Genus of plants in the conifer family Pinaceae
A
pine
is any
conifer
tree
or
shrub
in the
genus
Pinus
(
)
[1]
of the
family
Pinaceae
.
Pinus
is the sole
genus
in the
subfamily
Pinoideae
.
World Flora Online
accepts 187 species names of pines as current, with additional synonyms, making it the largest family among the conifers.
[2]
The
American Conifer Society
(ACS) and the
Royal Horticultural Society
accept 121 species. Pines are widely
distributed
in the
Northern Hemisphere
; they occupy large areas of
boreal forest
, but are found in many habitats, including the
Mediterranean Basin
.
The
lumber
from pine trees is called "pine"; it is one of the more extensively used types of lumber.
There are currently 818 named
cultivars
(or
trinomials
) recognized by the ACS.
[3]
It is a well-known type of
Christmas tree
.
Description
[
edit
]
Pine trees are
evergreen
, coniferous
resinous
trees
(or, rarely,
shrubs
) growing 3?80 metres (10?260 feet) 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 81.8 m (268 ft) tall
ponderosa pine
located in southern
Oregon
's
Rogue River-Siskiyou National Forest
.
[4]
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
(
P. longaeva
). One individual of this species, dubbed "
Methuselah
", 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.
[5]
An older tree, now cut down, was dated at 4,900 years old.
[6]
[7]
It was discovered in a grove beneath
Wheeler Peak
and it is now known as "
Prometheus
" after the
Greek immortal
.
[7]
The spiral growth of branches, needles, and
cones
scales
are arranged
in
Fibonacci number
ratios.
[8]
[9]
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
soil fertility
and vigour of the trees.
Bark
[
edit
]
The
bark
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
buds
at the tip of the year's new
shoot
, but others are multinodal, producing two or more whorls of branches per year.
Foliage
[
edit
]
Pines have four types of
leaf
:
- Seed leaves (
cotyledons
) on seedlings are borne in a whorl of 4?24.
- Juvenile leaves, which follow immediately on seedlings and young plants, are
2?6 centimetres (
3
⁄
4
?
2
+
1
⁄
4
inches) long, single, green or often blue-green, and arranged spirally on the shoot. These are produced for six months to five years, rarely longer.
- Scale leaves, similar to bud scales, are small, brown and not photosynthetic, and arranged spirally like the juvenile leaves.
- Needles, the adult leaves, are green (
photosynthetic
) 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
[
edit
]
Pines are
monoecious
, having the male and female cones on the same tree.
[10]
: 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 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. 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
,
[11]
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
P. rigida
.
Etymology
[
edit
]
The modern English name "pine" derives from Latin
pinus
, which some have traced to the Indo-European base
*p?t-
‘resin’ (source of English
pituitary
).
[12]
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
Fohre
? but in modern English,
fir
is now restricted to
fir
(
Abies
) and
Douglas-fir
(
Pseudotsuga
).
Taxonomy
[
edit
]
Pines are
gymnosperms
. The genus is divided into two subgenera based on the number of
fibrovascular bundles
in the needle. The subgenera can be distinguished by cone, seed, and leaf characters:
- Pinus
subg.
Pinus
, the yellow, or hard pine group, generally with harder wood and two or three needles per
fascicle
.
[13]
The subgenus is also named
diploxylon
, on account of its two fibrovascular bundles.
- Pinus
subg.
Strobus
, the white, or soft pine group. Its members usually have softer wood and five needles per fascicle.
[13]
The subgenus is also named
haploxylon
, on account of its one fibrovascular bundle.
Phylogenetic evidence indicates that both subgenera have a very ancient divergence from one another.
[14]
Each subgenus is further divided into sections and subsections.
Many of the smaller groups of
Pinus
are composed of closely related species with recent divergence and history of hybridization. This results in low morphological and genetic differences. This, coupled with low sampling and underdeveloped genetic techniques, has made taxonomy difficult to determine.
[15]
Recent research using large genetic datasets has clarified these relationships into the groupings we recognize today.
Phylogeny
[
edit
]
Pinus
is the largest genus of the
Pinaceae
, the pine family, which first appeared in the
Jurassic
period.
[16]
Based on recent
Transcriptome
analysis,
Pinus
is most closely related to the genus
Cathaya
, which in turn is closely related to
spruces
. These genera, with
firs
and
larches
, form the pinoid
clade
of the Pinaceae.
[14]
Pines first appeared during the Early Cretaceous, with the oldest verified fossil of the genus being
Pinus yorkshirensis
from the
Hauterivian
-
Barremian
boundary (~130-125 million years ago) from the
Speeton Clay
, England.
[17]
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.
[18]
As the pines have diversified, gene transfer between different species has created a complex history of genetic relatedness.
Jin et al. 2021
[19]
|
Stull et al. 2021
[14]
|
Pinus
|
subgenus
Strobus
|
section
Parrya
|
|
|
section
Quinquefoliae
|
|
|
|
|
subgenus
Pinus
|
section
Pinus
|
subsection
Pinaster
|
|
|
subsection
Pinus
|
|
|
|
|
section
Trifoliae
|
|
|
|
|
|
|
|
|
|
|
|
|
subsection
Contorta
|
|
|
subsection
Australes
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Pinus
|
(
Strobus
)
|
section
|
|
Parrya
|
section
|
|
|
|
|
subsection
|
P. krempfii
|
Krempfii
|
subsection
|
|
|
|
|
|
|
|
|
series
|
|
Cembrae
|
series
|
|
Pumilae
|
|
|
|
|
|
Strobus
|
|
|
|
Quinquefoliae
|
|
|
(
Pinus
)
|
section
|
subsection
|
|
Pinaster
|
subsection
|
|
|
|
|
series
|
|
Tropicales
|
series
|
|
Tabuliformes
|
|
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|
Pinus
|
|
Pinus
|
section
|
|
|
|
|
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|
|
subsection
|
series
|
|
Leiophyllae
|
series
|
|
Attenuatae
|
|
Attenuata
|
subsection
|
|
Australes
|
|
|
|
|
|
Trifoliae
|
|
|
|
|
|
Distribution and habitat
[
edit
]
Pines are native to the
Northern Hemisphere
, and to a few parts from the tropics to
temperate
regions in the
Southern Hemisphere
. Most regions of the Northern Hemisphere host some
native species
of pines; they occupy large areas of
Boreal forest
, and are found all around the Mediterranean Basin. 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
[
citation needed
]
to as far south as 12°N.
[20]
Pines may be found in a very large variety of environments, ranging from semi-arid desert to rainforests, from sea level up to 5,200 m (17,100 ft), from the coldest to the hottest environments on Earth. They often occur in mountainous areas with favorable soils and at least some water.
[21]
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
[22]
and threaten native ecosystems.
Ecology
[
edit
]
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.
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.
[23]
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.
[24]
Pine
pollen
may play an important role in the functioning of
detrital
food webs
.
[25]
Nutrients from pollen aid detritivores in development, growth, and maturation, and may enable fungi to decompose nutritionally scarce litter.
[25]
Pine pollen is also involved in moving plant matter between terrestrial and aquatic ecosystems.
[25]
Wildlife
[
edit
]
Pine needles serve as food for various
Lepidoptera
(
butterfly
and
moth
) species. 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
. 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 the
Symphytan
species
pine sawfly
, and
goats
.
[26]
Uses
[
edit
]
Lumber and construction
[
edit
]
Pines are among the most commercially important tree species, valued for their timber and
wood pulp
throughout the world.
[27]
[28]
In temperate and tropical regions, they are fast-growing
softwoods
that grow in relatively dense stands. Commercial pines are grown in
plantations
for timber that is denser and therefore more durable than spruce (
Picea
). Pine wood is widely used in high-value carpentry items such as furniture, window frames, panelling, floors, and roofing, and the resin of some species is an important source of
turpentine
.
Because pine wood has no insect- or decay-resistant qualities after logging, in its untreated state it is generally recommended for indoor construction purposes only (indoor
drywall
framing, for example). For outside use, pine needs to be treated with copper azole,
chromated copper arsenate
or other suitable
chemical preservative
.
[29]
Ornamental uses
[
edit
]
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.
[30]
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.
[31]
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.
[32]
Farming
[
edit
]
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.
[33]
A 30-year-old commercial pine tree grown in good conditions in Arkansas will be about 0.3 m (1 ft) in diameter and about 20 m (66 ft) high. After 50 years, the same tree will be about
0.5 m (
1
+
1
⁄
2
ft) in diameter and 25 m (82 ft) high, and its wood will be worth about seven times as much as the 30-year-old tree.
This however depends on the region, species and silvicultural techniques. In New Zealand, a plantation's maximum value is reached after around 28 years with height being as high as 30 m (98 ft) and diameter
0.5 m (
1
+
1
⁄
2
ft), with maximum wood production after around 35 years (again depending on factors such as site, stocking and genetics). Trees are normally planted 3?4 m apart, or about 1,000 per hectare (100,000 per square kilometre).
[34]
[35]
[36]
[37]
Food and nutrients
[
edit
]
The seeds (pine nuts) are generally edible; the young male cones can be cooked and eaten, as can the bark of young twigs.
[38]
Some species have large pine nuts, which are harvested and sold for cooking and baking. They are an essential ingredient of
pesto
alla genovese
.
The soft, moist, white inner bark (
cambium
) beneath the woody outer bark is edible and very high in vitamins
A
and
C
.
[3]
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
.
[39]
Adirondack Indians got their name from the
Mohawk Indian
word
atiru:taks
, meaning "tree eaters".
[39]
A
tea
is made by steeping young, green pine needles in boiling water (known as
tallstrunt
in Sweden).
[39]
In eastern Asia, pine and other conifers are accepted among consumers as a beverage product, and used in teas, as well as wine.
[40]
In Greece, the wine
retsina
is flavoured with Aleppo pine resin.
Pine needles from
Pinus densiflora
were found to contain 30.54 milligram/gram of
proanthocyanidins
when extracted with hot water.
[41]
Comparative to ethanol extraction resulting in 30.11 mg/g, simply extracting in hot water is preferable.
In
traditional Chinese medicine
,
pine resin
is used for burns, wounds and dermal complaints.
[42]
Culture
[
edit
]
Pines have been a frequently mentioned tree throughout history, including in literature, art, and in religious texts.
The pine is a particular motif in Chinese art and literature, which sometimes combines painting and poetry in the same work. Some of the main symbolic attributes of pines in Chinese art and literature are longevity and steadfastness: the pine retains its green needles through all the seasons. Sometimes the pine and cypress are paired. At other times the pine, plum, and bamboo are considered as the "
Three Friends of Winter
".
[43]
Literature
[
edit
]
Writers of various nationalities and ethnicities have written of pines. Among them,
John Muir
,
[44]
Dora Sigerson Shorter
,
[45]
Eugene Field
,
[46]
Bai Juyi
,
[47]
Theodore Winthrop
,
[48]
and Rev. George Allan D.D.
[49]
Pines are often featured in art, whether painting and
fine art
,
[50]
drawing,
[51]
photography, or
folk art
.
Religious texts
[
edit
]
Pine trees, as well as other conifers, are mentioned in some verses of the
Bible
, depending on the translation. In the
Book of Nehemiah
8:15, the
King James Version
gives the following translation:
[52]
"And that they should publish and proclaim in all their cities, and in Jerusalem, saying, Go forth unto the mount, and fetch olive branches, and
pine branches
[emphasis added], and myrtle branches, and palm branches, and branches of thick trees, to make
booths
, as it is written."
However, the term here in Hebrew (?? ???) 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 (???? in Hebrew), and other translations use "pine" for the word translated as "box" by the King James (????? in Hebrew).
Some botanical authorities believe that the Hebrew word "????" (b?r?sh), which is used many times in the Bible, designates
P. halepensis
, or in
Hosea
14:8
[53]
which refers to fruit,
Pinus pinea
, the stone pine.
[54]
The word used in modern Hebrew for pine is "??????" (oren), which occurs only in Isaiah 44:14,
[55]
but two manuscripts have "???" (
cedar
), a much more common word.
[56]
See also
[
edit
]
References
[
edit
]
- ^
"Pinus".
Medical Definition of PINUS
.
Merriam-Webster Medical Dictionary
.
- ^
"
Pinus
(L.)"
.
World Flora Online
. The World Flora Online Consortium. 2022
. Retrieved
14 August
2022
.
- ^
a
b
"Pinus / pine | Conifer Genus"
.
American Conifer Society
. Retrieved
1 March
2022
.
- ^
a
b
Fattig, Paul (23 January 2011).
"Tallest of the tall"
.
Mail Tribune
. Medford, Oregon. Archived from
the original
on 23 September 2012
. Retrieved
27 January
2011
.
- ^
Ryan, Michael; Richardson, David M. (December 1999). "The Complete Pine".
BioScience
.
49
(12): 1023?1024.
JSTOR
1313736
.
- ^
Miranda, Carolina A. (28 February 2015).
"Follow-up: More tales of the Prometheus tree and how it died"
.
Los Angeles Times
. Retrieved
16 October
2020
.
- ^
a
b
Eveleth, Rose
(15 November 2012).
"How One Man Accidentally Killed the Oldest Tree Ever"
.
Smithsonian
.
Smithsonian Institution
. Retrieved
16 October
2020
.
- ^
Zeng, Lanling; Wang, Guozhao (2009).
"Modeling golden section in plants"
.
Progress in Natural Science
.
19
(2): 255?260.
doi
:
10.1016/j.pnsc.2008.07.004
.
The ratio between two pine needles is 0.618 [...] the angle between the two neighbors is about 135° and the angle between the main stem and each branch is close to 34.4° which is the golden section of 90°
- ^
Bracewell, Ronald; Rawlings, John.
"Pinus (Pine) Notes"
.
Trees of Stanford
. Retrieved
2 February
2020
.
- ^
Judd, W.S.; Campbell, C.S.; Kellogg, E.A.; Stevens, P.F.; Donoghue, M.J. (2002).
Plant systematics, a phylogenetic approach
(2 ed.). Sinauer Associates.
ISBN
0-87893-403-0
.
- ^
Tomback, Diana F.
(June 1982). "Dispersal of Whitebark Pine seeds by Clark's Nutcracker: a mutualism hypothesis".
The Journal of Animal Ecology
.
51
(2): 451?467.
Bibcode
:
1982JAnEc..51..451T
.
doi
:
10.2307/3976
.
JSTOR
3976
.
- ^
"Where Are You From? - Credo Reference"
.
credoreference.com
.
- ^
a
b
Barnes, Burton Verne; Wagner, Warren Herbert (January 2004).
Michigan Trees: A Guide to the Trees of the Great Lakes Region
. University of Michigan Press. pp. 81?.
ISBN
978-0-472-08921-5
.
Archived
from the original on 2016-05-11.
- ^
a
b
c
Stull, Gregory W.; Qu, Xiao-Jian; Parins-Fukuchi, Caroline; Yang, Ying-Ying; Yang, Jun-Bo; Yang, Zhi-Yun; Hu, Yi; Ma, Hong; Soltis, Pamela S.; Soltis, Douglas E.; Li, De-Zhu (July 19, 2021).
"Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms"
.
Nature Plants
.
7
(8): 1015?1025.
doi
:
10.1038/s41477-021-00964-4
.
ISSN
2055-0278
.
PMID
34282286
.
S2CID
236141481
.
Archived
from the original on January 10, 2022
. Retrieved
January 10,
2022
.
- ^
Flores-Renteria, Lluvia; Wegier, Ana; Ortega Del Vecchyo, Diego; Ortiz-Medrano, Alejandra; Pinero, Daniel; Whipple, Amy V.; et al. (December 2013). "Genetic, morphological, geographical and ecological approaches reveal phylogenetic relationships in complex groups, an example of recently diverged pinyon pine species (Subsection Cembroides)".
Molecular Phylogenetics and Evolution
.
69
(3): 940?9.
doi
:
10.1016/j.ympev.2013.06.010
.
PMID
23831459
.
- ^
Ran, Jin-Hua; Shen, Ting-Ting; Wu, Hui; Gong, Xun; Wang, Xiao-Quan (December 2018). "Phylogeny and evolutionary history of Pinaceae updated by transcriptomic analysis".
Molecular Phylogenetics and Evolution
.
129
: 106?116.
doi
:
10.1016/j.ympev.2018.08.011
.
PMID
30153503
.
S2CID
52110440
.
- ^
Ryberg, Patricia E.; Rothwell, Gar W.; Stockey, Ruth A.; Hilton, Jason; Mapes, Gene; Riding, James B. (2012). "Reconsidering Relationships among Stem and Crown Group Pinaceae: Oldest Record of the Genus
Pinus
from the Early Cretaceous of Yorkshire, United Kingdom".
International Journal of Plant Sciences
.
173
(8): 917?932.
doi
:
10.1086/667228
.
S2CID
85402168
.
- ^
Hernandez-Leon, Sergio; Gernandt, David S.; Perez de la Rosa, Jorge A.; Jardon-Barbolla, Lev (2013-07-30).
"Phylogenetic relationships and species delimitation in pinus section trifoliae inferrred from plastid DNA"
.
PLOS ONE
.
8
(7): e70501.
Bibcode
:
2013PLoSO...870501H
.
doi
:
10.1371/journal.pone.0070501
.
PMC
3728320
.
PMID
23936218
.
- ^
Jin, Wei-Tao; Gernandt, David S.; Wehenkel, Christian; Xia, Xiao-Mei; Wei, Xiao-Xin; Wang, Xiao-Quan (May 2021).
"Phylogenomic and ecological analyses reveal the spatiotemporal evolution of global pines"
.
PNAS
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edit
]
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