|
Wildlife, Animals, and Plants
|
|
Introductory
SPECIES: Pinus albicaulis | Whitebark Pine
ABBREVIATION :
PINALB
SYNONYMS :
NO-ENTRY
SCS PLANT CODE :
PIAL
COMMON NAMES :
whitebark pine
alpine pine
mountain pine
scrub pine
white pine
TAXONOMY :
The scientific name for whitebark pine is Pinus albicaulis Engelm.
There are no known hybrids between whitebark pine and other pines,
although a hybrid is suspected between whitebark pine and limber pine
(Pinus flexilis) near Jefferson City, Montana [15]. These two species
are very similar in habit and vegetative features but differ chiefly by
their ovulate (female) cones [9,23].
LIFE FORM :
Tree
FEDERAL LEGAL STATUS :
No special status
OTHER STATUS :
NO-ENTRY
COMPILED BY AND DATE :
Kathy Ahlenslager January 1987
LAST REVISED BY AND DATE :
Kathy Ahlenslager January 1988
AUTHORSHIP AND CITATION :
Ahlenslager, Kathleen E. 1987. Pinus albicaulis. In: Remainder of Citation
DISTRIBUTION AND OCCURRENCE
SPECIES: Pinus albicaulis | Whitebark Pine
GENERAL DISTRIBUTION :
The distribution of whitebark pine is split into two broad sections.
Western populations of whitebark pine extend from about 55 degrees N. in
western British Columbia, along the lower part of the Fraser River,
south into Washington and along the Cascade Mountains, southward through
the high mountains of Washington and Oregon into California. In
northern California, whitebark pine is scattered in isolated
populations, but farther south in the Sierra Nevada of central
California it is more continuous to its southern limit near Mount
Whitney about 37 degrees N. [4,9,23]. Eastern populations occur
southward from about 55 degrees N. in central Alberta, Canada, and
follow the northern Rocky Mountains south into western Montana and
central Idaho. Stands are extensive in northwestern Wyoming. Except
for disjunct populations in northeastern Nevada (about 41 degrees N.),
the southern and eastern limit of whitebark pine is the Wind River
Mountains of Wyoming [4,9,23]. Whitebark pine does not occur south of
the Wyoming Basin.
The distribution of whitebark pine is strongly influenced by Clark's
nutcrackers, which are important in the dispersal of seeds and
establishment of seedlings [32,43].
ECOSYSTEMS :
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES23 Fir - spruce
FRES26 Lodgepole pine
FRES44 Alpine
STATES :
CA ID MT OR UT WA WY AB BC
ADMINISTRATIVE UNITS :
BIHO CRLA GLAC GRTE KICA LAVO
MORA NOCA OLYM SEQU YELL YOSE
BLM PHYSIOGRAPHIC REGIONS :
1 Northern Pacific Border
2 Cascade Mountains
4 Sierra Mountains
8 Northern Rocky Mountains
9 Middle Rocky Mountains
11 Southern Rocky Mountains
KUCHLER PLANT ASSOCIATIONS :
K004 Fir - hemlock forest
K008 Lodgepole pine - subalpine forest
K015 Western spruce - fir forest
SAF COVER TYPES :
205 Mountain hemlock
206 Engelmann spruce - subalpine fir
208 Whitebark pine
209 Bristlecone pine
210 Interior Douglas-fir
219 Limber pine
256 California mixed subalpine
SRM (RANGELAND) COVER TYPES :
NO-ENTRY
HABITAT TYPES AND PLANT COMMUNITIES :
In western North America whitebark pine is a dominant or codominant
species in many high-elevation forests. In the Rocky Mountains, eastern
Cascades, and Blue Mountains, it is a minor component in mixed stands of
Engelmann spruce (Picea engelmannii) and subalpine fir (Abies
lasiocarpa). It is found with mountain hemlock (Tsuga mertensiana) in
the Cascades and British Columbia Coast Ranges. In the upper subalpine
forests of California, it is associated with subalpine fir, lodgepole
pine (Pinus contorta), western white pine (P. monticola), foxtail pine
(P. balfouriana), and limber pine (P. flexilis) [1,3,22]. Publications
using whitebark pine in classification schemes are listed below.
Coniferous forest habitat types of the Wind River Mountains, Wyoming. [42]
Forest habitat types of eastern Idaho-western Wyoming. [43]
Forest habitat types of Montana. [40]
Forest habitat types of northern Idaho: a second approximation. [8]
Forest vegetation of eastern Washington and northern Idaho. [11]
Steppe vegetation of Washington. [10]
VALUE AND USE
SPECIES: Pinus albicaulis | Whitebark Pine
WOOD PRODUCTS VALUE :
Whitebark pine is slow growing and after about 250 years reaches small
to moderately large sizes, depending on the site. The bole is generally
short and often of poor form for timber. In general, there is little
interest in whitebark pine for commercial timber, although it is
harvested in some localities in small amounts for lumber, posts, poles,
and firewood. Yields of merchantable timber are very low, about 10 to
20 feet per acre per year (0.7-1.4 m/ha/yr) in high-elevation stands
where it forms a short tree with large branches [3,8,14,19,25,40,48].
Higher yields of up to 40 feet per acre per year (48 m/ha/yr) have been
reported in better subalpine forest sites.
The best sites for treeline growth of whitebark pine in Montana are in
subalpine fir and sedge (Luzula hitchcockii) habitat types, Menziesia
(Menziesia ferruginea) phase. On such sites, whitebark reaches 70 to
100 feet (21-30 m) tall and 20 to 30 inches (50-70 cm) in d.b.h. at 250
to 300 years of age. When growing in association with whitebark pine,
Engelmann spruce grows larger and is the primary objective of management
[3].
Where whitebark pine is a component of better sites in spruce-fir
forests in Canada and the Rocky Mountains of the western United States,
whitebark pine produces good-quality timber with lumber properties
similar to western white pine. However, whitebark pine is graded lower
because of its darker appearance [3,14].
In some mountain valleys of southwestern Alberta between 5,000 and 6,000
feet (1,520-1,830 m), whitebark pine trees grow larger than associated
lodgepole pine and are harvested. The wood density of whitebark pine is
similar to local Douglas-fir (Pseudotsuga menziesii, but heavier than
hybrid spruce, subalpine fir, and lodgepole pine [12,15].
IMPORTANCE TO LIVESTOCK AND WILDLIFE :
Compared with other temperate coniferous forests, whitebark pine forests
are highly productive in terms of forage. Whitebark pine seeds are
present in early spring (from caches) and late fall when other wildlife
foods are scarce or low in digestibility. Seeds are wingless, large,
and high in calories. Although seed crops vary annually, they are an
important food source for numerous birds, rodents, and bears. Cones are
indehiscent and retain mature seeds. Thus, they can be harvested
efficiently [30,47].
The distribution of whitebark pine is strongly influenced by the
dispersal and establishment of seeds by Clark's nutcrackers. The
interaction between the two species is a result of coevolution and is
mutualistic. Clark's nutcrackers have evolved a sublingual throat pouch
in which to carry seeds to sites where they cache them about 2.5 inches
(1 cm) below the soil surface in groups of one to five. Birds harvest
and cache seeds in the late summer and fall for use during the following
winter and spring. Additional birds which feed on whitebark pine seeds
include William's sapsucker, white-headed woodpecker, mountain
chichadee, white-breasted nuthatch, Cassin's finch, red crossbill, pine
grosbeak, and blue grouse [20,33,44,45,46].
Bears in the Yellowstone area regularly eat pine seeds in the spring
(March to June) and fall (September and October). Most whitebark pine
seed eaten by grizzly and black bears are from red squirrel cone caches.
Rodents, such as red squirrels, Douglas' squirrels, ground squirrels,
and chipmunks, store large quantities of intact cones in middens at the
base of trees or underground in caches. Although deer mice cannot gnaw
the cones, they eat and cache loose seeds [26,46].
Whitebark pine survives where tree growth is limited and provides hiding
and thermal cover for wildlife. Cavity-nesting birds use tree trunks
and snags. Mule deer, elk, and predatory animals also use whitebark
habitat [40,45].
PALATABILITY :
The foliage of whitebark pine is not browsed to any extent by animals.
Blue grouse do eat the buds and needles of it in the winter [Arno pers.
comm. 1987]. The degree of use shown by livestock and wildlife species
for whitebark pine in Montana and Wyoming is rated as follows [13]:
MT WY
Cattle Poor Poor
Sheep Poor Poor
Horses Poor Poor
Pronghonr ---- Poor
Elk Poor Fair
Mule deer Poor Fair
White-tailed deer ---- Fair
Small mammals Fair Good
Small nongame birds Good Good
Upland game birds Good Good
Waterfowl ---- Poor
NUTRITIONAL VALUE :
Whitebark pine is rated poor in energy value and poor in protein value
[13].
COVER VALUE :
The degree to which whitebark pine provides environmental protection
during one or more seasons for wildlife species in Montana and Wyoming
is rated as follows [13]:
MT WY
Pronghorn ---- Poor
Elk Good Fair
Mule deer Good Fair
White-tailed deer ---- Poor
Small mammals Good Fair
Small nongame birds Good Good
Upland game birds Good Fair
Waterfowl ---- Poor
VALUE FOR REHABILITATION OF DISTURBED SITES :
Growth of whitebark pine in Montana, North Dakota, and Wyoming is
reported to be good on gentle to steep slopes. Its potential for
erosion control is low to medium. Short-term revegetation potential is
rated low; long-term potential is medium [13].
Whitebark pine has limited use in the rehabilitation of disturbed sites.
Vegetation recovery is slow in the exposed, dry, rocky sites where it is
found, and soil erosion can prevent complete restoration. Its survival
at high elevations indicates that it has potential for use in land
reclamation projects. Its slow growth and low seed germination are
handicaps in its use in reclamation [14,41,43].
OTHER USES AND VALUES :
Besides providing cover and food for numerous animals and birds, the
principal use of whitebark pine sites is watershed protection. In
western Montana whitebark pine habitats are the major sources of
streamflow. On steep terrain, trees help to stabilize snow, soil, and
rocks [3,14,19,23,30,40,43].
Seeds and inner tree bark of whitebark pine were included in the diets
of some Indians [20,28]. The species is also used to a limited extent
as an ornamental [28]. Although the open character of whitebark pine
sites appeals to many recreationists, the sites are often fragile and
degrade rapidly even with relatively low levels of recreation use [18].
MANAGEMENT CONSIDERATIONS :
Watershed protection and enhancement are of primary management
importance in whitebark pine areas. In addition, the slow rate of
vegetation recovery requires dispersed and low-impact recreation if the
aesthetic appeal of whitebark pine forests is to be maintained
[8,14,40].
Whitebark pine is replaced by more shade-tolerant species on many sites.
This can result from fire suppression, as well as damage by mountain
pine beetle and white pine blister rust. Unless these stands are
rejuvenated, seed crops of this pine species will diminish.
Rejuvenation may be accomplished by prescribed burning, seed plantings,
and propagation of rust-resistant trees. The recovery of grizzly bears
and their use of pine seeds is also a consideration for the management
of whitepark pine sites [3,14].
Damaging agents: Whitebark pine is very susceptible to white pine
blister rust (Cronartium ribicola). Large areas of whitebark pine in
northern Idaho and northwestern Montana have died from blister rust
[Arno pers. comm. 1987]. Infection occurs where adequate moisture
permits infestation of currants and gooseberries (Ribes spp.), alternate
hosts for the rust. White pine blister rust kills the upper
(cone-bearing) branches of large trees and sometimes entire trees.
Blister rust has a severe impact on saplings, thus often preventing
whitebark pine replacement [Arno pers. comm. 1987]. Several other
diseases also infect trees, generally with minor consequences [3,14,19].
Whitebark pine trees are also killed by the mountain pine beetle
(Dendroctonus ponderosae), often after being weakened by rust. The
inner bark of larger trees supports the growth of larvae which girdle
the trees. Numerous other pests infect whitebark pines, including dwarf
mistletoes (Arceuthobium spp.) and saprophytes [3,44].
BOTANICAL AND ECOLOGICAL CHARACTERISTICS
SPECIES: Pinus albicaulis | Whitebark Pine
GENERAL BOTANICAL CHARACTERISTICS :
Whitebark pine is a slow growing, long-lived, ectomycorrhizal, native
conifer characteristic of treeline. Trees often reach 400 to 700 years
of age. The oldest known cored tree is 750 years old and is in Mount
Robson Provincial Park, British Columbia [2,3,13,18,19,37].
Trees in well-developed stands are 50 to 70 feet (15-20 m) tall and 24
to 36 inches (60-90 cm) in d.b.h. Growing at the uppermost limits of
growth, trees usually are dwarfed or contorted. At upper treeline this
species takes on a spreading krummholz growth form and grows in isolated
cushions of "alpine scrub" 1 to 3 feet (0.3-1 m) tall [3,39].
In Crowsnest Forest, Alberta, the largest whitebark pine is at 107 feet
(37 m) high and 31 inches (79 cm) in d.b.h. [12]. The largest reported
whitebark pine in the United States is in the Sawtooth Range of central
Idaho and is 69 feet (21 m) in height and 9.5 feet (2.9 m) in d.b.h.
[3,41]. On good sites in the Flathead National Forest of Montana,
dominant whitebark pines are often 80 to 100 feet (24-30 m) tall [Arno
pers. comm. 1987].
Whitebark pine trees commonly have two or more trunks that are often
partially fused at the base. Electrophoretic evidence reveals that two
or more trunks of what appears to be a single tree are indeed separate
trees with distinct genotypes. This supports the idea that several
mature trees can arise from single seed caches [36] and that seeds
cached by Clark's nutcrackers are instrumental in the establishment of
trees [48]. Trees develop a deep and spreading root system on most
sites [3,19].
RAUNKIAER LIFE FORM :
Phanerophyte
REGENERATION PROCESSES :
The minimum seed-bearing age of whitebark pine trees is between 20 and
30 years, and the interval between large seed crops is 3 to 5 years. On
most sites significant amounts of seed occur only on trees greater than
80 years of age [47,Arno pers. comm. 1987]. Large seed crops are
produced at irregular intervals, interrupted by smaller crops and crop
failures [32]. Cone production fluctuates widely between years, and
variations in seed crops may play an important role in the intial
establishment of a stand.
Greater than 95 percent of the whitebark pine seed crop is harvested by
animals [Arno pres. comm. 1987]. The primary disperser of seeds is the
Clark's nutcracker. The seed-caching activities of nutcrackers in the
Sierra Nevada are reported in detail by Tomback [44,45,46,47] and in the
Absaroka and Teton Ranges of Wyoming by Lanner [33,22]. Evidence for
nutcrackers facilitating the regeneration and spread of whitebark pine
includes the rapid regeneration of trees on burned or clearcut areas,
the clumping of seedlings, and the disjunct occurrence of populations.
Besides Clark's nutcrackers, other vertebrates also harvest, feed on,
and cache whitebark pine seeds. However, they do not possess the
behavior to systematically disperse and cache the seeds. Rodents
disperse fewer seeds than nutcrackers, in shorter distances from parent
trees, and in sites less suitable for germination. Also, seed caches of
rodents are larger and have lower potential for successful tree
establishment. In addition, not many seeds are left to germinate from
rodent caches because the olfactory sense of rodents may allow them to
find and use caches more efficiently than nutcrackers [3,30,45,46,47].
Clark's nutcrackers are effective dispersers of seeds in part because of
the long distances they travel [45]. A nutcracker can carry as many as
150 seeds in its sublingual throat pouch and can store 850 seeds per
day, usually in caches of four to five seeds. Over a 42-day period one
bird may cache 32,000 seeds. They bury the seeds to 1.2 inches (3 cm)
in depth, which is suitable for germination. Nutcrackers store three to
five times their energetic requirements, so more seeds are buried than
are recovered. These seeds, along with abandoned caches, can germinate
and produce new trees.
Squirrel pressures on seed crops and blister rust damage have caused a
reduction in whitebark pine populations in Idaho and Washington [11].
Regeneration of whitebark pine is sporadic. In addition to consistant
losses from vertebrates, Eggers [14] reported that in the Rocky
Mountains those seeds that do survive have low germination rates. In
contrast, Tomback [46] reported a good survival rate of seeds from
caches in the Sierra Nevada, with 56 percent for the first year and 25
percent by the fourth year. Seedling survival rate after a fire was 25
percent [46].
At treeline in Jasper National Park, Alberta, it was found that higher
than average mean minimum summer temperature is one of the main factors
triggering seedling establishment for whitebark pine. In contrast,
there was little association between mean summer precipitation and
seedling germination [24]. Seeds from high-elevation krummholz stands
of whitebark pine have less germination capacity than do those from
lower elevations [48].
The viable period for seeds in cold storage, 0 to 5 degrees Fahrenheit
(-17 to 15 deg C), is 8 years [30]. Cold stratification of 90 to 120
days for fresh seeds is recommended by some researchers [32]. A
germination success of about 30 percent has been reported for seeds
soaked in cold water for 1 to 2 days, then stored damp in plastic bags
at 33 to 41 degrees Fahrenheit (0.5-5.0 deg C) for 90 to 120 days [15].
Seed immaturity and physical dormancy were cited for poor germination in
seeds collected near Cranbrook, British Columbia, at 7,200 feet (2,200
m). The growing season is short at high elevations, leaving little time
for seeds to mature. The lack of development of embryos was partially
overcome by exposing hydrated seeds to a temperature of 68 degrees
Fahrenheit (20 deg C) for 30 to 60 days. A temperature of 36 degrees
Fahrenheit (2 deg C) for 60 to 90 days broke dormancy. It was found
that poor germination resulted from the restraint imposed by the seed
coat which inhibited the protrusion of the radicle root. These
researchers concluded that it was not an inability of seeds to imbibe
water [35].
Removal of seed coats did not confer an advantage in germination, as
germination was 30 percent regardless of whether the coats were clipped
or entirely removed [35]. Sterilization only slightly increased
germination of seeds which received cold stratification. Germination
was significantly improved with a combined warm and cold treatment [35].
In another study, researchers found conflicting evidence, where the
limiting factor for germination could indeed be the restriction of water
uptake by the seed coat. When seeds were cut and exposed to germination
paper the germination rate was 61 percent. The rate was increased to 91
percent with cold stratification plus clipping [41].
SITE CHARACTERISTICS :
Whitebark pine grows on dry rocky sites on high mountains between 6,000
and 10,000 feet (1,800 and 3,030 m). It is characteristic of treeline
where it forms dense krummholz thickets. In Banff and Jasper National
Parks at treeline about 6,500 to 7,500 feet (2,000-2,300 m), whitebark
pine trees are dwarfed and isolated on dry exposed sites. Here at the
northern end of its range, it is a minor component of treeline.
Whitebark pine is an important component of high-elevation forests in
Idaho, Montana, and Wyoming between 5,900 and 10,500 feewt (1,800-3,200
m). In high-elevation forests in the Cascades of southern Oregon and
northern California between 8,000 and 9,500 feet (2,440 and 2,900 m),
whitebark pine is a major component of treeline [2,3].
Whitebark pine occurs at elevations as low as 5,000 feet (1,470 m) in
British Columbia and in the Cascades of Washington. The lowest reported
natural occurrence of whitebark pine is 3,600 feet (1,100 m) on Mt.
Hood in Oregon. In the southern Sierra Nevada, it commonly occurs up to
11,500 feet (340 m) in elevation [Arno pers. comm. 1987].
The dispersal of whitebark pine seeds by Clark's nutcrackers strongly
affects the distribution and abundance of this species. Trees occur on
dry rocky, subalpine slopes, and exposed ridges. Stands are generally
open with an undergrowth of low shrubs, forbs, and grasses [2,9,23].
Sites where whitebark pine occurs as a climax are drier than those where
it is seral.
Whitebark pine is important in areas where the mean annual precipitation
is 24 to 70 inches (600-1,800 mm) [3]. The climate is characterized by
cool summers and cold winters with deep snowpack. Trees have high frost
resistance and low shade tolerance.
Trees are found predominately on acidic substrates, although they also
have been reported on calcareous ones. Most soils under whitebark pine
stands are Inceptisols. The growth of whitebark pine in Montana and
Wyoming is reported as good on sandy-loam and loam, fair on gravels and
clay loams, and poor on clay [3,13,14,19,43].
SUCCESSIONAL STATUS :
In upper elevation subalpine forests, whitebark pine is generally seral
and competes with and is replaced by more shade-tolerant trees.
Subalpine fir, a very shade-tolerant species, is the most abundant
associate and most serious competitor of whitebark pine. Although
whitebark pine is more shade tolerant than lodgepole pine and subalpine
larch (Larix lyallii), it is less shade tolerant than Engelmann spruce
and mountain hemlock (Tsuga mertensiana). Whitebark pine is the
potential climax species on high exposed treeline sites and
exceptionally dry sites [2,3,14,47]. It sometimes acts as a pioneer
species in the invasion of meadows and burned areas [17,19]. On dry,
wind-exposed sites, the regeneration of whitebark pine may require
several decades, even though it is often the first tree to become
established [3,15,17,48].
The distribution of seral whitebark pine is strongly affected by the
dispersal of seeds by Clark's nutcrackers. The fact that bird
dispersion of seed occurs allows whitebark pine to be more widespread as
a seral species. The dispersal of seeds by them throughout subalpine
habitats is partly responsible for the status of whitebark pine as a
pioneer and postfire invader [43].
SEASONAL DEVELOPMENT :
Whitebark pine produces pollen during June and July. Cones mature in
late August and September of the second year. Female cones remain
closed until they fall to the ground, where they disintegrate by decay
and predation by animals. Few cones are available to fall, as most are
harvested in the trees by Clark's nutcrackers and red squirrels [3,14].
FIRE ECOLOGY
SPECIES: Pinus albicaulis | Whitebark Pine
FIRE ECOLOGY OR ADAPTATIONS :
The vulnerability of whitebark pine to fire is reduced by the open
structure of its stands and the dry, exposed habitats with meager
undergrowth in which it grows. Whitebark pine is favored by severe,
stand-replacing fires which burn shade-tolerant associated trees. Where
succession to shade-tolerant species is relatively rapid, fires are
important in moist sites for whitebark pine perpetuation [2].
Evidence from palynological and firescar studies shows that fires were
infrequent (fire intervals from 50 to 300 years) in whitebark pine
communities from the last Ice Age glaciation to the early 1900's. Since
1965 only a few thousand acres of seral whitebark pine have burned.
Largely due to fire suppression, less than 1 percent of the seral
whitebark habitat types have burned since then. At this rate the
"average" stand would burn every 3,000 years or longer [1,17].
With the lengthening of fire intervals, older stands become more
susceptible to pine beetle epidemics, which advance succession toward
dominance by shade-tolerant species. In addition, fire may stimulate
the growth of currents and gooseberries, the alternate hosts for white
pine blister rust, and thus the spread of the rust into whitebark pine
trees [2].
POSTFIRE REGENERATION STRATEGY :
Tree without adventitious-bud root crown
Initial-offsite colonizer (off-site, initial community)
FIRE EFFECTS
SPECIES: Pinus albicaulis | Whitebark Pine
IMMEDIATE FIRE EFFECT ON PLANT :
The moderately fire-resistant whitebark pine is favored by both creeping
surface or ground fires and severe fires. Both types of fire kill more
shade-tolerant and fire-sensitive asssociate species of whitebark pine,
such as subalpine fir. Hot surface fires that heat the cambium cause
fire injury or death to these thin-barked trees. Fires of low to
moderate severity can climb into trees if fuel ladders and downfall are
present, thus increasing the potential of torching. Most fires
occurring where whitebark pines grow are ignited by lightning and do not
spread very rapidly or cause severe tree injury [1,2].
DISCUSSION AND QUALIFICATION OF FIRE EFFECT :
NO-ENTRY
PLANT RESPONSE TO FIRE :
The regeneration of whitebark pine in small openings is probably the
result of surface fires. In contrast, the perpetuation of whitebark
pine on moist sites where succession to shade-tolerant species is
relatively rapid is probably due to severe fires. The occurrence of
whitebark pine and Engelmann spruce in subalpine basins and on moist
north slopes is probably the result of fire [2].
Postfire seedling survivor rate is reported as 25 percent. A maximum of
150 years is required for afforestation at current rates. Fires create
sites conducive to the planting of seeds by Clark's nutcrackers and for
seedling establishment [1,2,17,46,47].
DISCUSSION AND QUALIFICATION OF PLANT RESPONSE :
NO-ENTRY
FIRE MANAGEMENT CONSIDERATIONS :
NO-ENTRY
REFERENCES
SPECIES: Pinus albicaulis | Whitebark 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. 1986. Whitebark pine cone crops--a diminishing source
of wildlife food?. Western Journal of Applied Forestry. 3: 92-94. [341]
3. Arno, Stephen F.; Hoff, Raymond J. 1990. Pinus albicaulis Engelm.
whitebark pine. 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: 268-279. [13390]
4. Bailey, D. K. 1975. Pinus albicaulis. Curtis's Botanical Magazine.
180(3): 140-147; 1975. [376]
5. Barbour, Michael G.; Major, Jack, eds. 1977. Terrestrial vegetation of
California. New York: John Wiley & Sons. 1002 p. [388]
6. 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]
7. Cole, David N. 1982. Vegetation of two drainages in Eagle Cap
Wilderness, Wallowa Mountains, Oregon. Res. Pap. INT-288. Ogden, UT:
U.S. Department of Agriculture, Forest Service, Intermountain Forest and
Range Experiment Station. 42 p. [658]
8. Cooper, Stephen V.; Neiman, Kenneth E.; Steele, Robert; Roberts, David
W. 1987. Forest habitat types of northern Idaho: a second approximation.
Gen. Tech. Rep. INT-236. Ogden, UT: U.S. Department of Agriculture,
Forest Service, Intermountain Research Station. 135 p. [867]
9. Cronquist, Arthur; Holmgren, Arthur H.; Holmgren, Noel H.; Reveal, James
L. 1972. Intermountain flora: Vascular plants of the Intermountain West,
U.S.A. Vol. 1. New York: Hafner Publishing Company, Inc. 270 p. [717]
10. Daubenmire, R. 1970. Steppe vegetation of Washington. Technical Bulletin
62. Pullman, WA: Washington State University, College of Agriculture,
Washington Agricultural Experiment Station. 131 p. [733]
11. Daubenmire, Rexford F.; Daubenmire, Jean B. 1968. Forest vegetation of
eastern Washington and northern Idaho. Technical Bulletin 60. Pullman,
WA: Washington State University, Agricultural Experiment Station. 104 p.
[749]
12. Day, R. J. 1967. Whitebark pine in the Rocky Mountains of Alberta.
Forestry Chronicle. 1967 September: 278-283. [766]
13. Dittberner, Phillip L.; Olson, Michael R. 1983. The plant information
network (PIN) data base: Colorado, Montana, North Dakota, Utah, and
Wyoming. FWS/OBS-83/86. Washington, DC: U.S. Department of the Interior,
Fish and Wildlife Service. 786 p. [806]
14. Eggers, Douglas E. 1986. Management of whitebark pine as potential
grizzly bear habitat. In: Contreras, Glen P.; Evans, Keith E.,
compilers. Proceedings--Grizzly Bear Habitat Symposium; 1985 April
30-May 2; Missoula, MT. General Techinical Report INT-207. Missoula, MT:
U.S. Deptment of Agriculture, Forest Service, Intermountain Research
Station: 170-175. [851]
15. Ericson, John E. 1965. A suspected hybrid between Pinus ablicaulis
Engelm. and Pinus flexilis James. Proceedings of the Montana Academy of
Sciences. (25): 58- 59. [871]
16. Eyre, F. H., ed. 1980. Forest cover types of the United States and
Canada. Washington, DC: Society of American Foresters. 148 p. [905]
17. Fischer, William C.; Clayton, Bruce D. 1983. Fire ecology of Montana
forest habitat types east of the Continental Divide. Gen. Tech. Rep.
INT-141. Ogden, UT: U.S. Department of Agriculture, Forest Service,
Intermountain Forest and Range Experiment Station. 83 p. [923]
18. Forcella, Frank. 1978. Flora and chorology of the Pinus
albicaulis--Vaccinium scoparium association. Madrono. 25: 139-150.
[935]
19. Forcella, F.; Weaver, T. 1977. Biomass and productivity of the subalpine
Pinus albicaulis--Vaccinium scoparium association in Montana, USA.
Vegetatio. 35(2): 95-105. [934]
20. Forcella, Frank; Weaver T. 1980. Food production in the Pinus
albicaulis--Vaccinum scoparium association. Proceedings of the Montana
Academy of Science. 39: 73-80. [936]
21. 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]
22. Hall, Frederick C. 1973. Plant communities of the Blue Mountains in
eastern Oregon and southeastern Washington. R6-Area Guide 3-1. Portland,
OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest
Region. 82 p. [1059]
23. Hitchcock, C. Leo; Cronquist, Arthur; Ownbey, Marion. 1969. Vascular
plants of the Pacific Northwest. Part 1: Vascular cryptograms,
gymnosperms, and monocotyledons. Seattle, WA: University of Washington
Press. 914 p. [1169]
24. Hutchins, H. E.; Lanner, R. M. 1982. The central role of Clark's
nutcracker in the dispersal and establishment of whitebark pine.
Oecologia. 55: 192-201. [1228]
25. Kasper, J. B.; Szabo, T. 1970. The physical and mechanical properties of
whitebark pine. Forestry Chronicle. August: 315-316. [1310]
26. Kendall, Katherine Clement. 1981. Bear use of pine nuts. Bozeman, MT:
Montana State University; 1981. 27 p. M.S. thesis. [1331]
27. Kozlowski, T. T.; Ahlgren, C. E., eds. 1974. Fire and ecosystems. New
York: Academic Press. 542 p. [1374]
28. Krochmal, Arnold; Krochmal, Connie. 1982. Uncultivated nuts of the
United States. Agriculture Information Bulletin 450. Washington, DC:
U.S. Department of Agriculture, Forest Service. 89 p. [1377]
29. Kessell, Stephen R.; Potter, Meredith W. 1980. A quantitative succession
model for nine Montana forest communities. Environmental Management.
4(3): 227-240. [1336]
30. 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]
31. 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]
32. Lanner, Ronald M. 1980. Avian seed dispersal as a factor in the ecology
and evolution of limber and whitebark pines. In: Dancik, Bruce;
Higginbotham, Kenneth, eds. Proceedings, 6th North American forest
biology workshop; 1980 August 11-13; Edmonton, AB. Edmonton, AB:
University of Alberta: 15-48. [1404]
33. Lanner, Ronald M. 1982. Adaptations of whitebark pine for seed dispersal
by Clark's nutcracker. Canadian Journal of Forest Research. 12: 391-402.
[1403]
34. Leadem, Carole L. 1986. Seed dormancy in three Pinus species on the
Inland Mountain West. In: Shearer, Raymond D., compiler.
Proceedings--conifer tree seed in the Inland Mountain West symposium;
1985 August 5-6; Missoula, MT. Gen. Tech. Rep. INT-203. Ogden, UT: U.S.
Department of Agriculture, Forest Service, Intermountain Research
Station: 117-124. [1429]
35. Linhart, Yan B.; Tomback, Diana F. 1985. Seed dispersal by nutcrackers
causes multi-trunk growth form in pines. Oecologia. 67: 07-110. [1459]
36. Luckman, B. H.; Jozsa, L. A.; Murphy, P. J. 1984. Living
seven-hundred-year-old Picea engelmannii and Pinus alibcaulis in the
Canadian Rockies. Arctic and Alpine Research. 16(4): 419-422. [1481]
37. Lueck, Dennis. 1980. Ecology of Pinus albicaulis on Bachelor Butte,
Oregon. Corvallis, OR: Oregon State University; 1980. 90 p. M.S. thesis.
Abstract. [1483]
38. 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]
39. Miracle, Len. 1983. Record whitebark pine. Intermountain Reporter.
Ogden, UT: U.S. Department of Agriculture, Forest Service. May: 1-2.
[1662]
40. Pfister, Robert D.; Kovalchik, Bernard L.; Arno, Stephen F.; Presby,
Richard C. 1977. Forest habitat types of Montana. Gen. Tech. Rep.
INT-34. Ogden, UT: U.S. Department of Agriculture, Forest Service,
Intermountain Forest and Range Experiment Station. 174 p. [1878]
41. Pitel, J. A.; Wang, B. S. P. 1980. A preliminary study of dormancy in
Pinus albicaulis seeds. Canadian Forestry Service, Bi-monthly Research
Notes: January/February: 4-5. [1892]
42. Reed, Robert M. 1976. Coniferous forest habitat types of the Wind River
Mountains, Wyoming. American Midland Naturalist. 95(1): 159-173. [1950]
43. Steele, Robert; Cooper, Stephen V.; Ondov, David M.; [and others]. 1983.
Forest habitat types of eastern Idaho-western Wyoming. Gen. Tech. Rep.
INT-144. Ogden, UT: U.S. Department of Agriculture, Forest Service,
Intermountain Forest and Range Experiment Station. 122 p. [2230]
44. Tomback, Diana F. 1977. Foraging strategies of Clark's nutcracker.
Living Bird. 16: 123-161; 1977. [2349]
45. Tomback, Diana F. 1981. Notes on cones and vertebrate-mediated seed.
Madrono. 28(2): 91-94; 1981. [2348]
46. Tomback, Diana F. 1982. Dispersal of whitebark pine seeds by Clark's
nutcracker: a mutualism hypothesis. Journal of Animal Ecology. 51:
451-467. [2346]
47. Tomback, Diana F. 1986. Post-fire regeneration of krummholz whitebark
pine: A consequence of nutcracker seed caching. Madrono. 33(2): 100-110.
[2347]
48. Weaver, T.; Dale, D. 1974. Pinus albicaulis in central Montana:
Environment, vegetation and production. American Midland Naturalist.
92(1): 222-230; 1974. [2470]
Index
Related categories for Species: Pinus albicaulis
| Whitebark Pine
|
 |
