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Bolander Beach Pine

Pinus contorta Douglas ex Loudon

Comments

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Pinus contorta is fire successional over most of its range and is characterized by prolific seeding and high seed viability in disturbed habitats, often resulting in extremely slow-growing, overly dense stands. Some authors consider it to consist of 4 races; these have been given various infraspecific ranks, but perhaps they are more conventionally treated as 3 varieties.
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Flora of North America Vol. 2 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Description

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Shrubs or trees to 50m; trunk to 0.9m diam., straight to contorted; crown various according to genetic race. Bark brown to gray- or red-brown, platy to furrowed. Lower branches often descending, the upper spreading or ascending. Twigs slender, orange to red-brown, aging darker brown, rough. Buds narrowly to broadly ovoid, dark red-brown, to 1.2cm, slightly resinous. Leaves 2 per fascicle, spreading or ascending, persisting 3--8 years, 2--8cm ´ 0.7--2(--3)mm, twisted, yellow-green to dark green, all surfaces with fine stomatal lines, margins finely serrulate, apex blunt to acute or narrowly acuminate; sheath 0.3--0.6(--1)cm, persistent. Pollen cones ellipsoid to cylindric, 5--15mm, orange-red. Seed cones maturing in 2 years or variably serotinous, variably persistent, spreading to reflexed, often curved, nearly symmetric or variably asymmetric, lanceoloid to ovoid before opening, broadly ovoid to nearly globose when open, 2--6cm, tan to pale red-brown, lustrous, nearly sessile or on stalks to 1cm; apophyses nearly rhombic, variously elongate, cross-keeled, often mammillate toward outer cone base and on inside above middle; umbo central, depressed-triangular, prickle barely elongate to stubby or slender and to 6mm. Seeds compressed, obovoid; body ca. 5mm, red-brown, mottled with black, or all black; wing 10--14mm. 2 n =24 (variety not indicated).
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Flora of North America Vol. 2 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Comprehensive Description

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Pinus contorta, or lodgepole pine, is a common North American gymnosperm of the family Pinaceae. In Canada it is native to the Yukon, the Northwest Territories, British Columbia, Alberta, and Saskatchewan and, in the US, Alaska, Washington, Oregon, Idaho, Montana, South Dakota, Wyoming, Nevada, Utah, Colorado and California (USDA 2016).

There are five varieties of lodgepole pine found in different geographical regions: the shore pine (Pinus contorta var. contorta) that typically inhabits coastal areas, the Sierra lodgepole (Pinus contorta var. murrayana) which is found in the Sierra Nevada, the Rocky Mountain lodgepole (Pinus contorta var. latifolia) which is native to the Rocky Mountain range, the Bolander pine (Pinus contorta var. bolanderi) which is found in parts of California, and the Yukon pine (Pinus contorta var. yukonensis) of the Yukon Territories (Lotan and Critchfield 2004; USDA 2016).

Pinus contorta is very adaptable and can grow in many different environments, from rocky high elevations to sandy coastlines (British Columbia, 2016). While lodgepole pines are not considered to be invasive, they are one of the first plants to colonize an area after a fire or in cases of slope instability and floods (NPS 2016). Lodgepole pines tolerate a range of growing conditions and a wide variety of soils if they are well drained, although they prefer soils that are moist.

Lodgepole pines are most identifiable by their needles and cones. They typically have dark green needles that come in pairs and are about 1 to 3 inches long. Reproduction can usually start about 5­10 years into the life of the tree. The male cones are small and grow in bunches at the tips of the branches. The female cones, which take about 2 years to mature, will sometimes open once fertilized to release the seeds, or they can stay closed (a process called serotiny) until they are under high heat from a forest fire. The female cones are egg shaped, about 2 inches in length, and have a very strong outer protective layer (Lotan and Critchfield 2004). Pinus contorta can live up to 200 years and it can take up to 2 years for reproduction. The trees tend to be about 50 to 75 feet tall and are usually very thin, with an 8 to 12 inch trunk diameter and a narrow crown (NPS 2016). The species is shade intolerant, meaning that any branches that are left in the shade and do not receive sunlight will wither and fall off the tree. This is why the branches of lodgepole pines often do not start until higher up on the tree when growing around other trees. The bark is thin, reddish brown to grey, and finely scaled (British Columbia 2016).

References

  • British Columbia. 2016. Lodgepole Pine. Ministry of Forests, Lands, and Natural Resource Operations (www.for.gov.bc.ca. Accessed: May 15, 2016.
  • Lotan, James E., and William B. Critchfield. 2004. "Pinus contorta Dougl. ex Loud." United States Forest Service, Silvics Manual, Vol. 1 (www.na.fs.fed.us. Accessed: May 15, 2016).
  • NPS. 2016. "Lodgepole Pine." US National Park Service, Department of the Interior (www.nps.gov. Accessed: May 15, 2016).
  • USDA, NRCS. 2016. "Plant Profile for Pinus contorta (lodgepole pine)." The PLANTS Database (http://plants.usda.gov, Accessed: May 15, 2016). National Plant Data Team, Greensboro, NC 27401-4901 USA.

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Authors: Martha Rubardt and Rose Thompson; Editor: Gordon L. Miller, Ph.D.; Seattle University EVST 2100 - Natural History: Theory and Practice
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Associated Forest Cover

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Lodgepole pine grows both in extensive, pure stands, and in association with many western conifers. The forest cover type Lodgepole Pine (Society of American Foresters Type 218) (26) exists as a pure (80 percent or more) component of basal area stocking, as a majority (50 percent or more), or as a plurality (20 percent or more). The cover type includes all recognized subspecies of Pinus contorta.

Lodgepole pine is a component in 27 of the 55 SAF western forest cover types. In the Northern Interior (Boreal) group it is represented in White Spruce (Type 201), White Spruce-Aspen (Type 251), White Spruce-Paper Birch (Type 202), Paper Birch (Type 252), and Black Spruce (Type 204).

It is a component in all six high elevation cover types: Mountain Hemlock (Type 205), Engelmann Spruce-Subalpine Fir (Type 206), Red Fir (Type 207), Whitebark Pine (Type 208), Bristlecone Pine (Type 209), and California Mixed Subalpine (Type 256). At middle elevations in the interior it is a minor component of seven other types: Interior Douglas-Fir (Type 210), Western Larch (Type 212), Grand Fir (Type 213), Western White Pine (Type 215), Blue Spruce (Type 216), Aspen (Type 217), and Limber Pine (Type 219). In the North Pacific forests, it is a component in Coastal True Fir (Type 226), Western Redcedar-Western Hemlock (Type 227), Western Redcedar (Type 228), Douglas-Fir-Western Hemlock (Type 230), Port-Orford-Cedar (Type 231), and Redwood (Type 232). At low elevations in the interior it is associated with Interior Ponderosa Pine (Type 237) and in the South Pacific forests it is a component of Jeffrey Pine (Type 247).

Lodgepole pine, with probably the widest range of environmental tolerance of any conifer in North America, grows in association with many plant species (30,50,59,60). The lodgepole pine forest type is the third most extensive commercial forest type in the Rocky Mountains.

Lodgepole pine's successional role depends upon environmental conditions and extent of competition from associated species. Lodgepole pine is a minor seral species in warm, moist habitats and a dominant seral species in cool dry habitats. It is often persistent even on cool and dry sites and can attain edaphic climax at relatively high elevations on poor sites. Fire regimes have played a role in this successional continuum, especially where repeated fires have eliminated a seed source for other species (27). Lodgepole pine may even overwhelm a site with seed stored in serotinous cones. It has four basic successional roles (50):

Minor Seral- A component of even-aged stands rapidly being replaced by shade-tolerant associates in 50 to 200 years.

Dominant Seral- The dominant cover type of even-aged stands with a vigorous understory of shade-tolerant species that will replace lodgepole pine in 100 to    200 years.

Persistent- The dominant cover type of even-aged stands with little evidence of replacement by shade-tolerant species.

Climax- The only tree species capable of growing in a particular environment; lodgepole pine is self-perpetuating.

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Climate

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Lodgepole pine grows under a wide variety of climatic conditions (52). Temperature regimes vary greatly. Minimum temperatures range from 7° C (45° F) on the coast to -57° C (-70° F) in the Northern Rocky Mountains. Maximum temperatures range from 27° C (80° F) along the coast and at high elevations to well over 38° C (100° F) at low elevations in the interior. Average July minimums frequently are below freezing at high elevations. Lodgepole seedlings are relatively resistant to frost injury in some locations (16,42) and often survive in "frost-pockets" where other species do not.

At low elevations in the interior, lodgepole pine grows in areas receiving only 250 mm (10 in) of mean annual precipitation, whereas it receives more than 500 mm (200 in) along the northern coast. Many interior sites often are low in summer rainfall. Seasonal distribution of precipitation is significant; snowfall supplies most of the soil water used for rapid growth in early summer. Temperatures are frequently favorable for germination after snowmelt, and germination occurs rapidly. Lodgepole is very intolerant of shade and generally grows best in full sunlight.

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Damaging Agents

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The mountain pine beetle (Dendroctonus ponderosae) is the most severe insect pest of lodgepole pine. The epidemics that periodically occur in many lodgepole pine stands seriously affect the sustained yield and regulation of managed stands.

Adult beetles attack lodgepole pine in July or August, introducing bluestain fungi (8). The beetles construct egg galleries in the phloem where larvae feed and together with the fungi, girdle and kill the tree. Larvae overwinter in the tree, complete development, and emerge as adult beetles in the spring.

Harvesting has been considered as a means of preventing mountain pine beetle epidemics (19). Silvicultural practices in an integrated program for controlling losses to mountain pine beetle have been suggested (9,18). No mortality occurred in heavily thinned stands in Oregon where vigor ratings were high (44).

The mountain pine beetle has played an historic role in the dynamics of lodgepole pine ecosystems. By periodically invading stands and creating large amounts of fuels, which are eventually consumed by fire, creating favorable conditions for regeneration (12,39), the beetle has increased the probability that lodgepole pine will reoccupy the site at the expense of other species.

Another aggressive bark beetle that attacks lodgepole pine is the pine engraver (Ips pini). Ips commonly develops in logging slash, especially slash that is shaded and does not dry quickly. Prompt slash disposal is an effective control measure. Ips also can build up in windthrows.

Other insects that can be damaging local pests are the lodgepole terminal weevil (Pissodes terminalis), which can be destructive to elongating terminal leaders; larvae of the Warren's collar weevil (Hylobius warreni), which girdles roots and the root collar; larvae of the weevil Magdalis gentilis, which mine branches; various sucking insects, such as the pine needle scale (Chionaspis pinifoliae), the black pineleaf scale (Nuculaspis californica), and the spruce spider mite (Oligonychus ununguis); and several defoliating insects, among which are the lodgepole sawfly (Neodiprion burkei), the lodgepole needle miner (Coleotechnites milleri), the sugar pine tortrix (Choristoneura lambertiana), the pine tube moth (Argyrotaenia pinatubana), and the pandora moth (Coloradia pandora) (7).

Dwarf mistletoe (particularly Arceuthobium americanum) is the most widespread and serious parasite affecting lodgepole pine (11,29). A. americanum seeds are forcibly ejected from the fruit for distances as great as 9 m (about 30 ft). The sticky seeds adhere to the foliage of potential host trees. The proportion of trees visibly infected can double each 5 years between the ages of 10 and 25, with nearly a third of the trees infected at age 25 (29).

Rate of spread in young stands is about 0.3 to 0.5 m (1.0 to 1.5 ft) per year, with the fastest rate in dense stands. In many areas, more than 50 percent of lodgepole pine forests are infected. Dwarf mistletoe infection results in reduced diameter and height growth, increased mortality, reduced wood quality, decreased seed production, and overall decreased vigor.

Both harvesting and fire can greatly lessen the rate of spread and rates of infection. Effective control can be accomplished by clearcutting and locating boundaries of the unit to minimize reinfection from surrounding stands. Fire can effectively limit spread of dwarf mistletoe by eliminating sources of infection and establishing vast acreages of dwarf mistletoe-free areas.

Lodgepole pine is subject to attack by many fungal pathogens (33). These fungi are responsible for reduced growth and considerable cull and mortality. They also contribute in no small measure to the large amounts of logging residues that commonly occur when lodgepole pine is harvested.

One of the most serious diseases in lodgepole pine is a stem canker caused by Atropellis piniphila. Cankered stems are usually useless for lumber or posts and poles. Stem cankers of rust fungi cause extensive mortality, growth loss, and cull in lodgepole pine. Of these comandra blister rust (Cronartium comandrae) isthe most serious. The western gall rust (Peridermium harknessii) is especially damaging; trunk cankers can cause cull in logs and can kill seedlings and saplings. Because this rust does not require an alternate host, it can directly reinfect pines. Other fungi attack lodgepole pine and may cause serious losses in wood production. Examples are needle casts (such as Elytroderma deformans and Lophodermella concolor); root rots (such as Armillaria mellea and Heterobasidion annosum); and wood decays (such as Phellinus pini and Peniophora pseudo-pini).

Seed and seedling diseases are not usually damaging, although locally several mold fungi are associated with seed losses in germination, and rotting and damping-off can affect young seedlings.

Because of its relatively thin bark, lodgepole pine is more susceptible to fire than Douglas-fir and many other associates. It is less susceptible than Engelmann spruce or subalpine fir. Mortality from beetle epidemics often creates large amounts of jackstrawed fuel, which ignites easily from lightning and other sources and hampers fire control efforts.

Chinook winds following extremely cold weather occasionally cause red belt injury, particularly in Canada and Montana. Defoliation of trees is common and mortality can occur over large areas. Heavy snow can break or bend trees, particularly in dense stands with narrow crowns and intense root competition. Thinning can contribute to snow breakage, particularly if previously dense stands are opened suddenly.

Animals can cause considerable damage in thinned stands in some areas. Porcupines were attracted to thinned and fertilized stands in Montana. Pocket gophers often cover small seedlings under their entrance mounds and "winter-casts." They also feed on or clip both roots and tops. Gopher populations often explode as vegetation increases in open areas.

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Flowering and Fruiting

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Male and female strobili generally are home separately on the same tree in this monoecious species, with female flowers most often at the apical end of main branches in the upper crown, and male flowers on older lateral branches of the lower crown. The reddish purple female flowers grow in whorls of two to five and are 10 to 12 mm (0.4 to 0.5 in) long. The pale yellow to yellowish orange male flowers are crowded clusters of catkins at the base of new shoots and are 8 to 14 mm (0.3 to 0.6 in) long. It is not uncommon to find a dominance of maleness or femaleness on individual trees.

Pollen generally matures in mid-May to mid-July (table 1) (20,52). The time at which pollen matures appears to be related to elevation and climate.

Table 1- Time of pollen shedding in natural stands of lodgepole pine (20,52, modified)
Stand location
Elevation¹ Years observed Date of peak shedding m ft Vancouver, BC - - 2 Middle to late May Northwestern Washington 150 500 10 May 12 Mendocino White Plains, California - - 1 June 9 Northern Cascades 1200 4,000 - Mid-June Northern Idaho; western Montana - - 10 June 13 Central and eastern Washington and Oregon 790 to 1300 2,600 to 4,250 - June 13 Southeastern Alberta (subalpine forest) - - 10 June 22 Sierra Nevada, California 1820 6,000 3 June 22 Central Montana; Yellowstone region - - 10 June 25 Northern Utah 2190 7,200 2 July 12 Southern Idaho 2070 6,800 1 July 7 Northern Idaho; western Montana 670 to 1265 2,200 to 4,150 10 June 6 Eastside Montana; Yellowstone National Park 975 to 2060 3,200 to 6,750 10 June 17 ¹Dash indicates data are not available. Seed cones usually mature in August, September, or October, more than a year after pollination. Inland forms and high elevation stands apparently mature earlier than coastal forms or low elevation stands. Cones open in early September in the Northern Rocky Mountains. Cone maturity is indicated by a change in color from purple-green to light brown (54).

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Genetics

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This summary is based on a recent review of the literature on the genetics of lodgepole pine (20). The ability of some strains of lodgepole pine to grow well on poor sites and in cold climates has interested European foresters for many years. Much of what is known about the genetic diversity of the species has been learned from provenance tests, mostly in northwestern Europe. These tests have established that much of the variation observed in natural stands of lodgepole pine has a genetic basis.

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Growth and Yield

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Growth and yield of lodgepole pine is greatly affected by stand density (31) as well as by environmental factors (2,6,22,46). In fact, site index curves have been developed with corrections for effects of stand density.

Maximum yield in the Rocky Mountains was 280 m³/ha (20,000 fbm/acre) at a density of 1,980 trees per hectare (800/acre), but only 21 m³/ha (1,500 fbm/acre) at a density of 4,450/ha (1,800/acre), assuming 5 fbm/ft³; original figures were in board feet (27).

In extreme cases 70-year-old stands with 247,000 trees/ha (100,000 trees/acre) averaged only 1.2 m (4 ft) in height and less than 2.5 cm (1 in) in diameter at ground level.

Yields of 168 to 224 m³/ha (about 12,000 to 16,000 fbm/acre) can be found in old-growth Rocky Mountain lodgepole pine. Yields of more than 336 m³/ha (about 24,000 fbm/acre) are the result of a fortuitous combination of favorable initial stocking, good site quality, and absence of mountain pine beetle and dwarf mistletoe.

Relationships among age, stocking levels, and development in natural stands were summarized for medium sites in Montana and Idaho (site index 22.9 m or 75 ft at 100 years) (table 2). Under light to moderate stocking, live crowns are 25 to 60 percent of total height.

Table 2- Relationships among stand age and stocking level, and tree development and typical yield in natural stands of lodgepole pine, summarized for medium sites in Montana and Idaho (site index 22.9 m or 75 ft at base age of 100 years)¹
Age
Stocking Average height of dominants Average stand diameter
Total cubic volume
Merchantable volume yr trees/ha trees/acre m ft cm in m³/ha ft³/acre m³/ha ft³/acre fbm/acre   20   1,240    500   5.5 18   8.6   3.4   16.1    230 - 19,770 8,000   3.0 10   4.1   1.6   28.0    400 -   50   1,180    479 12.5 41 16.5  6.5 144.9 2,070 130.2 1,860   5,100 15,200 6,150   9.1 30   6.9   2.7 165.9 2,370 - - -   80   1,030    418 18.0 59 20.6   8.1 285.6 4,080 266.0 3,800 12,100   7,500 3,034 14.6 48   9.1   3.6 280.0 4,000 - - - 110      850    344 22.3 73 23.6   9.3 385.7 5,510 363.3 5,190 18,200   4,600 1,861 18.9 62 11.4   4.5 357.0 5,100 273.0 3,900   8,400 140      680    275 25.3 83 26.7 10.5 448.7 6,140 426.3 6,090 23,200   3,070 1,243 22.3 73 14.0   5.5 416.5 5,950 301.0 4,300 10,300 ¹Compiled from unpublished yield tables furnished by D.M. Cole, USDA Forest Service, Intermountain Research Station, Bozeman, MT. Cubic volumes are from trees 11.4 cm (4.5 in) in d.b.h. to a 7.6 cm (3 in) diameter top. Board foot volumes are from trees larger than 16.5 cm (6.5 in) in d.b.h. to a 15.2 cm (6 in) diameter top. Mature sizes vary greatly between stands. In the Rocky Mountains, most trees at 140 years of age were 18 to 33 cm (about 7 to 13 in) in d.b.h. and 18 to 25 m (about 60 to 80 ft) in height (27).

Trees in the Blue Mountains of Oregon average 30 cm (about 12 in) in d.b.h. and 23 m (about 75 ft) tall at 100 years of age. Sierra Nevada trees the same age are larger, averaging 42 cm (about 16 to 17 in) in d.b.h. and 28 to 30 m (about 90 to 100 ft) tall. Coastal trees are smaller but vary greatly. Mature trees range from 15 to 50 cm (about 6 to 20 in) in d.b.h. and only 6 to 12 m (20 to 40 ft) tall. Dwarf lodgepole pines are only about a meter (2 to 5 ft) tall and are found along the coast in Mendocino County, CA. This small size is thought to be caused by a highly acid hardpan.

Growth of lodgepole pine is often so stagnant that stand culture is not practical. Early management and control of stocking greatly affects growth and yield of lodgepole pine stands (17). Average annual growth in old-growth unmanaged stands in the central Rocky Mountains only was 0.4 to 0.6 m³/ha (about 25 to 40 fbm/acre) because of large numbers of small trees and a high incidence of dwarf mistletoe (4). (Calculations assume 5 fbm/ft³; original figures were in board feet). Annual net growth may be increased to 2.1 to 5.6 m³/ha (about 150 to 400 fbm/acre) by controlling stand density and reducing dwarf mistletoe infection (5,25).

Control of stand density offers the greatest opportunity for increasing productivity of any readily available management practice.

Culmination of total cubic volume occurs as early as 40 years in severely stagnated stands, and between 50 and 80 years for overstocked, but not greatly stagnated, stands. Merchantable volume culmination in stands of the latter type occurs between 110 and 140 years, depending on merchantability standards.

Thinning of young overstocked and stagnating stands can restore growth potential and redirect it into merchantable-size products. With more complete utilization (lower merchantable d.b.h. and top diameters), most of the yield increase possible from thinning is attained with the first entry, a stocking control thinning (17).

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Reaction to Competition

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Lodgepole pine is very intolerant of shade and competition from other plant species. Occasionally seedlings become established under a forest canopy, but these individuals rarely do well. In spite of its shade intolerance, lodgepole pine maintains itself in dense stands for long periods, often for 100 years or more.

In the absence of fire, lodgepole pine is usually succeeded by its more tolerant associates, such as Engelmann spruce and subalpine fir. Succession proceeds at variable rates, however, and is particularly slow in some high elevation forests.

Pure stands of lodgepole pine persist for varying lengths of time. In northern Idaho and central Oregon, stands begin to break up at 80 to 100 years, while stands at higher elevations, such as in Montana, southern Idaho, Utah, and Wyoming, last for several hundred years. Pure stands in and around Yellowstone National Park contain 300- to 400-year-old trees, with several groups of younger even-aged trees. These stands no doubt originated as even-aged stands but have been breaking up for more than two centuries.

The ability of lodgepole pine to regenerate at the expense of other species is due not only to cone serotiny but also to seed viability, germinative energy, early rapid growth, and ability to survive a wide variety of microsite and soil situations (39).

Compared to its associates, lodgepole pine is intermediate in its needs for water, requiring more than Douglas-fir and ponderosa pine and less than Engelmann spruce and subalpine fir. On some sites, lodgepole pine appears to compete well for water, however, and grows where other species may be excluded because of lack of water (45,57); on others it appears to be tolerant of high water tables (14,43). It is also intermediate in its tolerance to extremes of temperature (27).

Lodgepole pine shows good response to thinning at an early age (17). Heavily stocked stands must be thinned before stagnation occurs. The best age for thinning varies with site and density. Poor sites and overstocked stands particularly must be thinned as early as age 10.

Diameter growth acceleration is usually greatest in heavy thinnings; cubic volume and basal area growth are usually greatest in light thinnings (27). Although mechanical thinning, as with bulldozer strips, is a convenient alternative, obtaining a proper response (36) is difficult.

At older ages, growth response is strongly correlated with crown size, vigor, and amount of release provided (27). Attempts at partial cutting of mature and over-mature stands have resulted in little gain or even negative net volume growth (1,28).

Lodgepole pine can be maintained best in a vigorous, productive forest by using a silvicultural method that regenerates even-aged stands (38). This often may be accomplished by clearcutting and by relying upon natural regeneration or planting. Planting provides an excellent opportunity for initial stocking control and/or genetic improvement.

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Rooting Habit

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The root system of lodgepole pine varies considerably in form, depending on soil type. Root growth is particularly important during the critical first year. Root growth of 12.7 to 15.2 cm (5 to 6 in) was reported for seedlings growing on prepared seedbeds in Montana and Idaho (34). First-season seedlings had an average root depth of only 9.6 cm (3.8 in) on scarified, unshaded seedbeds in the central Rocky Mountains (47). Seedlings growing near grass competition usually do not penetrate beyond 5 or 6 cm (about 2 in).

Taproots and vertical sinkers are common, but where a hardpan or water table is encountered, the taproot may die, bend, or assume a horizontal position. Planting may affect root configuration. Taproots of seedlings planted with "J-roots" often grow horizontally for many years before sinkers develop.

Because of its shallow root system, lodgepole pine is susceptible to windfall, particularly after stands are opened by harvesting. Windfirmness varies with stand density, soil conditions, and topography. Shallow roots are common above hardpan or in shallow, rocky soils.

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Seed Production and Dissemination

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Lodgepole pine produces viable seed at an early age, commonly 5 to 10 years; germination percentage is as high as that of seed borne by mature trees. Pollen flowers have been observed on 2-0 seedlings in the Lucky Peak Nursery near Boise, ID.

Lodgepole pine is a prolific seed producer. Good crops can be expected at 1- to 3-year intervals, with light crops intervening. The cones withstand below freezing temperatures and are not generally affected by cone- and seed-feeding insects. Only squirrels and coreid bugs are significant seed predators. Seed production should not be taken for granted, however. Complete seed crop failures have occurred at 2800 m (9,200 ft) in northwest Wyoming for 2 to 4 years in a row (42).

Cone production of individual dominant and codominant trees can vary from a few hundred to a few thousand per tree (37). Cones are persistent, and serotinous (closed) cones accumulate for decades. Annual production may run from 173,000 to 790,000 seeds per hectare (70,000 to 320,000/acre) with half to one-third available for annual seedfall (27). An annual seedfall of 99,000 to 222,000 seeds per hectare (40,000 to 90,000/acre) was found in central Montana (58). These figures might be considered typical for interior lodgepole pine where some portion of the trees are of the serotinous type. In Oregon, where the nonserotinous cone habit is prevalent, seedfall ranged from about 35,000 to over 1.2 million/ha (14,000 to 500,000/acre) (21). Most years seedfall was on the order of hundreds of thousands per hectare. Where stored seeds are in the millions per hectare (in closed cones), the number of seeds stored is probably 10 times that of seeds produced annually (37).

Although the number of fully developed seeds per cone varies from as few as 1 to 2 to as many as 50, a normal average for large cone lots in the Rocky Mountains is from 10 to 24 seeds per cone (42). Sierra Nevada populations range from 5 to 37 seeds per cone (20).

The serotinous cone habit varies over wide geographic areas as well as locally (37). Serotinous cones are not common in eastern Oregon, rare in coastal populations, and absent in the Sierra Nevada and southern California and Baja California populations (20). Although common in the Rocky Mountains, this cone habit varies considerably (37). Many stands in the Rockies have less than 50 percent serotinous-cone trees.

Lodgepole pine has long been regarded as a fire-maintained subclimax type. Its ability to regenerate in extremely dense stands to the exclusion of other species can be attributed to the closed cone habit. Millions of seeds per hectare are held in reserve for many years and are readily available to germinate on the seedbed prepared by fire. Recent evidence seems to indicate that fire selects strongly for the closed cone habit (49).

Serotinous cones do not open at maturity because of a resinous bond between the cone scales. The bonds break with temperatures between 45° and 60° C (113° to 140° F) (48), and cone scales are then free to open hygroscopically. Large quantities of seeds are thus available for regenerating a stand following fire. Closed cones at or near the soil surface (less than 30 cm or about 12 in) are also subjected to temperatures from insolation sufficient to open them and may provide seed in harvested areas. Some seeds may be damaged by fire, however, particularly in fires burning in logging slash.

Seeds stored in serotinous cones on the tree remain viable for years. Apparently, prolonged viability can be maintained so long as cones or seeds are not in contact with the ground. Once cones are on the ground, cones open. Damping-off fungi may infect the seed, rodents may feed on the seeds, or germination may occur; for the most part, seeds are not stored in the soil.

Lodgepole pine has relatively small seeds for pine. Seed weights vary considerably, ranging from 2.3 mg (0.04 grains) per seed in the Interior of Canada to 11.4 mg (0.18 grains) per seed in the Sierra Nevada (20). Lodgepole pine seeds average about 298,000 cleaned seeds per kilogram (135,000/lb) for varieties contorta, 258,000/kg (117,000/lb) for murrayana, and 207,000/kg (94,000/lb) for latifolia (54). Density of seedfall 20 m (66 ft) from the timber edge is only 10 to 30 percent of that at the timber edge for stands in the Rocky Mountains (fig. 1) (42). Dispersal of sufficient seed to adequately restock an area often is only about 60 m (200 ft) (23,38). Prevailing winds, thermal effects, or scudding on the snow may disperse seeds far beyond these distances, however.


Figure 1- Sound seed per hectare as a function of distance
from the nearest timber edge.


The annual seedfall from nonserotinous cones helps in restocking relatively minor disturbances in the stand, in maintaining the presence of lodgepole pine in mixed stands, and in expanding conifers into other vegetative types. Seldom do we find stands without some trees of the open-coned type. The efficacy of this seed source can be seen in the dense stands of lodgepole pine along road cuts, powerline rights-of-way, and ditches or where disturbance occurs near lodgepole pine stands.

Studies of seedfall have shown variation in the number of seeds released soon after cone maturation, but most seeds (80 to 90 percent) are released before the following growing season (27).

Where large amounts of seed are stored in serotinous cones, a most effective means of seed dispersal in clearcuts is from cones attached to the slash and those knocked from the slash and scattered over the forest floor during slash disposal. Many cones on or near the ground are opened by normal summer soil surface temperatures (35). In Montana 83 percent of the cones on the ground opened the first year on south slopes compared to 40 percent on north slopes. Maximum seed release from serotinous cones near the ground takes place during the first year of exposure. In fact, cones may open after the first few minutes of exposure to temperatures high enough to break the resinous bonds.

In slash, serotinous cones that are well above the ground behave like those on a tree- they remain closed, and stored seeds remain viable for years.

Seeds in unopened cones and those released from the slash may also be lost to rodents, fungi, and other destructive agents. Seeds from closed cones are usually available only for the first growing season following harvest, but stocking from open-cone seed sources can continue to increase for several years.

Slash disposal on areas where regeneration is planned from serotinous cones must be carefully planned and executed. Seed supply will be largely destroyed if slash to be burned is piled before cones have had a chance to open (38). Piling slash should be delayed until sufficient cones have opened to assure adequate stocking. Piling then scatters seeds and opened cones and helps prepare the seedbed. Piling slash after germination can also decrease stocking because young seedlings are trampled or buried.

Broadcast burning may hasten release of seeds from cones not in a position to open from high soil-surface temperatures. Some seeds will be destroyed, however; the amount will vary with fire intensity.

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Seedling Development

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Germination under field conditions is good if climate and seedbed are favorable. Best germination occurs in full sunlight and on bare mineral soil or disturbed duff, free of competing vegetation. Germination is epigeal. Temperatures fluctuating between 8° and 26° C (47° and 78° F) favor germination. Adequate soil moisture is required for germination and survival during the critical few weeks following germination (34,51,55). In southwest Montana and southeast Idaho, 75 to 90 percent of a season's total germination occurred during the 2 weeks following snowmelt in late June (34), when the soil was saturated and temperatures were favorable. Germination can be delayed if cones do not open during the previous summer.

Although lodgepole pine germinates well on most organic seedbeds, such materials tend to dry faster than mineral soil and seedlings often die in this seedbed. Lodgepole pine seeds have little need for stratification and germination depends largely upon temperature (20). At optimum temperatures and moisture, almost 100 percent of the seeds germinate rapidly.

Both shading and competition inhibit germination and survival. Newly germinated seedlings are relatively insensitive to temperature extremes. Because residual overstory following partial cutting usually does not provide the most favorable conditions for regeneration, clearcutting is generally recommended. On some areas, however, lodgepole pine has established itself in the shade of lightly cut or uneven-aged stands and may persist for many years in the understory. Some of these trees eventually may establish a crown sufficient to permit reasonable growth.

Drought is a common cause of mortality among first-year seedlings; losses vary with soil type and seedbed condition. Greatest losses occur on soils with low water-holding capacity, and duff and litter. Well decomposed organic material, incorporated in the soil, enhances seedling survival, however. Disturbed mineral soil seedbeds generally produce the best germination and survival (34,40,41). Shading has been demonstrated to help under drought conditions in Wyoming (10).

Drought losses usually decline considerably after the first growing season. First-year seedlings are particularly vulnerable because of a relatively shallow root system (34,47).

Young, succulent seedlings may die because of high soil surface temperatures (13). By 2 to 4 weeks of age, seedlings are able to withstand soil surface temperatures higher than 60° C (140° F), which commonly occur at high elevation sites. Freezing temperatures may kill seedlings either directly or by frost heaving. In much of the range of lodgepole pine, however, frosts occur regularly throughout the growing season and seedlings from different sources vary in frost resistance (16). The amount of frost heaving varies considerably by soil type, location, and year of occurrence but can cause significant losses.

Lodgepole pine seedlings are poor competitors and competition from grass is often most detrimental. The Douglas-fir/pinegrass habitat type is one of the most difficult sites for lodgepole pine regeneration, particularly if the regeneration effort is delayed until a firm sod cover is established.

Grazing animals, particularly cattle, can cause seedling mortality by trampling. Sheep actually seek the succulent new "candles" in the spring and nibble needles and small branches if other feed is not abundant.

A common problem of regenerating lodgepole pine stands is overstocking, which results in stagnation at early ages. Many sites are stocked with tens of thousands and even hundreds of thousands of trees per hectare.

If trees are well distributed, stocking should not exceed 1,240 to 1,980 stems per hectare (500 to 800/acre) between 5 years and 20 years of age (17). Proper distribution and full utilization of the site, however, may require establishment of 2,470/ha (1,000/acre) and thinning to obtain proper spacing. There is also potential for significant genetic gains from selection of elite trees when thinning.

An average height of 3.6 m (12 ft) and d.b.h. of 5 cm (2 in) on fully stocked 20-year-old stands was found on above average sites in Montana (27). Average heights of 2.0 m (6.7 ft), 4.2 m (13.8 ft), and 7.6 m (24.9 ft) were found on low, medium, and high sites in 20-year-old stands in the Foothills Section of Alberta (for density class 1,240 stems per hectare or 500/acre at 70 years of age) (32).

Lodgepole pine height growth begins earlier than any of its associates except other pines and larch (53).

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Soils and Topography

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Lodgepole pine grows on soils that vary widely but are usually moist. Growth is best where soil parent materials are granites, shales, and coarse-grained lavas (24,27); other soils have developed from glacial till of widely varying composition, Recent, Tertiary, and Oligocene alluvium and colluvium (from such sources as quartzites and argillites), limestone of the Belt geologic series, pumice, and volcanic ash. Lodgepole pine is seldom found on the generally drier soils derived from limestone. In Canada, however, extensive stands occur on calcareous glacial tills (56). Glacial drift provides a balance of moisture and porosity on which the species seems to thrive, as in Alberta, where it grows better on glacial tills than on alluvial soils or lacustrine deposits. In Montana, highly calcareous soils derived from dolomitic limestone usually do not support lodgepole pine, subalpine fir (Abies lasiocarpa), and Engelmann spruce (Picea engelmannii), although they do support Rocky Mountain Douglas-fir (Pseudotsuga menziesii var. glauca). Nevertheless, soils developed in colluvium from other types of limestone and calcareous glacial till do support stands of lodgepole pine.

Extensive stands of lodgepole pine (var. latifolia) occur on soils classified as Inceptisols or Alfisols in the interior forests. Although the species commonly grows on Andepts and does well on these soils in some areas, the Boralfs and Ochrepts probably support better tree development and more extensive stands. Frequently lodgepole pine soils on Boralfs and Ochrepts have cryic soil temperature regimes. In the Blue Mountains of Oregon lodgepole pine does well on Andepts, where it is nearly always found on volcanic ash or alluvial material overlying residual basaltic soils, at elevations between 910 and 2130 m (3,000 and 7,000 ft). The ash cap soils are deeper and hold more moisture than the residual soils.

The coastal form of lodgepole pine (var. contorta) is often found on Histosols (peat bogs or muskegs) in southeastern Alaska, British Columbia, and western Washington, and on dry, sandy, or gravelly sites farther south along the coast on Inceptisols, Alfisols, and Ultisols.

Soil properties and soil moisture often favor lodgepole pine locally over other species. Lodgepole pine grows on wet flats and poorly drained soils in the Cascade Range in Washington and Oregon, and the Sierra Nevada in California. Soils with underlying hardpan support lodgepole pine to the exclusion of such species as ponderosa pine (Pinus ponderosa), redwood (Sequoia sempervirens), or Douglas-fir in the Sierra Nevada, eastern Oregon, and Mendocino County, CA. Lodgepole pine also grows on level sites with and without high water tables in central Oregon where frost tolerance during germination allows its establishment to the exclusion of other species. Extensive stands are found in these areas on well drained sites above 1600 m (5,250 ft), with patterns of occurrence attributed to past fires.

On infertile soils, lodgepole pine is often the only tree species that will grow. Nevertheless, experiments have demonstrated significant growth increase from fertilization, particularly nitrogen (15).

Lodgepole pine thrives in a wide variety of topographic situations. It grows well on gentle slopes and in basins, but good stands are also found on rough and rocky terrain and on steep slopes and ridges, including bare gravel. Northern and eastern slopes are more favorable than southern and western aspects (3).

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Special Uses

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Lodgepole pine is not only an important timber species but is also a major tree cover in many scenic and recreational areas and on critical watersheds. It provides many acres of wildlife habitat and is associated with many grazing allotments throughout its range. It is important to local communities throughout the West.

Lodgepole pine is used for framing, paneling, posts, corral poles, utility poles, railroad ties, and pulpwood. As new developments such as structural particleboard become practical, the rapid juvenile growth of the species will be an advantage where gross cubic volume becomes important. Even now, with properly designed machinery, it is economically harvested, and this harvesting, properly done, can enhance watershed, forage, wildlife habitat, and scenic and recreational values.

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Vegetative Reproduction

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Lodgepole pine can be grafted successfully, but results vary depending upon the clone (20). Natural sprouting has been observed on the Bitterroot National Forest in Montana. Branches not severed often become leaders on stumps left in thinning operations.

Lodgepole pine cuttings are relatively easy to root. Adventitious roots have been developed artificially from 8-year-old lodgepole pine (by air-layering) after treatment with either indole-acetic or indole-butyric acid (17).

Callus tissue cultures and liquid cell suspensions have been produced from seedling hypocotyl tissue, excised embryos, and actively growing shoots.

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Distribution

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Lodgepole pine is an ubiquitous species with a wide ecological amplitude. It grows throughout the Rocky Mountain and Pacific coast regions, extending north to about latitude 64° N. in the Yukon Territory and south to about latitude 31° N. in Baja California, and west to east from the Pacific Ocean to the Black Hills of South Dakota. Forests dominated by lodgepole pine cover some 6 million ha (15 million acres) in the Western United States and some 20 million ha (50 million acres) in Canada.


- The native range of lodgepole pine.

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Brief Summary

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Pinaceae -- Pine family

James E. Lotan and William B. Critchfield

Lodgepole pine (Pinus contorta) is a two-needled pine of the subgenus Pinus. The species has been divided geographically into four varieties: P. contorta var. contorta, the coastal form known as shore pine, coast pine, or beach pine; P. contorta var. bolanderi, a Mendocino County White Plains form in California called Bolander pine; P. contorta var. murrayana in the Sierra Nevada, called Sierra lodgepole pine or tamarack pine; and P. contorta var. latifolia, the inland form often referred to as Rocky Mountain lodgepole pine or black pine. Although the coastal form grows mainly between sea level and 610 m (2,000 ft), the inland form is found from 490 to 3660 m (1,600 to 12,000 ft).

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Physical Description

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Tree, Shrub, Evergreen, Monoecious, Habit erect, Trees without or rarely having knees, Tree with bark rough or scaly, Young shoots 3-dimensional, Buds resinous, Leaves needle-like, Leaves alternate, Needle-like leaf margins finely serrulate (use magnification or slide your finger along the leaf), Leaf apex acute, Leaf apex obtuse, Leaves < 5 cm long, Leaves > 5 cm long, Leaves < 10 cm long, Leaves yellow-green above, Leaves yellow-green below, Leaves not blue-green, Needle-like leaves triangular, Needle-like leaves twisted, Needle-like leaf habit erect, Needle-like leaves per fascicle mostly 2, Needle-like leaf sheath persistent, Twigs glabrous, Twigs viscid, Twigs not viscid, Twigs without peg-like projections or large fascicles after needles fall, Berry-like cones orange, Woody seed cones < 5 cm long, Woody seed cones > 5 cm long, Seed cones bearing a scarlike umbo, Umbo with missing or very weak prickle, Umbo with obvious prickle, Bracts of seed cone included, Seeds red, Seeds brown, Seeds black, Seeds winged, Seeds unequally winged, Seed wings prominent, Seed wings equal to or broader than body.
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Pinus contorta

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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. It is common near the ocean shore and in dry montane forests to the subalpine, but is rare in lowland rain forests. Like all pines (member species of the genus Pinus), it is an evergreen conifer.

Subspecies

There are four subspecies of Pinus contorta, and one of them is sometimes considered to have two varieties.[3] The subspecies are sometimes treated at the rank of variety.[2][4][5]

Description

Depending on subspecies, Pinus contorta grows as an evergreen shrub or tree. The shrub form is krummholz and is approximately 1 to 3 meters (3 to 10 ft) high. The thin and narrow-crowned tree can grow 40 to 50 m (130 to 160 ft) high and achieve up to 2 m (7 ft) in diameter at chest height.[4] 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.[17][18][19]

The species name is contorta because of the twisted, bent pines found at coastal areas (shore pine)[20] and the tree's twisted needles.[8][21][22] Pinus contorta is occasionally known under several English names: black pine, scrub pine, and coast pine.[23][24] 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.[20] Shore pine bark is somewhat thick and corky, fissuring into a checkered pattern.[20] Some lodgepole pines have been reported in low elevations with features closer to those of the shore pine, including the bark.[20]

Tamarack pine can grow up to centuries old and lodgepole pines in Yellowstone Park have survived over 300 years.[20]

Foliage

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The needles are 4 to 8 cm (1+12 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 millimeters (34 and 1+14 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 centimeters (1+12 to 3 in) long and 0.9 to 2 mm (132 to 332 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.[18] The foliage of lodgepole pine is yellow-green as compared to shore pine, which is dark green.[20]

Cones

The cones of lodgepole and shore pine begin to be produced when the trees are about ten years old.[20] The cones are 3–7 cm (1–3 in) long, with prickles on the scales.[20]

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.[25] The variation in their serotiny has been correlated with wildfires and mountain pine beetle attacks.[26] The cones of the coastal Pacific subspecies, P. contorta subsp. contorta, are typically non-serotinous,[24] and those of the inland Pacific subspecies, P. contorta subsp. murrayana, are completely non-serotinous.[27]

Distribution

Pinus contorta occurs from upper, dry montane forests to the subalpine region of western North America.[28][29][17] 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.[20] 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.[20] Further south, the species can be found in higher elevations up to 3,350 m (10,990 ft) above sea level, particularly in southern Colorado.[20] It is rare in lowland rain forests.[29][17] Shore pine can be found in very infertile soils in coastal regions from Southeast Alaska to Northern California.[20]

Lodgepole and shore pine can be found intermingled (and apparently hybridized) north of Puget Sound.[20] Less dependent on fire, tamarack pine can be found in California's upper mountains and mingled with lodgepole in Oregon.[20] Pinus contorta can be found in the closed-cone pine forest of coastal California.

Ecology

Pinus contorta is a fire-dependent species, requiring wildfires to maintain healthy populations of diverse ages. The thin bark of the lodgepole pine minimizes its defense to fire, although 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.[30] It otherwise fares poorly in crowds as other species are more shade tolerant, although some (e.g. firs) are more susceptible to fire.[20]

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

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.[30]

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.[31]

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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.[32][33]

Porcupines consume the inner bark of lodgepole pine.[20]

Threats

Larger members of the species are attacked by mountain pine beetle, which it fights with pitch but can be overwhelmed.[20] 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.[20] Exceptional cold can kill some of the beetles.[20]

A study released in 2011 concluded that Pinus contorta could experience significant reductions in distribution due to climate change by the late 21st century.[34][35]

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.

Uses

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

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.[20] Lodgepole pine was used by European settlers to build log cabins.[20] Logs are still used in rural areas as posts, fences, lumber, and firewood.[20] Shore pine pitch has historically been used as glue.[20]

Tree plantations of Pinus contorta have been planted extensively in Norway, Sweden, Ireland and the UK for forestry, such as timber uses. In Iceland it is used for reforestation and afforestation purposes.[36] 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.[20]

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.[37] The gum of shore pine was used medicinally as well as for chewing.[20]

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.[38][39]

Emblem

Lodgepole pine is the provincial tree of Alberta, Canada.

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. Retrieved 11 November 2021.
  2. ^ a b c "Pinus contorta". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA).
  3. ^ Conifer Specialist Group (1998). "Pinus contorta var. bolanderi". IUCN Red List of Threatened Species. 1998. Retrieved 12 May 2006.old-form url
  4. ^ a b Kral, Robert (1993). "Pinus contorta". In Flora of North America Editorial Committee (ed.). Flora of North America North of Mexico (FNA). Vol. 2. New York and Oxford. Retrieved 12 September 2010 – via eFloras.org, Missouri Botanical Garden, St. Louis, MO & Harvard University Herbaria, Cambridge, MA.
  5. ^ "Pinus contorta". World Checklist of Selected Plant Families (WCSP). Royal Botanic Gardens, Kew – via The Plant List.
  6. ^ Jepson Flora Project (ed.). "Pinus contorta subsp. bolanderi". Jepson eFlora. The Jepson Herbarium, University of California, Berkeley.
  7. ^ "Pinus contorta var. bolanderi". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA).
  8. ^ a b c Farjon, Aljos (2010). A Handbook of the World's Conifers. Vol. 2. Leiden, Netherlands: Koninklijke Brill NV. pp. 654–655. ISBN 978-90-04-17718-5.
  9. ^ "Pinus contorta ssp. contorta". Calflora. Berkeley, California: The Calflora Database.
  10. ^ "Pinus contorta var. contorta". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA).
  11. ^ 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.
  12. ^ "Pinus contorta var. murrayana". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA).
  13. ^ "Pinus contorta ssp. murrayana". Calflora. Berkeley, California: The Calflora Database.
  14. ^ "Pinus contorta var. latifolia". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA).
  15. ^ USDA, NRCS (n.d.). "Pinus contorta". The PLANTS Database (plants.usda.gov). Greensboro, North Carolina: National Plant Data Team. Retrieved 24 January 2015.
  16. ^ 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.
  17. ^ a b c Giblin, David, ed. (2015). "Pinus contorta". WTU Herbarium Image Collection. Burke Museum, University of Washington. Retrieved 24 January 2015.
  18. ^ a b Schütt, Weisgerber; Schuck, Lang; Stimm, Roloff (2008). Lexikon der Nadelbäume. Hamburg, Germany: Nikol. pp. 365–367. ISBN 978-3-933203-80-9.
  19. ^ 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.
  20. ^ a b c d e f g h i j k l m n o p q r s t u v w x y Arno, Stephen F.; Hammerly, Ramona P. (2020) [1977]. Northwest Trees: Identifying & Understanding the Region's Native Trees (field guide ed.). Seattle: Mountaineers Books. pp. 61–70. ISBN 1-68051-329-X. OCLC 1141235469.
  21. ^ "Plants and Trees: lodgepole pine". U.S. Forest Service. Retrieved 12 August 2014.
  22. ^ "Pinus contorta var. contorta : Shore Pine". Oregon State University. Retrieved 12 August 2014.
  23. ^ "Forests of Crater Lake National Park: Lodgepole Pine (Pinus Contorta)". Crater Lake Institute. Retrieved 12 August 2014.
  24. ^ a b 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.
  25. ^ 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.
  26. ^ 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.
  27. ^ 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.
  28. ^ 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.
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Pinus contorta: Brief Summary

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

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