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 cembra conservation in Europe
Swiss stone pine shows high polymorphism, indicating high genetic diversity across its range (Gugerli, Rüegg, and Vendramin, 2009). Some research revealed around 93% of genetic variation in Swiss stone pine is within populations, while some natural Swiss stone pine populations in the Alps showed an excess of homozygotes, possibly indicating self-fertilization and mating between related trees (Teodosiu, 2007). However, Swiss stone pine typically shows substantial genetic variation and low inbreeding (Tóth et al., 2019).
Populations in the Alps and Carpathians showed low genetic differentiation between each other; however, Carpathian populations showed a higher level of genetic diversity (Ulber, Gugerli, and Bozic, 2004; Tóth et al., 2019). Across the species range there is low differentiation between populations and no clear differentiation between northern and southern populations (Teodosiu, 2007). However, there is some evidence of regional-scale genetic structuring (Tóth et al., 2019).
Altitudinal variation in growth characteristics has been observed in Swiss stone pine (Ulber, Gugerli, and Bozic, 2004). However, climatic heterogeneity and geographic and genetic isolation are key drivers of differentiation within marginal populations (Tóth et al., 2019).
Gene exchange between populations of Swiss stone pine is heavily dependent on seed dispersal by birds such as the nutcracker (Nucifraga caryocatactes L.) (Teodosiu, 2007). At high elevations, seed dispersal by the nutcracker and Swiss stone pine’s competitiveness ensures preservation of the species gene pool, while at lower altitudes the limited activity of nutcrackers in dense forests may result in reduced genetic diversity in Swiss stone pine (Teodosiu, 2007).
Swiss stone pine has heavy seeds and strong habitat requirements to be competitive, which limits its seed dispersal and ability to colonize new environments; this can also cause bottlenecking in isolated and marginal populations, thus reducing genetic diversity (Tóth et al., 2019).
The bibliographic review was conducted by James Chaplin of the EUFORGEN Secretariat in August 2024.
Marginal and isolated populations such as those in the northern margin of the Swiss Alps have lower genetic variation and higher structuring and differentiation than populations in the central Swiss Alps (Ulber, Gugerli, and Bozic, 2004; Tóth et al., 2019). This differentiation is due to their isolation and the more-varied climatic environments they are found in (Tóth et al., 2019). Fragmented populations of Swiss stone pine are often associated with weak gene flow, genetic drift, and higher genetic differentiation (Tóth et al., 2019).
Swiss stone pine survived in multiple glacial refugia in the Alps, colonizing the central Alps from a single south-eastern Alpine refugium after the last glacial maximum (Gugerli, Rüegg, and Vendramin, 2009; Tóth et al., 2019). Reduced genetic diversity in Swiss stone pine from east to west and low genetic diversity and high differentiation in marginal populations are likely the result of postglacial colonization creating genetic drift and reducing gene flow from founder events at the leading range edge (Gugerli, Rüegg, and Vendramin, 2009; Tóth et al., 2019). Swiss stone pine can live for 500 to 1 000 years, with long generation intervals, which limits its opportunity to genetically differentiate over short periods; this may explain low differentiation between populations in the Alps and Carpathians (Tóth et al., 2019).
The bibliographic review was conducted by James Chaplin of the EUFORGEN Secretariat in August 2024.
Swiss pine has weak competitiveness and human activity has fragmented and reduced the size of populations, reducing gene flow. This increases inbreeding and the risk of losing genetic diversity in the species, which already inhabits unstable areas where avalanches, forest fires, landslides, or unusual climatic events can lead to further dramatic losses of genetic diversity (Ulber, Gugerli, and Bozic, 2004). Many Swiss stone pine stands have a highly skewed age distribution because of a lack of recruitment, often due to grazing from domestic and wild animals (Ulber, Gugerli, and Bozic, 2004). Weak competitiveness also reduces the capability of the species to spread (Ulber, Gugerli, and Bozic, 2004).
Small and isolated populations of Swiss stone pine with unique adaptations are of special interest for conservation (Ulber, Gugerli, and Bozic, 2004). Swiss stone pine is used for protection and aesthetic functions, so stands can be managed for multiple goals including genetic conservation. Dense, multilayered, uneven-aged forests with a clustered structure meet these demands best (Ulber, Gugerli, and Bozic, 2004).
Natural regeneration should be encouraged in stands for dynamic in situ gene conservation (Ulber, Gugerli, and Bozic, 2004). Removing patches of very dense vegetation cover and ensuring the presence of nutcrackers will help encourage natural regeneration (Ulber, Gugerli, and Bozic, 2004). When planting or direct seeding is used, reproductive material must only be well-adapted, acclimatized, and sufficiently variable material from local sites (Ulber, Gugerli, and Bozic, 2004).
The bibliographic review was conducted by James Chaplin of the EUFORGEN Secretariat in August 2024.
Genetic Characterisation of Pinus cembra and its GCUs
Availability of FRM
FOREMATIS
Pinus cembra - Technical guidelines for genetic conservation and use for Swiss stone pine
Publication Year: 2004The main strategy of genetic conservation of Swiss stone pine should be a dynamic in situ conservation. However, complementary dynamic or static ex situ conservation may be appropriate in some cases.
In situ conservation can be done within natural forest reserves, in gene conservation units specifically managed for the purpose, as well as in forests managed mainly for other purposes. Generally, abundant and large stands of interfertile autochthonous stone pine trees are best to assure genetic adaptability and adaptedness in the long term. However, small and isolated populations of P. cembra can be of special interest if particular adaptations can be assumed.
Normally, gene conservation can be matched with other goals of the stand management. As many of the P. cembra forests are expected to fulfil protection and/or aesthetic functions, a continuous tree cover is needed. A sufficiently dense, multilayered, uneven-aged forest with a clustered structure meets these demands best. Small-scale silvicultural practices such as the group selection method or the cluster selection method are recommended for both wood production and protection purposes. Where stone pine is naturally mixed with other tree species it is advisable to retain the mixture.
The crucial point in any case for a dynamic in situ gene conservation is the regeneration of the stand. Natural regeneration is considered to be the best way for gene conservation.
If there are suitable germination sites and seed trees within the nutcracker's dispersal range, regeneration of P. cembra occurs naturally. Sometimes a very dense cover of grasses or Alpine roses can be an obstacle for seedling growth, and the removal of patches of this vegetation can be useful. If necessary, animals affecting the young plants have to be excluded or individual protection measures must be taken. Forest gaps smaller in diameter than the height of the surrounding stand tend to accumulate snow during the winter and become snowfree only late in spring. Therefore, the dimensions of regeneration gaps should be 1–4 times the stand height depending on the site conditions.
For artificial regeneration in existing stands or for highelevation afforestations with P. cembra, reproductive material must be chosen carefully owing to extreme site conditions. Only well-adapted and sufficiently variable material originating from similar sites ensures a long-term success for both production and protection purposes. For plantations near the tree limit, the provenance of the material should lie within a 100-metre altitude range relative to the planting site. It is suggested to breed the plants in montane altitudes and to transplant them into a nursery located above 1500 m asl to ensure hardy acclimatization. Direct seeding has proven to be a satisfactory alternative to planting. As a consequence of the erratic seed production and the slow growth process, the planning of a plantation should start well in advance. Seed orchards at lower altitudes containing high-elevation provenances can facilitate a more regular supply of seeds.
Seed orchards can serve as a means of dynamic ex situ conservation of P. cembra if the number of progenies is sufficiently high (at least 50 trees proposed per origin population), especially in the case of endangered small or relic populations. However, it is preferable to conduct dynamic ex situ conservation in the vicinity of the original site and to use the local material. This is what the nutcracker tends to do when it "salvages" the species from accessible spots to rocky outcrops where the seedlings may survive.
Static ex situ conservation of reproductive material of P. cembra for gene conservation purposes is advisable only in case of emergency and the material should be recultivated as soon as possible.
Contacts of experts
NA
Further reading
Dzialuk, A., Chybicki, I., Gout, R., Mączka, T., Fleischer, P., Konrad, H., Curtu, A.L., Sofletea, N., and Valadon, A. 2014. No reduction in genetic diversity of Swiss stone pine (Pinus cembra L.) in Tatra Mountains despite high fragmentation and small population size. Conservation Genetics, 15: 1433–1445.
Wojnicka-Półtorak, A., Celiński, K., Chudzińska, E., Prus-Głowacki, W., and Niemtur, S. 2015. Genetic resources of Pinus cembra L. marginal populations from the Tatra Mountains: Implications for Conservation. Biochemical Genetics, 53: 49–61.
References
Gugerli, F., Rüegg, M., and Vendramin, G.G. 2009. Gradual decline in genetic diversity in Swiss stone pine populations (Pinus cembra) across Switzerland suggests postglacial re-colonization into the Alps from a common eastern glacial refugium. Botanica Helvetica, 119: 13–22.
Teodosiu, M. 2007. Genetic diversity and differentiation in Swiss stone pine (Pinus cembra L.). Annals of Forest Research, 50: 7–16.
Tóth, E.G., Tremblay, F., Housset, J.M., Bergeron, Y., and Carcaillet, C. 2019. Geographic isolation and climatic variability contribute to genetic differentiation in fragmented populations of the long-lived subalpine conifer Pinus cembra L. in the western Alps. BMC Evolutionary Biology, 19: 190. https://doi.org/10.1186/s12862-019-1510-4.
Ulber, M., Gugerli, F., and Bozic, G. 2004. EUFORGEN Technical Guidelines for genetic conservation and use for Swiss stone pine (Pinus cembra). Rome, International Plant Genetic Resources Institute. 6 pages.
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