Pinus pinea
Stone pine

Stone pine (Pinus pinea) is a coniferous tree species native to the Mediterranean region. However, due to widespread cultivation its native distribution is not apparent (Carrasquinho and Gonçalves, 2013). Thriving in dry, sunny areas with high temperatures, it is found across southern Europe, growing in various climatic conditions in small and scattered populations from sea level up to 1 000 m above sea level (Jaramillo‐Correa et al., 2020). Stone pine prefers well-drained, sandy, acidic soils but is tolerant of coastal conditions, frost, and calcareous soils. Despite its wide distribution, old-growth forests are scarce and typically associated with coastal dunes or former estuarine marshes (Mutke et al., 2019).

Humans have utilized stone pine for centuries, primarily for its edible seeds. The tree is a commercially important species even today. The wood is of low quality but used locally for furniture, while the resin is tapped and used for rosin. Other uses include stabilization of sand dunes and as an ornamental in warmer regions in Europe.

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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 pinea conservation in Europe

Genetic diversity and variation

Stone pine is genetically depauperate, with less genetic diversity than other pine species because of demographic collapse and a severe genetic bottleneck (Fady, Fineschi, and Vendramin, 2004; Jaramillo‐Correa et al., 2020). The genetic bottleneck in stone pine is likely the result of natural climatic fluctuations during the Quaternary (Vendramin et al., 2008). Severe population bottlenecking was likely not a single event but a trend of population collapse that continued through the Holocene and reduced ancestral stone pine populations to 1%–10% of their original size (Vendramin et al., 2008; Jaramillo‐Correa et al., 2020). The presence of one haplotype right across the range suggests that it may have survived in only a single population at some point in its history (Vendramin et al., 2008).

Gene flow may be restricted by stone pine’s reliance on seed dispersal by birds and the fact that cones take three years to mature (Jaramillo‐Correa et al., 2020). Restricted gene flow means mutations appearing after the species’ genetic decline are only locally dispersed and are more likely to be lost by genetic drift (Jaramillo‐Correa et al., 2020). Stone pine has maintained a low level of diversity while spreading across diverse and fragmented regions, which implies either mutations in the species are rare or that expansion was recent (Vendramin et al., 2008). Despite bottlenecking and low diversity, stone pine has managed to survive; this could be because the species shows a significant amount of genetic variation in adaptive traits considering its low genetic diversity, allowing it to expand into diverse habitats (Vendramin et al., 2008). For example, research has shown significant variation among provenances of stone pine for growth traits such as total height and growth rate (Carrasquinho and Gonçalves, 2013).

Genetic distribution and clustering 

Despite its wide distribution, stone pine is known since the late 1970s to be genetically very uniform, when studies revealed that certain proteins were unexpectedly identical across many individuals and populations. There is no description in the literature of geographical races, ecotypes, or cultivars; however, one variety, P. pinea var. fragilis, which produces soft-shelled seeds, may be the result of a single mutation (Fady, Fineschi, and Vendramin, 2004). There is also no evidence of any strong geographic structure in adaptive traits such as vigour (Fady, Fineschi, and Vendramin, 2004). Studies have identified only four haplotypes around the Mediterranean; most populations contain a single haplotype, while Spanish populations contain another, and Lebanese populations contain two more (Fady, Fineschi, and Vendramin, 2004; Vendramin et al., 2008). The haplotypes observed in Spain and Lebanon are likely later mutations that occurred after the species’ genetic bottleneck.

 

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

Cultivation and human intervention

There is evidence that people were eating stone pine nuts 300 000 years ago, and that it has been cultivated for more than 6 000 years (Jaramillo‐Correa et al., 2020). The split between the eastern Mediterranean and Iberian populations probably took place about 9 500 years ago, around the same time as the earliest archaeological evidence for stone pine cultivation. Rapid and widespread range expansion and dispersion in the Holocene was probably human mediated as natural dispersion is low and colonization occurred very quickly and coincides with the beginning of widespread cultivation around 3 000 years ago (Mutke et al., 2019; Jaramillo‐Correa et al., 2020).

 

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

Threats

Despite low genetic diversity stone pine is not considered a threatened species. It is rarely attacked by pests and diseases, although some insects are known to cause damage (Fady, Fineschi, and Vendramin, 2004). Forest fires are the main threat to stone pine genetic diversity, but stone pine is considerably less fire-sensitive than other species, given its thick bark and high crown devoid of low branches (Fady, Fineschi, and Vendramin, 2004). Fire is also necessary for natural stone pine regeneration.

Stone pine is highly sensitive to air pollution, especially when combined with other environmental stresses such as drought (Fady, Fineschi, and Vendramin, 2004). Widespread cultivation using the same genetic material has also likely contributed to the species’ reduced genetic diversity (Fady, Fineschi, and Vendramin, 2004).

Management

Analysis of stone pine genetic variability is inconsistent, hence research to help inform management would be beneficial (Carrasquinho and Gonçalves, 2013). Because of its uniform and low genetic diversity, much of the genetic variability within stone pine can be captured within a small number of individuals (Jaramillo‐Correa et al., 2020). However, research is still needed to determine whether current Mediterranean forests are really well adapted and truly natural, and to understand the history and ecology of this species (Fady, Fineschi, and Vendramin, 2004).

Establishment of in situ conservation networks, where selected populations are allowed to naturally regenerate without introduction of exotic material, is recommended (Fady, Fineschi, and Vendramin, 2004). Prioritization should be given to areas where extensive populations currently exist or with ecological extremes (Fady, Fineschi, and Vendramin, 2004). As many seed trees as possible should be left to maximize outcrossing and pollen flow in natural regeneration, including severely damaged/burned trees, to overcome the species low genetic diversity (Fady, Fineschi, and Vendramin, 2004). Stands should also implement fire- and overgrazing-protection measures.

 

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

Genetic Characterisation of Pinus pinea and its GCUs

Availability of FRM

FOREMATIS

EUFORGEN’s publications

Technical guidelines for genetic conservation and use

Pinus pinea - Technical guidelines for genetic conservation and use for Italian stone pine

Publication Year: 2004
Author: Fady, B.; Fineschi, S.; Vendramin, G.G.

The conservation of forest genetic resources in the Mediterranean basin is a very complex task, as ecological and socioeconomic conditions are highly variable among countries. Because of their history of overexploitation since agriculture emerged some 10 000 years ago, assessing whether current Mediterranean forests are really well adapted and truly natural is challenging, although necessary for any careful conservation strategy.
This is the case for P. pinea, in particular. A number of scientific gaps should be filled. The past history and ecology of this species need to be understood to outline the areas of autochthony. Knowing its current adaptive diversity is also a prerequisite to outlining its potential distribution area and the consequences it may suffer from environmental changes. As in other forest tree species, implementing an in situ conservation network where selected populations are allowed to naturally regenerate without introduction of exotic material is recommended. Regions of autochthony such as Spain and the eastern Mediterranean, areas where ecological conditions are extreme (high altitude, low rainfall, high salinity etc.), and areas where extensive populations currently exist, should be the primary targets for such a network.

Appropriate silvicultural and management strategies should include the leaving of the highest possible number of seed trees before regeneration to promote maximum outcrossing and pollen flow. This might mean not cutting severely burned trees after wild fires. It should also include, in areas that are not designated for seed production, letting natural selection (rather than managed thinning) sort out young trees after regeneration. Wild fires and overgrazing being the most important risks for P. pinea forests, fire protection and social measures that might reduce these hazards should also be addressed for the effective conservation of this typically Mediterranean pine.

The conservation of forest genetic resources in the Mediterranean basin is a very complex task, as ecological and socioeconomic conditions are highly variable among countries. Because of their history of overexploitation since agriculture emerged some 10 000 years ago, assessing whether current Mediterranean forests are really well adapted and truly natural is challenging, although necessary for...
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References

Carrasquinho, I. and Gonçalves, E. 2013. Genetic variability among Pinus pinea L. provenances for survival and growth traits in Portugal. Tree Genetics & Genomes, 9: 855–866.

Fady, B., Fineschi, S., and Vendramin, G.G. 2004. EUFORGEN Technical Guidelines for genetic conservation and use for Italian stone pine (Pinus pinea). Rome, International Plant Genetic Resources Institute. 6 pages.

Jaramillo‐Correa, J.P., Bagnoli, F., Grivet, D., Fady, B., Aravanopoulos, F.A., Vendramin, G.G., and González‐Martínez, S.C. 2020. Evolutionary rate and genetic load in an emblematic Mediterranean tree following an ancient and prolonged population collapse. Molecular Ecology, 29(24): 4797–4811.

Mutke, S., Vendramin, G.G., Fady, B., Bagnoli, F., and González-Martínez, S.C. 2019. Molecular and quantitative genetics of stone pine (Pinus pinea). In: D. Nandwani, ed. Genetic diversity in horticultural plants, pp. 61–84. Sustainable Development and Biodiversity, vol 22. Cham, Switzerland, Springer.

Vendramin, G.G., Fady, B., González-Martínez, S.C., Hu, F.S., Scotti, I., Sebastiani, F., Soto, A., and Petit, R.J. 2008. Genetically depauperate but widespread: the case of an emblematic Mediterranean pine. Evolution, 62(3): 680–688.