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— CH. 1 · INTRODUCTION —

Pine

9 min listen · Ch. 1 of 7
7 sections
  • Pinus is the oldest surviving genus of its family, with fossil evidence stretching back to the Early Cretaceous period, roughly 125-130 million years ago. That first pine emerged in what is now England, preserved in a layer of rock called the Speeton Clay. Today the genus has spread across virtually every habitat in the Northern Hemisphere, from sea-level coasts to elevations of 5,200 metres, from the Arctic circle in Norway down past the equator into Sumatra. A single individual tree in the White Mountains of California, nicknamed "Methuselah", has stood for around 4,800 years, making it one of the oldest living organisms on Earth. So what exactly is a pine? How did one genus come to dominate so much of the planet's landscape? And why do we flavour wine with its resin, name entire peoples after eating its bark, and compose orchestral symphonies in its honour?

  • The tallest known pine, a sugar pine called Pinus lambertiana, reaches 83.45 metres and stands in Yosemite National Park. At the other end of the scale, the Siberian dwarf pine and the Potosi pinyon represent the genus's smallest members. Most species fall somewhere between 15 and 45 metres tall. The structure of these trees is precise in ways that have fascinated mathematicians: the spirals of branches, needles, and cone scales are arranged according to Fibonacci number ratios, the same sequence that appears in sunflower heads and nautilus shells.

    Branches emerge from the trunk in what botanists call pseudo-whorls, tight spirals that look to the eye like rings appearing from a single point. Many pines produce one such ring per year, allowing a rough estimate of a tree's age just by counting the branch tiers. Others are multinodal, pushing out two or more rings of branches each year.

    The bark tells its own story. Most species carry thick, scaly bark that protects the heartwood within. The inner bark, known as the cambium, is soft and white, and it is high enough in vitamins A and C to have served as a food source for Indigenous peoples. That fact gave the Adirondack Indians their name: the Mohawk word atirú:taks means "tree eaters", a reference to this cambium-eating practice.

  • A pine needle is not simply a leaf. It belongs to a cluster called a fascicle, with anywhere from one to seven needles per bundle, though two to five is the usual range. Each fascicle grows from a small bud anchored in the axil of a scale leaf, and the needles can persist for anywhere from 1.5 to 40 years depending on the species. If an animal eats the growing tip of a shoot, the fascicles just below the damage can generate a replacement bud and restore the lost growth.

    Reproduction in pines is a slow affair. Female cones take between 1.5 and 3 years to mature after pollination, with actual fertilisation delayed a full year. At maturity, these woody structures range from 3 to 60 centimetres in length. Most release their seeds when the cones open naturally, but some species have developed a more dramatic strategy called serotiny. In pyriscence, the most common form, resin glues the cone scales shut until a wildfire melts it. Species such as Pinus radiata and Pinus muricata rely on this mechanism. The fire clears the forest floor of competing vegetation and the seeds drop onto freshly burned, open ground, where they can establish with little competition.

    For the whitebark pine, the seeds are not released by heat at all. They are locked inside cones until a bird, specifically species like nutcrackers, physically breaks the cones open. The bird caches the seeds as a food store, and those it fails to retrieve may germinate. Red squirrels also feed on Pinus sylvestris seeds in Scotland, though unlike the birds, they do little to disperse the seeds widely.

  • Transcriptome analysis places Pinus most closely related to the genus Cathaya, which in turn sits closest to Picea, the spruces. Together with firs and larches, these genera form what researchers call the pinoid clade of the Pinaceae family.

    Within the genus itself, the evolutionary story is tangled. Wind pollination, long lifespans, overlapping generations, large population sizes, and weak reproductive barriers between species have all encouraged hybridisation across millions of years. Gene transfer between different species has created a history of genetic relatedness that is genuinely difficult to unravel. Two separate phylogenies published in 2021, one by Jin and colleagues and one by Stull and colleagues, arrive at different conclusions, illustrating how contested the internal relationships still are.

    Taxonomists divide the genus into two main subgenera. Pinus subg. Pinus, the hard or yellow pine group, carries cones with resin seals on the scales and generally harder wood. Pinus subg. Strobus, the soft pine and pinyon pine group, lacks that resin seal and tends toward softer wood. The two subgenera diverged from one another in the deep past, according to phylogenetic evidence. World Flora Online currently accepts 134 species-rank taxa within the genus, making it the largest genus among all the conifers. The greatest diversity of pine species on Earth is found in Mexico.

  • Pine wood has driven economies for centuries. It grows fast in relatively dense stands across temperate and tropical regions, producing softwood that is denser and more durable than spruce. The wood appears in furniture, window frames, flooring, panelling, and roofing because of its abundance and relatively low cost. When treated with compounds such as copper azole or chromated copper arsenate, it can withstand outdoor conditions; untreated, it is best confined to indoor framing, since it offers little resistance to insects or decay.

    The pine bolete, a basidiomycete fungus called Boletus pinophilus, forms ectomycorrhizal associations with several pine species including Pinus cembra, Pinus nigra, and Pinus sylvestris, trading minerals for sugars through the root system. Pine pollen also feeds detritivores and can help fungi break down nutrient-poor plant litter on the forest floor.

    Not all pine relationships are mutually beneficial. The pine processionary moth is one of the most serious pests of commercial pine forestry: its caterpillars can strip a tree of all its needles. The pine beauty moth preys on mature stands, and the sawfly Diprion pini poses a particular threat to Pinus sylvestris plantations. Nematode attacks can cause pine wilt disease, which kills infected trees quickly. Pinus radiata, introduced across the Southern Hemisphere for its timber value, has become invasive in some regions, establishing itself in habitats where it was never intended to grow.

  • Pine nuts, the seeds carried on the female cones, appear in cuisines across the world. The most famous preparation is pesto alla genovese, which uses them as a core ingredient. Young male cones can also be cooked and eaten, and the cambium can be dried and ground into a powder used as a flour substitute or as a thickener in soups and stews. Some Swedes brew what is called tallstrunt, a herbal tea made from young green needles steeped in boiling water. In Greece, the wine retsina is flavoured with resin from Pinus halepensis, the Aleppo pine.

    Beyond food, turpentine oil extracted from pine resin or pine wood has long served as a solvent in paints, resins, and varnishes. In traditional Chinese medicine, pine resin treats burns, wounds, and skin complaints. Chinese ink sticks for calligraphy are made from pine soot, which produces a matt black ink when mixed with water. Latvian designer Tamara Orjola has demonstrated a more contemporary application: using pine needles to create biodegradable paper, furniture, textiles, and dyestuffs. In Latin America, weavers have long incorporated pine needles into baskets.

  • In ancient Egypt, an image of the god Osiris was placed inside a cavity carved into a pine tree. Ancient Greek tradition linked pines to the goddess Pitthea, while in Rome the tree was worshipped during festivals honouring Attis and Cybele. The Greek god Dionysus carried the pine cone as a symbol of fertility: his followers bore a staff called a thyrsus, topped with a pine cone. Thousands of kilometres away and centuries later, the Buryat people of Siberia revered groves of Pinus sylvestris, and ancient Celtic Druids lit fires of the same species to mark the midwinter solstice.

    In Chinese art and literature, the pine appears repeatedly as a symbol of longevity and steadfastness, prized because it keeps its green needles through winter. It is paired sometimes with the cypress and sometimes with the plum and bamboo in a grouping called the "Three Friends of Winter".

    In 1924, the Italian composer Ottorino Respighi completed his tone poem Pines of Rome. Its four movements each portray a pine-clad location in the city: the Villa Borghese gardens, the setting near a catacomb, the Janiculum Hill, and the Appian Way. A 2021 study cataloguing pines in painting lists over a hundred works by artists including Paul Cezanne, Jean-Baptiste-Camille Corot, Akseli Gallen-Kallela, James William Giles, and Tom Thomson. Respighi's score remains one of the most performed orchestral pieces of the early twentieth century, a reminder that a genus rooted in the Cretaceous still shapes human culture today.

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Common questions

What is the oldest known pine tree?

The oldest known individual pine is a Great Basin bristlecone pine (Pinus longaeva) in the White Mountains of California, nicknamed "Methuselah", estimated to be around 4,800 years old. An older individual near Wheeler Peak was dated at 4,900 years but has since been cut down.

How tall can pine trees grow?

Most pine species reach between 15 and 45 metres tall. The tallest known individual is an 83.45-metre sugar pine (Pinus lambertiana) in Yosemite National Park. The smallest species are the Siberian dwarf pine and the Potosi pinyon.

What is serotiny in pine trees?

Serotiny is a strategy in which pine cones remain sealed for years until an environmental trigger causes them to open and release seeds. The most common form, pyriscence, uses resin to keep cones shut until a forest fire melts it, as seen in Pinus radiata and Pinus muricata.

Where do pine trees grow and how many species exist?

Pines are native to the Northern Hemisphere, with the highest species diversity in Mexico. World Flora Online accepts 134 species-rank taxa, making Pinus the largest genus among the conifers. One species, Pinus merkusii, crosses the equator into Sumatra at 2 degrees south.

What are pine trees used for besides timber?

Pine nuts are used in foods including pesto alla genovese, and the inner bark can be eaten raw or dried as a flour substitute. Pine resin flavours the Greek wine retsina and yields turpentine oil. Pine soot is used to make Chinese calligraphy ink sticks, and pine needles have been woven into baskets in Latin America.

What is the cultural significance of the pine tree in world history?

Pines held religious significance in ancient Egypt (associated with Osiris), ancient Greece (linked to Pitthea and Dionysus), and ancient Rome (worshipped in festivals of Attis and Cybele). The Buryat people of Siberia and Celtic Druids also revered Pinus sylvestris. In Chinese tradition the pine symbolises longevity, and in 1924 Italian composer Ottorino Respighi dedicated his tone poem Pines of Rome to pine-clad settings in the city.

All sources

80 references cited across the entry

  1. 1JournalPalaeoflora of Southern Africa. Prodromus of South African Megafloras Devonian to Lower CretaceousJ. M., H. M. Anderson — 1985
  2. 3NewsTallest of the tallPaul Fattig — 23 January 2011
  3. 5JournalThe Complete PineMichael Ryan et al. — December 1999
  4. 6NewsFollow-up: More tales of the Prometheus tree and how it diedCarolina A. Miranda — 28 February 2015
  5. 7MagazineHow One Man Accidentally Killed the Oldest Tree EverRose Eveleth — Smithsonian Institution — 15 November 2012
  6. 8JournalModeling golden section in plantsLanling Zeng et al. — 2009
  7. 9Pinus (Pine) NotesRonald Bracewell et al.
  8. 10BookRoyal Horticultural Society Dictionary of GardeningMacmillan Press; Stockton Press — 1992
  9. 11BookPlant systematics, a phylogenetic approachW.S. Judd et al. — Sinauer Associates — 2002
  10. 12JournalDispersal of Whitebark Pine seeds by Clark's Nutcracker: a mutualism hypothesisDiana F. Tomback — June 1982
  11. 13BookConifersKeith Rushforth — Christopher Helm Publishers — 1987
  12. 15BookTrees and Bushes in Wood and HedgerowHelge Vedel et al. — Methuen — 1960
  13. 16BookSpecies plantarum: exhibentes plantas rite cognitas, ad genera relatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas. Tomus I & II.Carolus Linnaeus — Impensis Laurentii Salvii — 1753
  14. 17BookEcology and Biogeography of PinusR. A. Price et al. — Cambridge University Press — 1998
  15. 19JournalPhylogeny and evolutionary history of Pinaceae updated by transcriptomic analysisJin-Hua Ran et al. — December 2018
  16. 20JournalReconsidering Relationships among Stem and Crown Group Pinaceae: Oldest Record of the Genus Pinus from the Early Cretaceous of Yorkshire, United KingdomPatricia E. Ryberg et al. — 2012
  17. 22JournalPhylogenetic relationships and species delimitation in Pinus section Trifoliae inferred from plastid DNASergio Hernández-León et al. — 30 July 2013
  18. 23JournalGenetic, morphological, geographical and ecological approaches reveal phylogenetic relationships in complex groups, an example of recently diverged pinyon pine species (Subsection Cembroides)Lluvia Flores-Rentería et al. — December 2013
  19. 24JournalPhylogenomic and ecological analyses reveal the spatiotemporal evolution of global pinesWei-Tao Jin et al. — May 2021
  20. 26JournalPhylogeny and classification of PinusDavid S. Gernandt et al. — 2005
  21. 29Genetic diversity and conservation of Mexican forest treesC. Wehenkel et al. — US Department of Agriculture
  22. 31JournalCurrent and Potential Spatial Distribution of Six Endangered Pine Species of Mexico: Towards a Conservation StrategyMartin Enrique Romero-Sanchez et al. — 2018
  23. 33The forest biomeUniversity of California Museum of Paleontology
  24. 35BookGeographic Distribution of the Pines of the WorldWilliam B. Critchfield et al. — USDA Forest Service — 1966
  25. 36JournalInsights on the persistence of pines (Pinus species) in the Late Cretaceous and their increasing dominance in the AnthropoceneSurendra P. Singh et al. — 21 September 2018
  26. 37Pinus ssp. (tree), General ImpactInvasive Species Specialist Group — 13 March 2006
  27. 38Pine Trees30 August 2020
  28. 39JournalThe Interaction of Fire, Fuels and Climate across Rocky Mountain ForestsTania Schoennagel et al. — 2004
  29. 41JournalResprouting as a key functional trait: how buds, protection and resources drive persistence after fireP. J. Clarke et al. — January 2013
  30. 42BookNational Wildlife Federation Field Guide to Trees of North AmericaMoore et al. — Sterling — 2008
  31. 44JournalGenetic Variation Related to High Elevation Adaptation Revealed by Common Garden Experiments in Pinus yunnanensisYan-Qiang Sun et al. — 11 February 2020
  32. 46JournalThe history and control of the pine beauty moth, Panolis flammea (D. & S.) (Lepidoptera: Noctuidae), in Scotland from 1976 to 2000B. J. Hicks et al. — 2001
  33. 49JournalLa Processionnaire du PinCatherine Bonnet et al. — INRA — 2008
  34. 50JournalQuaternary history and contemporary patterns in a currently expanding speciesCarole Kerdelhué et al. — 2009
  35. 54Diprion piniJuly 2015
  36. 55BookMade for each other: A symbiosis of birds and pinesRonald M. Lanner — Oxford University Press — 1996
  37. 56BookWhy birds matter: avian ecological function and ecosystem servicesDiana F. Tomback — University of Chicago Press — 2016
  38. 57JournalPatterns of exploitation of annually varying Pinus sylvestris cone crops by seed-eaters of differing dispersal abilityRon W. Summers — 2011
  39. 59JournalNationalization and globalization trends in the wild mushroom commerce of Italy with emphasis on porcini (Boletus edulis and allied species)Nicola Sitta et al. — 2008
  40. 60BookMushrooms, Humans and Nature in a Changing World: Perspectives from Ecological, Agricultural and Social SciencesMatteo Gallardi — Springer — 2020
  41. 65Timber treatmentweathertight.org.nz — 18 October 2010
  42. 66BookThe Complete Guide to Edible Wild PlantsSkyhorse Publishing — 2009
  43. 67Pesto's official recipeCampionato Mondiale Pesto al Mortaio
  44. 69BookField Guide to Edible Wild PlantsBradford Angier — Stackpole Books — 1974
  45. 70JournalAntibrowning and antimicrobial activities of the water-soluble extract from pine needles of Cedrus deodaraWei-Cai Zeng et al. — March 2011
  46. 71BookThe Oxford Companion to WineOxford University Press — 2006
  47. 72JournalExtraction of turpentine essential oil from Pinus pinaster ait: Comparison of yield and composition between conventional- or microwave assisted-hydro-distillation and vacuum distillationPascale Chalier et al. — 2024
  48. 73Turpentine Production and ProcessingNew Zealand Institute of Chemistry
  49. 74Culture TrunksCollege of Liberal Arts, University of Texas
  50. 75JournalChemical composition, antimicrobial, insecticidal, phytotoxic and antioxidant activities of Mediterranean Pinus brutia and Pinus pinea resin essential oilsZeynep Ulukanli et al. — December 2014
  51. 79The March of Time: Respighi's Pines of RomeCalvin Dotsey — Houston Symphony — 7 January 2020
  52. 80JournalLes Pins vus par les peintresJean Pinon — 2021