To learn more about the map elements, please download the "Pan-European strategy for genetic conservation of forest trees"
This distribution map has been developed by the European Commission Joint Research Centre (partly based on the EUFORGEN map) and released under Creative Commons Attribution 4.0 International (CC-BY 4.0)
Caudullo, G., Welk, E., San-Miguel-Ayanz, J., 2017. Chorological maps for the main European woody species. Data in Brief 12, 662-666. DOI: https://doi.org/10.1016/j.dib.2017.05.007
The following experts have contributed to the development of the EUFORGEN distribution maps:
Fazia Krouchi (Algeria), Hasmik Ghalachyan (Armenia), Thomas Geburek (Austria), Berthold Heinze (Austria), Rudi Litschauer (Austria), Rudolf Litschauer (Austria), Michael Mengl (Austria), Ferdinand Müller (Austria), Franz Starlinger (Austria), Valida Ali-zade (Azerbaijan), Vahid Djalal Hajiyev (Azerbaijan), Karen Cox (Belgium), Bart De Cuyper (Belgium), Olivier Desteucq (Belgium), Patrick Mertens (Belgium), Jos Van Slycken (Belgium), An Vanden Broeck (Belgium), Kristine Vander Mijnsbrugge (Belgium), Dalibor Ballian (Bosnia and Herzegovina), Alexander H. Alexandrov (Bulgaria), Alexander Delkov (Bulgaria), Ivanova Denitsa Pandeva (Bulgaria), Peter Zhelev Stoyanov (Bulgaria), Joso Gracan (Croatia), Marilena Idzojtic (Croatia), Mladen Ivankovic (Croatia), Željka Ivanović (Croatia), Davorin Kajba (Croatia), Hrvoje Marjanovic (Croatia), Sanja Peric (Croatia), Andreas Christou (Cyprus), Xenophon Hadjikyriacou (Cyprus), Václav Buriánek (Czech Republic), Jan Chládek (Czech Republic), Josef Frýdl (Czech Republic), Petr Novotný (Czech Republic), Martin Slovacek (Czech Republic), Zdenek Špišek (Czech Republic), Karel Vancura (Czech Republic), Ulrik Bräuner (Denmark), Bjerne Ditlevsen (Denmark), Jon Kehlet Hansen (Denmark), Jan Svejgaard Jensen (Denmark), Kalev Jðgiste (Estonia), Tiit Maaten (Estonia), Raul Pihu (Estonia), Ülo Tamm (Estonia), Arvo Tullus (Estonia), Aivo Vares (Estonia), Teijo Nikkanen (Finland), Sanna Paanukoski (Finland), Mari Rusanen (Finland), Pekka Vakkari (Finland), Leena Yrjänä (Finland), Daniel Cambon (France), Eric Collin (France), Alexis Ducousso (France), Bruno Fady (France), François Lefèvre (France), Brigitte Musch (France), Sylvie Oddou-Muratorio (France), Luc E. Pâques (France), Julien Saudubray (France), Marc Villar (France), Vlatko Andonovski (FYR Macedonia), Dragi Pop-Stojanov (FYR Macedonia), Merab Machavariani (Georgia), Irina Tvauri (Georgia), Alexander Urushadze (Georgia), Bernd Degen (Germany), Jochen Kleinschmit (Germany), Armin König (Germany), Armin König (Germany), Volker Schneck (Germany), Richard Stephan (Germany), H. H. Kausch-Blecken Von Schmeling (Germany), Georg von Wühlisch (Germany), Iris Wagner (Germany), Heino Wolf (Germany), Paraskevi Alizoti (Greece), Filippos Aravanopoulos (Greece), Andreas Drouzas (Greece), Despina Paitaridou (Greece), Aristotelis C. Papageorgiou (Greece), Kostas Thanos (Greece), Sándor Bordács (Hungary), Csaba Mátyás (Hungary), László Nagy (Hungary), Thröstur Eysteinsson (Iceland), Adalsteinn Sigurgeirsson (Iceland), Halldór Sverrisson (Iceland), John Fennessy (Ireland), Ellen O'Connor (Ireland), Fulvio Ducci (Italy), Silvia Fineschi (Italy), Bartolomeo Schirone (Italy), Marco Cosimo Simeone (Italy), Giovanni Giuseppe Vendramin (Italy), Lorenzo Vietto (Italy), Janis Birgelis (Latvia), Virgilijus Baliuckas (Lithuania), Kestutis Cesnavicius (Lithuania), Darius Danusevicius (Lithuania), Valmantas Kundrotas (Lithuania), Alfas Pliûra (Lithuania), Darius Raudonius (Lithuania), Robert du Fays (Luxembourg), Myriam Heuertz (Luxembourg), Claude Parini (Luxembourg), Fred Trossen (Luxembourg), Frank Wolter (Luxembourg), Joseph Buhagiar (Malta), Eman Calleja (Malta), Ion Palancean (Moldova), Dragos Postolache (Moldova), Gheorghe Postolache (Moldova), Hassan Sbay (Morocco), Tor Myking (Norway), Tore Skrøppa (Norway), Anna Gugala (Poland), Jan Kowalczyk (Poland), Czeslaw Koziol (Poland), Jan Matras (Poland), Zbigniew Sobierajski (Poland), Maria Helena Almeida (Portugal), Filipe Costa e Silva (Portugal), Luís Reis (Portugal), Maria Carolina Varela (Portugal), Ioan Blada (Romania), Alexandru-Lucian Curtu (Romania), Lucian Dinca (Romania), Georgeta Mihai (Romania), Mihai Olaru (Romania), Gheorghe Parnuta (Romania), Natalia Demidova (Russian Federation), Mikhail V. Pridnya (Russian Federation), Andrey Prokazin (Russian Federation), Srdjan Bojovic (Serbia) , Vasilije Isajev (Serbia), Saša Orlovic (Serbia), Rudolf Bruchánik (Slovakia), Roman Longauer (Slovakia), Ladislav Paule (Slovakia), Gregor Bozič (Slovenia), Robert Brus (Slovenia), Katarina Celič (Slovenia), Hojka Kraigher (Slovenia), Andrej Verlič (Slovenia), Marjana Westergren (Slovenia), Ricardo Alía (Spain), Josefa Fernández-López (Spain), Luis Gil Sanchez (Spain), Pablo Gonzalez Goicoechea (Spain), Santiago C. González-Martínez (Spain), Sonia Martin Albertos (Spain), Eduardo Notivol Paino (Spain), María Arantxa Prada (Spain), Alvaro Soto de Viana (Spain), Lennart Ackzell (Sweden), Jonas Bergquist (Sweden), Sanna Black-Samuelsson (Sweden), Jonas Cedergren (Sweden), Gösta Eriksson (Sweden), Markus Bolliger (Switzerland), Felix Gugerli (Switzerland), Rolf Holderegger (Switzerland), Peter Rotach (Switzerland), Marcus Ulber (Switzerland), Sven M.G. de Vries (The Netherlands), Khouja Mohamed Larbi (Tunisia), Murat Alan (Turkey), Gaye Kandemir (Turkey), Gursel Karagöz (Turkey), Zeki Kaya (Turkey), Hasan Özer (Turkey), Hacer Semerci (Turkey), Ferit Toplu (Turkey), Mykola M. Vedmid (Ukraine), Roman T. Volosyanchuk (Ukraine), Stuart A'Hara (United Kingdom), Joan Cottrell (United Kingdom), Colin Edwards (United Kingdom), Michael Frankis (United Kingdom), Jason Hubert (United Kingdom), Karen Russell (United Kingdom), C.J.A. Samuel (United Kingdom).
Status of Pinus sylvestris conservation in Europe
The species has a lot of genetic differences within populations, but not as many between populations (Vasilyeva et al., 2021). This is to be expected for a species living in many different environments over a wide range and with easy gene flow (wind-dispersed pollen), making populations of trees more genetically similar. This is different from species that prefer specific habitats and are spread out, which can have larger genetic differences between their populations. Small differences between populations may be due to widespread commercial use in Europe's forests, where commercially valuable trees are chosen, making populations more homogeneous (Vasilyeva et al., 2021; Danusevicius et al., 2023).
Analysis of populations of trees in the Apennine mountains found they are genetically distinct to populations in the Alps or other places, even though they have less genetic variation because of isolation and small population size (Scalfi et al., 2009). This might be because the northern Apennines may have been an additional Pleistocene refugial area. These populations might allow a greater variety of genes to be conserved, as they show greater variation between populations than expected for the species, due to their higher fragmentation (Scalfi et al., 2009; Danusevicius et al., 2023).
The bibliographic review was conducted by James Chaplin of the EUFORGEN Secretariat in August 2024.
Current patterns of genetic distribution may be due to the last glacial maximum (LGM) in Europe, which pushed Scots pine to glacial refugia in Spain, Italy, and Türkiye (Dering et al., 2017; Sannikov et al., 2020). The currant distribution of the species in northern Europe is the result of postglacial dispersion and colonization from these Pleistocene refugia (Dering et al., 2017; Sannikov et al., 2020). Genetic structure today is mainly due to this movement from the south to the north (Dering et al., 2017). Decreased genetic structure north and eastward may be because colonization after the LGM in northern Europe occurred from a single mitotype from select refugia.
Research on populations that stayed in glacial refugia, such as in Italy, reveals that they have less genetic variety than the northern European populations but are genetically distinct from northern populations and each other (Dering et al., 2017). Trees in the warmer south might not have as much genetic variety because they have smaller populations; however, these populations are genetically distinct from northern populations and each other, thus possessing greater genetic diversity overall (Scalfi et al., 2009; Wachowiak et al., 2010). Some data suggests there are five main genetic groups of Scots pine, located in east and central Europe, Finland, Iberia, Italy, Türkiye, and a widespread group across Europe (NIBIO et al., 2020).
The bibliographic review was conducted by James Chaplin of the EUFORGEN Secretariat in August 2024.
When selecting genetic conservation units for Scots pine along a cline, ecological information should be preferred over neutral markers. Genetic conservation units should be large enough to buffer against outside geneflow, and nearby genetically degraded stands should be avoided or removed (Mátyás, Ackzell, and Samuel, 2004). For isolated groups of trees, a starting area of 10 hectares is enough, and it can be made bigger in the future with successive regenerations. The species requires human interaction to prevent ecological succession, and artificial regeneration is needed in most conditions; however, natural regeneration should be used as much as possible. In these cases, cones from 50 or more trees (with an equal quantity of seed from each tree) should be collected to sample genetic resources properly (Mátyás, Ackzell, and Samuel, 2004). Direct sowing should be preferred to planting, and selective removal of trees should be confined to malformed individuals. Priorities for specific genetic conservation measures will differ regionally and should be visualized in the context of locally applied forest management practices.
The bibliographic review was conducted by James Chaplin of the EUFORGEN Secretariat in August 2024.
Genetic Characterisation of Pinus sylvestris and its GCUs
Availability of FRM
Pinus sylvestris - Technical guidelines for genetic conservation and use for Scots pine
Publication Year: 2003Conservation priorities
Because Scots pine is a species with an extremely wide distribution and occupying a broad range of habitats, genetic conservation seems to be a task of low priority. However, the need to address genetic resources of Scots pine is supported by the widely proven genetic diversity between populations, the effects of century-long cultivation and the expected environmental changes at the margins of the distribution.
As Scots pine is one of Europe’s most important tree species under forest management, the anthropogenic influence is obvious. Both survey and recording of native local (autochthonous) stands are important for gene conservation. These records could include various identification data; molecular markers become increasingly useful for this task.
Long-term provenance tests have proved the value and importance of locally adapted populations. This is valid primarily for extreme site conditions (higher altitudes, coastal environments, extreme boreal conditions, rocky or semiarid sites). Preserved populations on these sites exhibit less plasticity when transferred to other conditions, but are usually superior locally. Special care should be taken, therefore, to select representative populations for conservation on such sites. Native stands selected for gene conservation will also serve as ‘population standards’ when compared with man-made forests.
As with populations on extreme sites, isolated outliers might have been exposed to specific selection pressures or drift and may carry rare alleles. Such populations should be carefully protected and steps taken to collect forest reproduction material at the sites. Local material should be used for regeneration and material from endangered sites should also be established in ex situ conservation stands.
Expected climate change will first affect the populations at the southern fringes of the distributional range. These populations are often remarkably vigorous and tolerant and may be of value for future breeding. Here also ex situ measures should be applied to safeguard long-term survival.
The long tradition of artificial regeneration may have developed landraces that could also be targets for gene conservation efforts. These populations usually represent diverse, rather plastic genetic resources, valuable for future breeding and reproduction.
Establishment and management of gene conservation units
When selecting gene conservation units along a continuous cline, ecological information should be preferred to neutral markers. In the absence of drift, in a contiguous distribution range adaptively different populations may be expected at distances where annual mean temperature differs by a minimum of 1.0–1.5°C (equal to ca. 200 km in a flat landscape).
The size of gene conservation units of Scots pine should be sufficiently large to compensate for and buffer against outside geneflow: 100 ha should be considered the minimum. Nearby occurrences of genetically degraded or otherwise unsuitable stands should be either avoided or removed. A conservation unit should consist of numerous adjoining stands of various age, provided their origin is the same. In areas of scattered occurrence, initial size may be 10 ha as a minimum, which can be increased during successive regenerations.
In many instances the pioneer character of Scots pine demands human interaction to prevent ecological succession. As far as possible, natural regeneration should be applied; this is less problematic on drier or poorer sites. Regeneration of admixed species should be tolerated for ecological reasons. The light demand of the species does not allow the development of a very complex stand structure, but this is not necessary as evenaged stands may hold equal diversity. Regeneration felling should be carried out stepwise, allowing for recruitment from numerous seed years. Scots pine is genetically rather insensitive to the type of regeneration cutting used. However, if the influx of outside pollen were minimized (a goal which can be met with only partial success), shelterwood cutting would be preferred to other regeneration regimes. Fencing of the unit has to be considered where high game density threatens natural regeneration processes.
In certain cases artificial regeneration may be necessary (e.g. for ex situ conservation). To sample genetic resources properly, cones from 50 or more well-distributed trees should be collected (preferably in a good seed year). The quantity of seed from each tree must be equal in order to get a balanced participation in the final seed lot. Mixing of repeated seed harvests is beneficial, and no seed sorting or grading must be applied.
Direct sowing should be preferred to planting. If possible, planting should be carried out with higher density than usual to allow for more natural selection.
Intermediate low-intensity fellings and management should maintain a relatively dense stand structure. Selective removal of trees should be confined to malformed individuals; otherwise a broad variation of phenotypes should be allowed.
In summary, priorities for specific gene conservation measures will differ regionally. Preservation of genetic resources of Scots pine should be visualized in the context of locally applied forest management practices (especially control of seed sources for artificial regeneration), the extent of protected or unmanaged areas and the occurrence, density or fragmentation of the species at the landscape level, together with actual threats and risks. The urgency to set up gene conservation units will be much higher in an area with fragmented remnants of local populations surrounded by planted forests of uncontrolled origin than in a region where sustainable forestry relying on natural regeneration and local seed sources is practised.
Contacts of experts
NA
Further reading
Wójkiewicz, B., Cavers, S., and Wachowiak, W. 2016. Current approaches and perspectives in population genetics of Scots pine (Pinus sylvestris L.). Forest Science, 62(3): 343–354.
Donnelly, K., Cottrell, J., Ennos, R.A., Vendramin, G.G., A'Hara, S., King, S., Perry, A., Wachowiak, W., and Cavers, S. 2017. Reconstructing the plant mitochondrial genome for marker discovery: a case study using Pinus. Molecular Ecology Resources, 17(5): 943–954.
Wachowiak, W., Żukowska, W.B., Perry, A., Lewandowski, A., Cavers, S., and Łabiszak, B. 2023. Phylogeography of Scots pine in Europe and Asia based on mtDNA polymorphisms. Journal of Systematics and Evolution, 61(2): 315–327.
References
Danusevicius, D., Rajora, O.P., Kavaliauskas, D., Baliuckas, V., and Augustaitis, A. 2023. Genetic diversity and fine-scale spatial genetic structure of unmanaged old-growth versus managed second-growth populations of Scots pine (Pinus sylvestris L.) in Lithuania. European Journal of Forest Research, 142: 773–793.
Dering, M., Kosiński, P., Wyka, T.P., Pers-Kamczyc, E., Boratyński, A., Boratyńska, K., Reich, P.B., Romo, A., Zadworny, M., Żytkowiak, R., and Oleksyn, J. 2017. Tertiary remnants and Holocene colonizers: Genetic structure and phylogeography of Scots pine reveal higher genetic diversity in young boreal than in relict Mediterranean populations and a dual colonization of Fennoscandia. Diversity and Distributions, 23(5): 540–555.
Mátyás, C., Ackzell, L., and Samuel, C.J.A. 2004. EUFORGEN technical guidelines for genetic conservation and use for Scots pine (Pinus sylvestris). Maccarese, Italy, Bioversity International.
NIBIO and WSL, INRAE, AUTH, UU, CSIC, Biodiversity, CNR, Skogforsk, INIA, NERC, THUNEN, ASP., 2020. GenTree, Optimizing the management and sustainable use of forest genetic resources in Europe, Deliverable D1.5 Report characterizing the genetic diversity of the European Conservation Network and monitoring strategies. rep. Research and Innovation action: GA no. 676876, pp. 47–53.
Sannikov, S.N., Petrova, I.V., Egorov, E.V., and Sannikova, N.S. 2020. Searching for and revealing the system of Pleistocene refugia for the species Pinus sylvestris L. Russian Journal of Ecology, 51: 215–223.
Scalfi, M., Piotti, A., Rossi, M., and Piovani, P. 2009. Genetic variability of Italian southern Scots pine (Pinus sylvestris L.) populations: the rear edge of the range. European Journal of Forest Research, 128: 377–386.
Vasilyeva, Y., Chertov, N., Nechaeva, Y., Sboeva, Y., Pystogova, N., Boronnikova, S., and Kalendar, R. 2021. Genetic structure, differentiation and originality of Pinus sylvestris L. populations in the east of the East European Plain. Forests, 12(8): 999. https://doi.org/10.3390/f12080999
Wachowiak, W., Salmela, M.J., Ennos, R.A., Iason, G., and Cavers, S. 2011. High genetic diversity at the extreme range edge: nucleotide variation at nuclear loci in Scots pine (Pinus sylvestris L.) in Scotland. Heredity, 106(5): 775–787.