Pinus contorta

Pinus contorta, with the common names lodgepole pine and shore pine, and also known as twisted pine,[2] and contorta pine,[2] is a common tree in western North America.[3] It is common near the ocean shore and in dry montane forests to the subalpine, but is rare in lowland rain forests.[4][5] Like all pines (member species of the genus Pinus), it is an evergreen conifer.

Lodgepole pine
Pinus contorta subsp. contorta in Anacortes Community Forest Lands, Washington
Scientific classification
Kingdom: Plantae
Clade: Tracheophytes
Division: Pinophyta
Class: Pinopsida
Order: Pinales
Family: Pinaceae
Genus: Pinus
Subgenus: P. subg. Pinus
Section: P. sect. Trifoliae
Subsection: P. subsect. Contortae
Species:
P. contorta
Binomial name
Pinus contorta
Subspecies

4, see text

Distribution map:
  • Pinus contorta subsp. contorta
  • P. contorta subsp. latifolia
  • P. contorta subsp. murrayana

Subspecies

There are four subspecies of Pinus contorta, and one of them is sometimes considered to have two varieties.[6] The subspecies are sometimes treated at the rank of variety.[2][7][8]

ConeImageSubspeciesCommon Name & DescriptionDistribution
Pinus contorta subsp. bolanderiBolander'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.[9]endemic to Mendocino County on the northwestern California coast;[10] near threatened by fires and development
Pinus contorta subsp. contortashore pinePacific Coast, southern Alaska to northwest California.[11][12][13]
Pinus contorta subsp. murrayanatamarack pine, or Sierra lodgepole pine[14]Cascade Range from Washington 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).[15][11][16]
Pinus contorta subsp. latifolialodgepole pineRocky Mountains, Colorado to Yukon and Saskatchewan; aspen parkland and boreal forests.[17][18][19]

Description

Depending on subspecies, Pinus contorta grows as an evergreen shrub or tree. The shrub form is krummholz and is approximately 1 to 3 m (3 to 10 ft) high. The thin and narrow-crowned tree is 40 to 50 m (130 to 160 ft) high and can achieve up to 2 m (7 ft) in diameter at chest height.[7] 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.[5][20][21]

The species name is contorta because of the twisted, bent pines found at coastal areas and the tree's twisted needles.[11][22][23] Pinus contorta is occasionally known under several English names: black pine, scrub pine, and coast pine.[24][25] P. contorta subsp. latifolia will hybridise with the closely related jack pine (Pinus banksiana).

Needles and buds
The needles are 4 to 8 cm (1 1⁄2 to 3 in) long in fascicles of two, alternate on twigs. The female cones are 3 to 7 cm (1 to 3 in) long with sharp-tipped scales.

The egg-shaped growth buds are reddish-brown and between 20 and 30 mm (3⁄4 and 1 1⁄4 in) 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 4 to 8 cm (1 1⁄2 to 3 in) long and 0.9 to 2 mm (0.04 to 0.08 in) 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 2,000 m (6,600 ft), 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.[20]

Cones

The cones are 3–7 centimetres (1–3 in) long. The cones have prickles on the scales.

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.[26] The variation in their serotiny has been correlated with wildfires and mountain pine beetle attacks.[27] The cones of the coastal Pacific subspecies, P. contorta subsp. contorta, are typically non-serotinous,[25] and those of the inland Pacific subspecies, P. contorta subsp. murrayana, are completely non-serotinous.[28]

Ecology

Pinus contorta is a fire-dependent species, requiring wildfires to maintain healthy populations of diverse ages. The bark of the lodgepole pine is fairly thin, minimizing the tree's defense to fire; however, the heat of fire opens the cones to release the seeds. This allows the species to regenerate and maintain its place in the forest habitat.[29] One plant community in which Pinus contorta is found is the closed-cone pine forest of coastal California.

P. contorta subsp. latifolia forest 23 years before (above) and 10 years after (below) the Yellowstone fires of 1988

Excessive wildfire prevention disrupts the fire ecology. The stands are usually so densely populated that the trees self-thin, or out-compete each other, leaving dead trees standing. These become a dry ladder fuel that can accelerate the fire to the crown of living trees. When the fire reaches the crowns of the trees, it can jump from tree to tree and becomes relatively unstoppable.

The natural fire regime for this species is primarily driven by climate. The fires occur most often after years of drought. Pinus contorta occurs from the upper montane to the subalpine region. These types of forests experience a lot of moisture in the form of snow in the winter due to their altitude. The density of the tree stand also inhibits the establishment of an understory. With all of that being said, the likelihood of a surface fire occurring is rare. Thus, infrequent but severe fires dominate this species.[29]

An example of the climate that plays a huge role in the fire regime of Pinus contorta 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. So when the AMO +, ENSO − and PDO −, there is going to be a drought and likely a severe subalpine fire.[30]

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.[31][32]

Threats

This species is attacked by blue stain fungus (Grosmannia clavigera), distributed by the mountain pine beetle from its mouth.

A study released in 2011 concluded that Pinus contorta could experience significant reductions in distribution due to climate change by the late 21st century.[33][34]

Uses
Pinus contorta subsp. murrayana near summit of Pywiack Dome in Yosemite National Park.

Construction

Tree plantations of Pinus contorta have been planted extensively in Norway, Sweden, Ireland and UK for forestry, such as timber uses. It is also commonly used for pressure treated lumber throughout North America.

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.[35]

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 (ssp. contorta) 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.[36][37]

Emblem

Lodgepole pine is the Provincial tree of Alberta, Canada.

Invasive species

Pinus contorta is 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.

References
  1. Farjon, A. (2013). "Pinus contorta". IUCN Red List of Threatened Species. 2013: e.T42351A2974612. doi:10.2305/IUCN.UK.2013-1.RLTS.T42351A2974612.en.
  2. "Pinus contorta". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA).
  3. Moore, Gerry; Kershner, Bruce; Craig Tufts; Daniel Mathews; Gil Nelson; Spellenberg, Richard; Thieret, John W.; Terry Purinton; Block, Andrew (2008). National Wildlife Federation Field Guide to Trees of North America. New York: Sterling. p. 91. ISBN 978-1-4027-3875-3.
  4. Sullivan, Steven. K. (2015). "Pinus contorta". Wildflower Search. Retrieved 24 January 2015.
  5. Giblin, David, ed. (2015). "Pinus contorta". WTU Herbarium Image Collection. Burke Museum, University of Washington. Retrieved 24 January 2015.
  6. Conifer Specialist Group (1998). "Pinus contorta var. bolanderi". IUCN Red List of Threatened Species. 1998. Retrieved 12 May 2006.
  7. Kral, Robert (1993). "Pinus contorta". In Flora of North America Editorial Committee (ed.). Flora of North America North of Mexico (FNA). 2. New York and Oxford. Retrieved 12 September 2010 – via eFloras.org, Missouri Botanical Garden, St. Louis, MO & Harvard University Herbaria, Cambridge, MA.
  8. "Pinus contorta". World Checklist of Selected Plant Families (WCSP). Royal Botanic Gardens, Kew – via The Plant List.
  9. Jepson Flora Project (ed.). "Pinus contorta subsp. bolanderi". Jepson eFlora. The Jepson Herbarium, University of California, Berkeley.
  10. "Pinus contorta var. bolanderi". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA).
  11. Farjon, Aljos (2010). A Handbook of the World's Conifers. 2. Leiden, Netherlands: Koninklijke Brill NV. pp. 654–655. ISBN 978-90-04-17718-5.
  12. "Pinus contorta ssp. contorta". Calflora: Information on California plants for education, research and conservation, with data contributed by public and private institutions and individuals, including the Consortium of California Herbaria. Berkeley, California: The Calflora Database – via www.calflora.org.
  13. "Pinus contorta var. contorta". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA).
  14. Chase, J. Smeaton (1911). "Pinus murrayana (Tamarack, Lodge-pole-pine)". Cone-bearing Trees of the California Mountains. Eytel, Carl (illustrations). Chicago: A.C. McClurg & Co. p. 36. LCCN 11004975. OCLC 3477527.
  15. "Pinus contorta var. murrayana". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA).
  16. "Pinus contorta ssp. murrayana". Calflora: Information on California plants for education, research and conservation, with data contributed by public and private institutions and individuals, including the Consortium of California Herbaria. Berkeley, California: The Calflora Database – via www.calflora.org.
  17. "Pinus contorta var. latifolia". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA).
  18. "Pinus contorta". Natural Resources Conservation Service PLANTS Database. USDA. Retrieved 24 January 2015.
  19. Johnson, Kershaw; MacKinnon, Pojar (1995). Plants of the Western Boreal Forest and Aspen Parkland. Edmonton AB: Lonepine Publishing. p. 27. ISBN 978-1-55105-058-4.
  20. Schütt, Weisgerber; Schuck, Lang; Stimm, Roloff (2008). Lexikon der Nadelbäume. Hamburg, Germany: Nikol. pp. 365–367. ISBN 978-3-933203-80-9.
  21. Klinkenberg, Brian, ed. (2014). "Pinus contorta". E-Flora BC: Electronic Atlas of the Plants of British Columbia [eflora.bc.ca]. Lab for Advanced Spatial Analysis, Department of Geography, University of British Columbia, Vancouver. Archived from the original on 26 February 2015. Retrieved 24 January 2015.
  22. "Plants and Trees: lodgepole pine". U.S. Forest Service. Retrieved 12 August 2014.
  23. "Pinus contorta var. contorta : Shore Pine". Oregon State University. Retrieved 12 August 2014.
  24. "Forests of Crater Lake National Park: Lodgepole Pine (Pinus Contorta)". Crater Lake Institute. Retrieved 12 August 2014.
  25. Cope, Amy B. (1993). "Pinus contorta var. contorta". Fire Effects Information System (FEIS). US Department of Agriculture (USDA), Forest Service (USFS), Rocky Mountain Research Station, Fire Sciences Laboratory – via https://www.feis-crs.org/feis/.
  26. Anderson, Michelle D. (2003). "Pinus contorta var. latifolia". Fire Effects Information System (FEIS). US Department of Agriculture (USDA), Forest Service (USFS), Rocky Mountain Research Station, Fire Sciences Laboratory – via https://www.feis-crs.org/feis/.
  27. Feduck, Mike. "The genetic basis of cone serotiny in Pinus contorta as a function of mixed-severity and stand-replacement fire regimes". bioRxiv 10.1101/023267.
  28. Cope, Amy B. (1993). "Pinus contorta var. murrayana". Fire Effects Information System (FEIS). US Department of Agriculture (USDA), Forest Service (USFS), Rocky Mountain Research Station, Fire Sciences Laboratory – via https://www.feis-crs.org/feis/.
  29. Schoennagel, Tania; Thomas Veblen (2004). "The Interaction of Fire, Fuels and Climate across Rocky Mountain Forests". BioScience. 54 (7): 661–76. doi:10.1641/0006-3568(2004)054[0661:TIOFFA]2.0.CO;2. ISSN 0006-3568.
  30. Kauffman, J. Boone (August 2004). "Death Rides the Forest: Perceptions of Fire, Land Use and Ecological Restoration of Western Forests" (PDF). Conservation Biology. 18 (4): 878–82. doi:10.1111/j.1523-1739.2004.545_1.x. Retrieved 24 February 2010.
  31. Paul, L.R.; Chapman, B.K.; Chanway, C.P. (2007). "Nitrogen Fixation Associated with Suillus tomentosus Tuberculate Ectomycorrhizae on Pinus contorta var. latifolia". Annals of Botany. 99 (6): 1101–1109. doi:10.1093/aob/mcm061. PMC 3243579. PMID 17468111.
  32. Chapman, W.K.; Paul, L.R. (2012). "Evidence that Northern Pioneering Pines with Tuberculate Mycorrhizae are Unaffected by Varying Soil Nitrogen Levels". Microbial Ecology. 64 (4): 964–72. doi:10.1007/s00248-012-0076-0. PMC 3474912. PMID 22677953.
  33. Coops, Nicholas C.; Waring, Richard H. (March 2011). "A process-based approach to estimate lodgepole pine (Pinus contorta Dougl.) distribution in the Pacific Northwest under climate change". Climatic Change. 105 (1–2): 313–328. Bibcode:2011ClCh..105..313C. doi:10.1007/s10584-010-9861-2. S2CID 54177131.
  34. Rudolf, John Collins (28 February 2011). "Climate Change Takes Toll on the Lodgepole Pine". Green: A Blog About Energy and the Environment. Retrieved 1 March 2011.
  35. "NAEB". umich.edu.
  36. "RHS Plantfinder – Pinus contorta 'Chief Joseph'". Retrieved 2 May 2018.
  37. "AGM Plants – Ornamental" (PDF). Royal Horticultural Society. July 2017. p. 78. Retrieved 2 April 2018.
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