Pine

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Pine tree
Pinus densiflora Kumgangsan.jpg
Korean red pine (Pinus densiflora), North Korea
Scientific classification Red Pencil Icon.png
Kingdom: Plantae
Clade: Tracheophytes
Division: Pinophyta
Class: Pinopsida
Order: Pinales
Family: Pinaceae
Subfamily: Pinoideae
Genus: Pinus
L.
Subgenera

See List of Pinus species for complete taxonomy to species level. See list of pines by region for list of species by geographic distribution.

Contents

Pinus range.png
Range of Pinus

A pine is any conifer in the genus Pinus ( /ˈpns/ [1] ) of the family Pinaceae. Pinus is the sole genus in the subfamily Pinoideae. The Plant List compiled by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accepts 126 species names of pines as current, together with 35 unresolved species and many more synonyms. [2] Pine may also refer to the lumber derived from pine trees; pine is one of the more extensively used types of wood used as lumber.

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). [3] Before the 19th century, pines were often referred to as firs (from Old Norse fura, by way of Middle English firre). In some European languages, Germanic cognates of the Old Norse name are still in use for pines—in Danish fyr, in Norwegian fura/fure/furu, Swedish fura/furu, Dutch vuren, and German Föhre—but in modern English, fir is now restricted to fir (Abies) and Douglas fir (Pseudotsuga).

Description

Illustration of needles, cones, and seeds of Scots pine (Pinus sylvestris) Illustration Pinus sylvestris0 new.jpg
Illustration of needles, cones, and seeds of Scots pine (Pinus sylvestris)

Pine trees are evergreen, coniferous resinous trees (or, rarely, shrubs) growing 3–80 m (10–260 ft) tall, with the majority of species reaching 15–45 m (50–150 ft) tall. [4] The smallest are Siberian dwarf pine and Potosi pinyon, and the tallest is an 81.79 m (268.35 ft) tall ponderosa pine located in southern Oregon's Rogue River-Siskiyou National Forest. [5]

Pines are long lived and typically reach ages of 100–1,000 years, some even more. The longest-lived is the Great Basin bristlecone pine, Pinus longaeva. One individual of this species, dubbed "Methuselah", is one of the world's oldest living organisms at around 4,600 years old. This tree can be found in the White Mountains of California. [6] 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" after the Greek immortal.[ citation needed ]

The spiral growth of branches, needles, and cone scales may be arranged in Fibonacci number ratios. [7] [8] 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 foresters a means to evaluate fertility of the soil and vigour of the trees.

Bark

The bark of most pines is thick and scaly, but some species have thin, flaky bark. [9] The branches are produced in regular "pseudo whorls", actually a very tight spiral but appearing like a ring of branches arising from the same point. Many pines are uninodal, producing just one such whorl of branches each year, from buds at the tip of the year's new shoot, but others are multinodal, producing two or more whorls of branches per year.

Foliage

Pines have four types of leaf:

Cones

Pine cone A pine cone.jpg
Pine cone

Pines are mostly monoecious, having the male and female cones on the same tree, though a few species are sub-dioecious, with individuals predominantly, but not wholly, single-sex. 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 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 (wind-dispersed), but some are larger and have only a vestigial wing, and are bird-dispersed (see below). 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, [10] ), 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 a resin binds the cones shut until melted by a forest fire, for example in Pinus rigida .

Evolutionary history

Conifers evolved about 300 million years ago, and pines perhaps around 153 million years ago. The genus Pinus is thought to have diverged from other pines approximately 95 million years ago.

Pinus is the largest genus of the Pinaceae, the pine family, which is the oldest and largest conifer family. It dates back to 206 million years ago. [11] Based on recent transriptome analysis, Pinus is most closely related to the genus Cathaya , which in turn is closely related to spruces. These genera, with firs and larches, form the pinoid clade of the Pinaceae.

The evolutionary history of the genus Pinus has been complicated by hybridization. Pines are prone to inter-specific breeding. Wind pollination, long life spans, overlapping generations, large population size, and weak reproductive isolation make breeding across species more likely. [12] As the pines have diversified, gene transfer between different species has created a complex history of genetic relatedness.

Taxonomy, nomenclature and codification

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

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. [14] Recent research using large genetic datasets has clarified these relationships into the groupings we recognize today.

Distribution

Monterey Pine in Sydney, Australia, which were introduced to the region in the late 19th century. Prospectsydneypineforest.jpg
Monterey Pine in Sydney, Australia, which were introduced to the region in the late 19th century.

Pines are native to the Northern Hemisphere, and in a few parts of the tropics in the Southern Hemisphere. Most regions of the Northern Hemisphere (see List of pines by region) host some native species of pines. One species (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.[ citation needed ]

Pines may be found in a very large variety of environments, ranging from semi-arid desert to rainforests, from sea level up to 5,200 metres (17,100 ft), from the coldest to the hottest environments on Earth. They often occur in mountainous areas with favorable soils and at least some water. [15]

Various species have been introduced to temperate and subtropical regions of both hemispheres, where they are grown as timber or cultivated as ornamental plants in parks and gardens. A number of such introduced species have become naturalized, and some species are considered invasive in some areas [16] and threaten native ecosystems.

Ecology

Pine forest in Vagamon, southern Western Ghats, Kerala (India) Vagamon Pine Forest.jpg
Pine forest in Vagamon, southern Western Ghats, Kerala (India)

Pines grow well in acid soils, some also on calcareous soils; most require good soil drainage, preferring sandy soils, but a few (e.g. lodgepole pine) can tolerate poorly drained wet soils. A few are able to sprout after forest fires (e.g. Canary Island pine). Some species of pines (e.g. bishop pine) need fire to regenerate, and their populations slowly decline under fire suppression regimens.

Several species are adapted to extreme conditions imposed by elevation and latitude (e.g. Siberian dwarf pine, mountain pine, whitebark pine, and the bristlecone pines). The pinyon pines and a number of others, notably Turkish pine and gray pine, are particularly well adapted to growth in hot, dry semidesert climates.[ citation needed ]

The seeds are commonly eaten by birds, such as grouse, crossbills, jays, nuthatches, siskins, and woodpeckers, and by squirrels. Some birds, notably the spotted nutcracker, Clark's nutcracker, and pinyon jay, are of importance in distributing pine seeds to new areas. Pine needles are sometimes eaten by some Lepidoptera (butterfly and moth) species (see list of Lepidoptera that feed on pines), the Symphytan species pine sawfly, and goats.[ citation needed ]

Pine pollen may play an important role in the functioning of detrital food webs. [17] Nutrients from pollen aid detritivores in development, growth, and maturation, and may enable fungi to decompose nutritionally scarce litter. [17] Pine pollen is also involved in moving plant matter between terrestrial and aquatic ecosystems. [17]

Uses

Lumber and construction

Logging Pinus ponderosa, Arizona, USA John Deere 2054 DHSP forestry swing machine, Kaibab National Forest 1.jpg
Logging Pinus ponderosa , Arizona, USA

Pines are among the most commercially important tree species valued for their timber and wood pulp throughout the world. [18] [19] In temperate and tropical regions, they are fast-growing softwoods that grow in relatively dense stands, their acidic decaying needles inhibiting the sprouting of competing hardwoods. Commercial pines are grown in plantations for timber that is denser and therefore more durable than spruce (Picea). Pine wood is widely used in high-value carpentry items such as furniture, window frames, panelling, floors, and roofing, and the resin of some species is an important source of turpentine.

Because pines have no insect- or decay-resistant qualities after logging, untreated they are generally recommended for construction purposes as indoor use only (indoor drywall framing, for example). For outside use, pine needs to be treated with copper azole, chromated copper arsenate or other suitable chemical preservative. [20] .

Ornamental uses

"Pine Clouds", 1903 painting on fan by Wu Ku-hsiang 'Pine Clouds', 1903 painting by Wu Ku-hsiang.jpg
"Pine Clouds", 1903 painting on fan by Wu Ku-hsiang

Many pine species make attractive ornamental plantings for parks and larger gardens with a variety of dwarf cultivars being suitable for smaller spaces. Pines are also commercially grown and harvested for Christmas trees. 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. [21] Pine needles are also used for making decorative articles such as baskets, trays, pots, etc, and during the U.S. Civil War, the needles of the longleaf pine "Georgia pine" were widely employed in this. [22] 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 products including paper, furniture, textiles and dye. [23]

Wildlife

Pine beauty moth (Panolis flammea) on pine needles Panolis.flammea.7102.jpg
Pine beauty moth (Panolis flammea) on pine needles

Pine needles serve as food for various Lepidoptera. Several species of pine are attacked by nematodes, causing pine wilt disease, 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 nuts.

Farming

Forchem tall oil refinery in Rauma, Finland. Forchem Rauma 2.jpg
Forchem tall oil refinery in Rauma, Finland.

When grown for sawing timber, pine plantations can be harvested after 30 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, while most older ones are good enough for saw timber.

The final wood quality can be improved by pruning small branches at ages 5, 7, and 9. Pruning usually goes up to a height of 6 metres (20 ft). This results in smooth timber with no knots, which is considerably more valuable. [24]

A 30-year-old commercial pine tree grown in good conditions will be about 0.3 m (1.0 ft) in diameter and about 20 m (66 ft) high. After 50 years, the same tree will be about 0.5 m (1.6 ft) in diameter and 25 m (82 ft) high, and its wood will be worth about seven times as much as the 30-year-old tree. [25]

Trees are planted 3–4 m apart, or about 1000 per hectare (100,000 per km2).

Food and nutrients

Edible seeds of the Korean pine (Pinus koraiensis) KoreanPineSeeds.jpg
Edible seeds of the Korean pine ( Pinus koraiensis )

Some species have large seeds, called pine nuts, that are harvested and sold for cooking and baking. They are an essential ingredient of pesto alla genovese.

The soft, moist, white inner bark (cambium) beneath the woody outer bark is edible and very high in vitamins A and C.[ citation needed ] 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 Indian word atirú:taks, meaning "tree eaters".

A tea made by steeping young, green pine needles in boiling water (known as tallstrunt in Sweden) is high in vitamins A and C.[ citation needed ] In eastern Asia, pine and other conifers are accepted among consumers as a beverage product, and used in teas, as well as wine. [26]

Pine needles from Pinus densiflora were found to contain 30.54 mg/g of proanthocyanidins when extracted with hot water. [27] Comparative to ethanol extraction resulting in 30.11 mg/g, simply extracting in hot water is preferable.

In traditional Chinese medicine (TCM), pine resin is used for burns, wounds and dermal complaints. [28]

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, [29] Dora Sigerson Shorter, [30] Eugene Field, [31] the Chinese, [32] Theodore Winthrop, [33] and Rev. George Allan D.D. [34]

Art

Under the Pines, Evening, Claude Monet (1888) (Philadelphia Museum of Art) Under the Pines PMA(15) (16982300531).jpg
Under the Pines, Evening, Claude Monet (1888) (Philadelphia Museum of Art)
By Camille Pissarro. Pissarro - kew-gardens-crossroads-near-the-pond-1892.jpg
By Camille Pissarro.

Pines are often featured in art, whether painting and fine art, [35] drawing, [36] photography, or folk art.

Religious texts

Pine trees, as well as other conifers, are mentioned in The Bible. In Nehemiah 8:15, the King James Version renders the following translation: [37]

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

Pines are also mentioned in Isaiah 41:

"17: When the poor and needy seek water, and there is none, and their tongue faileth for thirst, I the LORD will hear them, I the God of Israel will not forsake them. 18: I will open rivers in high places, and fountains in the midst of the valleys: I will make the wilderness a pool of water, and the dry land springs of water. 19: I will plant in the wilderness the cedar, the shittah tree, and the myrtle, and the oil tree; I will set in the desert the fir tree, and the pine, and the box tree together: 20: That they may see, and know, and consider, and understand together, that the hand of the LORD hath done this, and the Holy One of Israel hath created it."

And in Isaiah 60:

"13: 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."

Chinese culture

Many Chinese ink sticks are made out of pine: the wood being burnt and the soot collected as an ingredient to provide the coloring Hu Kai Wen Long Xiang Feng Wu Zhu Xing Mo .jpg
Many Chinese ink sticks are made out of pine: the wood being burnt and the soot collected as an ingredient to provide the coloring

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". [38] 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

Related Research Articles

Douglas fir species of tree

Pseudotsuga menziesii is an evergreen conifer species in the pine family, Pinaceae. It is native to western North America and is known as Douglas fir, Douglas-fir, Oregon pine, and Columbian pine. Despite its common name, it is not a true fir. There are three varieties: coast Douglas-fir, Rocky Mountain Douglas-fir and Mexican Douglas fir.

<i>Cedrus</i> Genus of plants (coniferous trees)

Cedrus, common English name cedar, is a genus of coniferous trees in the plant family Pinaceae. They are native to the mountains of the western Himalayas and the Mediterranean region, occurring at altitudes of 1,500–3,200 m in the Himalayas and 1,000–2,200 m in the Mediterranean.

Pinophyta Division of plants

The Pinophyta, also known as Coniferophyta or Coniferae, or commonly as conifers, are a division of vascular land plants containing a single extant class, Pinopsida. They are gymnosperms, cone-bearing seed plants. All extant conifers are perennial woody plants with secondary growth. The great majority are trees, though a few are shrubs. Examples include cedars, Douglas firs, cypresses, firs, junipers, kauri, larches, pines, hemlocks, redwoods, spruces, and yews. As of 1998, the division Pinophyta was estimated to contain eight families, 68 genera, and 629 living species.

Scots pine Species of conifer in the family Pinaceae

Scots pine is a species of pine that is native to Eurasia, ranging from Western Europe to Eastern Siberia, south to the Caucasus Mountains and Anatolia, and north to well inside the Arctic Circle in Fennoscandia. In the north of its range, it occurs from sea level to 1,000 m (3,300 ft), while in the south of its range it is a mountain tree, growing at 1,200–2,600 m (3,900–8,500 ft) altitude. It is readily identified by its combination of fairly short, blue-green leaves and orange-red bark.

Pinaceae family of plants

The Pinaceae, pine family, are trees or shrubs, including many of the well-known conifers of commercial importance such as cedars, firs, hemlocks, larches, pines and spruces. The family is included in the order Pinales, formerly known as Coniferales. Pinaceae are supported as monophyletic by their protein-type sieve cell plastids, pattern of proembryogeny, and lack of bioflavonoids. They are the largest extant conifer family in species diversity, with between 220 and 250 species in 11 genera, and the second-largest in geographical range, found in most of the Northern Hemisphere, with the majority of the species in temperate climates, but ranging from subarctic to tropical. The family often forms the dominant component of boreal, coastal, and montane forests. One species, Pinus merkusii, grows just south of the equator in Southeast Asia. Major centres of diversity are found in the mountains of southwest China, Mexico, central Japan, and California.

<i>Tsuga</i> genus of plants

Tsuga is a genus of conifers in the subfamily Abietoideae. The common name hemlock (tree) is derived from a perceived similarity in the smell of its crushed foliage to that of the unrelated plant poison hemlock. Unlike the latter, Tsuga species are not poisonous.

<i>Pinus strobiformis</i> species of plant

Pinus strobiformis, commonly known as southwestern white pine, Mexican white pine or Chihuahua white pine, is a medium-sized white pine tree whose native habitat is in southwestern United States and Mexico. It is typically a high-elevation pine growing mixed with other conifers.

<i>Pinus wallichiana</i> species of plant

Pinus wallichiana is a coniferous evergreen tree native to the Himalaya, Karakoram and Hindu Kush mountains, from eastern Afghanistan east across northern Pakistan and north west India to Yunnan in southwest China. It grows in mountain valleys at altitudes of 1800–4300 m, reaching 30–50 m (98–164 ft) in height. It favours a temperate climate with dry winters and wet summers. In Pashto, it is known as Nishtar.

<i>Pinus virginiana</i> species of plant

Pinus virginiana, the Virginia pine, scrub pine, Jersey pine, is a medium-sized tree, often found on poorer soils from Long Island in southern New York south through the Appalachian Mountains to western Tennessee and Alabama. The usual size range for this pine is 9–18 m, but can grow larger under optimum conditions. The trunk can be as large as 0.5 m diameter. This tree prefers well-drained loam or clay, but will also grow on very poor, sandy soil, where it remains small and stunted. The typical life span is 65 to 90 years.

<i>Pinus ponderosa</i> Species of large pine tree in North America

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 most widely distributed pine species in North America.

Conifer cone Reproductive organ on conifers

A cone is an organ on plants in the division Pinophyta (conifers) that contains the reproductive structures. The familiar woody cone is the female cone, which produces seeds. The male cones, which produce pollen, are usually herbaceous and much less conspicuous even at full maturity. The name "cone" derives from the fact that the shape in some species resembles a geometric cone. The individual plates of a cone are known as scales.

<i>Pseudolarix</i> monotypic genus of deciduous conifers in the family Pinaceae

Pseudolarix is a monotypic genus of coniferous trees in the pine family Pinaceae. The sole species, Pseudolarix amabilis, is commonly known as the golden larch, though it is not a true larch (Larix), being more closely related to Keteleeria, Abies and Cedrus. It is native to eastern China, occurring in small areas in the mountains of southern Anhui, Zhejiang, Fujian, Jiangxi, Hunan, Hubei and eastern Sichuan, at altitudes of 100–1,500 m (328–4,921 ft). Golden larch is sometimes known under an old scientific name Pseudolarix kaempferi, but this may cause confusion with Larix kaempferi, the Japanese larch.

<i>Pinus roxburghii</i> species of plant, Chir Pine

Pinus roxburghii, commonly known as chir pine or longleaf Indian pine, is a species of pine, native to the Himalayas. It was named after William Roxburgh.

<i>Pinus kesiya</i> species of plant, Khasi pine

Pinus kesiya is one of the most widely distributed pines in Asia. Its range extends south and east from the Khasi Hills in the northeast Indian state of Meghalaya, to northern Thailand, Philippines, Burma, Cambodia, Laos, southernmost China, and Vietnam. It is an important plantation species elsewhere in the world, including in southern Africa and South America.

<i>Pinus aristata</i> Species of plant

Pinus aristata, the Rocky Mountain bristlecone pine, is a long-living species of bristlecone pine tree native to the United States. It appears in the Rocky Mountains in Colorado and northern New Mexico, with isolated populations in the San Francisco Peaks in Arizona and the Kaibab National Forest north of the Grand Canyon. It is usually found at very high altitudes, from 7,000–13,000 feet (2,100–4,000 m), in cold, dry subalpine climate conditions, often at the tree line, although it also forms extensive closed-canopy stands at somewhat lower elevations.

<i>Pinus durangensis</i> species of plant, Durango Pine

Pinus durangensis, the Durango pine, is a pine tree species endemic to the Sierra Madre Occidental mountain range of Northwestern Mexico.

<i>Pinus maximartinezii</i> species of plant

Pinus maximartinezii, called Martinez pinyon, big-cone pinyon or maxipiñon, is a pine in the pinyon pine group, native to central Mexico. The range is highly localised, confined to a small area of the southern Sierra Madre Occidental in southern Zacatecas. It occurs at moderate altitudes, from 1800–2400 m and 21° North latitude, in warm and temperate, dry climate conditions.

<i>Pinus massoniana</i> species of plant

Pinus massoniana is a species of pine, native to Taiwan, and a wide area of central and southern China, including Hong Kong, and northern Vietnam, growing at low to moderate altitudes, mostly below 1,500 m but rarely up to 2,000 m altitude.

<i>Pinus greggii</i> species of plant

Pinus greggii, or Gregg's pine, is a small to medium high pine tree native to eastern Mexico, found in two distinct regions. It has an open crown and long and slender branches. The needles are in bundles of three with an average length of 11 cm. The cones are clustered in groups of 5 – 10. The branches en upper trunk is smooth. Pinus greggii is introduced in several countries.

<i>Pseudotsuga menziesii <span style="font-style:normal;">var.</span> menziesii</i> a variety of Douglas-fir in the Pacific Northwest

Pseudotsuga menziesii var. menziesii, also known as Coast Douglas-fir, Pacific Douglas-fir, Oregon pine, or Douglas spruce, is an evergreen conifer native to western North America from west-central British Columbia, Canada southward to central California, United States. In Oregon and Washington its range is continuous from the Cascades crest west to the Pacific Coast Ranges and Pacific Ocean. In California, it is found in the Klamath and California Coast Ranges as far south as the Santa Lucia Mountains with a small stand as far south as the Purisima Hills, Santa Barbara County. In the Sierra Nevada it ranges as far south as the Yosemite region. It occurs from near sea level along the coast to 1,800 metres (5,900 ft) in the California Mountains. Further inland, coast Douglas-fir is replaced by Rocky Mountain or interior Douglas-fir. Interior Douglas-fir intergrades with coast Douglas-fir in the Cascades of northern Washington and southern British Columbia.

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