Illustration of tree, foliage, cone, and seed [Matt Strieby, 2023].
Pollen cones on a tree in Serranía de Ronda, Spain [Jose Angel Campos Sandoval, 2007].
Seed cones on a tree in Serranía de Ronda, Spain [Jose Angel Campos Sandoval, 2007].
Fertile seed cones on a naturalized specimen growing on coastal Oregon sand dunes [C. J. Earle, 2021.05.25].
Seeds with attached wings (Steve Hurst @ USDA-NRCS PLANTS Database, accessed 2007.07.20).
Conservation Status
(subsp. escarena & pinaster)
(subsp. renoui)
Pinus pinaster
Maritime pine, cluster pine [English]; pinheiro bravo [Portuguese]; pi marítim, pi pinastre [Catalan]; pino marítimo, pino resinero, pino rodeno [Spanish].
Three subspecies:
See POWO for the extensive synonymy, which includes over 40 names; of these only Pinus maritima, an invalid name, is reasonably common.
This is the type species of Pinus subsect. Pinaster Loudon, a mostly Mediterranean group of 7 species. It is nearly the basal name in the subsection, sister to a clade that includes all the other species except P. heldreichii, with a most recent common ancestor dating to about the Eocene/Oligocene boundary (Jin et al. 2021). The subspecies are separated by minor morphological and ecological differences; the most disparate is subsp. renoui in North Africa, while subsp. escarena on the Mediterranean coast and subsp. pinaster on the Atlantic coast are very similar (Farjon 2010). However, analysis of mitochondrial haplotypes has revealed considerable genetic structure, indicating greatest genetic diversity among populations of central and southeast Spain, with very low diversity in the western Iberian Peninsula and in Morocco. These results also support the existence of glacial refugia for the species in southeastern Spain, coastal Portugal, and coastal North Africa, with haplotypes from Africa occurring in Italy but not in more westerly areas (Bucci et al. 2007). A complementary analysis of nuclear DNA markers groups the populations into a Moroccan, an Iberian, and a French-Italian group, emphasizing the dispersal barriers created by the Pyrenees and the Strait of Gibraltar (Naydenov et al. 2014). The nuclear analysis also estimates the time to the most recent common ancestor at approximately 700,000 years ago, suggesting that the species has maintained these disparate genetic groups across multiple glacial-interglacial cycles.
Monoecious evergreen trees to 40 m tall, with a commonly somewhat sinuous stem (straight in some areas, notably Corsica and NW Africa) and a usually open crown of regular, candelabra-like upswept branches, broad ovoid-conic when young, becoming irregular and denser with age. The bark is very thick, scaly or plated, fissured, patterned red-brown, blackish and buff. Branching is mostly uninodal, but occasionally multinodal on vigorous young trees. Shoots are stout, 7-15 mm thick, buff to yellow-brown, rough. Foliage buds are large, cylindric to ovoid-acute, with red-brown scales having long free tips, revolute, fringed with white hairs. Adult leaves, retained 1.5-3 years, are 12-25 cm long in fascicles of two (with some fascicles of 3), with a persistent 2 cm sheath. They are green to yellow-green, sometimes tinged glaucous, spreading, very stout, about 2-2.2 mm thick, with serrulate margins, and fine lines of stomata on all faces. Juvenile leaves are strongly glaucous, 3-6 cm long, on slender (3 mm) shoots, and are grown for the first 2-5 years of life. Cones are slightly deflexed on short stout stalks, symmetrical, hard, heavy, ovoid-conic, (7-) 9-18 (-20) cm long, 4-6 cm broad when closed, ripening rich glossy chestnut-brown in April two years after pollination, and opening the same summer or up to 10 years later, to 7-11 cm broad. The scales are stout, thick, woody, and stiff; the apophysis is 10-16 × 18-26 mm wide, rhomboid, with a strong raised transverse ridge; the umbo is dorsal, blackish grey, stout, 4-7 mm wide and 3 mm high. The seeds are shiny blackish brown above, matt grey below, 7-11 × 5 mm with an 18-25 × 10 mm wing, buff with numerous straight dark brown streaks, the wing easily removed from the seed (Frankis, M.P., pers. comm. 1999.02.10). See García Esteban et al. (2004) for a detailed characterization of the wood anatomy.
Subsp. escarena are trees to 40 m tall with leaves to 25 cm long and 2 mm thick, with seed cones to 22 cm long (Farjon 2010).
Subsp. pinaster are trees to 30 m tall with leaves 10-20 cm long and seed cones 10-18 cm long (Farjon 2010).
Subsp. renoui are trees to 25 m tall with a dense, domed crown. Leaves are 12-20 cm long, slender, 1.5-2 mm thick, with 10-15 cm long seed cones (Farjon 2010).
Algeria, France, Italy, Malta, Monaco, Morocco, Portugal, Spain, Tunisia, and United Kingdom: Gibraltar. Occurs at elevations as low as sea level in France, rising to 900 m in Corsica, 1500-1700 m in southeast Spain, and 2000–2200 m in the Atlas mountains of Morocco. It also occupies very diverse soils, derived from silicates, carbonates, or ultramafics, and on dunes. It mainly grows in relatively xeric woodlands that, if more mesic, would become broad-leaved woodlands; and it forms mixed woodlands dominated typically by Pinus spp. or Quercus spp. (Olmedo‑Cobo et al. 2019 and sources therein). It is the most abundant conifer in Portugal, with a distribution estimated at 7,133 km2 in 2015 (ICNF 2019). Since the 1950's, P. pinaster has also been so widely planted that plantations are now found throughout most of its native range, in France as far north as Brittany, and on all continents but Antarctica. It has been designated an exotic invasive species in parts of South America and Africa (Charco et al. 2014). Within Europe, its original native distribution is not precisely known, since humans have been growing it for timber and resin for thousands of years; records from earlier times are known from only a few locales.
Distribution of the Mediterranean species of subsect. Pinaster. Polygons provided by Caudullo et al. (2024). P. pinaster shown in gray. Click on any polygon to see the species represented.
Subsp. escarena is found around the W Mediterranean in Spain (including the Balearic Islands), France (including Corsica), Italy (including Sardinia and Sicily), and Malta (Farjon 2010). Widely distributed with many populations, it has been assessed as of "Least Concern" for conservation.
Subsp. pinaster has an Atlantic distribution, including coastal France, Portugal, and parts of Spain; it has also been widely planted in interior locations and such trees may not be confidently assignable to subsp. escarena or pinaster (Farjon 2010). Widely distributed with many populations, it has been assessed as of "Least Concern" for conservation.
Subsp. renoui is in Morocco and near the coast in NE Algeria and Tunisia (Farjon 2010). It has always been rare and in 2011 was assessed as "Endangered" due to the presence of only two small, relict, widely-separated populations; ongoing logging and habitat degradation to an unknown extent; and the risk of hybridization with plantations of the European subspecies (Farjon 2013).
Bannister and Neuner (2001) describe P. pinaster as hardy to Zone 8 (cold hardiness limit between -12.1°C and -6.7°C). However, a common-garden experiment by Prada et al. (2017) found high intraspecific variation in cold-hardiness and other intraspecific traits: "Two continental Iberian populations showed high cold tolerance and slower growth... The coastal populations displayed the opposite behavior, while the continental Moroccan population presented a unique combination of traits. We confirmed trade-offs between cold-hardiness and growth at the population level, but not within populations."
Some populations of this species are well-adapted to tolerance of low-intensity fire. Adaptations to fire include thick bark, serotinous cones, and tolerance of high levels of crown scorch, but these characteristics are highly variable between populations. In the mountains of Spain, where P. pinaster is a forest dominant species, natural ignition density from lightning exceeds the country’s average by a factor of 27. However, fire risk has been exacerbated by the widespread use of P. pinaster to reforest abandoned agricultural lands and establish timber plantations, which due to their extent and uniform structure represent a high risk of catastrophic wildfire; thus there is an ongoing management focus on techniques to manage and limit the impacts of wildfire on the P. pinaster landscape (Fernandes and Rigolot 2007, Niccoli et al. 2019).
Abies pinsapo and Pinus pinaster form a natural vegetation type on the ultramafic-derived soils of the Sierra Bermeja of southern Spain, which is the largest ultramafic body in western Europe. Evidence from soil charcoal indicates this association has persisted for millennia (Olmedo‑Cobo et al. 2019).
The largest recorded specimen appears to have been 4.78 m in girth (152 cm dbh) when last measured in 2013; it grows near Talayuela, Spain. The tallest is stated to be 38.1 m tall (Landes, France), measured 2021 (Dominique Beziat email 2021.02.09). There is a 2017 record of a 39.2 m tall tree near Braga, Portugal (Dominique Beziat email 2020.11.29). The tallest ever recorded, according to a chief technician of the Forests National Office (France), was a 42 m tall tree in the forest of the Landes (S.W. of France) that fell during a storm in December, 1999 (Dominique Beziat email, 2020.11.28).
The oldest known living specimen, 165 years, was documented in a tree-ring chronology covering the period 1821-1985 (crossdated after 1836), collected in east-central Spain by Klaus Richter (doi.org/10.25921/4y97-4x15). The next-oldest tree, 162 years, was documented in a tree-ring chronology covering the period 1843-2004 (crossdated after 1866), collected in northern Morocco by Ramzi Touchan (doi.org/10.25921/j049-eh91). This site was used in a dendroclimatic drought reconstruction (Touchan et al. 2011). Only three chronologies have been developed for this species; considerably older trees may exist.
P. pinaster is one of the most important forest trees in France, Portugal and Spain. It is used for wood and resin production, recreation, and soil protection. In the Atlantic region rotation ages of 40-50 years are common and the species is heavily exploited for pulp and paper production, construction, chipboard, flooring and pallets. In the Mediterranean region rotation ages vary from 80 to 120 years; trees in Corsica and some mountainous areas of central Spain produce a high quality timber, while usage in some areas (plains of Castille and several southern populations in southern Spain) is limited because the trees are very crooked (Alía and Martín 2003). The species has also been widely planted in southwestern Australia (Ritson and Sochacki 2003).
Pinus pinaster was introduced to the Landes de Gascogne under Napoleon III, where it now forms the largest artificial forest in Europe, covering 8200 km2 on the Atlantic coast of SW France. The area was originally planted (from 1789 onward) for land reclamation, with a huge area of shifting sand dunes threatening fertile farmland futher inland. The area is now, however, of major economic importance to the timber industry (Frankis pers. comm. 1999.02.10, Theraroz 2024).
Historically, the species was heavily exploited in Spain for resin production from the 1840s until the 1970s, when low prices ended the industry. During this period, P. pinaster was the pine most often tapped for resin, which became one of the most important non-timber products. Trees historically scarred for resin production are still common in Spain (Génova et al. 2014).
The species has been widely planted in other parts of the world with a Mediterranean climate, and is now naturalised in South Africa and elsewhere (Price et al. 1998); in recent years planting outside of Europe has declined considerably as P. radiata provides larger crops of better quality timber in the same conditions (Frankis, M.P., pers. comm. 1999.02.10).
P. pinaster has been used in a variety of dendrochronology studies, starting in 1982. Some of these studies have looked at the species' utility in reconstructing past climate variation, and most of the others have looked at ecological problems such as stand development, productivity, fire ecology, and the impacts of insect defoliators. Génova et al. (2014) present a brief review of these studies.
The oldest recorded specimen is 210 years, crossdated, for a tree sampled by Didier Bert and Frédéric Lagane at Pin Cazau, Landes de Gascogne Forest, France (Brown 2025).
The epithet is from the Italian "pinastro", a common name for the tree.
Aiton, W. 1789. Hortus Kewensis vol. 3, p. 367. Available: www.biodiversitylibrary.org/item/23434, accessed 2011.05.21.
Alía, Ricardo, and Sonia Martín. 2003. Technical Guidelines for Genetic Conservation and Use, Pinus Pinaster. Euforgen.
Brown, Peter. 2025. OLDLIST, a database of old trees. https://www.rmtrr.org/oldlist.htm, accessed 2025.03.01.
Bucci, Gabriele, Santiago C. González‐Martínez, Grégoire Le Provost, Christophe Plomion, Maria Margarida Ribeiro, Federico Sebastiani, Ricardo Alía, and Giovanni Giuseppe Vendramin. 2007. Range‐wide phylogeography and gene zones in Pinus pinaster Ait. revealed by chloroplast microsatellite markers. Molecular Ecology 16(10):2137–53. https://doi.org/10.1111/j.1365-294X.2007.03275.x.
Charco, J., M. Becerra, C. Santabárbara, et al. 2014. Árboles y arbustos autóctonos de Andalucía. Centro de Investigaciones Ambientales del Mediterráneo-CIAMED, Madrid.
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Farjon, A. 2013. Pinus pinaster subsp. renoui. The IUCN Red List of Threatened Species 2013: e.T63554A3127306. https://dx.doi.org/10.2305/IUCN.UK.2013-1.RLTS.T63554A3127306.en, accessed 2025.02.12.
Fernandes, Paulo M., and Eric Rigolot. 2007. The fire ecology and management of maritime pine (Pinus pinaster Ait.). Forest Ecology and Management 241(1–3):1–13. https://doi.org/10.1016/j.foreco.2007.01.010.
Ferrer-Gallego, P. Pablo. 2023. Type designation of the Maritime pine Pinus pinaster (Pinaceae). Phytotaxa 587(3):294-300. https://doi.org/10.11646/phytotaxa.587.3.7.
Gaussen, H., V.H. Heywood, and A.O. Chater. Pinus. In: T.G. Tutin et al. 1993 Flora Europaea 2nd edition, Vol. 1. Cambridge.
Génova, Mar, Leocadia Caminero, and Javier Dochao. 2014. Resin tapping in Pinus pinaster: effects on growth and response function to climate. European Journal of Forest Research 133:323–33.
Greuter, W., H.M. Burdet, and G. Long. 1984. Med.Checklist - A critical inventory of vascular plants of the circum-mediterranean countries, Vol. 1. Geneva.
ICNF. 2019. Inventário Florestal Nacional: 2015 Relatório Final. Lisboa: ICNF. 284 p. Available: https://www.icnf.pt/api/file/doc/c8cc40b3b7ec8541, accessed 2025.01.23.
Jin, Wei-Tao, David S. Gernandt, Christian Wehenkel, Xiao-Mei Xia, Xiao-Xin Wei, and Xiao-Quan Wang. 2021. Phylogenomic and ecological analyses reveal the spatiotemporal evolution of global pines. Proceedings of the National Academy of Sciences 118(20): e2022302118. doi.org/10.1073/pnas.2022302118.
Maire, R. 1952. Flore de l'Afrique Du Nord. Paris: Lechevalier. 1:145. Available: Scribd, accessed 2025.02.12.
Naydenov, Krassimir D., Alexander Alexandrov, Vlado Matevski, Kole Vasilevski, Michel K. Naydenov, Veselka Gyuleva, Christopher Carcaillet, Nadya Wahid, and Salim Kamary. 2014. Range-wide genetic structure of maritime pine predates the last glacial maximum: evidence from nuclear DNA. Hereditas 151(1):1–13. https://doi.org/10.1111/j.1601-5223.2013.00027.x.
Niccoli, Francesco, Assunta Esposito, Simona Altieri, and Giovanna Battipaglia. 2019. Fire severity influences ecophysiological responses of Pinus pinaster Ait. Frontiers in Plant Science 10539. https://doi.org/10.3389/fpls.2019.00539.
Olmedo-Cobo, José Antonio, Raquel Cunill-Artigas, and José Gómez-Zotano. 2019. The native status of Pinus pinaster on Serpentine Soils: Charcoal Analysis and Palaeoenvironmental History in Sierra Bermeja (Southern Iberian Peninsula, Spain).” Vegetation History and Archaeobotany 28, no. 4 (July 2019): 417–32. https://doi.org/10.1007/s00334-018-0701-z.
Prada, Eva, José Climent, Ricardo Alía, and Raquel Díaz. 2016. Life-history correlations with seasonal cold hardiness in maritime pine. American Journal of Botany 103(12):2126-2135.
Richter, K. 1890. Plantae Europeae 1:1. Available: Biodiversity Heritage Library, accessed 2025.02.12.
Ritson, P., and S. Sochacki. 2003. Measurement and prediction of biomass and carbon content of Pinus pinaster trees in farm forestry plantations, southwestern Australia. Forest Ecology and Management 175:103–117.
Theraroz, Adélaïde. 2024. Maritime pine genetic resources: geographical variability, selection pressures and future adaptation. Ph.D. dissertation, Université de Bordeaux.
Touchan, Ramzi, Kevin J. Anchukaitis, David M. Meko, Said Attalah, Christopher Baisan, Ali Aloui. 2011. Spatiotemporal drought variability in northwestern Africa over the last nine centuries. Climate Dynamics 37(1):237-252. doi: 10.1007/s00382-010-0804-4
Elwes and Henry 1906-1913 at the Biodiversity Heritage Library. This series of volumes, privately printed, provides some of the most engaging descriptions of conifers ever published. Although they only treat species cultivated in the U.K. and Ireland, and the taxonomy is a bit dated, still these accounts are thorough, treating such topics as species description, range, varieties, exceptionally old or tall specimens, remarkable trees, and cultivation. Despite being over a century old, they are generally accurate, and are illustrated with some remarkable photographs and lithographs.
The species account at Threatened Conifers of the World.
Last Modified 2025-03-01
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