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Subalpine Meadows of Mount Rainier National Park,
Washington, U.S.A.
of Colorado
different locations in the subalpine zone of Mount
Rainier National Park, with substantial temporal
variation in regeneration of subalpine fir (Abies
lasiocarpa). Recruitment in subalpine meadows has
been continuous on the west side of Mount Rainier
since about 1930, but has occurred in short,
discrete periods on the east side. Variation in
snowpack from west to east on the mountain has a
substantial impact on climatic factors that limit
tree establishment. Warm, dry summer climate
facilitates tree establishment on the west side
where snowpacks are generally very high; cool, wet
summer climate enhances tree establishment on the
east side where snowpacks are lower. Density of
tree establishment is significantly greater in
heath-shrub (ericaceous) vegetation than in other
vegetation types. Within heath-shrub vegetation
types, tree establishment is highest at lower
elevations, on topographic convexities, and in
plant communities dominated by Phyllodoce
empetriformis. Survival of subalpine fir
seedlings during the first 3 yr after germination
is significantly greater in heath-shrub vegetation
than other vegetation types. If the climate
becomes warmer and drier during the next century,
continued rapid regeneration of trees can be
expected in subalpine meadows on the west side of
Mount Rainier National Park. This may result in
displacement of wildflower meadows that are a
attraction for park visitors. A better
understanding of climatic and environmental
limitations on tree establishment will assist
resource managers in developing sound management
strategies for subalpine ecosystems
vary among subalpine meadows within the park, 2)
whether factors limiting tree establishment vary
over space and time, and 3) what might we expect
the meadows to look like, with respect to trees,
in the future. Increasing understanding of
current tree establishment patterns and local
factors affecting tree establishment will improve
out predictions of the future composition of the
subalpine zone. This information can then be used
to develop scientifically-based resource
management decisions in response to potential
climatic change and increasing demands for human
use of park resources.
subalpine meadows within Mount Rainier National
Park (Fig. 2). The park is located on the western
slope of the Cascade Range, 100 km southeast of
the Seattle -Tacoma metropolitan area. It
encompasses 95,389 ha and extends from low
elevation, old growth forest (530 m) through
subalpine and alpine communities to the summit of
Mount Rainier at 4400 m. Climate is temperate
maritime with cool, wet winters and mild, dry
summers. Most of the annual precipitation falls as
snow between October and May. Limited climatic
data indicate that precipitation generally is
higher on the westside of the park, and increases
with elevation up to about 3000 m. Subalpine
parkland covers approximately 23% of the park.
Meadow vegetation of this zone can be described by
five broad vegetation types (Henderson, 1974). (1)
heath-shrub types dominated by one or two
ericaceous species, including Phyllodoce
empetriformis, P. glanduliflora, Cassiope
mertensiana, and Vaccinium deliciosum, (2) lush
herbaceous vegetation dominated by tall perennials
including Valeriana sitchensis, Lupinus
latifolius, and Veratrum viride, (3) low
herbaceous vegetation dominated by Potentilla
flabellifolia and Antennaria lanata, often with
lesser amounts of Carex nigricans, (4) wet sedge
types in low, wet areas dominated by C. nigricans,
C. spectabilis, Aster alpigenus, and Antennaria
lanata, and (5) dry grass vegetation found on
well-drained sites common on the east side of the
park, dominated by Festuca viridula and Lupinus
latifolius. The dominant tree species in the
subalpine zone are Abies lasiocarpa, Tsuga
mertensiana, and Chamaecyparis nootkatensis. Pinus
albicaulis and Picea engelmannii are present on
drier sites on the eastside of the park. Human
activities have had limited influence on the
subalpine ecosystem of Mount Rainier National
Park. Grazing by sheep and cattle occurred in
several areas on the eastside of the park in the
early 1890s prior to the establishment of the
Pacific Forest Reserve in 1893. Once the reserve
was created, it became illegal for stock to graze
on federal lands, but it still continued to some
extent due to lack of enforcement (Martinson,
1966). After the establishment of the park in
1899, grazing was limited to cattle in two areas
during 1917-1919, use of milk cows in Paradise
(1905-1910), and sheep for 2 yr (1931-1932) on the
east boundary (McIntyre, 1952). Plant removal and
soil erosion was so great from these practices
that a survey completed in 1944 recommended that
no additional grazing permits be issued in the
event of a war emergency (Stagner, 1944). More
recent influences on subalpine ecosystems include
tree removals from the Paradise region of the
park, as well as increased recreational use of
this area (Rochefort and Gibbons, 1993; Rochefort
and Peterson, 1993).
five subalpine meadows of Mount Rainier National
Park in 1991. Spray Park, Paradise, Sunrise, Grand
Park, and Burnt Park (Fig. 2). Sites were selected
based on geographic location and weather patterns
within the park. Paradise and Sunrise were
selected because they were surveyed in the 1960s
and we wanted to see if more recent periods of
establishment had occurred since that time
(Franklin et al, 1966; Franklin et al., 1971). The
three additional meadows were chosen to expand the
spatial/geographic scope of the Franklin study and
to describe variation in patterns of establishment
within the park. We concentrated on west versus
east (rainshadow) sides of the park because
weather patterns (precipitation and
temperature)and vegetation vary significantly
between the two sides. Two to five strip transects
were established randomly within the predominant
vegetation types of each meadow (Table i). Strip
transects were used to estimate tree density
because visual observation indicated that tree
density decreased with distance from clumps of
large trees (trees approximately 20--30 m tall and
200 yr old). Transects were established in areas
where there were no visible signs of fire (fire
scars or charred trunks) or human use (tree
cutting or bare ground) and in sites that had no
recorded history of grazing, tree cutting or
development. Strip transects were 3 m wide and
extended 60 m from the outer edge of mature tree
clumps. Transects were divided into 3 m x 5 m
blocks for ease in tallying trees. Within each
block, tree species were identified, tree height
and basal diameter were recorded, and every other
tree was sampled for age determination. Seedlings
and sap-lings (<5 cm basal diameter) were
collected for age determination by cutting below
the root collar. Seedlings were aged by counting
terminal bud scars, and saplings were aged by
counting rings on sanded basal disks; both counts
were conducted under a dissecting microscope.
Seedlings were generally trees with a basal
diameter less than 2 mm, height less than 12 cm,
and less than 15 yr old. Larger trees were sampled
by collecting a core close to the root collar.
Cores were finely sanded and annual rings counted
under a dissecting microscope. Only tree cores
that included the center of the tree were included
for aging. All trees were aged by two individuals,
and discrepancies were resolved by additional
counts. Analysis of our 1991 data revealed that
more trees established in heath-shrub vegetation
types than other vegetation types surveyed.
Therefore, we wanted to look more closely at
seedling survival patterns among vegetation types.
Abies lasiocarpa seedlings germinating in 1992
were monitored for 3 yr to determine if survival
rates of seedlings germinating in heath-shrub
vegetation (Phyllodoce empetriformis, P.
glanduliflora, Cassiope mertensiana) were
significantly different from those germinating in
other vegetation types. Eleven paired sites were
established at the Paradise study site. In each
site, 20 to 50 seed-lings were tagged and
monitored for survival from July 1992 until
October 1994. Surveys of heath-shrub communities
were conducted in 1992 to determine if tree
establishment within them is random with respect
to landscape position and other environmental
parameters. Forty-five circular plots (12.6 m2)
were randomly established in heath-shrub
vegetation at the Paradise study site (1640 to
1920 m elevation). Random sample points were
selected by placing a grid over a vegetation map
of the Paradise meadow (approximately 390 ha). A
random number generator was used to select
coordinates for potential plots; all random
locations falling within mapped ericaceous
vegetation types were sampled. Within each plot,
all trees were counted, identified to species, and
height measured. Dominant plant association,
slope, aspect, topography (convex, flat, concave),
and landscape position (ridge, midslope, bench,
valley bottom) were recorded. Vegetation was
classified into one of four plant community types
based on the dominant species: (1) Phyllodoce
empetriformis / Vaccinium deliciosum, (2) P.
empetriformis / Lupinus latifolius, (3) P.
empetriformis / Cassiope mertensiana, (4) P.
empetriformis.
data were summarized by 5-yr intervals for the
1930-1990 time period. Five-year periods were used
to examine these relationships because successful
tree establishment depends on climatic factors
during at least 3 yr after germination (Cui and
Smith, 1991; Jakubos and Romme, 1993). In
addition, we recognized that germination dates
could be miscalculated by several years due to
missing rings and the difficulty recognizing
terminal bud scale scars (Henderson, 1974; Little
et al., 1994; Miller, 1995). Data analysis focused
on the 1930-1990 period, for which climatic data
was nearly complete. Stepwise multiple
regression was used to examine the relationship
between climate and tree establishment. Numbers of
trees established were summed for each period, and
climatic variables were averaged. Counts of tree
establishment were transformed using a square root
transformation to stabilize the variance. Climatic
variables included monthly average temperature
(May through October) and total precipitation (May
through September) (Paradise Ranger Station, Mount
Rainier National Park database), and monthly
Palmer Drought Severity Index (PDSI) (state of
Washington Division 4 data from the National
Climatic Data Center database). Spring PDSI was
the average value for May-June, while summer PDSI
was the average value for July-August. The effect
of snowpack on tree establishment was investigated
by using snow depth at Paradise on 15 May
(Paradise Ranger Station, Mount Rainier National
Park database). Data missing from these records
were previously estimated by Little (1992) using
standard techniques (Paulhus and Kohler, 1952;
MacDonald, 1957). Selection of climate variables
for analysis was based on the assumption that seed
germination and survival of seedlings are most
influenced by growing season length, temperature,
and precipitation. Relationships with winter
weather were not examined because snow generally
covers seed-lings from November through May.
Tree density by vegetation type was compared using
data from all established strip transects. Blocks
along each transect were numbered from 1 to 12
indicating their distance from mature trees. The
number of blocks sampled in each vegetation type
was then tallied by distance class. A chi-square
analysis was performed to determine if the
sampling distribution among vegetation types was
homogeneous with respect to distance from a tree
clump (seed source). Tree density for each 3 m X 5
m subplot was tallied and categorized by
vegetation type. Data were transformed using the
log (x f I) transformation (Zar, 1984) because the
variances were positively skewed. Analysis of
variance was then used to compare tree densities
within four vegetation types: heath-shrub, lush
herbaceous, low herbaceous, and dry grass. The wet
sedge vegetation type did not have a large enough
sample size to include in the analysis. Following
rejection of the null hypothesis of equal mean
tree densities among the four vegetation types,
multiple comparisons were made using the Tukey HSD
test (p = 0.05). Seedling survival was analyzed
by performing a paired t-test comparing percent
survival of seedlings inside and outside heather
at annual intervals. Mean differences between 11
paired plots were used for analysis in 1992 and
1993, but only 9 pairs were used in 1994 due to
removal of tags by animals that summer
Discriminant analysis was used to identify
important factors associated with successful tree
establishment in heather communities. Predictor
variables were categorized using a binary
dependent variable (i.e., "O" if no A. lasiocarpa
were observed in the plots, "1" if one or more A.
lasiocarpa were observed). Prediction potential
of the classification criteria was evaluated using
a cross-validation (jackknife) procedure (SAS,
1988). Discriminant coefficients were examined to
identify the direction of the relationship between
predictor variables and the dependent variable.
The class means were then compared with an ANOVA
to identify significant differences.
Subalpine Meadows of Mount Rainier National Park,
Washington, U.S.A.
of Colorado
Washington
and west side meadows
meters
to a more complete description of the vegetation
type within the text of the article.
site along with the number that were aged for the
study
themselves on each of the five study sites
intervals
each site for a specific 5 year interval
associated with tree establishment.
transformed number of trees established and
climatic variables
and west side meadows
regression analysis
coefficient
each site
within four vegetation types
part of the study
type
are statistically different at the p = .001 level
communities and non-heath communities
inventoried
counts on first date of inventory
Subalpine Meadows of Mount Rainier National Park,
Washington, U.S.A.
suitability of this information for a particular
purpose. Original data elements were compiled
from various sources. This information may be
updated, corrected, or otherwise modified without
notification. For additional information about
this data contact the author(s).
article.
Cascadia Field Station Box 352100
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