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A pine is any
conifer Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...
tree or
shrub A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...
in the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
''Pinus'' () of the family
Pinaceae The Pinaceae (), or pine family, are conifer trees or shrubs, including many of the well-known conifers of commercial importance such as Cedrus, cedars, firs, Tsuga, hemlocks, Pinyon_pine, piñons, larches, pines and spruces. The family is incl ...
. ''Pinus'' is the sole genus in the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...
Pinoideae. ''
World Flora Online World Flora Online is an Internet-based compendium of the world's plant species. Description The World Flora Online (WFO) is an open-access database, launched in October 2012 as a follow-up project to The Plant List, with the aim of publishi ...
'' accepts 134 species-rank taxa (119 species and 15
nothospecies #REDIRECT Hybrid name In botanical nomenclature, a hybrid may be given a hybrid name, which is a special kind of botanical name, but there is no requirement that a hybrid name should be created for plants that are believed to be of hybrid origin. ...
) of pines as current, with additional synonyms, and ''
Plants of the World Online Plants of the World Online (POWO) is an online taxonomic database published by the Royal Botanic Gardens, Kew. History Following the Convention on Biological Diversity, the Royal Botanic Gardens in Kew launched Plants of the World Online i ...
'' 126 species-rank taxa (113 species and 13 nothospecies), making it the largest genus among the conifers. The highest species diversity of pines is found in Mexico. Pines are widely
distributed Distribution may refer to: Mathematics *Distribution (mathematics), generalized functions used to formulate solutions of partial differential equations *Probability distribution, the probability of a particular value or value range of a varia ...
in the
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the equator. For other planets in the Solar System, north is defined by humans as being in the same celestial sphere, celestial hemisphere relative to the invariable plane of the Solar ...
; they occupy large areas of
boreal forest Taiga or tayga ( ; , ), also known as boreal forest or snow forest, is a biome characterized by pinophyta, coniferous forests consisting mostly of pines, spruces, and larches. The taiga, or boreal forest, is the world's largest land biome. I ...
, but are found in many habitats, including the Mediterranean Basin, and dry tropical forests in southeast Asia and Central America. Wood from pine trees is one of the most extensively used types of
timber Lumber is wood that has been processed into uniform and useful sizes (dimensional lumber), including beams and planks or boards. Lumber is mainly used for construction framing, as well as finishing (floors, wall panels, window frames). ...
, and some pines are widely used as
Christmas tree A Christmas tree is a decorated tree, usually an evergreen pinophyta, conifer, such as a spruce, pine or fir, associated with the celebration of Christmas. It may also consist of an artificial tree of similar appearance. The custom was deve ...
s.


Description

Pine trees are
evergreen In botany, an evergreen is a plant which has Leaf, foliage that remains green and functional throughout the year. This contrasts with deciduous plants, which lose their foliage completely during the winter or dry season. Consisting of many diffe ...
, coniferous
resin A resin is a solid or highly viscous liquid that can be converted into a polymer. Resins may be biological or synthetic in origin, but are typically harvested from plants. Resins are mixtures of organic compounds, predominantly terpenes. Commo ...
ous
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, e.g., including only woody plants with secondary growth, only ...
s (or, rarely,
shrub A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...
s) growing tall, with the majority of species reaching tall. The smallest are
Siberian dwarf pine ''Pinus pumila'', commonly known as the Siberian dwarf pine, dwarf Siberian pine, dwarf stone pine, Japanese stone pine, or creeping pine, is a tree in the family Pinaceae native plant, native to northeastern Asia and the Japan, Japanese isles. ...
and
Potosi pinyon ''Pinus culminicola'', commonly known as Potosí pinyon or Potosí Piñón, is a pine in the pinyon pine group, native and endemic to northeast Mexico. The range is highly localised, confined to a small area of high summits in the northern Sier ...
, and the tallest is an tall
sugar pine ''Pinus lambertiana'' (commonly known as the sugar pine or sugar cone pine) is the tallest and most massive pine tree and has the longest Conifer cone, cones of any conifer. It is native to coastal and inland mountain areas along the Pacific coa ...
located in
Yosemite National Park Yosemite National Park ( ) is a List of national parks of the United States, national park of the United States in California. It is bordered on the southeast by Sierra National Forest and on the northwest by Stanislaus National Forest. The p ...
. 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'' (commonly referred to as the Great Basin bristlecone pine, intermountain bristlecone pine, or western bristlecone pine) is a long-living species of bristlecone pine tree found in the higher mountains of California, Nevada, and ...
(''P. longaeva''). One individual of this species, dubbed "
Methuselah Methuselah (; ''Məṯūšélaḥ'', in pausa ''Məṯūšālaḥ'', "His death shall send" or "Man of the javelin" or "Death of sword"; ''Mathousalas'') was a biblical patriarch and a figure in Judaism, Christianity, and Islam. He is clai ...
", is one of the world's oldest living organisms at around 4,800-years old. This tree can be found in the White Mountains of California. An older tree, now cut down, was dated at 4,900-years old. It was discovered in a grove beneath Wheeler Peak (Nevada), Wheeler Peak and it is now known as "Prometheus (tree), Prometheus" after the Prometheus, Greek immortal. The spiral growth of branches, needles, and Conifer cone, cones scales Phyllotaxis, are arranged in Fibonacci number ratios. File:Big bristlecone pine Pinus longaeva.jpg, Ancient ''Pinus longaeva'', California File:Tree Types and Barks 004.jpg, ''Pinus taeda'' bark File:Pine needles.png, Pine needles File:Pinus radiata cone.jpg, ''Pinus radiata'' female cone


Bark

The Bark (botany), 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 plant stem, 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 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 Fascicle (botany), 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 apical meristem, 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 monoecious, having the male and female cones on the same tree. 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. The seeds are mostly small and winged, and are Anemochory, anemochorous (wind-dispersed), but some are larger and 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, 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 melted by a forest fire, for example in ''Pinus radiata, P. radiata'' and ''Pinus muricata, P. muricata''; the seeds are then released after the fire to colonise the burnt ground with minimal competition from other plants.


Naming


Etymology

The modern English name "pine" derives from Latin ''pinus'', traced to the Indo-European base ''*pīt-'' 'resin'. Before the 19th century, pines were often called firs, a name now applied to another genus, ''Fir, Abies''. In some European languages, Germanic cognates of the Old Norse name are still in use for pines, as in Danish language, Danish ''fyr'', Swedish language, Swedish ''fura/furu'', and German language, German ''Föhre''.


Taxonomic history

The genus ''Pinus'' was named by Carl Linnaeus in 1753. ''Pinus sylvestris'', the Scots pine, was later chosen as the type species.


Evolution


Taxonomy

Pines are gymnosperms. The genus is divided into two subgenera based on the number of Vascular bundle, 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: *Pinus subg. Pinus, ''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). The subgenus has also been called ''diploxylon'', on account of its two fibrovascular bundles. *Strobus, ''Pinus'' subg. ''Strobus'', synonym (biology), 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). The subgenus has also been called ''haploxylon'', on account of its single fibrovascular bundle. Phylogenetic evidence indicates that both subgenera have a very ancient divergence from one another. 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 hybridisation. This results in low morphological and genetic differences. This, coupled with low sampling and underdeveloped genetic techniques, has made taxonomy difficult to determine. Recent research using large genetic datasets has clarified these relationships into the groupings often accepted today.


Phylogeny

''Pinus'' is the largest genus of the
Pinaceae The Pinaceae (), or pine family, are conifer trees or shrubs, including many of the well-known conifers of commercial importance such as Cedrus, cedars, firs, Tsuga, hemlocks, Pinyon_pine, piñons, larches, pines and spruces. The family is incl ...
, the pine family, which first appeared in the Jurassic period. Based on recent 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. 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 Formation, Speeton Clay, England. However, there are possible records from the Jurassic. The evolutionary history of the genus ''Pinus'' has been complicated by Hybrid (biology), 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. Two recent phylogenies are given below; the differences between them, and other published phylogenies, demonstrate these complications:


Distribution and habitat

Pines are native to the
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the equator. For other planets in the Solar System, north is defined by humans as being in the same celestial sphere, celestial hemisphere relative to the invariable plane of the Solar ...
, and to a few parts from the tropics to temperate climate, temperate regions in the Southern Hemisphere. Most regions of the Northern Hemisphere host some native species of pines; they occupy large areas of Boreal forest, and are found all around the Mediterranean Basin. The northernmost is Pinus sylvestris, Scots pine, reaching just north of 70° N in Stabbursdalen National Park in Norway; Google Maps shows geolocated images with pines at 70° 09' N. One species (Pinus merkusii, Sumatran pine) crosses the equator in Sumatra to 2°S. In North America, various species occur in regions at latitudes from as far north as 66° N to as far south as 12°N. Pines may be found in a very large variety of environments, ranging from semi-arid desert to rainforests, from sea level up to , from the coldest to the hottest environments on Earth. They often occur in mountainous areas with favourable soils and at least some water. Various species have been introduced to temperate and subtropics, subtropical regions of both hemispheres, where they are grown as
timber Lumber is wood that has been processed into uniform and useful sizes (dimensional lumber), including beams and planks or boards. Lumber is mainly used for construction framing, as well as finishing (floors, wall panels, window frames). ...
or cultivated as ornamental plants in parks and gardens. A number of such introduced species have become naturalisation (biology), naturalised, and some species are considered invasive species, invasive in some areas and threaten native ecosystems.


Ecology

Pines grow well in acid soils, some also on calcareous soils; most require good soil drainage, preferring sandy soils, but a few (e.g. Pinus contorta, lodgepole pine) can tolerate poorly drained wet soils. A few are able to sprout after forest fires (e.g. Pinus canariensis, Canary Island pine). Some species of pines (e.g. Pinus muricata, Bishop pine) 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. Several species are adapted to extreme conditions imposed by elevation and latitude (e.g. Siberian dwarf pine, Pinus mugo, mountain pine, whitebark pine, and the bristlecone pines). The pinyon pines and a number of others, notably Pinus brutia, Turkish pine and Pinus sabiniana, gray pine, are particularly well adapted to growth in hot, dry desert, semidesert climates. Pine pollen may play an important role in the functioning of Detritivore, detrital food webs. Nutrients from pollen aid detritivores in development, growth, and maturation, and may enable fungi to decompose nutritionally scarce litter. Pine pollen is also involved in moving plant matter between terrestrial and aquatic ecosystems.


Wildlife

Pine needles serve as food for various Lepidoptera (butterfly and moth) species. Several species of pine are attacked by nematodes, causing Bursaphelenchus xylophilus, pine wilt disease, which can kill some quickly. Some of these Lepidoptera species, many of them moths, specialise in feeding on only one or sometimes several species of pine. Beside that many species of birds and mammals shelter in pine habitat or feed on pine nuts. The seeds are commonly eaten by birds, such as grouse, crossbills, jays, nuthatches, siskins, and woodpeckers, and by squirrels. Some birds, notably Nutcracker (bird), nutcrackers and pinyon jays, are of major importance in distributing pine seeds to new areas. Pine needles are sometimes eaten by the Symphytan species pine sawfly, and goats.


Uses


Timber and construction

Pines are among the most commercially important tree species, valued for their timber and Pulp (paper), wood pulp throughout the world. 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.Turpentine is extracted from the wood of some species of pine. As 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). It is commonly used in Canadian Lumber Standard graded wood. For outside use, pine needs to be treated with copper azole, chromated copper arsenate or other suitable Wood preservation#Chemical preservatives, chemical preservative.


Ornamental uses

Many pine species make attractive ornamental plantings for parks and larger gardens with a variety of dwarf cultivars being suitable for smaller spaces. There are currently 818 named cultivars (or trinomial nomenclature, trinomials) recognised by the American Conifer Society ACS. Pines are also commercially grown and harvested for
Christmas tree A Christmas tree is a decorated tree, usually an evergreen pinophyta, conifer, such as a spruce, pine or fir, associated with the celebration of Christmas. It may also consist of an artificial tree of similar appearance. The custom was deve ...
s. Pine cones, among the largest and most durable of all conifer cones, are craft favourites. Pine boughs, appreciated especially in wintertime for their pleasant smell and greenery, are popularly cut for decorations. Pine needles are also used for making decorative articles such as baskets, trays, pots, etc., and during the American Civil War, U.S. Civil War, the needles of the Pinus palustris, longleaf pine "Georgia pine" were widely employed in this. This originally Native American skill is now being replicated across the world. Pine needle handicrafts are made in the US, Canada, Mexico, Nicaragua, and India. Pine needles are also versatile and have been used by Latvian designer Tamara Orjola to create different Biodegradation, biodegradable products including paper, furniture, textiles and dye.


Forestry

When grown for sawlogs, pine plantations can be harvested after 25 years, with some stands being allowed to grow up to 50 or more years (the wood value increases more quickly as the trees age). In colder and drier climates, growth is slower, and harvesting can be at much older ages. 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 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. A 30-year-old commercial pine tree grown in good conditions in Arkansas will be about in diameter and about high. After 50 years, the same tree will be about in diameter and 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 and diameter , 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).


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. Some species have large pine nuts, which are harvested and sold for cooking and baking. They are an ingredient of ''pesto alla genovese''. The soft, moist, white inner bark (vascular cambium, cambium) beneath the woody outer bark is edible and very high in vitamins vitamin A, A and vitamin C, C. 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. Adirondack Indians got their name from the Mohawk people, Mohawk Indian word ''atirú:taks'', meaning "tree eaters". A tea is made by steeping young, green pine needles in boiling water (known as ''tallstrunt'' in Sweden). In eastern Asia, pine and other conifers are accepted among consumers as a beverage product, and used in teas, as well as wine. 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. In traditional Chinese medicine, pine resin is used for burns, wounds and skin complaints.


Culture

Pines have been a frequently mentioned tree throughout history, including in literature, art, and in religious texts. The pine is a particular 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 "Three Friends of Winter".


Literature

Writers of various nationalities and ethnicities have written of pines. Among them, John Muir, Dora Sigerson Shorter, Eugene Field, Bai Juyi, Theodore Winthrop, and Rev. George Allan D.D.


Art

Pines are often featured in art, whether painting and fine art, drawing, photography, or folk art.


Religious texts

Trees which may be pines or other conifers are mentioned in some verses of the Bible. In the Book of Nehemiah 8:15, the King James Version translates Hebrew "עץ שמן" (etz shman), 'oil tree', as pine, and the unknown type of tree of Hebrew "תדהר" in Isaiah 60:13 similarly. Some botanical authorities believe that the Hebrew word "ברוש" (bərōsh), "cypress", which is used many times in the Bible, properly designates ''Pinus halepensis'', the Aleppo or Jerusalem pine, or in Book of Hosea, Hosea 14:8 which refers to fruit, ''Pinus pinea'', the stone pine. The word used in modern Hebrew for pine is "אֹ֖רֶן" (oren), which occurs only in Isaiah 44:14, but two manuscripts have "ארז" (cedrus, cedar), a much more common word.Biblia Hebraica Stuttgartensia ad loc.


See also

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


References


Bibliography

* * * * * * * * *


External links


40 Species of Pine Trees You Can Grow
by The Spruce * , covers Californian species
Pinus in Flora of North America

Pinus in the USDA Plants Database

Conifer Database
{{Authority control Pinaceae Pinus, Pinus