Approaches to the Conservation of Forest Genetic Resources in Europe in the Context of Climate Change

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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 Approaches to the conservation of forest EUFORGEN genetic resources in Europe in the context of climate change

Colin T. Kelleher, Sven M.G. de Vries, Virgilijus Baliuckas, Michele Bozzano, Josef Frýdl, Pablo Gonzalez Goicoechea, Mladen Ivankovic, Gaye Kandemir, Jarkko Koskela, Czesław Kozioł, Mirko Liesebach, Andreas Rudow, Lorenzo Vietto and Peter Zhelev Stoyanov

Approaches to the conservation of forest genetic resources in Europe in the context of climate change

Bioversity International is a global research-for-development organization. We have a vision – that agricultural biodiversity nourishes people and sustains the planet. We deliver scientific evidence, management practices and policy options to use and safeguard agricultural and tree biodiversity to attain sustainable 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, productivity 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 cooperation 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 the implementation 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 experiences, 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 Coordinators nominated by the member countries. The EUFORGEN Secretariat is hosted by Bioversity International. Further information on EUFORGEN can be found at www.euforgen.org.

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Citation: Kelleher, C. T., de Vries, S.M.G., Baliuckas, V., Bozzano, M., Frýdl , J., Gonzalez Goicoechea, P., Ivankovic, M., Kandemir, G., Koskela, J., Kozioł, C., Liesebach, M., Rudow, A., Vietto, L., and Zhelev Stoyanov P. 2015. Approaches to the Conservation of Forest Genetic Resources in Europe in the Context of Climate Change. European Forest Genetic Resources Programme (EUFORGEN), Bioversity International, Rome, Italy. xiv+46 pp.

Layout: Ewa Hermanowicz Cover photos: Michele Bozzano

ISBN 978-92-9255-032-5

This publication has been printed using certified paper and processes so as to ensure minimal environmental impact and to promote sustainable forest management.

Authors

Colin T. Kelleher Gaye Kandemir National Botanic Gardens, Dublin Ministry of Forest and Water Affairs, Ankara Ireland Turkey

Sven M.G. de Vries Jarkko Koskela Centre for Genetic Resources, Bioversity International, Rome Wageningen University and Research Italy Centre, Wageningen the Netherlands Czesław Kozioł Kostrzyca Forest Gene Bank, Miłków Virgilijus Baliuckas Poland Institute of Forestry, Lithuanian Research Centre for Agriculture, Girionys Mirko Liesebach Lithuania Thünen Institute of Forest Genetics, Grosshansdorf Michele Bozzano Germany Bioversity International, Rome Italy Andreas Rudow Institute of Terrestrial Ecosystems, Josef Frýdl ETH Zürich Forestry and Game Management Research Switzerland Institute, Jílovišteˇ Czech Republic Lorenzo Vietto Council for Agricultural Research and Pablo Gonzalez Goicoechea Economics, Research Unit for Intensive NEIKER-Tecnalia, Vitoria-Gasteiz Wood Production (CREA-PLF) Spain Casale Monferrato Italy Mladen Ivankovic Croatian Forest Research Institute, Peter Zhelev Stoyanov Jastrebarsko University of Forestry, Sofia Croatia Bulgaria

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iv P r e f a c e

Preface

During the past two decades, the impacts of climate change on forests and the role of forests in mitigating climate change have been debated intensively. Globally, deforestation has contributed significantly to climate change by releasing carbon dioxide into the atmosphere and reducing the production of oxygen. Deforestation still con- tinues in many parts of the world and a number of international initiatives have been developed to promote and implement sustainable forest management, especially in developing countries. In Europe, forests have been increasing both in terms of area and growing stock during the past 50 years. European forests have thus acted as a carbon sink as they have been recovering from the past centuries of deforestation and overharvesting.

In recent years, European policymakers have discussed various options for adaptation and mitigation to climate change and developed various policies to enhance the role of forests and the forest sector in mitigating the impacts of climate change. Most countries have incorporated climate change aspects into their national forest programmes and national action plans for biodiversity conservation. Several countries have also developed cross-sectoral national adaptation strategies to climate change. Unfortu- nately, these measures have mostly neglected the role of forest genetic resources in the adaptation of forests to climate change. Furthermore, although climate change is also considered as a threat to biodiversity conservation, most conservation efforts have focused on species and habitat diversity and paid little attention to genetic diversity, the fundamental basis of all biological diversity.

Within the framework of the European Forest Genetic Resources Programme (EUFORGEN), the implications of climate change for the conservation and use of forest genetic resources have been increasingly discussed during recent years. EUFORGEN was established in 1994 to coordinate pan-European collaboration on forest genetic resources as part of the FOREST EUROPE process (earlier the Ministerial Conference on the Protection of Forests in Europe). During Phase IV (2010-2014), EUFORGEN had three objectives, 1) promote appropriate use of forest genetic resources as part of sustainable forest management to facilitate adaptation of forests and forest management to 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. EUFORGEN has brought together scientists, managers and policymakers to discuss various issues related to forest genetic resources and to develop pan-European approaches for better management of these resources.

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The present report presents the findings and recommendations of the EUFORGEN working group on climate change and conservation of forest genetic resources. The report presents the current state of knowledge on the implications of climate change for conserving forest genetic resources and provides recommendations for further action. The working group met twice; the first meeting was hosted by Bioversity In- ternational in Maccarese, Italy on 18-20 June 2013 and the second one by the Centre for Genetic Resources in Wageningen, Netherlands on 4-6 February 2014. The Work- ing Group provided an update to the EUFORGEN Steering Committee during its ninth meeting held in Tallinn, Estonia, on 3-5 December 2013. The draft report was presented to the EUFORGEN Steering Committee for review during its 10th meeting which was held in Edinburgh, United Kingdom on 16-18 June 2014. Comments from the Steering Committee were addressed in the final report.

We acknowledge with thanks the contributions received from: Mladen Ivankovic (Croatia), Petr Novotný (Czech Republic), Václav Buriánek (Czech Republic), Mart Külvik (Estonia), Leena Yrjänä (Finland), Eric Collin (France), Alexis Ducous- so (France), Bruno Fady (France), Heino Wolf (Germany), Konstantinos Spanos (Greece), Aristotelis C. Papageorgiou (Greece), Attila Borovics (Hungary), Giovan- ni Giuseppe Vendramin (Italy), Maria Gras (Italy), Darius Danusevicius (Lithua- nia), Frank Wolter (Luxembourg), Tor Myking (Norway), Mari Mette Tollefsrud (Norway), Małgorzata Pałucka (Poland), Iwona Szyp-Borowska (Poland), Ladislav Paule (Slovakia), Dušan Gömöry (Slovakia), Robert Brus (Slovenia), Dragan Mat- ijašic (Slovenia), Santiago C. González-Martínez (Spain), Sanna Black-Samuelsson (Sweden), Peter Rotach (Switzerland), Murat Alan (Turkey) and Joan Cottrell (Unit- ed Kingdom).

vi C o n t e n t s

Contents

Authors iii Preface v Acronyms used in the text ix Executive summary xi Introduction 1 State of Knowledge 5 Predictions of climate change 5 General consequences of climate change for forests 6 Climate change consequences for forest genetic resources 8 Threatened tree species and populations 10 Knowledge of measures and existing strategies 16 Implications for conservation 22 Recommendations for management of genetic conservation units and establishment of additional units 25 Management of existing genetic conservation units 25 Establishment of additional genetic conservation units 27 Recommendations for complementary ex situ approaches 31 Prioritization of tree species and establishment of ex situ populations 31 Dynamic ex situ conservation 33 Static ex situ conservation measures 34 Adequate monitoring 34 Recommendations for research 35 Conclusions and overall recommendations 37 References 39 Appendix 1 45

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viii A c r o n y m s

Acronyms used in the text

CO2 carbon dioxide EU European Union EUFGIS European Information System on Forest Genetic Resources EUFORGEN European Forest Genetic Resources Programme FGR forest genetic resources FAO Food and Agriculture Organization of the United Nations IPCC Intergovernmental Panel on Climate Change IUFRO International Union of Forest Research Organizations IUCN International Union for Conservation of Nature UN United Nations UNFCCC United Nations Framework Convention on Climate Change

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x E x e c u t i v e S u m m a r y

Executive summary

Over the past decades, European countries have taken significant steps in conserving the genetic resources of forest trees. However, they have rarely taken into account the implications of climate change for the conservation of forest genetic resources. Climate change poses unique conservation challenges that require specific responses. For these reasons, the EUFORGEN Steering Committee established a working group on climate change and the conservation of forest genetic resources. This report presents the conclusions of the working group.

In Europe, forests have been expanding in terms of area and timber stock over the past 50 years and subsequently they have acted as a carbon sink while they have been recovering from previous eras of deforestation. National adaptation strategies to climate change and other policies have been formulated in many European countries to harness the potential of forests and the forestry sector for mitigating climate change. However, the impacts of climate change on forests, and especially on their genetic diversity have not been given a proper consideration in these policies.

The working group made several recommendations for action. These focus on 1) establishing additional genetic conservation units specifically to respond to climate change, 2) enhancing cooperation among countries and enlarging the pan-European collaboration on the conservation of forest genetic resources, 3) the need for continued and expanded monitoring and sharing of data, including the development of decision tools and red lists, and 4) further research on aspects of assisted migration and on marginal and peripheral tree populations.

Climate change and forest genetic diversity Global climate models predict an increase in average surface temperature for Europe, with winters becoming warmer in the north and summers in the south. The models also predict increases in winter precipitation in the north and decreases in the south, as well as decreases in summer precipitation in central and western Europe. Southern and western Europe are likely to experience warmer and dryer conditions with droughts starting earlier and lasting longer. Impacts will be most severe in the southern Iberian Peninsula, the Alps, the eastern Adriatic and southern Greece. In addition to these changes, the models also predict that extreme events will occur more often. While uncertainties remain, it can be concluded that the near future will see changes in the range of individual tree species, and that the species composition and the dynamics of forests will be affected.

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From the point of view of conservation, the most important areas to focus on will be at the edge of species’ ranges. While populations are migrating into areas that climate change has made more suitable, the leading edge is likely to suffer genetic narrowing because a few founder individuals, representing only a subset of the total genetic diversity, will be pioneers in new areas. At the same time the populations at the trailing edge are likely to experience fragmentation and a reduction in the number of individuals, with subsequent loss of genetic diversity.

Assessing and managing threats It remains uncertain exactly how tree species and populations will respond to climate change. Nevertheless, it is important to attempt to assess the impacts of climate change on tree populations using a combination of intrinsic factors, such as population structure, regenerative capacity and dispersal ability, and extrinsic factors, including biotic threats like pests, pathogens and species competition and abiotic threats such as fire or changes in land use. The working group considered a wide range of potential threats and developed a preliminary decision cascade tool to aid the identification and management of those populations most in need of conservation. The proposed decision cascade tool appended to the report is a proof of concept and will need further development. Such a tool will be helpful in drawing up a red list of threatened tree species in Europe, which should focus on populations rather than species and which is urgently needed to guide future conservation.

The working group noted that many European tree species, especially those threatened in southern Europe, are also present in North Africa and the islands of Macaronesia. These populations are perhaps even more likely to be negatively affected by climate change, with little to no chance of colonisation from the south. For this reason, the working group recommends that for the purpose of conservation of forest genetic resources, the pan-European collaboration should include these areas. Collaboration should be initiated to include countries containing populations of relevant species outside of Europe. A larger geographic coverage of the regional collaboration on forest genetic resources would also be useful for developing a red list of threatened tree populations.

Conservation approaches Many European countries currently have conservation strategies for forest genetic resources and efforts are under way to expand and strengthen these strategies. Current strategies, for example the EUFORGEN led pan-European network, do not always

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include populations that are under threat due to climate change. The report offers suggestions as to how to identify such units and recommends that these additional units be added to the European Information System on Forest Genetic Resources (EUFGIS) database, with an additional identifying field, so that information from them can be considered alongside all other conservation units in making strategic management decisions. There is also a possible role for assisted migration, especially for those species facing the most intense climate change. While there will clearly be problems to solve, assisted migration remains an important option to consider. Even with greater attention to the in situ conservation, there will still be a need for ex situ conservation. Collections of living trees, seeds, gametes and plant tissue may all be considered in different cases, and priorities assigned according to an agreed set of criteria. As with in situ conservation units, ex situ units should be monitored and be included and specifically identified in the EUFGIS database.

Research The report deals primarily with questions of management, but in considering these questions, gaps in scientific knowledge became apparent. Little is known about the concrete effects of climate change on genetic diversity of tree populations. In addition, there is a lack of knowledge about assisted migration and how it might affect the genetic composition of a species and the species composition of an ecosystem. Marginal and peripheral tree populations are also poorly studied. These populations need to be identified, characterised and studied. More research results on these topics would help to improve conservation in the context of climate change and influence future policymaking.

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xiv I ntroduction

Introduction

Climate change and its consequences The impacts of climate change on forests for the Earth and human societies have and biodiversity have been analysed been analysed and discussed at the in- by numerous modelling studies. The ternational, regional and national levels projected changes in climate are expect- for more than 20 years. In 1992, the Rio ed to shift and reduce the distribution Earth Summit adopted the UN Frame- range of major commercial tree species work Convention on Climate Change in Europe (e.g. Hanewinkel et al., 2012). (UNFCCC) which was designed to However, most modelling approaches stabilize greenhouse gas emissions at have considered species as static and a level that would prevent dangerous independent entities and overlooked changes in the global climate system. the importance of genetic diversity and The UNFCCC entered into force in 1994 phenotypic plasticity in adaptation to and a total of 195 countries have ratified climate change (e.g. Bellard et al., 2012). it, although greenhouse gas emissions Furthermore, studies on the impacts continue to increase. According to the of climate change on biodiversity have Intergovernmental Panel on Climate largely focused on the species and eco- Change (IPCC), the global average sur- system responses and neglected the role face temperature (combined land and of genetic diversity (Bellard et al., 2012). ocean temperatures) has increased by The incorporation of genetic diversi- nearly 1°C during the period 1901 to ty and phenotypic plasticity into spe- 2012 (IPCC 2013). The recent IPCC re- cies-distribution models has shown that port also notes that it is very likely that the predicted decreases and/or changes temperature will continue to increase in the distribution ranges of tree species throughout the 21st century in different due to climate change are likely to be parts of the world, including Europe. smaller than previously estimated (Ben- The latest projections indicate that sum- ito Garzón et al., 2011). mer temperature (June-August) will increase by 3°C to 4°C in most parts of Climate change will impact existing Europe by 2081 to 2100, and even 4°C to conservation networks or systems and 5°C in some places in the Mediterranean will influence how new systems are de- region (IPCC 2013). This will considera- signed. Conservation systems should bly alter the climatic conditions to which be based on a dynamic and large-scale European forests are currently adapted. approach to ensure that the target spe-

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cies are able to adapt to climate change thermore, the workshop recommended (Hannah 2010). In Europe, a pan-Euro- that the policymakers promote forest pean network of genetic conservation management practices which maintain units of forest trees is an example of such evolutionary processes of forest trees a conservation approach. Currently, this and support natural regeneration of pan-European network consists of 3214 forests, especially in areas where long- conservation units in 34 countries. The term natural regeneration is self-sus- documentation of this network and the tainable despite climate change. During launch of the EUFGIS Portal1 in 2010 EUFORGEN Phase III (2005 to 2009), were results of the earlier EUFORGEN the Conifers, Scattered Broadleaves and work that had been carried out over sev- Stand-forming Broadleaves Networks eral years. continued the discussion on the implications of climate change for the conser- In 2006, EUFORGEN and the Interna- vation of forest genetic resources while tional Union of Forest Research Organ- developing minimum requirements for izations (IUFRO) organized a workshop the genetic conservation units of these on climate change and forest genetic groups of tree species. In addition, the diversity (see Koskela et al., 2007). The Forest Management Network also dis- workshop noted that the impacts of cli- cussed the impacts of climate change on mate change on forests will vary in dif- the conservation and use of forest ge- ferent parts of Europe bringing both op- netic resources. Building on this work, a portunities and threats. The workshop EUFORGEN working group developed also stressed the key role that forest ge- a pan-European genetic conservation netic resources play in maintaining the strategy for forest trees in 2012-2013 (de resilience of forests against the threats Vries et al., 2015). and in harnessing the opportunities. One of the workshop recommendations In 2012, the EUFORGEN Steering Com- urged European policymakers and the mittee decided to establish a working forest sector to recognize the importance group to further review genetic con- of forest genetic diversity in mitigating servation methods (both in situ and ex the negative effects of climate change situ) in the context of climate change. on European forests by incorporating The working group was tasked to devel- the management of this diversity into op recommendations for the manage- national forest programmes and other ment of the conservation units and the relevant policies, programmes and strat- networks of the units, and to propose egies (e.g. national adaptation strategies complementary conservation measures. to climate change and national action More specifically, the Steering Commit- plans for biodiversity conservation). Fur- tee requested the working group to:

1 http://portal.eufgis.org/

2 I ntroduction

• Review relevant outputs of the previ- Furthermore, the Steering Committee ad- ous Forest Management Network vised that the working group should not • Review predictions of climate change only focus on the management of single and their consequences for conserva- units but also the pan-European network tion of FGR (e.g. abundance, compo- of genetic conservation units. The work- sition and distribution of forest tree ing group was also requested to address species and populations) both in situ and ex situ conservation in • Review findings on the most threat- climate change context, and analyse the ened tree species and populations level of duplication needed in conserva- • Develop recommendations for man- tion efforts. The working group was also agement of genetic conservation asked to explore the idea of establishing units conservation units outside the current • Develop complementary ex situ distribution ranges of tree species. The approaches following chapters of this report present • Present an update in detail the findings and recommenda- • Prepare a draft report tions of the working group.

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4 S t a t e o f K n o w l e d g e

State of Knowledge

Predictions of climate change variety of future emission scenarios the It is well documented that human activ- IPCC (2007a) has predicted temperature ities have significantly affected global increases of about 0.2˚C per decade and climate since the beginning of industri- a minimum of 0.1˚C per decade under a alisation which started around the 1750s. very optimistic scenario of emissions be- The Fifth Assessment Report of the Inter- ing maintained at the year 2000 levels. governmental Panel on Climate Change (IPCC 2013) echoed the finding of the Global climate models used in the IPCC fourth report (IPCC 2007a), according to Fourth Assessment Report (2007a) pre- which the concentration of atmospheric dicted an increase in average surface air carbon dioxide (CO2) has increased from temperature for Europe into the mid- 280 ppm in the pre-industrial era to 379 21st century (2041-2070) (Roeckner et al., ppm in 2005. The highest rate of increase 2003; Marsland et al., 2003). The models of CO2 concentration was detected in the predicted that warming during the win- period from 1995 to 2005. This increase ter period (December-February) will be exceeded the natural range over the last greatest in north-eastern Europe (more 650,000 years (180 to 300 ppm) and is re- than 3° C), while in the summer period ported to be primarily a result of burning (Jun-Aug),ˇ the largest increase in sur- fossil fuels and changes in land use, such face air temperatures can be expected in as increased urbanization and deforest- southern Europe, (e.g. in the Iberian Pen- ation. The global methane concentration insula) where temperatures could rise by has also increased significantly from 715 up to 4° C during this period (Brankovic ppb to 1,774 ppb in 2005. This again ex- et al., 2010). ceeds the natural range (320 to 790 ppb) and is considered to be due primarily The anticipated changes in winter precip- to agricultural development. Increased itation patterns in Europe are expected to

CO2 and methane concentrations have increase in areas north of 45 ° N latitude resulted in a marked greenhouse effect and decrease in areas south of 45 ° N and global warming beyond that ex- latitude. For summer, the same models pected through purely natural process- predict a reduction in precipitation in the es. These increases and their impacts are temperate latitudes of central and west- being used to predict future changes in ern Europe. Warmer and drier conditions a variety of climatic variables. Using a may cause prolonged droughts and ex-

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tend the fire season with increased forest dictions for Europe, we can expect con- fire risk, particularly in Mediterranean tinued changes in species ranges, species countries where droughts will probably composition and forest dynamics (e.g. start earlier and last longer. Countries in EC 2008). central Europe will probably experience the same number of hot days as those Forests generally occupy large areas and currently occurring in southern Europe. are populated by long-lived trees. Thus, The regions in Europe most affected are climate change represents a particular likely to be the southern Iberian Penin- set of challenges to the biology of forest sula, the Alps, the eastern Adriatic and trees. Within the lifetime of an individual southern Greece (IPCC 2007a). tree the conditions could become up to 3˚C warmer (based on a 150-year lifespan The negative effects of expected changes at an IPCC prediction of a 0.2˚C decadal in average temperature and precipitation increase in global temperature). Howev- will be accentuated by the increase in er, forest trees also have a particular ad- frequency of extreme events. The speed aptation capacity in comparison to other of climate change is expected to be rapid, plants, such as annual herbs, that may with the expectation of rearrangement help them to cope with climate change. of current global distribution of climatic Forest trees are known to demonstrate conditions within this century (Loarie extensive plasticity in their response to et al., 2009). Climate change and other climate change. This has been shown global changes will greatly influence the through changes in phenology, such as forest ecosystem in Europe (IPCC 2007b). bud burst, in response to temperature (Menzel et al., 2006). Local adaptations are also evident within relatively short General consequences of climate change for timescales (e.g. Jump et al., 2006; Savola- forests inen et al., 2011). It is important to note that, while climate change is occurring and measurable in terms of physical and biotic changes, Range shifts there remains great uncertainty as to its Forest areas are expected to expand in magnitude and eventual impacts. Predic- the north and north-east but contract in tions based on current data and trends the south (Kremer et al., 2012; Metzger allow us to speculate on future events, et al., 2004; IPCC 2007b). Shifts in biocli- but predictions, by their nature, contain matic envelopes are predicted to change an element of uncertainty. It is difficult the distributions of species (Kremer et al., to make accurate predictions in particu- 2012). The loss in the south and gain in lar for extreme weather events, such as the north is both a challenge and an op- storms. That being said, based on the pre- portunity. It is likely that there will be

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extreme pressures on southern tree spe- lead to changes in the abundance and cies and populations to adapt in order to distribution of a number of tree species survive. As tree species are long lived, in this region, such as cork oak (Quercus extinction in the short-term (100 years) suber) and Aleppo pine (Pinus halepensis) is unlikely (except in cases of rare species (Schröter et al., 2005; Ruiz-Labourdette and those with restricted distributions et al., 2013). In addition to the changes and specialised habitats). However, there in species composition through replace- will be shifts in species’ ranges and tree ment, there may be new opportunities populations will have to adapt to new for hybridisation of formerly isolated physical conditions. The northern expan- species. Hybridisation is common in de- sion should occur through the spread of ciduous oak species and, with changes existing adaptations (e.g. temperate tree in ranges, new species interactions will species moving into tundra as it warms). occur. This can have an economic effect However, the southern tree populations if, for example, species restricted to the will have to adapt to new conditions, Mediterranean (such as Quercus fagin- such as unprecedented temperature in- ea and Q. fraineto) move northwards creases and drought. In addition to range and hybridize with the highly valuable shifts at the extremes, altitudinal shifts are temperate oaks (Q. petraea and Q. robur) also expected. Models predict that alpine (Valbuena-Carabaña et al. 2005; Kremer, species are extremely sensitive to climate 2010). change (Schröter et al., 2005). A particular concern will also be the fragmentation of a species range into disjointed popula- Competition and disease tions and this is predicted to occur more In addition to abiotic conditions new bi- in the southern and marginal popula- otic challenges, such as competition from tions. Fragmented populations will also other species and exposure to diseases, occur along with a northern expansion will negatively impact the survival of of populations. However, recolonization populations. It is suggested that forests through fragmented populations was of native conifers may be replaced by a feature of postglacial colonisation of deciduous trees in western and central many tree species (e.g. Lowe et al., 2006). Europe (Maracchi et al., 2005; Koca et al., 2006). Along with the displacement of species, there will be an exposure to Species composition new diseases and pathogens. As a con- As already illustrated from the changes sequence of climate change, many spe- in species ranges, species new to north- cies of phytophagous insects of northern ern latitudes will give rise to new species temperate forests, including some of the compositions and communities. Drought most important forest pests, have also in the Mediterranean region is likely to been observed to expand their range

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northward and upward, and this could which are likely due to the effects of affect the forest ecosystems through their climate change over the last 50-100 years, activity (Battisti 2008). Moreover, chang- have been observed in recent decades in ing climatic conditions can also influence Populus, Quercus, Pinus, Tsuga and Nyssa the spread of infectious pathogens such in the eastern United States (Johnson and as fungi and bacteria and add stress to Abrams, 2009). forest trees making them more susceptible (Kliejunas, 2011). This is an area that needs further study, and is certainly of interest in terms of the economic use of forests. Another Adaptation and changes in performance study comparing growth rates of mul- With the loss and expansion of species tiple Quercus species in Britain showed range edges there will be a subsequent species-specific responses, with southern loss and potential expansion of European (non-native) oaks performing locally adapted tree populations. The better than the native oaks (Sanders et al., adaptability of populations can affect 2014). This is of significance in terms of their performance. The productivity selecting future timber species but also in of forests in the Mediterranean region terms of species dynamics. is closely related to precipitation patterns, and drought in particular can decrease productivity (Cotrufo et Climate change consequences for forest al., 2011). However, in contrast to the genetic resources effects in southern populations, primary Forest genetic resources are fundamental to productivity and biomass will likely the diversity and adaptation of forest trees increase in northern populations due to and their populations. FGR are crucial to longer growing seasons (IPCC, 2007). A maintaining the adaptive capacity in re- review of the effects of climate change on sponse to climate change. The consequenc- primary productivity in forests globally es of climate change on FGR will be heavily shows a net increase in the primary dependent on the size and distribution of productivity since the middle of the 20th the current tree populations and indeed on century (Boisvenue and Running 2006). the biology of the species. A widespread The impacts on wood production and common species is less likely to be as sus- wood quality remain uncertain, although ceptible to the impacts of climate change experiments have shown changes in comparison to a species that has a nar- in wood chemistry (increased starch row distribution and small populations. compared to cellulose) and morphology The overall consequences are expected to (larger annual rings) in young birch be changes in allele frequencies and allele due to elevated CO2 (e.g., Kostiainen et compositions as a result of altered forest al., 2006). Changes in tree-ring growth, dynamics. Changes are also predicted at

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the edges of distibution ranges and within pansion. Studies on the genetic impacts of the core areas of the ranges. ice ages have shown a variety of patterns and also point to the lack of a “one size Despite the uncertain consequences of cli- fits all” approach (e.g. Lascoux et al., 2004). mate change on FGR, one likely outcome While current patterns of genetic structure is an overall reduction in variation at least are a result of dispersal after the last glacial in the short-term. Novel variation via spon- maximum and in particular of dispersal taneous mutation occurs continually in a into suitable territory that is unoccupied population and is acted on by natural or ar- (Hewitt 2000), a considerably different tificial selection to generate a better-adapt- pattern should be expected when species ed progeny. However, studies to date show ranges shift into occupied habitats. At the that the potential adaptation rate is much northern edge of the range founder effects slower than the pace at which the climate is from stochastic dispersal events resulting currently changing (Aitken et al., 2008) and in isolated populations are expected to in- that there is a prospect of an immediate re- fluence FGR (Hampe and Petit, 2005). As duction in our genetic resources in light of more of the landscape is currently occupied current climate change projections. by different habitats, the pattern is likely to continue in the form of a fragmented mo- Although there is a degree of plasticity in saic compared to the homogenisation and trees, natural selection of favourable traits admixture that occurred following the last will occur throughout the range, but this is glacial maximum. Isolated populations likely to be more intense at the range edg- can result in rapid adaptation to local con- es. This should be evident from changes in, ditions with a subsequent change in allele for example, allele frequencies of phenol- frequencies of the selected genes. Another ogy-related genes across clines. Data are element influencing and in some cases hin- currently limited on the effects of climate dering adaptation in the northern popula- change on natural selection (Donnelly et tions is prevalence of pollen from southern al., 2012). A change in allele frequencies populations dispersing northwards by pre- correlated with an altitudinal temperature vailing winds. This can affect the fitness of gradient has been shown for Fagus sylvatica the populations as the southern pollen is (Jump et al., 2006) and a growing number bringing genes adapted to different condi- of studies are investigating changes due to tions, which may not suit the northern envi- climate change. The responses are going ronment (Savolainen et al., 2007). However, to be very different at the different parts of in the long term this may be advantageous the range – the edges (northern/southern/ in adapting to a warming climate. In con- marginal) and the core area. trast, a species with limited seed dispersal will have a greater likelihood of local adap- In the north an important factor is the ge- tation (Kremer et al., 2012). netic landscape that is created from this ex-

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At the southern edge, extinction and frag- Assessment of threats mentation of populations is predicted with Just as predictions of climate change the subsequent consequences of genetic are uncertain (see pages 6-7), there is drift and restricted genepools. Southern also great uncertainty concerning the species will face increasing population biological response of tree species and fragmentation and reduction in population their populations and the resultant numbers. Populations in the south will impacts on FGR. To assess a threat a face unprecedented warmer conditions number of issues need to be taken into and they possibly do not have the neces- account. The IUCN uses population de- sary adaptation potential to survive these mographics and population ranges to changes (Aitken et al., 2008). Some of the assess the global threatened status of a southernmost populations are likely to be- species and this is also a useful model to come extinct, resulting in an overall reduc- apply on a regional basis (IUCN, 2012). tion in FGR. Potter and Crane (2010) have proposed a risk assessment system that includes Within the core of the distribution the most intrinsic and external factors. Intrinsic likely outcome will be a change in the spe- factors are demographic and ecolog- cies composition through changes in forest ical variables that are specific to the dynamics with a possible reduction in ge- species itself (e.g. population structure netic diversity. and density, fragmentation, regenerative capacity, dispersal ability, habitat Natural selection will of course be part of affinities and genetic variation). These a dynamic process where current climatic intrinsic factors are generally known envelopes of species are modified in the or can be estimated and can be used to north and south, but this is a more long- assess adaptive potential or potential term component of adaptation and indeed vulnerability. External factors (namely speciation. competition with other species, pest and pathogen threats and habitat pressures such as fire) are less easy to quan- Threatened tree species and populations tify and are therefore associated with a Threat occurs through three main pro greater degree of uncertainty. cesses: 1) direct elimination of individuals (e.g. overexploitation, grazing, pathogens, fire), 2) elimination of habitat (e.g. destruc- Adaptive potential tion, pollution, environmental change), 3) Threat to species or populations is a genetic impoverishment (e.g. by fragmen- multifaceted phenomenon. There are tation, genetic drift) (e.g. Potter and Crane, two natural possibilities to avoid the 2010). Climate change can influence all threat of extinction of a species, when three processes. climate change is one of the main drivers:

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adaptation and migration – both pollen have a comprehensive set of markers for and seed (e.g. Kremer 2007, Aitken et predicting adaptive potential. In many al., 2008). cases the only option is to use landscape and habitat variability within a species/ Adaptation responses comprise 1) population range as a proxy indicator plasticity (tolerance on the individual of adaptive potential. The adaptive level), 2) epigenetic processes (from responses of forest trees and ecosystems one generation to the next, e.g. to environmental changes and erosion Madlung, 2004) and 3) genetic variation of biodiversity has been studied within (evolutionary divergence and natural the EVOLTREE Network of Excellence selection at a population or species level). which was launched as a European In trees genetic differentiation in adaptive initiative in April 2006. Candidate genes traits is generally high. This indicates have been catalogued for phenological strong diversifying selection in the past and drought-related traits in important (Savolainen et al., 2007; Alberto et al., tree families such as Salicaceae, Fagaceae 2013). Recent studies generally observe a and Pinaceae (Kremer et al., 2011). high degree of plasticity for tree species as well as an element of epigenetic adaptive Migration responses involve evading processes, mostly in relation to limiting the threat through dispersal. Migration factors such as drought tolerance and is dependent on the species capacity in frost hardiness (Schueler et al. 2014) and terms of pollen and seed production or also in relation to phenology (Menzel et vegetative dispersal, the permeability of al., 2006). Adaptation capacity on all levels the landscape matrix to dispersal, and (plasticity, epigenetics, and genetics) the availability of suitable habitat to col- has the potential to influence species onise. While future migration away from persistence. A major issue with regard to temperature extremes in the south is gen- quantifying adaptive potential is the lack erally assumed to be predominantly lati- of data. In particular, the independence tudinal, it can also involve an altitudinal of neutral genetic variation and adaptive component where vertical buffers such as genetic variation (Holderegger et al., mountains exist. In fact, without vertical 2006) means that most of the available buffers, migration needs to be much fast- genetic data does not help to predict er to evade the threats (Loarie et al., 2009; adaptative potential. The development Ordonez & Williams, 2013). On the oth- of adaptive genetic markers coding for er hand, low-frequency episodic events ecological traits (by, for example, genome of long-distance dispersal can accelerate scanning and phenotype/genotype the populations’ migration responses association-studies) is ongoing (e.g. (Alberto et al., 2013b). In the particular Holderegger et al., 2008; Manel et al., 2010; case of species specialised to riverine Kremer et al., 2011) but we do not yet habitats, such as black poplar (Populus ni-

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gra), potential migration through natural Distribution of populations dissemination of seeds to more suitable Specific demographic and distribution areas is restricted because of their limited patterns provide general indications for capacity to migrate both in latitude and actual or potential threats. Differences altitude along the constrained range of in distributions will be a key factor in the riparian corridors (Villar et al., 2010). the susceptibility of a species to climate change. Therefore a rough estimation Predictions of ecological ranges from bi- and classification of specific distribu- oclimatic models show that a high pro- tion types of populations is necessary. portion of the area within a potential According to Frey (2003) populations distribution range of a species is not re- can be: a) in the main distribution zone alised by the species due to the influence (mainly conjunct populations), b) part of community ecology (e.g. competition of a side distribution zone (mainly dis- by established climax species and co-mi- junct populations), or c) within the relict grators, cooperation with seed vectors) zone (outside of the current distribution (e.g. Savolainen et al., 2007; Schueler et range). These categories are separate to al., 2014). In periods of environmental a species distribution and are particular- change all tree species are likely to suffer ly relevant when estimating impacts on lags in performance. Thus, interspecific tree populations or genetic conservation competition may be reduced, facilitat- units. These distribution types are impor- ing persistence under suboptimal condi- tant in assessing threat, as a population tions. Multi-species migration includes within the main distribution zone (the an immense variety of ecological interac- core) will probably be less threatened tions and thus leads to great uncertainty than those on the periphery of a range. (Kremer 2007, Aitken et al., 2008).

In general, rarity and low genetic varia- Estimating specific threat types tion are used as indicators of threat. How- The primary aim of the conservation ever, threat processes can lead to genetic approaches presented in this report is differentiation and thus local adaptation. to maintain adaptive variation across An evolutionary perspective on species the distribution range of forest trees in diversity shows that taxa are dynamic, Europe. Thus we have to focus on those not static. In some cases current divisions regions that will most likely be the most of subspecies and populations as distinct affected. These regions contain subpopu- species will probably not match future lations which host important intraspecific reality (e.g. white oak species in Europe, genetic variation. The focus on marginal Petit et al., 2002). and peripheral tree populations is being highlighted in a current EU initiative (COST Action FP1202 Strengthening con-

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servation: a key issue for adaptation of mar- species and correlates to high poten- ginal/peripheral populations of forest trees tial for long distance gene flow in trees to climate change in Europe (MaP-FGR)). (Hamrick, 2004; Alberto et al., 2013b). A special interest lies on marginal populations at the rear edges of species dis- In contrast, the rare and endemic tributions (Hampe and Petit, 2005). The Spanish fir (Abies pinsapo) in Southern IPCC (2007a) indicated that in Europe, Spain and northern Morocco is already the southern Iberian Peninsula, the Alps, showing signs of decline in genetic the eastern Adriatic and southern Greece variation (Liepelt et al., 2010). Another are those regions that will be most affect- example is the rare endemic Sicilian ed by climate change. The main threat in fir (Abies nebrodensis), which shows these areas is that of temperature increase low levels of neutral genetic variation and susceptibility to drought. It is these (Parducci et al., 2001). Due to its close areas that maintain populations at the relation to the European silver fir southern edge of the range of many spe- (Abies alba), the remaining population cies. Another important set of marginal of Sicilian fir has to be considered as a populations is that of islands. marginal population at the edge of the distribution range. A similar case is the Based on the IUCN threatened status Sicilian subpopulation of Turkey oak criteria of low population numbers, re- (Quercus cerris subsp. gussonei, locally duction in abundance and restricted dis- interpreted as Quercus gussonei, Sala et tributions (IUCN, 2012), marginal tree al., 2011). populations of some widespread species should be considered threatened. Low or declining abundance. This is Although the scale is different, the main the principal IUCN criterion for threat threat type indicators and thresholds are because this is relatively easy to evalu- similar for both threatened species and ate by demographic studies. Population threatened populations. It is possible to thresholds for minimal viable popula- assess a number of indicators for actual tions and a decline in population over and future threats. three generations or 100 years (for long living tree species) are taken into account (IUCN 2012). Indicators for general threats Low genetic variation. Compared to Decline can consist of dieback of adult herbaceous annuals and perennials, tree trees, such as drought-induced mor- species tend to have higher neutral ge- tality events at the southern edges of netic diversity within populations, while distribution (e.g. Atlas cedar (Cedrus genetic differentiation among popula- atlantica) dieback in Morocco and Al- tions is lower in more than 90% of tree geria, or Scots pine (Pinus sylvestris)

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dieback in Turkey) as well as in arid tation effects start when distances be- inner alpine valleys of Switzerland tween tree populations reach 50-100 km. (Allen et al., 2010). Other causes of die- Rare endemic species with limited dis- back can relate to introduced patho- tribution patterns often have a high lev- gens, e.g. cork oak (Quercus suber) de- el of fragmentation, such as Spanish fir cline by Pythophtora tinctoria infection (Abies pinsapo) with separation in differ- in southern Spain (Gómez-Aparicio, ent relict populations. Depending on the 2013). taxonomic interpretation of Trojan fir it is a single strongly isolated population Lack of regenerative capacity and (Abies equi-trojani) or a highly fragment- dispersal ability. Lack of regeneration ed marginal population of Turkish fir can also lead to declines in abundance. (Abies bornmuelleriana subsp. equi-trojani) Regeneration failure over time leads to or even of Caucasian fir (Abies nordman- unsustainable age structure in a stand niana subsp. equi-trojani). and inevitably to abundance decline. At the southern edge of distribution regeneration failure due to drought Indicators for threat triggered by climate change can occur long before adult trees are High susceptibility to drought stress. affected (highly susceptible youth There are numerous indications of in- growth phases). Another cause can be creases in the frequency of extreme overgrazing by red deer, as in the case drought events in recent decades, and of core populations of yew (Taxus bac- also of the high impact of drought on cata) and northern marginal popula- the decline of species and populations tions of the service tree (Sorbus domes- (e.g. Allen et al., 2010; Choat et al., 2012). tica) in Switzerland (Rudow, 2001), or The species which are most predisposed cattle in the case of Italian maple (Acer are those having marginal populations opalus subsp. granatense) in southern at the southern distribution edges (for Spain (Gómez-Aparicio et al., 2005). example Atlas cedar in Morocco and Al- geria or Scots pine in Central Anatolia, Seed set and dispersal ability are cru- Turkey). There are several variables cor- cial to the survival of populations and related with drought such as the aridi- species. These are both critically af- ty index, mean annual precipitation or fected by pollinators and dispersers. maximum temperature. Moreover, extreme drought events are often accom- High degree of fragmentation. Frag- panied and reinforced by disturbance mentation can increase vulnerabili- regimes such as existing or new insect ty and genetic erosion. Based on the pests (gradation of bark beetles, proces- long-distance gene flow in many tree sionary moths) or by wildfires (e.g. Al- species (Kremer et al., 2012), fragmen- len et al., 2010).

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Migration impediments. Physical fea- due to weak competitive capacity in tures of the landscape can act as barri- other areas (e.g. Pyrus pyraster in the ers to potential migration routes along Jura mountain range, Pinus heldreichii in zonal gradients. The Mediterranean sea the Balkans). is a barrier for northward shifts of North African or island populations of Medi- Another limitation to rare azonal habi- terranean species (e.g. Cedrus atlantica tats is related to specializations in wet- in Atlas mountains, Cedrus libani subsp. land sites. There are typical examples brevifolia or Quercus alnifolia in Cyprus) limited to moors (e.g. Pinus mugo subsp. or for east-west-shifts from one coast/ rotundata) or to alluvial forest (e.g. Ul- peninsula/islands to the other. mus laevis, Populus nigra, Alnus cordata and Platanus orientalis). Even if azonal Another barrier type consists of large flat wetland habitats are less susceptible to regions on continental plates for species drought pressure, migration is highly or populations already using mountain impeded, especially in moors and zones ranges as vertical buffers adjacent or with highly fragmented alluvial forests within these plains, e.g. the Castilla-La due to intensive watershed manage- Mancha in Central Spain, the Central ment and river regulation (Barsoum and Anatolia in Turkey, the Pannonian basin Hughes, 1998). in Hungary, and North-east European and Russian plains. Threat intensity Ecological factors. A number of eco- increases if vertical buffers are exhausted logical factors can be associated with (for example, Larix decidua and Pinus or exacerbated by climate change. Cli- cembra in the Carpathians). mate change has been implicated in the increase of invasive species, pathogens Limitation to azonal rare habitats. and diseases (e.g. Simberloff, 2000). A species or population limited to With changes in climate there will be rare azonal mountainous habitat can subsequent changes in species perfor- fragment and this can potentially lead mance, thus leading to inter-species to a threatened status (for example competition, not only with invasive al- most boreal tree species in Central ien species, but also with native species Europe, most deciduous tree species in that are favoured, for example, by drier the Mediterranean region). A different soils. Another particular case of ecolog- geology of mountain regions compared ical indicators will be the increased risk to the surrounding zones can have of fire in particular in southern region. similar effects. Many examples of This will favour those species that are secondary species are known, e.g. adapted to fire – such as seed germina- tree species, which are found only on tion after a fire. calcareous soils of mountain ranges,

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Overview of threatened species and populations preliminary list be further developed There is no red list focussing on and finalized as a separate EUFORGEN threatened European tree species, much activity in the future. Such a list can be less a list including tree populations developed using the criteria discussed threatened by climate change. The IUCN in the section above. World List of Threatened Trees (Oldfield et al., 1998) only provides fragmentary It is also important to consider those tree information concerning species in species which form part of the isolated Europe. The European Red List of Macaronesian flora (Laurisilva) in the vascular plants (Bilz et al., 2011) focuses adjacent Atlantic islands. The Laurisilvae on plants listed in policy documents, represent the last relicts of tertiary Med- crop wild relatives and aquatic plants. iterranean (Canary Islands, Madeira) or The lists from policy documents sub-Mediterranean (Azores) evergreen date to 1991 and earlier. Currently a broadleaves forest flora (Schäfer 2003). limited number of tree species that are The Macaronesian flora consists of many potentially vulnerable to climate change endemic species, which by nature of are ranked in the category ‘endangered’ their isolation and fragmentation have (e.g. Abies pinsapo, Cedrus atlantica); most to be considered as potentially threat- are still in the category ‘least concern’ ened or threatened species (e.g. several (e.g. Pinus sylvestris, Populus nigra, Alnus Laurus and Ilex species, Picconia excelsa, cordata, Pinus brutia, Pinus pinea, Platanus Ocotea foetens, Arbutus canariensis). orientalis, Juniperus thurifera) or ‘not assessed yet’ (e.g. Ulmus laevis, Abies alba subs. nebrodensis, Liquidambar orientalis, Knowledge of measures and existing Quercus alnifolia, Acer sempervirens) strategies (IUCN, 2014). An up-to-date overview of threatened and potentially threatened Background species and populations of European Conservation of FGR faces several trees, including a focus on marginal challenges, with climate change be- populations and adjacent regions of ing the latest. There is an urgent need Europe (Maghreb/North Africa, East to strengthen efforts among European Mediterranean/Near East, Caucasus/ countries to conserve FGR, particularly Alborz) is needed. This requires a those in marginal populations. Europe systematic evaluation of all tree taxa is a complex region where the distribu- across their distribution ranges. A tion ranges of tree species extend across preliminary list of threatened tree large geographical areas, with marked species and populations was developed environmental variation. As biological during the preparation of this report. distributions do not coincide with na- However, it is recommended that this tional boundaries, the regional cooper-

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ation among countries both in sustain- at species and landscape levels only, able forest management and in FGR while a better connection between forest conservation is crucial. genetic conservation networks and other biodiversity objectives is needed (Lefèvre et al., 2013). The discussion Incorporating FGR into national and EU policies on the integration of conservation National forest programmes are and use of FGR as part of sustainable important policy tools, which can forest management into these policies support the integration of FGR and strategies is underway within conservation into actions at the practical EUFORGEN. In the face of climate forest management level. National change, the European Information forest programmes are now in place System on Forest Genetic Resources in many European countries (Lefèvre, (EUFGIS), launched in September 2007; Rusanen et al., 2007; Hubert and 2010, has been created with the specific Cottrell, 2007; Graudal et al., 1995; Behm objective of supporting countries et al., 1997; Tessier du Cros, 2001). Within in these efforts. It could promote many programmes the conservation useful linkages between national focus is to maintain variability in FGR conservation programmes and adaptive traits (e.g. Graudal et al., 1995; biodiversity conservation efforts; it can Myking, 2002). Various conservation also support linkages between applied strategies have been revised based on conservation and research activities. the expected impacts of climate change but there is still much to be implemented in forest management practices. Forest Strategies managers are often unaware of the importance of using high-quality forest In situ approaches reproductive material and of the genetic consequences of management practices. Conservation networks. In view of the The appropriate use of FGR could predicted climate change, long-term con- indeed support the resilience of forests, servation of forest genetic diversity pro- mitigate the risks and facilitate the vides the insurance for sustainable forest adaptation of forests to climate change. management. The heritable genetic vari- In this regard, it would be worthwhile ation and the intensity of selection play a if conservation and use of FGR were critical role in the evolutionary response incorporated into national adaptation of species and populations to a changing strategies to climate change and national environment. For this the first priority biodiversity action plans. National of a conservation strategy should be to biodiversity action plans generally focus maintain high levels of diversity within on conservation of biological diversity the genetic conservation units and spe-

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cies across Europe (Koskela et al., 2007). ecological characteristics and their vulnera- Within EUFORGEN, priority is given to bility to climate change (see de Vries et al., a dynamic conservation of forest genetic 2015). resources in situ and ex situ. A pan-Europe- an network of selected genetic conservation Monitoring. Continuous monitoring is an units for various tree species has been cre- important element in a strategy to conserve ated according to pan-European minimum FGR. In particular, recent EUFORGEN ef- requirements and data standards of these forts to develop and undertake genetic units (Koskela et al., 2013). The aim of this monitoring could prove to be useful over network is to conserve adaptive genetic time. Field inventories and continuous diversity present across the range of Euro- monitoring incorporating ecological and pean conditions in which each tree species genetic data would provide a baseline to occurs. Geo-referenced data on genetic con- assess how well genetic diversity is actual- servation units based on 26 data standards ly conserved. This would show how well at the unit level (geographical area) and both adaptive and neutral genetic diversi- 18 data standards at the population level ty is maintained through time, and would (target tree species within a unit) has been reveal changes in species composition, the documented into the EUFGIS database. As occurrence of indicator species and com- of May 2015 the network consisted of 3,214 petitors of target species and in turn give units, with 4,061 populations of 100 tree an indication of the consequences of man- species. The EUFGIS database has been agement practices and/or environmental shown to be particularly useful in identify- changes (Aravanopoulos, 2011). The data ing gaps in the conservation network, thus generated by monitoring could be stored helping to create a more comprehensive set in the EUFGIS portal and used to revise of units for genetic conservation. management plans.

The conserved adaptive diversity of forest In situ management. Since there are threats trees throughout their distribution ranges to the integrity of FGR from a number of will provide the raw material by which causes and given the uncertainty of climate adaptation through natural selection will predictions, a comprehensive spectrum of occur during climate change. Using the possible conservation strategies and man- EUFGIS information system as a tool and agement measures, should be considered a climatic zoning of Europe (Metzger et al., in the context of climate change. 2013) as a proxy for characterizing adaptive diversity conserved in the genetic con- For most forest tree species, management servation units across the continent, a core plans for the conservation units allow network of dynamic conservation units silvicultural interventions directed can be selected for different species accord- towards the support of natural ing to their geographical distribution, their regeneration both in terms of quality

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and quantity of regenerating material. variation (higher initial diversity) and Applying given critical values for the promoting natural regeneration by number and density of seed trees, silvicultural interventions (tending, shortening of the regeneration time, thinning, selectively removing poor regulating competition by other tree quality individuals); 2) planting a species and controlling invasive species mixture of provenances alongside should also be taken into consideration. the current population using the A good level of genetic diversity and best climate predictions to guide the reduced consanguinity in the regenerated choice of provenances (using also a seedlings should be a management proportion of local stock might be objective. Natural regeneration should useful as a safeguard); and 3) using be the preferred means but if this fails to assisted migration by planting a single occur, assisted regeneration can be carried provenance or species for locations out using local seed lots to maintain local that are predicted to experience high phenotypic identity (Lefèvre, 2007). rates of climate change. This option will involve the transfer of genetic Three different strategies have recently resources from populations under been proposed to enhance resilience of particular climate pressures into areas forest stands to climate change in central predicted to have future conditions Europe (Bolte et al., 2009). This includes equivalent to the extant conditions. For 1) ‘Conservation of forest structures’ example, transferring drought-adapted by silvicultural intervention for older populations from the south to increase stands located in areas predicted to the adaptability of populations in the have low impacts from climate change; north, which would otherwise have 2) ‘active adaptation’ by thinning, an increased susceptibility to drought re-spacing and choice of alternative in the future (O’Neill et al., 2008). This species are proposed for stands where will be addressed in more detail in the the impacts are anticipated to be severe recommendations section of this report. and 3) ‘passive adaptation’ for stands of low value (ecological and economic) In the case of restoration programmes that will rely on natural evolution facing an evolutionary perspective should be the future climate shift. To help British considered (Rice and Emery, 2003). The forests to achieve higher levels of genetic exclusive use of local material may not diversity to accommodate climate allow for a rapid adaptation to the pre- change, another three potential strategies dicted climate scenarios (Harris et al., have been proposed using either natural 2006). However, to prevent introduction regeneration or planting and each has a of invasive forest pathogens, restrictions different mix of risks and benefits (Hubert on introductions should be developed. and Cottrell, 2007): 1) maintaining genetic The introduction of new species can

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negatively impact native biodiversity unknown genetic variability or charac- (extinction of local tree species, mat- terized genetically by phenotypic traits ing among invasive and native species, or molecular markers. They tend to be loss of stability among the native pop- expensive to establish and to maintain. ulation) (Chornesky and Randall, 2003; Dynamic conservation can take place in Cleeland and Mooney, 2001). Restora- ex situ stands when natural selection oc- tion and afforestation (ecological res- curs at a site and when artificially plant- toration) of degraded agricultural land ed trees (species, provenances, families) has also been considered an important can be regenerated from seeds without mitigation response to climate change. much intervention (Amaral, 2004). If the Afforestation and reforestation imple- original population is sufficiently sam- mented for carbon sequestration to mit- pled and the stand is large enough (min- igate global warming under the Kyoto imum viable population size) (FAO, Protocol could likely result in planta- 1992), these stands could provide sourc- tions with limited genetic diversity or es of reproductive material for commer- containing fast growing exotic species cial forestry. Although costly, multi-site that could have further negative impact stands can ensure further adaptations to on biodiversity (Caparros and Jacque- a range of different environmental con- mont, 2003). ditions and prevent unexpected losses of genetic material (Geburek and Turok, As a pre-emptive option in some cases, 2005). artificial in situ units can be created to serve as founder populations for new Assisted migration consists of trans- establishments, as in the case of riparian ferring species or populations from a species (e.g. Populus nigra) (Rotach, 2001). vulnerable site to a new site that is predicted to be more suitable under future Ex situ approaches climate projections. Migration can comprise long-distance transfers into more Dynamic ex situ. Conservation of FGR favourable regions or indeed bolstering is likely to become more complicated a threatened population with exter- with rapidly changing climate. There- nal genetic material. However, it can fore, actions for ex situ conservation also include shorter, more incremental will become increasingly important as a transfers, to support natural migration complement to, or substitute for, in situ via the establishment of migration cor- conservation (St. Clair and Hove, 2011). ridors and ‘stepping stones’. For exam- Generally defined as planted forests es- ple, isolated forest areas surrounded tablished outside the original habitat of by agricultural land in the plains could the genetic resources, ex situ conserva- be better connected through establish- tion stands may be genetic resources of ment of small stands or linear treelines/

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hedgerows to link them and to facilitate lumbia is the first jurisdiction to have more long distance gene flow. Prove- modified seed transfer guidelines spe- nance tests can be used to determine re- cifically in response to climate change sponse curves of seed sources to differ- (Loss et al., 2011a). ent climates. Examples including those based on Norway spruce (Picea abies), loblolly pine (Pinus taeda) and other Static ex situ. Seed orchards, clone southern pine species, which showed banks and clonal archives are examples reduced growth of between 5 and 10% of static ex situ conservation units, in compared to genetically adapted seed that no changes will naturally occur in sources (Schmidtling, 1994). Assisted the genetic structure of the collection. migration remains a controversial con- These plantings are established with the cept that evokes discussion within the sole purpose of preserving the genetic conservation community (McLachlan et diversity of a valuable population, to al., 2007). In Europe, the scientific com- safeguard endangered species that oth- munity agrees that for some species erwise might be lost or to conserve/in- growing in areas that will experience crease the genetic diversity of rare spe- most intense climate change, assisted cies of those with scattered distribution. migration offers a potentially cost effective strategy for climate-change mitiga- Collections of trees in arboreta or botanic tion, if used in addition to traditional gardens can also contribute to the main- conservation strategies and implemen- tenance of unique and rare genotypes tation of conservation genetics practices of different tree species, despite the fact (Loss et al., 2011a). Assisted migration or that in most cases these collections un- relocation is likely to be widely adopted der-represent the within-species genet- as a response strategy but it has not been ic variability due to the low numbers implemented yet through any policy of individuals that are present (Hurka, measures (Loss et al., 2011b). However, it 1994). Collections in botanic gardens, will likely have many problems such as although limited, are also useful as re- synecology (e.g. asynchrony or absence search resources and also for propaga- of specific pollinators, specific seed vec- tion (e.g. Kay et al., 2011). tors, specific mycorrhiza) and maladaptation (e.g. frost hardiness problems For species in imminent danger of ex- in northward migration). Guidelines tinction or with declining populations, for provenance transfer and seed zones which fail to produce or whose seeds are need to be adapted to account for north- recalcitrant, cryopreservation, vegetative ward migration of species and for pos- propagation and in vitro culture offer the sible assemblages in space and time only safe and cost-effective techniques (Hemery, 2007). In Canada, British Co- for their conservation, even if only a lim-

21 F G R C o n s e r v a t i o n a n d C l i m a t e C h a n g e

ited amount of genetic diversity can be the response of biota to it are dynamic. preserved (Theilade et al., 2004). In vitro For example, bioclimatic envelopes will conservation can be effectively used only change as the climate changes and species for collection and storage of particular adapt. As a consequence, species distri- genotypes of problematic species that butions and the genetic compositions of are propagated vegetatively (Engelman, the species will also change. Monitoring, 1997). in particular genetic monitoring, will be a key component of the conservation efforts. The dynamic nature of the change Implications for conservation and response needs to be built into future The conservation initiatives outlined in approaches to conserve FGR in Europe this report are dependent on a number of and beyond. In terms of changes in ge- logistical and technical issues. netic composition, it is important to know the different stakeholders’ views of con- The logistical issues are primarily those of servation. From a forest manager’s per- resources – cost and capacity. As conser- spective, the most appealing FGR benefit vation of FGR is a cross-border initiative, is that which gives rise to trees that pro- it will involve collaboration of state bodies duce good quality timber. However, from to bring about the effective conservation of a purely conservation point of view, the species and particularly threatened pop- adaptive capacity of a species takes pri- ulations into the future. The costs of as- ority over good quality timber. There is sessing threats, ground-truthing of units, a potential bias and resultant change in designating and managing units and allele frequencies depending on the selec- continual future monitoring need to be tion rationale. Thus, a focus on the adap- addressed at national and pan-European tive potential must be maintained when levels. As outlined in this report the south- conserving the units. The dynamic situa- ern tree populations are those most under tion should also be borne in mind when threat from climate change. However, the considering potential invasive species investment in conservation of these popu- and diseases. This is also important when lations may be key to the species survival dealing with populations at the vanguard as a whole and beneficial beyond the cur- of the species distributions. These popu- rent geography of these populations. lations will colonize new areas and possibly disrupt existing habitats. The technical issues are dependent on the biological reality of climate change It should also be recognised that it may and the response of tree populations and not be possible to save everything. For species. A matter of most obvious concern example, suitably sized populations is the dynamic nature of the conserva- are needed to maintain adaptive tion efforts needed. Climate change and variation. However, this may not be

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possible in small relict populations species with distribution restricted that have been subjected to extreme to the southernmost areas could face drought. It will therefore be necessary extinction. Assisted migration or ex situ to prioritise the units selected based conservation units in appropriate areas on limited resources. The focus should may be the only possible option for be on marginal and in particular, rear- dynamic conservation of such species edge populations. Populations at the and populations. rear-end of species distributions, and

23 F G R C o n s e r v a t i o n a n d C l i m a t e C h a n g e

24 M a n ag e m e n t o f C o n s e r v a t i o n U n i t s

Recommendations for management of genetic conservation units and establishment of additional Units

Any conservation strategy directed seed sources and genetic diversity. towards reducing the negative impact These units have been identified in of climate change should aim to reduce many countries, but only recently have the risk of poor performance or mass they been combined into a European- mortality. The uncertain nature of wide network. The units included climate change and the long generation in the EUFGIS database are selected intervals of forest trees put them at on the basis of a set of minimum particular risk for maladaptation to requirements that ensure populations climate change (St. Clair and Howe, are of sufficient size for inclusion in a 2007). Genetic diversity is critical for conservation network (Koskela et al., long-term forest sustainability as it 2013). However, the rationale behind provides the raw material for adaptation selecting conservation units must and natural selection. Forest managers remain fluid and should be periodically might consider options that aim to assessed in the context of climate change. maintain the existing genetic diversity Bioclimatic regions across European in forest stands or, in such cases where countries will undergo contraction or it is particularly impoverished, even to expansion of their ranges. In particular, increase it. To manage units effectively some will shift towards warmer and it is proposed to develop a decision drier conditions. Because the current cascade tool to guide forest managers to network of conservation units is based gauge suitable actions. An initial draft of on bioclimatic regions, a revision of the such a tool is provided in Appendix 1. network may be necessary in order to It is proposed to develop this through maintain the current design of one unit further EUFORGEN actions and to try per bioclimatic region per country (as and incorporate predictions based on per the pan European core network, current scientific knowledge. de Vries et al., 2015.). The addition of units specifically threatened by climate change addresses this in the short term. Management of existing genetic conservation units Genetic conservation units represent Genetic conservation units are the most valuable set of forest stands, populations considered to be particularly often containing remnants of the origi- valuable for the management of forest nal autochthonous populations of forest

25 F G R C o n s e r v a t i o n a n d C l i m a t e C h a n g e

trees. In line with the overall forest man- a useful framework in which to mon- agement goals for genetic conservation itor change. They can be used to assess units, forest management should pro- the progress of a population and can be mote natural processes. Such approach- used to determine levels of intervention, es should make the most use of man- such as thinning for increased regenera- agement techniques that mimic natural tion or removal of competing species. A disturbances, including the maximal use scheme for genetic monitoring is already of processes that induce, stimulate, and proposed by another EUFORGEN report support genetic diversity. In cases where (Aravanopoulos et al., 2015). natural regeneration does not occur, then artificial regeneration with a local Promote active management in the seed source should be used. units. After assessing the status of a population, the first option should be to The following criteria should be used to maintain the population in situ where aid a decision cascade which focuses on possible through active management. the maintenance of adaptive variation. Forest management should compensate for the effect of a changing climate until Monitoring of vitality and natural re- proper migration strategies are defined. generation. Monitoring is essential to determine the level of change that is oc- Favour units that contain altitudinal and curring. A set of parameters for assess- other ecological gradients. When selecting ing genetic risk has been suggested by units for the conservation network, priority Potter and Crane (2010). A modified ver- should be given to large units that contain sion of the list of parameters is present- significant altitudinal or ecological varia- ed in Table 1. These parameters provide tion. A larger altitudinal variation increas-

Intrinsic factors External factors Specific issues Particular to the species/ Outside of the “control” of population the species/population

Population structure/density Pest and pathogens Endemism

Regenerative capacity Competitive species Conservation status

Dispersal ability (seed/pollen) Invasive species Marginal/ range-edge populations

Habitat affinities Pollinators/dispersers

Genetic variation

Table 1. Parameters for assessing genetic risks (Potter & Crane, 2010, adapted)

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es variability of environmental conditions should be accepted that species/popula- and associated adaptive genetic diversity. If tions will be exposed to genetic changes a unit spans multiple habitats, for example that cannot be avoided (e.g., hybridiza- soil types, this is also more valuable than tions after secondary contacts), that some a more homogenous unit. For tree species losses of genetic diversity will be inev- with populations that are widely distrib- itable, and that the main objective of the uted but scattered, large units, involving conservation network is the conservation small groups of populations are preferable. of forest genetic resources, as opposed to In contrast, for those rare species which the conservation of the units per se. A re- only exist as small populations, conserva- view of the existing conservation network tion units can include a limited number of shows that once they have been com- trees in the core zone, which often repre- pleted by the countries involved, a good sents a sub-population that is available on coverage of the genetic resources from the site. the most important tree species in Europe is attained (de Vries et al., 2015). Howev- er, the pan-European core network has Establishment of additional genetic not been specifically designed to combat conservation units the risks to FGR associated with climate change and further actions are needed if Selection of genetic conservation units for we want the core network to retain large climate change vulnerability parts of the species genetic diversity in the The threats and uncertainties posed by near future (up to 2075 to 2100). Most ur- climate change upon the conservation gently, we should address the following: network make it necessary to incorporate several changes into the established criteria for the management of conservation Select additional units to represent marginal tree units, which have been reviewed above. populations Among those modifications, some will in- National authorities should pay particular volve changes in organization (e.g. to es- attention to marginal populations, as they tablish specific databases for conservation may contain specific adaptations to envi- units aimed at mitigating climate-change ronments that lie at the limits of the bio- effects, or whether to increase monitor- logical envelopes of each particular tree ing), while others may require analytical species. Southern marginal populations at changes (e.g. to apply an environmental/ the bioclimatic limits of the species distri- regional approach instead of the prevail- butions face unprecedented changes and ing focus on the species networks), and will most likely become extinct without others could be considered “experimen- intervention (Aitken et al., 2008). tal” due to uncertainties faced during the process. While applying such changes, it

27 F G R C o n s e r v a t i o n a n d C l i m a t e C h a n g e

Add units of newly established or migrated tree to be modified for the additional units in populations response to needs due to climate change. Long-distance dispersal events have a Either a relaxing of the criteria or a mod- large influence in the genetic make-up of ification to allow for particular conser- new populations colonizing the northern vation status may need to be included. limits of the species distribution areas Traceability of such relaxed conditions (LeCorre and Kremer, 2012). These rare might prove useful for future strategies. events create islands of related individu- Thus the network could be composed of: als with reduced genetic diversity, which will greatly contribute to colonizing more Core Units. Populations of autochtho- northerly territories. There is a risk of ac- nous origin, located within the main dis- cumulating deleterious mutations in these tribution range, in typical environments. populations (e.g. Peischl et al., 2013), thus supplementing these populations with Marginal Units. Populations of autoch- additional genetic material could help to thonous origin, located at the periphery overcome a potential genetic bottleneck. of the distribution range, in potentially atypical environments. These units could incorporate both single popula- Allow additional units to be added to the core tion and meta-population approaches network of conservation units in order to maintain (or increase) their The utility of new conservation units adaptive potential. specifically selected to address climate change will be greater if they are inte- Migrated Units. Populations of non-au- grated within the existing core network. tochthonous origin, located within the However, some kind of flag (new data- future predicted distribution envelope base field) may be necessary in order to of the species, typically the result of as- track such populations for any future sisted migration. strategy. This flagging would be important to identify the reasons why these units were chosen. A periodic revision Monitor the additional units of the country X zones core units will be Monitoring of the marginal and addi- necessary to adapt to future changes (see tional units may require additional ef- de Vries et al., 2015). forts by country authorities. For southernmost populations, the detection of early signs of decline might be vital if Core units versus additional units silvicultural regimes are intended to The additional units can be added as have any significant contributions to subsets of the core network. The concept populations’ adaptation. Furthermore, and criteria of the core network will need biotic interactions could pose additional

28 M a n ag e m e n t o f C o n s e r v a t i o n U n i t s

risks to genetic resources, even for those Species that are favoured by climate change populations that are well within the cli- In spite of threats to FGR, climate change mate envelopes (for example, if there is will favour the spread of some species (e.g. a lag between the migration of pests and Quercus ilex will likely increase its distribu- trees as a consequence of shorter gener- tion range in countries such as France). Even ation times). For northernmost marginal for such species, the rear-end and relict pop- populations, the detection of colonizing ulations should receive special attention in events may provide early clues into the order to avoid losses of valuable FGR. As most appropriate environments for fu- mentioned above, the front end will lose ture colonization. genetic diversity at least during the initial phases of expansion. However, there is the possibility that populations from different Exploring regional approaches to the glacial refugia could merge in some are- conservation of FGR as, creating new genetic diversity through Instead of the species-focused networks mixing of allopatric populations. In turn, in place, a regional approach could be this could create segregation of adaptive applied for some species, in particular traits leading to increased adaptation to cli- species complexes or taxonomically un- mate change. It is in such areas of genotypic certain groups, such as Pinus halepensis merging that the establishment of new con- or white oaks of the Mediterranean servation units might be most profitable. basin.

Review of scientific advances Taxonomic issues The recommendations and methods in It is important to note that species are this report need to be updated as appro- not static entities and that, for example, priate. A review of scientific advances hybridisation can lead to speciation. with a focus on the potential impact of The case of oaks in southern Europe il- climate change on conservation of FGR lustrates the potential for hybridisation should be implemented in the future. and formation of species. Hybridisation Such a review should facilitate communi- in certain systems could be assessed if cation between scientists and managers additional units are chosen in “hybrid for an early incorporation of new find- zones”. ings into the conservation policies.

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30 E x S i t u A pp r o a c h e s

Recommendations for complementary ex situ approaches

In some cases it will not be possible to Prioritization of tree species and maintain in situ units as the conditions establishment of ex situ populations will change too much to allow a tree population to survive. In addition, some Prioritization populations may hold particular vari- In order to optimize conservation efforts, ants or adaptive potential that need to the prioritization of species and the co- be maintained. In these cases, ex situ ordination of conservation measures on conservation measures will be neces- a pan-European level are necessary. The sary. As mentioned previously, the de- prioritization of species must consider cision cascade tool (Appendix 1) should 1) necessity of conservation according to help managers to determine when ex the importance of species (e.g. ecological situ approaches are most necessary. and economical values), 2) urgency of conservation according to the types and Ex situ conservation is defined here as intensities of threat (as described earli- “the conservation of components of bi- er), and 3) feasibility according to the ex- ological diversity outside their natural pected costs and impacts of measures in habitats” (see article 2 of the Convention relation to the available budgets. on Biological Diversity). Ex situ collections include whole plant collections, Ex situ measures are often closely relat- zygotes, gametes and somatic tissue. ed to the urgency of action. Having an Of course there are fundamental differ- overview of (potentially) threatened ences between whole plant collections, species and populations of European such as seed orchards and collections trees would allow earlier intervention such as cryopreserved seeds or embry- for ex situ measures. Such a systematic os. The preferred option is to maintain a and taxonomically synchronized “Red dynamic situation, one in which natural List” of threatened European tree species processes, such as gene flow and natu- and populations has to be established ral selection, are taking place. Static ex and maintained on a long-term basis. situ measures are those that instead of Suggestions have been made within this promoting a dynamic situation hold the report as to the criteria for establishing a material in stasis – such as cryopreserva- red list – those related to IUCN criteria tion of tissues. and also some specific to impacts pre-

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dicted based on climate change. During ecological distance of mother trees, d) the preparation of this report the au- number of harvest years. Seed collection thors found that a concurrent initiative protocols should aim to capture maximal is in place as part of the COST Action genetic diversity in artificial regeneration FP1202 to prepare a list of marginal in order to optimise the potential for ge- populations under threat. Adjacent re- netic and epigenetic adaptation. gions of Europe have to be taken into account, most of all North Africa, Near 2) Ex situ population size: the presence East and the Caucasus. These regions at of a large number of regenerating indi- the southern edge of the Mediterrane- viduals is crucial in order to provide the an climatic zone are most sensitive to best opportunity for natural selection species distribution shifts triggered by processes to operate. climate warming. If they lose their current tree species they will be exposed If possible, ex situ conservation units to desertification, because there are no should be established with at least 5,000 alternative more southerly tree species seedlings/saplings collected from pop- available to migrate northwards. ulations that contain at least 500 adult individuals. Such ex situ conservation units should be based on seed collected Establishment of ex situ populations across at least two seasons from at least As most tree species have high adaptive 50 spatially separated mother trees grow- potential (see section ‘Adaptive poten- ing across the range of ecological condi- tial’ on page 11) this feature is important tions within the site (Skrøppa, 2005). For to consider and harness in the context of scattered, rare and endangered species ex situ conservation measures. Therefore that have populations with low num- all measures dealing with artificial re- bers of trees these minimum standards generation, i.e. most of the ex situ meas- may have to be reduced. Alternatively, ures, but also assisted regeneration in in joining of fragmented subpopulations situ conservation units, must ensure that with neighbouring subpopulations in ex maximal variability of the original popu- situ units could be considered in order lation is captured by 1) using appropriate to achieve minimal viable population seed collection strategies and 2) establish- thresholds. ing optimally sized ex situ populations. As a general precaution on establishing 1) Seed collection strategies: best prac- units, the risk of introducing pests and tice in seed harvest takes into account pathogens should also be considered. and fulfils minimum requirements con- Since prevention and early detection are cerning a) population size, b) number of the most cost effective strategies, phy- mother trees, c) maximum spatial and tosanitary measures (post-entry quaran-

32 E x S i t u A pp r o a c h e s

tines and border controls for biosafety) practice are uncertain and bring with it should be taken seriously to safeguard associated risks and impacts. biodiversity (Clarke, 2008). In order to minimise unforeseeable implications, the establishment of ex situ Dynamic ex situ conservation conservation units originating from as- Dynamic ex situ conservation involves sisted migration should be performed the establishment of populations out- in gradual steps over relatively short side their natural habitats with the focus geographical and ecological distances. on facilitating and maintaining natural These distances should not exceed a regeneration in the population. In some maximum threshold equivalent to the cases it will not be possible to maintain change expected within the next 20-30 populations at their current location and years, or within 25% of one rotation time so assisted migration may be the only op- (e.g. O’Neill, 2013; Wang et al., 2010a tion for dynamic conservation (Thomas, and 2010b). Existing provenance trials 2011). Assisted migration can involve hu- in Europe could be studied to build a man-mediated movement of a popula- consensus of the potential suitability of tion via the introduction of a provenance assisted migration in various cases (e.g. or species into a new region or of facilitat- Alberto et al., 2013b). Depending on the ing natural migration. Within the context goal of the transfer, pure stands of in- of this report, assisted migration discus- troduced material can be set up where a sions will focus on movement of a pop- single unit is threatened and needs to be ulation beyond that which is expected to conserved, while mixed stands of intro- occur under short-term natural circum- duced and local material can be estab- stances. The focus is also on maintaining lished to encourage the natural process- genetic diversity within European tree es in other cases. species rather than introduced commer- cially important species. It is recommended to establish at least two ex situ units for each valuable conser- Assisted migration is considered by vation unit since the greatest effort is in some authors as a potentially important collection and growing of reproductive climate change adaptation strategy (Mil- material and the risk of losing one poplar et al., 2007; Campbell et al., 2009), but ulation can be high. Once a unit is dupli- should only be seen as a complementary cated, natural processes are still ongoing measure or as a last resort because gene- in the original unit, and these should be cology information is still lacking for safeguarded and monitored. most of the forest tree species (Eskelin et al., 2011) and the implications of this The use of static concepts such as seed zones and provenance regions will need

33 F G R C o n s e r v a t i o n a n d C l i m a t e C h a n g e

to be reviewed to ensure that managers uals in stasis, other static ex situ conser- have access to sufficient genetic variation vation measures should be taken, such in their planting stock. As forest repro- as preservation of individuals in botanic ductive material may need to be moved gardens, artificial populations in seed or- across national boundaries for planting chards and genotype collections or circa purposes seed transfer standards may re- situm conservation in cultural land and quire modification in response to climate settlement areas (e.g. Dawson et al., 2013). change to ensure that plantations have the potential to adapt to future environmental conditions. Adequate monitoring Future ex situ conservation units have to be included in the monitoring system in Static ex situ conservation measures order to assess their development and For some species the preferred dynamic viability under the conditions at the cho- conservation approach will not be possi- sen sites. It is suggested to have a par- ble and in these cases, the only alterna- ticular “flag” in the EUFGIS database tive is to maintain the species in stasis to identify ex situ units. Particular focus in a seed bank or gene bank (e.g. cryo- should be placed on the genetic moni- preservation). In general, the earlier the toring of the migration units. Ex situ dy- intervention, the more effective it is and namic units will give valuable data on in most cases also the more economical. future performance of species. Parallel to the preservation of individ-

34 R e c o m m e n d a t i o n s f o r R e s e a r c h

Recommendations for research

It is vital to increase the current scientif- significant amount of provenance trials ic understanding regarding the impacts that have been conducted throughout and adaptability of tree species and pop- Europe. These can be used to assess ulations to climate change. Although potential responses of species to large the recommendations in this report pri- scale movement. Through further marily deal with management issues, it research we can better estimate impacts is important to highlight some research or future responses. gaps. Little is known about the impacts of climate change on genetic diversity, The second area highlighted in this re- with only a few studies to date showing port is that of marginal and peripheral concrete effects. Aside from individual populations. These populations need to species and plant community responses be identified and characterised to enable to environmental changes, two particu- predictions of future threatened regions. lar areas are highlighted in this report. Research should be carried out to determine response and impacts within these Firstly, there is the lack of knowledge marginal populations. Various types of surrounding assisted migration marginality, including altitudinal mar- and its potential impacts on genetic ginality, are particularly important in composition and species composition. populations within the core distribution One obvious source of data is the of a species range.

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36 C o n c l u s i o n s a n d R e c o m m e n d a t i o n s

Conclusions and overall recommendations

Conclusions Recommendations Genetic diversity of forest trees in Eu- • European countries should continue rope is likely to be reduced as a result of populating the EUFGIS database in climate change. This review shows that order to identify gaps in regions and marginal populations in southern areas species. are particularly at risk. The most likely • Add additional units to EUFGIS threats are drought and increased tem- specifically flagged as FGR for cli- perature in southern Europe. Climatic mate change units. A particular fo- changes are likely to result in altered cus should be placed on marginal species ranges and species composition. populations and particularly those The species biology will determine to with known limitations in terms some extent the level of impact of cli- of ecological plasticity or genetic mate change on the populations; for diversity. example, species with large popula- • A resolution from FOREST EUROPE tions and broad geographic ranges are is needed regarding the movement expected to be less affected than those of the material between countries with small populations and limited ge- for conservation purposes. ographic ranges. • The area covered by pan-Europe- an collaboration on FGR should be The consensus is that the current pan-Eu- enlarged to include, for example, ropean network was not specifically de- Macronesian areas and other areas signed to conserve FGR under climate with high endemism. These areas change. Thus, additional measures will harbour high levels of endemism be needed to conserve those FGR that and unique genotypes. are most threatened, e.g. small marginal • Collaboration with North African populations living under extreme climat- countries should be sought on com- ic conditions. A number of approaches mon species. It is likely that com- were reviewed, but a key point is that mon species will currently exist there is very little empirical data availa- under more “advanced” climatic ble. Thus, best practice can only be based conditions in North Africa in com- on predictions and estimates. parison to Europe.

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• A decision cascade tool should be • More research is needed to assess further developed and used in the the potential effects and impacts of future. The tool presented within this assisted migration. Existing prove- report is an exemplar rather than a nance trials can be used as a starting finished product. This could be pro- point, but further research is nec- gressed in the form of a particular essary to understand the potential EUFORGEN initiative. adaptation of trees to future climate • The development of species and pop- change scenarios. ulation/region red lists should be • Research is also needed to assess considered as a EUFORGEN initia- the impacts of climate change on tive. A number of examples are given marginal and peripheral popula- within this report, but this needs to tions, in particular in relation to their be more comprehensively reviewed adaptability. and used as a continual work-in-progress. Part of this is being undertak- en by the COST Action FP1202.

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Appendix 1

POTENTIAL DECISION CASCADE FOR GENE CONSERVATION UNDER CLIMATE CHANGE: MEASURES, INDICATORS (vers.1.2: Rough draft to give an idea of the recommended decision cascade, detailed indicators for measures have to be elaborated)

o = obligatory indicator (AND); xx/yy/zz = alternative indicators (OR); >%/>>%/>>>%= level of population decline %

Level/ Measures Indicators Step - - - Relevant forest genetic Relevant forest resource (national data) Conservation unit minimum (EUFGIS, requirements Koskela et al 2013) network Criteria for core et al 2015) (deVries Criteria for conservation unit genetic monitoring (Aravanopoulos et al 2015) Criteria for complementary genetic monitoring of the units (this report) Relatve number of repro declining (% ducing trees per 10 years) Absolute number of repro declining under ducing trees < minimum requirement Specific pest, invasive neophyte over Lack of regeneration > 10 years Migration barriers, exhaust ed vertical buffers for genetic High probability drift Danger of extinction 0 general prevention from habitat destruc- o tion (or rehabilitation) restriction against introduction o of pest, invasive neophyte etc. 1. in situ conservation in genetic conservation units unit establishment o EUFGIS record o unit demographic monitoring o unit included in core network o unit target of genetic monitoring x x 2. in situ silvicultural measures in genetic conservation units regulation of competition or o >% y y pathogenes artificial regeneration o o 3. in situ replacement/reorganisation of genetic conservation units replacement by existing equiva- o o >> % y y o lent unit within country x zone duplication of unit within o o >> % y y o surroundings/habitat recombining duplicates of similar/near unit within o o >>> % y y o o surroundings/habitat 4. ex situ assisted migration of genetic conservation units duplication of unit in direction x x >>> % y y o z z of expected change (2x) recombining duplicates of unit in x x >>> % y y o o direction of expected change (2x) 5. ex situ preservation in field genotype collections in conservation orchards, botanical garden x x o o o networks 6. ex situ preservation in stasis seed bank, cryoconservation, in x x o o o o vitro conservation

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46 EUFORGEN