| Sugar pine | |
|---|---|
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Scientific classification | |
| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Gymnospermae |
| Division: | Pinophyta |
| Class: | Pinopsida |
| Order: | Pinales |
| Family: | Pinaceae |
| Genus: | Pinus |
| Subgenus: | P. subg.Strobus |
| Section: | P.sect. Quinquefoliae |
| Subsection: | P.subsect. Strobus |
| Species: | P. lambertiana |
| Binomial name | |
| Pinus lambertiana | |
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| Natural range ofPinus lambertiana | |
Pinus lambertiana (commonly known as thesugar pine) is the tallest and most massivepine tree and has among the longestcones of anyconifer. It is native to coastal and inland mountain areas along thePacific coast ofNorth America, as far north asOregon and as far south asBaja California in Mexico.
The sugar pine is the tallest and largestPinus species, commonly growing to 40–60 meters (130–195 ft) tall, exceptionally to 82 m (269 ft) tall, with a trunk diameter of 1.2–2.5 m (3 ft 11 in – 8 ft 2 in), exceptionally 3.5 m (11 ft 6 in).[2] The tallest recorded specimen is 83.45 m (273 ft 9 in) tall, is located inYosemite National Park, and was discovered in 2015.[3] The second tallest recorded was "Yosemite Giant", an 82.05 m (269 ft 2 in) tall specimen in Yosemite National Park, which died from abark beetle attack in 2007. Yosemite National Park also has the third tallest, measured to 80.5 m (264 ft 1 in) tall as of June 2013; theRim Fire affected this specimen, but it survived. The next tallest known living specimens grow in southernOregon; one in Umpqua National Forest is 77.7 m (254 ft 11 in) tall and another in Siskiyou National Forest is 77.2 m (253 ft 3 in) tall.
The bark ofPinus lambertiana ranges from brown to purple in color and is 5–10 centimeters (2–4 in) thick.[2] The upper branches can reach out over 8 m (26 ft).[2] Like all members of thewhite pine group (Pinus subgenusStrobus), theleaves ("needles") grow infascicles ("bundles") of five[2] with adeciduous sheath. They are 5–11 cm (2–4+1⁄4 in) long.[4] Sugar pine is notable for having among the longestcones of any conifer, mostly 10–50 cm (4–20 in) long,[2][5] although they can occasionally reach as much as 56 cm (22 in).[6][7] Although the cones of theCoulter pine are more massive; the unripe weight of 1–2 kilograms (2.2–4.4 lb) makes the sugar pine's cones perilous projectiles when chewed off by squirrels.[2] Theseeds are1–2 cm (1⁄2–3⁄4 in) long, with a2–3-centimeter (3⁄4–1+1⁄4-inch) long wing[5] that aids their dispersal by wind. Sugar pine never grows in pure stands, always in a mixed forest, and isshade tolerant in its youth.[8]
The sugar pine occurs in Oregon andCalifornia in theWestern United States southward toBaja California, specifically in theCascade Range,Sierra Nevada,Coast Ranges, andSierra San Pedro Martir. It is generally more abundant towards the south and can be found between 500 and 1,500 m (1,600 and 4,900 ft) above sea level.[2]
The massive 31 gigabase mega-genome of sugar pine has been sequenced in 2016 by the large PineRefSeq consortium.[9] This makes the genome one of the largest sequenced and assembled so far.[9]
The transposable elements that make up the megagenome are linked to the evolutionary change of the sugar pine. The sugar pine contains extended regions ofnon-coding DNA, most of which is derived fromtransposable elements. The genome of the sugar pine represents one extreme in all plants, with a stable diploid genome that is expanded by the proliferation of transposable elements, in contrast to the frequent polyploidization events inangiosperms.[10]
In late stage ofembryonal development, the sugar pine embryo changes from a smooth and narrowparaboloid to a less symmetric structure. This configuration is caused by a transverse orientation of division planes in the upper portion of the embryo axis. The root initial zone is established, and theepicotyl develops as an anlage flanked by regions of that define the cotyledonary buttresses. At this stage, the embryo is composed of thesuspensor, root initials androot cap region,hypocotyl-shoot axis, and theepicotyl. The upper (distal) portion of the embryo, which gives rise to thecotyledons and the epicotyl, is considered to be theshoot apex.[11]
Theshoot apex has the following four zones:[12]
NaturalistJohn Muir considered sugar pine to be the "king of the conifers". The common name comes from the sweet resin, whichNative Americans used as a sweetener.[13] John Muir found it preferable tomaple sugar.[14] It is also known as the great sugar pine. The scientific name was assigned byDavid Douglas in 1826,[2] in honor of the London botanistAylmer Bourke Lambert.[15]
The large size and high nutritional value of the sugar pine seeds are appealing to many species. Yellow pine chipmunks (Neotamias amoenus) and Steller's jays (Cyanocitta stelleri) gather and hoard sugar pine seeds. Chipmunks gather wind-dispersed seeds from the ground and store them in large amounts. Jays collect seeds by pecking the cones with their beaks and catching the seeds as they fall out. Although wind is a main dispersal agent of sugar pine seeds, animals tend to collect and store them before the wind can blow them far.[16]
Black bears (Ursus americanus) feed on sugar pine seeds in the fall months within the Sierra Nevada. Both sugar pine and oak species are currently in decline, directly affecting black bear food sources within the Sierra Nevada.[17]
The sugar pine has been severely affected by the white pine blister rust (Cronartium ribicola),[18] afungal pathogen accidentally introduced from Europe in 1909. A high proportion of sugar pines have been killed by the blister rust, particularly in the northern part of the species' range where blister rust has been present longer. The rust has also destroyed much of the Western white pine andwhitebark pine throughout their ranges.[19] TheU.S. Forest Service has a program for developing rust-resistant varieties of sugar pine and western white pine. Seedlings of these trees have been introduced into the wild. The Sugar Pine Foundation in theLake Tahoe Basin has been successful in finding resistant sugar pine seed trees. Blister rust is much less common in California, where sugar, Western white and whitebark pines still survive in great numbers.[20]
Sugar pine trees have been impacted by the mountain pine beetle (Dendroctonus ponderosae), which is native to western North America. The beetles lay their eggs inside of the tree and inhibit the tree's ability to defend itself against invading species. Beetle infestation can also cause nutrient deficiencies that slowly weaken the tree's overall health, making pines more susceptible to other threats such as fires andwhite pine blister rust.[21] Blister rust can weaken the tree and enable further infestation by mountain pine beetles.[22]
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The species is generallyresistant to fire because of its thick bark and because it clears away competing species.[2] However, its mortality has been directly linked to drier conditions and higher temperatures. Climate change presents a threat to species health: higher temperatures can decreaseresin levels within the trees, weakening defenses against pathogens. At the same time, warmer winters increase survival of pests and pathogens. The weakened or dying trees then provide fuel for forest fires, which may become more frequent and more intense with rising summer temperatures, particularly if coupled with drier conditions and stronger winds.[23]
Sugar pine trees are in slow decline due to several threats; white pine blister rust, mountain pine beetles, and climate change. Efforts to restore sugar pines and other white pine trees that have been impacted by invasive species, climate change, and fires have been undertaken by governmental and non-governmental entities. One nonprofit, the Sugar Pine Foundation, was created in 2004 to plant sugar pine seeds in the Sierra Nevada along the border of California and Nevada.[20] They plant seedlings grown from seeds collected from tree strains resistant to blister rust. The foundation's aim is to build in the a wild a sugar pine population that is resistant to white pine rust.[24]
According to David Douglas, who was guided to the (exceptionally thick) tree specimen he was looking for by a Native American,[2] some tribes ate the sweetish seeds. These were eaten raw and roasted, and also used to make flour or pulverized into a spread.[2] Native Americans also ate the inner bark.[2] The sweet sap orpitch was consumed, in small quantities due to its laxative properties,[25] but could also be chewed as gum.[2] Its flavor is thought largely to be derived from thepinitol it contains.[2]
In the mid-19th century, the trees were used liberally as lumber during theCalifornia Gold Rush. In modern times they are used in much lower quantities, being spared for high-end products as withWestern white pine.[2]
The odorless wood is also preferred for packing fruit, as well as storing drugs and other goods. Its straight grain also makes it a usefulorgan pipe material.[25] The wood was also long used for piano keys; in 1907 or 1908 the Connection piano-action maker Pratt, Read & Co. purchased "950,000 feet of clear sugar pine" for that use in & around Placerville, CA.[26]
In theAchomawicreation myth, Annikadel, the creator, makes one of the 'First People' by intentionally dropping a sugar pine seed in a place where it can grow. One of the descendants in thisancestry is Sugarpine-Cone man, who has a handsome son named Ahsoballache.[27]
After Ahsoballache marries the daughter of To'kis theChipmunk-woman, his grandfather insists that the new couple have a child. To this end, the grandfather breaks open a scale from a sugar pine cone, and secretly instructs Ahsoballache to immerse the scale's contents in spring water, then hide them inside a covered basket. Ahsoballache performs the tasks that night; at the next dawn, he and his wife discover the infantEdechewe near their bed.[27]
TheWasho language has a word for sugar pine,simt'á:gɨm, and also a word for "sugar pine sugar",nanómba.
