Pine

Pine Temporal range: Barremian–Recent PreꞒ Ꞓ O S D C P T J K Pg N
Pinus densiflora (Korean red pine), North Korea
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Gymnospermae
Division:	Pinophyta
Class:	Pinopsida
Order:	Pinales
Family:	Pinaceae
Subfamily:	Pinoideae
Genus:	Pinus L.
Type species
L.
Subgenera
Subgenus Strobus Subgenus Pinus See List of Pinus species for complete taxonomy to species level. See list of pines by region for list of species by geographic distribution.
Range of Pinus
Synonyms
Apinus de Necker ex Rydberg Caryopitys Small Cembra Opiz Ducampopinus Chevalier Haploxylon (Koehne) Komarov Leucopitys Nieuwland Pinea Wolf ex Opiz Strobus (Sweet ex Spach) Opiz

A pine is any conifer tree or shrub in the genus Pinus (/ˈpaɪnuːs/)^[1] of the family Pinaceae. Pinus is the sole genus in the subfamily Pinoideae.

World Flora Online, created by the Royal Botanic Gardens, Kew, and Missouri Botanical Garden, accepts 187 species names of pines as current, together with more synonyms.^[2] The American Conifer Society (ACS) and the Royal Horticultural Society accept 121 species.

Pines are commonly found in the Northern Hemisphere.

Pine may also refer to the lumber derived from pine trees; it is one of the more extensively used types of lumber. The pine family is the largest conifer family, and there are currently 818 named cultivars (or trinomials) recognized by the ACS.^[3] It is also a well-known type of Christmas tree.

Description

Pine trees are

Pines are long lived and typically reach ages of 100–1,000 years, some even more. The longest-lived is the

The spiral growth of branches, needles, and

of the soil and vigour of the trees.

Bark

The bark of most pines is thick and scaly, but some species have thin, flaky bark.^[10] The branches are produced in regular "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.

Foliage

Pines have four types of leaf:

• Seed leaves (cotyledons) on seedlings are borne in a whorl of 4–24.
• Juvenile leaves, which follow immediately on seedlings and young plants, are 2–6 centimetres (3⁄4–2+1⁄4 inches) 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 (
  growing tip
  is damaged (e.g. eaten by an animal), the needle fascicles just below the damage will generate a stem-producing bud, which can then replace the lost growth tip.

Cones

Pines are

, with actual fertilization 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.

The seeds are mostly small and winged, and are

.

Taxonomy

Pines are gymnosperms. The genus is divided into two subgenera based on the number of fibrovascular bundles in the needle. The subgenera can be distinguished by cone, seed, and leaf characters:

• Pinus subg. Pinus, the yellow, or hard pine group, generally with harder wood and two or three needles per fascicle.^[13]
  The subgenus is also named diploxylon, on account of its two fibrovascular bundles.
• Pinus subg. Strobus, the white, or soft pine group. Its members usually have softer wood and five needles per fascicle.^[13]
  The subgenus is also named haploxylon, on account of its one fibrovascular bundle.

Phylogenetic evidence indicates that both subgenera have a very ancient divergence from one another.[14] Each subgenus is further divided into sections and subsections.

Many of the smaller groups of Pinus are composed of closely related species with recent divergence and history of hybridization. This results in low morphological and genetic differences. This, coupled with low sampling and underdeveloped genetic techniques, has made taxonomy difficult to determine.^[15] Recent research using large genetic datasets has clarified these relationships into the groupings we recognize today.

Etymology

The modern English name "pine" derives from Latin pinus, which some have traced to the Indo-European base *pīt- ‘resin’ (source of English pituitary).

(Pseudotsuga).

Phylogeny

Pinus is the largest genus of the Pinaceae, the pine family, which first appeared in the Jurassic period.^[17] Based on recent Transcriptome analysis, Pinus is most closely related to the genus Cathaya, which in turn is closely related to spruces. These genera, with firs and larches, form the pinoid clade of the Pinaceae.^[14] Pines first appeared during the Early Cretaceous, with the oldest verified fossil of the genus being Pinus yorkshirensis from the Hauterivian-Barremian boundary (~130-125 million years ago) from the Speeton Clay, England.^[18]

The evolutionary history of the genus Pinus has been complicated by hybridization. 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.^[19] As the pines have diversified, gene transfer between different species has created a complex history of genetic relatedness.

Jin et al. 2021[20]

Stull et al. 2021[14]

Pinus subgenus Strobus section Parrya subsection Nelsonii P. nelsonii subsection Balfouriana P. aristata P. balfouriana P. longaeva subsection Cembroides P. pinceana P. maximartinezii P. rzedowskii P. quadrifolia P. monophylla P. culminicola P. discolor P. remota P. edulis P. cembroides section Quinquefoliae subsection Gerardiana P. bungeana P. squamata P. gerardiana subsection Krempfii P. krempfii subsection Strobus P. peuce P. lambertiana P. strobus P. chiapensis P. monticola P. flexilis P. strobiformis P. ayacahuite P. albicaulis P. koraiensis P. sibirica P. cembra P. parviflora P. wallichiana P. armandii P. dabeshanensis P. fenzeliana P. mastersiana P. bhutanica P. morrisonicola P. wangii P. kwangtungensis P. dalatensis subgenus Pinus section Pinus subsection Pinaster P. heldreichii P. pinaster P. halepensis P. brutia P. pinea P. canariensis P. roxburghii subsection Pinus P. latteri P. merkusii P. resinosa P. nigra P. densiflora P. sylvestris P. mugo P. uncinata P. massoniana P. thunbergii P. luchuensis P. kesiya P. yunnanensis P. densata P. henryi P. tabuliformis P. hwangshanensis P. taiwanensis section Trifoliae subsection Sabiniana P. jeffreyi P. coulteri P. torreyana P. sabiniana subsection Ponderosae P. washoensis P. ponderosa P. hartwegii P. engelmannii P. durangensis P. arizonica P. devoniana P. montezumae P. pseudostrobus P. douglasiana P. maximinoi subsection Attenuata P. radiata P. attenuata P. muricata subsection Contorta P. contorta P. banksiana P. virginiana P. clausa subsection Australes P. pungens P. serotina P. rigida P. glabra P. taeda P. echinata P. elliottii P. palustris P. caribaea P. oocarpa P. tecunumanii P. greggii P. patula P. teocote P. lawsonii P. herrerae P. pringlei P. leiophylla P. lumholtzii P. praetermissa P. jaliscana P. luzmariae P. georginae

Pinus (Strobus) section subsection P. nelsonii Nelsonii subsection P. aristata P. balfouriana P. longaeva Balfouriana subsection P. monophylla P. juarezensis P. quadrifolia P. pinceana P. maximartinezii P. rzedowskii P. edulis P. discolor P. johannis P. culminicola P. cembroides P. remota Cembroides Parrya section subsection P. bungeana P. gerardiana Gerardiana subsection P. krempfii Krempfii subsection series P. peuce Macedonia series P. strobus P. flexilis P. ayacahuite P. strobiformis Strobi series P. parviflora P. monticola P. cembra P. lambertiana P. albicaulis P. dalatensis Cembrae series P. pumila P. wallichiana P. koraiensis P. sibirica P. squamata P. morrisonicola P. armandii P. fenzeliana P. kwangtungensis P. bhutanica P. wangii Pumilae Strobus Quinquefoliae (Pinus) section subsection P. heldreichii P. halepensis P. brutia P. pinaster P. pinea P. canariensis P. roxburghii Pinaster subsection series P. densata P. latteri P. kesiya P. nigra P. resinosa P. sylvestris P. mugo P. uncinata Pinus series P. tropicalis P. hwangshanensis P. massoniana P. merkusii Tropicales series P. densiflora P. thunbergii P. taiwanensis P. yunnanensis P. luchuensis P. tabuliformis Tabuliformes Pinus Pinus section subsection P. virginiana P. contorta P. clausa Contorta subsection P. maximinoi P. montezumae P. sabiniana P. torreyana P. ponderosa P. douglasiana P. yecorensis P. cooperi P. durangensis P. nubicola P. coulteri P. jeffreyi P. washoensis P. arizonica P. engelmannii P. devoniana P. hartwegii P. pseudostrobus Ponderosae subsection series P. patula P. leiophylla Leiophyllae series P. tecunumanii P. muricata P. radiata P. attenuata P. glabra Attenuatae Attenuata subsection series P. herrerae P. caribaea P. palustris P. echinata P. occidentalis Palustres P. praetermissa series P. serotina P. rigida P. pungens P. taeda Taeda series P. elliottii P. teocote P. greggii P. lumholtzii P. jaliscana P. luzmariae P. georginae P. maestrensis P. pringlei P. lawsonii P. cubensis P. oocarpa Oocarpae Australes Trifoliae

Distribution and habitat

Monterey pine in Sydney, Australia, which were introduced

to the region in the late 19th century.

Pines are native to the

Pines may be found in a very large variety of environments, ranging from semi-arid desert to rainforests, from sea level up to 5,200 m (17,100 ft), from the coldest to the hottest environments on Earth. They often occur in mountainous areas with favorable soils and at least some water.[22]

Various species have been introduced to temperate and

and threaten native ecosystems.

Ecology

Pines grow well in acid soils, some also on

) need fire to regenerate, and their populations slowly decline under fire suppression regimens.

Pine trees are beneficial to the environment since they can remove carbon dioxide from the atmosphere. Although several studies have indicated that after the establishment of pine plantations in grasslands, there is an alteration of carbon pools including a decrease of the soil organic carbon pool.^[24]

Several species are adapted to extreme conditions imposed by elevation and latitude (e.g. Siberian dwarf pine,

Pine pollen may play an important role in the functioning of detrital food webs.^[26] Nutrients from pollen aid detritivores in development, growth, and maturation, and may enable fungi to decompose nutritionally scarce litter.^[26] Pine pollen is also involved in moving plant matter between terrestrial and aquatic ecosystems.^[26]

Wildlife

Pine needles serve as food for various

.

The seeds are commonly eaten by birds, such as grouse, crossbills, jays, nuthatches, siskins, and woodpeckers, and by

Uses

Lumber and construction

Pines are among the most commercially important tree species valued for their timber and wood pulp throughout the world.^[28][29] In temperate and tropical regions, they are fast-growing softwoods that grow in relatively dense stands, their acidic decaying needles inhibiting the sprouting of competing hardwoods.^{[citation needed]} 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, and the resin of some species is an important source of turpentine.

Because pine wood has no insect- or decay-resistant qualities after logging, in its untreated state it is generally recommended for indoor construction purposes only (indoor drywall framing, for example). For outside use, pine needs to be treated with copper azole, chromated copper arsenate or other suitable chemical preservative.^[30]

Ornamental uses

Many pine species make attractive ornamental plantings for

Farming

When grown for sawing timber, pine plantations can be harvested after 25 years, with some stands being allowed to grow up to 50 (as the wood value increases more quickly as the trees age). Imperfect trees (such as those with bent trunks or forks, smaller trees, or diseased trees) are removed in a "thinning" operation every 5–10 years. Thinning allows the best trees to grow much faster, because it prevents weaker trees from competing for sunlight, water, and nutrients. Young trees removed during thinning are used for pulpwood or are left in the forest, while most older ones are good enough for saw timber.[34]

A 30-year-old commercial pine tree grown in good conditions in Arkansas will be about 0.3 m (1 ft) in diameter and about 20 m (66 ft) high. After 50 years, the same tree will be about 0.5 m (1+1⁄2 ft) in diameter and 25 m (82 ft) high, and its wood will be worth about seven times as much as the 30-year-old tree. This however depends on the region, species and silvicultural techniques. In New Zealand, a plantation's maximum value is reached after around 28 years with height being as high as 30 m (98 ft) and diameter 0.5 m (1+1⁄2 ft), with maximum wood production after around 35 years (again depending on factors such as site, stocking and genetics). Trees are normally planted 3–4 m apart, or about 1,000 per hectare (100,000 per square kilometre).^{[35][36][37][38]}

Food and nutrients

The seeds (pine nuts) are generally edible; the young male cones can be cooked and eaten, as can the bark of young twigs.^[39] Some species have large pine nuts, which are harvested and sold for cooking and baking. They are an essential ingredient of pesto alla genovese.

The soft, moist, white inner bark (

A tea is made by steeping young, green pine needles in boiling water (known as tallstrunt in Sweden).^[40] In eastern Asia, pine and other conifers are accepted among consumers as a beverage product, and used in teas, as well as wine.^[41] In Greece, the wine retsina is flavoured with Aleppo pine resin.

Pine needles from Pinus densiflora were found to contain 30.54 milligram/gram of proanthocyanidins when extracted with hot water.^[42] Comparative to ethanol extraction resulting in 30.11 mg/g, simply extracting in hot water is preferable.

In

Culture

Pines have been a frequently mentioned tree throughout history, including in literature, paintings and other art, and in religious texts.

Literature

Writers of various nationalities and ethnicities have written of pines. Among them, John Muir,^[44] Dora Sigerson Shorter,^[45] Eugene Field,^[46] Bai Juyi,^[47] Theodore Winthrop,^[48] and Rev. George Allan D.D.^[49]

Art

Algonquin Park

, Ontario

Pines are often featured in art, whether painting and fine art,^[50] drawing,^[51] photography, or folk art.

Religious texts

Pine trees, as well as other conifers, are mentioned in some verses of the Bible, depending on the translation. In the Book of Nehemiah 8:15, the King James Version gives the following translation:^[52]

"And that they should publish and proclaim in all their cities, and in Jerusalem, saying, Go forth unto the mount, and fetch olive branches, and pine branches [emphasis added], and myrtle branches, and palm branches, and branches of thick trees, to make booths, as it is written."

However, the term here in Hebrew (עץ שמן) means "oil tree" and it is not clear what kind of tree is meant. Pines are also mentioned in some translations of Isaiah 60:13, such as the King James:

"The glory of Lebanon shall come unto thee, the fir tree, the pine tree, and the box together, to beautify the place of my sanctuary; and I will make the place of my feet glorious."

Again, it is not clear what tree is meant (תדהר in Hebrew), and other translations use "pine" for the word translated as "box" by the King James (תאשור in Hebrew).

Some botanical authorities believe that the Hebrew word "ברוש" (bərōsh), which is used many times in the Bible, designates

, the stone pine. ^[54] The word used in modern Hebrew for pine is "אֹ֖רֶן" (oren), which occurs only in Isaiah 44:14,^[55] but two manuscripts have "ארז" (cedar), a much more common word.^[56]

Chinese culture

The pine is a motif in Chinese art and literature, which sometimes combines painting and poetry in the same work. Some of the main symbolic attributes of pines in Chinese art and literature are longevity and steadfastness: the pine retains its green needles through all the seasons. Sometimes the pine and cypress are paired. At other times the pine, plum, and bamboo are considered as the "

: interestingly enough, one of the main ingredients for Chinese ink has been pine soot.

See also

• El Pino (The Pine Tree)
• Pine barrens
• Pine-cypress forest
• Pine Tree Flag
• Tree of Peace

References

Bibliography

• Farjon A (2005). Pines (2nd ed.). Leiden: E. J. Brill.
  ISBN 90-04-13916-8
  .
• Little Jr EL, Critchfield WB (1969). Subdivisions of the Genus Pinus (Pines). Misc. Publ. 1144 (Superintendent of Documents Number: A 1.38:1144) (Report). US Department of Agriculture.
• Richardson DM, ed. (1998). Ecology and Biogeography of Pinus. Cambridge, England: Cambridge University Press. p. 530.
  ISBN 0-521-55176-5
  .
• Sulavik SB (2007). Adirondack; Of Indians and Mountains, 1535-1838. Fleischmanns, NY: Purple Mountain Press. pp. 244 pages.
  ISBN 978-1-930098-79-4
  .
• Mirov NT (1967). The Genus Pinus. New York, NY: Ronald Press.
• "Classification of pines". The Lovett Pinetum Charitable Foundation. Archived from the original on 2004-06-03. Retrieved 2004-01-17.
• Mirov NT, Stanley RG (1959). "The Pine Tree". Annual Review of Plant Physiology. 10: 223–238.
  doi:10.1146/annurev.pp.10.060159.001255
  .
• Philips R (1979). Trees of North America and Europe. New York, NY: Random House, Inc.
  ISBN 0-394-50259-0
  .
• Earle, Christopher J., ed. (2018). "Pinus". The Gymnosperm Database.

External links

Wikimedia Commons has media related to Pinus.

• 40 Species of Pine Trees You Can Grow by The Spruce
• Jepson eFlora, The Jepson Herbarium, University of California, Berkeley, covers Californian species
• Pinus in Flora of North America
• Pinus in the USDA Plants Database
• Conifer Database