Pinus subsect. Balfourianae
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The term bristlecone pine covers three
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of
pine 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. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accepts ...
tree (family
Pinaceae The Pinaceae, or pine family, are conifer 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 kn ...
, 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. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden ...
'', subsection ''Balfourianae''). All three species are long-lived and highly resilient to harsh weather and bad soils. One of the three species, ''Pinus longaeva'', is among the longest-lived life forms on Earth. The oldest of this species is more than 4,800 years old, making it the oldest known individual of any species. Despite their potential age and low reproductive rate, bristlecone pines, particularly ''Pinus longaeva'', are usually a first-succession species, tending to occupy new open ground. They generally compete poorly in less-than-harsh environments, making them hard to cultivate. In gardens, they succumb quickly to root rot. They do very well, however, where most other plants cannot even grow, such as in rocky
dolomitic Dolomite () is an anhydrous carbonate mineral composed of calcium magnesium carbonate, ideally The term is also used for a sedimentary carbonate rock composed mostly of the mineral dolomite. An alternative name sometimes used for the dolomiti ...
soils in areas with virtually no rainfall. Bristlecone pines grow in scattered
subalpine Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures fall as elevation increases, causing the ecosystem to stratify. This stratification is a crucial f ...
groves at high altitude in arid regions of the
Western United States The Western United States (also called the American West, the Far West, and the West) is the region comprising the westernmost states of the United States. As American settlement in the U.S. expanded westward, the meaning of the term ''the We ...
. Bristlecones, along with all related species in class Pinopsida, are cone-bearing
seed plant A spermatophyte (; ), also known as phanerogam (taxon Phanerogamae) or phaenogam (taxon Phaenogamae), is any plant that produces seeds, hence the alternative name seed plant. Spermatophytes are a subset of the embryophytes or land plants. They inc ...
s commonly known as conifers; the name comes from the prickles on the female cones.


Species and range

There are three closely related species of bristlecone pines: *
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 ...
(''Pinus longaeva'') in
Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to it ...
,
Nevada Nevada ( ; ) is a state in the Western region of the United States. It is bordered by Oregon to the northwest, Idaho to the northeast, California to the west, Arizona to the southeast, and Utah to the east. Nevada is the 7th-most extensive, ...
and eastern
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the m ...
. The famous longest-lived species; often the term ''bristlecone pine'' refers to this tree in particular. * Rocky Mountain bristlecone pine (''Pinus aristata'') in
Colorado Colorado (, other variants) is a state in the Mountain states, Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the wes ...
,
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ke ...
and
Arizona Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a state in the Southwestern United States. It is the 6th largest and the 14th most populous of the 50 states. Its capital and largest city is Phoenix. Arizona is part of the Fou ...
. The most populous species; capable of forming closed canopies and, unlike the other two, is commonly cultivated. *
Foxtail pine ''Pinus balfouriana'', the foxtail pine, is a rare high-elevation pine that is endemic to California, United States. It is closely related to the Great Basin and Rocky Mountain bristlecone pines, in the subsection ''Balfourianae''. Descriptio ...
(''Pinus balfouriana'') with two disjunct populations found in the
Klamath Mountains The Klamath Mountains are a rugged and lightly populated mountain range in northwestern California and southwestern Oregon in the western United States. As a mountain system within both the greater Pacific Coast Ranges and the California Coast ...
(subspecies ''balfouriana'') and the southern Sierra Nevada (subspecies ''austrina''). A small outlying population was reported in southern Oregon, but was proven to have been misidentified. Forms the thickest groves of the three. At least some of the three species can hybridize in cultivation, but the ranges of wild populations do not overlap. The
Colorado River The Colorado River ( es, Río Colorado) is one of the principal rivers (along with the Rio Grande) in the Southwestern United States and northern Mexico. The river drains an expansive, arid watershed that encompasses parts of seven U.S. s ...
and
Green River Green River may refer to: Rivers Canada * Green River (British Columbia), a tributary of the Lillooet River *Green River, a tributary of the Saint John River, also known by its French name of Rivière Verte *Green River (Ontario), a tributary of ...
produce a gap between the ranges of ''P. longaeva'' and ''P. aristata'' and the northern
Owens Valley Owens Valley ( Numic: ''Payahǖǖnadǖ'', meaning "place of flowing water") is an arid valley of the Owens River in eastern California in the United States. It is located to the east of the Sierra Nevada, west of the White Mountains and Iny ...
provides a gap between the ranges of ''P. longaeva'' and ''P. balfouriana''.


Description

Bristlecone pines grow in isolated groves just below the
tree line The tree line is the edge of the habitat at which trees are capable of growing. It is found at high elevations and high latitudes. Beyond the tree line, trees cannot tolerate the environmental conditions (usually cold temperatures, extreme snow ...
, between elevation on dolomitic soils. The trees grow in soils that are shallow lithosols, usually derived from
dolomite Dolomite may refer to: *Dolomite (mineral), a carbonate mineral *Dolomite (rock), also known as dolostone, a sedimentary carbonate rock *Dolomite, Alabama, United States, an unincorporated community *Dolomite, California, United States, an unincor ...
and sometimes limestone, and occasionally sandstone or quartzite soils. Dolomitic soils are alkaline, high in
calcium Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar t ...
and
magnesium Magnesium is a chemical element with the symbol Mg and atomic number 12. It is a shiny gray metal having a low density, low melting point and high chemical reactivity. Like the other alkaline earth metals (group 2 of the periodic ta ...
, and low in
phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ear ...
. Those factors tend to exclude other plant species, allowing bristlecones to thrive. Because of cold temperatures, dry soils, high winds, and short growing seasons, the trees grow very slowly. Even the tree's needles, which grow in bunches of five, can remain on the tree for forty years, which gives the tree's terminal branches the unique appearance of a long bottle brush. The bristlecone pine's root system is mostly composed of highly branched, shallow roots, while a few large, branching roots provide structural support. The bristlecone pine is extremely drought tolerant due to its branched shallow root system, its waxy needles, and thick needle cuticles that aid in water retention. The wood is very dense and resinous, and thus resistant to invasion by insects, fungi, and other potential pests. The tree's longevity is due in part to the wood's extreme durability. While other species of trees that grow nearby suffer rot, bare bristlecone pines can endure, even after death, often still standing on their roots, for many centuries. Exposed wood on living and dead trees does not rot, but rather erodes like stone due to wind, rain, and freezing, which creates unusual forms and shapes. The bristlecone pine has an intrinsically low rate of reproduction and regeneration, and it is thought that under present climatic and environmental conditions the rate of regeneration may be insufficient to sustain its population. The species are on the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of nat ...
(IUCN) red list. The species are labeled under Least Concern (LC), the justification for this being that no subpopulations for Great Basin bristlecone pines are decreasing. Subpopulations seem to be increasing or remaining stable. Many bristlecone pine habitats have been protected, including the
Inyo National Forest Inyo National Forest is a United States National Forest covering parts of the eastern Sierra Nevada of California and the White Mountains of California and Nevada. The forest hosts several superlatives, including Mount Whitney, the highest po ...
's Ancient Bristlecone Pine Forest in the White Mountains of California and the
Great Basin National Park Great Basin National Park is an American national park located in White Pine County in east-central Nevada, near the Utah border, established in 1986. The park is most commonly entered by way of Nevada State Route 488, which is connected to U ...
in Nevada, where cutting or gathering wood is prohibited. The green pine needles give the twisted branches a bottle-brush appearance. The needles of the tree surround the branch to an extent of about one foot near the tip of the limb. The name bristlecone pine refers to the dark purple female cones that bear incurved prickles on their surface. The dark color of these cones helps to absorb heat. After maturity, which takes about two years, the cones will become brown in color. These ancient trees have a gnarled and stunted appearance, especially those found at high altitudes, and have reddish-brown bark with deep fissures. As the tree ages, much of its
vascular cambium The vascular cambium is the main growth tissue in the stems and roots of many plants, specifically in dicots such as buttercups and oak trees, gymnosperms such as pine trees, as well as in certain other vascular plants. It produces secondary xy ...
layer may die. In very old specimens, often only a narrow strip of living tissue connects the roots to a handful of live branches. Even though the trees' needles may age, they still remain functional in regulating water and by their ability to photosynthesize.


Oldest living organisms

Bristlecone pines are known for attaining great ages. The oldest bristlecone pine in the White Mountains is
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 f ...
, which has a verified age of years. It is located in the
Inyo National Forest Inyo National Forest is a United States National Forest covering parts of the eastern Sierra Nevada of California and the White Mountains of California and Nevada. The forest hosts several superlatives, including Mount Whitney, the highest po ...
in Eastern California. However, the specific location of Methuselah is a closely guarded secret. The other two species, ''Pinus balfouriana'' and ''Pinus aristata'', are also long-lived, though not to the extreme extent of ''P. longaeva''; specimens of both have been measured or estimated to be up to 3,000 years old. The longevity of the trees is believed to be related to the proportion of dead wood to live wood. This high ratio reduces respiration and water loss, thereby extending the life of the tree. Trees that reproduce by
cloning Cloning is the process of producing individual organisms with identical or virtually identical DNA, either by natural or artificial means. In nature, some organisms produce clones through asexual reproduction. In the field of biotechnology, c ...
can be considered to be much older than bristlecone pines. A colony of 47,000
quaking aspen ''Populus tremuloides'' is a deciduous tree native to cooler areas of North America, one of several species referred to by the common name aspen. It is commonly called quaking aspen, trembling aspen, American aspen, mountain or golden aspen, tr ...
trees (nicknamed " Pando"), covering in the
Fishlake National Forest Fishlake National Forest is a U.S. National Forest located in south central Utah. The namesake for the forest is Fish Lake, the largest freshwater mountain lake in the state. Wildlife Animals that inhabit this forest are elk, shrews, deer, b ...
in
Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to it ...
, United States, has been estimated to be 80,000 years old, although
tree ring Dendrochronology (or tree-ring dating) is the scientific method of dating tree rings (also called growth rings) to the exact year they were formed. As well as dating them, this can give data for dendroclimatology, the study of climate and atmos ...
samples date individual, above-ground trees at an average of about 130 years. Bristlecone pines are invaluable to dendroclimatologists, because they provide the longest continual climatically sensitive tree-ring chronologies on Earth. By cross-dating millennia-old bristlecone pine debris, some chronologies reach beyond 9,000 years before present. In addition, ratios of stable carbon isotopes from bristlecone pine tree rings are sensitive to past variations in moisture availability. This information can be used to reconstruct precipitation changes in the past. The Rocky Mountain population is severely threatened by an introduced fungal disease known as
white pine blister rust ''Cronartium ribicola'' is a species of rust fungus in the family Cronartiaceae that causes the disease white pine blister rust. Other names include: (French), (German), (Spanish). ''Cronartium ribicola'' is native to China, and was subseque ...
, and by
mountain pine beetle The mountain pine beetle (''Dendroctonus ponderosae'') is a species of bark beetle native to the forests of western North America from Mexico to central British Columbia. It has a hard black exoskeleton, and measures approximately , about the siz ...
s. Climate change may also affect the species as temperatures increased over a 30-year period throughout the southern Rocky Mountain range. These changes in climate would mostly affect trees in higher elevations. With these problems, the genetic diversity within the species has become a concern; old specimens of bristlecone pine, however, have survived previous warmer periods.


See also

*
List of longest-living organisms This is a list of the longest-living biological organisms: the individual(s) (or in some instances, clones) of a species with the longest natural maximum lifespans. For a given species, such a designation may include: # The oldest known individ ...


References


Citations


General sources

* *


External links

*
Gymnosperm Database

''Pinus longaeva''''Pinus aristata''''Pinus balfouriana''


{{Taxonbar, from=Q515556
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. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden ...
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. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden ...
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. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden ...
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. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden ...
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. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden ...
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. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden ...
Balfourianae Pinus taxa by common names Plant subsections Pine, Bristlecone Flora without expected TNC conservation status