Overview

Distribution

Range Description

USA: mainly California, in two disjunct (sub)populations in northern (Klamath Mts.) and in southern (Sierra Nevada) California. One locality (metapopulation) of the northern (sub)population is just across the state border in Oregon (Lanner 2007).
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More info for the term: Pleistocene

Foxtail pine is endemic to California. There is a 300-mile (480-km) gap in the distributions of the 2 subspecies. Southern foxtail pine occurs in the high southern Sierra Nevada of Fresno, Tulare, and Inyo counties. Its distribution lies within Sequoia and Inyo National Forests and Sequoia-Kings Canyon National Park. Highest population concentrations are in Sequoia-Kings Canyon National Park, the center of most research on southern foxtail pine. Northern foxtail pine occurs in the high North Coast and Klamath ranges in Siskiyou, Trinity, Shasta, and Tehama counties. Northern foxtail pine's distribution lies within the Klamath, Shasta, Trinity, and Mendocino National Forests, including the Marble Mountain, Yolla-Bolly-Middle Eel, and Trinity Alps Wilderness Areas [26,39,59,61]. Southern and northern foxtail pines have probably been separated since major upliftings of the Sierra Nevada during the early Pleistocene [8]. Inyo Valley, located between the southern Sierra Nevada and the White Mountains, creates a 20-mile-wide (32-km) gap between southern foxtail and Great Basin bristlecone pine populations [26]. The U.S. Geological Survey provides a distributional map of southern and northern foxtail pines.
  • 39. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. [21992]
  • 8. Bailey, D. K. 1970. Phytogeography and taxonomy of Pinus subsection Balfourianae. Annals of the Missouri Botanical Garden. 57: 210-249. [375]
  • 26. Critchfield, William B.; Little, Elbert L., Jr. 1966. Geographic distribution of the pines of the world. Misc. Publ. 991. Washington, DC: U.S. Department of Agriculture, Forest Service. 97 p. [20314]
  • 59. Lanner, Ronald M. 1999. Conifers of California. Los Olivos, CA: Cachuma Press. 274 p. [30288]
  • 61. Little, Elbert L., Jr. 1975. Rare and local conifers in the United States. Conservation Research Rep. No. 19. Washington, DC: U.S. Department of Agriculture, Forest Service. 25 p. [15691]

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National Distribution

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

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Regional Distribution in the Western United States

More info on this topic.

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 [16]:

1 Northern Pacific Border

2 Cascade Mountains

4 Sierra Mountains
  • 16. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434]

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States or Provinces

CA

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Localities documented in Tropicos sources

Pinus balfouriana Balf.:
United States (North America)

Note: This information is based on publications available through Tropicos and may not represent the entire distribution. Tropicos does not categorize distributions as native or non-native.
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Physical Description

Morphology

Description

Trees to 22m; trunk to 2.6m diam., erect or leaning; crown broadly conic to irregular. Bark gray to salmon or cinnamon, platy or irregularly deep-fissured or with irregular blocky plates. Branches contorted, ascending to descending; twigs red-brown, aging gray to drab yellow-gray, glabrous or puberulent, young branches resembling long bottlebrushes because of persistent leaves. Buds ovoid-acuminate, red-brown, 0.8--1cm, resinous. Leaves 5 per fascicle, upcurved, persisting 10--30 years, 1.5--4cm ´ 1--1.4mm, mostly connivent, deep blue- to deep yellow-green, abaxial surface without median groove but usually with 2 subepidermal but evident resin bands, adaxial surfaces conspicuously whitened by stomates, margins mostly entire to blunt, apex broadly acute to acuminate; sheath 0.5--1cm, soon forming rosette, shed early. Pollen cones ellipsoid, ca. 6--10mm, red. Seed cones maturing in 2 years, shedding seeds and falling soon thereafter, spreading, symmetric, lance-cylindric with conic base before opening, broadly lance-ovoid or ovoid to cylindric or ovoid-cylindric when open, 6--9(--11)cm, purple, aging red-brown, nearly sessile; apophyses much thickened, rounded, larger toward cone base; umbo central, usually depressed; prickle absent or weak, to 1mm, resin exudates amber. Seeds ellipsoid to narrowly obovoid; body to 10mm, pale brown, mottled with deep red; wing 10--12mm. 2 n =24.
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Description

More info for the terms: dehiscent, density, tree

Foxtail pine is a native conifer. It is a low-growing pine, generally 20 to 50 feet (6-15 m) tall, but occasionally reaching 72+ feet (22+ m) in height [39,76]. The champion tree is a northern foxtail pine on the Trinity National Forest that measures 76 feet (23 m) in height, 34 feet (10 m) in spread, and 26.3 feet (8 m) in circumference [2]. Foxtail pine's trunk is usually single-stemmed. Unlike other North American conifers, foxtail pine rarely assumes krummholz form at high elevations; instead, it retains a straight bole [8,10,82,83]. Bark of mature foxtail pine is "exceptionally thick" [8]: nearly as thick as that of ponderosa pine (Pinus ponderosa). Bark thickness measurements of 1.85 inches (4.70 cm) [45] and 2.4 to 3.1 inches (6-8 cm) [8] are reported for mature foxtail pines. Mature tree crowns are 8.5 feet (2.6 m) or less in diameter. Branches are short and thick in diameter [39,76]. Branching habit is sparse in southern foxtail pine [59]; southern foxtail pine also tends to be self-pruning. Northern foxtail pine branches tend to be thicker, and may extend to the ground [6,39,67] (the photos in the Introductory section illustrate this difference). Foxtail is a 5-needle pine. The Balfourianae complex is unique among pines in that about half of their branches originate from within the needle fascicles [24,59]. Needle clusters are thickly set toward the branch ends, resembling foxtails [58]. Needles persist for 5 to 7+ years [64,70]; trees at lower elevations tend to retain needles longer than trees at timberline [64]. Female cones are 2.4 to 7.5 inches (6-19 cm) long, dehiscent, and have tiny prickles. Seeds are small (~0.3 inch (8 mm) long), with detachable seed wings about 3 times longer than the seeds [39,76]. Northern foxtail pines tend to have heavier cones and larger seeds with longer seed wings than southern foxtail pines [66].

Morphological differences between southern foxtail pine, northern foxtail pine, and Great Basin bristlecone pine are slight. Southern foxtail pine has thinner bark that tends to grow in square plates (pictured above right) compared to northern foxtail pine, which has relatively thicker bark that tends to grow in narrow ridges. Southern foxtail pine retains its needles longer than northern foxtail pine [39]. Northern foxtail pine tends to have a fuller crown and suffer from less cambial die-back than southern foxtail pine [59]. Great Basin bristlecone pine is distinguished from foxtail pines by having relatively longer cone prickles (2-6 mm) compared to foxtail pines (<1mm). Distributions of Great Basin, southern foxtail, and northern foxtail pines do not overlap [39,61], so distinguishing among them in the field is easy.

Stand structure: Foxtail pine communities are typically open, with a sparse understory and scattered woody debris. Arid, high-elevation conditions allow woody debris to persist for many years without decaying [44]. In Sequoia-Kings Canyon National Park, southern foxtail pine grows in widely spaced woodlands in its upper elevational range and is often the only tree species. At lower elevations it forms a more dense forest, either in mixed or monospecific stands [10,12,81,82]. The foxtail pine-alpine ecotone is usually abrupt as a result of foxtail pine's inability to form krummholz [63]. Northern foxtail pine communities tend toward greater density then southern foxtail pine communities [82]. In the Klamath Ranges, stand densities of northern foxtail pine communities ranged from a minimum of  51 trees/ha in the Yolla Bolly Mountains to a maximum of 381 trees/ha in the Trinity Mountains [30]. Stand densities of southern foxtail pine communities in Sequoia-Kings Canyon National Park range from 50 trees/ha to 600 trees/ha [30,63,80,82]. Ryerson [82] found a mean stand density of 100 trees/ha on sites across southern foxtail pine's distribution.

Age class: Age class structure within foxtail pine stands appears mixed [30,68]. Few studies have been conducted on age class distributions in foxtail pine. In a study across the Klamath Ranges, Eckert and Sawyer [30] found northern foxtail pines less than 100 years of age were most common (>50% relative density). A few very old trees (around 1,000 years of age) were scattered within all the study sites. In a study across southern foxtail pine's distribution in the Sierra Nevada, Ryerson [82] found most trees were in the 350- to 500-year-old class, followed by trees less than 200 years old, and trees older than 800 years, respectively.

Foxtail pine is a very long-lived conifer, although it does not approach the extreme ages of bristlecone pines. Foxtail pine occurs on wetter sites than bristlecones; consequently, foxtail pines show relatively faster growth, develop heart rot, and die more quickly than bristlecone pines [8]. Foxtail pine has advanced heart rot by 1,000 years of age. The oldest foxtail pine on record (as of 2004) is a 3,400-year-old southern foxtail pine [82]. Northern foxtail pines occur in wetter habitats then southern foxtail pines and are shorter lived, attaining maximum ages of about 1,600 years [30,68,82].

Physiology: Its relative inability to withstand cold may partially explain foxtail pine's narrow distribution compared to its more widely distributed high-elevation associate, whitebark pine. Poor ability to form krummholz limits foxtail pine's ability to withstand ice blasting [13]. Its seedlings are less resistant to freezing than whitebark pine seedlings [82].

  • 39. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. [21992]
  • 8. Bailey, D. K. 1970. Phytogeography and taxonomy of Pinus subsection Balfourianae. Annals of the Missouri Botanical Garden. 57: 210-249. [375]
  • 45. Jackson, James F.; Adams, Dean C.; Jackson, Ursula B. 1999. Allometry of constitutive defense: a model and a comparative test with tree bark and fire regime. The American Naturalist. 153(6): 614-632. [31152]
  • 59. Lanner, Ronald M. 1999. Conifers of California. Los Olivos, CA: Cachuma Press. 274 p. [30288]
  • 61. Little, Elbert L., Jr. 1975. Rare and local conifers in the United States. Conservation Research Rep. No. 19. Washington, DC: U.S. Department of Agriculture, Forest Service. 25 p. [15691]
  • 10. Bancroft, Larry. 1979. Fire management plan: Sequoia and Kings Canyon National Parks. San Francisco, CA: U.S. Department of the Interior, National Park Service, Western Region. 190 p. [11887]
  • 12. Barbour, Michael G. 1988. Californian upland forests and woodlands. In: Barbour, Michael G.; Billings, William Dwight, eds. North American terrestrial vegetation. Cambridge; New York: Cambridge University Press: 131-164. [13880]
  • 30. Eckert, Andrew J.; Sawyer, John O. 2002. Foxtail pine importance and conifer diversity in the Klamath Mountains and southern Sierra Nevada, California. Madrono. 49(1): 33-45. [42265]
  • 44. Holland, Robert F. 1986. Preliminary descriptions of the terrestrial natural communities of California. Sacramento, CA: California Department of Fish and Game. 156 p. [12756]
  • 63. Lloyd, Andrea H.; Graumlich, Lisa J. 1997. Holocene dynamics of treeline forests in the Sierra Nevada. Ecology. 78(4): 1199-1210. [27723]
  • 64. Lloyd, Andrea. 1998. Growth of foxtail pine seedlings at treeline in the southeastern Sierra Nevada, California, U.S.A. Ecoscience. 5(2): 250-257. [48110]
  • 67. Mastroguiseppe, R. J.; Mastroguiseppe, J. D. 1980. A study of Pinus balfouriana Grev. & Balf. (Pinaceae). Systematic Botany. 5(1): 86-104. [1546]
  • 68. Mastroguiseppe, Ronald J. 1972. Geographic variation in foxtail pine, Pinus balfouriana Grev. & Balf. Humbolt, CA: California State University, Humboldt. 98 p. Thesis. [1548]
  • 76. Munz, Philip A. 1974. A flora of southern California. Berkeley, CA: University of California Press. 1086 p. [4924]
  • 80. Roy, D. Graham; Vankat, John L. 1999. Reversal of human-induced vegetation changes in Sequoia National Park, California. Canadian Journal of Forest Research. 29(4): 399-412. [36282]
  • 81. Rundel, Philip W.; Parsons, David J.; Gordon, Donald T. 1977. Montane and subalpine vegetation of the Sierra Nevada and Cascade Ranges. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley & Sons: 559-599. [4235]
  • 82. Ryerson, A. Diane. 1983. Population structure of Pinus balfouriana Grev. & Balf. along the margins of its distribution area in the Sierran and Klamath regions of California. Sacramento, CA: California State University. 197 p. Thesis. [48204]
  • 83. Ryerson, Diane. 1984. Krummholz foxtail pines. Fremontia. 11(4): 30. [47919]
  • 6. Arno, Stephen F.; Gyer, Jane. 1973. Discovering Sierra trees. Yosemite National Park, CA: Yosemite Natural History Association. 89 p. [48231]
  • 13. Barbour, Michael G.; Major, Jack, eds. 1977. Terrestrial vegetation of California. New York: John Wiley & Sons. 1002 p. [388]
  • 24. Connor, Kristina F.; Lanner, Ronald M. 1987. The architectural significance of interfoliar branches in Pinus subsection Balfourianae. Canadian Journal of Forest Research. 17(3): 269-272. [47907]
  • 58. Lanner, Ronald M. 1996. Made for each other: a symbiosis of birds and pines. New York: Oxford University Press. 160 p. [29914]
  • 66. Mastroguiseppe, R. J. 1968. Geographic variation in foxtail pine. Progress Report. Placerville, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station; Institute of Forest Genetics. 15 p. On file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. [48235]
  • 70. McCune, Bruce. 1988. Ecological diversity in North American pines. American Journal of Botany. 75(3): 353-368. [5651]
  • 2. American Forests. 2004. Foxtail pine: Pinus balfouriana. In: National register of big trees, [Online]. Available: http://www.americanforests.org/resources/bigtrees/ [2004, June 14]. [48210]

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Physical Description

Tree, 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 entire (use magnification), Leaf apex acute, Leaves < 5 cm long, Leaves < 10 cm long, Leaves grey-green, Leaves not blue-green, Leaves white-striped, Needle-like leaves somewhat rounded, Needle-like leaves not twisted, Needle-like leaf habit erect, Needle-like leaves per fascicle mostly 5, Needle-like leaf sheath early deciduous, Twigs glabrous, 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, Umbo with obvious prickle, Bracts of seed cone included, Seeds red, Seeds brown, Seeds winged, Seeds wingless, Seeds unequally winged, Seed wings prominent, Seed wings narrower than body, Seed wings equal to or broader than body.
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Stephen C. Meyers

Source: USDA NRCS PLANTS Database

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Diagnostic Description

Synonym

Pinus balfouriana var. austrina (R.Mastrogiuseppe & J.Mastrogiuseppe) Silba; P. balfouriana subsp. austrina R.Mastrogiuseppe & J.Mastrogiuseppe
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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
Pinus balfouriana occurs in the subalpine to alpine zones of the Klamath Mountains (the northern [sub]population) and of the southern Sierra Nevada (the southern [sub]population). In the north it is found at altitudes of between 1,600 m and 2,400 m a.s.l., in the south between 2,900 m and 3,700 m. Stands of this pine are very open and occur on dry, rocky, exposed high slopes and ridges, usually devoid of other significant vegetation. Stands may be pure or mixed with P. albicaulis, sometimes Juniperus occidentalis grows with it, too. Regeneration and growth are extremely slow and stands commonly look as if entirely composed of veteran trees of great age. Regeneration is probably episodal and may be linked with climatic cycles. Unlike its even longer lived 'cousin' P. longaeva, growing only 35-40 km to the east of the Sierra Nevada, little is known about the exact ages of some of the oldest trees, but they are likely to be more than 2000 years old.

Systems
  • Terrestrial
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Habitat characteristics

More info for the terms: density, serpentine soils, tree, ultramafic soils

Foxtail pine is adapted to harsh environmental conditions [59,70]. Its long life span, slow growth, and persistent needles are typical of conifer species adapted to stressful habitats [70]. Foxtail pine  holds an ecological position between whitebark pine and limber pine: it is less tolerant of cool, short growing seasons than whitebark pine and less tolerant of warm, arid growing conditions than limber pine [44,86]. North of southern foxtail pine's distribution in the Sierra, freezing temperatures occur all year long [82]. Foxtail pine communities are most common on "safe sites," such as ultramafic soils and dry granite fields, that few other conifer species can tolerate [30,59]. Slope varies from gentle to as much as 55% [82,83].

Southern foxtail pine grows on well-drained, decomposed granite and granite boulder fields. Southern foxtail pine does not occur on serpentine or other ultramafic soils, which are rare in the high Sierra Nevada [51,54,82]. It is more common on the drier, eastern side of the Sierra Nevada, while whitebark pine is more common on the west slope [10,82]. Climate in the southern Sierra Nevada is mediterranean, with cold winters and warm, dry summers [67,86]. Annual precipitation on the east slope ranges from 20 to 30 inches (500-750 mm) [13]. Southern foxtail pine occurs from 8,900 to 12,000 feet (2,700-3,700 m) elevation [39]. Tree damage from ice- and sandstorms is common [67]. Highest density of southern foxtail pine occurs on north-facing slopes; least density is on south slopes. Percent slope across southern foxtail pine's range averages less than 33% [82].

Habitat of northern foxtail pine is even more restricted than that of southern foxtail pine. The Klamath Ranges are geologically complex, consisting of steep elevational gradients and a variety of parent rock materials that strongly influence plant community boundaries [51]. Climate is mediterranean, but is strongly moderated by the maritime influence of the nearby Pacific Ocean [67]. Annual precipitation averages from 49 to 60 inches (1,250-1,750 mm) [13]. Northern foxtail pine occurs from 6,900 to 8,200 feet (2,100-2,500 m) elevation [39]. There are relatively few high-elevation peaks in the Klamath Ranges; therefore, northern foxtail pine tends to segregate into small populations on isolated "sky islands" [77]. Substrates on which northern foxtail pine grows include gabbro, granodiorite, limestone, schist, and most commonly, serpentine [51,54,59,86]. Because most associated conifers (except Jeffrey pine) are less tolerant to them, serpentine soils can partially ameliorate the elevational restriction and lower northern foxtail pine's elevational distribution. Northern foxtail pine tends to grow in large, monospecific stands when on serpentine soils. On other substrates it is generally found in small stands (a few hundred trees) on ridge crests, mountain tops, and steep, south- or west-facing slopes [30,68,77,86]. Populations on serpentine soils are more likely to occur on all aspects, including valley bottoms and lake shores [77]. Percent slope ranged from 15-32% on 4 sites in the Klamath Ranges [82].

  • 39. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. [21992]
  • 59. Lanner, Ronald M. 1999. Conifers of California. Los Olivos, CA: Cachuma Press. 274 p. [30288]
  • 10. Bancroft, Larry. 1979. Fire management plan: Sequoia and Kings Canyon National Parks. San Francisco, CA: U.S. Department of the Interior, National Park Service, Western Region. 190 p. [11887]
  • 30. Eckert, Andrew J.; Sawyer, John O. 2002. Foxtail pine importance and conifer diversity in the Klamath Mountains and southern Sierra Nevada, California. Madrono. 49(1): 33-45. [42265]
  • 44. Holland, Robert F. 1986. Preliminary descriptions of the terrestrial natural communities of California. Sacramento, CA: California Department of Fish and Game. 156 p. [12756]
  • 54. Laacke, Robert J. 1990. Abies magnifica A. Murr. California red fir. In: Burns, Russell M.; Honkala, Barbara H., technical coordinators. Silvics of North America. Volume 1. Conifers. Agric. Handb. 654. Washington, DC: U.S. Department of Agriculture, Forest Service: 71-79. [13370]
  • 67. Mastroguiseppe, R. J.; Mastroguiseppe, J. D. 1980. A study of Pinus balfouriana Grev. & Balf. (Pinaceae). Systematic Botany. 5(1): 86-104. [1546]
  • 68. Mastroguiseppe, Ronald J. 1972. Geographic variation in foxtail pine, Pinus balfouriana Grev. & Balf. Humbolt, CA: California State University, Humboldt. 98 p. Thesis. [1548]
  • 77. Oline, David K.; Mitton, Jeffry B.; Grant, Michael C. 2000. Population and subspecific genetic differentiation in the foxtail pine (Pinus balfouriana). Evolution. 54(5): 1813-1819. [48101]
  • 82. Ryerson, A. Diane. 1983. Population structure of Pinus balfouriana Grev. & Balf. along the margins of its distribution area in the Sierran and Klamath regions of California. Sacramento, CA: California State University. 197 p. Thesis. [48204]
  • 83. Ryerson, Diane. 1984. Krummholz foxtail pines. Fremontia. 11(4): 30. [47919]
  • 86. Sawyer, John O.; Thornburgh, Dale A. 1977. Montane and subalpine vegetation of the Klamath Mountains. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley & Sons: 699-732. [685]
  • 13. Barbour, Michael G.; Major, Jack, eds. 1977. Terrestrial vegetation of California. New York: John Wiley & Sons. 1002 p. [388]
  • 51. Kruckeberg, Arthur R. 1984. California serpentines: flora, vegetation, geology, soils and management problems. Publications in Botany Volume 48. Berkeley, CA: University of California Press. 180 p. [12482]
  • 70. McCune, Bruce. 1988. Ecological diversity in North American pines. American Journal of Botany. 75(3): 353-368. [5651]

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Key Plant Community Associations

More info for the terms: cover, serpentine soils, shrub, shrubs, tree

Southern foxtail pine is the dominant conifer in upper-elevation
subalpine communities of the south-central Sierra Nevada [12,80,106]. It is the
most abundant subalpine conifer in Sequoia-Kings Canyon National Park
[10]. Southern foxtail pine often occurs in pure
stands [63]. California red fir (Abies magnifica var. magnifica), Sierra juniper
(Juniperus occidentalis ssp. australis), Sierra lodgepole pine (Pinus contorta var. murrayana),
Jeffrey pine (P.
jeffreyi), and limber pine (P. flexilis) may associate on low-elevation foxtail pine sites, while whitebark pine
(P. albicaulis) may associate on high-elevation sites [81,82,106]. The most common
shrub associates in
southern foxtail pine communities are bush chinquapin (Chrysolepis sempervirens),
oceanspray (Holodiscus discolor), wax currant (Ribes cereum), gooseberry currant (R. montigenum),
Parish's snowberry
(Symphoricarpos rotundifolius var. parishii), and curlleaf mountain-mahogany
(Cercocarpus ledifolius) [82,83]. Pinemat manzanita (Arctostaphylos nevadensis),
green manzanita (A. patula), and bitter cherry (Prunus emarginata) may also be present [10,82].

In Sequoia-King Canyon National Park, Vankat [106] found less than one-fourth of southern foxtail pine communities
contained shrubs. Herbaceous cover averaged around 10%, although some stands showed
as much as 65% herbaceous cover. Tree cover was almost entirely foxtail pine; a few stands also had Sierra lodgepole pine.
Bush chinquapin and
oceanspray were the most common shrubs. Shrub associates are more common on marginal foxtail pine
sites than in Sequoia-Kings Canyon, which is prime foxtail habitat. Southern monardella (Monardella australis)
is the most consistent herbaceous associate across southern foxtail pine's range [82]. At their lowest elevations,
southern foxtail pine communities merge with upper-elevation Sierra lodgepole pine or, more rarely,
east-slope Jeffrey pine
communities. Southern foxtail pine communities often
form a mosaic with subalpine meadows [15]. At their highest elevations, southern foxtail pine communities merge into
alpine meadows and fell-fields [44].
The Klamath Ranges support some of the most diverse plant communities in North
America [86], and northern foxtail pine contributes to this diversity. Northern foxtail communities are
typically more diverse compared to southern foxtail pine communities [67]. Eckert and Sawyer [30] found that
along a latitudinal gradient, diversity of northern foxtail pine communities increased
to the north. Lowest diversity occurred in the southern Yolla Bolly Mountains, the
southernmost of the Klamath ranges, while highest diversity occurred in the
Trinity and Marble mountains, the northernmost of the Klamath ranges.
Northern foxtail pine generally dominates on subalpine serpentine soils.
Whitebark pine or mountain hemlock (Tsuga mertensiana) may
associate on upper-elevation sites, although they are more common on nonserpentine
soils [30,82]. Northern foxtail pine communities on
serpentine, other ultramafic,
or dry soils often form a mosaic with whitebark
pine-mountain hemlock or mountain hemlock-Brewer spruce (Picea
breweriana) communities that occur on nonserpentine or wetter,
north-slope soils [30,44,54,63]. Jeffrey pine and Sierra lodgepole pine may associate on
dry (south and west aspects), mid-subalpine sites (<2,200 m), while Shasta red fir (Abies magnifica var.
shastensis) and western white pine may associate on
wetter, mid-subalpine sites [30,38,44,54,67].
Western white pine or white fir (A. concolor) associate on lowest-elevation
subalpine sites [23,86].  Incense-cedar (Calocedrus decurrens) and
coast Douglas-fir (Pseudotsuga menziesii var. menziesii) may be
occasional associates at these lower elevations [67,82]. Northern foxtail pine
communities merge with coast Douglas-fir, Sierra lodgepole pine, or mixed-conifer forest
communities at their lowest
elevations and with alpine fell-field or
alpine meadow communities at highest
elevations [44,63].
In the Trinity Mountains on the Siskiyou-Trinity county line, northern foxtail pine occurs
in pure stands at timberline.
Shasta red fir, western white pine, and Jeffrey pine associate on mid-subalpine
sites. Trinity
buckwheat (Eriogonum alpinum), pinemat manzanita, big sagebrush (Artemisia tridentata), and
huckleberry oak (Quercus
vaccinifolia) occur in the understory. Ground-layer associates include cobwebby paintbrush (Castilleja arachnoidea),
Cascade aster (Eucephalus
ledophyllus), spreading phlox (Phlox diffusa), and bottlebrush
squirrel (Elymus elymoides) [23,86]. Unusually diverse northern foxtail
communities exist on China and Russian peaks, where northern foxtail pine
associates with Jeffrey pine, incense-cedar, and Pacific Douglas-fir [85].
Holland [44], Rundel and others [81], and Sawyer and Thornburgh [86] provide vegetation typings describing foxtail pine
communities.
  • 10. Bancroft, Larry. 1979. Fire management plan: Sequoia and Kings Canyon National Parks. San Francisco, CA: U.S. Department of the Interior, National Park Service, Western Region. 190 p. [11887]
  • 12. Barbour, Michael G. 1988. Californian upland forests and woodlands. In: Barbour, Michael G.; Billings, William Dwight, eds. North American terrestrial vegetation. Cambridge; New York: Cambridge University Press: 131-164. [13880]
  • 15. Benedict, Nathan B.; Major, Jack. 1982. A physiographic classification of subalpine meadows of the Sierra Nevada California. Madrono. 29(1): 1-12. [8268]
  • 23. Coleman, Robert G.; Kruckeberg, Arthur R. 1999. Geology and plant life of the Klamath-Siskiyou Mountain Region. Natural Areas Journal. 19(4): 320-340. [33090]
  • 30. Eckert, Andrew J.; Sawyer, John O. 2002. Foxtail pine importance and conifer diversity in the Klamath Mountains and southern Sierra Nevada, California. Madrono. 49(1): 33-45. [42265]
  • 38. Harvey, H. Thomas; Mastrogiuseppe, Ronald J. 1971. Foxtail pine on Sirretta Peak, California. Madrono. 21(3): 152. [47920]
  • 44. Holland, Robert F. 1986. Preliminary descriptions of the terrestrial natural communities of California. Sacramento, CA: California Department of Fish and Game. 156 p. [12756]
  • 54. Laacke, Robert J. 1990. Abies magnifica A. Murr. California red fir. In: Burns, Russell M.; Honkala, Barbara H., technical coordinators. Silvics of North America. Volume 1. Conifers. Agric. Handb. 654. Washington, DC: U.S. Department of Agriculture, Forest Service: 71-79. [13370]
  • 63. Lloyd, Andrea H.; Graumlich, Lisa J. 1997. Holocene dynamics of treeline forests in the Sierra Nevada. Ecology. 78(4): 1199-1210. [27723]
  • 67. Mastroguiseppe, R. J.; Mastroguiseppe, J. D. 1980. A study of Pinus balfouriana Grev. & Balf. (Pinaceae). Systematic Botany. 5(1): 86-104. [1546]
  • 80. Roy, D. Graham; Vankat, John L. 1999. Reversal of human-induced vegetation changes in Sequoia National Park, California. Canadian Journal of Forest Research. 29(4): 399-412. [36282]
  • 81. Rundel, Philip W.; Parsons, David J.; Gordon, Donald T. 1977. Montane and subalpine vegetation of the Sierra Nevada and Cascade Ranges. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley & Sons: 559-599. [4235]
  • 82. Ryerson, A. Diane. 1983. Population structure of Pinus balfouriana Grev. & Balf. along the margins of its distribution area in the Sierran and Klamath regions of California. Sacramento, CA: California State University. 197 p. Thesis. [48204]
  • 83. Ryerson, Diane. 1984. Krummholz foxtail pines. Fremontia. 11(4): 30. [47919]
  • 85. Sawyer, John O.; Keeler-Wolf, Todd. 1995. A manual of California vegetation. Misc. Report. Sacramento, CA: California Native Plant Society Press. 412 p. [48236]
  • 86. Sawyer, John O.; Thornburgh, Dale A. 1977. Montane and subalpine vegetation of the Klamath Mountains. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley & Sons: 699-732. [685]
  • 106. Vankat, John Lyman. 1970. Vegetation change in Sequoia National Park, California. Davis, CA: University of California. 197 p. Dissertation. [43459]

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Habitat: Rangeland Cover Types

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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 term: cover

SRM (RANGELAND) COVER TYPES [90]:

None
  • 90. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. [23362]

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Habitat: Cover Types

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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 [31]:

204 Mountain hemlock

207 Red fir

208 Whitebark pine

218 Lodgepole pine

219 Limber pine

256 California mixed subalpine
  • 31. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905]

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Habitat: Plant Associations

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

KUCHLER [53] PLANT ASSOCIATIONS:

K007 Red fir forest

K008 Lodgepole pine-subalpine forest
  • 53. Kuchler, A. W. 1964. United States [Potential natural vegetation of the conterminous United States]. Special Publication No. 36. New York: American Geographical Society. 1:3,168,000; colored. [3455]

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Habitat: Ecosystem

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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):

ECOSYSTEMS [34]:

FRES23 Fir-spruce

FRES26 Lodgepole pine
  • 34. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; Lewis, Mont E.; Smith, Dixie R. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998]

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Habitat & Distribution

Timberline and alpine meadows; of conservation concern; 1500--3500m; Calif.
Creative Commons Attribution Non Commercial Share Alike 3.0 (CC BY-NC-SA 3.0)

© Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA

Source: Missouri Botanical Garden

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General Ecology

Fire Management Considerations

More info for the terms: natural, prescribed natural fire

Southern foxtail pine communities are managed with prescribed natural fire (wildland fire for resource benefit) [91]. Fire may extend foxtail pine's distribution downslope on some sites [48], although the postfire interactions of California subalpine conifers have had too little study to predict postfire results with confidence. Fire may encourage foxtail pine recruitment and growth on existing foxtail pine sites by releasing nutrients from slow-decaying woody debris. On those relatively rare foxtail pine sites with closed canopies, fire would encourage foxtail pine recruitment by creating an open mineral seedbed.
  • 91. Skinner, Carl N.; Chang, Chi-ru. 1996. FIRE REGIMES, past and present. In: Status of the Sierra Nevada. Sierra Nevada Ecosystem Project: Final report to Congress. Volume II: Assessments and scientific basis for management options. Wildland Resources Center Report No. 37. Davis, CA: University of California, Centers for Water and Wildland Resources: 1041-1069. [28975]
  • 48. Kiefer, M. 1991. Forest age structure, species composition, and fire disturbance in the southern Sierra Nevada subalpine zone. Unpublished report [submitted to the Sequoia Natural History Association]. Tucson, AZ: University of Arizona, Laboratory of Tree-Ring Research. [48233]

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Plant Response to Fire

More info for the terms: fire management, natural

Foxtail pine's thick bark helps protect it from damage from surface fires. Sparse, large-diameter branches discourage torching and fire spread. Little is known about foxtail pine's response after the fire has passed. In the 1979 Fire Management Plant for Sequoia-Kings Canyon National Park, Bancroft [10] wrote "Very little research has been conducted on the effect of fire on natural regeneration in subalpine forests." This remains true of foxtail pine and other subalpine forests today. Research is needed on the fire ecology of foxtail pine.

Kiefer [48] found foxtail pine recruitment in Sequoia-Kings Canyon National Park was uneven-aged and did not appear to be correlated with fire history. This was in sharp contrast to associated Sierra lodgepole pine, whose recruitment dated from past fires. Kiefer suggested that climate may play a more important role in foxtail pine recruitment than fire.

  • 10. Bancroft, Larry. 1979. Fire management plan: Sequoia and Kings Canyon National Parks. San Francisco, CA: U.S. Department of the Interior, National Park Service, Western Region. 190 p. [11887]
  • 48. Kiefer, M. 1991. Forest age structure, species composition, and fire disturbance in the southern Sierra Nevada subalpine zone. Unpublished report [submitted to the Sequoia Natural History Association]. Tucson, AZ: University of Arizona, Laboratory of Tree-Ring Research. [48233]

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Immediate Effect of Fire

More info for the term: surface fire

Lightning damage to foxtail pines is common, especially to trees in the upper subalpine zone [82]. Low-severity surface fire leaves basal scars on foxtail pines and kills some trees [48,82].
  • 82. Ryerson, A. Diane. 1983. Population structure of Pinus balfouriana Grev. & Balf. along the margins of its distribution area in the Sierran and Klamath regions of California. Sacramento, CA: California State University. 197 p. Thesis. [48204]
  • 48. Kiefer, M. 1991. Forest age structure, species composition, and fire disturbance in the southern Sierra Nevada subalpine zone. Unpublished report [submitted to the Sequoia Natural History Association]. Tucson, AZ: University of Arizona, Laboratory of Tree-Ring Research. [48233]

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Post-fire Regeneration

More info for the terms: adventitious, initial off-site colonizer, tree

POSTFIRE REGENERATION STRATEGY [94]:
Tree without adventitious bud/root crown
Initial off-site colonizer (off-site, initial community)
  • 94. Stickney, Peter F. 1989. Seral origin of species originating in northern Rocky Mountain forests. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 10 p. [20090]

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Fire Ecology

More info for the terms: basal area, duff, fire intensity, fire occurrence, fire regime, fire-return interval, fuel, litter, mean fire-return interval, resistance, severity, tree

Fire adaptations: Foxtail pine has many characteristics of a fire survivor [87]. Some of its morphological characteristics are similar to ponderosa pine, a highly fire-adapted species [4]. Like ponderosa pine, foxtail pine is a long-lived tree with a large-diameter bole, thick bark, and large-diameter branches [8,45]. Branches are generally sparse and self-pruning in southern foxtail pine, although thin branching and a self-pruning habit are less common in northern foxtail pine [39,59,67]. Few fire studies on foxtail pine have been conducted; however, Ryerson [82] found mature, fire-scarred southern foxtail pines throughout the tree's distribution. As further evidence of foxtail pine's ability to survive fire, Keifer [48] reported that in the Sierra lodgepole pine-southern foxtail pine ecotone in Sequoia-Kings Canyon National Park, foxtail pines were uneven-aged and showed multiple fire scars, while Sierra lodgepole pines were even-aged and showed no evidence of scarring. More fire history studies are needed on foxtail pine.

Foxtail pine seedlings pioneer on burned sites. The seeds are small, light, and have large wings [39,70,76], suggesting the possibility of foxtail pine seed dispersal onto burns from on- and off-site parent trees. In Sequoia-Kings Canyon National Park, Ryerson [82] found southern foxtail pine seedlings on 2 burned sites. On the 1st burn, seedlings established near 4 lightning-killed, mature trees. On the 2nd burn, foxtail pine seedlings grew in openings created when fire burned across a ridgetop. Further studies are needed on patterns of foxtail pine seed dispersal and seedling establishment after fire.

FIRE REGIMES: Fires are infrequent, and are generally of low severity, in subalpine regions of the southern Sierra Nevada. Scant litter production and discontinuous fuels do not promote fire spread in foxtail pine communities. Fire intensity tends to decrease when lower-elevation fires burn into southern foxtail pine. Fire spread slows; or, fires may extinguish due to lack of fuels [10,20,21,49,65]. Although foxtail pine sites receive more lightning strikes than lower-elevation forests, ignitions are uncommon [102,103]. Rocky, highly dissected foxtail pine habitats rarely sustain large fires. In a fire history study, Keifer [48] found frequent fire in Sierra lodgepole pine, but only occasional fires in southern foxtail pine sites. The National Park Service [101] classifies fire occurrence as "very low" in subalpine conifer zones of Sequoia-King Canyon National Park, with a mean fire-return interval of 187 years and a maximum recorded fire-return interval of 508 years. Caprio and Lineback [21] found southern foxtail and whitebark pine communities of Sequoia-Kings Canyon National Park had the longest return fire intervals of all plant communities in the Park. Estimated area burned in southern foxtail pine communities averaged 145 acres/year (168 ha/yr) with a mean fire-return interval of 187 years. Estimated burn area extended to 153 acres/year (62 ha/year) under the maximum mean fire-return interval of 508 years. Fire scar data from 2 watersheds show few fires in foxtail-whitebark pine communities of Sequoia-Kings Canyon National Park from 1700 to 2000: 7 on north aspects and 3 on south aspects [19]. Differences in fire-return intervals between aspects were not significant [18].

The fire ecology of upper subalpine zones of California is poorly understood [101]. This is particularly true for northern foxtail pine, for which fire ecology and fire regime information are nearly absent. Thornburgh [95] found white fir-mountain hemlock communities of the Marble Mountains, where northern foxtail pine is an associate, experience a regime of mixed low- and moderate-severity fires. Fire effects and postfire recruitment of foxtail pine were not reported. Further documentation and research are needed on the fire ecology of foxtail pine and other subalpine communities of  California.

Occasionally, large, stand-replacing fires occur in southern foxtail pine [91]. For example, The 1949 Kern Canyon 2 Fire burned 1,100 acres (445 ha) of southern foxtail and Jeffrey pine habitat in Sequoia-Kings Canyon National Park. Ignited by lightning on 13 July, it was controlled by 31 July. Southwesterly winds up to 40 mph (64 km/hr) caused crowning and spotting. Steep, rugged terrain contributed to fast fire spread and resistance to control [10].

Fuels: Foxtail pine snags and woody debris are highly resinous, and are slow to decay in high-elevation habitats. In Sequoia-Kings Canyon National Park, downed foxtail pines that have been dead for over 1,000 years still retain medium-sized (>0.8-inch (2-cm) diameter) or larger branches [62]. Foxtail pine communities are not typically highly flammable though, because woody fuels are limited and discontinuous [65,91], and litter is sparse [81]. Live fuels are also scant. A 1978 fuel inventory in Sequoia-Kings Canyon National Park showed a mean of 10 tons/acre in foxtail and other subalpine types [10]. Basal area and litter quantity decreased with elevation, although litter quality (N:C ratio) increased with elevation [64]. The live understory is typically sparse in foxtail pine communities. Lloyd and Graumlich [63] found less than 1 plant/m² in southern foxtail pine understories in Sequoia-Kings Canyon National Park. Van Wagtendonk and others [105] reported the following fuelbed characteristics for southern foxtail pine:

Woody fuel depth Litter depth Duff depth Litter & duff depth
1.24 cm 0.19 cm 1.60 cm 1.79 cm

Quantitative measures of physical fuel properties such as surface-to-volume ratios are used in fuel models. By fuel size class, van Wagtendonk and others [104] provide mean surface-to-volume ratio, diameter and squared quadratic mean diameter, and angles of inclination tables for southern foxtail pine and other Sierra Nevada conifers.

The following table provides fire-return intervals for plant communities where foxtail pine occurs.  For further information, see the FEIS summary on the dominant species listed below.

Community or Ecosystem Dominant Species Fire-Return Interval Range (years)
whitebark pine Pinus albicaulis 50-200 [1,3]
Sierra lodgepole pine Pinus contorta var. murrayana 35-200 [5]
mountain hemlock Tsuga mertensiana 35 to > 200 [5]
Fire-return intervals for these species vary widely; trends in variation are noted in the species reviews.
  • 5. Arno, Stephen F. 2000. Fire in western forest ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 97-120. [36984]
  • 1. Agee, James K. 1994. Fire and weather disturbances in terrestrial ecosystems of the eastern Cascades. Gen. Tech. Rep. PNW-GTR-320. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 52 p. (Everett, Richard L., assessment team leader; Eastside forest ecosystem health assessment; Hessburg, Paul F., science team leader and tech. ed., Volume III: assessment). [22991]
  • 39. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. [21992]
  • 3. Arno, Stephen F. 1976. The historical role of fire on the Bitterroot National Forest. Res. Pap. INT-187. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 29 p. [15225]
  • 8. Bailey, D. K. 1970. Phytogeography and taxonomy of Pinus subsection Balfourianae. Annals of the Missouri Botanical Garden. 57: 210-249. [375]
  • 45. Jackson, James F.; Adams, Dean C.; Jackson, Ursula B. 1999. Allometry of constitutive defense: a model and a comparative test with tree bark and fire regime. The American Naturalist. 153(6): 614-632. [31152]
  • 59. Lanner, Ronald M. 1999. Conifers of California. Los Olivos, CA: Cachuma Press. 274 p. [30288]
  • 10. Bancroft, Larry. 1979. Fire management plan: Sequoia and Kings Canyon National Parks. San Francisco, CA: U.S. Department of the Interior, National Park Service, Western Region. 190 p. [11887]
  • 62. Lloyd, Andrea H. 1997. Response of tree-line populations of foxtail pine (Pinus balfouriana) to climate variation over the last 1000 years. Canadian Journal of Forest Research. 27(6): 936-942. [47903]
  • 63. Lloyd, Andrea H.; Graumlich, Lisa J. 1997. Holocene dynamics of treeline forests in the Sierra Nevada. Ecology. 78(4): 1199-1210. [27723]
  • 64. Lloyd, Andrea. 1998. Growth of foxtail pine seedlings at treeline in the southeastern Sierra Nevada, California, U.S.A. Ecoscience. 5(2): 250-257. [48110]
  • 67. Mastroguiseppe, R. J.; Mastroguiseppe, J. D. 1980. A study of Pinus balfouriana Grev. & Balf. (Pinaceae). Systematic Botany. 5(1): 86-104. [1546]
  • 76. Munz, Philip A. 1974. A flora of southern California. Berkeley, CA: University of California Press. 1086 p. [4924]
  • 81. Rundel, Philip W.; Parsons, David J.; Gordon, Donald T. 1977. Montane and subalpine vegetation of the Sierra Nevada and Cascade Ranges. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley & Sons: 559-599. [4235]
  • 82. Ryerson, A. Diane. 1983. Population structure of Pinus balfouriana Grev. & Balf. along the margins of its distribution area in the Sierran and Klamath regions of California. Sacramento, CA: California State University. 197 p. Thesis. [48204]
  • 4. Arno, Stephen F. 1988. Fire ecology and its management implications in ponderosa pine forests. In: Baumgartner, David M.; Lotan, James E., compilers. Ponderosa pine: The species and its management: Symposium proceedings; 1987 September 29 - October 1; Spokane, WA. Pullman, WA: Washington State University, Cooperative Extension: 133-139. [9410]
  • 70. McCune, Bruce. 1988. Ecological diversity in North American pines. American Journal of Botany. 75(3): 353-368. [5651]
  • 87. Schwilk, Dylan W.; Ackerly, David D. 2001. Flammability and serotiny as strategies: correlated evolution in pines. Oikos. 94(2): 326-336. [41219]
  • 95. Thornburgh, Dale A. 1995. The natural role of fire in the Marble Mountain Wilderness. In: Brown, James K.; Mutch, Robert W.; Spoon, Charles W.; Wakimoto, Ronald H., technical coordinators. Proceedings: symposium on fire in wilderness and park management; 1993 March 30 - April 1; Missoula, MT. Gen. Tech. Rep. INT-GTR-320. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 273-274. [26229]
  • 102. van Wagtendonk, J. W. 1991. Spatial analysis of lightning strikes in Yosemite National Park. In: Andrews, Patricia L.; Potts, Donald F., eds. Proceedings, 11th conference on fire and forest meteorology; 1991 April 16-19; Missoula, MT. SAF Publication 91-04. Bethesda, MD: Society of American Foresters: 605-611. [48238]
  • 103. van Wagtendonk, Jan W. 1991. GIS applications in fire management and research. In: Nodvin, Stephen C.; Waldrop, Thomas A., eds. Fire and the environment: ecological and cultural perspectives: Proceedings of an international symposium; 1990 March 20-24; Knoxville, TN. Gen. Tech. Rep. SE-69. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station: 212-214. [48237]
  • 104. van Wagtendonk, Jan W.; Benedict, James M.; Sydoriak, Walter M. 1996. Physical properties of woody fuel particles of Sierra Nevada conifers. International Journal of Wildland Fire. 6(3): 117-123. [29412]
  • 105. van Wagtendonk, Jan W.; Benedict, James M.; Sydoriak, Walter M. 1998. Fuel bed characteristics of Sierra Nevada conifers. Western Journal of Applied Forestry. 13(3): 73-84. [28859]
  • 20. Caprio, Anthony C.; Graber, David M. 2000. Returning fire to the mountains: Can we successfully restore the ecological role of pre-Euroamerican FIRE REGIMES to the Sierra Nevada? In: Cole, David N.; McCool, Stephen F.; Borrie, William T.; O'Loughlin, Jennifer, comps. Wilderness science in a time of change conference--Volume 5: wilderness ecosystems, threats, and management; 1999 May 23-27; Missoula, MT. Proceedings RMRS-P-15-VOL-5. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 233-241. [40571]
  • 91. Skinner, Carl N.; Chang, Chi-ru. 1996. FIRE REGIMES, past and present. In: Status of the Sierra Nevada. Sierra Nevada Ecosystem Project: Final report to Congress. Volume II: Assessments and scientific basis for management options. Wildland Resources Center Report No. 37. Davis, CA: University of California, Centers for Water and Wildland Resources: 1041-1069. [28975]
  • 18. Caprio, A. C. 2000. Reconstructing attributes of pre-European settlement fire at a watershed scale, Sequoia and Kings Canyon National Parks. In: Ecological Society of America: Proceedings, 85th annual meeting; 2000 August 6-10; Snowbird, UT, [Online]. Available: ttp://abstract.co.allenpress.com/pwet/esa2000/ [2004, June 21]. [48207]
  • 19. Caprio, A. C. 2001. Temporal and spatial dynamics of pre-Euro-American fire at a watershed scale, Sequoia and Kings Canyon National Parks. In: Sugihara, N. G.; Morales, M. E.; and Morales, T. J., eds. Emerging policies and new paradigms: Proceedings of the conference on fire management; 1999 November 16-19; San Diego, CA. Association for Fire Ecology Misc. Publication No. 2. [Place of publication unknown]: Association for Fire Ecology: 1-17. [48208]
  • 21. Caprio, Anthony C.; Lineback, Pat. 2002. Pre-twentieth century fire history of Sequoia and Kings Canyon National Park: A review and evaluation of our knowledge. In: Sugihara, Neil G.; Morales, Maria; Morales, Tony, eds. Fire in California ecosystems: integrating ecology, prevention and management: Proceedings of the symposium; 1997 November 17-20; San Diego, CA. Misc. Pub. No. 1. [Place of publication unknown]: Association for Fire Ecology: 180-199. [46205]
  • 48. Kiefer, M. 1991. Forest age structure, species composition, and fire disturbance in the southern Sierra Nevada subalpine zone. Unpublished report [submitted to the Sequoia Natural History Association]. Tucson, AZ: University of Arizona, Laboratory of Tree-Ring Research. [48233]
  • 49. Kilgore, Bruce M. 1981. Fire in ecosystem distribution and structure: western forests and scrublands. In: Mooney, H. A.; Bonnicksen, T. M.; Christensen, N. L.; [and others], technical coordinators. Proceedings of the conference: FIRE REGIMES and ecosystem properties; 1978 December 11-15; Honolulu, HI. Gen. Tech. Rep. WO-26. Washington, DC: U.S. Department of Agriculture, Forest Service: 58-89. [4388]
  • 65. Lotan, James E.; Alexander, Martin E.; Arno, Stephen F.; [and others]. 1981. Effects of fire on flora: A state-of-knowledge review: Proceedings of the national fire effects workshop; 1978 April 10-14; Denver, CO. Gen. Tech. Rep. WO-16. Washington, DC: U.S. Department of Agriculture, Forest Service. 71 p. [1475]
  • 101. U.S. Department of the Interior, National Park Service, Sequoia & Kings Canyon National Parks. 2004. Sequoia and Kings Canyon fire management plan: Fire and fuels management plan, [Online]. Sequoia & Kings Canyon National Parks (Producer). Available: http://www.nps.gov/seki/fire/ffmp/seki_ffmp_fmp.htm [2004, June 14]. [48206]

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Successional Status

More info on this topic.

More info for the terms: basal area, cover, mesic, succession, tree

Foxtail pine is shade intolerant, requiring open, sunny locations throughout its life cycle [8,9,82,86]. Foxtail pine pioneers on serpentine and high-elevation subalpine sites [30]. It competes poorly on nutrient-rich, mesic, and low-subalpine sites [77,86]. Foxtail pine is generally noninvasive [79]; however, it has extended its distribution into the California red fir zone in times of global cooling [82]. On high-elevation, ultramafic or dry granitic sites, foxtail pine is not threatened by successional replacement by shade-tolerant conifers such as California red fir and mountain hemlock because no other tree is as well adapted to the harsh sites that foxtail pine occupies [14,30]. On more favorable sites, successional replacement of foxtail pine by mountain hemlock and firs may be occurring. Research is needed on successional patterns in foxtail pine communities on mesic, nonserpentine sites.

A resurvey in Sequoia-Kings Canyon National Park showed that in 27 years, southern foxtail pine basal area and cover increased 8% and 16%, respectively. The changes were entirely due to foxtail pine diameter growth; in 27 years there had been no foxtail pine mortality, and no ingrowth of foxtail pine or other tree species, on the study plots [80]. To date (2004), there are no studies of succession in foxtail pine communities following fire, avalanche, or other disturbances. Studies documenting postdisturbance recruitment and succession in foxtail  pine communities are needed.

  • 8. Bailey, D. K. 1970. Phytogeography and taxonomy of Pinus subsection Balfourianae. Annals of the Missouri Botanical Garden. 57: 210-249. [375]
  • 30. Eckert, Andrew J.; Sawyer, John O. 2002. Foxtail pine importance and conifer diversity in the Klamath Mountains and southern Sierra Nevada, California. Madrono. 49(1): 33-45. [42265]
  • 77. Oline, David K.; Mitton, Jeffry B.; Grant, Michael C. 2000. Population and subspecific genetic differentiation in the foxtail pine (Pinus balfouriana). Evolution. 54(5): 1813-1819. [48101]
  • 80. Roy, D. Graham; Vankat, John L. 1999. Reversal of human-induced vegetation changes in Sequoia National Park, California. Canadian Journal of Forest Research. 29(4): 399-412. [36282]
  • 82. Ryerson, A. Diane. 1983. Population structure of Pinus balfouriana Grev. & Balf. along the margins of its distribution area in the Sierran and Klamath regions of California. Sacramento, CA: California State University. 197 p. Thesis. [48204]
  • 86. Sawyer, John O.; Thornburgh, Dale A. 1977. Montane and subalpine vegetation of the Klamath Mountains. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley & Sons: 699-732. [685]
  • 9. Baker, Frederick S. 1949. A revised tolerance table. Journal of Forestry. 47: 179-181. [20404]
  • 14. Benedict, Nathan B. 1982. Mountain meadows: stability and change. Madrono. 29(3): 148-153. [11083]
  • 79. Rejmanek, Marcel; Richardson, David M. 1996. What attributes make some plant species more invasive? Ecology. 77(6): 1655-1661. [27088]

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Regeneration Processes

More info for the terms: fresh, natural, presence, selection

Environmental interactions that foster foxtail pine recruitment are poorly understood. Climate and water balance may be the primary factors driving foxtail pine establishment, with best recruitment during periods of warm, wet winters and cool summer temperatures [33,36,37,62]. Dendrochronologists studying a 3,500-year history of southern foxtail pine recruitment and death rates in Sequoia-Kings Canyon National Park found 2 periods of poor southern foxtail pine recruitment. The 1st was during a drought lasting decades (950-550 years BP); the other was an extended period of below-average temperatures (450-50 years BP). Death rate was relatively stable over the last 1,000 years, averaging about 0.05% per capita. Death appeared to be due to local or endogenous factors except during periods of extreme climate fluctuation. Mortality spiked during the periods of extended drought and extended cold. Overall recruitment rate was slightly higher than death rate, averaging around 0.06% per capita [62,63]. As of this writing (2004), field studies on the seed germination and seedling establishment stages of foxtail pine's life cycle are lacking. Further studies are needed on the regeneration requirements and life cycle of foxtail pine.

Barriers to regeneration: Domestic livestock grazing may adversely affect foxtail pine regeneration in areas where grazing is still practiced. Vankat [106] found southern foxtail pine in Sequoia-Kings Canyon National Park showed a pulse of recruitment from 1890-1895. That period coincides with a period of reduced domestic sheep grazing in the southern Sierra Nevada.

White pine blister rust (Cronartium ribicola) affects the ability of 5-needle pines to reproduce by killing cone-bearing branch tips. An infected northern foxtail pine population on the Klamath National Forest (see Other Management Considerations) shows poor recruitment, although it is uncertain at this time if blister rust is responsible. Levels of blister rust infection in foxtail pine are being monitored [99].

Breeding system: Allozyme surveys show that genetic diversity is low in foxtail pine compared to other pine species. There is more genetic differentiation among than within populations. Interpopulation genetic diversity is particularly pronounced in northern foxtail pine, which tends to have small (300-600 individuals), isolated populations, and restricted between-population gene flow. Natural selection for serpentine tolerance, global warming (see Other Management Considerations, Climate), and genetic drift have likely contributed to northern foxtail pine's low genetic diversity [77].

Pollination: Foxtail pine is wind pollinated [59].

Seed production: Foxtail pine 1st produces cones at 20 to 50 years of age [52,82]. The cone cycle (development through maturity) takes 5 to 6 years  [28]. There is usually a 5- to 6-year interval between large cone crops [52]. Environmental conditions promoting large crops are undocumented (as of 2004).

Seed dispersal: Foxtail pine seed is dispersed by wind [58,59]. How long seed is retained in the cone, and whether it survives fire and disperses from cones onto burns, is poorly documented (as of 2004). Likewise, average range of dispersal for wind-blown foxtail pine seed is unknown, making it difficult to predict the potential for long-range foxtail pine seed dispersal onto burns or other open seedbeds.

Although Clark's nutcrackers disperse bristlecone pine seeds, there have been no sightings of the birds dispersing the smaller seeds of foxtail pine [60,70]. Trees growing from Clark's nutcracker caches often have multiple, genetically distinct stems [58,59]. The typical single-stemmed habit [8,10,82] of foxtail pine suggests that Clark's nutcracker dispersal and caching is unusual. Ryerson [83], however, noted the presence of  a few multiple-stemmed trees throughout foxtail pine's distribution, suggesting the possibility of Clark's nutcracker seed dispersal and caching. Genetic identities of multiple-stemmed foxtail pine "individuals" have not been determined. Further investigation is needed on mechanisms of seed dispersal for foxtail pine.

Seed banking: No information is available on this topic.

Germination: Seeds require stratification [25,28]. Fresh, stratified southern foxtail pine seed collected in Sequoia-Kings Canyon National Park showed 86% germination. After 9.4 years in cold storage, the same seed lot showed 72% germination [75].

Seedling establishment/growth: Based on limited information, foxtail pine seedling establishment appears to be episodic, occurring during periods of mild, wet winters [62,82].

Foxtail pine is a slow-growing conifer [59,70]. Best growth of southern foxtail pine occurs in years with relatively warm, wet winters and cool summers [33,36,37,62]. Studies on growth rates of foxtail pine are limited. One study found relative height growth rates of 0.2 to 0.9 inch (0.5-2.3 cm) per year for seedlings in Sequoia-Kings Canyon National Park. Seedlings in open, high-elevation sites tended to grow taller than seedlings in lower-elevation, forested areas [92]. For mature trees, another Sequoia-Kings Canyon study found trees at lower elevations (<8,200 feet (2,500 m)) had greater relative growth rates compared to trees at high elevations (>9,800 feet (3,000 m)). Relative growth rates were 6.7-9.1 inches/100 years compared to 2.4-3.1 inches/100 years (17-23 cm/100 yrs vs. 6-8 cm/100 yrs), at low and high elevations, respectively [82].

  • 8. Bailey, D. K. 1970. Phytogeography and taxonomy of Pinus subsection Balfourianae. Annals of the Missouri Botanical Garden. 57: 210-249. [375]
  • 59. Lanner, Ronald M. 1999. Conifers of California. Los Olivos, CA: Cachuma Press. 274 p. [30288]
  • 10. Bancroft, Larry. 1979. Fire management plan: Sequoia and Kings Canyon National Parks. San Francisco, CA: U.S. Department of the Interior, National Park Service, Western Region. 190 p. [11887]
  • 25. Critchfield, William B. 1977. Hybridization of foxtail and bristlecone pines. Madrono. 24(4): 193-244. [713]
  • 33. Garfin, Gregg M. 1998. Relationships between winter atmospheric circulation patterns and extreme tree growth anomalies in the Sierra Nevada. International Journal of Climatology. 18(7): 725-740. [40071]
  • 36. Graumlich, Lisa J. 1991. Subalpine tree growth, climate, and increasing CO2: an assessment of recent growth trends. Ecology. 72(1): 1-11. [47892]
  • 37. Graumlich, Lisa J. 1993. A 1000-year record of temperature and precipitation in the Sierra Nevada. Quaternary Research. 39(2): 249-255. [29155]
  • 60. Lanner, Ronald M.; Hutchins, Harry E.; Lanner, Harriette A. 1984. Bristlecone pine and Clark's nutcracker: probable interaction in the White Mountains, California. Great Basin Naturalist. 44(2): 357-360. [48202]
  • 62. Lloyd, Andrea H. 1997. Response of tree-line populations of foxtail pine (Pinus balfouriana) to climate variation over the last 1000 years. Canadian Journal of Forest Research. 27(6): 936-942. [47903]
  • 63. Lloyd, Andrea H.; Graumlich, Lisa J. 1997. Holocene dynamics of treeline forests in the Sierra Nevada. Ecology. 78(4): 1199-1210. [27723]
  • 77. Oline, David K.; Mitton, Jeffry B.; Grant, Michael C. 2000. Population and subspecific genetic differentiation in the foxtail pine (Pinus balfouriana). Evolution. 54(5): 1813-1819. [48101]
  • 82. Ryerson, A. Diane. 1983. Population structure of Pinus balfouriana Grev. & Balf. along the margins of its distribution area in the Sierran and Klamath regions of California. Sacramento, CA: California State University. 197 p. Thesis. [48204]
  • 83. Ryerson, Diane. 1984. Krummholz foxtail pines. Fremontia. 11(4): 30. [47919]
  • 92. Steinhoff, R. J. 1972. White pines of western North America and Central America. In: Bingham, Richard: Hoff, Raymond J., tech. coords. In: Biology of rust resistance in forest trees: Proceedings of a NATO/IUFRO Advanced Study Institute; 1969 August 17-24; Washington, DC. Misc. Pub. 1221. U.S. Department of Agriculture, Forest Service: 215-232. [30287]
  • 106. Vankat, John Lyman. 1970. Vegetation change in Sequoia National Park, California. Davis, CA: University of California. 197 p. Dissertation. [43459]
  • 52. Krugman, Stanley L.; Jenkinson, James L. 1974. Pinus L. pine. In: Schopmeyer, C. S., tech. cood. Seeds of woody plants in the United States. Agric. Handb. 450. Washington, D.C.: U.S. Department of Agriculture, Forest Service: 598-638. [37725]
  • 58. Lanner, Ronald M. 1996. Made for each other: a symbiosis of birds and pines. New York: Oxford University Press. 160 p. [29914]
  • 70. McCune, Bruce. 1988. Ecological diversity in North American pines. American Journal of Botany. 75(3): 353-368. [5651]
  • 75. Mirov, N. T. 1961. Composition of gum turpentines of pines. Tech. Bull. No. 1239. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station. 158 p. [22164]
  • 28. Dumroese, R. K.; Landis, T. D., Wenny, D. L. 1998. Raising forest tree seedlings at home: simple methods for growing conifers of the Pacific Northwest from seeds. Station Contribution Number 860, [Online]. Moscow, ID: University of Idaho (Producer). Available: http://www.uidaho.edu/seedlings/howtogrow/manual-menu.htm [2004, June 17]. [48212]
  • 99. U.S. Department of Agriculture, Forest Service, Pacific Southwest Region, Forest Pest Management. 1995. Forest pest conditions in California --1995. [Online]. In: Forest health protection. California Forest Pest Council (Producer). Available: http://www.fs.fed.us/r5/spf/publications1995.htm [2004, June 24]. [48213]

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Growth Form (according to Raunkiær Life-form classification)

More info on this topic.

More info for the term: phanerophyte

RAUNKIAER [78] LIFE FORM:
Phanerophyte
  • 78. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843]

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Life Form

More info for the term: tree

Tree

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Broad-scale Impacts of Fire

Fire and climate may play an interactive role in determining the California red fir-foxtail pine ecotone. California red firs (Abies magnifica var. magnifica and A. m. var. shastensis) have thicker bark and are more fire-tolerant than most firs [7], but are not as fire-tolerant as foxtail pine. Warming climate and more frequent fires may promote foxtail pine invasions into lower-elevation California red fir communities [82].
  • 82. Ryerson, A. Diane. 1983. Population structure of Pinus balfouriana Grev. & Balf. along the margins of its distribution area in the Sierran and Klamath regions of California. Sacramento, CA: California State University. 197 p. Thesis. [48204]
  • 7. Atzet, Thomas; McCrimmon, Lisa A. 1990. Preliminary plant associations of the southern Oregon Cascade Mountain province. Grants Pass, OR: U.S. Department of Agriculture, Forest Service, Siskiyou National Forest. 330 p. [12977]

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Life History and Behavior

Cyclicity

Phenology

More info on this topic.

Pollen dispersal and pollination of new foxtail pine cones occurs in July and August. Mature cones open and disperse seed in September and October [27,52]. Common garden studies show foxtail and bristlecone pines open their cones later in the season than other North American pine species [25]. Little is known of foxtail pine's seed biology and the phenological development of seedlings. Further work is needed in this area.
  • 25. Critchfield, William B. 1977. Hybridization of foxtail and bristlecone pines. Madrono. 24(4): 193-244. [713]
  • 52. Krugman, Stanley L.; Jenkinson, James L. 1974. Pinus L. pine. In: Schopmeyer, C. S., tech. cood. Seeds of woody plants in the United States. Agric. Handb. 450. Washington, D.C.: U.S. Department of Agriculture, Forest Service: 598-638. [37725]
  • 27. Duffield, J. W. 1953. Pine pollen collection dates--annual and geographic variation. For. Res. Notes No. 85. Berkeley, CA: U.S. Department of Agriculture, Forest Service, California Forest and Range Experiment Station. 9 p. [17970]

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Molecular Biology and Genetics

Molecular Biology

Barcode data: Pinus balfouriana

The following is a representative barcode sequence, the centroid of all available sequences for this species.


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Statistics of barcoding coverage: Pinus balfouriana

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 32
Specimens with Barcodes: 37
Species With Barcodes: 1
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
NT
Near Threatened

Red List Criteria

Version
3.1

Year Assessed
2013

Assessor/s
Farjon, A.

Reviewer/s
Thomas, P. & Stritch, L.

Contributor/s

Justification
Based on comprehensive sampling of specimens held in herbaria in California and elsewhere, and the fact that very few if any stands of this species would have an area of occupancy (AOO) larger than 4 km² an AOO of only 136 km² was calculated. There are two main areas, separated by nearly 500 km, but the northern area has two locations, one with a much smaller subpopulation than the other. Fragmentation and number of locations therefore also fall within the threshold for Endangered; however, the population appears to be stable at present and there is no evidence of past decline within the last few hundred years. Climate change and air pollution (the latter only relevant to the southern subpopulation) are potential threats. It is therefore appropriate to list this species as Near Threatened (almost qualifies for listing under criterion B2ab).
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The World Conservation Union's Species Survival Commission (IUCB-SSC) lists foxtail pine as a lower risk, conservation-dependent species [45].
  • 45. Jackson, James F.; Adams, Dean C.; Jackson, Ursula B. 1999. Allometry of constitutive defense: a model and a comparative test with tree bark and fire regime. The American Naturalist. 153(6): 614-632. [31152]

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National NatureServe Conservation Status

United States

Rounded National Status Rank: N4 - Apparently Secure

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NatureServe Conservation Status

Rounded Global Status Rank: G4 - Apparently Secure

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Population

Population
The total number of mature trees is not known, but the population is fragmented, first by a gap of nearly 500 km between the northern and southern (sub) populations, and second by the fact that individual stands are (widely) scattered within these two areas. Numbers of trees in each of these stands vary between a few score to many hundreds. Regeneration and growth to maturity are extremely slow in most stands and may be episodic; this means that if little or no regeneration is observed in a stand at present this will not be evidence of decline. A tree that lives for several millennia only needs to produce offspring a few times in that whole period to replace itself and maintain the population.

Population Trend
Stable
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Threats

Major Threats
This species may be at risk in the long term from climate change if this is continuing to accelerate, possibly bringing in competitors and/or pathogens it may not be able to cope with. At present, both disjunct populations are well protected within National Parks and National Forest Wilderness Areas and unaffected by long-term effects of fire suppression in forests as the trees usually occur in remote subalpine locations where such measures have not been undertaken. It would be profitable to study the (aut-)ecology of this species in more detail in order to be able to estimate risks under various climate change scenarios. The southern (sub)population in Kings Canyon N.P. and Sequoia N.P. are subject to air pollution from major urban centres such as Los Angeles. The effect, if any, on this species is as yet unknown and needs to be researched.
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Management

Conservation Actions

Conservation Actions
This species is mostly present within protected areas, including famous national parks like Kings Canyon and Sequoia N.P.
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Management considerations

More info for the terms: interference, resistance, tree

Damaging bioagents:
Foxtail pine is highly susceptible to white pine blister rust,
a usually fatal fungal disease that affects 5-needle pines [40,71,84].
In the greenhouse, Hoff and others [40] inoculated 18 species of Eurasian and
North American 5-needle pine seedlings with blister rust. Of the 18 species,
foxtail and southwestern white pine (Pinus strobiformis) showed least
resistance to blister rust. All of the 92 inoculated foxtail pines became
infected. Other studies show a 75-100% seedling infection rate [93]. Although the
mediterranean climate of California once protected all but the northernmost populations of 5-needle pines from blister
rust, that is no longer true. Some northern foxtail pines on the Scott River
District of the Klamath National Forest have blister rust [96,97,99].
As of this writing (2004), blister rust has not been detected in southern
foxtail pine, although western white pine and sugar pine (P. lambertiana)
in Sequoia-Kings Canyon National Park are infected [84,99].

Blister rust-infected trees may take from 2 years to decades to succumb, but
infection is always fatal [41,42].
Gooseberries and currants (Ribes spp.) are the primary host of white
pine blister rust. Life cycle of white pine blister rust is complex.
Gitzendanner and others [35]
and McDonald and Hoff [71]
provide details of the rust's life history and ecology. Hoff [41]
provides a diagnostic guide to aid managers in recognizing symptoms of blister
rust infection in white pines. There are no known methods of controlling blister
rust
[47]. Fungicide application, pruning infected tree branches, and/or
removing Ribes spp. have neither eliminated nor controlled white pine
blister rust [22,71], and such treatments have undesirable ecological effects
[47]. For further information on management of white pine blister rust, see Samman and
others [84].

Some northern foxtail pines on the Scott River District show phenological
resistance to blister rust. Identification and breeding programs for these
genetically valuable, blister-rust resistant individuals are crucial to an
integrated strategy for protecting and restoring foxtail and other white pines
[40,71,84]. Breeding programs for  blister rust-resistant foxtail pines
are being implemented [99].
Other damaging bioagents:
Foxtail pines are susceptible to
mountain pine beetle attacks [107]. Two rare species of Pityophthorus
bark beetles may feed primarily on foxtail pine [17]. While contributing to
biodiversity, little is known of the impacts of these Pityophthorus bark beetles to foxtail
pine. Limber pine dwarf-mistletoe (Arceuthobium cyanocarpum) occasionally
infects foxtail pine [50,69,73]. A fungal needle cast (Lophodermium
durilabrum) has caused minor damage to northern foxtail pines in the
Marble Mountains [72].
Climate
affects foxtail pine's elevational range. For most of the period for
which tree records are available (~3,500 years), southern foxtail pine
has existed above present treeline [62,63]. For example, Vankat
[106] found dead stands of foxtail pine above present timberline (10,800-
11,200 feet (3,300-3,400 m))
on the Kern River Watershed in Sequoia-Kings Canyon National Park. These "ghost
forests" may be relicts of foxtail pines that died during a period of global
warming [56]. Lloyd
and Graumlich [63] documented 3 episodes where southern foxtail pine expanded
upslope. Although the data are somewhat unclear [55,89], these
expansions appear to have occurred during relatively warm, wet periods. Presently, southern foxtail pine is expanding its
range both upslope and laterally into subalpine meadows and previously untreed
east slopes. This expansion has been explained as a response to global
warming [62,64], or due to a combination of factors including global warming, low conifer
diversity (and consequent lack of growth interference for foxtail pine in the upper elevations
of the southern Sierra Nevada), and stochasticity [68,82,88].
Northern foxtail pine is threatened by global warming. Already restricted to
a relatively few high-elevation peaks, there are no higher-elevation refugia for
the Klamath Mountains subspecies to migrate to. Many northern foxtail pine populations
are being "squeezed off the tops of mountains that are insufficiently high to
provide suitable habitat" [77].

  • 17. Bright, Donald E., Jr. 1971. New species, new synonymies and new records of bark-beetles from Arizona and California. The Pan-Pacific Entomologist. 47(1): 63-70. [48106]
  • 22. Carlson, Clinton E. 1978. Noneffectiveness of Ribes eradication as a control of white pine blister rust in Yellowstone National Park. Rep. No. 78-18. Missoula, MT: U.S. Department of Agriculture, Forest Service, Northern Region, State & Private Forestry, Forest Insect & Disease Management. 6 p. [22749]
  • 40. Hoff, R.; Bingham, R. T.; McDonald, G. I. 1980. Relative blister rust resistance of white pines. European Journal of Forest Pathology. 10(5): 307-316. [1177]
  • 41. Hoff, Ray J. 1992. How to recognize blister rust infection on whitebark pine. Res. Note INT-406. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 7 p. [19509]
  • 42. Hoff, Ray; Hagle, Susan. 1990. Diseases of whitebark pine with special emphasis on white pine blister rust. In: Schmidt, Wyman C.; McDonald, Kathy J., compilers. Proceedings--symposium on whitebark pine ecosystems: ecology and management of a high-mountain resource; 1989 March 29-31; Bozeman, MT. Gen Tech. Rep. INT-270. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 179-190. [5936]
  • 47. Keane, Robert E.; Arno, Stephen F. 2001. Restoration concepts and techniques. In: Tomback, Diana F.; Arno, Stephen F.; Keane, Robert E., eds. Whitebark pine communities: Ecology and restoration. Washington, DC: Island Press: 367-400. [36711]
  • 50. Kimmey, J. W. 1957. Dwarfmistletoes of California and their control. Tech. Pap. No. 19. Berkeley, CA: U.S. Department of Agriculture, Forest Service, California Forest and Range Experiment Station. 12 p. [16464]
  • 55. LaMarche, Valmore C., Jr. 1974. Paleoclomatic inferences from long tree-ring records. Science. 183(4129): 1043-1048. [1391]
  • 56. LaMarche, Valmore C., Jr.; Mooney, Harold A. 1967. Altithermal timberline advance in western United States. Nature. 213(5080): 980-982. [1394]
  • 62. Lloyd, Andrea H. 1997. Response of tree-line populations of foxtail pine (Pinus balfouriana) to climate variation over the last 1000 years. Canadian Journal of Forest Research. 27(6): 936-942. [47903]
  • 63. Lloyd, Andrea H.; Graumlich, Lisa J. 1997. Holocene dynamics of treeline forests in the Sierra Nevada. Ecology. 78(4): 1199-1210. [27723]
  • 64. Lloyd, Andrea. 1998. Growth of foxtail pine seedlings at treeline in the southeastern Sierra Nevada, California, U.S.A. Ecoscience. 5(2): 250-257. [48110]
  • 68. Mastroguiseppe, Ronald J. 1972. Geographic variation in foxtail pine, Pinus balfouriana Grev. & Balf. Humbolt, CA: California State University, Humboldt. 98 p. Thesis. [1548]
  • 69. Mathiasen, Robert L.; Daugherty, Carolyn M. 2001. Susceptibility of foxtail pine and western white pine to limber pine dwarf mistletoe in northern California. Western Journal of Applied Forestry. 16(2): 58-60. [38729]
  • 71. McDonald, Geral I.; Hoff, Raymond J. 2001. Blister rust: an introduced plague. In: Tomback, Diana F.; Arno, Stephen F.; Keane, Robert E., eds. Whitebark pine communities: Ecology and restoration. Washington, DC: Island Press: 193-220. [36703]
  • 72. Miller, Douglas R. 1969. Lophodermium durilabrum found on foxtail pine in California. Plant Disease Reporter. 53(4): 271. [47897]
  • 73. Miller, Douglas R.; Bynum, H. H. 1965. Dwarfmistletoe found on foxtail pine in California. Plant Disease Reporter. 49(8): 647-648. [47898]
  • 77. Oline, David K.; Mitton, Jeffry B.; Grant, Michael C. 2000. Population and subspecific genetic differentiation in the foxtail pine (Pinus balfouriana). Evolution. 54(5): 1813-1819. [48101]
  • 82. Ryerson, A. Diane. 1983. Population structure of Pinus balfouriana Grev. & Balf. along the margins of its distribution area in the Sierran and Klamath regions of California. Sacramento, CA: California State University. 197 p. Thesis. [48204]
  • 84. Samman, Safiya; Schwandt, John W.; Wilson, Jill L. 2003. Managing for healthy white pine ecosystems in the United States to reduce the impacts of white pine blister rust. Report R1-03-118. Missoula, MT: U.S. Department of Agriculture, Forest Service. 10 p. [47202]
  • 88. Scuderi, Louis A. 1987. Glacier variations in the Sierra Nevada, California, as related to a 1200-year tree-ring chronology. Quaternary Research. 27(3): 220-231. [48112]
  • 89. Scuderi, Louis A. 1993. A 2000-year tree ring record of annual temperatures in the Sierra Nevada Mountains. Science. 259(5100): 1433-1436. [47934]
  • 93. Stephen, B. R. 1985. Resistance of five-needle pines to blister rust. Allgemeine Forstzeitschrift. 28: 695-697. [30158]
  • 97. Tomback, Diana F.; Kendall, Katherine C. 2001. Biodiversity losses: the downward spiral. In: Tomback, Diana F.; Arno, Stephen F.; Keane, Robert E., eds. Whitebark pine communities: Ecology and restoration. Washington, DC: Island Press: 243-262. [36705]
  • 106. Vankat, John Lyman. 1970. Vegetation change in Sequoia National Park, California. Davis, CA: University of California. 197 p. Dissertation. [43459]
  • 107. Wood, Stephen A. 1963. A revision of the bark beetle genus Dendroctonus Erichson (Coleptera: Scolytidae). The Great Basin Naturalist. 23(1-2): 1-117. [48203]
  • 35. Gitzendanner, Matthew A.; White, Eleanor E.; Foord, Bret M.; [and others]. 1996. Genetics of Cronartium ribicola. III. Mating system. Canadian Journal of Botany. 74(22): 1952-1859. [28084]
  • 96. Tomback, Diana F. 2003. Whitebark pine: status, trends, strategies in the U.S.A. In: Parks Canada whitebark and limber pine workshop: Workshop proceedings; 2003 February 18-19; Calgary, AB. Ottawa: Parks Canada: 5-7. Available: http://www.whitebarkfound.org/PDF_files/WBPProceedings.pdf [2004, June 3]. [47867]
  • 99. U.S. Department of Agriculture, Forest Service, Pacific Southwest Region, Forest Pest Management. 1995. Forest pest conditions in California --1995. [Online]. In: Forest health protection. California Forest Pest Council (Producer). Available: http://www.fs.fed.us/r5/spf/publications1995.htm [2004, June 24]. [48213]

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Relevance to Humans and Ecosystems

Benefits

Importance to Livestock and Wildlife

More info for the term: cover

Foxtail pine boughs probably provide shelter for wildlife. Chipmunks and birds probably eat the seeds, but little is known about wildlife use of foxtail pine habitats. Research is needed on the ecology of foxtail pine communities.

Palatability/nutritional value: Foxtail pine seeds are palatable and nutritious, but they are not large compared to most 5-needle pines [98]:

Species Mean seed weight
Great Basin bristlecone pine 25 mg
foxtail pine 27 mg
limber pine 93 mg
whitebark pine 175 mg

Cover value: No information is available on this topic.

  • 98. Tomback, Diana F.; Linhart, Yan B. 1990. The evolution of bird-dispersed pines. Evolutionary Ecology. 4: 185-219. [17534]

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Other uses and values

Foxtail pine is an ecologically important species. The open structure of foxtail pine stands slows snowmelt and helps retain snowpack. Foxtail pine also helps stabilize soil on steep subalpine slopes [84,92].

As a long-lived conifer, foxtail pine is a valuable species for dendrochronological and related climate studies [33,36,37,88,89].

Wood Products: Foxtail pine is rarely harvested [82] and is not commercially important [92].

  • 33. Garfin, Gregg M. 1998. Relationships between winter atmospheric circulation patterns and extreme tree growth anomalies in the Sierra Nevada. International Journal of Climatology. 18(7): 725-740. [40071]
  • 36. Graumlich, Lisa J. 1991. Subalpine tree growth, climate, and increasing CO2: an assessment of recent growth trends. Ecology. 72(1): 1-11. [47892]
  • 37. Graumlich, Lisa J. 1993. A 1000-year record of temperature and precipitation in the Sierra Nevada. Quaternary Research. 39(2): 249-255. [29155]
  • 82. Ryerson, A. Diane. 1983. Population structure of Pinus balfouriana Grev. & Balf. along the margins of its distribution area in the Sierran and Klamath regions of California. Sacramento, CA: California State University. 197 p. Thesis. [48204]
  • 84. Samman, Safiya; Schwandt, John W.; Wilson, Jill L. 2003. Managing for healthy white pine ecosystems in the United States to reduce the impacts of white pine blister rust. Report R1-03-118. Missoula, MT: U.S. Department of Agriculture, Forest Service. 10 p. [47202]
  • 88. Scuderi, Louis A. 1987. Glacier variations in the Sierra Nevada, California, as related to a 1200-year tree-ring chronology. Quaternary Research. 27(3): 220-231. [48112]
  • 89. Scuderi, Louis A. 1993. A 2000-year tree ring record of annual temperatures in the Sierra Nevada Mountains. Science. 259(5100): 1433-1436. [47934]
  • 92. Steinhoff, R. J. 1972. White pines of western North America and Central America. In: Bingham, Richard: Hoff, Raymond J., tech. coords. In: Biology of rust resistance in forest trees: Proceedings of a NATO/IUFRO Advanced Study Institute; 1969 August 17-24; Washington, DC. Misc. Pub. 1221. U.S. Department of Agriculture, Forest Service: 215-232. [30287]

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Wikipedia

Foxtail pine

Pinus balfouriana (foxtail pine) is a rare pine that is endemic to California, United States. The two disjunct populations are found in the Klamath Mountains (subspecies balfouriana) and the southern Sierra Nevada[2] (subspecies austrina). A small outlying population was reported in southern Oregon, but was proven to have been misidentified.[3]

Description[edit]

P. balfouriana is a tree to 10–20 m (33–66 ft) tall, exceptionally 35 m (115 ft), and up to 2 m (7 ft) in trunk diameter. Its leaves are needle-like, in bundles of five (or sometimes four, in the southern Sierra) with a semi-persistent basal sheath, and 2–4 cm (0.8–1.6 in) long, deep glossy green on the outer face, and white on the inner faces; they persist for 10–15 years. The cones are 6–11 cm (2 38 - 4 516 in) long, dark purple ripening red-brown, with soft, flexible scales each with a one millimetre central prickle.

A Foxtail Pine in the southern Sierra Nevada.

Distribution[edit]

P. balfouriana occurs in the subalpine forest at an elevation of 1,950–2,750 m (6,400–9,020 ft) in the Klamath Mountains, and at 2,300–3,500 m (7,500–11,500 ft) in the Sierra Nevada. In the Sierra Nevada, Foxtail pines are limited to the area around Sequoia and Kings Canyon National Parks. In both areas, it is often a tree line species.

Age[edit]

It is thought that P. balfouriana can live up to 3000 years in the Sierra Nevada, although the highest currently proven age is 2110 years. In the Klamath Mountains, ages are only known to about 1000 years.

Related species[edit]

P. balfouriana is closely related to the bristlecone pines, being classified in the same subsection Balfourianae; it has been hybridised with the Great Basin Bristlecone Pine in cultivation, though no hybrids have ever been found in the wild.

References[edit]

  1. ^ Farjon, A. (2011). "Pinus balfouriana". IUCN Red List of Threatened Species. Version 3.1. International Union for Conservation of Nature. Retrieved 2013-11-10. 
  2. ^ 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. 83. ISBN 1-4027-3875-7. 
  3. ^ Kauffmann, Michael E. (2012). Conifer Country. Kneeland, CA: Backcountry Press. ISBN 9780578094168.OCLC 798852130.

Further reading[edit]

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Notes

Comments

Pinus balfouriana is the true "foxtail pine." In leaf character it is hardly, if at all, distinguishable from P . longaeva , but its strongly conic-based cones with distinctly shorter-prickled, sunken-centered umbos at once distinguish it from that species. 

 Plants shown to be genetically distinct from the type (differences in chemistry, form, foliage, cone orientation, and seeds) have been called Pinus balfouriana subsp. austrina R.Mastrogiuseppe & J.Mastrogiuseppe. As in several other species or species complexes in Pinus , however, there is a problem with a character gradient involving related taxa. The evidence presented by D.K. Bailey (1970) and later by R.J. Mastrogiuseppe and J.D. Mastrogiuseppe (1980) could as well be used to indicate that P . balfouriana (with its two infraspecific taxa) and P . longaeva represent a single species of three subspecies or three varieties. The more conservative view of Bailey is followed here.

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© Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO, 63110 USA

Source: Missouri Botanical Garden

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Names and Taxonomy

Taxonomy

More info for the term: natural

The scientific name of foxtail pine is Pinus balfouriana Grev. & Balf.
(Pinaceae). There are 2 subspecies [8,32,39,47,68]:

Pinus balfouriana subsp. austrina R.J. & J.D. Mastrogiuseppe 
southern foxtail pine

Pinus balfouriana subsp. balfouriana  northern foxtail pine
Foxtail pine, Great Basin bristlecone pine (P. longaeva), and Rocky Mountain bristlecone pine (P. aristata)
share a common ancestor [83,109]. Taxa within the foxtail-bristlecone pine
complex (Pinus, subgenus Strobus, section
Parrya Mayr, subsection Balfourianae Englm.) are distinguished by growth
form, bark, and differences in chemical composition
[8,25,68,76]; however,
these characters intergrade [32,39,68].
Foxtail and bristlecone pines readily produce fertile hybrids in the laboratory [93,109].
Disjunct distributions, and possibly other factors, prevent natural hybridization among the 4 taxa. Southern
foxtail and Great Basin bristlecone pine populations seem geographically close enough for
limited pollen dispersal
(see General Distribution); yet to
date (2004), southern foxtail ×
Great Basin bristlecone pine hybrids have not been found in the field
[8,60].
  • 39. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. [21992]
  • 8. Bailey, D. K. 1970. Phytogeography and taxonomy of Pinus subsection Balfourianae. Annals of the Missouri Botanical Garden. 57: 210-249. [375]
  • 25. Critchfield, William B. 1977. Hybridization of foxtail and bristlecone pines. Madrono. 24(4): 193-244. [713]
  • 47. Keane, Robert E.; Arno, Stephen F. 2001. Restoration concepts and techniques. In: Tomback, Diana F.; Arno, Stephen F.; Keane, Robert E., eds. Whitebark pine communities: Ecology and restoration. Washington, DC: Island Press: 367-400. [36711]
  • 60. Lanner, Ronald M.; Hutchins, Harry E.; Lanner, Harriette A. 1984. Bristlecone pine and Clark's nutcracker: probable interaction in the White Mountains, California. Great Basin Naturalist. 44(2): 357-360. [48202]
  • 68. Mastroguiseppe, Ronald J. 1972. Geographic variation in foxtail pine, Pinus balfouriana Grev. & Balf. Humbolt, CA: California State University, Humboldt. 98 p. Thesis. [1548]
  • 76. Munz, Philip A. 1974. A flora of southern California. Berkeley, CA: University of California Press. 1086 p. [4924]
  • 83. Ryerson, Diane. 1984. Krummholz foxtail pines. Fremontia. 11(4): 30. [47919]
  • 93. Stephen, B. R. 1985. Resistance of five-needle pines to blister rust. Allgemeine Forstzeitschrift. 28: 695-697. [30158]
  • 32. Flora of North America Association. 2000. Flora of North America north of Mexico. Volume 2: Pteridophytes and gymnosperms, [Online]. Flora of North America Association (Producer). Available: http://hua.huh.harvard.edu/FNA/ [2004, May 27]. [36990]

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Common Names

foxtail pine

southern foxtail pine

northern foxtail pine

Sierra foxtail pine

Klamath foxtail pine

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