Pine

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A pine is any conifer in the genus Pinus (Template:IPAc-en<ref>Template:Cite encyclopedia</ref>) of the family Pinaceae. Pinus is the sole genus in the subfamily Pinoideae. The species are evergreen trees or shrubs with their leaves in bunches, usually of 2 to 5 needles. The seeds are carried on woody cones, with two seeds to each cone scale.

Pines are widely distributed in the Northern Hemisphere; they occupy large areas of taiga (boreal forest), but are found in many habitats, including the Mediterranean Basin, and dry tropical forests in southeast Asia and Central America. Some are fire-resistant or fire-dependent.

Pine trees provide one of the most extensively used types of timber. The seeds are used to make dishes such as pesto, while retsina wine is flavoured with pine resin.

Description

Tree

Pine trees are evergreen, coniferous resinous trees (or, rarely, shrubs) growing Template:Convert tall, with the majority of species reaching Template:Convert tall.<ref name="Fattig-2011">Template:Cite news</ref> The smallest are Siberian dwarf pine and Potosi pinyon, and the tallest is an Template:Convert tall sugar pine (Pinus lambertiana) located in Yosemite National Park.<ref name="Associated Press-2021">Template:Cite news</ref>

Pines are long lived and typically reach ages of 100–1,000 years, some even more. The longest-lived is the Great Basin bristlecone pine (Pinus longaeva). One individual in the White Mountains of California, dubbed "Methuselah", is among the world's oldest living organisms at around 4,800-years old.<ref name="Ryan-1999">Template:Cite journal</ref> An older tree near Wheeler Peak, now cut down, was dated at 4,900-years old.<ref>Template:Cite news</ref><ref name="Eveleth-2012">Template:Cite magazine</ref>

The spirals of branches, needles, and cone scales are arranged in Fibonacci number ratios.<ref>Template:Cite journal</ref><ref>Template:Cite web</ref>

Bark

The bark of most pines is thick and scaly, but some species have thin, flaky bark. The branches are produced in "pseudo-whorls", actually a very tight spiral but appearing like a ring of branches arising from the same point. Many pines are uninodal, producing just one such whorl of branches each year, from buds at the tip of the year's new shoot, but others are multinodal, producing two or more whorls of branches per year.<ref name="RHS-Dict-1992">Template:Cite book</ref>

Foliage

Pines have four types of leaf:<ref name="RHS-Dict-1992"/>

  • Seed leaves (cotyledons) on seedlings are borne in a whorl of 4–24.
  • Juvenile leaves, which follow immediately on seedlings and young plants, are Template:Convert long, single, green or often blue-green, and arranged spirally on the shoot. These are produced for six months to five years, rarely longer.
  • Scale leaves, similar to bud scales, are small, brown and not photosynthetic, and arranged spirally like the juvenile leaves.
  • Needles, the adult leaves, are green (photosynthetic) and bundled in clusters called fascicles. The needles can number from one to seven per fascicle, but generally number from two to five. Each fascicle is produced from a small bud on a dwarf shoot in the axil of a scale leaf. These bud scales often remain on the fascicle as a basal sheath. The needles persist for 1.5–40 years, depending on species. If a shoot's growing tip is damaged (e.g. eaten by an animal), the needle fascicles just below the damage generate a stem-producing bud, which can then replace the lost growth tip.

Cones

Pines are monoecious, having the male and female cones on the same tree.<ref name="Judd-2002">Template:Cite book</ref>Template:Rp The male cones are small, typically 1–5 cm long, and only present for a short period (usually in spring, though autumn in a few pines), falling as soon as they have shed their pollen. The female cones take 1.5–3 years (depending on species) to mature after pollination, with actual fertilisation delayed one year. At maturity, the female cones are 3–60 cm long. Each cone has numerous spirally-arranged scales, with two seeds on each fertile scale; the scales at the base and tip of the cone are small and sterile without seeds.<ref name="RHS-Dict-1992"/>

The seeds (pine nuts) are mostly small and winged, and are anemochorous (wind-dispersed). Some are larger, have only a vestigial wing, and are bird-dispersed. Female cones are woody and sometimes armed to protect developing seeds from foragers. At maturity, the cones usually open to release the seeds. In some of the bird-dispersed species, for example whitebark pine,<ref>Template:Cite journal</ref> the seeds are only released by the bird breaking the cones open. In others, the seeds are stored in closed cones for many years until an environmental cue triggers the cones to open, releasing the seeds. This is called serotiny. The most common form of serotiny is pyriscence, in which resin binds the cones shut until the resin is melted by a forest fire, for example in P. radiata and P. muricata. The seeds are then released after the fire, enabling them to colonise the burnt ground with minimal competition from other plants.<ref name="RHS-Dict-1992"/><ref name="Rushforth-1987">Template:Cite book</ref>

Naming

The modern English name "pine" derives from Latin pinus, traced to the Indo-European base *pīt- 'resin'.<ref>Template:Cite web</ref> Before the 19th century, pines were often called firs, a name now applied to another genus, Abies. In some European languages, Germanic cognates of the Old Norse name are still in use for pines, as in Danish fyr and German Föhre.<ref>Template:Cite book</ref> The genus Pinus was named by Carl Linnaeus in 1753. Pinus sylvestris, the Scots pine, was later chosen as the type species.<ref>Template:Cite book</ref><ref>Template:Cite book</ref>

Evolution

Fossil history

The Pinaceae, the pine family, first appeared in the Jurassic period.<ref>Template:Cite journal</ref> The genus Pinus first appeared during the Early Cretaceous; the oldest verified fossil is Pinus yorkshirensis from the Hauterivian-Barremian boundary (~130-125 million years ago) from the Speeton Clay, England.<ref>Template:Cite journal</ref> However, there are possible records of the genus from the Jurassic.<ref>Template:Cite web</ref>

External phylogeny

Based on transcriptome analysis, Pinus is most closely related to the genus Cathaya, which in turn is closely related to the genus Picea, the spruces. These genera, with firs and larches, form the pinoid clade of the Pinaceae.<ref name="Stull-2021"/>

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Internal phylogeny

The evolutionary history of the genus Pinus has been complicated by hybridisation. Pines are prone to inter-specific breeding. Wind pollination, long life spans, overlapping generations, large population size, and weak reproductive isolation make breeding across species more likely. As the pines have diversified, gene transfer between different species has created a complex history of genetic relatedness.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> Research using large genetic datasets has clarified these relationships.<ref name="Gernandt-2005"/> Two 21st century phylogenies are given below; the differences between them demonstrate these complications:

Jin et al. 2021<ref>Template:Cite journal</ref> Stull et al. 2021<ref name="Stull-2021"/>

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Taxonomy

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Pines are gymnosperms. The genus is divided into two subgenera based on the number of fibrovascular bundles in the needle, and the presence or absence of a resin seal on the scales of the mature cones before opening. The subgenera can be distinguished by cone, seed, and leaf characters:<ref name="RHS-Dict-1992"/>

  • Pinus subg. Pinus, the yellow, or hard pine group, with cones with a resin seal on the scales, and generally with harder wood; the needle fascicles mostly have a persistent sheath (two exceptions, Pinus leiophylla and Pinus lumholtzii, have deciduous sheaths).<ref name="RHS-Dict-1992"/>
  • Pinus subg. Strobus, syn. Pinus subg. Ducampopinus, the white or soft pine, and pinyon pine groups, with cones without a resin seal on the scales, and usually have softer wood; the needle fascicles mostly have a deciduous sheath (one exception, Pinus nelsonii, has a persistent sheath).<ref name="RHS-Dict-1992"/>

Phylogenetic evidence indicates that the subgenera diverged anciently from one another.<ref name="Stull-2021">Template:Cite journal</ref> Each subgenus is further divided into sections and subsections.<ref name="Gernandt-2005">Template:Cite journal</ref>

World Flora Online accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as current, with additional synonyms,<ref name="World Flora Online-2022">Template:Cite web</ref> and Plants of the World Online 126 species-rank taxa (113 species and 13 nothospecies),<ref name="Plants of the World Online-2013">Template:Cite web</ref> making it the largest genus among the conifers. The highest species diversity of pines is found in Mexico.<ref>Template:Cite web</ref><ref>Template:Cite journal</ref><ref>Template:Cite journal</ref>

Distribution

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Pines are native to the Northern Hemisphere, with the most species in North America, some in Asia, and a few in Europe. Only two species, Pinus sylvestris and Pinus sibirica, occur in more than one of those regions (Asia and Europe).<ref name="Nobis-2012">Template:Cite journal</ref> They occupy large areas of boreal forest (taiga) in latitudes between 50° and 60° N; about a third of this biome is in North America and Scandinavia, the rest in Siberia.<ref name="UCMP">Template:Cite web</ref> The northernmost species is Scots pine, reaching just north of 70° N in Stabbursdalen National Park in Norway;<ref name="Stabbursdalen Nasjonalpark">Template:Cite web</ref> One species, Pinus merkusii, crosses the equator in Sumatra to 2°S.<ref name="Critchfield-1966">Template:Cite book</ref> In North America, various species occur in regions at latitudes from as far north as 66° N<ref name="Critchfield-1966"/> to as far south as 12°N.<ref>Template:Cite journal</ref>

Various species have been introduced to temperate and subtropical regions of both hemispheres, where they are grown as timber or cultivated as ornamental plants in parks and gardens. A number of such introduced species have become naturalised, and species such as Pinus radiata are considered invasive in some regions.<ref name="Global Invasive Species Database-2006">Template:Cite web</ref>

Ecology

Environmental factors

Pines grow in a very large variety of environments, ranging from semi-arid desert to rainforests, from sea level up to Template:Convert, from the coldest to the hottest environments on Earth. They often occur in mountainous areas with favourable soils.<ref>Template:Cite web</ref>

Pinus contorta is a fire-dependent species, requiring wildfires to maintain healthy populations of diverse ages.<ref name="Schoennagel-2004">Template:Cite journal</ref> Pinus canariensis is highly fire-resistant,<ref>Template:Cite web</ref> with adaptations such as growing epicormic sprouts after losing its needles in a fire.<ref>Template:Cite journal</ref> Some species such as Pinus muricata need fire to open their cones, allowing them to disperse their seeds.<ref name="Block, Andrew-2008">Template:Cite book</ref> Other pines such as Pinus mugo<ref name="Solár-2013">Template:Cite journal</ref> and Pinus yunnanensis can grow at high elevation.<ref name="Sun-2020">Template:Cite journal</ref> Some pines, such as Pinus sabiniana, are adapted to growth in hot, dry semidesert climates.<ref>Template:Cite web</ref>

Species interactions

Pine needles serve as food for the caterpillar larvae of several moth species including the pine beauty, a pest of mature stands of pine trees,<ref>Template:Cite journal</ref> and the pine hawk-moth, a large species which causes only occasional damage.<ref>Template:Cite web</ref> Some moths, notably the pine processionary, whose caterpillars can completely defoliate pine trees,<ref>Template:Cite web</ref><ref name="Bonnet-2008">Template:Cite journal</ref><ref name="Kerdelhué-2009">Template:Cite journal Template:Open access</ref> and the pine-tree lappet, are serious pests of commercial forestry.<ref>Template:Cite web</ref>

Several species of pine are attacked by nematodes, causing pine wilt disease, which can quickly kill trees.<ref>Bursaphelenchus xylophilus, Pine Wilt Nematode. Nematology. University of Nebraska, Lincoln.</ref><ref name="Bursaphelenchus">Bursaphelenchus xylophilus. Template:Webarchive Nemaplex. UC Davis.</ref> The sawfly Diprion pini is likewise a serious commercial pest of pine forestry, especially of Pinus sylvestris.<ref name="CAPS-2015">Template:Cite web</ref> Some birds such as nutcrackers are specialist feeders on pine seeds, and are important in distributing the seeds widely.<ref>Template:Cite book</ref><ref>Template:Cite book</ref> Crossbills rely on Pinus sylvestris seeds in Scotland, and similarly help significantly to disperse the seeds, whereas red squirrels feed on the seeds but do little for seed dispersal.<ref name="Summers-2011">Template:Cite journal</ref> Pine pollen may contribute to food webs involving detritivores. Nutrients from pollen aid detritivores in development, growth, and maturation, and may enable fungi to decompose plant litter which is low in nutrients.<ref name="Filipiak-2016">Template:Cite journal</ref> The edible<ref name="Sitta-2008">Template:Cite journal</ref> basidiomycete fungus Boletus pinophilus (pine bolete) forms an ectomycorrhizal association with pines such as P. cembra, P. nigra, and P. sylvestris.<ref name="Gallardi-2020">Template:Cite book</ref>

Uses

Timber

Pines are among the most commercially important tree species, valued for their timber and wood pulp throughout the world.<ref>Template:Cite news</ref><ref>Template:Cite web</ref> In temperate and tropical regions, they are fast-growing softwoods that grow in relatively dense stands. Commercial pines are grown in plantations for timber that is denser and therefore more durable than spruce (Picea). Pine wood is widely used in high-value carpentry items such as furniture, window frames, panelling, floors, and roofing due to its abundance and low cost.<ref>Wiemann, M. C. (2010). Characteristics and Availability of Commercially Important Woods. In Wood handbook: Wood as an engineering material (pp. 2-2-2–45). Chapter, Forest Products Laboratory; For sale by the Supt. of Docs., U.S. Govt. https://research.fs.usda.gov/treesearch/37440</ref>

As pine wood has little resistance to insects or decay after logging, in its untreated state it is generally recommended for indoor construction purposes only, such as indoor drywall framing. It is commonly used in Canadian Lumber Standard graded wood.<ref name="Jenkins-2023">Template:Cite web</ref> For outside use, pine needs to be treated with copper azole, chromated copper arsenate or other suitable chemical preservatives.<ref>Template:Cite web</ref>

Ornamental uses

Many pine trees make attractive ornamental plantings for parks and larger gardens, while dwarf cultivars are suitable for smaller spaces. There are at least 818 named cultivars (or trinomials) recognised by the American Conifer Society ACS.<ref name="American Conifer Society"/>

Food

The seeds (pine nuts) are generally edible; the young male cones can be cooked and eaten, as can the bark of young twigs.<ref>Template:Cite book</ref> Some species have large pine nuts, which are harvested and sold for cooking and baking. They are an ingredient of pesto alla genovese.<ref name="CMPM">Template:Cite web</ref>

The soft, moist, white inner bark (cambium) beneath the woody outer bark is edible and very high in vitamins A and C.<ref name="American Conifer Society">Template:Cite web</ref> It can be eaten raw in slices as a snack or dried and ground up into a powder for use as an ersatz flour or thickener in stews, soups, and other foods, such as bark bread.<ref name="Angier-1974">Template:Cite book</ref> The use of pine cambium gave the Adirondack Indians their name, from the Mohawk Indian word atirú:taks, meaning "tree eaters".<ref name="Angier-1974"/>

A herbal tea is made by steeping young, green pine needles in boiling water (known as tallstrunt in Sweden).<ref name="Angier-1974" /> In eastern Asia, pine and other conifers are accepted among consumers as a beverage product, and used in teas, as well as wine.<ref name="Zeng-2011">Template:Cite journal</ref> In Greece, the wine retsina is flavoured with resin from Pinus halepensis (Aleppo pine).<ref name="Robinson-2006">Template:Cite book</ref>

Other uses

Turpentine oil, traditionally used as a solvent in paints, resins and varnishes, is extracted from pine resin<ref name="Chalier-2024">Template:Cite journal</ref> or pine wood.<ref>Template:Cite web</ref> Pine needles are woven into baskets in Latin America.<ref>Template:Cite web</ref> In traditional Chinese medicine, pine resin is used for burns, wounds and skin complaints.<ref>Template:Cite journal</ref> Chinese ink sticks for calligraphy are often made of pine soot, producing a matt black ink when mixed with water.<ref>Template:Cite web</ref> Pine needles have been used by Latvian designer Tamara Orjola to create biodegradable products including paper, furniture, textiles and dyestuffs.<ref>Template:Cite web</ref>

Culture

File:Villa Borghese Gardens 1 (5895905875).jpg
A movement of Respighi's tone poem Pines of Rome depicts the stone pines of the Villa Borghese gardens.<ref name="Dotsey 2020"/>

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In ancient Egypt, the god Osiris was honoured with an image placed in a cavity inside a pine tree. In ancient Greece, the goddess Pitthea was linked with pines, while in ancient Rome, the tree was worshipped in the festival of the god Attis and the goddess Cybele.<ref name="Trees for Life"/> The Greek god of wine, Dionysus (also called Bacchus), was associated with pine as a symbol of fertility, and his devotees carried a stick topped with a pine cone (a thyrsus), a phallic symbol.<ref name="Trees for Life"/> The Buryat people of Siberia revered groves of Pinus sylvestris, while ancient Celtic Druids marked the midwinter solstice with fires of the same species.<ref name="Trees for Life">Template:Cite web</ref>

In 1924, the Italian composer Ottorino Respighi completed his tone poem Pines of Rome. Each of its four movements depicts a pine-clad setting in the city of Rome, namely the Villa Borghese gardens, near a catacomb, on the Janiculum Hill, and along the Appian Way.<ref name="Dotsey 2020">Template:Cite web</ref>

Pines are often featured in paintings. A 2021 study lists over a hundred works: many are by artists from the Mediterranean region, such as Paul Cézanne and Jean-Baptiste-Camille Corot; Northern Europe, such as Akseli Gallen-Kallela and James William Giles; and North America, with works by Tom Thomson and others. The paintings often depict Pinus pinea by the Mediterranean sea; other species include P. sylvestris and P. pinaster.<ref name="Pinon-2021">Template:Cite journal</ref> The pine is a particular motif in Chinese art and literature, which sometimes combines painting and poetry in the same work. The pine symbolises longevity and steadfastness, as it retains its green needles throughout the year. Sometimes the pine and cypress are paired. At other times the pine, plum, and bamboo are considered as the "Three Friends of Winter".<ref>Eberhard, Wolfram (2003 [1986 (German version 1983)]), A Dictionary of Chinese Symbols: Hidden Symbols in Chinese Life and Thought. London, New York: Routledge. Template:ISBN, sub "Pine".</ref>

See also

References

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Bibliography

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