Index of Species Information

SPECIES:  Abies lasiocarpa

Introductory

SPECIES: Abies lasiocarpa
AUTHORSHIP AND CITATION : Uchytil, Ronald J. 1991. Abies lasiocarpa. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].
ABBREVIATION : ABILAS ABILASA ABILASL SYNONYMS : Abies balsamea ssp. lasiocarpa (Hook.) Boivin Abies balsamea var. fallax (Engelm.) Boivin SCS PLANT CODE : ABLA ABLAA COMMON NAMES : subalpine fir alpine fir western balsam fir balsam white balsam balsam fir white fir Rocky Mountain fir pino real blanco cork-bark fir corkbark fir Arizona fir TAXONOMY : The genus Abies consists of about 40 species of evergreen trees found in the northern hemisphere. Nine species of Abies, including subalpine fir, are native to the United States [75]. The currently accepted scientific name of subalpine fir is Abies lasiocarpa (Hook.) Nutt. [75]. Subalpine fir is widely distributed and exhibits geographic variation. Two varieties are recognized based on morphological differences [75]: Abies lasiocarpa var. arizonica - corkbark fir Abies lasiocarpa var. lasiocarpa - subalpine fir Subalpine fir hybridizes with balsam fir (A. balsamea) where their ranges overlap in the Canadian Rockies [41]. LIFE FORM : Tree FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY

DISTRIBUTION AND OCCURRENCE

SPECIES: Abies lasiocarpa
GENERAL DISTRIBUTION : Subalpine fir is the mostly widely distributed fir in North America, spanning more than 32 degrees of latitude [11].  It occurs chiefly in mountainous areas from the Yukon interior near treeline and along the coast of southeastern Alaska south through western Alberta and British Columbia to southern Colorado and scattered mountain ranges of Arizona and New Mexico [54,75].  In the western portion of its range, subalpine fir does not occur along the western slope of the Coast Range in southern British Columbia or along the Coast Ranges of Washington and Oregon but does occur on Vancouver Island and in the Olympic Mountains of Washington [11].  It occurs on both slopes of the Cascade Mountains as far south as southern Oregon [11].  The two varieties are distributed as follows [11,75]: var. lasiocarpa - almost the same as the species, but not in central and southeastern Arizona.  var. arizonica - from central Colorado to southwestern New Mexico, and in southeastern and central Arizona.  Subalpine fir and corkbark fir occur together in scattered mountain ranges in southwestern Colorado, northern, western, and southwestern New Mexico, and in the high mountains of Arizona [11]. ECOSYSTEMS :    FRES20  Douglas-fir    FRES22  Western white pine    FRES23  Fir - spruce    FRES24  Hemlock - Sitka spruce    FRES25  Larch    FRES26  Lodgepole pine    FRES44  Alpine STATES :      AK  AZ  CO  ID  MT  NV  NM  OR  UT  WA      WY  AB  BC  YT BLM PHYSIOGRAPHIC REGIONS :     1  Northern Pacific Border     2  Cascade Mountains     5  Columbia Plateau     6  Upper Basin and Range     8  Northern Rocky Mountains     9  Middle Rocky Mountains    11  Southern Rocky Mountains    12  Colorado Plateau KUCHLER PLANT ASSOCIATIONS :    K004  Fir - hemlock forest    K012  Douglas-fir forest    K015  Western spruce - pine forest    K018  Pine - Douglas-fir forest    K020  Spruce - fir - Douglas-fir forest    K021  Southwestern spruce - fir forest    K052  Alpine meadows and barren SAF COVER TYPES :    201  White spruce    202  White spruce - paper birch    205  Mountain hemlock    206  Engelmann spruce - subalpine fir    208  Whitebark pine    209  Bristlecone pine    210  Interior Douglas-fir    212  Western larch    213  Grand fir    215  Western white pine    216  Blue spruce    217  Aspen    218  Lodgepole pine    219  Limber pine    223  Sitka spruce    224  Western hemlock    226  Coastal true fir - hemlock    253  Black spruce - white spruce SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Forests in which subalpine fir attains climax dominance or codominance are widespread throughout the mountains of western North America.  The subalpine fir series generally occupies cold, high elevation mountain forests.  Engelmann spruce (Picea engelmannii) is usually associated with subalpine fir.  It occurs as either a climax codominant or as a persistent, long-lived seral species in most subalpine fir habitat types. Published classification schemes listing subalpine fir as a dominant part of the vegetation in habitat types (hts), community types (cts), plant associations (pas), ecosystem associations (eas), site types (sts) or dominance types (dts) are presented below: Area                    Classification          Authority AK: -----               general veg. cts        Viereck & Dyrness 1980 AZ: San Francisco       forest, alpine &       Peaks RNA           meadow cts            Rominger & Paulik 1983 AZ, NM: -----           forest & woodland hts   Layser & Schubert 1979         Apache, Gila,           Cibola NFs    forest hts              Fitzhugh & others 1987         s of Mogollon           Rim           forest hts              Develice & Ludwig 1983b n AZ, n NM              forest hts              Larson & Moir 1987 CO: Arapaho &       Roosevelt NFs     forest hts              Hess & Alexander 1986     Gunnison &       Uncompahgre NFs   forest hts              Komarkova & others 1988     Routt NF            forest hts              Hoffman & Alexander 1980     White River-        grassland, shrubland,       Arapaho NF          & forestland hts      Hess & Wasser 1982     White River NF      forest hts              Hoffman & Alexander 1983 w CO                    riparian pas            Baker 1989a ID: Sawtooth, White       Cloud, Boulder,       & Pioneer Mtns    general veg. cts        Schlatterer 1972 c ID                    forest hts              Steele & others 1981 n ID                    forest hts              Cooper & others 1987 se ID                   aspen cts               Mueggler & Campbell 1986 e ID, w WY              forest hts              Steele & others 1983 MT: -----               forest hts              Pfister & others 1977     -----               riparian dts            Hansen & others 1988 c, e MT                 riparian cts, hts       Hansen & others 1990 nw MT                   riparian hts, cts       Boggs & others 1990 sw MT                   riparian rst, cts, hts  Hansen & others 1989 NM: Cibola NF           forest hts              Alexander & others 1987     Lincoln NF          forest hts              Alexander & others 1984 n NM, s CO              forest hts              Develice & Ludwig 1983a n NM, s CO              forest hts              Develice  & others 1986 OR: Wallowa-Whitman NF  steppe & forest pas     Johnson & Simon 1987     Eagle Cap       Wilderness        general veg. cts        Cole 1982 OR, WA: -----           general veg. cts        Franklin & Dyrness 1973         Blue Mtns       general veg. pas        Hall 1973 UT: -----               aspen cts               Mueggler & Campbell 1986 c, s UT                 forest hts              Youngblood & Mauk 1985 n UT                    forest hts              Mauk & Henderson 1984 WA: Okanogan NF         forest pas              Williams & Lillybridge 1983     Mount Rainier NP    forest pas              Franklin & others 1988        North Cascades NP   forest pas              Agee & Kertis 1987 e WA, n ID              forest hts, cts         Daubenmire & Daubenmire 1968 WY: Bridger-Teton NF    aspen cts               Youngblood & Mueggler 1981     Medicine NF         forest hts              Alexander & others 1986     Bighorn Mtns        forest hts              Hoffman & Alexander 1976     Wind River Mtns     forest hts              Reed 1976 USFS R-2                general veg. pas        Johnston 1987 USFS R-2                general veg. hts,pas    Wasser & Hess 1982 USFS R-4                aspen cts               Mueggler 1988 w-c AB                  forest cts              Corns 1983 BC: -----               grassland, forest hts   McLean 1970     -----               general veg. eas        Pojar & others 1984 nw BC                   forest eas              Haeussler & others 1985           

MANAGEMENT CONSIDERATIONS

SPECIES: Abies lasiocarpa
WOOD PRODUCTS VALUE : Subalpine fir wood is odorless, light-weight, soft, and low in bending and compressive strength [11].  It is easy to work, glues well, and holds nails and screws fairly well.  The wood is primarily used for products such as lumber for home construction and for prefabricated wood products [9].  Subalpine fir has excellent pulping properties [9].  Use for poles and pilings requires large amounts of preservatives because the wood decays rapidly [108]. IMPORTANCE TO LIVESTOCK AND WILDLIFE : Subalpine-fir-dominated stands generally do not produce enough forage for livestock but do provide browse and cover for large and small wildlife species.  Mule deer, elk, moose, woodland caribou, black bear, and grizzly bear often use subalpine fir habitats as summer range [10,23,113]].  Subalpine fir forests are generally not suitable winter range for deer and elk because of heavy snowpack, but some lower elevation subalpine fir habitat types are used by moose and woodland caribou during the winter [23,61,89].  Subalpine fir forests support numerous species of small mammals and birds.  The snowshoe hare, flying squirrel, red squirrel, porcupine, pine marten, fisher, lynx, and several species of mice, voles, chipmunks, and shrews all inhabit subalpine fir forests [26,104,113].  Numerous species of birds nest and feed in subalpine fir forests, including several woodpeckers, flycatchers, kinglets, nuthatches, juncos, thrushes, chickadees, crossbills, the pine siskin, owls, and grouse [104,113]. The young growth of subalpine fir is sometimes eaten by mule deer, elk, bighorn sheep, and snowshoe hares, but it is not an important food item. Subalpine fir comprises only a small portion of the summer diet of mountain goats but can be a major food source in the winter and spring [99].  Throughout much of Montana, Idaho, and Wyoming, subalpine fir is an important winter food of moose [89].  On moose winter range near Jackson Hole, Wyoming, an average of 13 to 18 percent of small subalpine fir trees were browsed by moose, and 44 to 78 percent of the branches on trees browsed were utilized [61].  In Yellowstone National Park, grizzly bears sometimes strip the bark of subalpine fir to feed on the underlying cambium [19].  The winter diet of blue grouse consists primarily of conifer needles.  These grouse often winter in subalpine stands and may feed heavily on the needles and buds of subalpine fir [103,113]. Subalpine fir seeds are eaten by several species of small mammals and birds.  Red squirrels eat seeds from cached subalpine fir cones [71]. Fir seeds are also eaten by chipmunks and mice.  Several birds, including chickadees, nuthatches, crossbills, the pine siskin, and the Clark's nutcracker remove and eat the seeds from fir cones [49,77]. Because subalpine fir seeds are large, comprising about 26 percent of a cone's weight, they are an energy-efficient food source for small birds [49].  Small birds may make considerable use of fir seeds, but their foraging is scattered and sporadic throughout subalpine forests [49]. PALATABILITY : The palatability of subalpine fir to domestic livestock is low [30]. Its palatability to big game animals is generally low also, but in some locations it is highly palatable to moose and mountain goats during winter and spring [89,99].  The seeds are palatable to numerous small mammal species [77].  Red squirrels generally eat subalpine fir seeds after other cached conifer seeds have been consumed [71].  The needles are highly palatable to blue grouse. The relish and degree of use shown by livestock and wildlife species for subalpine fir in several western states is rated as follows [30,61,89,96]:                        CO    MT    WY    UT    ID Cattle                poor  poor  poor  poor  ---- Sheep                 poor  poor  poor  poor  ---- Horses                poor  poor  poor  poor  ---- Pronghorn             ----  ----  poor  poor  ---- Elk                   ----  poor  poor  poor  ---- Moose                 ----  good  good  ----  fair Mule deer             ----  poor  poor  poor  ---- White-tailed deer     ----  ----  poor  ----  ---- Small mammals         ----  fair  fair  good  ---- Small nongame birds   ----  ----  fair  good  ---- Upland game birds     ----  fair  fair  good  ---- Waterfowl             ----  ----  poor  poor  ---- NUTRITIONAL VALUE : Subalpine fir is low in protein value but fair in energy value [30]. Percent composition of subalpine fir browse collected near Jackson Hole, Wyoming, was as follows [61]:   date              crude        ether       crude      nitrogen collected          protein      extract      fiber    free extract  11/25              5.57         7.53        20.19      50.26 A study in Montana found the following concentration of elements in subalpine fir needles and twigs [110]:       1-yr-old green needles    twigs < 0.25 inch (0.64 cm) in diameter      (micrograms/gram [mean])           (micrograms/gram [mean]) Ca             9722                               5840 Cu                7.4                                7.9 Fe               64                                182 K              5553                               7031 Mg              819                               1038 Mn             1020                                587 N             10690                               4962 Na              103                                124 P              1450                               2254 Zn               43                                  5           (percent [mean])                    (percent [mean]) Ash               3.5                                3.5 COVER VALUE : Big game:  Subalpine fir habitat types provide excellent hiding cover for deer, elk, mountain goats, moose, and bear [10,113].  Certain low elevation subalpine fir forests may be used by elk during calving, and high elevation subalpine fir forests by bighorn sheep during lambing and lamb rearing [113].  Dense stands provide cool summertime shade for big game animals [71].  In Yellowstone National Park, grizzly bear daybeds are often found in subalpine fir stands [19]. Small mammals and birds:  Small subalpine firs provide good year-round hiding cover.  Dense thickets of small trees are often nearly impenetrable and provide hiding places for small mammals such as snowshoe hares and porcupines [13,71].  Blue grouse often overwinter in subalpine trees and rely almost exclusively on them for escape cover [103].  Subalpine fir snags are used by numerous cavity-nesting birds, but are generally less preferred than those of associated conifers [105]. The degree to which subalpine fir provides environmental protection during one or more seasons for wildlife species is as follows [30]:                        CO    MT    WY    UT Pronghorn             ----  ----  poor  poor Elk                   good  fair  good  good Mule deer             good  fair  good  good White-tailed deer     ----  ----  poor  ---- Small mammals         good  good  good  good Small nongame birds   good  fair  good  good Upland game birds     good  good  good  fair Waterfowl             ----  ----  poor  poor VALUE FOR REHABILITATION OF DISTURBED SITES : Subalpine fir can be planted on disturbed sites within forest vegetation types where it naturally occurs [92].  It is generally recommended for cool and moist sites within subalpine areas [120].  Its erosion control potential is listed as medium in Utah and Montana, and high in Colorado [30].  Because this wide-ranging tree exhibits a large degree of genetic variation, seed or nursery stock for rehabilitation projects should come from a local source.  Transplanting nursery stock is generally more successful than direct seeding [92].  Seedlings exhibit very slow initial growth and are therefore usually outplanted as 2- to 3-year-old seedlings [41].  Wild seedlings may also be transplanted [120].  A maximum spacing of 10 x 10 feet (3 x 3 m) has been recommended for seedlings or transplants [120].  Methods for collecting, processing, testing, storing, and planting subalpine fir seeds have been discussed in detail [32,41]. OTHER USES AND VALUES : Subalpine fir is sometimes used as a landscape plant to produce screenings or windbreaks [114].  In the Pacific Northwest it is sometimes transplanted into rock gardens or simulated subalpine settings [69]. Native Americans used various parts of subalpine fir for numerous purposes.  A hair tonic was prepared by mixing powdered needles with deer grease.  Finely ground needles were also sprinkled on open cuts. Sticky resin collected from the bark was boiled and used as an antiseptic for wounds or as a tea for colds.  Boughs were placed in rooms for their aroma, and pulverized needles were used as a body scent or as perfume for clothing [53]. Resin from the bark is used in the optical industry and in laboratories as a cement for lenses and microscope slides [71]. OTHER MANAGEMENT CONSIDERATIONS : Timber harvest:  Shelterwood and individual tree selection silvicultural methods favor subalpine fir over Engelmann spruce, lodgepole pine (Pinus contorta), and Douglas-fir (Pseudotsuga menziesii); clearcutting and group selection cutting favor subalpine fir over Pacific silver fir (Abies amabilis), grand fir (A. grandis), and mountain hemlock (Tsuga mertensiana) where they grow together [11].  The seed tree method is generally not used because of the susceptibility of subalpine fir to windthrow [11].  In the Rocky Mountains, clearcutting and shelterwood cutting have been the most commonly used harvesting methods in old-growth Engelmann spruce-subalpine fir stands because these stands tend to be even-aged and overmature [8].  Uneven-aged silviculture can pose a problem because residual subalpine fir trees damaged during thinning operations are susceptible to attack by decay fungi. Silvicultural systems and cutting methods for managing subalpine fir have been discussed in detail [7,8,9,11]. Pests and diseases:  Subalpine fir is attacked by numerous insects.  The most destructive seem to be the western spruce budworm, western balsam bark beetle, and balsam woolly aphid [11].  Subalpine fir is one of the most common hosts of the western spruce budworm.  This pest generally attacks low and middle elevation subalpine fir forests but is largely absent from high elevation forests [21].  The balsam woolly aphid has virtually eliminated subalpine fir from some stands in the Cascades [11].  Other insect pests include the Douglas-fir tussock moth, western black-headed budworm, and fir engraver beetle. Subalpine fir is susceptible to annosus root disease, caused by the fungus Heterobasidion annosum, which results in root and butt decay. Outbreaks of this disease are often centered around large 20-year-old or older fir stumps that contain the fungus' fruiting bodies [102]. Subalpine fir is most seriously affected by this disease in the northern and central Rocky Mountains [123], and is affected to a lesser extent in the Pacific Northwest [102].  Subalpine fir is susceptible to several other wood rotting fungi that cause heart, trunk, butt, or root rots, including brown stringy rot, red heart rot, red ring rot, shoestring rot, brown cubical rot, white spongy root rot, and white pocket rot [39].  Trees weakened by wood rots often become infested by fir engraver beetles and usually succumb to windfall and breakage [11].  Fir broom rust is another common problem in Engelmann spruce-subalpine fir stands and causes bole deformation, spike tops and wind breakage, and makes trees more susceptible to decay fungi [11]. Habitat for threatened and endangered species:  Old-growth subalpine fir stands in northern Idaho may provide critical habitat for woodland caribou [23].  Numerous subalpine fir habitat types, especially those containing huckleberries (Vaccinium spp.), provide critical habitat for grizzly bears [127].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Abies lasiocarpa
GENERAL BOTANICAL CHARACTERISTICS : Subalpine fir is a native, coniferous, evergreen tree.  It is the smallest of the eight species of fir native to the western United States.  Five growth forms, each apparently an adaptation to a particular environment, are described below [9,54]: 1.  The typical form is found throughout much of the subalpine zone. These trees have an extremely narrow and dense crown with short branches.  Trees growing in openings retain their lower branches, which often droop and extend down to the ground.  Trees growing in the overstory may be clear of lower branches for 20 to 30 percent of the tree's height. 2.  A somewhat broad-crowned, bullet-shaped tree is more typical of older specimens and drier climates.  3.  A mature tree with a layered apron is occasionally found in some areas. 4.  A flag form tree often occurs at timberline.  These individuals are characterized by an upright trunk that extends above a krummholzlike mat.  Branches on the trunk generally grow only along the leeward side of the trunk, giving the plant a flaglike appearance. 5.  The krummholz form is typical of alpine areas above timberline.  In these areas, because of cold temperatures and severe winds, subalpine fir grows in dwarfed, shrubby mats along the ground, and is often much broader than it is tall. The typical form often grows to heights of 60 to 100 feet (18-30 m), and trunk diameters reach 18 to 24 inches (46-61 cm) [39].  Trees up to 130 feet (40 m) tall and 30 inches (76 cm) in diameter have been found but are rare [39].  Subalpine fir grows very slowly; 150- to 200-year-old trees are usually only 10 to 20 inches (25-50 cm) in diameter [39]. Trees seldom live more than 250 years because they are very susceptible to heart rots [9]. Needles are blunt tipped, flattened, and 1 to 1.2 inches (2.5-3 cm) long [25,57].  Bark on young trees is thin, gray, and smooth, with numerous resin vesicles; on older trees it is shallowly fissured and scaly [9]. Corkbark fir is generally recognized by its creamy-white, thick, corky bark [25].  The root system generally is shallow but under favorable conditions may develop relatively deep laterals [9]. Subalpine fir is monoecious.  Single or small bunches of erect female cones occur in the upper part of the crown on the upper side of young branches.  Dense clusters of the smaller male cones occur lower on the crown on the underside of 1-year-old twigs.  Subalpine fir seeds are 0.23 to 0.28 inch (6-7 mm) long and have broad wings about 0.4 inch (1 cm) long [25,121].  Corkbark fir seeds are about 70 percent larger than subalpine fir seeds [39]. RAUNKIAER LIFE FORM :    Undisturbed State: Phanerophyte (mesophanerophyte)    Undisturbed State: Phanerophyte (microphanerophyte)    Undisturbed State: Phanerophyte (nanophanerophyte) Krummholz form    Burned or Clipped State: Therophyte REGENERATION PROCESSES : Cone and seed production:  Subalpine fir can begin producing cones when 20 years old and 4 or 5 feet (1.2-1.5 m) tall, but under closed forest conditions seed production is generally not significant until trees are older and taller [11].  Corkbark fir generally does not produce cones until about 50 years old [39].  Nearly all cones are produced on the uppermost part of the crown.  Maximum seed production is by dominant trees between 150 and 200 years old [39].  Yearly seed production is very erratic; good seed crops are produced every 3 to 5 years, with light crops or crop failures in between [39,40,54,86].  Corkbark fir is also a poor seed producer, having more crop failure years than good seed crop years [9].  Subalpine fir averages 34,800 seeds per pound (76,700/kg), while corkbark fir seeds average 22,300 per pound (49,150/kg) [41]. Seed predation:  Insect pests reduce seed yields by feeding on cones and seeds; however, the magnitude of loss is variable [11].  Red squirrels cut and cache large quantities of subalpine fir cones [9].  After dispersal, numerous small rodents and birds consume seeds from the ground. Dispersal:  Mature subalpine and corkbark fir seeds have a large wing and are dispersed primarily by wind in the fall as cones disintegrate on the tree.  Seeds travel primarily in the direction of prevailing winds, but upslope drafts can influence dispersal at low and middle elevations [9].  Studies in Colorado showed that about one-half of subalpine fir seeds dispersed into clearcuts fell within 100 feet (30 m) of the clearcut's windward edge, while the remainder fell within 260 feet (80 m) of the edge [86].  Some seeds are also dispersed by red squirrels which cut and cache cones before they disintegrate; seeds commonly germinate from these middens, forming thickets [71]. Germination and viability:  Seeds overwinter under or in snow.  This cold, moist stratification is required for germination [41]. Germination begins in the spring a few days after snowmelt and is usually completed within a few weeks [9,106].  Percent germination is low due to unsound seed; about 31 to 38 percent for subalpine fir and 26 to 33 percent for corkbark fir [39,41].  Under natural conditions seeds remain viable for 1 year [41].  Stratification procedures for stored seeds have been described in detail [41,74]. Seedling establishment and survival:  Seedlings establish best on mineral soil seedbeds but will also establish on other surfaces including litter, duff, and decaying wood [11].  Because Engelmann spruce requires a mineral soil seedbed, subalpine fir seedlings usually outnumber spruce seedlings in the understory of spruce-fir stands. Thus, even though it is short-lived, many ecologists consider subalpine fir better able to regenerate under climax conditions than Engelmann spruce.  Subalpine fir is very shade tolerant and easily establishes under a closed canopy.  Throughout the Rocky Mountains subalpine fir is reproducing abundantly under conditions of dense shade and is often abundant as seedlings and saplings in spruce-fir forests, even where Engelmann spruce dominates the overstory [11,67]. At higher elevations, seedling survival is sometimes greater on duff seedbeds because the duff helps protect seedlings from high-intensity summer rain storms and frost heaving [35].  At lower elevations, seedling densities are often greater on mineral soils [35].  In a spruce-fir forest in southeastern Wyoming, Knapp and Smith [67] found that 42 percent of subalpine fir seedlings were on litter deeper than 1 inch (2.5 cm), compared with only 5 percent of Engelmann spruce seedlings.  Subalpine fir is able to establish in duff because of its rapid root growth.  Comparing seedlings grown in a greenhouse, subalpine fir's taproot length (29 mm) 2 weeks after germination was over 200 percent greater than the taproot length of Engelmann spruce (9.4 mm) [67]. Growth:  Subalpine fir seedlings grow very slowly.  One-year-old seedlings are frequently less than 1 inch (2.5 cm) tall [11].  One study found 15-year-old seedlings averaged only 11 inches (28 cm) in height on burned-over slopes, 10 inches (25 cm) on cut-over dry slopes, and 6 inches (15 cm) on cut-over wet flats [11].  Under favorable conditions trees reach a height of 4 to 5 feet (1.2-1.5 m) in 20 to 40 years [11]. Under a closed canopy, trees 4 to 6 feet tall (1.2-1.8 m) are often 35 to 50 years old. Vegetative reproduction:  Near timberline subalpine fir frequently reproduces by layering, probably as a result of heavy snow, wind, and cold temperatures which restrict growth away from the ground.  Layering often results in clusters of subalpine fir growing near timberline [13]. Under closed forest canopy, reproduction by layering is negligible. SITE CHARACTERISTICS : Subalpine fir is a middle to upper elevation mountain conifer.  It generally occupies sites with a short growing season caused by cold winters, cool summers, frequent summer frosts, and heavy snowpack.  It forms extensive forests between warm and dry lower elevation forests of Douglas-fir, white fir (Abies concolor), grand fir, lodgepole pine, or blue spruce (Picea pungens) and higher elevation alpine tundra [23,29,112,125].  At its lower elevational limits, subalpine fir is often restricted to streambottoms, ravines, frosty basins, or north exposures.  It increasingly occupies westerly and easterly aspects with increasing elevation and may occupy all aspects at upper timberline [23,91]. Stand condition and associated conifers:  Throughout its range, subalpine fir is most commonly associated with Engelmann spruce.  These two species frequently occur as codominants forming widespread subalpine forests.  In the central and southern Rocky Mountains, Engelmann spruce commonly makes up 70 percent of overstory trees, with subalpine fir dominating the understory [9].  Within spruce-fir forests of this region, Engelmann spruce tends to be more important at higher elevations and on wetter sites, while subalpine fir is more abundant on drier lower elevation sites [9].  In the northern Rocky Mountains, subalpine fir typically dominates climax stands, but Engelmann spruce becomes increasingly important on moist, cool sites [23,91].  Other associates which vary by latitude and elevation are listed below [39]: Location              Elevation   Associates northern Rocky Mtns   low         western white pine (Pinus monticola),                                   Douglas-fir, western larch (Larix                                   occidentalis), grand fir, western hemlock                                   (Tsuga heterophylla), western redcedar                                   (Thuja plicata)                        high       lodgepole pine, subalpine larch (Larix                                   lyallii), whitebark pine (Pinus                                   albicaulis), mountain hemlock central Rocky Mtns     low        lodgepole pine, Douglas-fir, aspen                                   (Populus tremuloides), blue spruce                        high       whitebark pine, limber pine (Pinus                                   flexilis), bristlecone pine (P. aristata) southern Rocky Mtns    low        white fir, Douglas-fir, blue spruce, aspen                        high       corkbark fir Cascade Mtns           low        Pacific silver fir, mountain hemlock,                                   lodgepole pine                         high       mountain hemlock, whitebark pine Understory associates:  Understory vegetation is extremely variable, changing with elevation, exposure, and soil moisture.  Habitat type and plant association guides describe characteristic understory plants for differing sites. Elevation:  Alexander and others [11] described the following elevational ranges for subalpine fir: Coast Range of southeastern Alaska - subalpine fir is found from sea level to 3,500 feet (0-1,067 m).  Coast Range and interior plateaus of Yukon Territory and British Columbia - subalpine fir is found from 2,000 to 5,000 feet (610-1,524 m). Olympic and Cascade Mountains of Washington and Oregon - subalpine fir is generally found from 4,000 to 6,000 feet (1,219-1,829 m), but may be found as low as 2,000 feet (610 m) along cold streambottoms and on lava flows, and as high as 8,000 feet (2,438 m) on sheltered slopes. Rocky Mountains of British Columbia and Alberta south of the Peace River - subalpine fir is found from 3,000 to 7,000 feet (914-2,134 m) but is more abundant above 5,000 feet (1,524 m). Rocky Mountains of Montana and Idaho and associated ranges of eastern Oregon and Washington - subalpine fir grows from 2,000 to 11,000 feet (610-3,353 m) but is most common at 5,000 to 9,000 feet (1,524-2,743 m). Rocky Mountains of Wyoming, Utah, and Colorado - subalpine fir occurs from 8,000 to 11,500 feet (2,438-3,506 m) but is most common at 9,000 to 11,000 feet (2,743-3,353 m). Rocky Mountains of New Mexico and Arizona - subalpine fir occurs from 8,000 to 12,000 feet (2,438-3,658 m) but is usually found on north slopes from 9,500 to 11,000 feet (2,896-3,353 m). SUCCESSIONAL STATUS : In the Rocky Mountains, subalpine fir is a shade-tolerant climax species favored by long fire-free intervals.  Its seedlings outcompete spruces, lodgepole pine, and Douglas-fir when light intensities are less than 50 percent of full sunlight, but cannot compete with these conifers under brighter light [11].  In Montana and Idaho and in the mountains of eastern Washington and eastern Oregon, subalpine fir often forms pure stands at climax, but it may also mix with Engelmann spruce, which, although considered to be seral to subalpine fir, outlives it and persists to climax.  In the Rocky Mountains north and south of Montana and Idaho, Engelmann spruce and subalpine fir may codominate at climax [11]. Throughout much of the Cascade Mountains subalpine fir grows as a shade-intolerant, seral species and is gradually replaced by more shade-tolerant associates such as Pacific silver fir, grand fir, and mountain hemlock [43].  It often invades recently disturbed areas with lodgepole pine.  It also pioneers harsh sites on raw geologically young surfaces such as lava flows and talus slopes and on climatically harsh sites near timberline [43]. In areas where subalpine fir is a climax dominant, succession following disturbance varies depending upon the severity and type of disturbance, elevation, and the availability of conifer seeds.  Subalpine fir may establish immediately following disturbances if mature trees survive to provide seeds and seral species such as lodgepole pine and aspen are scarce.  Near treeline, it may take 100 years or more for subalpine fir to establish seedlings following fire because an increase in herbaceous species prevents seeds from reaching mineral soil and the harsh climate kills many seedlings that do establish [18,109].  Aspen and lodgepole pine are the most common seral species.  They often form pure stands and completely dominate low and middle elevation stands within the subalpine fir zone following large fires [5,68].  These species grow rapidly and quickly overtop any subalpine fir seedlings that may establish at the same time.  Aspen stands can sometimes persist for decades or even centuries when conifer seed trees are eliminated [29].  When lodgepole pine establishes immediately following stand-destroying fires, it often forms even-aged dense stands that dominate for 100 to 300 years. Because it is very shade tolerant, subalpine fir eventually establishes under the pine canopy, usually within 100 years, and attains dominance as the pine stand begins to break up [90,98]. In many of the warmer and lower elevation subalpine fir habitat types, subalpine fir has not achieved climax dominance because of repeated fires which favor shade-intolerant seral conifers.  Many of these habitat types are in midsuccessional stages.  Lodgepole pine, western larch, western white pine, or Douglas-fir dominate the overstory, but subalpine fir seedlings and saplings occur in the understory [93,113]. SEASONAL DEVELOPMENT : Subalpine fir requires 2 years to complete its reproductive cycle [115]. Cones are initiated in the spring of the first year as microscopic primordia within vegetative buds.  Bud differentiation occurs in midsummer, and separate seed-cone and pollen-cone buds develop until each becomes dormant in the fall [115].  During the spring of the second year, cone buds resume growth and conelets are recognizable in the early spring.  During the second year, reproduction phenology generally proceeds as follows: Phenological event  Location      Timing of event           Reference flowering           sw MT, nw WY   mid-June - early July     cones full size         "               late August          seeds dispersed         "         early Sept - early Oct     [41,101] flowering           nw MT, n ID   mid-June - early July      cones full size         "         late July - early Aug      seeds dispersed         "               mid-Sept             [41,101] flowering               OR         late May - early July     seed dispersal          "             early Oct               [41] male bud burst      Linn, OR         early to mid-May        female bud burst        "            mid to late May         pollen shed             "                 June               seed dispersal begins   "              early October          [45] flowering        AZ, San Fran.Peaks     late June            cone ripening           "           mid-Sept - early Oct     seed dispersal          "           late Sept - early Oct     [41]

FIRE ECOLOGY

SPECIES: Abies lasiocarpa
FIRE ECOLOGY OR ADAPTATIONS : Plant adaptations to fire:  Subalpine fir is very fire sensitive and generally suffers high mortality even from low intensity fires.  It relies on wind-dispersed seeds which readily germinate on fire-prepared seedbeds to colonize burned areas.  The occasional mature tree which survives fire, those escaping fire in small, unburned pockets, and trees adjacent to burned areas provide seeds to colonize burned sites.  In subalpine habitats, scattered subalpine fir trees often escape fire because of discontinuous fuels, broken and rocky terrain, and the moist and cool environment [78,87,91]. Fire regime:  Subalpine fir habitat types vary from cold and wet at higher elevations to warm and moist or cool and dry at lower elevations. This environmental gradient influences the mean fire return interval (MFRI).  Relatively dry lower elevation subalpine fir habitat types have more frequent and less intense fires than moist middle and upper elevation subalpine fir habitat types [12,91].  Such forests in the Bitterroot National Forest in Montana have a MFRI of 17 to 28 years [14].  Fires at this frequency kill subalpine fir and keep these forests dominated by seral conifers such as lodgepole pine, Douglas-fir, or western larch.  Moist, middle and upper elevation subalpine fir habitat types, however, generally experience high intensity stand-replacing fires at intervals of 100 years or more.  Mean fire return intervals for middle and upper elevation subalpine fir habitat types in several areas are presented below: Location                  Community dominants           MFRI    Reference                                                        (years) Kananaskis Park, AB      subalpine fir, spruce,          90       [12]                          lodgepole pine         northern Cascades, WA    subalpine fir                 154        [2] northern Cascades, WA    subalpine fir-lodgepole pine  109        [2] Olympic NP, WA           subalpine fir                 150        [116] Yellowstone NP, WY       subalpine fir                 300-350    [98] Coram Exp. Forest, nw MT western larch, Douglas-fir,   117-146    [129]                          lodgepole pine, subalpine fir Fuels and fire behavior:  The fuel structure in subalpine-fir-dominated stands promotes highly destructive stand-destroying fires.  Fuel loads in subalpine fir stands are greater than in lower elevation montane stands because the cool and moist environment slows the decomposition of organic matter allowing fuels to accumulate more rapidly [1].  Fuel beds tend to be irregular, with over twice as much fuel accumulating under the narrow-crowned trees as between them [116].  The needles are small and fine and form a compact fuel bed in which fire spreads slowly [34]. These concentrated, slow burning fuels frequently produce flames high enough to reach subalpine fir's low-growing dead branches [116].  Thus crowning is common in subalpine fir stands. Once a crown fire begins, it spreads easily because subalpine fir has a tendency to grow in dense stands and has highly flammable foliage.  A lightning strike on May 7, 1987, in a subalpine fir-mountain hemlock stand in Mount Rainier National Park started a crown fire even though the ground was still partially snow covered.  The fire spread slowly through the tree crowns by (1) igniting lichens draped along the fine branches, (2) preheating and igniting the foliage, and (3) spreading to a nearby tree by igniting its lichens [62]. POSTFIRE REGENERATION STRATEGY :    crown-stored residual colonizer; short-viability seed in on-site cones    secondary colonizer; off-site seed carried to site after year 2    off-site colonizer; seed carried by wind; postfire years 1 and 2

FIRE EFFECTS

SPECIES: Abies lasiocarpa
IMMEDIATE FIRE EFFECT ON PLANT : Subalpine fir is one of the least fire-resistant western conifers.  It is very susceptible to fire because it has (1) thin bark that provides little insulation for the cambium, (2) bark which ignites readily, (3) shallow roots which are susceptible to soil heating, (4) low-growing branches, (5) a tendency to grow in dense stands, (6) highly flammable foliage, and (7) moderate to heavy lichen growth [37,111]. Subalpine fir forests are normally subject to highly destructive crown fires that occur at 100-year or longer intervals.  Such fires typically kill all subalpine fir trees.  Subalpine fir is also very susceptible to surface fires because fine fuels which are often concentrated under mature trees burn slowly and girdle the thin-barked bole [34]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Following fire, subalpine fir reestablishes via seeds dispersed by wind from trees surviving in protected pockets or from trees adjacent to burned areas.  Subalpine fir readily establishes on burned mineral soil seedbeds [107].  Ash does not affect germination, but if it is deep, it can prevent a seedling's roots from reaching mineral soil [85]. Although seedling establishment is often favored by shade, it will establish in full sunlight following fire [87]. The rate of establishment is quite variable, and depends on the proximity of the seed source (because the heavy seeds are dispersed over short distances) and seed production during the year of the fire and immediate postfire years.  In general, subalpine fir seedling establishment is very slow in areas suffering large, continuous crown fires but is relatively rapid on small burned-over areas where a seed source is nearby [90,124,128].  Three years after a late August wildfire in northern Colorado, in a dense, mature stand composed of Engelmann spruce, subalpine fir, and lodgepole pine, subalpine fir had established 15,200 seedlings per acre (37,500/ha) on small burns that were less than one-tenth of an acre in size.  But on areas within the middle of the main burn, subalpine fir had established only 12 seedling per acre (30/ha) 3 years after the fire [16].  In Colorado, Peet [90] found a 75-year-old burn that had few conifer seedlings even though an old-growth subalpine fir-Engelmann spruce stand was 218 yards (200 m) away. Reinvasion into large burns is slow because much of the seed source is destroyed.  However, sometimes sporadic survivors provide a limited seed source so that a small number of seedlings establish quickly following fire.  When this occurs, large quantities of seeds are dispersed several decades later as the early invading seedlings mature and reach cone bearing age [128]. On areas where subalpine fir is abundant and lodgepole pine scarce before burning, subalpine fir establishes quickly following fire if sufficient numbers of seed trees survive or are near the burn.  However, if lodgepole pine is present prior to burning, it usually seeds in aggressively and assumes a dominant role because it quickly overtops any fir seeding in at the same time [34].  Subalpine fir can be suppressed for several decades in seral lodgepole stands which develop following fire; one-hundred-year-old individuals may be only 3 feet (0.9 m) tall [90].  It may take 50 to 150 years after a fire for substantial subalpine fir establishment under dense lodgepole pine stands [18,109,128]. In the Olympic Mountains, tree seedling establishment following fires in closed mountain hemlock-subalpine fir forests was higher during wet growing seasons than during dry growing seasons.  Establishment rates were higher near the edge of a fire or near survivors than in areas removed from a seed source [4].  On many burned areas, subalpine fir did not establish seedlings for several years because of poor seed crops. On some burns there was a lag time of 40 to 50 years after fire before there was substantial seedling establishment.  This was a result of early invading trees maturing and dispersing seeds [4]. High elevation subalpine fir stands that have burned often remain open for several decades or more [18,31].  The harsh environment near treeline makes it difficult for tree seedlings to establish and survive [18].  Grasses and sedges may form a mat in subalpine meadows which prevents tree seeds from reaching mineral soil [109]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : For further information on subalpine fir response to fire, see Fire Case Studies. Hamilton's Research Project Summary and Research Papers (Hamilton 2006a, Hamilton 2006b)provide information on prescribed fire and postfire response of plant community species, including subalpine fir, that was not available when this species review was originally written. FIRE MANAGEMENT CONSIDERATIONS : Subalpine fir is very fire sensitive and is often killed even by surface fires.  Following timber harvest, on sites where subalpine fir is not a preferred species, light surface fires may be used to kill subalpine fir and promote the establishment of other conifers [93].  Fuels remain moist in many high elevation subalpine fir habitat types during most of the year, leaving only a short time period during certain years when prescribed burning can take place [63,91]. Subalpine fir seeds germinate poorly in soils under burned slash piles [130] but readily germinate on mineral soil seedbeds prepared by broadcast burning [16,107].

FIRE CASE STUDIES

SPECIES: Abies lasiocarpa
FIRE CASE STUDY CITATION : Uchytil, Ronald J., compiler. 1991. Effects of a prescribed crown fire on Table Mountain, Washington, on subalpine fir. In: Abies lasiocarpa. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ []. REFERENCE : Woodard, Paul Michael. 1977. Effects of prescribed burning on two different-aged high-elevation plant communities in eastern Washington. Seattle, WA: University of Washington. 228 p. Dissertation. [124]. SEASON/SEVERITY CLASSIFICATION : fall/severe STUDY LOCATION : The study took place in the Cascade Mountains in central Washington approximately midway between the towns of Ellensburg and Wenatchee. PREFIRE VEGETATIVE COMMUNITY : The study area is within the subalpine fir (Abies lasiocarpa) zone.  Two adjacent but different-aged stands were burned.  One area was a young, lodgepole pine (Pinus contorta) thicket, the other was a old, decadent stand of lodgepole pine with subalpine fir and Engelmann spruce (Picea engelmannii) attaining dominance.  Each is described below. Lodgepole pine thicket - Lodgepole pine is the overstory dominant, but subalpine fir and Engelmann spruce are also principal components. Subalpine fir and Engelmann spruce are codominants in the intermediate size classes.  Subalpine fir is the only tree regenerating under the canopy.  Understory plants include elk sedge (Carex geyeri), Hood sedge (C. hoodii), heartleaf arnica (Arnica cordifolia), broadleaf arnica (A. latifolia), bigleaf lupine (Lupinus polyphyllus), and dwarf bilberry (Vaccinium myrtillus). Snag area - This is a decadent lodgepole pine stand, with subalpine fir and Engelmann spruce dominating the overstory canopy.  Fir and spruce codominate the intermediate size classes, but subalpine fir is the only tree regenerating under the canopy.  Predominant understory plants include subalpine fir, elk sedge, broadleaf arnica, and mosses (Rhacomitrium canascens, Polytridum commune). TARGET SPECIES PHENOLOGICAL STATE : NO-ENTRY SITE DESCRIPTION : The burned site is a southwest facing gentle slope at an elevation of 5,596 to 5,776 feet (1,706-1,761 m).  A total of 27 acres (10.9 ha) were burned.  Climate:  The climate is typical of most areas within the subalpine zone.  The winters are cold and wet and the summers cool and dry.  Frost and freezing temperatures can occur during any month of the year.  Over 70 percent of precipitation falls as snow between October and March. Soil and duff:  The two stands have similar soils, but the soil in the snag area is more fertile and better developed.  In both areas soils are derived from basalt residium, have a clay-loam texture, and average 6 inches (15.2 cm) deep.  The effective rooting depth was about 20 inches (51 cm) in the thicket area, and 20 to 40 inches (51-102 cm) in the snag area.  On both areas, duff was generally from 1 to 4 inches (0.4-1.6 cm) thick.  The mean depth of duff was 2.3 inches (5.94 cm) on the thicket area, and 1.9 inches (4.92 cm) on the snag area.  Fuel loading:  Prior to burning, mean fuel loads were as follows:           fuels                    thicket area          snag area                                 tons/acre tonnes/ha  tons/acre tonnes/ha dead and down wood  0.0-0.25 inch (0.0-0.6 cm)        0.6       1.3        0.9       2.0  0.26-0.99 inch (0.61-2.5 cm)      2.1       4.6        2.9       6.4  1.0-3.0 inches (2.6-7.6 cm)       4.4       9.8        5.1      11.4  > 3.0 inches (7.6 cm) rotten     17.4      38.9       30.9      69.2  > 3.0 inches (7.6 cm) solid      17.0      38.1       50.1     112.2 litter                            41.4      92.7       89.8     201.2 duff                              32.5      72.8       30.0      67.1 FIRE DESCRIPTION : The fire was ignited with drip torches on September 30, 1975, taking approximately 1 hour and 20 minutes to ignite the entire 27 acres (10.9 ha).  Actual fire prescriptions at the time of ignition were as follows: ambient air temperature = 60 to 63 degrees F (16-17 C) relative humidity = 19-21 percent wind = calm gusts to 15.6 miles/hr (26 km/hr), from the south-southwest days since last rain = 15 fine fuel moisture content = 13 percent Within about 10 minutes after ignition, trees began to crown out. Fire behavior for each area is summarized below: Snag area - Crowning occurred throughout most of the snag area.  Flame heights were estimated to be 125 feet (38 m) by one observer, and 50 feet (15 m) above the tops of 90 foot (27 m) crowns by two other observers.  The fire consumed all small downed and dead wood from 0.0 to 1 inch (0-2.5 cm) in diameter, as well as needles and small twigs on live standing trees.  Ninety-six percent of down and dead fuels less than 3 inches (7.6 cm) in diameter were consumed.  In general, 90 to 100 percent of the duff layer was removed.  Many trees less than 3 inches (7.6 cm) in diameter at the base were completely consumed, and nearly all standing snags were blown down or burned down.  Where crowning occurred, the only thing that remained immediately following the fire was reddened soil, ash-covered soil, large-diameter logs, and dead trees. Lodgepole thicket - Fire within the lodgepole thicket was much less severe.  The crown fire which occurred within the snag area stopped when it met the boundary of the lodgepole thicket.  Dead and down fuels less than 3 inches (7.6 cm) in diameter were reduced by 70 percent.  Dead and down fuels greater than 3 inches (7.6 cm) in diameter were reduced 34 percent.  Duff was reduced about 25 percent. FIRE EFFECTS ON TARGET SPECIES : Where crowning occurred subalpine fir suffered 100 percent mortality. Fire burning within the crown of live trees consumed all needles and small twigs leaving scorched skeletons.  Many live trees less than 3 inches (7.6 cm) in diameter at the base were completely consumed. One year after this fire, subalpine fir seedlings established from seed produced by unburned trees adjacent to the burned area.  However, stocking was considered low and attributed to a poor seed crop. Seedling establishment was restricted to areas along the margin of the burn. Subalpine fir pre and postfire frequency and cover are summarized below:                          thicket area              snag area                       prefire   postfire        prefire   postfire frequency             100%        38%            100%       12% relative ground cover  37%        12%             20%        4% FIRE MANAGEMENT IMPLICATIONS : High intensity crown fires which are common in subalpine fir forests, kill all or nearly all seed trees within a burned area.  If subalpine fir is to naturally regenerate on this type of burn, seeds must come from adjacent unburned stands.  Because subalpine fir seeds are dispersed over relatively short distances, initial seedling establishment is restricted to the burn's edge.  Subalpine fir cone production can be erratic from year to year, with the best regeneration occurring during good seed crop years.

REFERENCES

SPECIES: Abies lasiocarpa
REFERENCES : 1. A. D. Revill Associates. 1978. Ecological eff. of fire and its mgmt. in Canada's national parks: a synthesis of the literature. Vols 1&2. Lit. Rev. & Annot. Bibliography. Ottawa, ON: Parks Canada, National Parks Branch, Natural Resources Division. 345 p. [3416] 2. Agee, James K.; Finney, Mark; DeGouvenain, Roland. 1990. Forest fire history of Desolation Peak, Washington. Canadian Journal of Forest Research. 20: 350-356. [11035] 3. Agee, James K.; Kertis, Jane. 1987. Forest types of the North Cascades National Park Service Complex. Canadian Journal of Botany. 65: 1520-1530. [6327] 4. Agee, James K.; Smith, Larry. 1984. Subalpine tree reestablishment after fire in the Olympic Mountains, Washington. Ecology. 65(3): 810-819. [6102] 5. Alexander, Billy G., Jr.; Fitzhugh, E. Lee; Ronco, Frank, Jr.; Ludwig, John A. 1987. A classification of forest habitat types of the northern portion of the Cibola National Forest, New Mexico. Gen. Tech. Rep. RM-143. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 35 p. [4207] 6. Alexander, Billy G., Jr.; Ronco, Frank, Jr.; Fitzhugh, E. Lee; Ludwig, John A. 1984. A classification of forest habitat types of the Lincoln National Forest, New Mexico. Gen. Tech. Rep. RM-104. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 29 p. [300] 7. Alexander, Robert R. 1974. Silviculture of subalpine forests in the central and southern Rocky Mountains: the status of our knowledge. Res. Pap. RM-121. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 88 p. [8223] 8. Alexander, Robert R. 1986. Silvicultural systems and cutting methods for old-growth spruce-fir forests in the central and southern Rocky Mountains. Gen. Tech. Rep. RM-126. 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