Associated Forest Cover
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Brewer spruce grows with a wide variety of associated plants and
vegetation types in the montane and subalpine forests of the Klamath
region. Although it grows throughout the region, its range is one of
local, disjunct populations of various sizes. In some areas, it is an
occasional climax tree species in mixed stands dominated by California red
fir (Abies magnifica), white fir (A. concolor), or
mountain hemlock (Tsuga mertensiana). In other areas, it grows in
small, pure, dense stands on north-, east-, and west-facing slopes. It is
also found as individuals invading seral pine stands and montane
chaparral. In the Siskiyou Mountains' Brewer spruce seedlings and saplings
are found in montane chaparral on all aspects. It is associated with
Sadler oak Quercus sadleriana), huckleberry oak (Q.
vaccinifolia), and greenleaf manzanita (Arctostaphylos patula).
Some of the best-developed stands of Brewer spruce are located on
moderate east- and west-facing slopes with, deep soil. These forests in
the California red fir/Sadler oak habitat have a 70 to 80 percent canopy
cover. Density of trees over 10 cm (4 in) in d.b.h. is 125 to 320 Brewer
spruce per hectare (50 to 130/acre), 30 to 95 white fir per hectare (12 to
39/acre), 10 to 70 Douglas-fir (Pseudotsuga menziesii) per hectare
(4 to 28/acre), 0 to 10 sugar pine (Pinus lambertiana) per hectare
(0 to 4/acre), and 5 to 60 California red fir per hectare (2 to 24/acre).
The total basal area ranges from 35 to 54 m²/ha (153 to 235 ft²/acre),.
Dense reproduction is present in the tolerant conifers: Brewer spruce,
California red fir, and white fir; Brewer spruce has about one-third the
total number of seedlings under 180 cm (70 in) in height.
Brewer spruce is an element of the following vegetation habitat types of
the Klamath region (2,12,13,14). Plants of major importance are listed for
each type.
Abies concolor zone, Siskiyou Mountains-
Abies concolor/Vaccinium membranaceum (white fir/thinleaf
huckleberry) habitat type. Brewer spruce is a minor climax species. Other
shrubs: Sadler oak.
Abies concolor/Pachistima myrsinites (white fir/Oregon boxwood)
habitat type. Brewer spruce is often a codominant climax species. Other
trees: Douglas-fir and sugar pine. Other shrubs: Sadler oak, Oregongrape
(Berberis nervosa). Other herbs: western prince's-pine (Chimaphila
umbellata), rattlesnake plantain (Goodyera oblongifolia), and
vanillaleaf (Achlys triphylla).
Chamaecyparis lawsoniana/Rhododendron occidentale (Port-Orford-cedar/western
azalea) habitat type. Brewer spruce is often a minor climax species. Other
trees: western white pine (Pinus monticola). Other shrubs: Sadler
oak.
Abies concolor zone, central Klamath region-
Abies concolor/Chimaphila umbellata (white fir/western
prince's-pine) habitat type. Brewer spruce occasionally occurs as a minor
climax species. Other trees: Douglas-fir, sugar pine, ponderosa pine (Pinus
ponderosa), and incense-cedar (Libocedrus decurrens). Other
shrubs: western hazel (Corylus cornuta), wood rose (Rosa
gymnocarpa), and snowberry (Symphoricarpos hesperius).
Abies magnifica zone, Siskiyou Mountains-
Abies magnifica/Quercus sadleriana (California red fir/Sadler
oak) habitat type. Brewer spruce often occurs as a minor climax species.
Other trees: white fir, Douglas-fir, western white pine, and sugar pine.
Other shrubs: thinleaf huckleberry.
Abies magnifica/Arctostaphylos nevadensis (California red
fir/pine mat manzanita) habitat type. Brewer spruce occasionally occurs as
a codominant climax species. Other trees: western white pine. Other
shrubs: Sadler Oak and greenleaf manzanita.
Picea breweriana/Quercus vaccinifolia (Brewer spruce/huckleberry
oak) habitat type. Brewer spruce occurs as a codominant climax species.
Other trees: western white pine, California red fir, Douglas-fir, and
incense-cedar. Other shrubs: greenleaf manzanita, pine mat manzanita, and
Sadler oak.
Picea breweriana/Quercus sadleriana (Brewer spruce/Sadler oak)
habitat type. Brewer spruce occurs as the dominant climax species. Other
trees: western white pine and white fir. Other shrubs: huckleberry oak and
thinleaf huckleberry.
Abies magnifica zone, central and eastern Klamath region-
Abies magnifica/Leucothoe davisiae (California red fir/mountain
laurel) habitat type. Brewer spruce is an occasional minor climax species.
Other trees: white fir, lodgepole pine (Pinus contorta), western
white pine, and mountain hemlock (Tsuga mertensiana). Other
shrubs: swamp current (Ribes lacustre), mountain alder (Alnus
tenuifolia), and mountain ash (Sorbus californica). Other
herbs: groundsel (Senecio triangularis), queenscup (Clintonia
uniflora), starflower (Trientalis latifolia), trillium (Trillium
ovatum), and false Solomon's seal (Smilacina stellata).
Abies magnifica/Linnaea borealis (California red fir/twinflower)
habitat type. Brewer spruce is an occasional codominant climax species in
open forest stands. Other trees: Douglas-fir, white fir, western white
pine, mountain hemlock, sugar pine, ponderosa pine, western yew (Taxus
brevifolia), incense-cedar, Engelmann spruce (Picea engelmannii),
lodgepole pine, and subalpine fir (Abies lasiocarpa). Other
shrubs: Sadler oak, wood rose, and snowberry. Other herbs: queenscup.
Abies magnifica/Quercus vaccinifolia (California red
fir/huckleberry oak) habitat type. Brewer spruce is an occasional minor
climax species. Other trees: white fir, Douglas-fir, sugar pine, lodgepole
pine, and western white pine. Other shrubs: greenleaf manzanita, pine mat
manzanita, and bush chinkapin (Castanopsis sempervirens).
Tsuga mertensiana zone, Siskiyou Mountains-
Tsuga mertensiana/Vaccinium membranaceum (mountain
hemlock/thinleaf huckleberry) habitat type. Brewer spruce is a codominant
climax species. Other trees: California red fir, western white pine, and
Alaska-cedar (Chamaecyparis nootkatensis).
Tsuga mertensiana zone, central and eastern Klamath region-
Tsuga mertensiana/Phyllodoce empetriformis (western
hemlock/heather) habitat type. Brewer spruce is an occasional codominant
climax species. Other trees: California red fir, white fir, western white
pine, and lodgepole pine. Other shrubs: Labrador-tea (Ledum
glandulosum).
Tsuga mertensiana/Quercus vaccinifolia (western
hemlock/huckleberry oak) habitat type. Brewer spruce is of minor
importance as a climax species. Other trees: California red fir and
western white pine. Other shrubs: pine mat manzanita, bush chinkapin, and
greenleaf manzanita.
Brewer spruce is a minor component in three forest cover types (4):
Mountain Hemlock (Society of American Foresters Type 205), Red Fir (Type
207), and California Mixed Subalpine (Type 256).
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Climate
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The range of Brewer spruce is primarily influenced by the maritime
climate of the Pacific Ocean: cool, wet winters and warm, relatively dry
summers. The climate changes drastically, however, from west to east
across the range of Brewer spruce, which is 113 km (70 mi). On the most
westerly ridge occupied by Brewer spruce, average annual precipitation is
more than 2800 mm (110 in); farther east, it is less than 1000 mm (39 in).
Most of the precipitation falls as winter rain or snow; less than 5
percent occurs in the summer. The amount of -snowfall and accumulated
snowpack varies greatly from year to year and geographically across the
range. A few stands of Brewer spruce in valleys receive no snow some
years, whereas other stands accumulate up to 4 to 5 m (13 to 16 ft) (5).
Humidity is high in the western part of the range and relatively low in
the eastern part. Summer fog is common along the western-most ridges and
valleys. Summer thunderstorms are common in the eastern portion of the
range. Temperatures also vary widely. In the western portion, the mean
temperature in January is 5° C (41° F); in July, 11° C (52°
F). On the eastern edge of the range, the mean temperature is -1° C
(30° F) in January and 20° C (68° F) in July.
The varied climate indicates that Brewer spruce has an ecological
amplitude that should enable it to obtain a wider and more contiguous
distribution. Its sensitivity to fire seems to have restricted its range
(13).
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Damaging Agents
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The shallow root system of Brewer spruce makes
it more susceptible than its associates to windthrow. In some areas, the
high incidence of root rot (Heterobasidion annosum) further lowers
its resistance to wind. Thin bark and long weeping branches make Brewer
spruce susceptible to fire. Areas predictably occupied by Brewer spruce
are limited to fire-resistant open forests on north-facing slopes or rocky
ridges. A series of major forest fires in 1987 burned throughout the range
of Brewer spruce. Most of the small, pure stands on north slopes were not
damaged. In the more prevalent mixed stands, low-intensity ground fires
killed the Brewer spruce and white fir, but the thick-barked Douglas-fir,
pines, and California red fir survived. The recovery of Brewer spruce may
take decades or centuries because of the extent of these fires. The
increased potential of forest fires and inability of Brewer spruce
seedlings to tolerate high moisture stress may result in its rapid
extinction. If global warming occurs (11), it could threaten the existence
of localized tree species such as Brewer spruce.
As a small tree, Brewer spruce has enough flexibility to bend under the
weight of heavy snow. It develops a pistol butt as the tree matures.
Comparatively little damage from insects or fungi has been recorded for
Brewer spruce (3). The Cooley spruce gall adelgid (Adelges cooleyi) is
common but does little damage (6). Seed chalcids (Megastigmus spp.)
have been observed in mature seeds. In some areas, 36 percent of the
Brewer spruce was parasitized by the dwarfmistletoe Arceuthobium
campylopodum (8). Brewer spruce is intolerant of industrial fumes.
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Flowering and Fruiting
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Brewer spruce is monoecious. Strobilus
buds form in early summer and develop in the fall. The dark purple male
strobili are borne the following spring in axils of needles of the
previous year's shoots, on branches throughout the tree. The
pollen-bearing strobili are from 19 to 32 mm, (0.75 to 1.25 in) long and
about 13 mm (0.5 in) in diameter. Pollen is shed in early summer; the male
strobili dry and fall soon after pollen is shed. Female strobili are borne
at the tips of primary branches in the upper two-thirds of the crown. They
develop into erect, dark green, cylindrical conelets that are 38 mm (1.5
in) long and 13 mm (0.5 in) thick when receptive. The female conelets are
apparently receptive at the time pollen is shed. The location of female
and male flowers throughout the tree, concurrent with timing of strobilus
development, apparently encourages selfing of Brewer spruce located singly
or in small, isolated stands. After pollination, the conelets turn down
and mature the same season into dark brown cones 8 to 15 cm (3 to 6 in)
long. The fruit matures from September to October; dissemination
immediately follows.
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Genetics
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Little if any variability has been observed throughout the restricted
range of Brewer spruce. In the eastern Klamath area, Brewer spruce grows
adjacent to Engelmann spruce without hybridization (12).
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Growth and Yield
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After the seedling stage height growth of
Brewer spruce is slower than its common associates, Douglas-fir, white
fir, red fir, and western white pine. In several stands in the Siskiyou
Mountains, annual height growth of sapling and pole-size Brewer spruce
averaged 0.15 m (0.5 ft).
Most mature stands of Brewer spruce consist of a wide range of ages and
sizes. Numerous stands contain Brewer spruce up to 117 cm (46 in) in
diameter, the largest 125 cm (49.3 in) in d.b.h. and 48.8 m (160 ft) in
height. The biggest Brewer spruce listed by the American Forestry
Association (1) has a circumference of 4.17 m (13.67 ft) at 1.37 m (4.5
ft) above the ground and is 51.8 m (170 ft) tall; it is located in the
Siskiyou National Forest in southwest Oregon. The general structure of a
mixed species, all-aged stand (Sadler oak habitat type) is shown in table
1.
Table 1- Stand structure of mixed species, all-aged
Brewer spruce stands (Sadler oak habitat type)
Diameter class
Brewer spruce
White
fir
Red
fir
Douglas-
fir
Western white pine
Sugar
pine
cm
trees/ha
3 to 29
430
506
82
69
-
-
30 to 59
114
32
27
2
7
-
60 to 89
17
15
15
-
2
-
90 to 119
2
-
7
-
-
-
120+
-
-
2
2
-
2
in
trees/acre
1 to 11
174
205
33
28
-
-
12 to 23
46
13
11
1
3
-
24 to 35
7
6
6
-
1
-
35 to 47
1
-
3
-
-
-
47+
-
-
1
1
-
1
Little volume or yield information is available for these Brewer spruce
stands. The total basal area of the few stands sampled averages 47 m²/ha
(205 ft²/acre), with a current annual increment of 2 m²/ha (9 ft²/acre)
(15).
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Reaction to Competition
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Brewer spruce is tolerant of shade at
all ages and is very competitive. In the Abies magnifica zone on
mesic to xeric sites, it is more competitive than mountain hemlock,
Port-Orford-cedar, white fir, Douglas-fir, Alaska-cedar, incense-cedar,
sugar pine, western white pine, ponderosa pine, lodgepole pine, and
Jeffrey pine. California red fir is considered slightly more competitive
on mesic sites because of its faster height growth and longer lifespan.
Brewer spruce is well adapted to growth at cool temperatures during the
growing season. Its light compensation point is less than 2 percent of
full sunlight. Brewer spruce can withstand considerable soil drought but
is extremely sensitive to high evaporation demands. The stomata close
under high evaporation, halting photosynthesis (17).
Brewer spruce is best managed on mesic sites characterized by the
presence of Sadler oak. It grows best in mixed-species stands with
uneven-aged management.
Brewer spruce can be planted under montane chaparral dominated by Sadler
oak, huckleberry oak, and greenleaf manzanita. It has the ability to grow
well under competition for soil moisture and light.
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Rooting Habit
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Brewer spruce has a shallow root system on all
soils; however, on deeper soils, a few vertical roots may extend several
meters in depth.
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Seed Production and Dissemination
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Seed production starts when
the trees are from 20 to 30 years old. Actual production has not been
determined. Observations indicate that mature Brewer spruce trees are fair
seed producers; crops occur at 2-year intervals, and some trees produce
cones each year.
The seeds are 3 to 5 mm (0.1 to 0.2 in) long; the wing is four times as
long as the seed. The seeds are the largest of the North American spruces:
134,500/kg (61,000/lb), with a range of 112,500 to 163,000/kg (51,000 to
74,000/lb). The relatively large wing aids dissemination of the seed by
the wind. Cones and seeds do not appear to be a preferred food for
rodents.
Seeds may be stored for 5 to 17 years in sealed containers at low
temperatures, 1° to 3° C (33° to 38° F), at a moisture
content of 4 to 8 percent. A cold, moist stratification of 30 days
increases germination. Germination of sound seed ranges from 50 to 96
percent; the reported average is 88 percent (15).
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Seedling Development
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The germination of Brewer spruce is
epigeal, seedlings becoming established on all types of natural seedbeds:
decaying logs, forest humus, loose soil from upturned roots, and leaf
litter under brush fields. Throughout the range of Brewer spruce, natural
regeneration is abundant under dense Brewer spruce-white fir stands. These
stands contain an average of 1,360 Brewer spruce and 3,460 white fir
seedlings per hectare (550 and 1,400/acre) less than 15 cm (6 in) in
height. Brewer spruce seedlings cannot survive strong sunlight. The
shallow, slow-growing root system causes the seedlings to be susceptible
to the high moisture stress and temperatures of exposed sites. Brewer
spruce seedlings are usually lacking in clearcuts, even when these are
adjacent to stands containing large cone bearing trees.
Seedlings are small, with four to seven cotyledons Initial growth is
slow; the epicotyl height growth is less than 6 mm (0.24 in) the first
season.
Subsequent growth of seedlings is slow but quite variable. Under dense
stands, the age of seedling 1.37 m (4.5 ft) tall ranges from 27 to 82
years Brewer spruce seedlings growing in south-facing montane chaparral
were from 25 to 40 years old when they were 1.37 m (4.5 ft) tall. Small
Brewer spruce survive overstory removal.
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Soils and Topography
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Brewer spruce grows on a wide variety of geologic parent materials and
soils. It is found on soils developed from sedimentary, granitic,
serpentine, and metavolcanic rocks. Most of the large stands of Brewer
spruce are found on shallow, rocky, undeveloped soils of the order
Entisols; a few stands are on deeper, well-developed soils. Throughout its
range, Brewer spruce is never found in areas where the soils are saturated
during the growing season. It appears to be generally restricted from the
more fertile soils by competition from true firs (Abies spp.).
Brewer spruce is more abundant on less fertile soils.
Brewer spruce is found on most topographic locations-ridgetops, north-
and south-facing slopes, benches, and valley bottoms. The only habitat
restriction is boggy or wet areas. The apparently preferred location is
the steep, north-facing slopes where the largest stands are located. In
the western Siskiyou Mountains, these locations are north slopes near the
tops of the ridges, but in the eastern Salmon Mountains, the largest
stands are on middle, north-facing slopes. Brewer spruce is found from
elevations of 700 to 2100 in (2,300 to 6,900 ft) in the western Siskiyou
Mountains and from 1370 to 2290 in (4,500 to 7,500 ft) in the eastern
Klamath region.
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Special Uses
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The wood of Brewer spruce has no special uses. Harvested trees are
normally mixed with other species and utilized as low grade lumber. In
Europe, it has been considered one of the most popular of all ornamental
conifers (10).
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Vegetative Reproduction
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Layering has no been observed in
natural stands of Brewer spruce Artificial propagation is best from seed
(10).
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Distribution
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Brewer spruce is found only in the mountains of northwestern California
and southwestern Oregon near the Pacific coast between latitudes 40°
50' N. and 42° 40' N. The best developed stands are on the high
ridges of the western Siskiyou Mountains in California and Oregon. Other
concentrations are found on high ridges and in upper valleys of the
Marble, Salmon, and Trinity Mountains of California (7). Throughout the
rest of the range, Brewer spruce grows as a single tree and as scattered
small populations in valleys and on ridgetops (9,12,16).
- The native range of Brewer spruce
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Brief Summary
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Pinaceae -- Pine family
Dale Thornburgh
Brewer spruce (Picea breweriana), named for its first collector,
is often considered "mysterious" because this attractive conifer
is found on seldom-visited high mountain ridges and steep north slopes.
Its other common name, weeping spruce, is derived from the distinctive
feature of many rope-like branchlets that hang in a fringe from all but
the topmost slender horizontal limbs. This branching habit results in many
knots in the wood, which has little commercial importance.
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