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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 single ...
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, including only woody plants with secondary growth, plants that are ...
or
shrub A shrub (often also called a 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 ...
in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
''Pinus'' () of the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
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, larches, pines and spruces. The family is included in the order Pinales, ...
. ''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 subfamily botanical names with "-oideae", and zoologi ...
Pinoideae. The
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 publishing an ...
created by the
Royal Botanic Gardens, Kew Royal Botanic Gardens, Kew is a non-departmental public body in the United Kingdom sponsored by the Department for Environment, Food and Rural Affairs. An internationally important botanical research and education institution, it employs 1,100 ...
and
Missouri Botanical Garden The Missouri Botanical Garden is a botanical garden located at 4344 Shaw Boulevard in St. Louis, Missouri. It is also known informally as Shaw's Garden for founder and philanthropist Henry Shaw. Its herbarium, with more than 6.6 million spe ...
accepts 187 species names of pines as current, together with more synonyms. The
American Conifer Society The American Conifer Society was founded in 1983 to help educate the public about conifers, which are cone-bearing plants. The Society is governed by a board of directors with representation from each of the Society's four regions. The Society pu ...
(ACS) and the
Royal Horticultural Society The Royal Horticultural Society (RHS), founded in 1804 as the Horticultural Society of London, is the UK's leading gardening charity. The RHS promotes horticulture through its five gardens at Wisley (Surrey), Hyde Hall (Essex), Harlow Carr (Nort ...
accept 121 species. Pines are commonly found 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 as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
. ''Pine'' may also refer to the
lumber Lumber is wood that has been processed into dimensional lumber, including beams and planks or boards, a stage in the process of wood production. Lumber is mainly used for construction framing, as well as finishing (floors, wall panels, wi ...
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
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture, ...
s (or trinomials) recognized by the ACS.


Description

Pine trees are
evergreen In botany, an evergreen is a plant which has foliage that remains green and functional through more than one growing season. This also pertains to plants that retain their foliage only in warm climates, and contrasts with deciduous plants, which ...
, coniferous
resin In polymer chemistry and materials science, resin is a solid or highly viscous substance of plant or synthetic origin that is typically convertible into polymers. Resins are usually mixtures of organic compounds. This article focuses on natu ...
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, including only woody plants with secondary growth, plants that are ...
s (or, rarely,
shrub A shrub (often also called a 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 ...
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 to northeastern Asia and the Japanese isles. It shares the common ...
and
Potosi pinyon ''Pinus culminicola'', commonly known as Potosí pinyon or Potosí Piñón, is a pine in the pinyon pine group, native and endemism, endemic to northeast Mexico. The range is highly localised, confined to a small area of high summits in the north ...
, and the tallest is an tall
ponderosa pine ''Pinus ponderosa'', commonly known as the ponderosa pine, bull pine, blackjack pine, western yellow-pine, or filipinus pine is a very large pine tree species of variable habitat native to mountainous regions of western North America. It is the ...
located in southern
Oregon Oregon () is a U.S. state, state in the Pacific Northwest region of the Western United States. The Columbia River delineates much of Oregon's northern boundary with Washington (state), Washington, while the Snake River delineates much of it ...
's
Rogue River-Siskiyou National Forest A rogue is a person or entity that flouts accepted norms of behavior. Rogue or rogues may also refer to: Companies * Rogue Ales, a microbrewery in Newport, Oregon * Rogue Arts, a film production company * Rogue Entertainment, a software comp ...
. 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 () ( he, מְתוּשֶׁלַח ''Məṯūšélaḥ'', in pausa ''Məṯūšālaḥ'', "His death shall send" or "Man of the javelin" or "Death of Sword"; gr, Μαθουσάλας ''Mathousalas'') was a biblical patriarch and a ...
", 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 and it is now known as "
Prometheus In Greek mythology, Prometheus (; , , possibly meaning "forethought")Smith"Prometheus". is a Titan god of fire. Prometheus is best known for defying the gods by stealing fire from them and giving it to humanity in the form of technology, know ...
" after the Greek immortal. The spiral growth of branches, needles, and
cones A cone is a three-dimensional geometric shape that tapers smoothly from a flat base (frequently, though not necessarily, circular) to a point called the apex or vertex. A cone is formed by a set of line segments, half-lines, or lines conn ...
scales may be arranged in
Fibonacci number In mathematics, the Fibonacci numbers, commonly denoted , form a sequence, the Fibonacci sequence, in which each number is the sum of the two preceding ones. The sequence commonly starts from 0 and 1, although some authors start the sequence from ...
ratios. The new spring shoots are sometimes called "candles"; they are covered in brown or whitish bud scales and point upward at first, then later turn green and spread outward. These "candles" offer
forester A forester is a person who practises forestry, the science, art, and profession of managing forests. Foresters engage in a broad range of activities including ecological restoration and management of protected areas. Foresters manage forests to ...
s a means to evaluate
fertility Fertility is the capability to produce offspring through reproduction following the onset of sexual maturity. The fertility rate is the average number of children born by a female during her lifetime and is quantified demographically. Fertili ...
of the soil and vigour of the trees.


Bark

The
bark Bark may refer to: * Bark (botany), an outer layer of a woody plant such as a tree or stick * Bark (sound), a vocalization of some animals (which is commonly the dog) Places * Bark, Germany * Bark, Warmian-Masurian Voivodeship, Poland Arts, ...
of most pines is thick and scaly, but some species have thin, flaky bark. 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
bud In botany, a bud is an undeveloped or embryonic shoot and normally occurs in the axil of a leaf or at the tip of a stem. Once formed, a bud may remain for some time in a dormant condition, or it may form a shoot immediately. Buds may be spec ...
s at the tip of the year's new
shoot In botany, a plant shoot consists of any plant stem together with its appendages, leaves and lateral buds, flowering stems, and flower buds. The new growth from seed germination that grows upward is a shoot where leaves will develop. In the spri ...
, but others are multinodal, producing two or more whorls of branches per year.


Foliage

Pines have four types of
leaf A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
: * Seed leaves (
cotyledon A cotyledon (; ; ; , gen. (), ) is a significant part of the embryo within the seed of a plant, and is defined as "the embryonic leaf in seed-bearing plants, one or more of which are the first to appear from a germinating seed." The numb ...
s) 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 Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in c ...
) 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 will generate a stem-producing bud, which can then replace the lost growth tip.


Cones

Pines are
monoecious Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system alongside gynomonoecy, andromonoecy and trimonoecy. Monoecy is conne ...
, 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 Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
. The female cones take 1.5–3 years (depending on species) to mature after
pollination Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds ...
, 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
anemophilous Anemophily or wind pollination is a form of pollination whereby pollen is distributed by wind. Almost all gymnosperms are anemophilous, as are many plants in the order Poales, including grasses, sedges, and rushes. Other common anemophilous plan ...
(wind-dispersed), but some are larger and have only a vestigial wing, and are
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
-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 ''Pinus albicaulis'', known by the common names whitebark pine, white bark pine, white pine, pitch pine, scrub pine, and creeping pine, is a conifer tree native to the mountains of the western United States and Canada, specifically subalpine ...
, 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 Serotiny in botany simply means 'following' or 'later'. In the case of serotinous flowers, it means flowers which grow following the growth of leaves, or even more simply, flowering later in the season than is customary with allied species. Havi ...
. The most common form of serotiny is pyriscence, in which a resin binds the cones shut until melted by a forest fire, for example in '' P. rigida''.


Taxonomy

Pines are
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό ...
s. 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 Fascicle or ''fasciculus'' may refer to: Anatomy and histology * Muscle fascicle, a bundle of skeletal muscle fibers * Nerve fascicle, a bundle of axons (nerve fibers) ** Superior longitudinal fasciculus *** Arcuate fasciculus ** Gracile fas ...
. 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. 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, having diverged during the late
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
. 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. 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''). Before the 19th century, pines were often referred to as firs (from
Old Norse Old Norse, Old Nordic, or Old Scandinavian, is a stage of development of North Germanic languages, North Germanic dialects before their final divergence into separate Nordic languages. Old Norse was spoken by inhabitants of Scandinavia and t ...
''fura'', by way of
Middle English Middle English (abbreviated to ME) is a form of the English language that was spoken after the Norman conquest of 1066, until the late 15th century. The English language underwent distinct variations and developments following the Old English p ...
''firre''). In some European languages, Germanic cognates of the Old Norse name are still in use for pines — in
Danish Danish may refer to: * Something of, from, or related to the country of Denmark People * A national or citizen of Denmark, also called a "Dane," see Demographics of Denmark * Culture of Denmark * Danish people or Danes, people with a Danish ance ...
''fyr'', in
Norwegian Norwegian, Norwayan, or Norsk may refer to: *Something of, from, or related to Norway, a country in northwestern Europe *Norwegians, both a nation and an ethnic group native to Norway *Demographics of Norway *The Norwegian language, including the ...
''fura/fure/furu'',
Swedish Swedish or ' may refer to: Anything from or related to Sweden, a country in Northern Europe. Or, specifically: * Swedish language, a North Germanic language spoken primarily in Sweden and Finland ** Swedish alphabet, the official alphabet used by ...
''fura/furu'',
Dutch Dutch commonly refers to: * Something of, from, or related to the Netherlands * Dutch people () * Dutch language () Dutch may also refer to: Places * Dutch, West Virginia, a community in the United States * Pennsylvania Dutch Country People E ...
''vuren'', and
German German(s) may refer to: * Germany (of or related to) **Germania (historical use) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizens of Germany, see also German nationality law **Ger ...
''Föhre'' — but in modern English, ''fir'' is now restricted to fir (''Abies'') and
Douglas-fir The Douglas fir (''Pseudotsuga menziesii'') is an evergreen conifer species in the pine family, Pinaceae. It is native to western North America and is also known as Douglas-fir, Douglas spruce, Oregon pine, and Columbian pine. There are three va ...
(''Pseudotsuga'').


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, larches, pines and spruces. The family is included in the order Pinales, ...
, the pine family, which first appeared in the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
period. Based on recent
Transcriptome The transcriptome is the set of all RNA transcripts, including coding and non-coding, in an individual or a population of cells. The term can also sometimes be used to refer to all RNAs, or just mRNA, depending on the particular experiment. The t ...
analysis, ''Pinus'' is most closely related to the genus ''
Cathaya ''Cathaya'' is a genus in the pine family, Pinaceae, with one known living species, ''Cathaya argyrophylla''. ''Cathaya'' is a member of the subfamily Laricoideae, most closely related to ''Pseudotsuga'' and ''Larix''. A second species, ''C. nan ...
'', which in turn is closely related to
spruce A spruce is a tree of the genus ''Picea'' (), a genus of about 35 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal (taiga) regions of the Earth. ''Picea'' is the sole genus in the subfami ...
s. These genera, with
fir Firs (''Abies'') are a genus of 48–56 species of evergreen coniferous trees in the family (biology), family Pinaceae. They are found on mountains throughout much of North America, North and Central America, Europe, Asia, and North Africa. The ...
s and
larch Larches are deciduous conifers in the genus ''Larix'', of the family Pinaceae (subfamily Laricoideae). Growing from tall, they are native to much of the cooler temperate northern hemisphere, on lowlands in the north and high on mountains furt ...
es, form the pinoid
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
of the Pinaceae. Pines first appeared during the Early Cretaceous, with the oldest verified fossil of the genus is ''
Pinus yorkshirensis ''Pinus yorkshirensis'' is an extinct species of pine tree. The fossil pine cone came from Hauterivian and Barremian-aged sedimentary rocks located in the Speeton Clay in Yorkshire (hence the species epithet). Discovery and naming The type spe ...
'' from the
Hauterivian The Hauterivian is, in the geologic timescale, an age in the Early Cretaceous Epoch or a stage in the Lower Cretaceous Series. It spans the time between 132.9 ± 2 Ma and 129.4 ± 1.5 Ma (million years ago). The Hauterivian is preceded by the Va ...
-
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma (million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is preceded ...
boundary (131–129 million years ago) from the
Speeton Clay The Speeton Clay Formation (SpC)Speeton Clay Formation
- hybridization Hybridization (or hybridisation) may refer to: *Hybridization (biology), the process of combining different varieties of organisms to create a hybrid *Orbital hybridization, in chemistry, the mixing of atomic orbitals into new hybrid orbitals *Nu ...
. Pines are prone to inter-specific breeding. Wind pollination, long life spans, overlapping generations, large population size, and weak
reproductive isolation The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, behaviors and physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensure that any offspring ...
make breeding across species more likely. As the pines have diversified, gene transfer between different species has created a complex history of genetic relatedness. The following cladogram shows the phylogenetic relationships between the pine species as described in 2021. , label2=subsection ''Pinaster'', 2= , label2=subgenus ''Strobus'', 2= , label2=subsection ''Gerardiana'', 2= , label2=section ''Parrya'', 2=} , 2=


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 as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
, and to a few parts from the tropics to
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
regions in the Southern Hemisphere. Most regions of the Northern Hemisphere host some
native species In biogeography, a native species is indigenous to a given region or ecosystem if its presence in that region is the result of only local natural evolution (though often popularised as "with no human intervention") during history. The term is equ ...
of pines. One species (
Sumatran pine ''Pinus merkusii'', the Merkus pine or Sumatran pine, is a pine native to the Malesia region of southeast Asia, and the only one that occurs naturally south of the equator. Description ''Pinus merkusii'' is a medium-sized to large tree, reachin ...
) 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 favorable soils and at least some water. Various species have been introduced to temperate and
subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Geographical z ...
regions of both hemispheres, where they are grown as
timber Lumber is wood that has been processed into dimensional lumber, including beams and planks or boards, a stage in the process of wood production. Lumber is mainly used for construction framing, as well as finishing (floors, wall panels, wi ...
or cultivated as ornamental plants in parks and gardens. A number of such introduced species have become naturalized, and some species are considered invasive in some areas and threaten native ecosystems.


Ecology

Pines grow well in acid soils, some also on
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an adje ...
soils; most require good soil drainage, preferring sandy soils, but a few (e.g.
lodgepole pine ''Pinus contorta'', with the common names lodgepole pine and shore pine, and also known as twisted pine, and contorta pine, is a common tree in western North America. It is common near the ocean shore and in dry montane forests to the subalpine, ...
) can tolerate poorly drained wet soils. A few are able to sprout after forest fires (e.g.
Canary Island pine ''Pinus canariensis'', the Canary Island pine, is a species of gymnosperm in the conifer family Pinaceae. It is a large, evergreen tree, native and endemic to the outer Canary Islands of the Atlantic Ocean. Description ''Pinus canariensis'' is a ...
). Some species of pines (e.g.
bishop pine ''Pinus muricata'', the bishop pine, is a pine with a very restricted range: mostly in California, including several offshore Channel Islands, and a few locations in Baja California, Mexico. It is always on or near the coast. In San Luis Obispo ...
) 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,
mountain pine ''Pinus mugo'', known as bog pine, creeping pine, dwarf mountain pine, mugo pine, mountain pine, scrub mountain pine, or Swiss mountain pine, is a species of conifer, native to high elevation habitats from southwestern to Central Europe and Sou ...
, whitebark pine, and the
bristlecone pine The term bristlecone pine covers three species of pine tree (family Pinaceae, genus ''Pinus A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. T ...
s). The pinyon pines and a number of others, notably
Turkish pine ''Pinus brutia'', commonly known as the Turkish pine, is a species of pine native to the eastern Mediterranean region. The bulk of its range is in Turkey. Turkish pine is also known by several other common names: Calabrian pine (from a naturali ...
and
gray pine ''Pinus sabiniana'' (sometimes spelled ''P. sabineana''), with vernacular names including towani pine, foothill pine, gray pine, bull pine, and digger pine, is a pine endemic to California in the United States. Some sources discourage using the n ...
, are particularly well adapted to growth in hot, dry
semidesert A semi-arid climate, semi-desert climate, or steppe climate is a dry climate sub-type. It is located on regions that receive precipitation below potential evapotranspiration, but not as low as a desert climate. There are different kinds of semi-ar ...
climates. Pine
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
may play an important role in the functioning of
detrital Detritus (; adj. ''detrital'' ) is particles of rock derived from pre-existing rock through weathering and erosion.Essentials of Geology, 3rd Ed, Stephen Marshak, p G-7 A fragment of detritus is called a clast.Essentials of Geology, 3rd Ed, Stephen ...
food web A food web is the natural interconnection of food chains and a graphical representation of what-eats-what in an ecological community. Another name for food web is consumer-resource system. Ecologists can broadly lump all life forms into one ...
s. 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 Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
(
butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...
and
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
) species. Several species of pine are attacked by
nematodes The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a broa ...
, causing pine
wilt disease A wilt disease is any number of diseases that affect the vascular system of plants. Attacks by fungi, bacteria, and nematodes can cause rapid killing of plants, large tree branches or even entire trees. Wilt diseases in woody plants tend to fal ...
, which can kill some quickly. Some of these Lepidoptera species, many of them moths, specialize 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 nut Pine nuts, also called piñón (), pinoli (), pignoli or chilgoza (), are the edible seeds of pines (family Pinaceae, genus ''Pinus''). According to the Food and Agriculture Organization, only 29 species provide edible nuts, while 20 are trade ...
s. The seeds are commonly eaten by birds, such as grouse, crossbills, jays, nuthatches, siskins, and woodpeckers, and by
squirrel Squirrels are members of the family Sciuridae, a family that includes small or medium-size rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrels. Squ ...
s. Some birds, notably the
spotted nutcracker The spotted nutcracker, Eurasian nutcracker, or simply nutcracker (''Nucifraga caryocatactes'') is a passerine bird slightly larger than the Eurasian jay. It has a much larger bill and a slimmer looking head without any crest. The feathering over ...
,
Clark's nutcracker Clark's nutcracker (''Nucifraga columbiana''), sometimes referred to as Clark's crow or woodpecker crow, is a passerine bird in the family Corvidae, native to the mountains of western North America. The nutcracker is an omnivore, but subsists mai ...
, and
pinyon jay The pinyon jay (''Gymnorhinus cyanocephalus'') is a species of jay, and is the only member of the genus ''Gymnorhinus''. Native to Western North America, the species ranges from central Oregon to northern Baja California, and eastward as far as ...
, are of importance in distributing pine seeds to new areas. Pine needles are sometimes eaten by the
Symphyta Sawflies are the insects of the suborder Symphyta within the order Hymenoptera, alongside ants, bees, and wasps. The common name comes from the saw-like appearance of the ovipositor, which the females use to cut into the plants where they lay ...
n species
pine sawfly The Diprionidae are a small family of conifer Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division c ...
, and
goat The goat or domestic goat (''Capra hircus'') is a domesticated species of goat-antelope typically kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the a ...
s.


Uses


Lumber and construction

Pines are among the most commercially important tree species valued for their timber and
wood pulp Pulp is a lignocellulosic fibrous material prepared by chemically or mechanically separating cellulose fibers from wood, fiber crops, waste paper, or rags. Mixed with water and other chemical or plant-based additives, pulp is the major raw mate ...
throughout the world. In temperate and tropical regions, they are fast-growing
softwood file:Pinus sylvestris wood ray section 1 beentree.jpg, Scots Pine, a typical and well-known softwood Softwood is wood from gymnosperm trees such as conifers. The term is opposed to hardwood, which is the wood from angiosperm trees. The main diff ...
s that grow in relatively dense stands, their acidic decaying needles inhibiting the sprouting of competing hardwoods. Commercial pines are grown in
plantation A plantation is an agricultural estate, generally centered on a plantation house, meant for farming that specializes in cash crops, usually mainly planted with a single crop, with perhaps ancillary areas for vegetables for eating and so on. The ...
s 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 Turpentine (which is also called spirit of turpentine, oil of turpentine, terebenthene, terebinthine and (colloquially) turps) is a fluid obtained by the distillation of resin harvested from living trees, mainly pines. Mainly used as a special ...
. 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 Drywall (also called plasterboard, dry lining, wallboard, sheet rock, gypsum board, buster board, custard board, and gypsum panel) is a panel made of calcium sulfate dihydrate (gypsum), with or without additives, typically extruded between thick ...
framing, for example). For outside use, pine needs to be treated with copper azole,
chromated copper arsenate Chromated copper arsenate (CCA) is a wood preservative containing compounds of chromium, copper, and arsenic, in various proportions. It is used to impregnate timber and other wood products, especially those intended for outdoor use, in order to pr ...
or other suitable chemical preservative.


Ornamental uses

Many pine species make attractive ornamental plantings for
park A park is an area of natural, semi-natural or planted space set aside for human enjoyment and recreation or for the protection of wildlife or natural habitats. Urban parks are urban green space, green spaces set aside for recreation inside t ...
s and larger
garden A garden is a planned space, usually outdoors, set aside for the cultivation, display, and enjoyment of plants and other forms of nature. The single feature identifying even the wildest wild garden is ''control''. The garden can incorporate both ...
s with a variety of dwarf
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture, ...
s being suitable for smaller spaces. Pines are also commercially grown and harvested for
Christmas tree A Christmas tree is a decorated tree, usually an evergreen conifer, such as a spruce, pine or fir, or an artificial tree of similar appearance, associated with the celebration of Christmas. The custom was further developed in early modern ...
s. Pine cones, the largest and most durable of all conifer cones, are craft favorites. 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
U.S. Civil War The American Civil War (April 12, 1861 – May 26, 1865; also known by other names) was a civil war in the United States. It was fought between the Union ("the North") and the Confederacy ("the South"), the latter formed by states th ...
, the needles of the
longleaf pine The longleaf pine (''Pinus palustris'') is a pine species native to the Southeastern United States, found along the coastal plain from East Texas to southern Virginia, extending into northern and central Florida. In this area it is also known as ...
"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
biodegradable Biodegradation is the breakdown of organic matter by microorganisms, such as bacteria and fungi. It is generally assumed to be a natural process, which differentiates it from composting. Composting is a human-driven process in which biodegradati ...
products including paper, furniture, textiles and dye.


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. 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 essential ingredient of ''
pesto Pesto () is a sauce that traditionally consists of crushed garlic, European pine nuts, coarse salt, basil leaves, and hard cheese such as Parmigiano-Reggiano (also known as Parmesan cheese) or Pecorino Sardo (cheese made from sheep's milk), al ...
alla genovese''. The soft, moist, white inner bark (
cambium A cambium (plural cambia or cambiums), in plants, is a tissue layer that provides partially undifferentiated cells for plant growth. It is found in the area between xylem and phloem. A cambium can also be defined as a cellular plant tissue from w ...
) beneath the woody outer bark is edible and very high in vitamins A and C. It can be eaten raw in slices as a snack or dried and ground up into a powder for use as an
ersatz An ersatz good () is a substitute good, especially one that is considered inferior to the good it replaces. It has particular connotations of wartime usage. Etymology ''Ersatz'' is a German word literally meaning ''substitute'' or ''replacement ...
flour or thickener in stews, soups, and other foods, such as
bark bread Bark bread is a traditional food made with cambium (phloem) flour. It has a history of use as famine food. History Bark bread seems to be a primarily Scandinavian tradition. Mention of it is found in medieval literature, and it may have an even ...
. Adirondack Indians got their name from the Mohawk Indian word ''atirú:taks'', meaning "tree eaters". A
tea Tea is an aromatic beverage prepared by pouring hot or boiling water over cured or fresh leaves of '' Camellia sinensis'', an evergreen shrub native to East Asia which probably originated in the borderlands of southwestern China and northe ...
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 Retsina ( el, Ρετσίνα) is a Greek white (or rosé) resinated wine, which has been made for at least 2,000 years. Its unique flavor is said to have originated from the practice of sealing wine vessels, particularly amphorae, with Alepp ...
is flavoured with Aleppo pine resin. Pine needles from ''
Pinus densiflora ''Pinus densiflora'', also called the Japanese red pine, the Japanese pine, or Korean red pine, is a species of pine tree native to East Asia. Distribution and habitat ''P. densiflora'' has a home range that includes Japan, the Korean Peninsula, ...
'' were found to contain 30.54 milligram/gram of
proanthocyanidin Proanthocyanidins are a class of polyphenols found in many plants, such as cranberry, blueberry, and grape seeds. Chemically, they are oligomeric flavonoids. Many are oligomers of catechin and epicatechin and their gallic acid esters. More complex ...
s when extracted with hot water. Comparative to ethanol extraction resulting in 30.11 mg/g, simply extracting in hot water is preferable. In
traditional Chinese medicine Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of action ...
,
pine resin In polymer chemistry and materials science, resin is a solid or highly viscous substance of plant or synthetic origin that is typically convertible into polymers. Resins are usually mixtures of organic compounds. This article focuses on natu ...
is used for burns, wounds and dermal complaints.


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 John Muir ( ; April 21, 1838December 24, 1914), also known as "John of the Mountains" and "Father of the National Parks", was an influential Scottish-American naturalist, author, environmental philosopher, botanist, zoologist, glaciologist, a ...
,
Dora Sigerson Shorter Dora Maria Sigerson Shorter (16 August 1866 – 6 January 1918) was an Irish poet and sculptor, who after her marriage in 1895 wrote under the name Dora Sigerson Shorter. Life She was born in Dublin, Ireland, the daughter of George Sigerson, a ...
,
Eugene Field Eugene Field Sr. (September 2, 1850 – November 4, 1895) was an American writer, best known for his children's poetry and humorous essays. He was known as the "poet of childhood". Early life and education Field was born in St. Louis, Missour ...
,
Bai Juyi Bai Juyi (also Bo Juyi or Po Chü-i; ; 772–846), courtesy name Letian (樂天), was a renowned Chinese poet and Tang dynasty government official. Many of his poems concern his career or observations made about everyday life, including as g ...
,
Theodore Winthrop Major Theodore Woolsey Winthrop (September 22, 1828 – June 10, 1861) was a writer, lawyer, and world traveller. He was one of the first Union officers killed in the American Civil War. Biography Winthrop was born in New Haven, Connecticut, ...
, and Rev. George Allan D.D.


Art

Pines are often featured in art, whether painting and
fine art In European academic traditions, fine art is developed primarily for aesthetics or creative expression, distinguishing it from decorative art or applied art, which also has to serve some practical function, such as pottery or most metalwork ...
, drawing, photography, or
folk art Folk art covers all forms of visual art made in the context of folk culture. Definitions vary, but generally the objects have practical utility of some kind, rather than being exclusively decorative art, decorative. The makers of folk art a ...
.


Religious texts

Pine trees, as well as other conifers, are mentioned in some verses of the
Bible The Bible (from Koine Greek , , 'the books') is a collection of religious texts or scriptures that are held to be sacred in Christianity, Judaism, Samaritanism, and many other religions. The Bible is an anthologya compilation of texts of a ...
, depending on the translation. In the
Book of Nehemiah The Book of Nehemiah in the Hebrew Bible, largely takes the form of a first-person memoir concerning the rebuilding of the walls of Jerusalem after the Babylonian exile by Nehemiah, a Jew who is a high official at the Persian court, and the dedic ...
8:15, the
King James Version The King James Version (KJV), also the King James Bible (KJB) and the Authorized Version, is an Bible translations into English, English translation of the Christian Bible for the Church of England, which was commissioned in 1604 and publis ...
gives the following translation:
"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'' mphasis added and myrtle branches, and palm branches, and branches of thick trees, to mak
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 '' P. halepensis'', or in
Hosea In the Hebrew Bible, Hosea ( or ; he, הוֹשֵׁעַ – ''Hōšēaʿ'', 'Salvation'; gr, Ὡσηέ – ''Hōsēé''), son of Beeri, was an 8th-century BCE prophet in Israel and the nominal primary author of the Book of Hosea. He is the ...
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" (
cedar Cedar may refer to: Trees and plants *''Cedrus'', common English name cedar, an Old-World genus of coniferous trees in the plant family Pinaceae *Cedar (plant), a list of trees and plants known as cedar Places United States * Cedar, Arizona * ...
), a much more common word.


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 "
Three Friends of Winter The Three Friends of Winter is an art motif that comprises the pine, bamboo, and plum. . The Chinese celebrated the pine, bamboo and plum together, as they observed that these plants do not wither as the cold days deepen into the winter season ...
". Eberhard, Wolfram (2003
986 (German version 1983) Year 986 ( CMLXXXVI) was a common year starting on Friday (link will display the full calendar) of the Julian calendar. Events By place Byzantine Empire * August 17 – Battle of the Gates of Trajan: Emperor Basil II leads a Byz ...
, ''A Dictionary of Chinese Symbols: Hidden Symbols in Chinese Life and Thought''. London, New York: Routledge. , ''sub'' "Pine".
Many Chinese art works and/or literature (some involving pines) have been done using paper, brush, and Chinese ink: interestingly enough, one of the main ingredients for Chinese ink has been pine soot.


See also

*
El Pino (The Pine Tree) El Pino ( en, The Pine Tree) is a large bunya pine located on the southeastern corner of Folsom Street and N. Indiana Street in East Los Angeles, California. The tree overlooks the Wellington Heights neighborhood of East Los Angeles and the Boy ...
*
Pine barrens Pine barrens, pine plains, sand plains, or pineland areas occur throughout the U.S. from Florida to Maine (see Atlantic coastal pine barrens) as well as the Midwest, West, and Canada and parts of Eurasia. Perhaps the most well known pine-barre ...
* Pine-cypress forest *
Pine Tree Flag The Tree Flag (or the Appeal to Heaven Flag) was one of the flags used during the American Revolution. The flag, which featured a pine tree with the motto "An Appeal to Heaven," or less frequently "An Appeal to God", was originally used by a squ ...
*
Tree of Peace The Iroquois Tree of Peace finds its roots in a man named Dekanawida, the peace-giver. The legends surrounding his place amongst the Iroquois (the Haudenosaunee) is based in his role in creating the Five Nations Confederacy, which consisted of ...


References


Bibliography

* * * * * * * * *


External links


40 Species of Pine Trees You Can Grow
by The Spruce *, covers Californian species
Pinus in Flora of North AmericaPinus in the USDA Plants Database
{{Authority control Pinaceae