Apine is anyconifer in thegenusPinus (/ˈpaɪ.nəs/[2]) of the familyPinaceae.Pinus is the sole genus in thesubfamilyPinoideae. The species are evergreentrees or shrubs with their leaves inbunches, usually of 2 to 5 needles. The seeds are carried on woody cones, with two seeds to each cone scale.
Pine trees provide one of the most extensively used types oftimber. The seeds are used to make dishes such aspesto, whileretsina wine is flavoured with pine resin.
Thebark 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, frombuds at the tip of the year's newshoot, but others are multinodal, producing two or more whorls of branches per year.[10]
Many pines, likePinus taeda, have thick bark that flakes into scales.
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 (photosynthetic) and bundled in clusters calledfascicles. 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'sgrowing tip is damaged (e.g. eaten by an animal), the needle fascicles just below the damage generate a stem-producing bud, which can then replace the lost growth tip.
Pines aremonoecious, having the male and female cones on the same tree.[11]: 205 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 theirpollen. The female cones take 1.5–3 years (depending on species) to mature afterpollination, 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.[10]
The seeds (pine nuts) are mostly small and winged, and areanemochorous (wind-dispersed). Some are larger, have only a vestigial wing, and arebird-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 examplewhitebark pine,[12] 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 calledserotiny. The most common form of serotiny ispyriscence, in which resin binds the cones shut until the resin is melted by a forest fire, for example inP. radiata andP. muricata. The seeds are then released after the fire, enabling them to colonise the burnt ground with minimal competition from other plants.[10][13]
The modern English name "pine" derives from Latinpinus, traced to the Indo-European base*pīt- 'resin'.[14] Before the 19th century, pines were often called firs, a name now applied to another genus,Abies. In some European languages, Germanic cognates of the Old Norse name are still in use for pines, as inDanishfyr andGermanFöhre.[15] The genusPinus was named byCarl Linnaeus in 1753.Pinus sylvestris, the Scots pine, was later chosen as thetype species.[16][17]
Based ontranscriptome analysis,Pinus is most closely related to the genusCathaya, which in turn is closely related to the genusPicea, the spruces. These genera, withfirs andlarches, form the pinoidclade of the Pinaceae.[21]
The evolutionary history of the genusPinus has been complicated byhybridisation. Pines are prone to inter-specific breeding. Wind pollination, long life spans, overlapping generations, large population size, and weakreproductive 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.[22][23] Research using large genetic datasets has clarified these relationships.[24] Two 21st century phylogenies are given below; the differences between them demonstrate these complications:
Pines aregymnosperms. The genus is divided into two subgenera based on the number offibrovascular 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:[10]
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 andPinus lumholtzii, have deciduous sheaths).[10]
Pinus subg.Strobus,syn.Pinus subg.Ducampopinus, the white or soft pine, and pinyon pine groups, with cones without a resin seal on the scales, and usually have softer wood; the needle fascicles mostly have a deciduous sheath (one exception,Pinus nelsonii, has a persistent sheath).[10]
Phylogenetic evidence indicates that the subgenera diverged anciently from one another.[21] Each subgenus is further divided into sections and subsections.[24]
World Flora Online accepts 134 species-rank taxa (119 species and 15nothospecies) of pines as current, with additional synonyms,[26] andPlants of the World Online 126 species-rank taxa (113 species and 13 nothospecies),[27] making it the largest genus among the conifers. The highest species diversity of pines is found in Mexico.[28][29][30]
Pines are native to theNorthern Hemisphere, with the most species in North America, some in Asia, and a few in Europe. Only two species,Pinus sylvestris andPinus sibirica, occur in more than one of those regions (Asia and Europe).[31] They occupy large areas ofboreal forest (taiga) in latitudes between 50° and 60° N; about a third of thisbiome is in North America andScandinavia, the rest inSiberia.[32] The northernmost species isScots pine, reaching just north of 70° N inStabbursdalen National Park in Norway;[33] One species,Pinus merkusii, crosses the equator inSumatra to 2°S.[34] In North America, various species occur in regions at latitudes from as far north as 66° N[34] to as far south as 12°N.[35]
Various species have been introduced to temperate andsubtropical regions of both hemispheres, where they are grown astimber or cultivated as ornamental plants in parks and gardens. A number of such introduced species have becomenaturalised, and species such asPinus radiata are consideredinvasive in some regions.[36]
Pines grow 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 favourable soils.[37]
Pine needles serve as food for thecaterpillar larvae of severalmoth species including thepine beauty, a pest of mature stands of pine trees,[45] and thepine hawk-moth, a large species which causes only occasional damage.[46] Some moths, notably thepine processionary, whose caterpillars can completely defoliate pine trees,[47][48][49] and thepine-tree lappet, are serious pests of commercial forestry.[50]
Pine-tree lappet (Dendrolimus pini) caterpillars on Scots pine with damage to needles
Several species of pine are attacked bynematodes, causingpine wilt disease, which can quickly kill trees.[51][52]The sawflyDiprion pini is likewise a serious commercial pest of pine forestry, especially ofPinus sylvestris.[53] Some birds such asnutcrackers are specialist feeders on pine seeds, and are important in distributing the seeds widely.[54][55]Crossbills rely onPinus sylvestris seeds in Scotland, and similarly help significantly to disperse the seeds, whereasred squirrels feed on the seeds but do little for seed dispersal.[56] Pinepollen may contribute tofood webs involvingdetritivores. Nutrients from pollen aid detritivores in development, growth, and maturation, and may enable fungi to decompose plant litter which is low in nutrients.[57] The edible[58]basidiomycete fungusBoletus pinophilus (pine bolete) forms anectomycorrhizal association with pines such asP. cembra,P. nigra, andP. sylvestris.[59]
Mass ofDiprion pini sawfly caterpillars on pine tree
Pines are among the most commercially important tree species, valued for their timber andwood pulp throughout the world.[60][61] In temperate and tropical regions, they are fast-growingsoftwoods that grow in relatively dense stands. Commercial pines are grown inplantations 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.[62]
As pine wood has little resistance to insects or decay after logging, in its untreated state it is generally recommended for indoor construction purposes only, such as indoordrywall framing. It is commonly used inCanadian Lumber Standard graded wood.[63] For outside use, pine needs to be treated with copper azole,chromated copper arsenate or other suitablechemical preservatives.[64]
The seeds (pine nuts) are generally edible; the young male cones can be cooked and eaten, as can the bark of young twigs.[66] Some species have large pine nuts, which are harvested and sold for cooking and baking. They are an ingredient ofpesto alla genovese.[67]
The soft, moist, white inner bark (cambium) beneath the woody outer bark is edible and very high in vitaminsA andC.[65] It can be eaten raw in slices as a snack or dried and ground up into a powder for use as anersatz flour or thickener in stews, soups, and other foods, such asbark bread.[68] The use of pine cambium gave the Adirondack Indians their name, from theMohawk Indian wordatirú:taks, meaning "tree eaters".[68]
Aherbal tea is made by steeping young, green pine needles in boiling water (known astallstrunt in Sweden).[68] In eastern Asia, pine and other conifers are accepted among consumers as a beverage product, and used in teas, as well as wine.[69] In Greece, the wineretsina is flavoured with resin fromPinus halepensis (Aleppo pine).[70]
Turpentine oil, traditionally used as asolvent inpaints,resins andvarnishes, is extracted from pine resin[71] or pine wood.[72] Pine needles are woven into baskets in Latin America.[73] Intraditional Chinese medicine,pine resin is used for burns, wounds and skin complaints.[74] Chinese ink sticks forcalligraphy are often made of pinesoot, producing a matt black ink when mixed with water.[75] Pine needles have been used by Latvian designer Tamara Orjola to createbiodegradable products including paper, furniture, textiles and dyestuffs.[76]
In ancient Egypt, the godOsiris was honoured with an image placed in a cavity inside a pine tree. In ancient Greece, the goddess Pitthea was linked with pines, while in ancient Rome, the tree was worshipped in the festival of the godAttis and the goddessCybele.[78] The Greek god of wine,Dionysus (also called Bacchus), was associated with pine as a symbol of fertility, and his devotees carried a stick topped with a pine cone (athyrsus), aphallic symbol.[78] TheBuryat people of Siberia revered groves ofPinus sylvestris, while ancient CelticDruids marked the midwinter solstice with fires of the same species.[78]
Pines are often featured in paintings. A 2021 study lists over a hundred works: many are by artists from the Mediterranean region, such asPaul Cézanne andJean-Baptiste-Camille Corot; Northern Europe, such asAkseli Gallen-Kallela andJames William Giles; and North America, with works byTom Thomson and others. The paintings often depictPinus pinea by theMediterranean sea; other species includeP. sylvestris andP. pinaster.[79] The pine is a particular motif in Chinese art and literature, which sometimes combines painting and poetry in the same work. The pine symbolises longevity and steadfastness, as it retains its green needles throughout the year. Sometimes the pine and cypress are paired. At other times the pine, plum, and bamboo are considered as the "Three Friends of Winter".[80]
Amaenad (follower ofDionysus) holding athyrsus topped with a pine cone. Roman, 120–140 AD
Pines on Mount Tai,Ike no Taiga, Japan, 18th century
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