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Image of Pinus flexilis var. reflexa Engelm.
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Limber Pine

Pinus flexilis E. James

Associations

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Foodplant / parasite
erumpent aecium of Cronartium ribicola parasitises stem of Pinus flexilis
Remarks: season: 3-6
Other: major host/prey

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Comments

provided by eFloras
Pinus flexilis , much branched with a strongly tapering trunk, is little utilized because of its form and relative inaccessibility. It reportedly forms intermediates with P . strobiformis where the two overlap. The fresh-cut wood has the odor of turpentine.
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Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
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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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Trees to 26m; trunk to 2m diam., straight to contorted; crown conic, becoming rounded. Bark gray, nearly smooth, cross-checked in age into scaly plates and ridges. Branches spreading to ascending, often persistent to trunk base; twigs pale red-brown, puberulous (rarely glabrous), slightly resinous, aging gray, smooth. Buds ovoid, light red-brown, 0.9--1cm, resinous; lower scales ciliolate along margins. Leaves 5 per fascicle, spreading to upcurved and ascending, persisting 5--6 years, 3--7cm ´ 1--1.5mm, pliant, dark green, abaxial surface with less conspicuous stomatal bands than adaxial surfaces, adaxial surfaces with strong, pale stomatal bands, margins finely serrulate, apex conic-acute to acuminate; sheath 1--1.5(--2)cm, shed early. Pollen cones broadly ellipsoid-cylindric, ca. 15mm, pale red or yellow. Seed cones maturing in 2 years, shedding seeds and falling soon thereafter, spreading, symmetric, lance-ovoid before opening, cylindro-ovoid when open, 7--15cm, straw-colored, resinous, sessile to short-stalked, apophyses much thickened, strongly cross-keeled, umbo terminal, depressed. Seeds irregularly obovoid; body 10--15mm, brown, sometimes mottled darker, wingless or nearly so. 2 n =24.
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Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
bibliographic citation
Flora of North America Vol. 2 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
source
Flora of North America @ eFloras.org
editor
Flora of North America Editorial Committee
project
eFloras.org
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eFloras

Habitat & Distribution

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High montane forests, often at timberline; (1000--)1500--3600m; Alta., B.C.; Ariz., Calif., Colo., Idaho, Mont., Nebr., Nev., N.Mex., N.Dak., Oreg., S.Dak., Utah, Wyo.
license
cc-by-nc-sa-3.0
copyright
Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
bibliographic citation
Flora of North America Vol. 2 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
source
Flora of North America @ eFloras.org
editor
Flora of North America Editorial Committee
project
eFloras.org
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visit source
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eFloras

Synonym

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Apinus flexilis (E.James) Rydberg
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copyright
Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA
bibliographic citation
Flora of North America Vol. 2 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
source
Flora of North America @ eFloras.org
editor
Flora of North America Editorial Committee
project
eFloras.org
original
visit source
partner site
eFloras

Broad-scale Impacts of Plant Response to Fire

provided by Fire Effects Information System Plants
More info for the terms: fuel, fuel moisture, prescribed fire, woodland

In 1976 spring prescribed burning was conducted in open-canopy limber pine stands in the Little Belt Mountains of central Montana at about 5,500 feet (1,675 m) [55]. Further general site descriptions appear in Keown 1982 [56]. Air temperatures ranged from 55 to 65 degrees Fahrenheit (13-18
oC). Relative humidity was 20% to 40%, and winds were calm to 25
miles per hour (40 km/h). Fuel moisture was 7%. The management objective was to improve understory browse and forage.
Limber pine mortality at postfire year 1 was 20% in grassy stands and as high as 80% in shrubby
stands [55].

The Research Project Summary Response of vegetation to prescribed burning in a Jeffrey pine-California
black oak woodland and a deergrass meadow at Cuyamaca State Park, California
, provides information on prescribed
fire and postfire responses of many plant community species including limber pine.

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bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Common Names

provided by Fire Effects Information System Plants
limber pine

Rocky Mountain white pine
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bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Cover Value

provided by Fire Effects Information System Plants
More info for the term: cover

The degree to which limber pine provides cover for wildlife species is as follows [26]:
CO MT ND UT WY
Pronghorn ---- ---- Poor Poor Fair
Elk ---- ---- ---- Good ----
Mule deer ---- ---- ---- Good ----
White-tailed deer ---- ---- Good ---- Good
Small mammals Good ---- ---- Good Good
Small nongame birds Good ---- Good Good Good
Upland game birds ---- Good ---- Good Good
Waterfowl ---- ---- ---- ---- Poor
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bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Description

provided by Fire Effects Information System Plants
Limber pine is a slow growing, long-lived species, sometimes taking several hundred years to reach maturity [26,75]. Mature trees may
exceed 1000 years of age [53,96,127]. Limber pine stands are broadly even-aged [89], though populations also occur in uneven-aged stands
and on very harsh sites as widely spaced, isolated individuals [96,102,112]. Trees often have an irregular or
multi-stem growth form, and rarely reach over 50 feet (15 m) [11,120,127]. At high elevations they sometimes form krummholz [11,127]. Trunks may reach 6.5 feet (2 m) in diameter [41]. The species is cold and drought tolerant. Trees are ectomycorrhizal, have deep taproots,
and are very windfirm [33,120].



Cones of limber pine are cylindrical, 3 to 6 inches (8-15 cm) long. They release
their seeds if not preyed upon (see Regeneration Processes). The seeds are large
(7-12 mm long) and sometimes have a vestigial wing [21,63,74].
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bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Distribution

provided by Fire Effects Information System Plants
Limber pine occurs from Alberta and British Columbia south to California, Arizona, and New Mexico. It is scattered widely across the Great Basin in Utah,
Nevada, and into Colorado, Wyoming, and Montana. Isolated populations occur in
the Dakotas [65,69,86,104,106] and Nebraska [30,109].
The U.S. Geological Survey provides a distributional map of limber pine. The Whitebark and Limber Pine Information System
provides distributional information at the stand level.
license
cc-publicdomain
bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Fire Ecology

provided by Fire Effects Information System Plants
More info for the terms: fire regime, fire-return interval, fuel, series, woodland

The thin bark of young limber pine trees does not protect them from even
low-severity fires. Because the bark at the base of older trees is often 2 inches (5 cm) thick, these trees can withstand stem scorch from low-severity
fires. Terminal buds are somewhat protected from the heat associated
with crown scorch by the tight clusters of needles around them [1,29,53,85,127].

FIRE REGIMES:

Wildfires are less frequent in limber pine communities than in other conifer habitats because of limited productivity and fuel accumulation associated with poor soil development, short growing seasons, and late snowmelt [29,53,78,85,96,117,127].
Keeley and Zedler [53] categorized 38 pines within a series of 5 fire predictability regimes. They include limber pine among those pines growing in areas with very low site (and therefore fuel) productivity and unpredictable fire return intervals of up to 1000 years. Where enough biomass accumulates to carry fires, limber pine may be cached by Clark's nutcrackers and establish in burned sites previously dominated by other conifers [53,66].

Fire-return intervals for some associated communities or ecosystems in which limber pine occurs are listed below. Find further fire regime information for the plant communities in which this
species may occur by entering the species name in the FEIS home page under "Find FIRE REGIMES".

Community or Ecosystem Dominant Species Fire-Return Interval Range (years)


sagebrush steppe Artemisia tridentata/Pseudoroegneria spicata 20-70 [15]
mountain big sagebrush Artemisia tridentata var. vaseyana 20-60 [5,16]
Wyoming big sagebrush Artemisia tridentata var. wyomingensis 10-70 (40**) [119,126]
curlleaf mountain-mahogany* Cercocarpus ledifolius 13-1000 [6,95]
mountain-mahogany-Gambel oak scrub Cercocarpus ledifolius-Quercus gambelii

western juniper Juniperus occidentalis 20-70 
Rocky Mountain juniper Juniperus scopulorum

Engelmann spruce-subalpine fir Picea engelmannii-Abies lasiocarpa 35 to > 200 
pinyon-juniper Pinus-Juniperus spp.
whitebark pine* Pinus albicaulis 50-200 [15]
Rocky Mountain lodgepole pine* Pinus contorta var. latifolia 25-300+ [3,90]
Sierra lodgepole pine* Pinus contorta var. murrayana 35-200

Jeffrey pine Pinus jeffreyi 5-30
Rocky Mountain ponderosa pine* Pinus ponderosa var. scopulorum 2-10 
Arizona pine Pinus ponderosa var. arizonica 2-10 [15]
quaking aspen (west of the Great Plains) Populus tremuloides 7-120 [15,34,77]
mountain grasslands Pseudoroegneria spicata 3-40 (10)** [3]
Rocky Mountain Douglas-fir* Pseudotsuga menziesii var. glauca 25-100 
oak-juniper woodland (Southwest) Quercus-Juniperus spp. 15]


*fire-return interval varies widely; trends in variation are noted in the species review

**mean

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bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Fire Management Considerations

provided by Fire Effects Information System Plants
Fischer and Clayton [29] suggest that limber pine growing in open stands can be
maintained by periodic fires that reduce the undergrowth. Where limber
pine and Douglas-fir codominate, fire can be a thinning agent that slightly
favors limber pine over Douglas-fir in the younger age
classes.
license
cc-publicdomain
bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Growth Form (according to Raunkiær Life-form classification)

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More info on this topic.

More info for the term: phanerophyte

RAUNKIAER [88] LIFE FORM:




Phanerophyte


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bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Habitat characteristics

provided by Fire Effects Information System Plants
More info for the terms: cover, grassland, litter, tree

Limber pine grows across a wider range of elevations that any other tree species in the central Rocky Mountains [94], inhabiting some of the driest sites capable of supporting trees [11,85,102,111]. In many high-elevation sites it occupies or forms the upper treeline [30,69,83,94,122], but in
northern parts of its range it is found at low elevations along plains grassland
edges [22,94,96]. It typically occurs on steep, rocky, well-drained, windswept, and nutrient-poor sites on exposed ridges and summits [1,10,11,32,42]. Limber pine is often reported growing on calcareous soil [11,19,85]. It is also reported on soils derived from many other types of parent material [12,17,23,42,68,101].




Ground cover and litter accumulation in limber pine stands are often sparse, accumulating only under individual trees [11,127]. Severe sheet erosion of fine particles often occurs from summer convection storms over sparsely vegetated sites. Snowpack accumulations on limber pine sites may be light as a result of high insolation and winter winds [127].



Site preference often separates limber pine and whitebark pine, which is ecologically similar in many respects [112]. Limber pine has a wider geographical distribution and altitudinal range than whitebark pine. Relative to whitebark pine, limber pine occurs on warm, dry sites at
low and middle elevations. Where their ranges overlap, the 2 species sometimes grow
together on droughty soils. Occasionally, limber pine grows at higher elevations than whitebark pine. South of the range of whitebark pine in California, Colorado, Nevada, and southern Wyoming, the more drought-resistant limber pine replaces whitebark pine and may form the alpine treeline [74,112,121].



Elevations reported in the literature for limber pine are as follows:


7,500 to 11,000 feet (2,290-3,350 m) in California [80]

5,000 to 12,500 feet (1,500-3,800 m) in Colorado [27,42,49]

4,000 to 6,000 feet (1,200-1,800 m) in Montana [85,91]

6,500 to 11,500 feet (2,000-3,500 m) in Nevada [114]

5,000 to 7,000 feet (1,500-2,100 m) in Oregon [19]

6,000 to 11,600 feet (1,830-3,540 m) in Utah [123]

license
cc-publicdomain
bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Habitat: Cover Types

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More info on this topic.

This species is known to occur in association with the following cover types (as classified by the Society of American Foresters):

More info for the term: cover

SAF COVER TYPES [28]:





206 Engelmann spruce-subalpine fir

208 Whitebark pine

209 Bristlecone pine

210 Interior Douglas-fir

217 Aspen

218 Lodgepole pine

219 Limber pine

220 Rocky Mountain juniper

237 Interior ponderosa pine

239 Pinyon-juniper

256 California mixed subalpine

license
cc-publicdomain
bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Habitat: Ecosystem

provided by Fire Effects Information System Plants
More info on this topic.

This species is known to occur in the following ecosystem types (as named by the U.S. Forest Service in their Forest and Range Ecosystem [FRES] Type classification):

More info for the term: shrub

ECOSYSTEMS [31]:





FRES20 Douglas-fir

FRES21 Ponderosa pine

FRES23 Fir-spruce

FRES26 Lodgepole pine

FRES29 Sagebrush

FRES34 Chaparral-mountain shrub

FRES35 Pinyon-juniper

license
cc-publicdomain
bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Habitat: Plant Associations

provided by Fire Effects Information System Plants
More info on this topic.

This species is known to occur in association with the following plant community types (as classified by Küchler 1964):

More info for the terms: forest, woodland

KUCHLER [60] PLANT ASSOCIATIONS:





K008 Lodgepole pine-subalpine forest

K011 Western ponderosa forest

K012 Douglas-fir forest

K015 Western spruce-fir forest

K017 Black Hills pine forest

K018 Pine-Douglas-fir forest

K019 Arizona pine forest

K020 Spruce-fir-Douglas-fir forest

K021 Southwestern spruce-fir forest

K022 Great Basin pine forest

K023 Juniper-pinyon woodland

K033 Chaparral

K037 Mountain-mahogany-oak scrub

K038 Great Basin sagebrush

K046 Desert: vegetation largely lacking

license
cc-publicdomain
bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Habitat: Rangeland Cover Types

provided by Fire Effects Information System Plants
More info on this topic.

This species is known to occur in association with the following Rangeland Cover Types (as classified by the Society for Range Management, SRM):

More info for the terms: cover, shrubland, woodland

SRM (RANGELAND) COVER TYPES [99]:




109 Ponderosa pine shrubland

209 Montane shrubland

210 Bitterbrush

402 Mountain big sagebrush

412 Juniper-pinyon woodland

413 Gambel oak

415 Curlleaf mountain-mahogany

504 Juniper-pinyon pine woodland


license
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bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Immediate Effect of Fire

provided by Fire Effects Information System Plants
Limber pine is often killed by fire because of its relatively thin bark. Keeley and Zedler [53] argue that the lack of evolution of thick, fire-resistant bark in this species is a result of very long and unpredictable fire return intervals in the unproductive sites where it occurs. The degree of stem scorch usually determines the extent of fire injury to trees. Young trees are usually killed by any fire that scorches their stems. Mature trees with thicker bark can survive [29]. The vulnerability of this species to fire is reduced by the open stand structure, sparse fuels, and sparse undergrowth of limber pine communities [85,101].
license
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bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Importance to Livestock and Wildlife

provided by Fire Effects Information System Plants
The large, wingless seeds of limber pine have high energy content.
Pine "nuts" provide critical food for rodents and birds, which cache the
seeds for later use. Other small mammals and birds benefit from these caches. Bears also feed from caches
[62,66]. Sites with limber pine provide key winter range for deer and elk [85].
Bighorn sheep use open stands on ridges. Difficult access and low grass production result in low forage
value of limber pine stands for livestock [1,42].

license
cc-publicdomain
bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Key Plant Community Associations

provided by Fire Effects Information System Plants
More info for the terms: association, herbaceous, shrub, shrubs, tree

Plant community associates of limber pine are described below by state.



California: In the Sierran subalpine, limber pine grows in association with Sierra lodgepole pine (Pinus contorta var. murrayana), bush chinquapin (Chrysolepis sempervirens), greenleaf manzanita (Arctostaphylos patula), curlleaf mountain-mahogany (Cercocarpus ledifolius), and whitethorn ceanothus (Ceanothus cordulatus) [81]. In montane areas of southern California, limber pine is reported with white fir (Abies concolor), Jeffrey pine (P. jeffreyi), singleleaf pinyon (P. monophylla), Sierra lodgepole pine, whitebark pine (P. albicaulis), foxtail pine (P. balfouriana), Great Basin bristlecone pine (P. longaeva), western juniper (Juniperus occidentalis),
curlleaf mountain-mahogany, and big sagebrush (Artemisia tridentata) [68,108].




Colorado: Tree associates include interior ponderosa pine (P. ponderosa var. scopulorum), Rocky Mountain lodgepole pine (P. c. var. latifolia), subalpine fir (A. lasiocarpa), Engelmann spruce (Picea engelmannii), Rocky Mountain Douglas-fir (Pseudotsuga menziesii var. glauca), white fir, whitebark pine, Rocky Mountain bristlecone pine (P. aristata), and quaking aspen (Populus tremuloides). Associated understory species include bearberry (Arctostaphylos uva-ursi), common juniper (J. communis), big sagebrush, purple pinegrass (Calamagrostis purpurascens), spike fescue (Leucopoa kingii), and Thurber fescue (Festuca thurberi) [27,38,42,49,87,100].




Idaho: In Craters of the Moon National Monument, limber pine is commonly associated with antelope bitterbrush (Purshia tridentata), rubber rabbitbrush (Chrysothamnus nauseosus), and mountain big sagebrush (A. t. var. vaseyana). Associated grasses include Sandberg bluegrass (Poa secunda), bottlebrush squirreltail (Elymus elymoides), and Indian ricegrass (Achnatherum hymenoides) [12,23]. In the mountains of east-central Idaho, limber pine grows in association with Douglas-fir, subalpine fir, and whitebark pine [17].



Montana: Associated tree and shrub species include Rocky Mountain Douglas-fir,
Rocky Mountain lodgepole pine, Engelmann spruce, whitebark pine, subalpine fir, quaking aspen, common juniper, creeping juniper (J. horizontalis), and Rocky Mountain juniper (J. scopulorum). Associated shrubs include snowberry (Symphoricarpos spp.), Wood's rose (Rosa woodsii), and russet buffaloberry (Shepherdia canadensis). Associated grasses include Idaho fescue (Festuca idahoensis), rough rescue (F. altaica), and bluebunch wheatgrass (Pseudoroegneria spicata) [5,55,83,85,91,107].



New Mexico: In the Sandia Mountains limber pine occurs with Rocky Mountain Douglas-fir, white fir,
quaking aspen, Engelmann spruce, and corkbark fir (A. l. var. arizonica) [7]. It co-occurs with southwestern white pine (P. strobiformis) in the Sangre de Cristo Mountains [116].



Nevada and Utah: Limber pine commonly occurs in association with Rocky Mountain bristlecone, interior
ponderosa (P. ponderosa var. scopulorum), Rocky Mountain lodgepole, whitebark, Jeffrey, and singleleaf pinyon pines. It also occurs with Engelmann spruce, white fir, subalpine fir, quaking aspen, common juniper, and Utah juniper (J. osteosperma). Associated shrubs include Rocky mountain maple (Acer glabrum), Gambel oak (Quercus gambelii), and multiple species of sagebrush, mountain-mahogany, ceanothus, currant (Ribes spp.), manzanita (Arctostaphylos spp.), and snowberry [9,37,82,114].



Oregon: In the Wallowa Mountains of eastern Oregon limber pine commonly occurs with Douglas-fir and Rocky Mountain juniper. Herbaceous associates include western yarrow (Achillea millefolium), sagebrush fleabane (Erigeron austiniae), and silverleaf phacelia (Phacelia hastata) [19].



South Dakota: A population of limber pine in the Black Hills is associated with
interior ponderosa pine and white spruce (Picea glauca). Understory species include bearberry and common juniper [50].



Wyoming: Associated species reported for northwestern Wyoming include Rocky
Mountain lodgepole pine, Engelmann spruce, whitebark pine, Rocky Mountain Douglas-fir, subalpine fir, Rocky Mountain juniper, and common juniper [13,24,70].



Published classifications that include limber pine as an indicator or
dominant species are presented below:



Arizona [67,79]

California [51,92]

Colorado [8,20,25,42,49,57]

Idaho [102,103]

Montana [85,91]

North Dakota [32]

New Mexico [20,25,67,79]

Utah [41,73,127]

Wyoming [1,102,124]
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bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Life Form

provided by Fire Effects Information System Plants
More info for the term: tree

Tree

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bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Management considerations

provided by Fire Effects Information System Plants
More info for the term: cone

Management of limber pine forests associated with Douglas-fir typically
favors the growth of the economically important Douglas-fir. Of
primary management importance in these areas is watershed protection and
enhancement. The slow rate of vegetation recovery in areas where limber pine occurs requires dispersed,
low-impact recreation to maintain the aesthetic appeal of these forests [25]. Forage
productivity can be increased by periodic surface fires [29].



Limber pine trees are infected and killed by white pine blister rust (Cronartium ribicola) throughout
the tree's range. Ribes species are obligate alternate hosts of the rust [46,47,48,76,116]. Limber pine appears to have less resistance to blister rust than other North American white
pines (Strobi), with greenhouse infection levels as high as 98
to 100% [45,113]. In a 3-year greenhouse study of relative seedling
susceptibility to blister rust, limber pine mortality was 75% (n=348). In comparison, mortality in whitebark pine was 33%
(n=207) and 86% in southwestern white pine (n=323) [45].



Limber pine is susceptible to numerous other fungal diseases [101]. It can be heavily infected or killed by limber pine dwarf-mistletoe (Arceuthobium cyanocarpum)
[12,39,71,72], and is susceptible to infestation by mountain pine beetles, cone beetles, coneworms, and budworms [54,61,101].



The Whitebark and Limber Pine Information System
provides a database for storing and analyzing data on site characteristics,
stand structure, regeneration, and mortality and infection rates from white pine blister rust and
other damaging agents.
license
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bibliographic citation
Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Nutritional Value

provided by Fire Effects Information System Plants
Limber pine browse is rated fair in energy value and poor in protein value [26].
The seeds are highly nutritious, providing amino acids, lipids, and averaging
7,178 calories per gram [64].


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Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Occurrence in North America

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AZ CA CO ID MT NE NV
NM ND OR SD UT WY
AB BC

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Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Other uses and values

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Limber pine is used in the nursery trade
for landscaping [35,36,40].


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Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Palatability

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Although limber pine browse is unpalatable to large mammals, it
provides some food for birds and small mammals. The palatability of limber pine for livestock and wildlife has been rated as follows
[26]:
CO MT ND UT WY
Cattle Poor Poor Poor Poor Poor
Domestic sheep Poor Poor Poor Poor Poor
Horses Poor Poor Poor Poor Poor
Pronghorn ---- ---- Poor Poor Poor
Elk Poor Poor ---- Poor Fair
Mule deer Poor Poor Poor Poor Fair
White-tailed deer ---- ---- Poor ---- Fair
Small mammals ---- ---- ---- Good Good
Small nongame birds ---- ---- Poor Good Good
Upland game birds ---- ---- ---- Good Good
Waterfowl ---- ---- ---- ---- Poor
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Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Phenology

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More info for the term: phenology

Limber pine cones ripen from August to
September, and seeds are dispersed from September to October
[26,44,58,59]. Cones open in the fall. Observed dates for phenological events
of mature in limber pine east of the Continental Divide in Montana and Yellowstone National Park,
Wyoming, are given here [93]:



Shoots start: April 30 to June 6

Buds burst: April 30 to June 26

Pollen starts: June 20 to July 14

Pollen ends: July 4 to July 22

Shoots end: June 22 to August 5

Winter buds formed: June 11 to August 16

Cones full size: August 15 to August 16

Cones open (seed dispersal): August 23 to August 30



Also see Regeneration Processes regarding pollen phenology.
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Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Plant Response to Fire

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More info for the terms: forest, seed, tree

Regeneration of limber pine end Engelmann spruce was assessed in a high-altitude area disturbed by fire in 1905. The south-facing sites were on Niwot Ridge in Colorado's Roosevelt National Forest. The uppermost elevation of the burn reached slightly below the tree limit at about 11,000 feet (3,355 m). Although limber pine regeneration at the uppermost elevation is less than at lower elevations, limber pine colonization at all elevations began shortly after the fire. The author found no evidence that
treeline changed following the fire [98]. Postfire regeneration of limber pine
is a consequence of seed dispersal and caching by Clark's nutcrackers [53,66].

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Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Post-fire Regeneration

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More info for the terms: secondary colonizer, seed

POSTFIRE REGENERATION STRATEGY [105]:




Initial offsite colonizer (off-site, initial community)

Secondary colonizer (on-site or off-site seed sources)
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Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Regeneration Processes

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More info for the terms: fitness, grassland, meristem, mutualism, phenology, seed, tree

Limber pine reproduces entirely from seed; it does not layer lower branches in the soil [22,122].
Seeds are not effectively dispersed by wind. Small mammals and birds, especially Clark's nutcrackers and pinyon jays,
disperse limber pine seeds
[63,64,66,110,125]. The minimum seed-bearing age of limber pine ranges from 20 to 40 years.
There are 2 to 4 years between large seed crops [58,59,101]. Seeds from krummholz trees have
low germination potential [66]. 


Clark's nutcrackers have co-adapted an important mutualism with limber pine
and are the primary harvester and disperser of its seeds. Limber pine regeneration
on burns is largely from germinants of Clark's nutcrackers seed caches [63,64,66,110,125].
The birds begin harvesting seeds in late August, while the cones are still green and slightly closed. They remove the cones by pecking them loose,
fly them to perches, and peck between the scales to remove the seeds. As cones begin to open on the trees in September, Clark's nutcrackers remove exposed seeds.
An individual bird can store as many as 125 seeds in its sublingual pouch, then
flies to a cache area and deposits numerous caches from its pouchful of seeds. In a burned-over area in northern Utah,
Clark's nutcrackers cached an estimated 12,140 seeds per acre (30,000/ha) in 1 year [62,101,112]. 



Mating system: Limber pine seed dispersal by corvids leads to a genetic population structure different from that of wind-dispersed conifers with respect to patterns of gene flow and genetic relationships among neighboring trees. The seed caching
by birds influences the distribution, population age structure, and spacing of limber pine. Clusters of seedlings germinating from a single cache may generate
multi-stemmed growth forms that contain 2 or more distinct genotypes. A consequence of this growth form is
a tendency toward clumped stand structure. Because seeds within an individual
cache were often collected from a single parent tree, trees within clumps may be
more closely related compared to trees from neighboring clumps [64,110,113],
although multi-stemmed growth is most often a result of apical meristem damage
that results in several leaders on an individual tree [123]. Tomback and Linhart
[112] found that on 361 limber pine sites in Colorado, 30% showed clumping.
Several genetic studies have shown that from 0 to 82% of  individuals
within limber pine clumps are closely related [101,117,123]. On the Pawnee
National Grassland, clump members were related, on average, as nearly half-sibs.
Genetic consequences of this kinship include possible inbreeding. On the plus
side, closely related trees within clumps often form roots grafts, which may
increase survivorship and fitness of the entire clump [123].



Pollen phenology also influences gene flow. In Colorado, most sites that differ in elevation by more than 1,300 feet (400 m) in elevation do not have overlapping pollination periods, restricting
pollination between populations that are widely separated by elevation; however,
pollen transfer between intermediate populations and a high level of gene flow via
bird-dispersed seeds appear to maintain interpopulation gene flow [97].

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Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Regional Distribution in the Western United States

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This species can be found in the following regions of the western United States (according to the Bureau of Land Management classification of Physiographic Regions of the western United States):

BLM PHYSIOGRAPHIC REGIONS [14]:





4 Sierra Mountains

5 Columbia Plateau

6 Upper Basin and Range

7 Lower Basin and Range

8 Northern Rocky Mountains

9 Middle Rocky Mountains

10 Wyoming Basin

11 Southern Rocky Mountains

12 Colorado Plateau

15 Black Hills Uplift

16 Upper Missouri Basin and Broken Lands

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Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Successional Status

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More info for the terms: climax, seed, succession, xeric

According to Tomback and Linhart [112] limber pine (and whitebark pine) "are pioneering species that are either seral or topoedaphic climax species under different environmental conditions. In fact, seed dispersal by Clark's nutcrackers to outlying sites, treeline, and other harsh environments essentially increases the ecological niche breadth (in the Hutchinsonian sense) of these species. Clark's nutcrackers can maintain climax communities, colonize previously unforested sites, or initiate succession."



The later stages of succession in xeric subalpine forests vary due to differences in sites and seed availability. In the Colorado subalpine, Rebertus and others [89] studied conifer population age structure and succession on 3 burns greater than 100 years old. The sequence of conifer colonization appeared to be consistent: 1st limber pine, then Engelmann spruce, and later subalpine fir, with a delay between the 1st limber pine and
later subalpine fir of as long as 140 years. The authors suggested that the early advantage of limber pine was due to avian seed dispersal and exceptional drought tolerance in seedlings. Spatial analysis
suggested that limber pine facilitated the establishment of the other 2 species by providing shade or wind protection. On the xeric to slightly xeric sites, limber pine formed broadly even-aged, non-regenerating populations that were gradually replaced by the spruce and fir. On the most extreme sites, limber pine formed all-aged, self-maintaining populations with no evidence of replacement by the other species. The authors note that in lower elevation stands along the Front Range, limber pine is successional to Douglas-fir. In the even lower Pawnee National Grasslands of Colorado, limber pine forms all-aged, self-replacing populations. "Hence, many successional pathways could be operating at different sites or stages in stand development."

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Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Taxonomy

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The currently accepted scientific name of limber pine is Pinus flexilis James (Pinaceae). It is placed in
subgenus Strobus, subsection Strobi of Pinus
[2,21,30,43,44,52,104]. 


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Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Value for rehabilitation of disturbed sites

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More info for the terms: reclamation, restoration, seed, stratification

Because of its slow growth, limber pine has been used only to a limited extent in land reclamation projects [120]. Vegetation
recovery is slow on the exposed, hot, dry, rocky sites where it is found, and
soil erosion can prevent complete restoration. However, limber pine's drought tolerance and ability to survive at high elevations indicate that it has potential for use in revegetation projects [104].



Grossnickle and Reid [33] tested the feasibility of including limber pine seedlings in the reclamation of a high-elevation mining site in Colorado. The site was a molybdenum tailing pond buried in deep mine waste rock. One-year-old containerized
limber pine, lodgepole pine, and Engelmann spruce seedlings were inoculated with 3 species of ectomycorrhizal fungi prior to outplanting. Because of greenhouse colonization of seedling roots by a 4th
"wild" strain of ectomycorrhizal fungus, no uncolonized seedlings were outplanted. Some of the seedlings were also treated in the field with fertilizer or sewage sludge combined with wood chips. All seedlings were protected from wind and sun with cedar shingles.
During the 4th growing season, significant (p = 0.05) differences in seedling height among the fungal treatments were detected. The addition of the sewage sludge/wood chip slurry improved seedling height
of all 3 species in 1 of the fungal treatments. At the end of the 4th growing season, overall survival of limber pine, lodgepole pine, and Engelmann spruce seedlings was
60%, 52% and 62%, respectively. The authors noted that in all instances, seedling mortality appeared to be caused by unfavorable soil and climatic conditions and not by pathogen, insect, or animal damage.



Limber pine cones may yield 1,100 to 1,300 cleaned seeds per pound, and stored seed has been shown viable for at least 5 years. Freshly collected seeds may germinate without pretreatment, but cold, moist stratification
of up to 90 days improves germination [59].

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Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Wood Products Value

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The wood of limber pine has little commercial value. Its potential for
timber and fuelwood production is low, because the trees are slow growing with
irregular form [1,2,20,42,49,85,96,101]. Limber pine has been used locally for
mine props and railroad ties [102].
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Johnson, Kathleen A. 2001. Pinus flexilis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.fed.us/database/feis/plants/tree/pinfle/all.html

Associated Forest Cover

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As well as being dominant in the forest cover type Limber Pine (Society of American Foresters Type 219), limber pine is a minor component of the following (32); Engelmann Spruce-Subalpine Fir (Type 206), Whitebark Pine (Type 208), Bristlecone Pine (Type 209), Interior Douglas-Fir (Type 210), Aspen (Type 217), Lodgepole Pine (Type 218), and Interior Ponderosa Pine (Type 237).

In Canada, Montana, and central Idaho, limber pine forms pure stands at lower tree line or mixes with Douglas-fir (Pseudotsuga menziesii), and to a lesser extent, ponderosa pine (Pinus ponderosa) and Rocky Mountain juniper (Juniperus scopulorum). It also appears as a minor component in stands of lodgepole pine (Pinus contorta), Engelmann spruce (Picea engelmannii), and occasionally subalpine fir (Abies lasiocarpa). On some sites in Idaho and Montana, it is associated with whitebark pine (Pinus albicaulis). In Canada, it is sometimes found with white spruce (Picea glauca).

Southward into Wyoming, southern Idaho, and northern portions of Colorado, Utah, and Nevada, limber pine may dominate windswept slopes and ridges at upper or lower tree line or appear in stands of white fir (Abies concolor), lodgepole pine, and Douglas-fir. In this region, limber pine appears most often with Engelmann spruce, subalpine fir, and quaking aspen (Populus tremuloides), least often with ponderosa pine. In Wyoming, limber pine occasionally coexists with whitebark pine, particularly in the Wind River Range. The two species also coexist on a few sites in northeastern Nevada (5,25), but usually where their ranges overlap they occupy different soils.

Farther south in the remainder of its range, limber pine forms open stands near upper tree line, both separately and with Great Basin bristlecone pine (Pinus longaeva) (44) but less often with Rocky Mountain bristlecone pine (Pinus aristata) (6,18). It is also associated with whitebark pine on the east side of the Sierra Nevada (4). Occasionally, it mixes as a minor seral species with subalpine fir and white fir (23). Where limber pine would normally mix as a seral species with other conifers, as it does farther north, the closely related southwestern white pine (Pinus strobiformis) appears in these situations, but this species does not extend onto the dry windy sites where limber pine is climax (23).

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Climate

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Climatic data for actual limber pine habitat are quite scarce, but the general distribution of limber pine in Alberta, Montana, central Idaho, and east of the Continental Divide in Wyoming and Colorado, is in forested areas having a continental climate (2,3). This climate is typified by a relatively small amount of precipitation, with the wettest months during the growing season, very low humidity, and wide annual and diurnal temperature ranges. Winter conditions may be very cold, but relatively dry, and often include rapid fluctuations in temperature associated with chinook winds. Notable exceptions to this distribution are the small populations in eastern Oregon and adjacent Idaho, which lie within the Pacific maritime influence (3).

In the remainder of its distribution, limber pine grows in climates that tend to have either more evenly distributed yearly precipitation or a winter peak in precipitation along with summer convectional storms. Throughout its broad range, limber pine is mostly absent in areas strongly influenced by Pacific maritime weather patterns. Only at its southern limits in the mountains of eastern and southern California (10) does the pine encounter a strong pattern of proportionately high winter precipitation (3). The amount of precipitation, however, is relatively smaller than that of the Pacific Northwest.

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Damaging Agents

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Fire can easily kill young limber pines because of their thin bark. Fuel loads on most limber pine sites are too light, however, to generate severe fire damage, and most of the large trees normally survive. Porcupines feed on limber pine, especially in the winter months (11). Several insects attack the pine in various ways. In Montana, the budworm (Choristoneura lambertiana ponderosana) feeds on the new needles of limber pine (37). A cone moth (Dioryctria spp.) is presumed to have damaged limber pine seed in North Dakota (29). In northern Idaho, the woolly aphid (Pineus coloradensis) attacked limber pine seedlings growing in test plots, but the pine showed considerable resistance to this insect (16). Mountain pine beetle (Dendroctonus ponderosae) also occasionally attacks limber pine (1).

Limber pine is susceptible to several major diseases. Spongy root and butt rot (Armillaria mellea) and the red-brown butt rot (Phaeolus schweinitzii) attack limber pine over much of its range. The crumbly brown cubical rot (Fomitopsis pinicola) and red ring rot (Phellinus pini) commonly cause heart rot in mature and damaged trees (15,27). Limber pine is susceptible to white pine blister rust (Cronartium ribicola) and can suffer considerable mortality when susceptible species of the rust's alternate host (Ribes) are nearby. The limber pine dwarf mistletoe (Arceuthobium cyanocarpum) is a common parasite of this tree. Occasionally, lodgepole pine dwarf mistletoe (A. americanum) attacks limber pine, and the Douglas-fir dwarf mistletoe (A. douglasii) and southwestern dwarf mistletoe (A. vaginatum subsp. cryptopodium) occur as rare parasites (12). Several foliage diseases also attack this tree, the most damaging being brown-felt snow mold (Neopeckia coulteri) (15).

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Flowering and Fruiting

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Limber pine is monoecious-male and female strobili are borne separately on the same tree. As with most pines, male strobili predominate in the lower crown and female strobili most often develop at the apical end of main branches in the upper crown. Male strobili emerge from buds in the spring and are arranged in small clusters of indistinct spirals. They may be green or yellow to reddish purple but turn brown when mature and about to shed their pollen. Pollen is shed during June and July. Female strobili emerge from buds shortly after the male strobili and are green or red to purple. Cone scales flex and they remain receptive to pollen for only a relatively short time during June and July. After pollination, scales close and the strobili begin to develop slowly.

Fertilization takes place in the spring or early summer, about 13 months after pollination. Cones and seeds mature rapidly following fertilization. As they mature, cones change color from green to lustrous yellow. They are light brown when mature in August and September. Seed dispersal takes place during September and October (41).

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Genetics

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Population Differences Genetic variation exists within limber pine in a general north-south pattern, but the range of variability for any one trait is small. Some isolated populations in Wyoming, Nebraska, and Colorado appear to be more similar to those from more southern latitudes than to populations at the same latitude (36).

Races Three possible races of limber pine have been suggested, distinguished by height growth of the seedlings: (1) a northern race ranging from Alberta to north central Colorado and northern Utah and including the only sample from California; (2) a southeastern race that includes populations from the Wyoming-Nebraska border, east central Colorado, and north central New Mexico; and (3) a southwestern race in southern Utah and western Colorado (Nevada populations were not sampled) (43). Further study, however, found no geographically associated patterns or trends when a much wider variety of characteristics was analyzed from the same seed sources (36).

Hybrids Although zones of intergradation between limber pine and southwestern white pine are found in north central Arizona and north central New Mexico (36), no true hybrid populations of limber pine have been recorded. Limber pine has been crossed artificially with western white pine (Pinus monticola), southwestern white pine (P. strobiformis), Mexican white pine (P. ayacahuite), Himalayan pine (P. griffithii), eastern white pine (P. strobus), and possibly whitebark pine (P. albicaulis) (35).

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Growth and Yield

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The slow growth rate and poor form normally attributed to limber pine discourages commercial interest in its use for timber and there is little information regarding its growth and yield. There apparently has been no attempt, however, to plant this species on forest sites superior to those chosen by birds and rodents that cache the seed. Where occasional limber pines grow in more densely forested stands with other tree species, sapling and pole size trees are often straight and single stemmed. One study (29) suggests, however, that limber pine growth rates may be greater on exposed windy knolls than on warmer south slopes and more moist north slopes.

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Reaction to Competition

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Limber pine is relatively intolerant of shade and therefore seral to most of its associated trees, the exceptions being quaking aspen, Rocky Mountain juniper, and possibly ponderosa pine. It is also considered seral to bristlecone pine and Douglas-fir but will codominate with these species on severe dry sites (23). As a result, on most forest sites, limber pine normally acts as a pioneer species following fire or tree removal. Except on the most severe sites, where trees remain widely spaced, limber pine shows little evidence of maintaining its population in the presence of other conifers. It is most accurately classed as a species intolerant of shade.

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Rooting Habit

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Type of substrate undoubtedly influences the rooting habit of limber pine. On many of the very rocky sites where it grows, the root system must follow the pattern of rock fracturing. As a result, most limber pine are quite wind firm. In nurseries, where there are better soil conditions, it develops a more uniform root system and can be transplanted by the ball and burlap method if previously root pruned (7). Roots of limber pine are also known to associate with a mycorrhizal fungus (Gomphidius smithii) (40).

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Seed Production and Dissemination

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Large seed crops are produced every 2 to 4 years and seed numbers generally range from 7,050 to 15,000/kg (3,200 to 6,800/lb) and average 10,800/kg (4,900/lb) (38). Seed size varies geographically, with a tendency for increasingly larger seeds in more southerly latitudes (14,36). Although some trees produce seed having an ineffective vestigial wing (21), most limber pine seeds are wingless.

The seeds are disseminated largely by rodents and birds. Of the birds, Clark's nutcracker is most important; it can transport pine seed for at least 23 km (14 mi) from seed source to communal caching areas (42). It can carry up to 125 limber pine seeds per trip in a sublingual pouch and buries in the ground one to five seeds per cache at a depth of 2 to 3 cm (0.8 to 1.2 in). Estimates indicate that Clark's nutcrackers cached in 1 year about 30,000 seeds per hectare (12,140/acre), most of which were limber pine (22). The birds' preferred cache sites were windswept ridges and southerly aspects where snow does not accumulate and the ground is exposed early in the spring. The locations of most limber pine stands probably reflect the site preferences of dispersal agents rather than those of the pine, since its only other apparent means of dissemination is gravity.

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Seedling Development

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Germination is epigeal (41). Like seed size, rate of seedling development depends on the geographic source. In one study (36), 2-year-old nursery grown seedlings from Alberta averaged 4.3 cm (1.7 in), while those from New Mexico had reached 7.4 cm (2.9 in). Fall-sown seed, properly fertilized, produced a pencil-sized 2-0 seedling suitable for field planting (14). In the wild, many seedlings develop in clusters from Clark's nutcracker seed caches (20). The seedlings withstand this competition well and often retain the clumped habit into maturity.

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Soils and Topography

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In some areas, limber pine grows in greater numbers on certain soils, but the relationships vary geographically. In general, the substrates are Entisols. In Montana, limber pine grows mainly on calcareous substrates (26). Similarly, in eastern Idaho and western Wyoming, it grows mainly on soils derived from limestone or sandstone and is notably absent on adjacent granitic substrates (34), yet the population in South Dakota grows on soils derived from granitic rock (38). In central Idaho, limber pine is found largely on soils derived from sedimentary rocks; it is notably absent on granitic substrates, but grows in cracks of recent lava at Craters of the Moon National Monument (33). In eastern Oregon, a recently discovered population is on soils derived from serpentine (17). In Utah, it grows on soils developed from limestone, as on the Wasatch Plateau (9), and on soils derived from quartzites, shales and limestones of the Uinta Range (26). In southern Utah, it is most common on soils derived from sandstone and limestone (44). In northeastern Nevada, it also grows on various calcareous substrates (25), but in California, on substrates derived from granitic, obsidian, and pumice materials (30).

Limber pine grows on a variety of topographies, from gently rolling terrain to cliffs. It is most often found on rocky ridges and steep rocky slopes and can survive in extremely windswept areas at both lower and upper tree line.

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Special Uses

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Limber pine is seldom sought for timber, but small quantities are occasionally harvested along with more desirable species. The wood has been used for rough construction, mine timbers, railroad ties, and poles (2).

Although of marginal value for lumber, limber pine has other resource values. Its ability to grow on harsh sites often provides the only tree cover for wildlife. The large seeds are a nutritious food source for birds, rodents, and bears and were used as food by Native Americans and early pioneers (20,21,29). In areas where timber is scarce, limber pine may be an important source of fuelwood. Increasing demands for fuelwood could deplete the accessible dead trees and eventually conflict with wildlife needs for shelter and nesting cavities.

Limber pine's abilities to withstand severe wind and dry site conditions are desirable shelterbelt traits, but its slow growth rate may discourage its selection for that purpose. Young trees, however, can withstand considerable bending, a necessary trait for reforestation of snow avalanche paths, and much of the pine's natural habitat lies within avalanche areas. Some limber pine habitats are also valuable watersheds, and as a pioneer species, the pine is a logical choice for initial site protection and for increasing snowpack (39). The pine's characteristic branching pattern also adds to the esthetic appeal of the landscape, especially along ridge lines.

This tree's ability to endure very dry environments has allowed it to attain considerable age in some areas. One tree in southern California was found to be well over 1,000 years old (13); another in central Idaho was 1,650 years old (31). This feature makes limber pine a useful species in dendrochronologic studies.

Limber pine has potential as a Christmas tree, but its qualities are surpassed by southwestern white pine (14,43). Seedlings from several seed sources have grown too slowly for economical Christmas tree operations but have ornamental value as dwarfed trees and even bonsai (14). Some bonsai nurserymen also collect dwarfed limber pine from severe windy sites. As an ornamental, this species deserves more attention than current use would indicate. The ornamental trade has selected at least seven cultivated varieties: 'Columnaris'- a fastigiate form; 'Glauca' and 'Firmament'- both with exceptionally bluish-green foliage; 'Glenmore'- with longer, more silvery foliage; 'Nana'- a dwarf bushy form; 'Pendula'- with pendulous branches; and 'Tiny Temple'- a low growing form (7,19).

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Distribution

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Limber pine grows from Alberta and southeastern British Columbia to New Mexico, Arizona, and eastern California. Notable outliers of this general distribution are found in the western portions of North Dakota, South Dakota, and Nebraska, and in eastern Oregon and southwestern California.

In the northern half of its distribution, limber pine is generally found near lower tree line and on dry sites in the montane forests. Between the 45th and 40th parallels, it grows in both lower and upper elevation forests and anywhere in between on dry, windswept sites. Its position gradually shifts upward in more southerly latitudes, so that in southern portions of its distribution, limber pine is more common from upper montane to alpine tree line, with only minor occurrences in the lower forested zones. Because of this adaptability, limber pine ranges in elevation from about 870 m (2,850 ft) in North Dakota (29) to about 3810 m (12,500 ft) in Colorado (7).


- The native range of limber pine.

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Brief Summary

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Pinaceae -- Pine family

Robert Steele

Limber pine (Pinus flexilis), also known as white pine or Rocky Mountain white pine, is a long-lived, slow-growing tree of small to medium size. Its wood, light in weight, close-grained, and pale yellow, is used for rough construction, mine timbers, railroad ties, and poles. Its harvest is incidental to that of other, more desirable species.

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Physical Description

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Tree, Evergreen, Monoecious, Habit erect, Trees without or rarely having knees, Primary plant stem smooth, Tree with bark smooth, 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, Leaves < 5 cm long, Leaves > 5 cm long, Leaves < 10 cm long, Leaves not blue-green, Leaves white-striped, Needle-like leaves triangular, Needle-like leaves somewhat rounded, Needle-like leaves not twisted, Needle-like leaf habit erect, Needle-like leaf habit drooping, Needle-like leaves per fascicle mostly 5, Needle-like leaf sheath early deciduous, Twigs pubescent, 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, Seed cones bearing a scarlike umbo, Umbo with missing or very weak prickle, Bracts of seed cone included, Seeds brown, Seeds wingless.
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Pinus flexilis

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Pinus flexilis, the limber pine, is a species of pine tree-the family Pinaceae that occurs in the mountains of the Western United States, Mexico, and Canada. It is also called Rocky Mountain white pine.

A limber pine in Eagle Cap Wilderness, Oregon, has been documented as over 2,000 years old, and another one was confirmed at 1,140 years old.[4][5] Another candidate for the oldest limber pine was identified in 2006 near the Alta Ski Area in Utah; called "Twister", the tree was confirmed to be at least 1,700 years old and thought to be even older.[6]

Description

Its pliant branches gives it the common name "limber" and specific epithet flexilis. Its needles are about 8 centimeters (3+14 in) long and a dark, blueish green.[7] Its bark is heavily creased and dark grey. Its pale wood is lightweight and soft.

Pinus flexilis is typically a high-elevation pine, often marking the tree line either on its own, or with whitebark pine (Pinus albicaulis), either of the bristlecone pines, or lodgepole pine (Pinus contorta). In favorable conditions, it makes a tree to 20 metres (65 feet), rarely 25 m (80 ft) tall. On exposed tree line sites, mature trees are much smaller, reaching heights of only 5–10 m (15–35 ft).[8] In steeply-sloping, rocky, and windswept terrain in the Rocky Mountains of southern Alberta, limber pine is even more stunted, occurring in old stands where mature trees are consistently less than 3 m (10 ft) in height.[9]

One of the world's oldest living limber pine trees grows on the banks of the upper North Saskatchewan River at Whirlpool Point in Alberta. Recent measurements give a maximum girth of 185". In 1986, a core sample 10 cm was retrieved by two researchers who counted 400 rings. Extrapolating this data gives an age close to 3,000 years.

Similar species

Pinus flexilis is a member of the white pine group, Pinus subgenus Strobus, and like all members of that group, the leaves ('needles') are in fascicles (bundles) of five,[7] with a deciduous sheath. This distinguishes it from the lodgepole pine, with two needles per fascicle, and the bristlecone pines, which share five needles per fascicle but have a semi-persistent sheath.

Pinus albicaulis

Distinguishing limber pine from the related whitebark pine (P. albicaulis), also a white pine, is very much more difficult, and can only easily be done by the cones. In limber pine, the cones are 6–15 cm (2+14–6 in) long[7] where the species overlap, green when immature, and open to release the seeds; the scales are not fragile. In whitebark pine, the cones are 4–7 cm (1+122+34 in) long, dark purple when immature, and do not open on drying, but are fragile and are pulled apart by birds to release the seeds.[10] A useful clue is that whitebark pines almost never have intact old cones lying under them, whereas limber pines usually do. [11]

Pinus monticola

In the absence of cones, limber pine can also be hard to tell from Western white pine (P. monticola) where they occur together in the northern Rockies and the Sierra Nevada east slope. The most useful clue here is that limber pine needles are entire (smooth when rubbed gently in both directions), whereas Western white pine needles are finely serrated (feeling rough when rubbed gently from tip to base). Limber pine needles are also usually shorter, 4–7 cm (1+122+34 in) long, while western white pine needles are 5–10 cm (2–4 in), though the ranges overlap.

Distribution

The largest part of the limber pine's range is in the Rocky Mountains, from southwest Alberta[7] and southeastern British Columbia south through Colorado and New Mexico into the northern states of Mexico. It is also found through the Great Basin[7] states of Nevada and Utah, in the eastern Sierra Nevada and White Mountains of Northern California, and in the San Bernardino and San Gabriel Mountains of the Transverse Ranges in Southern California. Continuing south the species is found in the San Jacinto Mountains, Santa Rosa Mountains, and Hot Springs Mountain of the Peninsular Ranges. There are small disjunct populations in eastern Oregon, in western North Dakota and Nebraska,[12] and in the Black Hills of South Dakota.[2][8] It is found at a wide range of altitudes depending on the latitude, from 850 to 3,810 m (2,790 to 12,500 ft). In the northern half of its range, it grows in the montane zone near the lower tree line; in the middle of its range between the 45th and 40th parallels, it grows on windswept sites in the montane and subalpine zones; and in the southern part of its range, it grows mainly at high elevations in the subalpine zone near the upper tree line.[12] It can more often be found at the outer fringes of a forest than in the forest itself.[7]

Ecology

Pinus flexilis is an important source of food for several species, including red squirrels and Clark's nutcrackers, the latter being an important distributor of seeds.[13][7] There is evidence that limber pines co-evolved with Clark's nutcrackers, which are the primary dispersers of the seeds.[14][13][15] In a relic, low elevation population, seeds are also dispersed by small rodents.[16] American black bears and grizzly bears may raid squirrel caches for limber pine nuts.[7] Squirrels, Northern flickers, and mountain bluebirds often nest in the trees. There is some evidence that P. flexilis has a symbiotic relationship with nitrogen-fixing bacteria that inhabit the needles.[17]

The species is generally shade tolerant and resistant to fire, but does not thrive in dense habitats, instead occurring in areas relatively hostile to other species.[18][7]

Threats

Limber pine is susceptible to white pine blister rust,[7] caused by Cronartium ribicola, a fungus that was introduced accidentally from Europe. Limber pine mortality is high in many areas throughout its range, except Arizona, where it has not yet been found. However, there is little hope of controlling the blister rust in existing trees. Research is under way, locating and breeding from the occasional naturally resistant limber pines, and by studying the resistance mechanisms of the European and Asian white pines (e.g. Swiss pine, Macedonian pine), which are strongly resistant to the disease.

The tree has also been damaged by bark beetle epidemics, particularly at drought-affected low elevations.[7]

Cultivation

The popular cultivar P. flexilis 'Vanderwolf's Pyramid' is widely available as an ornamental tree for gardens. 'Vanderwolf's Pyramid' derives from P. reflexa, though it is usually listed in nursery catalogs under P. flexilis.

The Southwestern white pine is popular as a windbreak tree or an ornamental tree due to its drought tolerance. It is also grown as a Christmas tree, liked for the soft needles but with stiffer branches than an Eastern white pine.

Uses

The large seeds are edible,[19] and were reportedly consumed by Native Americans in Montana.[7]

See also

References

  1. ^ Schoettle, A.; Stritch, L. (2013). "Pinus flexilis". IUCN Red List of Threatened Species. 2013: e.T42363A2975338. doi:10.2305/IUCN.UK.2013-1.RLTS.T42363A2975338.en. Retrieved 19 November 2021.
  2. ^ a b Kral, Robert (1993). "Pinus flexilis". In Flora of North America Editorial Committee (ed.). Flora of North America North of Mexico (FNA). Vol. 2. New York and Oxford – via eFloras.org, Missouri Botanical Garden, St. Louis, MO & Harvard University Herbaria, Cambridge, MA.
  3. ^ "Pinus flexilis". Tropicos. Missouri Botanical Garden.
  4. ^ "Old Tree". Oregon Field Guide. 2010. Retrieved 2010-02-21.
  5. ^ Richard, Terry (September 24, 2012). "Ancient limber pine, likely Oregon's oldest living tree, draws twin brothers to Wallowas quest (photo essay, video)". The Oregonian. Retrieved 2015-02-18.
  6. ^ "Stay Flexible, Grow Old". BYU Magazine. Spring 2007. Retrieved 2015-02-18.
  7. ^ a b c d e f g h i j k l Arno, Stephen F.; Hammerly, Ramona P. (2020) [1977]. Northwest Trees: Identifying & Understanding the Region's Native Trees (field guide ed.). Seattle: Mountaineers Books. pp. 26, 43–49. ISBN 978-1-68051-329-5. OCLC 1141235469.
  8. ^ a b Moore, Gerry; Kershner, Bruce; Tufts, Craig; Mathews, Daniel; et al. (2008). National Wildlife Federation Field Guide to Trees of North America. New York: Sterling. p. 81. ISBN 978-1-4027-3875-3.
  9. ^ Letts, M.G.; Nakonechny, K.N.; Van Gaalen, K.E.; Smith, C.M. (2009). "Physiological acclimation of Pinus flexilis to drought stress on contrasting slope aspects in Waterton Lakes National Park, Alberta, Canada" (PDF). Canadian Journal of Forest Research. 39 (3): 629–641. doi:10.1139/X08-206. Archived from the original (PDF) on 2011-07-16.
  10. ^ "Pinus subgenus Strobus". Michael P. Frank's Cone Collection. Arboretum de Villardebelle.
  11. ^ Roady, Laura. "Whitebark Pine". Montana Outdoors. Montana Fish, Wildlife & Parks. Retrieved 15 March 2015.
  12. ^ a b Steele, Robert (1990). "Pinus flexilis". In Burns, Russell M.; Honkala, Barbara H. (eds.). Conifers. Silvics of North America. Washington, D.C.: United States Forest Service (USFS), United States Department of Agriculture (USDA). Vol. 1 – via Southern Research Station.
  13. ^ a b Siepielski, A. M.; Benkman, C. W. (June 2007). "Selection by a predispersal seed predator constrains the evolution of avian seed dispersal in pines". Functional Ecology. 21 (3): 611–618. doi:10.1111/j.1365-2435.2007.01261.x. ISSN 0269-8463.
  14. ^ Siepielski, Adam M.; Benkman, Craig W. (May 2007). "Convergent Patterns in the Selection Mosaic for Two North American Bird-Dispersed Pines". Ecological Monographs. 77 (2): 203–220. doi:10.1890/06-0929. ISSN 0012-9615.
  15. ^ Siepielski, Adam M.; Benkman, Craig W. (October 2008). "Seed Predation and Selection Exerted by a Seed Predator Influence Subalpine Tree Densities". Ecology. 89 (10): 2960–2966. doi:10.1890/08-0072.1. ISSN 0012-9658. PMID 18959333.
  16. ^ Tomback, Diana F.; Schoettle, Anna W.; Chevalier, Kristen E.; Jones, Cheri A. (2005-01-01). "Life on the edge for limber pine: Seed dispersal within a peripheral population". Écoscience. 12 (4): 519–529. doi:10.2980/i1195-6860-12-4-519.1. ISSN 1195-6860. S2CID 86330607.
  17. ^ Moyes, Andrew B.; Kueppers, Lara M.; Pett-Ridge, Jennifer; Carper, Dana L.; Vandehey, Nick; O'Neil, James; Frank, A. Carolin (2016-04-01). "Evidence for foliar endophytic nitrogen fixation in a widely distributed subalpine conifer" (PDF). New Phytologist. 210 (2): 657–668. doi:10.1111/nph.13850. ISSN 1469-8137. PMID 27000956.
  18. ^ Rebertus, A. J.; Burns, B. R.; Veblen, T. T. (August 1991). "Stand dynamics of Pinus flexilis -dominated subalpine forests in the Colorado Front Range". Journal of Vegetation Science. 2 (4): 445–458. doi:10.2307/3236026. JSTOR 3236026.
  19. ^ Whitney, Stephen (1985). Western Forests (The Audubon Society Nature Guides). New York: Knopf. pp. 407–08. ISBN 0-394-73127-1.

Further reading

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Pinus flexilis: Brief Summary

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Pinus flexilis, the limber pine, is a species of pine tree-the family Pinaceae that occurs in the mountains of the Western United States, Mexico, and Canada. It is also called Rocky Mountain white pine.

A limber pine in Eagle Cap Wilderness, Oregon, has been documented as over 2,000 years old, and another one was confirmed at 1,140 years old. Another candidate for the oldest limber pine was identified in 2006 near the Alta Ski Area in Utah; called "Twister", the tree was confirmed to be at least 1,700 years old and thought to be even older.

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