Pinus contorta

Pinus contorta, with the common names lodgepole pine, shore pine, twisted pine,<ref name="GRIN">Template:GRIN</ref> and contorta pine,<ref name=GRIN /> is an evergreen species of conifer tree. It is a common pine in western North America, found near the ocean shore and in dry montane forests to the subalpine, but is rare in lowland rain forests.

Description

Depending on subspecies, Pinus contorta grows as an evergreen shrub or tree. The shrub form is krummholz and is approximately Template:Convert high. The thin and narrow-crowned tree can grow Template:Convert high and achieve up to Template:Convert in diameter at chest height.<ref name="FNA">Template:EFloras</ref> The murrayana subspecies is the tallest. The crown is rounded and the top of the tree is flattened. In dense forests, the tree has a slim, conical crown. The formation of twin trees is common in some populations in British Columbia. The elastic branches stand upright or overhang and are difficult to break. The branches are covered with short shoots that are easy to remove.<ref name="WTU Herbarium2015">Template:Cite web</ref><ref name=Schutt>Template:Cite book</ref><ref name="Klinkenberg2014">Template:Cite web</ref>

The species name is contorta because of the twisted, bent pines (shore pine)<ref name=":0" /> found at coastal areas and the tree's twisted needles.<ref name=handbook /><ref name=boise>Template:Cite web</ref><ref>Template:Cite web</ref> Pinus contorta is occasionally known under several English names: black pine, scrub pine, and coast pine.<ref name=craterlake>Template:Cite web</ref><ref name="FEIS var contorta" /> P. contorta subsp. latifolia will hybridise with the closely related jack pine (P. banksiana).

The bark of lodgepole pine is thin, scaly and grayish brown.<ref name=":0" /> Shore pine bark is somewhat thick and corky, fissuring into a checkered pattern.<ref name=":0" /> Some lodgepole pines have been reported in low elevations with features closer to those of the shore pine, including the bark.<ref name=":0" />

Tamarack pine can grow up to centuries old and lodgepole pines in Yellowstone Park have survived over 300 years.<ref name=":0" />

File:Lodgepole Pine as Bonsai.jpg

Lodgepole pine being trained as bonsai. Notice the use of wire to position the branches of the tree. This is a yamadori (wild collected specimen) and has been styled by American bonsai artist Bjorn Bjorholm.

Foliage

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The needles are Template:Convert long in fascicles of two, alternate on twigs. The female cones are Template:Convert long with sharp-tipped scales.

The egg-shaped growth buds are reddish-brown and between Template:Convert long. They are short pointed, slightly rotated, and very resinous. Spring growth starts in beginning of April and the annual growth is completed by early July. The dark and mostly shiny needles are pointed and Template:Convert long and Template:Convert wide. The needle edge is weakly to strongly serrated. The needles are in pairs on short shoots and rotated about the shoots' longitudinal axes. In Alberta above Template:Convert, 1 to 5 needles occur per short shoot. A population with a high proportion of three-needled short shoots occurs in the Yukon. Needles live an average of four to six years, with a maximum of 13 years.<ref name=Schutt /> The foliage of lodgepole pine is yellow-green as compared to shore pine, which is dark green.<ref name=":0" />

Cones

The cones of lodgepole and shore pine begin to be produced when the trees are about ten years old.<ref name=":0" /> The cones are Template:Convert long, with prickles on the scales.<ref name=":0" />

Many populations of the Rocky Mountain subspecies, P. contorta subsp. latifolia, have serotinous cones. This means that the cones are closed and must be exposed to high temperatures, such as from forest fires, in order to open and release their seeds.<ref>Template:FEIS</ref> The variation in their serotiny has been correlated with wildfires and mountain pine beetle attacks.<ref name=feduck>Template:Cite bioRxiv</ref> The cones of the coastal Pacific subspecies, P. contorta subsp. contorta, are typically non-serotinous,<ref name="FEIS var contorta">Template:FEIS</ref> and those of the inland Pacific subspecies, P. contorta subsp. murrayana, are completely non-serotinous.<ref>Template:FEIS</ref> Sometimes cones will become buried by the continued growth of a branch. If the seeds are collected, they have germinated as much as 150 years after confinement.<ref>Template:Cite book</ref>

Distribution

Pinus contorta occurs from upper, dry montane forests to the subalpine region of western North America.<ref name="Moore2008">Template:Cite book</ref><ref name="Sullivan2015">Template:Cite web</ref><ref name="WTU Herbarium2015" /> It can be found on the western side of the Cascades, in inland British Columbia, and on the Rocky Mountains in Alberta, except where it is too high and dry.<ref name=":0" /> Lodgepole pine can tolerate relatively hostile environments such as high-elevation volcanic rock in Central Oregon (e.g. Crater Lake) and thin soils on the eastern slope of the Cascades.<ref name=":0" /> Further south, the species can be found in higher elevations up to Template:Convert above sea level, particularly in southern Colorado.<ref name=":0" /> It is rare in lowland rain forests.<ref name="Sullivan2015" /><ref name="WTU Herbarium2015" /> Shore pine can be found in very infertile soils in coastal regions from Southeast Alaska to Northern California.<ref name=":0" />

Lodgepole and shore pine can be found intermingled (and apparently hybridized) north of Puget Sound.<ref name=":0" /> Less dependent on fire, tamarack pine can be found in California's upper mountains and mingled with lodgepole in Oregon.<ref name=":0" /> Pinus contorta can be found in the closed-cone pine forest of coastal California.

Ecology

Pinus contorta is a fire-adapted species, often regenerating densely following wildfire. The species displays characteristically poor self pruning and thin bark, leading to the death of the tree following fire. Importantly however, the heat from fire results in the opening of their serotinous cones and release of seeds. This allows the species to regenerate well after fire and maintain its place in the forest habitat as an early successional species.<ref name="Schoennagel" /> Some populations demonstrate more significant reliance on other modes of regeneration rather than fire, with wind through, ice storms, and land slides also acting as stand replacing disturbances.<ref name=":0">Template:Cite book</ref> Template:Multiple image

The natural fire regime for this species is primarily driven by climate. The fires occur most often after years of drought. Forests in the upper montane to subalpine region experience much moisture in the winter via snow. The density of tree stands with the species inhibit the establishment of an understory (allowing ladder fuel to form), and surface fire is rare regardless. Thus, infrequent but severe fires dominate this species.<ref name="Schoennagel">Template:Cite journal</ref>

An example of the climate that plays a huge role in the fire regime of the species is quite complex. There are three different oscillations that play a major role in droughts. These are the Pacific decadal oscillation (PDO), Atlantic multidecadal oscillation (AMO) and El Niño (ENSO). A combination of these oscillations being in effect (+) or not in effect (−) have a global effect on the water available to these forests. The combination of AMO +, ENSO − and PDO − means there is going to be a drought and likely a severe subalpine fire.<ref>Template:Cite journal</ref>

File:Male cone of lodgepole pine.jpg

A cluster of pollen-bearing male cones at Mount San Antonio

Suillus tomentosus, a fungus, produces specialized structures called tuberculate ectomycorrhizae with the roots of lodgepole pine (Pinus contorta var. latifolia). These structures have been shown to be the location of concentrations of nitrogen-fixing bacteria which contribute a significant amount of nitrogen to tree growth and allow the pines to colonize nutrient-poor sites.<ref name=paul07>Template:Cite journal</ref><ref name=chapman12>Template:Cite journal</ref>

Porcupines consume the inner bark of lodgepole pine.<ref name=":0" />

Threats

Larger members of the species are attacked by mountain pine beetles, which it fights with pitch but can be overwhelmed.<ref name=":0" /> It is also affected by blue stain fungus (Grosmannia clavigera), which the mountain pine beetle carries in its mouth. Dwarf mistletoe also leeches off the species. Both the threat of pine beetles and dwarf mistletoe are curbed by wildfires, which occurred less in the 20th century due to firefighting. More recently, unthreatening lightning-sparked fires have been allowed to burn in wilderness areas in Idaho and Montana.<ref name=":0" /> Exceptional cold can kill some of the beetles.<ref name=":0" />

A study released in 2011 concluded that Pinus contorta could experience significant reductions in distribution due to climate change by the late 21st century.<ref>Template:Cite journal</ref><ref>Template:Cite web</ref>

As an invasive species

Pinus contorta is considered a serious invasive species of wilding conifer in New Zealand, along with several other western North American pine species. It is listed on the National Pest Plant Accord and is prohibited from sale, commercial propagation, and distribution.

Subspecies

There are four subspecies of Pinus contorta, and one of them is sometimes considered to have two varieties.<ref name="iucn-bolanderi">Template:Cite iucn</ref> The subspecies are sometimes treated at the rank of variety.<ref name=GRIN /><ref name="FNA" /><ref>Template:ThePlantList</ref>

Cone	Image	Subspecies	Common name	Distribution
File:Pinus contorta var bolanderi foliage immature cones.jpg	File:Mendocino Pygmy Forest in Van Damme State Park 3.jpg	Pinus contorta subsp. bolanderi	Bolander's beach pine, Bolander pine, Also treated as Pinus contorta subsp. contorta var. bolanderi, in which case what is treated here as Pinus contorta subsp. contorta is the variety Pinus contorta subsp. contorta var. contorta.<ref name="ReferenceA">Template:Jepson eFlora</ref>	endemic to Mendocino County on the northwestern California coast;<ref>Template:GRIN</ref> near threatened by fires and development
File:Pinus contorta 37636.JPG	File:Pinus contorta Pacific Rim NP 1.jpg	Pinus contorta subsp. contorta	shore pine	Pacific Coast, southern Alaska to northwest California.<ref name=handbook>Template:Cite book</ref><ref>Template:Calflora</ref><ref>Template:GRIN</ref>
File:Pinus contorta 8160.jpg	File:Pinus contorta Lassen.jpg	Pinus contorta subsp. murrayana	tamarack pine, or Sierra lodgepole pine<ref>Template:Cite book</ref>	Cascade Range from Oregon into northern California; the Sierra Nevada, the Transverse Ranges of southern California (including the San Bernardino Mountains, the Peninsular Ranges into northern Baja California, and the Spring Mountains of southern Nevada).<ref>Template:GRIN</ref><ref name=handbook /><ref>Template:Calflora</ref>
File:Pinus contorta Nederland CO.jpg	File:Pinus contorta subsp latifolia Thermopolis Wyoming.jpg	Pinus contorta subsp. latifolia	lodgepole pine	Rocky Mountains, Colorado to Yukon and Saskatchewan; aspen parkland and boreal forests.<ref>Template:GRIN</ref><ref name="PLANTS">Template:PLANTS</ref><ref>Template:Cite book</ref>

Uses

Construction

The common name "lodgepole pine" comes from the custom of Native Americans using the tall, straight trees to construct lodges (tepees) in the Rocky Mountain area.<ref name=":0" /> Lodgepole pine was used by European settlers to build log cabins.<ref name=":0" /> Logs are still used in rural areas as posts, fences, lumber, and firewood.<ref name=":0" /> Shore pine pitch has historically been used as glue.<ref name=":0" />

Tree plantations of Pinus contorta have been planted extensively in Norway, Sweden,<ref>Template:Cite news</ref> Ireland and the UK for forestry, such as timber uses. In Iceland it is used for reforestation and afforestation purposes.<ref>Template:Cite web</ref> It is also commonly used for pressure-treated lumber throughout North America.

Food

Native Americans consumed the inner bark of lodgepole pine to prevent starvation for themselves and their horses.<ref name=":0" /> The lodgepole pine cambium layer's edibility is dependent on thickness, consistency, and sweetness.<ref>Template:Cite journal</ref>

Medicinal

The indigenous peoples of the Pacific Northwest and of California used different parts of the plant internally and externally as a traditional medicine for various ailments.<ref>Template:Cite web</ref> The gum of shore pine was used medicinally as well as for chewing.<ref name=":0" />

Cultivation

Pinus contorta is cultivated as an ornamental tree by the horticulture industry. Plant nurseries grow Pinus contorta subsp. contorta and Pinus contorta subsp. murrayana for use in traditional and wildlife gardens, and as smaller selections of the native plant for natural landscaping. The shore pine's smaller varieties and cultivars are also used in container gardening, including as large bonsai specimens.

Cultivars of this species include:

• "Chief Joseph", a dwarf variety of Pinus contorta var. latifolia grown for its yellow winter needles
• "Spaan's Dwarf", a dwarf variety of Pinus contorta var. contorta that grows wider than it grows tall

The cultivar "Chief Joseph" has gained the Royal Horticultural Society's Award of Garden Merit.<ref name = RHSPF>Template:Cite web</ref><ref>Template:Cite web</ref>

Emblem

Lodgepole pine is the provincial tree of Alberta, Canada.<ref name=ABEmblems>Template:Cite web</ref>

External links

• Template:Jepson eFlora
• University of Wisconsin: Lodgepole forest webpage Template:Webarchive
• Virginia Tech dendrology website: Pinus contorta
• Guardian (U.K.) article: "Plague of beetles raises climate change fears for American beauty"
• Picea sitchensis – information, genetic conservation units and related resources. European Forest Genetic Resources Programme (EUFORGEN)
• Template:CalPhotos
• Template:Silvics
• Template:PFAF