenever introduced species (1) in situ and ex situ methods to conserve are used to replace local ecosystems, the genetic diversity of European forests; sufficient action should be taken at the (2) exchanges of reproductive materials; same time to conserve native flora and and (3) monitoring of progress in these fauna. fields. Subsequently, EUFORGEN was established in 1994. Resolution Helsinki Resolution H2 states that the S2 also recognized that the use of conservation of genetic resources of forest genetically-improved materials is of taxa should be recognized by countries as great importance for afforestation and an essential element of sustainable forest restocking, in particular where this is for management. Furthermore, Helsinki the purpose of the production of timber. Resolution H4 encouraged studies on Furthermore, Resolution S2 requested genetic variability of regionally important that countries keep records, at least for tree species in response to climate change public forests, of the exact identity of the and increased concentration of carbon reproduction materials used for planting dioxide, and on the degree and rate of and regeneration. evolutionary processes and adaptation.

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The General Declaration of the Lisbon European collaboration in this area. Conference (1998) preferred the use of Furthermore, it again urged countries native species and local provenances for to promote natural regeneration, and reforestation and afforestation. Annex 1 regeneration with native tree species and of Lisbon Resolution L2 introduced a provenances. quantitative indicator on changes in the proportions of stands managed for the In 2008, the Warsaw General Declaration conservation and utilization of forest recalled the maintenance, conservation, genetic resources (gene reserve forests, restoration and enhancement of the seed collection stands, etc.). According to biological diversity of forests, including the management guideline of Annex 2 of their genetic resources through Lisbon Resolution L2, adequate genetic, sustainable forest management. species and structural diversity should be encouraged and/or maintained to In 2011, the FOREST EUROPE process enhance stability, vitality and resistance decided to launch negotiations for a capacity of the forests to adverse Legally Binding Agreement on Forests environmental factors and to strengthen in Europe. An Intergovernmental natural regulation mechanisms. Negotiating Committee (INC) for this Furthermore, it stated that management agreement started its work in 2012 with plans should take account of endangered the aim to conclude the negotiations by or protected genetic in situ resources, 2013. The draft negotiated text, as it stands and that only those introduced species, after the resumed fourth negotiating provenances or varieties should be used session (Geneva, Switzerland, 7–8 whose impacts on the ecosystem and on November 2013), does not include any the genetic integrity of native species and direct reference to the use of FRM or the local provenances have been evaluated to conservation of forest genetic resources. avoid or minimize negative impacts. However, they are indirectly covered by the provisions of the draft agreement. One The Vienna General Declaration of the objectives of the draft agreement, (2003) recalls the importance of the for example, is to “to maintain, protect, maintenance, conservation, restoration restore and enhance forests, their health, and enhancement of biological diversity productivity, biodiversity, vitality and of forests, including their genetic resilience to threats and natural hazards, resources, both in Europe and on a global and their capacity to adapt to climate scale. According to Vienna Resolution change as well as their role in combating V4, countries should promote the desertification“. In October 2015, an conservation of forest genetic resources extraordinary Ministerial Conference as an integral part of sustainable forest will discuss those issues that could not management and continue the pan- be agreed by the INC.

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Regional activities – the Silva Mediterranea tries and included the Mediterranean example pines, firs, cedars and cypress as well Silva Mediterranea, which became cork oak (Besacier et al., 2011). The an FAO statutory body in 1948, is provenance trials have been invento- an international forum dedicated to ried recently for the establishment of Mediterranean forests. It incorporates a new database (Pichot, 2007; http:// countries from three continents w3.avignon.inra.fr/ForSilvaMed/), (Europe, northern Africa and Near developed by INRA-Avignon and Sil- East) that not only have very different va Mediterranea partners. This work legislation and environmental, social was partly carried out in the frame- and historical backgrounds, but also work of the TREEBREEDEX project different organizational structures (www.treebreedex.eu). and needs. Silva Mediterranea uses the OECD scheme as its main reference According to its work plan, the FGR for FRM certification, but some of Working Group of Silva Mediterra- the general criteria and guidelines nea will promote the development are supported and/or influenced and training of a new generation of by European Directive 1999/105/ researchers able to communicate and CE, especially in the Northern develop scientific and technical capaci- Mediterranean region. The FRM work ties to adapt FGR and forests to climate of Silva Mediterranea focuses on the change (Ducci, 2009; http://www.fao. creation of common criteria in order org/forestry/19318-0ebc0835b0ff- to facilitate international trade. For d94872e2249feaf1c10d6.pdf). For this example, Topak (1997) carried out an purpose, the Working Group prepared inventory of the basic material present recently a European Cooperation in in 17 Silva Mediterranea countries Science and Technology (COST) Action according to the OECD scheme. titled “Strengthening conservation: a Silva Mediterranea, and especially its key issue for adaptation of marginal/ Working Group on “Forest Genetic peripheral populations of forest tree Resources in the Mediterranean to climate change in Europe” (MaP/ Region”, promotes the use of existing FGR). This project was launched in knowledge and expertise through November 2012 and it involves several dissemination and training to match European countries in addition to the local needs. Silva Mediterranea partners, as well as international institutions, such as Bi- Silva Mediterranea established prov- oversity International, FAO, EFIMED enance trials in collaboration with and IUFRO. The project also aims at IUFRO in the 1970s. These trials were promoting the appropriate use of FRM located in nine Mediterranean coun- in the context of climate change.

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National provenance recommendations either all or only a sub-set of species in and support tools the national lists. As explained above, Council Directive 1999/105 only sets a framework to The form of publication of the recom- be subsequently addressed by the mendations is also different from country national legislation of the EU Member to country. They range from paper doc- States. Most national laws regulate only umentation to internet-based interactive production and commercialization, decision tools linking the planting site in but do not regulate the use of FRM. In an ecological zone with recommended order to secure the correct use of FRM, FRM available in a nursery. Most recom- some Member States have included in mendation systems are a combination of their national laws recommendations to documents and maps available on the forest owners on the use of FRM. In other internet. Member States, forest administrations make recommendations for the use In Germany, for example, the forest ad- of provenances in different regions. ministrations of the federal states pro- They mostly rely on the concept of vide forest owners with recommenda- provenance regions, which are areas tions for the use of provenances in differ- within which reproductive material can ent regions. They differ in the presenta- be transferred with little risk of being tion form between the states, but follow a poorly adapted to their new location. common consensus: the local provenance They are generally based on the results is preferentially recommended (e.g. for from provenance trials and the long- Bavaria see: http://www.stmelf.bayern. term experiences of practical forestry. de/wald/asp/014927/index.php). In general, climatic parameters are not given any preferential consideration. In France, recommendations for the use of French FRM are published under Provenance recommendations often http://agriculture.gouv.fr/conseils-du- have the role of a decision support, and tilisation-des-provenances-et-variet- forest owners are not obliged to follow es-forestieres. A revision of the French them. However, they can be binding recommendations is currently ongoing in in some countries under subsidiary the context of climate change. The seed schemes. Financial support for those tree company Vilmorin also gives rec- recommendations is crucial for success- ommendations for the utilization of seed ful implementation. European countries from Douglas fir seed orchards, based have reached different levels of imple- on site characteristics, budburst,­ growth mentation of these recommendations. performance (wood production) and ge- In many European countries, the prov- netic variability (see www.vilmorin-tree- enance recommendations are given for seeds.com).

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In Italy, several modes are used to provide alternative, material from selected stands guidelines and recommendations. These in northwestern­ Europe (Netherlands, include formal guidelines, technical north­west France, northern Germany papers issued based on the results of and Belgium) is indicated. Seed stock research programmes (http://www. from central and eastern continental Eu- ricercaforestale.it/), and networks of rope should not be planted, as it is poor- regional forest services and research ly adapted to the UK in terms of growth institutions coordinating relevant rate, survival, phenology and resistance activities. to foliar diseases. In some respects, the climatic dimension is taken into account In Slovenia, the Forest Act requires that by the statement that northward move- site-adapted FRM has to be used in ment of genetic material (by hundreds of any reforestation or afforestation with kilometres) may result in a gain in vigour planting and sowing. Accordingly, the compared with local-source provenanc- use of FRM in the same or in different es, whereas any southward movement is provenance regions and elevation considered ill advised. zones has recommendations defined by categories, from most appropriate, In Ireland, the Forest Service (Regulatory very appropriate, appropriate, less Authority) provides recommendations appropriate, appropriate only in for provenance choice based on results exceptional circumstances. The from an extensive series of provenance recommendations are part of the trials. Seed collected from registered regulation on delineation of provenance seed stands in Ireland should be used regions and its modification (Ur.l. RS when possible. A comprehensive table of 72/2003, 58/2012). Additionally, the acceptable seed origins is provided in the Slovenian Forestry Institute published guidelines, and for European species it is (in Slovenian) the EUFORGEN Technical required that the first choice is registered guidelines for conservation and use of Irish material. If this is unavailable, then forest genetic resources with Slovenian registered British (English or Welsh), recommendations for the use of FRM. French (north of Paris), Belgian, Dutch, Danish or German (north of Frankfurt) In the United Kingdom, the Forest- reproductive material from selected seed ry Commission publishes Information stands should be used. Notes, which provide recommendations for provenance selection for UK con- In Spain, the Ministry of Agriculture, ditions. The first choice of provenance Food and Environment, in collaboration should be a “selected” seed stand, pref- with the Spanish Forest Research Centre erably growing in the same region of (INIA-CIFOR), has published the regions provenance as the planting site. As an of provenances and recommendations

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for the use of FRM for those tree species planting purposes are made available. that are covered by the legislation The guidance process has three levels: at the national level. There are also the first level is a species recommenda- recommendations at the regional level, tion (including planting purpose), the such as those in Castilla y Leon, and second level consists of the recommenda- private forest owners should follow them tions for provenances and selected mate- if they want to receive grants from the rial (seed producing), and the third level public administration. is information on where to buy the plant material (i.e. availability of seed sources Some recommendation systems are in- in different nurseries). The system is a teractive decision tools, internet-based, web-based geographical interface, where linking plant sites in ecological zones users can locate their forest within 13 with recommended FRM available in ecological zones. The selection criteria nurseries. In Austria, an online platform are transparent and well documented. (www.herkunftsberatung.at) was created Information on possible subsidies is in- by the Research and Training Centre for cluded. The system also allows the user Forests, Natural Hazards and Landscape to provide comments in order to improve (BFW) in collaboration with the forest the system. administration to support forest owners in provenance selection. This online ser- Norway has an internet-based recom- vice is based on the national register of mendation system, which is hosted by approved seed sources, results of prove- the Norwegian Forest Seed Centre (www. nance trials in Austria and annually de- skogfroverket.no). Customers can choose clared seed collections. their planting location and elevation and will get suggestions for suitable seed lots, In Denmark, the Danish Nature Agency with differentiation between optimal and of the Ministry of Environment, in coop- usable (according to the internal Norwe- eration with the University of Copenha- gian transfer rules). After choosing a cer- gen, provides species and provenance tain seed lot, the customer will also get recommendations through a web-based information about which nursery is pro- tool that guides and helps users to se- ducing plants from that lot at that time. lect appropriate material suitable for the The suggestions for seed lots from stands planting purpose and locality (www. are made according to current legislation, plantevalg.dk). The system has two com- and no climate change effects are consid- ponents: a database where information ered. Recommendations for deployment about species and seed sources is avail- zones for improved orchard material are able, and a recommendation function made with a certain regard for possible through which guidelines for selection climatic changes. As the prediction of of species and seed sources for different future site conditions in Norway is very

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difficult because of the mountainous has an important role in maintaining landscape with steep elevation chang- resilience of forest ecosystems in the es within small distances, tree breeding face of threats associated with climate and the connected trials are considered change. A high level of genetic diversity a very important tool to explore and use in a population increases the probability the “genetic abilities” of the species in an that a proportion of the genotypes adaptation process to future climate. survive into the future by increasing the adaptability of that population. Sweden has a similar recommenda- Therefore, it is imperative that FRM with tion system, which is hosted by Skog- high levels of genetic diversity should be forsk (http://www.skogforsk.se/sv/ chosen to maintain genetic adaptability KunskapDirekt/Alla-Verktyg/Plant- at a particular site. This is particularly val/). Suggestions for suitable seed lots true at the rear edge of geographical are made with respect to future climate distributions, where local conditions scenarios. might already be ecologically extreme and where climate change effects are expected to be high. Conversely, FRM Scientific and practical considerations on the from these marginal populations can choice of provenances under climate change demonstrate local adaptation to xeric The success of tree planting efforts conditions and be a valuable source of depends fundamentally on the use FRM for reforestation under climate of appropriate FRM that is able to change (St. Clair and Howe, 2007; survive and thrive at the planting site. Mátyás, 1994; Robson et al., 2011). The decisions taken today on selecting suitable material for forest regeneration There are different approaches to must be made in the light of the climate preserve or increase genetic diversity projections for the next 30–200 years, and to prepare forests for the future. when the trees planted now will be One option could be to adopt a so called mature, even if this leads to stands “portfolio” approach and plant a mix that are adapted sub-optimally to the of provenances alongside the current current and future conditions. The use population, using climate change of appropriate FRM and the ability to predictions to guide the choice of verify the origin of FRM is therefore provenances for the future (e.g. Hubert of vital importance for our efforts to and Cottrell, 2007). A second, higher- prepare for and manage the effects of risk, approach would be to introduce climate change on European forests. FRM from a single provenance from a location with a climate similar to that Genetic diversity provides the raw predicted for the site. The assumption material for adaptation and it therefore is that the translocated provenance

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would contain genes more suitable for Island. In 1962, Haddock proposed survival in the future climate. A third seven seed collection zones for the entire option is to rely on natural regeneration British Columbia. This concept was later and harness the high genetic variability refined in the early 1970s, when 67 zones and phenotypic plasticity occurring for seed collection were delineated in all in most forest tree species, and use provinces (Ying and Yanchuk, 2006). forest management to create conditions The basis for the delineation was a where selection can occur on naturally preliminary approximation of ecological regenerated seedlings – the stage at classification of forest lands. A new which tree species experience the most revision, done in the early 1980s, also severe selection pressures. took into account results of provenance trial results from 24 seed zones. Simultaneously, for the first time, the Alternative approaches for seed “floating principle” of seed transfer was zone delineation and seed transfer introduced (Rehfeldt, 1983). It means that recommendations under climate change seed transfer limits are no more given Traditionally, the approach to seed by the fixed seed zone boundaries but transfer regulation on both legislative rather they are defined along adaptive and voluntary levels has been a clines along geographical distances. In geographical one, based on seed zones practice, this means that seedlings may or regions of provenances, which should be planted outside the seed zones in contain populations with identical or which the seed was collected, so long as similar adaptive features. This was the transfer is within the adaptive limits criticized by Mátyás (2007), for example, of the seed source based on a statistical who argued in favour of using climatic model (e.g. Wu and Ying, 2004). The criteria rather than geographical origin model defines the degree and direction as a basis for seed transfer. Moreover, of local optimality along longitude seed zones were often delineated based and elevation. Beside this scientific on environmental (mainly climatic) approach, the operational version homogeneity rather than genetic of seed transfer guidelines in British variation patterns or homogeneity of Columbia incorporates knowledge and adaptive responses demonstrated by experiences of local foresters to cover field trials. Therefore, new approaches specific situations across large forested to the delineation of seed zones or landscapes that cannot be predicted by regulation of seed transfer have models. appeared in recent years. The introduction of the climatic (eco- In Canada, the first seed collection zones logical) distance concept (Mátyás, 1994, were delineated in 1940 on Vancouver 1996) opened the possibility of applying

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climatic variables instead of geograph- based on genecological research using ical ones for transfer guidelines. With common-garden experiments (Johnson the improvement of statistical models et al., 2004; St. Clair and Howe, 2007; and software able to convert physical to St. Clair, Mandel and Vance-Borland, climate distance, Hutchinson (2004) and 2005). In nursery studies, dozens of Wang et al. (2006) included climate vari- adaptive traits and environmental ables in seed transfer rules. On the basis parameters were measured. To simplify of this, the seed transfer rules in British the analysis of the complex data sets, Columbia were re-examined and slight- these traits were combined into a few ly changed in light of climate change principle components that summarize (British Columbia Ministry of Forests, most of the variation. St. Clair, Mandel Lands and Natural Resource Opera- and Vance-Borland (2005) showed that tions, various dates). Now it is allowed two composite variables explain much to move most species in most areas 100 of the variation among individual traits to 200 m upwards in elevation. and among locations.

In the Pacific Northwest (Oregon and The first principle component, highly Washington States), seed zones were first related to minimum temperatures in established in the late 1960s, especially winter months and dates of last spring following requests from the expanding and first autumn frost, showed an east- tree improvement programmes for west cline associated with elevation and Douglas fir. A seed zone was defined by temperatures. Low temperatures appear a certain geographical area within which to have resulted in natural selection for seed can be collected and replanted, traits of earlier bud-set, presumably with the expectation that the resulting to avoid autumn frost, and faster stands will be adapted to the conditions emergence presumably to promote of the area. First seed zones were based seedling establishment as soon as on differences in climate and vegetation conditions are favourable in the spring. and they were subdivided by elevation (150 m or 500 ft) (Johnson et al., 2004). The second principle component was These zones were slightly revised in correlated with precipitation and 1973 (Tree Seed Zone Map, 1973). Based maximum temperatures in the summer on new results on adaptive traits from months, and shows a north-south cline. common-garden studies and provenance According to the authors, selection for trials, the tree seed zones were reviewed, earlier budburst­ could be a mechanism extended and made species-specific to ensure sufficient early growth before (Randall, 1996; Randall and Berrang, summer drought becomes a limiting 2002). The new recommendations for factor. The risk of maladaptation seed movement in this region are mainly was defined as the proportion of

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non-overlap between the normal under current climate as well as under distributions of the current population climate projection to 2080. Interestingly, and the population in the future populations showing the best (predicted based on climate scenarios) productivity indices originate from for the first composite variable. Based regions phylogenetically distinct from on this kind of studies, researchers can the core distribution area of Norway develop seed movement guidelines spruce, suggesting that population in the context of climate change. The history might explain part of the results of the above-mentioned study variation in climate response among suggest, for example, that Douglas fir populations. The authors suggested populations should be moved 450– that populations from currently warm 1130 m higher in elevation and 200– and drought-prone areas seem to be 540 km northward to match climates well adapted to the respective climate expected by the end of the 21st century. conditions, and may be appropriate Ukrainetz, O’Neill and Jaquish (2011) candidates for extended utilization in used this approach to develop both the future. seed zones and guidelines for “floating- principle” (focal-point) transfer for Enhancing the genetic potential of Picea glauca and P. engelmannii in British tree population to adapt to climate Columbia. They concluded that the change is also an option that needs to focal-point approach offers larger areas be considered. Savolainen et al. (2004) to which FRM can be transferred at a quantified genetic variation of growth given risk level of maladaptation. and hardiness traits within Scots pine populations in Finland. They In Austria, the outcomes of extensive found relatively high heritabilities of provenance experiments with Norway phenological traits and a strong genetic spruce established by K. Holzer were differentiation between northern and used to develop transfer guidelines southern populations (QST = 0.86). The for the Alpine region (Kapeller et al., authors used a simulation approach 2012). The authors used a population- to test whether Scots pine in northern response approach based on an Finland can change to the new predicted annual heat-moisture index, where optimum through migration and local the response functions were calculated selection during the next 100 years, both for individual populations assuming that the new phenotypic and clusters of populations based optimum can be deduced based on on the present provenance regions, the current match of temperature altitudinal belts and climatic data. sums and phenotypic means. The This approach allowed identification simulation showed that the adaptive of best-performing population groups response to climate change will lag

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behind the moving optimum. Although artificial regeneration with suitable seed no particular recommendations for sources can increase the proportion of human-mediated provenance transfer adapted genotypes. are given, the authors suggest that

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Scientific basis for Forest Reproductive Material transfer Guidelines

Methodological aspects of forest breeding zones) are generally based reproductive material transfer based on on climatic or bio­geo­graphical zoning provenance research rather than outcomes of provenance The molecular basis of adaptation has trials. The idea underlying this been the subject of intensive studies approach is that local adaptation is the during the last decade, using methods most important ecological force shaping of both forward and reverse genomics. genetic variation, and that climate and However, adaptive molecular markers photoperiod (i.e. factors associated are generally still under construction with the geographical location) are and have not been used for large- the main drivers of natural selection. scale mapping of genetic variation to There is, however, no unanimity about an extent comparable with neutral the reasonable size or number of such markers. Therefore, common-garden zones among countries. Neither there experiments, and especially provenance is a consensus about the principles and trials, remain the main source of methodologies on which they should information about adaptive processes be based. Sometimes empirical data and their outcomes in forest trees. provide no support for the existing zones (Isik, Keskin and McKeand, 2000). The main aim of the provenance research is the identification of Empirical information about the flourishing and sufficiently adapted behaviour of the transferred FRM in tree populations that can be used as terms of yield, growth, survival and seed sources for reforestation (König, tree quality has been used for a long 2005). Nevertheless, provenance trials time in decisions about appropriate also have other objectives, some of material for reforestation. They were them purely scientific, namely the based mainly on practical experience identification of the character (ecotypic (e.g. import of Slavonian oak to central or clinal) and main geographical trends Europe) or anecdotal information of variation of phenotypic traits. from provenance trials about the best-performing provenances. The The existing spatial frameworks for term ‘provenance’ is used here in a the recommendations on FRM transfer very narrow sense, denoting a local (regions of provenance, seed zones and population or a very small region (such

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as Polish spruce provenances Istebna or al., 1999). The response of a population Rycerka). However, a strong genotype- to transfer is expected to vary along the by-environment (G×E) interaction is ecological-distance gradient in a non- typical for forest trees, meaning that the linear manner. The maximum of the ‘best’ provenances need not necessarily response function determines the rate be the best everywhere. of transfer at which the performance of a population is the best. Various In modern provenance experiments, types of unimodal response functions numerous provenances have been have thus been used to model this repeatedly planted on a series of relationship (Gaussian, Weibull, beta, trial plots under a wide range of 2nd order polynomial). Typically, data environments. Such experimental set- from different test sites, even from ups allow testing of the reactions of different experiments, are pooled provenances to transfer: on the one together to develop a general transfer hand, determining the environment function, assuming that the shape and where a particular provenance peak locations of transfer curves do not performs best; on the other hand, vary significantly among environments choosing the optimum provenance for a (Rehfeldt et al., 1999). particular site. To derive climate-related responses of tree populations from Population response functions broadly designed provenance trials, describe the norm of reaction of an two primary approaches have been individual population to a range of test developed: general transfer functions, site environments. Methodology is very and population response functions similar to general transfer functions, (Aitken et al., 2008; Rehfeldt et al., 1999). but the provenances are not considered They can be used as an alternative jointly. Naturally, this approach basis for provenance recommendation requires that a provenance is repeatedly (Ukrainetz, O’Neill and Jaquish, 2011) planted over a representative number and may provide support for the of trial plots covering a sufficiently delineation of provenance regions. broad range of climates. Optimal transfer rates of populations may differ General transfer functions relate from each other, but they frequently fitness-related traits of the planted trees exhibit geographical continuity, and to geographical or climatic distances sometimes range-wide trends. They between provenances and common- can be thus used for the delineation of garden locations (i.e. the difference adaptively homogeneous areas, where of geographical or climatic variables populations exhibit similar reactions to between the site of origin and the site of climatic conditions. plantation (cf. Mátyás, 1994; Rehfeldt et

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The choice of proper climatic proxies is predicted the performance of seed the crucial issue in any approach based sources at unknown locations with on a mathematical modelling of tree- ordinary kriging based on provenance climate relationship. The reactions of tree data from tests sites, and generated populations on transfer need not follow seed-transfer guidelines by principal geographical gradients (longitude or component analysis of predicted latitude), and crude climate indicators, reaction norms. Hamann, Gylander such as mean annual temperatures and Chen (2011) applied multivariate or vegetation-season precipitations, regression trees to partition genetic do not necessarily describe properly variation, using a set of environmental those aspects of climate that are actual or geographical predictor variables drivers of adaptation. Various types of as partitioning criteria in a series of climatic indices have frequently been dichotomous splits of the genetic dataset, used for this purpose (cf. Mátyás and again based on provenance trial data. Yeatman 1992; Rehfeldt et al., 1999, St. Clair, Mandel and Vance-Borland 2002). The problem with these indices (2005) identified relationships between is their applicability in practice. No traits and environmental variables at forester would probably be willing to source locations by canonical correlation calculate, say, degree-days (>5°C)-to- analysis. Regression equations were annual precipitation ratio (Rehfeldt et then used to map genetic variation as a al., 2002) of the reforestation site and function of the environment, using GIS all potential FRM sources to decide tools. which FRM source is the proper one. However, such indices can be used in Inferences about natural populations geographical information system (GIS)- from provenance trials are associated based automated decision-support with several caveats. One applies to the systems incorporating geographical nature of what we call “provenance” distributions of climatic variables. and representativeness of provenance samples for the maternal population. Alternatively, transfer rules have been Seeds for establishing trials, especially generated by other statistical and when extensive international geostatistical approaches. Beaulieu, experiments are organized, may in some Perron and Bousquet (2004) and cases be drawn from commercial seed Campbell (1986) proposed relative risk lots. In that case, they might represent assessment, based on the extent of mis- a mixture of only a few open-pollinated match between the normal curves of families, sometimes collected from a the genotypic distributions of the local single stand, and typically represent and the transferred provenances at a the seed crop of a single year. Even plantation site. Hamann et al. (2000) when a seed crop is harvested from

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many mother trees, it may not properly vigour. It remains questionable as to represent the genetic structure of what extent inferences based on such the whole mother stand. In a year of measurements can be extrapolated a poor crop, only some of the trees to adult age, which is more relevant participate in fertilization. Moreover, from a practical point of view, unless genetic structures were shown to differ juvenile-mature correlations for the significantly also between crops of mast traits of interest were found to be strong years (Konnert and Behm, 2000). in previous experiments. Finally, the idea that provenance experiments may Another problem is associated with guide assisted migration as a measure the manner in which experiments for the mitigation of climate change are established. Typically, seedlings (Mátyás, 1996) relies on space-for-time are grown under optimal nursery substitution. We do not have enough conditions, and then planted in the field. knowledge about the validity of this Opportunities for natural selection at approach in this particular case. Even juvenile stages are largely circumvented taking into account all these limitations, by nursery practices. Early mortality is provenance experiments provide a huge modified by wide spacing, control of the resource for evolutionary ecology and competing vegetation, and protection climate change studies, and represent against large herbivores, rodents and the most reliable basis for practical pests. recommendations.

Older provenance trials usually did not include ecologically marginal sites and Case studies and transfer do not cover the whole breadth of a recommendations in selected species species’ distribution (Aitken et al., 2008). The following case studies are intended With increasing age, mortality cannot to illustrate the principles explained in be assessed because a part of trees need the previous sub-section, with examples to be removed by thinning, and the of those tree species for which enough number of standing trees may fall below knowledge has been gathered by the limit of representativeness. Most previous genetic studies and provenance studies are based on measurements at experiments at the range-wide, or at juvenile age. For instance, a thorough least regional, level. Consequently, the study of climate responses of Norway choice is biased towards widespread, spruce in Austria based on an extensive commercially important species, because experiment (389 provenances on 29 the research in the past primarily focused sites) was published by Kapeller et on this group of species. Although the al. (2012); however, height at 15 years selection of species is partly unbalanced was used here as a surrogate of growth from the point of view of life-history

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traits, it is at least balanced with respect motivated by the importance of Scots to distribution: boreal (Scots pine), pine as a valuable commercial forest temperate (European beech, silver fir) as tree species in large parts of its native well as Mediterranean species (Greek fir, distribution, which covers the whole Bornmüller’s fir, Nordman fir, Aleppo Euro-Siberian range, making Scots pine pine, Calabrian pine) are represented. one of the most widely distributed tree Douglas fir was also included as an species on earth. example of an introduced tree species that is widely used in European forestry. Towards the end of the 19th and the beginning of the 20th century, seed of forest trees was transferred across Scots pine (Pinus sylvestris) Europe without any regulation. Scots pine is a pioneer species that Substantial amounts of Scots pine seed spontaneously regenerates after major and cones were imported into Germany natural or human disturbances, if weed from Belgium, France, Austria, Hungary competition and grazing pressure and Russia (Lüdeman, 1961). In Sweden are low. The species grows mainly on and Livonia (present-day northern poorer, sandy soils, rocky outcrops and Latvia and southern Estonia), poor peat. The species is wind-pollinated and results were experienced with seed has both male and female flowers on the from central Europe, particularly seed same tree. Flowering is regular; female from ‘Pfälzer’ pine from southwest flowering starts at the age of 20–30 years. Germany, which since then has a bad Abundant male flowering appears some reputation (Langlet, 1938; Kurm, Meikar years later. Mast years are relatively and Tamm, 2003). This experience frequent but at the boreal forest limit underlined the importance of the origin seed maturation is impeded by the short of the reforestation material. For Scots growing season; mast years may occur pine, many provenance tests have as seldom as once or twice in 100 years been established (Giertych, 1991). The (Mátyás, Ackzell and Samuel, 2004). The results show a significant differentiation high migration potential of both pollen in growth among populations with and seed results in effective gene flow respect to the long-distance transfer of within the contiguous range, causing a FRM. In general, transfer of northern distinct, clinal pattern of variation within populations from 57–67°N southwards the species, at least for adaptive traits. results in reduced height growth and This is typically the case with growth stem volume (Giertych, 1979; Oleksyn and phenology characters, which are and Giertych, 1984). At the same time, determined primarily by temperature. populations originating from the The selection of Scots pine as one of central part of the species’ range in the model species for case studies was Europe grow as well or better than local

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provenances at northern latitudes. This The glacial history and Holocene indicates that the Central European migration of beech differ from most populations of Scots pine have a greater European widespread tree species. growth potential than the northern Although there were several glacial European populations. Statistical refugia in southern Europe, many modelling approaches have revealed refugial populations have not expanded, that climate change may lead to short- or have colonized only restricted areas, term plastic responses in contemporary because the spread was blocked by populations of Scots pine, including lowlands with big rivers. The major large losses in growth and productivity part of the present-day range of beech in the central and southern parts of (except Italy and southern Balkans) was eastern Europe, modest losses in growth colonized from a single source, located and productivity along the southern in Slovenia and Istria (Magri et al., periphery of the species’ distribution 2006). This explains the relatively low in Asia, and gains in productivity in differentiation for neutral nuclear loci the northern and eastern part of the generally observed in European beech. distribution range (Rehfeldt et al., The initial gene pools of beech subjected 2002). Results indicate that provenance to natural selection were relatively experiments can be efficiently used to homogeneous, except a gradual decline predict growth response reactions of of allelic richness towards the range FRM transferred to new environmental margins caused by recurrent founder conditions. effects during the Holocene migration (Comps et al., 2001), which was only partially compensated for by gene flow. European beech (Fagus sylvatica) Therefore, clinal patterns for several European beech is an example of phenotypic traits (especially phenology widespread broadleaved species with and cold resistance) observed in more or less continuous distribution common-garden experiments are in the centre of the range, although unlikely to result from isolation by small and isolated populations occur distance and can be attributed to at outer or inner range margins. As a adaptation. wind-pollinated species, it is expected to be predominantly outcrossing and to In the early 1990s, a large-scale exhibit high levels of gene flow, which international beech provenance has partly been confirmed by genetic experiment was established, containing studies (Merzeau et al., 1994). In spite 202 seed sources planted in two series of this, within-stand spatial genetic of plantations (field trials), comprising structures indicate strong isolation-by- 23 and 24 plantations, respectively. distance patterns (Chybicki et al., 2009). These trials are located in 21 European

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countries. The Apennine and Balkan functions. He found clear geographical peninsulas are strongly under- patterns for almost all combinations of represented in the collection of the response and the underlying climatic tested material in this experiment. and geographical variables. In general, correlations between optimum transfer During 2006–2010, an integrated rates and the underlying variables evaluation of the beech provenance were strong and significantly negative, experiment was accomplished within whereas optimum environments were a COST action. In general, trial sites nearly the same for all provenances. proved to have the most important This indicates that the extent of local effect on survival and growth (the adaptation is rather limited, probably respective variance components ranged due to big phenotypic plasticity of beech. from 30% to 70% of the total variance). The effect of seed source origin was Mátyás et al. (2011) found that the growth considerably smaller, amounting to only performance of beech populations at the 0% to 7%. At the same time, a significant xeric limit of the distribution generally provenance × trial-site interaction was deteriorates with transfer into more observed for most growth traits, with continental climate (as quantified by variance component of 4% to 21% (Alía the Ellenberg index). At the provenance et al., 2011). When averaged over the level, there is marked differentiation. sites, no clear geographical trend in the Maritime provenances show improving performance of individual provenances performance with warming and was identified for survival, and only drier conditions, while continental a weak correlation with latitude and populations display the opposite. The climate parameters of the site of origin results propose a relationship between was observed for height growth. There the climate of origin and the character of was also no clear spatial pattern of the response to changing climate. stability of the provenances (Finlay- Wilkinson joint regression model). Spring and autumn vegetative phenology Concerning height growth, it seems that are generally considered important there are populations that perform well fitness-related traits. A comprehensive over a broad range of environments. evaluation of leaf flushing done by Robson et al. (2011) confirmed that heat In spite of significant G×E interactions, accumulation during the winter and which obscure overall spatial trends spring (degree-hours >5°C) is the most in growth and survival, responses of important determinant of the timing provenance to transfer show quite clear of budburst,­ but photoperiod,­ chilling geographical patterns. Gömöry (2010) period and summer drought may play analysed provenance-level response a role as well. The total span of average

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flushing times of provenances was 11 Silver fir (Abies alba) to 12 days almost independently of Silver fir is an example of a conifer planting site. Irrespective of altitude, species that has suffered serious there is a general trend for provenances range reductions over the last from the southeast of Europe in the centuries, mainly through human Mediterranean and warm-continental influence (silvicultural preferences regions to flush early in comparison for monocultures of Norway spruce). with late-flushing provenances from Its natural range extends from 52°N the north and west of Europe, where (northern Germany) to 38°N (southern Oceanic influences on the climate are Italy) and from 22°E (eastern Romania) strong. to 03°W (western Pyrenees) (Cremer, 2009). Its distribution is characterized Gömöry and Paule (2011) showed by a central core in central and eastern that late-frost damage is almost Europe, and a fragmented marginal entirely determined by the bud­burst range with isolated populations. Silver date. Strong correlations between fir is a monoecious and wind-pollinated height/diameter growth and length conifer species and its seed are dispersed of vegetation season indicate that the mainly through wind. timing of leaf flushing and shedding results from an evolutionary trade-off In the mixed mountain forests of central between effective use of the vegetation Europe, silver fir exhibits a specific for photosynthesis and the avoidance role as a stabilizing element due to its of frost damage. If late frosts become deep seated root system and its ability frequent under climate change, transfer to regenerate and survive long periods of reproductive material from the under shade (Cremer, 2009; Schütt, West to the East may become a viable 1994). In the climate change context, mitigation option. Robson et al. (2011) silver fir is promoted by management suggested considering provenances plans in many parts of its natural from the southwest of Europe for distribution range as it is seen as a low future use under climate change, as risk alternative to the more vulnerable they are adapted to Mediterranean Norway spruce (Picea abies) (Brosinger, environments and yet flush relatively 2011). late, and thus may be candidates to withstand climate change without From numerous genetic studies, it is being susceptible to late frost. However, known that as a result of migration history these provenances are typically less after the last ice age, human influence on productive. forest ecosystems and natural selection processes, silver fir shows a high level of genetic differentiation at both a

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macro- and a micro-geographical scale (especially from Calabria) and a lower (Konnert and Bergmann, 1995; Konnert, resistance to more continental climate 1993; Vendramin et al., 1999; Liepelt conditions, compared with provenances et al., 2008). High genetic variation from the Alpine region. The stronger was found in Calabria (Bergmann, “oceanic” disposition of the Calabrian Gregorius and Larsen, 1990; Vicario et provenance Serra S. Bruno was also al., 1995; Camerano et al., 2012), whereas confirmed by a two-site test carried out populations at the northern edge of in Denmark (Hansen and Larsen, 2004) the natural distribution in northern on 13 provenances from southern Italy Bavaria, Thuringia and Saxonia show (Calabria), central Italy, Germany (Black very low genetic diversity (Konnert, Forest), Romania (Carpathians) and 1993; Llamas, 1994). In this area, silver Denmark. Height after 15 years, bud­ fir generally is of bad quality and shows burst and mortality varied significantly decline symptoms. Seed harvests in among provenances. In particular, in the these stands have, without exception, field trial that was well protected against a high proportion of empty seeds, late frost in the spring and early frost in indicating genetic depression. the fall by a shelter wood of larch, a high variance in height growth was observed. The large genetic differences are illustrated also by a variation in adaptive Between 1986 and 1989, a silver fir traits (Sagnard, Barberot and Fady, 2002) provenance trial was established in and growth behaviour in provenance southern Germany (Ruetz, Franke and trials (e.g. Wolf, 1994; Commarmot, 1995; Stimm, 1998). A total of 21 provenances Ruetz, Franke and Stimm, 1998; Ruetz, covering the entire natural distribution 2002). Even provenances originating range were planted in seven sites. from the same micro-geographical After 20 years, provenances from region (e.g. Calabria) differ significantly the Carpathians in Romania and in height growth and budburst­ date Slovakia showed the best growth (Hansen and Larsen, 2004). and the lowest mortality. Among the central European provenances, those The first provenance trial for silver from the Vosges Mountains in France fir was installed in 1905 by Engler in and from southwest Germany (Black Switzerland (see Pavari, 1951). In 1924, Forest), performed particularly well. Italy and France started an international Below-average growth was detected trial with 20 provenances from the in provenances from western France, Pyrenees, the Alps and the Apennines Italy, Macedonia and populations from (Pavari, 1951). Results showed a the northern border of the natural higher survival rate and resistance to distribution range in northeast Bavaria. drought effects in southern provenances Populations from Calabria and Serbia

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had high mortalities (Ruetz, 2002). Based species in introgression areas, such on results from genetic inventories as A. borisii-regis (Panetsos, 1990) or and provenance trials, silver fir from geographical variants, e.g. between the Carpathian region, especially from Abies bornmuelleriana and A. equi- Slovakia and Romania, is recommended trojani (Ata, 1990). Similar to Abies for planting to improve the situation of alba, the genetic differentiation of silver fir in southeast Germany. A prior Mediterranean firs generally exhibits condition is that material with provable a geographical pattern in both neutral identity is available (verification of the markers and phenotypic traits (Fady origin by means of genetic methods and Conkle, 1993; Parducci et al., 2001; is possible, and done at random). This Ducci et al., 2004). recommendation is based on practical reasons and it does not include any The distribution range of some considerations of climate change. Mediterranean firs is very fragmented and often reduced to single, scattered and isolated populations, which are Mediterranean firs characterized by a very small effective Eleven fir species, gathered under the size. Abies nebrodensis, A. pinsapo and term Mediterranean firs, occur in the A. numidica are the most endangered Mediterranean Region. They grow at species with regard to climate change, higher elevations, between 800 m and with an expected upward shift of 2800 m, depending on the latitude, altitudinal zones. from the Pyrenees, to the Alps, the Apennines, Sicily and the Balkans In the 1970s, several international (Abies alba Mill., Abies nebrodensis trials with Mediterranean firs were (Lojac.) Mattei, Abies borisii-regis set-up within the IUFRO network. Mattf., Abies cephalonica Loud.), to One of the first tests was established the Turkish Pontus and Near East with the participation of France, Italy (Abies bornmuelleriana Mattf., Abies and Greece, containing provenances of cilicica (Ant. et Kotschy) Carr., Abies Abies bornmuelleriana, A. equi-trojani, A. equi-trojani Aschers. et Sint.) Mattf.), nordmanniana and A. alba. At the same and to the Caucasus range (Abies time, FAO Silva Mediterranea expanded nordmanniana (Stev.) Spach.). The the trial with provenances of Abies western group of species extends to cephalonica, which varied considerably the Iberian peninsula (Abies pinsapo in survival and growth. Boiss) and to the Atlas mountains in Northern Africa (Abies marocana In Italy and Greece (Panetsos, 1990; Trabut., Abies numi­dica Carr.). These Ducci et al., 2011), A. cephalonica species are able to produce hybrid populations from Mainalon (Veti and

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Kapo), Parnassos (Koro and Mevr) and across the Mediterranean basin. The Cephalonia Island (Ceph) performed ability of both species to grow in the best, although the last-named was adverse climatic conditions of the sensitive to late spring frost. In southern Mediterranean region, combined France, the Greek provenances Mainalon with their fast growth on favourable and Parnon yielded the best results sites, make them very important and were therefore recommended for multipurpose species for forestry, reforestation (Fady, 1993). Furthermore, while their ability to endure forest fires A. cilicica has an interesting potential through specific mechanisms, render within the same bioclimatic range them irreplaceable for the special and as A. cephalonica. Even though A. sometimes delicate Mediterranean bornmuelleriana and A. equi-trojani are ecosystems (Aravanopoulos et al., of interest for southern France, their 2004; Alizoti, Bailian & Panetsos, 2004; cultivation should be restricted to acidic, Chambel, et al., 2013). high elevation and high rainfall sites. Provenance testing was launched in the Among the firs of the Pontus region, early 1970s in most of the Mediterranean A. bornmuelleriana proved to have countries, following international great vigour even in relatively arid initiatives coordinated by FAO and environments. A. bornmuelleriana IUFRO that resulted in the collection provenances from Kangal and Arag and of seed from 50 provenances (33 of the A. equi-trojani provenance Kazdag P. halepensis and 17 of P. brutia) and the showed good performances for both establishment of numerous provenance height and diameter growth rates, trials of the two species across the whole whereas provenances of A. nordmanniana Mediterranean basin. Later on, other presented the worst results. Due to international collaborative initiatives the popularity of A. nordmanniana exploited the already existing network and A. cephalonica as Christmas trees, and established also new trials for the reproductive material of these species is two species (FAO Silva Mediterranea, spread throughout Europe. IUFRO and EU projects). P. halepensis provenance trials have been established in Greece, Italy, France, Spain, Israel, Mediterranean pines of the halepensis group Morocco and Tunisia, while P. brutia (Pinus halepensis and Pinus brutia ) trials are mainly located in Turkey, Pinus halepensis and P. brutia are among Greece, France and Italy. The purpose the 11 species of pines growing in the of the provenance trials is not only the Mediterranean bioclimatic region. They evaluation of survival and growth of are pioneer, prominent, low-elevation the two species, but also their adaptive conifers with extensive distribution potential across different and sometimes

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limiting environmental conditions been reported by Climent et al. (2008), (Chambel et al., 2013). indicating that the eastern and northern populations are less precocious for Results of the above established cone bearing than southern Iberian and networks of trials indicate the existence North African provenances. Differences of high plasticity for growth traits for growth and adaptive traits have and significant G×E (genotype by been also revealed for P. brutia environment) interaction across sites. populations growing at different The relative ranking of provenances altitudinal gradients (Kaya and Isik, tends to remain rather stable, indicating 1997; Pichot and Vauthier, 2007). Eco­ that the G×E interaction is rather physio­logical studies carried out on quantitative and not qualitative P. halepensis provenances in harsh, (Chambel et al., 2013). G×E interaction near-desert, environmental conditions, for traits like stem form and frost showed the existence of significant resistance was lower than that for inter- and intra-provenance variation growth traits, both at a species and and indicated that populations at a provenance level. Results also exhibiting higher heterozygosity, indicate the existence of ample adaptive higher drought resistance and water- genetic diversity for both species, use efficiency survived and performed with P. halepensis provenances being better (Schiller and Atzmon, 2009; Klein in general more water-stress tolerant, et al., 2013). Regarding neutral genetic while P. brutia provenances being less diversity, high levels of differentiation susceptible to frost damage (Calamassi, has been recorded among populations, Falusi and Tocci, 1980; Falusi, Calamassi but rather low within populations, and Tocci, 1983; Calamassi, Falusi and with a tendency to increase from west Tocci, 1984; Grunwald and Schiller, to east. It has been suggested that the 1988; Atzmon, Moshe and Schiller, 2004; above-mentioned genetic patterns may Voltas et al., 2008). P. brutia has also have emerged due to a combination of been proven to be more resistant to the factors, such as the Pleistocene climate, insect Pissodes notatus (Ducci and Guidi, the species’ migration strategies, and 1998). Pinus halepensis exhibits a well- impact of wild fires (Fady, Semerci established north-east – south-west and Vendramin, 2003; Fady, 2012). cline for adaptive traits, with the north- Climate change is expected to result eastern provenances showing higher in temperature increase and more early growth rates and better stem forms pronounced and frequent droughts in than the southern Iberian and North the Mediterranean basin. In northern African provenances (Chambel et al., Mediterranean regions, though, 2013). Similar geographical trends for increase of growth or expansion of the early reproductive allocation have also distribution in the face of climate change

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has been postulated for P. halepensis Given all the above, it is recommended (Rathgeber et al., 2000; Thuiller, 2003). that P. halepensis FRM produced from At the edge of the distributions of both national provenances should be used, species a notable growth decline has based on the results obtained from the been recorded via dendrochronological networks of the existing experimental methods, i.e. for P. halepensis in the very trials. Genetic material proved to dry coastal areas of south-eastern Spain be tolerant to extreme, near-desert, (De Luis et al., 2007) and for P. brutia in conditions could be considered for some of the Aegean Sea Greek Islands planting in sites with potentially high (Körner, Sarris and Christodoulakis, temperature and drought indices. 2005; Sarris, Christodoulakis and Transfer and use of P. brutia FRM should Koerner, 2007). Frost damage can be be based on the species requirements another risk that primarily P. halepensis and adaptation limits, as well as on might encounter in mountainous or results obtained from the field trials. continental areas due to climate change, given the lack of hardiness of the species to cold temperatures and the expected Douglas fir (Pseudotsuga menziesii) abrupt temperature changes due to Douglas fir has one of the widest global warming (Fernández et al., 2003; natural ranges of any tree species and Climent et al., 2009). the largest south-to-north distribution of any commercially used conifer in Transfer of P. brutia FRM from Turkey North America, extending from 19°N is currently approved in several in Mexico to 55°N in western Canada. countries, i.e. Italy, France and Spain. Within this large geographical area Specifically in Italy, the material with strongly contrasting climatic imported is the one that exhibited conditions, Douglas fir occupies many the best survival and performance habitats. Populations are generally in the FAO/Silva Mediterranea field regarded as being closely adapted to trials. P. halepensis FRM is produced different environments. Two types or across the Mediterranean countries by varieties are recognized: the typical genetic material sampled from national coast or green variety (var. menziesii provenance regions and seed stands. or viridis) extends from Vancouver Use of introduced and maladapted Island and the coastal mountains of P. halepensis reproductive material (i.e. British Columbia along the Pacific Algerian and Italian provenances) for slope into California, and has a nearly afforestation purposes in south France continuous distribution from sea level resulted in high mortality due to frost to an elevation of around 1800 m. On the damage and cold winter temperatures eastern slope of the Cascades and in the (Tabeaud and Simon, 1993; Bedel, 1986). Rocky Mountains from northern British

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Columbia into the southwestern United and Bastien, 1992). In 1990, researchers States of America and Mexico, botanists from INRA-Orleans presented range- recognize the glauca variety, known also wide results based on data collected as “interior” or “blue” Douglas fir. This from 108 trials and from 20 institutions variety differs from menziesii in foliage in 15 countries (Breidenstein, Bastien colour, cone form, growth rate and and Roman-Amat, 1990). The results environmental requirements. It has a of this study demonstrate a better patchier distribution, especially in the adaptation and growth of coastal south and a wider elevation range, of provenances (var. menziesii) on nearly 500 to 3500 m. This variety mingles with all European sites. In central Europe, the coastal variety in southern British the best performing provenances Columbia and northeastern Washington originated from Washington State (Hermann, 1985). (below 600 masl), whereas provenances from northern Oregon grew fast only Douglas fir (Pseudotsuga menziesii) has on sites with a mild climate. In general, been grown in Europe for over 120 mortality was low to moderate on nearly years and it is one of the most important all sites (15–30%). Higher mortality was introduced species in many European observed on provenances from British countries. Beside site conditions and Columbia, whereas the Washington silvicultural treatment, the provenance State provenances showed the lowest is crucial for a successful introduction mortality. In nearly all tests, provenances of this non-native species. Until the of the glauca variety (interior Douglas late 1960s, in the majority of import fir) exhibited excellent survival in the cases, nothing was known about the first years (Kleinschmit et al., 1974, 1991) introduced provenances. In the mid- but poor growth performance in later 1960s, seed zones for Douglas fir were stages. established for the Pacific Northwest. At the same time (1967–1968), an In the Mediterranean region, and IUFRO provenance trial was started especially in Italy, several tens of with provenances from Oregon and thousands hectares of planted Douglas Washington State in the United States fir are growing in mountain ranges of America and from British Columbia with very good performances and in Canada. More than 120 provenances good technical characteristics. In this were planted in more than 100 field tests area, more than 120 provenances from in 33 countries all around the world, British Columbia to California, New but mainly in Europe. This is one of the Mexico and the interior were compared largest international provenance trials with provenances introduced earlier to installed for forest trees. Parts of the Italy. The results indicate that for Italy, numerous trials still exist (Kleinschmit the most appropriate provenances are

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from the Coastal Range in Oregon and The IUFRO provenance collection in California (higher elevations). All stimulated numerous other seed those provenances showed in general collections for Douglas fir in the 1970s. a low G×E interaction, very good For example, to refine provenance performance, high survival rate, low recommendations in southern percentage of forked trees, good stem Germany (Bavaria) more than twenty form and low number of branches. Also supplementary field tests were installed some Italian landraces ranked among between 1971 and 1976. These include the best, particularly the Calabrian provenances from which promising provenance Mercurella and the northern results are expected, including an Tuscany seed stand Acquerino (Ducci et altitudinal sampling up to 1000 masl in al., 2003). In Spain, the best materials some areas (Ruetz, 1987). As a result of came from north Oregon and south these trials, new guidelines for Douglas Washington from latitudes north of fir seed import and use were developed 45°N (Zas et al., 2003). based on seed zones and elevation (Ruetz, 1987). These guidelines also became part As a conclusion of these tests, of the provenance recommendations in recommendations for selecting Bavaria. For regions without summer provenances for different planting drought, provenances from the west sites throughout Europe are given. In slope of the Cascades in Washington central and eastern Europe, the choice from low and middle elevations are of Douglas fir reproductive material recommended. For the mildest regions is restricted to the middle-elevation of Bavaria and on dry sites, provenances zone of the Cascade range in northern from north Oregon and from the Washington State, as provenances from east slope of the Olympic Peninsula high latitude and elevation have slow in Washington State are preferred. growing rates. Only at sites with a harsh Provenances from the Washington coast climate, Douglas fir provenances from ranges have generally performed well high elevations could be interesting. In and have had little spring frost damage. southern and Mediterranean Europe, Thus they are recommended especially provenances from low elevations for sites susceptible to late spring frosts. in northern Oregon and southern Washington State seem to be of high interest.

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50 C h a l l e n g e s a n d O pp o r tu n i t i e s

Challenges and opportunities

Potential of tree breeding to meet for genetic improvement. Several conservation needs and climate change current breeding programmes include challenges drought tolerance (Butcher, 2007; Dalla- Tree breeding activities differ in their Salda et al., 2009), resistance to pests and intensity, focus and impact on forestry pathogens (whose incidence is expected among different regions of Europe. to increase under climate change), Intensive breeding programmes have vegetative and flowering phenology generally focused on a few forest tree or physiological characters as target species. Typically, breeding is organized traits. Recent development in forest as a recurrent cycle of selection, testing tree genomics, especially progressing and production. Intensive breeding may discovery of biologically relevant SNPs lead to the loss of diversity, but it also by means of association genetics may provides the opportunity to increase considerably accelerate progress in this diversity substantially by a proper respect (Hamanishi and Campbell, 2011; management of the genetic resources Neale and Kremer, 2011). Research is used. being conducted to identify candidate genes related to traits that will respond Breeding is one tool with which to to climate change (bud­burst, bud set, face climate change. The control of the winter hardiness, drought resistance genetic factor through tree improvement and water-use efficiency). However, is expected to produce trees that are the achievements obtained so far are better adapted to certain environmental not yet at the stage of introducing these conditions. As the combined effect of genes (by crossing or other means) into increasing temperatures and decreasing commercial varieties. precipitation, i.e. drought stress, is considered the major problem associated Phenotypic plasticity, the ability of a with climate change, planting of drought- genotype to produce different pheno- resistant trees may represent a mitigation types under different environments, option. Drought tolerance is a trait with may be considered a trait of its own, moderate heritability (Cumbie et al., 2011; which is heritable and may thus be sub- Newton et al., 1991) so there is potential ject to selection. Phenotypic plasticity

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is favoured by evolution in spatially Definitions for the stability of and temporally heterogeneous envi- phenotypic expression are nearly ronments. Phenotypic change may vary as numerous as their applications. not only in the amount and pattern, but In general, measures of phenotypic also in the speed of expression, revers- stability attempt to quantify a ibility, and ability to occur at different genotype’s tendency to exhibit developmental stages (Schlichting and constant phenotypic expression in Smith, 2002). Both spatial and tempo- different environments (Lynch and ral heterogeneity apply to forest trees, Walsh, 1998). In forestry, the patterns which form large populations cover- by which different FRMs (e.g. seed ing typically plenty of climatically and lots from selected seed stands, seeds edaphically differentiated micro­sites. from tested families in seed orchards As long-living organisms, trees are also or clones) respond to environmental exposed to environmental changes, conditions are primary determinants even during their lifespan. The term has of the FRM’s desirability in different been widely applied in forest genetics environmental conditions. Stability of literature, not always according to the FRM performance across a broad range original definition, which was the envi- of environmental conditions is usually ronmentally sensitive production of al- essential from breeders’ points of ternative phenotypes. The interpretation view. Removing unstable families is an of the term was increasingly becoming alternative to reduce the impact of G×E liberal and tree breeders often used the interaction in breeding populations term for expressing the breadth of site (Mata, Voltas and Zas, 2012). Many condition variation, successfully coped authors have suggested considering with by the population or genotype. It the genotypic stability across sites as is therefore advisable to use instead the a screening trait in selection processes term phenotypic stability to express the (Johnson, 1998). Selection based on ability to maintain fitness (growth su- stability parameters is also a safeguard periority) across a wide range of sites, decision regarding the current global often surpassing locally adapted popu- change scenarios. Selection for lations. Phenotypic stability is expressed specific adaptation at present may by reaction norms or transfer functions lead more easily to easing future calculated from comparative tests, such adaptation concerns in the deployed as provenance trials. The trait is genet- material (Ledig and Kitzmiller, 1992). ically determined and becomes increas- Removing the most interactive families ingly important in climatically changing indeed reduced G×E interaction, but environments. Breeding programmes achieving near-complete stability should prioritize the selection and im- in the breeding population would provement of this trait. require roguing up to 70% of the initial

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material. This would imply a too large There are at least two aspects of reduction in genetic variability, which breeding that must be emphasized in is, by far, not the best decision from a connection with climate change. First, sustainability point of view. Applying more attention needs to be paid to a low intensity selection for stability, the choice of target environments for that is, removing around one-third of the improved material. This requires the most interactive families should proper progeny testing, where field tests be the option of choice as it may are established not only on optimum substantially reduce interaction effects sites but also in climatically marginal while probably keeping a sufficient environments. Alternatively, field trials genetic pool for future activities in the may be replaced by indoor or nursery breeding programme. tests with manipulated temperature and watering regimes, whenever Selection is inevitably associated with appropriate. Development of rapid and the loss of genetic variation. Low- inexpensive early-testing procedures frequency genes, which may prove for drought tolerance is also desirable beneficial under future environments, in order to accelerate deployment may be lost in each breeding cycle of clones, families and provenances through both selection and genetic appropriate for future climates (Dvorak, drift associated with the small size 2012). The second aspect is preserving of breeding populations (Godt et sufficient genetic diversity within the al., 2001). In many countries and improved material. Lindgren, Gea and for many tree species, improved Jefferson (1996) developed the concept material represents substantial part of status number as a kind of effective of reforestation. This is the case for population size to monitor the loss of clonally propagated species such genetic diversity in seed orchards or as poplars and willows, with a few clonal mixtures. Gene markers can also clones cultivated on huge areas, but be used to monitor the potential loss of also commercially important conifers genes. Nevertheless, the risk of gene loss (Scots pine and European larch), is unavoidable, especially in advanced- where most seed stock is produced in generation breeding, although it can seed orchards. The available options be diminished by appropriate breeding for raising diversity within the design. Therefore, there is a need for reproductive material produced by breeding approaches that combine the seed orchards are supplemental mass achieving of breeding targets with gene pollination or supplementing seed lots conservation measures. harvested in seed orchards with seeds deriving from other origins or other One of the major challenges for tree seed orchards. breeding is that the high priority breed-

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ing goals set today may be of limited The main advantage of the MPBS is that it importance when it is time to harvest combines the capture of the total existing the expected gains, especially given the genetic variation with a satisfactory vari- uncertainty of environmental condi- ation within each sub-populations, and it tions, which may change dramatically allows the sub-populations to adapt to the over a rotation time and are beyond any prevailing environmental conditions, be- human control. Consequently, a suc- sides the fact that the disruptive selection cessful breeding programme should be and thus the speed of evolution might be designed to match the future changes in higher in a sub-population than in an ex- breeding objectives and environmental tensive population of thousands of trees. change. The Multiple Population Breed- ing System (MPBS) developed by G. According to Eriksson, Ekberg and Namkoong can fulfil such requirements, Clapham (2006), the MBPS has been as the breeding population is subdi- adopted in Sweden for silver birch, vided into smaller sub-populations of lodgepole pine, Norway spruce and Scots equal size that are preferentially plant- pine breeding programmes, and different ed over a broad span of site conditions, sub-populations of these species have while the target trait might be different been distributed across different sites and for the different sub-populations. The environmental conditions. The MPBS MPBS is hierarchically structured and strategy has also been adopted in many emphasizes inter-populational diversity other breeding programmes across the within an array of populations both in globe. the traits targeted for improvement as well as for environmental adaptabilities. The nucleus breeding system is another In this way multiple adaptive peaks can example of a long-term breeding system be simultaneously revealed, as within that has the aim of gene conservation each sub-population separate trait com- and long-term genetic gain. This system binations and site adaptabilities can be splits the breeding population into sub- selected for applying recurrent selection populations of unequal size. The smaller (Namkoong 1984; Eriksson, Namkoong nucleus contains fewer trees, while the and Roberts, 1993), while the system can larger part includes a significantly greater be modified to produce interpopulation­ number of trees. The most intensive hybrids. Conservation goals can also breeding efforts occur within the small be met through the MPBS as broad nucleus, while the conservation and long- sampling to establish multiple popula- term gain are secured in the larger sub- tions, and continuous development of population. Transfer of genetic material inter-populational variation can ensure from the larger population to the nucleus the existence of ample diversity to meet can minimize inbreeding within the changing environmental challenges. nucleus despite the loss of genetic gain.

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Marker-assisted breeding and genomics molecular biology, DNA analysis and approaches bio-informatics, which are all evolving During the past 20 years, great at an astonishing rate. Current genomic progress has been made in assessing approaches include transcriptomics neutral DNA variation in tree species – sequencing the expressed genes in using a range of molecular techniques an organism or specific tissue type that have been developed over that and producing expressed sequence period. Isozymes, RFLPs, RAPDs, tag (EST) libraries, which reduces AFLPs, micro­satellites and, more the genome complexity and hence recently, non-coding SNP markers, sequencing effort; proteomics and have all proved extremely valuable in metabolomics – analysing the proteins establishing the demographic patterns and primary and secondary metabolite of neutral genetic variation. From the profiles; and genetic mapping aimed at FRM viewpoint, this neutral variation understanding of genetic architecture. does not provide any information on Genetic maps are extremely useful as phenotypic or adaptive variation, loci controlling quantitatively inherited which is crucial when considering traits (QTL) have been already the transfer of material in the context identified in many forest trees for a of climate change. The next major variety of growth, wood quality, and technological advance is likely to be other economic and adaptive traits. harnessing the recent development However, such traits are generally of affordable, rapid and informative controlled by small contributions from diagnostic techniques to evaluate large a large number of genes and there is numbers of adaptive genes and genetic currently a move away from the QTL variation at both the individual and method towards the whole genome population levels. Understanding the scanning approach. adaptive genetic potential of forest tree populations is fundamentally The various genomic approaches and critically important for evaluating create synergies to provide valuable their vulnerability to climate change information regarding tree genome and suitability for transfer. organization and function. These approaches are only possible because A genomics approach aims to account of the progress that has been made for the way in which the total gene in DNA sequencing technologies. complex of an individual contributes to Next-generation sequencing (NGS) its phenotype. Genomics encompasses a technologies can produce thousands range of approaches that aim to integrate or millions of sequences at once and traditional genetic disciplines with the generate gigabytes of data in a matter emerging cutting edge technologies in of days at a much reduced cost.

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Rigault et al. (2011) used the power of have broad contiguous populations, yet NGS to confirm and enhance the Picea which demonstrate local adaptation, glauca gene catalogue by providing a they are excellent models for empirical deeper coverage for rare transcripts, by study in landscape genomics. extending many incomplete clusters and by augmenting the overall transcriptome A study that is underway at the coverage. With this availability of University of California in the United sequence information at thousands of States of America aims to generate putatively important genes or regulatory complete polymorphic transcriptome regions, the characterization of adaptive information from eight conifer species genetic diversity and the association across California, select genes showing with phenotypic trait variation is adaptive variation, and then design becoming more accessible. The progress dense (20 000 SNPs) Illumina Infinium that has been made with sequencing genotyping chips to genotype 2000 trees technologies is particularly important from each of the eight species (16 000 for studying the large and highly trees in total). The scale of this project repetitive genomes of conifer species. could not have been foreseen even 5 This has involved applying both the years ago. sequencing of lower complexity targets, i.e. transcriptome ESTs or restriction- A commercial whole-genome sequenc- site-associated DNA markers (RADs), ing service has been launched recent- and the whole-genome sequencing ly, with a two-week turnaround for approach. US$ 9 500. Third-generation sequencing technologies utilizing a nano-scale sen- One of the ultimate practical objectives sor to electronically read the sequence for tree genomics is to gain the ability of a single DNA molecule may make the to identify genes under selection due cost of human whole genome sequenc- to geo-climatic factors, determine their ing fall to below US$ 1 000. Progress of allelic diversity, and then apply that this sort would enable tree geneticists knowledge in management decisions to embrace the concept of genome-wide related to the use and transfer of genetic association studies (GWAS), involving resources. scanning large sample sizes with hun- dreds of thousands of SNP markers The discipline of landscape genomics has located throughout the genome with a recently emerged, combining population subsequent comparison of the frequen- genetics and landscape ecology to study cies of either single SNP alleles, gen- patterns of demographic and adaptive otypes, or multi-marker haplotypes, genetic variation across heterogeneous between phenotypically different co- landscapes. Since forest tree species often horts. This analysis identifies loci with

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statistically significant differences in Molecular markers for genetic allele or genotype frequencies between characterization of forest reproductive phenotypes, pointing to their role in the material trait. In contrast to candidate gene stud- Most forest tree species exhibit high ies, which select genes for studies based levels of genetic diversity that can on known or suspected mechanisms, be characterized using molecular GWAS permits a comprehensive scan of markers and used for identifying the the genome in an unbiased fashion, and origin of reproductive material raised thus has the potential to identify novel in nurseries prior to, or after planting, factors. should a need for confirmation arise. The majority of current molecular Genomic selection (Nakaya and Isobe, markers are selectively neutral (neutral 2011) is a form of marker-assisted variation results from differences selection (MAS) that selects favourable between genotypes that do not affect individuals based on genomic estimated their ability to survive and reproduce) breeding values (GEBVs). GEBVs are and they do not reflect the distribution also used in animal breeding, but have of adaptive variation, which will be not been a popular index in plant considered later. However, neutral breeding. They are defined as “the sum markers can provide information on the of the estimate of genetic deviation level of genetic diversity, colonization and the weighted sum of estimates of routes and lineages, in addition to a breed’s effects” which are predicted helping to confirm the origin of FRM or using phenotypic data from family to ensure that the trail of material from pedigrees. GEBV has become a feasible nursery to forest is genuine. approach with the recent advances in high-throughput genotyping platforms. The level of genetic diversity can be easily measured, typically with highly What once cost millions of dollars and variable microsatellites­ or SNP markers. took years to achieve may soon cost A simple example of this is the use of less than many standard diagnostic molecular markers to assess the genetic tests today. Whilst such technologies health of planting stock, i.e. whether may take a while to filter down to tree it is showing signs of inbreeding. In geneticists, it is clear that there will southern Germany, the genetic diversity be a paradigm shift in the amount of of all Douglas fir and silver fir stands sequence information generated. From are assessed using a suite of isozyme the FRM perspective, the challenge will and microsatellite­ markers. If the stand be to understand and exploit this huge shows lower levels of variation in amount of information in a meaningful comparison with other stands, it will way. be rejected as a seed source. Molecular

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markers are also used to determine Research efforts are currently focusing the level of purity of oak and larch. on studying variation in candidate genes As a further example, the inherent related to traits that will be important variability between individual trees can in response to climate change, such as be harnessed with molecular markers bud­burst (Derory et al., 2009), bud set to enable individual clone identification (Frewen et al., 2000) and water stress (e.g. cherry, apple and pear) and also (Perdiguero et al., 2011). For the time for the identification of hybrids (e.g. being, these approaches are limited to poplars (P. nigra × P. deltoides) and larch the search for candidate genes and the (L. kaempferi × L. decidua)). exploration of their diversity in extant populations (Gonzalez-Martinez et al., The choice of the most appropriate 2006). Although genomics approaches molecular method to trace the origin enable the assessment of genetic of FRM largely depends on the species differentiation simultaneously in a and the current information on the large number of loci, the influence of spatial distribution patterns of its polymorphisms on adaptive variation genetic diversity. For example, there cannot be evaluated without good is highly detailed information on the quality phenotypic data and extensive phylo­geo­graphical variation patterns breeding programmes. Current of chloroplast DNA haplotypes in evidence from genomics studies points European oaks (Petit et al., 2002) which to a complex multi-gene inheritance of makes chloroplast DNA markers an adaptive traits. excellent tool to track the origin of oak. The utility of uni-parentally inherited There is a clear underlying message variation is due to more restricted that the gathering of basic genetic seed dispersal. In many European tree information on tree species at all species, the highest levels of marker levels (individual trees, stands and diversity have been found in central countries) should be done with the Europe, which could be due to ancient knowledge that the greater the amount refugia or an admixture of genetically of information that is available, the differentiated lineages (Petit et al., 2003). greater the opportunity to use DNA markers accurately for the identification In the future, the identification of of origin of FRM, and to understand the adaptive DNA markers could be of current genetic health of tree species. great benefit for choosing suitable It must be remembered that trees do FRM material that will be best suited not follow political boundaries and to a particular climate scenario. There therefore the use of common marker is currently a great deal of ongoing­ sets with common standards between work in the field of adaptive variation. countries should be encouraged.

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Advances in phenotyping and phenomics of genomics, efficient techniques Traditionally, the set of traits assessed for assessing tree phenotypes were in field experiments with forest trees needed as key components of was limited to survival rate, growth, conventional breeding approaches phenology, frost resistance and tree- (Neale and Kremer, 2011). The ability architectural traits. Although these of rapid and cost-effective scoring of are the traits of primary interest for phenotypic traits on large numbers of a practical forester, their potential is trees is crucial, not only for practical limited for understanding adaptation breeding but also for revealing the processes and explaining response genetic background of complex patterns to environmental changes phenotypic traits. This also applies to over time (e.g. climate change) and conventionally assessed traits. Recent space (FRM transfer). development of new measuring devices and computer-aided field- Due to their size and longevity, data collection systems, such as forest trees are not suitable objects FieldMap, has allowed reliable and for scoring phenotypic traits in as rapid assessment of growth traits, a complex way as is possible with even in older trials. other organisms. In biology, the term ‘phenomics’ as an analogy to Nevertheless, deeper understanding genomics was recently coined for of the genetic basis of adaptation acquisition of high-dimensional processes and predicting responses phenotypic data on an organism- of tree populations to environmental wide scale (Houle, Govindaraju change requires broadening the and Omholt, 2010). Phenomics is an spectrum of traits assessed in all types emerging trans-discipline dedicated of indoor and outdoor experiments, to the systematic study of phenotypes including common-gardens, and using new methods for high-through­ finding high-throughput­ scoring put scoring, and focuses on linking procedures for these traits. These genetic variation assessed in genomic would include those traits listed below. and gene-expression studies to phenotypic variation. It has become a • Physiological traits. Recently, meth- recognized component of biological, odologies for monitoring a large medical and agricultural sciences. number of parameters characterizing plant carbon and water turnover have All recent research projects focusing developed considerably (Kramer et on tree genetics and breeding lay al., 2004; Rosenqvist and van Koot- emphasis on high-through­put en, 2003; Seibt et al., 2008) and sim- phenotyping. Long before the era ilar improvements are also expected

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in monitoring nutrient uptake, frost • Pest and pathogen resistance. Rapid resistance, drought tolerance, etc. At development of remote sensing and the same time, increased availability image analysis allows increasing the of high-resolution digital cameras efficiency of large-scale estimates of and image-processing software al- pest and pathogen damage in out- low documenting the growth pro- door trials (King, 2000). cess and reactions to time-dependent • Proteome and metabolome. stress factors, using measurements in Improved methods of analytical the full range of technically available organic chemistry allow identification wavelengths from infrared to X-ray of numerous proteins and secondary imaging, or other imaging methods, metabolites mediating the response such as magnetic resonance imaging of a genotype to a particular (MRI) or terahertz scanning (Eberius environment (Ossipov et al., 2008; and Lima-Guerra, 2009). Warren, Aranda and Cano, 2012). • Wood properties. Development of microscopy techniques associated Facilities allowing manipulation of the with computer-aided image analysis environment, such as phytotrons and has allowed identification of numer- climatic boxes, are available in many ous structural and ultrastr­ uctural laboratories throughout Europe. Recent wood traits (Fahlén and Salmén, international projects, such as NovelTree 2005; Xu et al., 2006). New techniques or Trees4Future, have undertaken have also appeared for the analysis inventories of such facilities and of wood chemistry (Selig et al., 2011; devices suitable for high-throughput Zhou, Taylor and Polle, 2011). Al- phenotyping, and allowed cross-linking though wood properties seem to be and international cooperation in their mainly of commercial importance, utilization. they affect tree stability, water bal- ance and other ecologically relevant traits.

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recommendations

FRM transfer is a valuable option for Local FRM will not always be the answer adapting forests to climate change to climate change challenges. As local Forest tree populations have (genetically) environmental conditions change, forest adapted over a long period (and are still managers should extend their options to adapting) to their respective habitats both local and non-local FRM. and, as a result of this adaptation, formed so-called provenances. Climate change is expected to alter forest habitat Use provenances instead of species in conditions in Europe at such a pace that assisted migration schemes the natural processes (selection, gene Science has repeatedly shown that flow, migration) that drive evolution adaptive genetic diversity within forest and adaptation will not act fast enough. tree species is often very large and yet it Therefore, human intervention in seems that, under the pressure of climate the form of FRM transfer (assisted change, forest management would migration) is a valuable option to adapt rather change species than provenances forests to climate change, especially when designing reforestation efforts. in those areas that are most severely Therefore, there is an urgent need to threatened by climate change. disseminate information and knowledge on the adaptive potential that is readily available from different FRM within tree Local is not always best species. Autochthonous local tree populations originate from complex selection processes acting on a restricted regional Transfer of FRM also has its limits gene pool. For various biotic and With increasing temperature and abiotic reasons, local populations do periods of drought there is an increasing not always demonstrate optimum demand from forestry for provenances fitness (for important forestry traits) from warmer southern regions. As compared with other FRM in common- long as extreme events such as late garden experiments. In addition, in frosts occur, these provenances can Europe, locally found populations often be recommended only in exceptional originate from historical FRM transfer cases, and the transfer of the material but for which passport data are lost. used has to be well documented. Short-

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rather than long-distance transfers will between control agencies in different often be more ecologically relevant and EU Member States and collaborating should be preferred. The conservation of countries are necessary. Recent local genetic resources should be taken technological breakthroughs, particularly into account when assisted migration is in molecular biology, make it possible being considered. Forest managers should for control agencies to easily and cost- protect particularly threatened FRM effectively monitor and trace FRM, and to (mostly peripheral populations from rear disseminate information to the end-users. edges of geographical distributions) that could be of use in other, more suitable, locations. The need for FRM documentation increases under climate change The best adapted FRM of today may Revision of transfer recommendations is not prove the best adapted FRM of necessary at the pan-European level tomorrow under climate change. By Provenance regions or seed zones keeping track of successes and failures in represent the spatial framework for FRM management decisions, forest managers transfer recommendations in Europe. will be able to adjust their strategies. Their number and size vary greatly Data on FRM – geographical origin, among countries because different harvesting conditions, genetic diversity approaches are used for their delineation. and production methods – are likely to FRM is increasingly traded across borders be key information sources for plantation but the national recommendations rarely efforts, so that forest managers should be offer help in deciding where the imported particularly keen to ask for and to keep a material should be used. Transfer record. recommendations should have a pan- European perspective and should also include climate change considerations. T ree breeding offers opportunities for forestry under climate change Breeding programmes have typically More stringent control of FRM is needed at all focused on improving yield, wood quality, production and marketing stages and resistance to pests and diseases. The legal frameworks for FRM production Resistance or tolerance to drought and marketing are well defined at EU and or increased water stress periods are OECD levels. However, major differences becoming a priority for new generations still persist at national level for controlling of breeding programmes, and so does the origin of FRM throughout the keeping large breeding populations. Such production chain. Harmonized control FRM can be tailored to managers’ needs, mechanisms and close cooperation from very specific to large portfolio uses.

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It is crucial to continue provenance molecular basis of adaptation and the research under climate change relationship between genetic variation Provenance experiments provide a in a large set of genes and phenotypes in wealth of resources for evolutionary environmental gradients still represent ecology and climate change studies, knowledge gaps. International and they are the most reliable basis for cooperation must be intensified in formulating practical recommendations this context, including collaborative for FRM. Existing networks of utilization of shared research facilities. provenance experiments need to be upgraded to include under-represented (e.g. peripheral) populations and sites, Dissemination of information on the value and need to be regularly monitored of FRM to forest owners, managers and and measured. Pan-European efforts policy-makers needs to be improved for collating, archiving and analysing The scientific community has extensive data at species-range level should knowledge and information on the be continued and intensified, so that potential that the use of FRM offers for results can be transferred into practical facilitating the adaptation of forests to recommendations for forest managers. climate change. Education and training for increasing awareness of stakeholders, such as forest owners, forest and forest Knowledge gaps should be filled on the habitat managers, and policy-makers, adaptation of forest trees needs to be continued and intensified. Understanding the adaptive genetic Ways to increase capacity building potential of forest tree populations and for using science-based knowledge their response to environmental changes and to foster an efficient and mutually in time and space is fundamentally rewarding science–management–policy and critically important for assessing dialogue, must be fully explored, more how FRM might be used in the face than ever before. of climate change. In particular, the

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Annex 1

List of European projects relevant for the use and transfer of forest reproductive material Project Framework Leader Short description LINKTREE BiodiverSa INIA LINKTREE uses a combination of high throughput (2009–2012) sequencing and/or genotyping of ecologically Spain relevant genes and quantitative genetics experiments to evaluate levels of standing genetic variation and signatures of selection in natural forests. (www.igv. fi.cnr.it/linktree) Trees4future FP7 - INRA Trees4Future is an Integrative European Research structures Infrastructure project that aims to integrate, develop 2011–2014 France and improve major forest genetics and forestry research infrastructures. (www.trees4future.eu) EUFORINNO FP7–Regpot SFI EUFORINNO addresses the strategy of SFI to become a reference centre for Central and South- Slovenia Eastern Europe in the European Forest Research and Innovation Area (EuFoRIA). Between the key areas of cooperation is also the control of FRM and the establishing of a forest gene bank. Map/FGR COST Action CREA SEL Strengthening the adaptation of forest genetic (2012–2015) resources (FGR) to climate change in marginal and/ Italy or peripheral populations of forest trees in Europe through appropriate conservation and management strategies NOVELTREE FP7 INRA NOVELTREE will provide an improved understanding of the biology of forest tree species and enable (2008–2012) France significant genetic improvement in the composition and characteristics of forest products in order to satisfy the needs (e.g. quality, quantity, sustainability, vulnerability) of consumers and of the forest-based sector. (www.noveltree.eu) FORGER FP7–KBBE ALTERRA The FORGER project will make available sound and integrated information on genetic resources to 2011–2015 Netherlands forest managers and policy makers by improving inventories on forest genetic resources in Europe, developing a common protocol to measure and monitor genetic diversity, analysing historic and current use and exchange of forest genetic resources over Europe, and analysing how forest genetic resources are affected by climate change and forest management practices. (www.fp7-forger.eu) TREEBREEDEX FP6– INRA The aim of the TREEBREEDEX project was to Infrastructures develop a European platform for research and 2006-2010 France technical collaboration on tree breeding. (http:// treebreedex.eu) Evaluation of COST Action vTI This COST Action had 22 countries participating. Beech Genetic E52 Its main objective was to evaluate jointly, for the Resources for Germany first time, 60 international beech provenance trials Sustainable 2006–2010 located in 19 European countries, a total of 200 Forestry provenances in all. EVOLTREE EU FP6 Network INRA Evoltree associates four major disciplines – of Excellence genomics, genetics, ecology and evolution – for 2006–2010 and France understanding, monitoring and predicting genetic ongoing as EFI diversity, ecosystem structures, dynamics and programme processes in European forests.

75 EUFORGEN