Pinus nigra
European black pine

Black Pine or Austrian Pine (Pinus nigra) is a fast-growing, robust, evergreen conifer (Scotti-Saintagne et al., 2023). It is native to the Mediterranean region and Asia Minor and has been introduced to the United States. The species is widespread but has a scattered range with often patchily distributed populations across different ecological environments and climatic conditions (Giovannelli et al., 2017; Scotti-Saintagne et al., 2023). It is mainly found in mountainous areas and is light-demanding but shade-tolerant, resistant to wind and drought, and adapted to dry and cold conditions.

Black pine is one of the most economically important native conifers in southern Europe. Its timber is durable, rich in resin, and easy to process, making it suitable for indoor flooring, general construction, and furniture-making. The species has a high adaptive plasticity, making it suitable for planting in a range of environments (Isajev et al., 2004). Additionally, due to its resistance to pollution and soil erosion, it is frequently utilized in reforestation projects as a windbreak or to control soil erosion and landslides, and as an ornamental tree in parks, urban and industrial contexts, and large gardens.

in situ genetic conservation unit
ex situ genetic conservation unit
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Acknowledgements

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 nigra conservation in Europe

Genetic diversity and variation

Genetic diversity in black pine is high but has been shown to be lower than other pine (Pinus) species such as Chinese red pine (Pinus tabulaeformis), Korean pine (Pinus koraiensis), Siberian pine (Pinus sibirica), and Scots pine (Pinus sylvestris), but higher than Qiaojia pine (Pinus squamata) (Rubio-Moraga et al., 2012). Genetic diversity in black pine is higher within populations than between populations, with research in Spain and Morocco showing only 9% of the total variation between black pine populations (Isajev et al., 2004; Rubio-Moraga et al., 2012).

Genetic distribution and clustering

Black pine shows some geographically related structuring and clustering of genetic diversity (Giovannelli et al., 2017). This could be the result of natural barriers, agricultural and forest harvesting, local adaptation, and management methods (Rubio-Moraga et al., 2012). However, in some populations geographic separation does not result in mating barriers (Isajev et al., 2004). Geographic distance was not correlated with genetic distance, and clustering of populations was not related to their geographical locations in Spanish and Moroccan populations; populations in Spain and Morocco did, however, show high differentiation, indicating some restriction in gene flow from separation, and such a pattern has been found in other parts of black pines range (Rubio-Moraga et al., 2012). High differentiation across black pine’s range occurred due to genetic drift, low levels of gene flow for a conifer, and the high number of Mediterranean glacial refugia in the Iberian, Italian, and Balkan peninsulas (Rubio-Moraga et al., 2012).

 

The bibliographic review was conducted by James Chaplin of the EUFORGEN Secretariat in August 2024.

Interspecific taxa dynamics

A fragmented distribution has led to a wide range of subspecies of the black pine, each with specific preferred habitat conditions in terms of soil and climate, and morphological variations. These are difficult to interpret and have resulted in several diverse classifications (Rubio-Moraga et al., 2012). Five subspecies of black pine are recognized (although there is no consensus, and some are sometimes regarded as distinct species): Pinus nigra ssp. nigra from the Alps; Pinus nigra. laricio from Corsica and Sicily; Pinus nigra ssp. pallasiana from Türkiye and Crimea; and Pinus nigra ssp. salzmannii (Salzmann’s pine) and Pinus nigra ssp. mauretanica from North Africa (Rubio-Moraga et al., 2012; Giovannelli et al., 2017). Corsican populations are the most differentiated and have a lack of genetic variation, likely because of their geographic isolation (Giovannelli et al., 2017). This makes identification of a single taxonomy for black pine difficult, especially as hybridization occurs easily (Isajev et al., 2004). However, morphological, and genetic markers have confirmed the common phylogenetic origin of all black pines (Isajev et al., 2004).

 

The bibliographic review was conducted by James Chaplin of the EUFORGEN Secretariat in August 2024.

Threats

Black pine is not recognized as a threatened species, although some endemic populations are a priority for conservation (Isajev et al., 2004). Widespread cultivation across Europe in the past two centuries, and the use of material from unknown sources means there has been a lot of mixing between local and imported gene pools across the tree’s distribution (Isajev et al., 2004). The same authors note that insects, funguses, and diseases are a threat, including European black pine shoot moth (Rhyacionia buoliana) and ‘red band disease’ (Dothistroma septospora). Forest fires, illegal cutting, and hybridization with planted black pine subspecies also threaten black pine’s genetic diversity, especially in fragmented populations (Isajev et al., 2004; Scotti-Saintagne et al., 2023).

Management

Inventories and testing of black pine to define its geographical distribution, conservation status, threats, and potential uses, and to identify genetic variability of populations are necessary across its range and at a regional level (Isajev et al., 2004; Rubio-Moraga et al., 2012). In situ conservation is advised and should be encouraged as both seed stands and gene conservation forests (Isajev et al., 2004). A network of experimental sites could be established with the sampling of marginal areas and different groups, such as the five identified genetic groups of Salzmann’s pine in France, serving as the basis for in situ gene conservation (Isajev et al., 2004; Scotti-Saintagne et al., 2023). However, cultivated, or exotic black pine varieties should not be planted close to autochthonous and naturalized stands, especially not next to localized or fragmented populations (Isajev et al., 2004).

 

The bibliographic review was conducted by James Chaplin of the EUFORGEN Secretariat in August 2024.

Genetic Characterisation of Pinus nigra and its GCUs

Availability of FRM

FOREMATIS

EUFORGEN’s publications

Technical guidelines for genetic conservation and use

Pinus nigra - Technical guidelines for genetic conservation and use for European black pine

Publication Year: 2003
Author: Isajev, V.; Fady, B.; Semerci, H.; Andonovski, V.

Because black pine of different origins has been extensively planted, it is now important to identify authochthonous populations. This undertaking should be carried out at the international level. In each country, an inventory should be made to define the geographical distribution of the species, its conservation status, threats and potential uses. Breeding activities provide valuable information by defining potential plantation, seed collection and transfer zones. In situ conservation activities should be encouraged separately as seed stands and gene conservation forests. Those do not serve the same goal and should not always be identical, especially to make the conservation of marginal populations possible. An international in situ network of 100–120 stands would seem appropriate to represent the natural ecological and genetic variability of black pines.

As intraspecific hybridization is easy among black pines, exotic or improved black pines should not be planted in the vicinity of autochthonous and naturalized stands. This is particularly true for localized and fragmented sub-species such as P.n. laricio, and is of extreme importance for subspecies that are threatened, such as P.n. salzmanii in France and P.n. mauretanica in North Africa. For these subspecies and other varieties of rare occurrence, ex situ conservation is appropriate and urgent. As a step in that direction, in 1999 a gene conservation forest was selected in Turkey for the rare P. nigra var. pyramidalis.

Information on the provenance and progeny trials established throughout Europe should be entered in a database. This network of experimental sites could be used for ex situ conservation of black pine. Marginal areas might need to be further sampled to strengthen this network and possibly planted as ex situ seed orchards to re-install depleted resources.

Because black pine of different origins has been extensively planted, it is now important to identify authochthonous populations. This undertaking should be carried out at the international level. In each country, an inventory should be made to define the geographical distribution of the species, its conservation status, threats and potential uses. Breeding activities provide valuable information...
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Further reading

Aguinagalde, I., Llorente, F., and Benito, C. 1997. Relationships among five populations of European black pine (Pinus nigra ARN.) using morphometric and isozyme markers. Silvae Genetica, 46(1): 1–5.

References

Giovannelli, G., Roig, A., Spanu, I., Vendramin, G.G., and Fady, B. 2017. A new set of nuclear microsatellites for an ecologically and economically important conifer: the European black pine (Pinus nigra Arn.). Plant Molecular Biology Reporter, 35: 379–388.

Isajev, V.,Fady, B., Semerci, H., and Andonovski, V. 2004. EUFORGEN Technical Guidelines for genetic conservation and use for European black pine (Pinus nigra). Rome, International Plant Genetic Resources Institute. 6 pages.

Rubio-Moraga, A., Candel-Perez, D., Lucas-Borja, M.E., Tiscar, P.A., Viñegla, B., Linares, J.C., Gómez-Gómez, L., and Ahrazem, O. 2012. Genetic diversity of Pinus nigra Arn. populations in southern Spain and northern Morocco revealed by inter-simple sequence repeat profiles. International Journal of Molecular Sciences, 13(5): 5645–5658.

Scotti-Saintagne, C., de Sousa Rodrigues, A., Roig, A., and Fady, B. 2023. A comprehensive strategy for the conservation of forest tree genetic diversity: an example with the protected Pinus nigra subsp. salzmannii (Dunal) Franco in France. Conservation Genetics, 25(2): 469–480.