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Introductory

SPECIES: Pinus balfouriana | Foxtail Pine
ABBREVIATION : PINBAL SYNONYMS : NO-ENTRY SCS PLANT CODE : PIBA PIBAA PIBAB COMMON NAMES : foxtail pine TAXONOMY : The currently accepted scientific name of foxtail pine is Pinus balfouriana Grev. & Balf. [8,13,18]. There are two subspecies [13,18]: P. b. ssp. austrina R. J. & J. D. Mastrogiuseppe P. b. ssp. balfouriana Grev. & Balf. Mastroguiseppe [13] published the name P. b. ssp. austrina for the southern Sierra Nevada group. In recognizing the southern Sierra Nevada populations as a distinct subspecies, the northern Klammath Mountains populations became P. b. ssp. balfouriana. In pollination experiments, the two subspecies exhibit almost full crossability. Southern Sierra Nevada Onion Valley female cones pollinated with pollen from the Klammath Mountains (Yolla Bolly) to the north produced 84 percent as much seed as they did with Onion Valley pollen. A crossability of 84 percent is fairly typical for two subspecies [5,13]. These two disjuncts have probably been separated since the Pleistocene. It is interesting that for some characteristics the southern Sierra Nevada subspecies is more similar to the Great Basin bristlecone pine (P. longaeva), but it is intermediate between the northern Klammath Mountains subspecies and Great Basin bristlecone pine for others [12,13]. Essential oils of wood and foliage support the morphological and chemical separation of foxtail and bristlecone pines. However, there may be a natural hybrid between them in the White Mountains of California. Cones from these trees resemble those of the foxtail pine. If this has occurred, then westerly winds may have carried pollen from the southern foxtail populations across the Owens Valley to the bristlecones in the White Mountains. When this cross was conducted in a laboratory, fairly high numbers of hybrid seed were produced [5,13,17]. LIFE FORM : Tree FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY COMPILED BY AND DATE : Kathy Ahlenslager, October 1986 LAST REVISED BY AND DATE : Kathy Ahlenslager, December 1987 AUTHORSHIP AND CITATION : Ahlenslager, Kathleen E. 1986. Pinus balfouriana. In: Remainder of Citation

DISTRIBUTION AND OCCURRENCE

SPECIES: Pinus balfouriana | Foxtail Pine
GENERAL DISTRIBUTION : Foxtail pine is endemic to California and occurs only at high elevations, on upper slopes and exposed ridges. It exists as two disjunct subspecies separated by about 300 air miles (92 km): in the Klammath Mountains of northwestern California and in the southern high Sierra Nevada. The nothern subspecies, balfouriana, occurs as several scattered stands in the Marble, Scott, and Yolla Bolly Mountains, as well as the Trinity Alps [5,8,12,13]. The southern subspecies, austrina, is centered on the upper South Fork of the Kern River drainage mostly in Sequoia and Kings Canyon National Parks, as well as the adjacent Inyo National Forest. On the east side of the Sierra Nevada, this subspecies is only 20 air miles (30 km) from the closely related bristlecone pine of the Inyo Mountains [5,8,12,13]. ECOSYSTEMS : FRES23 Fir - spruce FRES26 Lodgepole pine FRES28 Western hardwoods STATES : CA ADMINISTRATIVE UNITS : KICA SEQU BLM PHYSIOGRAPHIC REGIONS : 4 Sierra Mountains KUCHLER PLANT ASSOCIATIONS : K008 Lodgepole pine - subalpine forest SAF COVER TYPES : 205 Mountain hemlock 207 Red fir 256 California mixed subalpine SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Foxtail pine is not found as a dominant, codominant, or indicator species in any habitat type. In the Klammath Mountains, ssp. balfouriana occurs on ridges and upper southern and western slopes over 7,200 feet (2,200 m). Here it is associated with whitebark pine (Pinus albicaulis), mountain hemlock (Tsuga mertensiana), Shasta red fir (Abies magnifica var. shastensis), lodgepole pine (P. contorta ssp. murrayana), and Jeffrey pine (P. jeffreyi) [2,3,12,13]. In the southern Sierra Nevada, ssp. austrina is characteristic of treeline where it forms nearly pure forests. Although individual trees are widely spaced with little understory vegetation, there are occasional associations of lodgepole pine, whitebark pine, and limber pine (P. flexilis) [2,3,12,13].

VALUE AND USE

SPECIES: Pinus balfouriana | Foxtail Pine
WOOD PRODUCTS VALUE : NO-ENTRY IMPORTANCE TO LIVESTOCK AND WILDLIFE : The seeds of foxtail pine are edible to some wildlife. Trees often occur in clumps, probably from the caches of seed-eaters, such as Clark's nutcrackers. Small animals and birds may benefit from these seed caches [9]. PALATABILITY : NO-ENTRY NUTRITIONAL VALUE : NO-ENTRY COVER VALUE : Foxtail pine provides some cover for bird and small animal species [9]. VALUE FOR REHABILITATION OF DISTURBED SITES : Foxtail pine has little to no value for rehabilitation due to the Eextremely slow growth rate of seedlings and mature trees [1,5]. OTHER USES AND VALUES : Trees of this species are useful as habitat, food for wildlife, and for watershed protection. Foxtail pine is an interesting species due to its disjunct distribution and the size it reaches in harsh environments [13]. MANAGEMENT CONSIDERATIONS :

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Pinus balfouriana | Foxtail Pine
GENERAL BOTANICAL CHARACTERISTICS : Even at upper treeline, the typical growth habit of mature foxtail pines is erect and straight with a largely intact cambium, full crown, and retained leader. The sparse crowns of contorted trees are often only supported by a narrow strip of cambium, the living tissue of a tree. The effects of sand and ice-blasting by wind are seen in older trees with double "pick-a-back" growth form. The windward trunk is dead, but a narrow strip of cambium supports branches on the leeward side of the tree, the "piggy back" [12]. Trees of the northern foxtail pine, ssp. balfouriana grow erect or with an uphill lean. Individuals are commonly 30 to 60 feet (100-200 m) high. The tallest is recorded at 98 feet (32 m). One large relict of the southern foxtail pine, ssp. austrina, growing at 10,540 feet (3,200 m) on Alta Peak, Tulare County, California, is 6.7 feet (2 m) in d.b.h. and 78.5 feet (24 m) in height [3]. Trees retain needles 10 to 15 years. Although the southern Sierra Nevada foxtail pine usually grows erect, it is more often twisted. Trees retain needles 20 to 30 years [12]. Foxtail pines have deep and spreading root systems which probably occupy the openings between trees [2]. Since they grow as widely spaced individuals, the density of associated plants is low, with an absence of associated plant competition [13]. Foxtail pines and bristlecone pines (Pinus longaeva and P. aristata) share numerous interesting characteristics [13]: (1) occurrence at the upper limits of tree growth and a reduced rate of growth (2) a capacity for long life (3) continued growth after loss of large areas of tissue from drought, stress, wind injury, and soil erosion (4) heavy, dense, resin-impregnated wood with small, closely arranged water-conducting cells (tracheids) which provide resistance to moisture and decay (5) survival in an environment of drought, low temperatures, short growing season, daily temperature extremes, and poor soils (6) retention of needles for several years, which reduces their need for moisture and nutrients, as well as helps to carry a tree over several years of stress (7) relative safety from ground fire due to sparse ground cover and litter scarcity The longevity of foxtail pine is of interest because of the long life spans reported for its close relatives the bristlecone pines. In general, trees of northern foxtail pine do not live as long as those of southern foxtail pine. The greatest known age of the northern foxtail pine is 364 years. In comparison, the greatest known age of a southern foxtail pine is 682 years. Northern foxtail pine, growing in the moist Klammath Mountains, has a greater frequency of heartwood decay than southern foxtail, which grows in the drier Sierra Nevada [13]. It is often difficult to determine the exact age and growth rates of these trees because of asymmetric growth and heartwood decay. Tree ages can be estimated with increment bores. Based on ring counts and growth rates, the maximum attainable age for northern foxtail pine is estimated to be 1,300 to 1,500 years, while that of southern foxtail pine is 2,500 to 3,000 years [12,13]. RAUNKIAER LIFE FORM : Phanerophyte REGENERATION PROCESSES : The environmental and physiological factors contributing to the longevity of foxtail pines also work to lower their regeneration potential. Seedlings are slow growing [3]. With the caching of foxtail pine seeds by seed-eaters, seedlings often occur in dense clusters. The effects of this caching in the regeneration of foxtail pine appears beneficial, since there is a high frequency of clumped trees [6]. SITE CHARACTERISTICS : Subspecies balfouriana occurs as several scattered stands in the Marble, Scott, and Yolla Bolly mountains, as well as the Trinity Alps from 6,700 to 9,000 feet (2,000-2,400 m). Occasional individuals grow in mountain valleys as low as 6,400 feet (1,950 m). This Klammath Mountain subspecies is found on ridges, as well as upper southern and western slopes where trees form open stands at elevations over 7,200 feet (2,200 m). Northern foxtail pine occurs on several parent material types but is especially common on ultrabasic serpentine, schist, and granodiorite [2,5,12,13]. Subspecies austrina ranges from 8,500 to 12,000 feet (2,600-3,650 m) in elevation on the upper South Fork of the Kern River drainage and is characteristic of treeline. Occasional individuals are found as low as 7,700 feet (2,350 m). Trees grow in open stands on dry sunny ridges. The most commobn parent material is decomposed granite, which gives rise to shallow, coarse-textured soils, usually with boulders [2,5,12,13]. At its lower elevational limits, foxtail pine grows in pure stands or mixed with whitebark pine, lodgepole pine, Jeffrey pine, and western white pine (Pinus monticola). At higher elevations trees usually grow as widely spaced individuals in pure stands. Plants are intolerant of shade at all stages of growth. Ground cover under these trees is characterized by bare earth and rock with a few scattered herbs [3]. The growing conditions of ssp. balifouriana in the Klammath Mountains are less extreme than those of subsp. austrina in the southern Sierra Nevada. With average annual precipitation in the Klammath Mountains ranging from 50 to 60 inches (125-175 cm), subspecies balfouriana is found in the wettest habitat of any of the bristlecone or foxtail pines. In contrast, annual precipitation in the southern Sierra Nevada averages 20 to 30 inches (50-75 cm). Temperature extremes are great and winds severe. In both areas most of the precipitation occurs as winter snow. Summers are dry with low relative humidity [12,13]. SUCCESSIONAL STATUS : Foxtail pine is a climax species and persists at treeline for hundreds to thousands of years in the absence of disturbance and competition. It is highly adapted to its habitat of very shallow soils, slow primary succession, short growing season, and avalanches [2,13]. SEASONAL DEVELOPMENT : Foxtail pines flower from July to August. The period of flowering and coneopening is uniform for foxtail pine, and the two bristlecone pines. Ssp. balifouriana flowers somewhat later than than ssp. austrina [5,9]. Cones ripen from September to October with seed dispersal at this time. The minimum seed-bearing age for trees is 20 years with 5 to 6 years between large seed crops [9].

FIRE ECOLOGY

SPECIES: Pinus balfouriana | Foxtail Pine
FIRE ECOLOGY OR ADAPTATIONS : Foxtail pine occurs for the most part in the subalpine zone in habitats where fuels to carry fires are nonexistant. In this area low temperatures and a short growing season keep the production of organic matter low. Surface fires are infrequent, slow-burning, and of low intensity [15]. POSTFIRE REGENERATION STRATEGY : Tree without adventitious-bud root crown Initial-offsite colonizer (off-site, initial community)

FIRE EFFECTS

SPECIES: Pinus balfouriana | Foxtail Pine
IMMEDIATE FIRE EFFECT ON PLANT : Low intensity, infrequent fires do not have much effect on foxtail pines. Although the density of associated shrubs and herbs might be enough to carry periodic fires, most individuals grow in habitats where fuels are nonexistant. Even though fire injury provides an entrance channel for decay organisms, trees several hundred years of age have survived fire [13]. The charred remains of trees struck by lightning in the southern Sierra Nevada are evidence that periodic fires do occur, although they seldom spread over large treeline areas. This is due to the wide spacing of trees, 30 to 60 feet (10-20 m) apart and the lack of fuels from a forest floor composed primarily of mineral earth and rocks [1]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : NO-ENTRY DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : NO-ENTRY FIRE MANAGEMENT CONSIDERATIONS : NO-ENTRY

REFERENCES

SPECIES: Pinus balfouriana | Foxtail Pine
REFERENCES : 1. Arno, Stephen F. 1980. Forest fire history in the northern Rockies. Journal of Forestry. 78(8): 460-465. [11990] 2. Arno, Stephen F.; Hammerly, Ramona P. 1984. Timberline: Mountain and arctic forest frontiers. Seattle, WA: The Mountaineers. 304 p. [339] 3. Barbour, Michael G.; Major, Jack, eds. 1977. Terrestrial vegetation of California. New York: John Wiley & Sons. 1002 p. [388] 4. 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] 5. Critchfield, William B. 1977. Hybridization of foxtail and bristlecone pines. Madrono. 24(4): 193-244. [713] 6. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905] 7. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; [and others]. 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] 8. Griffin, James R.; Critchfield, William B. 1972. The distribution of forest trees in California. Res. Pap. PSW-82. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station. 118 p. [1041] 9. Krugman, Stanley L.; Jenkinson, James L. 1974. Pinaceae--pine family. In: Schopmeyer, C. S., technical coordinator. Seeds of woody plants in the United States. Agric. Handb. 450. Washington, DC: U.S. Department of Agriculture, Forest Service: 598-637. [1380] 10. Kuchler, A. W. 1964. Manual to accompany the map of potential vegetation of the conterminous United States. Special Publication No. 36. New York: American Geographical Society. 77 p. [1384] 11. Lyon, L. Jack; Stickney, Peter F. 1976. Early vegetal succession following large northern Rocky Mountain wildfires. In: Proceedings, Tall Timbers fire ecology conference and Intermountain Fire Research Council fire and land management symposium; 1974 October 8-10; Missoula, MT. No. 14. Tallahassee, FL: Tall Timbers Research Station: 355-373. [1496] 12. Mastroguiseppe, R. J.; Mastroguiseppe, J. D. 1980. A study of Pinus balfouriana Grev. & Balf. (Pinaceae). Systematic Botany. 5(1): 86-104. [1546] 13. Mastroguiseppe, Ronald J. 1972. Geographic variation in foxtail pine, Pinus balifouriana Grev. & Balf. Humbolt, CA: California State University, Humboldt. 98 p. M.S. thesis. [1548] 14. Mirov, N. T. 1967. The genus Pinus. New York: Ronald Press. 602 p. [1663] 15. Parsons, David J. 1981. The role of fire management in maintaining natural ecosystems. 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: 469-488. [5083] 16. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843] 17. Zavarin, Eugene; Snajberk, Karel; Bailey, Dana. 1976. Variability in the essentil oils of wood and foliage of Pinus aristata and Pinus longaeva. Biochemical Systematics and Ecology. 4: 81-92. [2690] 18. Kartesz, John T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. Volume II--thesaurus. 2nd ed. Portland, OR: Timber Press. 816 p. [23878] 19. 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; RWU 4403 files. 7 p. [20090] 20. U.S. Department of Agriculture, Soil Conservation Service. 1994. Plants of the U.S.--alphabetical listing. Washington, DC: U.S. Department of Agriculture, Soil Conservation Service. 954 p. [23104] 21. U.S. Department of the Interior, National Biological Survey. [n.d.]. NP Flora [Data base]. Davis, CA: U.S. Department of the Interior, National Biological Survey. [23119]

Index

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