''Pinus lambertiana'' (commonly known as the sugar pine or sugar cone pine) is the tallest and most massive

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. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as cu ...

tree and has the longest

cones In geometry, a cone is a three-dimensional figure that tapers smoothly from a flat base (typically a circle) to a point not contained in the base, called the ''apex'' or '' vertex''. A cone is formed by a set of line segments, half-lines, ...

of any

conifer Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...

. It is native to coastal and inland mountain areas along the

Pacific coast Pacific coast may be used to reference any coastline that borders the Pacific Ocean. Geography Americas North America Countries on the western side of North America have a Pacific coast as their western or south-western border. One of th ...

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

, as far north as

Oregon Oregon ( , ) is a U.S. state, state in the Pacific Northwest region of the United States. It is a part of the Western U.S., with the Columbia River delineating much of Oregon's northern boundary with Washington (state), Washington, while t ...

and as far south as

Baja California Baja California, officially the Free and Sovereign State of Baja California, is a state in Mexico. It is the northwesternmost of the 32 federal entities of Mexico. Before becoming a state in 1952, the area was known as the North Territory of B ...

in Mexico.

Description

Growth

The sugar pine is the tallest and largest ''Pinus'' species, commonly growing to tall, exceptionally to tall, with a trunk diameter of , exceptionally . The tallest recorded specimen is tall, is located in

Yosemite National Park Yosemite National Park ( ) is a List of national parks of the United States, national park of the United States in California. It is bordered on the southeast by Sierra National Forest and on the northwest by Stanislaus National Forest. The p ...

, and was discovered in 2015. The second tallest recorded was "Yosemite Giant", an tall specimen in Yosemite National Park, which died from a

bark beetle A bark beetle is the common name for the subfamily of beetles Scolytinae. Previously, this was considered a distinct family (Scolytidae), but is now understood to be a specialized clade of the "true weevil" family (Curculionidae). Although th ...

attack in 2007. Yosemite National Park also has the third tallest, measured to tall as of June 2013; the

Rim Fire Rim may refer to: *Rim (basketball), the hoop through which the ball must pass **Breakaway rim, a sprung basketball rim *Rim (coin), the raised edge which surrounds the coin design *Rim (crater), extending above the local surface *Rim (firearms), ...

affected this specimen, but it survived. The next tallest known living specimens grow in southern

: one in Umpqua National Forest is tall and another in Siskiyou National Forest is tall. Pinus lambertiana ProspectOR

The bark of ''Pinus lambertiana'' ranges from brown to purple in color and is thick. The upper branches can reach out over . Like all members of the white pine group (''Pinus'' subgenus ''Strobus''), the

leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...

("needles") grow in fascicles ("bundles") of five with a

deciduous In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...

sheath. They are long. Sugar pine is notable for having the longest

of any conifer, mostly long, exceptionally to long (although the cones of the Coulter pine are more massive); their unripe weight of makes them perilous projectiles when chewed off by squirrels. The

seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...

s are long, with a long wing that aids their dispersal by wind. Sugar pine never grows in pure stands, always in a mixed forest, and is

shade tolerant In ecology, shade tolerance is a plant's ability to tolerate low light levels. The term is also used in horticulture and landscaping, although in this context its use is sometimes imprecise, especially in labeling of plants for sale in nursery (ho ...

in its youth.

Distribution

The sugar pine occurs in Oregon and

California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...

in the

Western United States The Western United States (also called the American West, the Western States, the Far West, the Western territories, and the West) is List of regions of the United States, census regions United States Census Bureau. As American settlement i ...

southward to

, specifically in the

Cascade Range The Cascade Range or Cascades is a major mountain range of western North America, extending from southern British Columbia through Washington (state), Washington and Oregon to Northern California. It includes both non-volcanic mountains, such as m ...

Sierra Nevada The Sierra Nevada ( ) is a mountain range in the Western United States, between the Central Valley of California and the Great Basin. The vast majority of the range lies in the state of California, although the Carson Range spur lies primari ...

, Coast Ranges, and Sierra San Pedro Martir. It is generally more abundant towards the south and can be found between above sea level.

Genome

The massive 31 gigabase mega-genome of sugar pine has been sequenced in 2016 by the large PineRefSeq consortium. This makes the genome one of the largest sequenced and assembled so far. The transposable elements that make up the megagenome are linked to the evolutionary change of the sugar pine. The sugar pine contains extended regions of

non-coding DNA Non-coding DNA (ncDNA) sequences are components of an organism's DNA that do not encode protein sequences. Some non-coding DNA is transcribed into functional non-coding RNA molecules (e.g. transfer RNA, microRNA, piRNA, ribosomal RNA, and reg ...

, most of which is derived from

transposable element A transposable element (TE), also transposon, or jumping gene, is a type of mobile genetic element, a nucleic acid sequence in DNA that can change its position within a genome. The discovery of mobile genetic elements earned Barbara McClinto ...

s. 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 in

angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit ...

Embryonal growth

In late stage of

embryo An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...

nal development, the sugar pine embryo changes from a smooth and narrow

paraboloid In geometry, a paraboloid is a quadric surface that has exactly one axial symmetry, axis of symmetry and no central symmetry, center of symmetry. The term "paraboloid" is derived from parabola, which refers to a conic section that has a similar p ...

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 the

epicotyl An epicotyl is important for the beginning stages of a plant's life. It is the region of a seedling stem above the stalks of the seed leaves of an embryo plant. It grows rapidly, showing hypogeal germination, and extends the stem above the soil su ...

develops as an anlage flanked by regions of that define the cotyledonary buttresses. At this stage, the embryo is composed of the suspensor, root initials and

root cap The root cap is a type of tissue at the tip of a plant root. It is also called calyptra. Root caps contain statocytes which are involved in gravity perception in plants. If the cap is carefully removed the root will grow randomly. The root cap ...

region,

hypocotyl The hypocotyl (short for "hypocotyledonous stem", meaning "below seed leaf") is the stem of a germinating seedling, found below the cotyledons (seed leaves) and above the radicle (root). Eudicots As the plant embryo grows at germination, it send ...

-shoot axis, and the

. The upper (distal) portion of the embryo, which gives rise to the

cotyledon A cotyledon ( ; ; "a cavity, small cup, any cup-shaped hollow", gen. (), ) is a "seed leaf" – a significant part of the embryo within the seed of a plant – and is formally defined as "the embryonic leaf in seed-bearing plants, one or mor ...

s and the epicotyl, is considered to be the

shoot apex In cell biology, the meristem is a structure composed of specialized tissue found in plants, consisting of stem cells, known as meristematic cells, which are undifferentiated cells capable of continuous cellular division. These meristematic ce ...

Shoot apex

The

has the following four zones: #The apical initials produce all cells of the shoot apex through cell division. It is located at the top of the

meristem In cell biology, the meristem is a structure composed of specialized tissue found in plants, consisting of stem cells, known as meristematic cells, which are undifferentiated cells capable of continuous cellular division. These meristematic c ...

and the cells are larger in size compared to other cells on the surface layer. #The central mother cell generates the rib meristem and the inner layers of the peripheral tissue zone through cell division. It presents a typical

gymnosperm The gymnosperms ( ; ) are a group of woody, perennial Seed plant, seed-producing plants, typically lacking the protective outer covering which surrounds the seeds in flowering plants, that include Pinophyta, conifers, cycads, Ginkgo, and gnetoph ...

appearance and is characterized by cell expansion and unusual

mitosis Mitosis () is a part of the cell cycle in eukaryote, eukaryotic cells in which replicated chromosomes are separated into two new Cell nucleus, nuclei. Cell division by mitosis is an equational division which gives rise to genetically identic ...

that occurs in the central region. The rate of mitosis increases on its outer edge. #The peripheral tissue zone consists of two layers of cells that are characterized by dense cytoplasm and mitosis of high frequency. #Lastly, the rib meristem is a regular arrangement of vertical files of cells which mature into the pith of the axis.

Etymology

Naturalist

John Muir John Muir ( ; April 21, 1838December 24, 1914), also known as "John of the Mountains" and "Father of the national park, National Parks", was a Scottish-born American naturalist, author, environmental philosopher, botanist, zoologist, glaciologi ...

considered sugar pine to be the "king of the conifers". The common name comes from the sweet resin, which Native Americans used as a sweetener. John Muir found it preferable to

maple sugar Maple sugar is a traditional sweetener in Canada and the Northeastern United States, prepared from the sap of the maple tree ("maple syrup, maple sap"). Sources Three species of maple trees in the genus ''Acer (plant), Acer'' are predomina ...

. It is also known as the great sugar pine. The scientific name was assigned by David Douglas in 1826, in honor of his friend, a botanist friend from London:

Aylmer Bourke Lambert Aylmer Bourke Lambert (2 February 1761 – 10 January 1842) was a British botanist, one of the first fellows of the Linnean Society. Early life Aylmer Bourke Lambert was born at Bath, England, Bath, England on 2 February 1761, the son of E ...

Ecology

Wildlife

The large size and high nutritional value of the sugar pine seeds are appealing to many species. Yellow pine chipmunks (''

Neotamias amoenus The yellow-pine chipmunk (''Neotamias amoenus'') is a species of order Rodentia in the family Sciuridae. It is found in parts of Canada and the United States. These chipmunks are normally found in brush-covered areas, and in California, they in ...

'') 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. Black bears (''

Ursus americanus The American black bear (''Ursus americanus''), or simply black bear, is a species of medium-sized bear which is endemic to North America. It is the continent's smallest and most widely distributed bear species. It is an omnivore, with a diet v ...

'') 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.

Threats

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 and

white pine blister rust ''Cronartium ribicola'' is a species of rust (fungus), rust fungus in the family Cronartiaceae that causes the disease white pine blister rust. Other names include: (French), (German), (Spanish). Origin ''Cronartium ribicola'' is native to C ...

. Blister rust can weaken the tree and enable further infestation by mountain pine beetles. The sugar pine has been severely affected by the white pine blister rust (''

Cronartium ribicola ''Cronartium ribicola'' is a species of rust (fungus), rust fungus in the family Cronartiaceae that causes the disease white pine blister rust. Other names include: (French), (German), (Spanish). Origin ''Cronartium ribicola'' is native to C ...

''), a

fungal pathogen Pathogenic fungi are fungi that cause disease in humans or other organisms. Although fungi are eukaryotic, many pathogenic fungi are microorganisms. Approximately 300 fungi are known to be pathogenic to humans; their study is called "medical mycolo ...

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 and

whitebark pine ''Pinus albicaulis'', known by the common names whitebark pine, white bark pine, white pine, pitch pine, scrub pine, and creeping pine, is a conifer tree native to the mountains of the western United States and Canada, specifically subalpine ...

throughout their ranges. The

U.S. Forest Service The United States Forest Service (USFS) is an agency within the U.S. Department of Agriculture. It administers the nation's 154 national forests and 20 national grasslands covering of land. The major divisions of the agency are the Chief's ...

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 the

Lake Tahoe Lake Tahoe (; Washo language, Washo: ''dáʔaw'') is a Fresh water, freshwater lake in the Sierra Nevada of the Western United States, straddling the border between California and Nevada. Lying at above sea level, Lake Tahoe is the largest a ...

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. The species is generally resistant to fire because of its thick bark and because it clears away competing species. However, its mortality has been directly linked to drier conditions and higher temperatures. Climate change presents a threat to species health: higher temperatures can decrease

resin A resin is a solid or highly viscous liquid that can be converted into a polymer. Resins may be biological or synthetic in origin, but are typically harvested from plants. Resins are mixtures of organic compounds, predominantly terpenes. Commo ...

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.

Protective efforts

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. They plant seedlings grown from seeds collected from tree strains resistant to blister rust. The foundation's aim is to build a wild a sugar pine population that is resistant to white pine rust.

Uses

According to David Douglas, who was guided to the (exceptionally thick) tree specimen he was looking for by a Native American, some tribes ate the sweetish seeds. These were eaten raw and roasted, and also used to make flour or pulverized into a spread. Native Americans also ate the inner bark. The sweet sap or pitch was consumed, in small quantities due to its laxative properties, but could also be chewed as gum. Its flavor is thought largely to be derived from the pinitol it contains. In the mid-19th century, the trees were used liberally as lumber during the

California Gold Rush The California gold rush (1848–1855) began on January 24, 1848, when gold was found by James W. Marshall at Sutter's Mill in Coloma, California. The news of gold brought approximately 300,000 people to California from the rest of the U ...

. In modern times they are used in much lower quantities, being spared for high-end products as with Western white pine. The odorless wood is also preferred for packing fruit, as well as storing drugs and other goods. Its straight grain also makes it a useful

organ pipe An organ pipe is a sound-producing element of the pipe organ that resonator, resonates at a specific Pitch (music), pitch when pressurized air (commonly referred to as ''wind'') is driven through it. Each pipe is tuned to a note of the musical ...

material. 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."Buying Lumber for Keys." Music Trade Review 47:10 (5 September 1908), 35.

Folklore

In the Achomawi

creation myth A creation myth or cosmogonic myth is a type of cosmogony, a symbolic narrative of how the world began and how people first came to inhabit it., "Creation myths are symbolic stories describing how the universe and its inhabitants came to be. Cre ...

, 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 this

ancestry An ancestor, also known as a forefather, fore-elder, or a forebear, is a parent or ( recursively) the parent of an antecedent (i.e., a grandparent, great-grandparent, great-great-grandparent and so forth). ''Ancestor'' is "any person from ...

is Sugarpine-Cone man, who has a handsome son named Ahsoballache. After Ahsoballache marries the daughter of To'kis the

Chipmunk Chipmunks are small, striped rodents of subtribe Tamiina. Chipmunks are found in North America, with the exception of the Siberian chipmunk which is found primarily in Asia. Taxonomy and systematics Chipmunks are classified as four genera: '' ...

-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 infant Edechewe near their bed. The

Washo language Washo (or Washoe; endonym ) is an endangered Native American language isolate spoken by the Washo on the California–Nevada border in the drainages of the Truckee and Carson Rivers, especially around Lake Tahoe. While there were only 20 el ...

has a word for sugar pine, , and also a word for "sugar pine sugar", .

References

External links

U.C. Jepson Manual treatment for ''Pinus lambertiana''

US Forest Service – Dorena Genetic Resource Center
– USFS rust resistance program
The Sugar Pine Foundation
– ''The Sugar Pine and Western White Pine Restoration Program'' * *
Arboretum de Villardebelle: photo of a cone
{{Authority control lambertiana Trees of Northern America Flora of California Flora of the Sierra Nevada (United States) Natural history of the California Coast Ranges Natural history of the Transverse Ranges Edible nuts and seeds Plants used in traditional Native American medicine Least concern flora of the United States