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Rainier National Park
David L. Peterson, Temporal and Spatial
Distribution of Trees in Subalpine Meadows of
Mount Rainier National Park, Washington, U.S.A.,
Arctic and Alpine Research, Vol. 28, No. 1, 1996,
pp. 52-59
persistent and widespread species in mountainous
areas of the Pacific Northwest. Although these
species are quite common, little information is
available on the population dynamics or
reproductive biology of the species. In National
Parks, increasing recreational demands escalates
the potential for damage to these populations.
This study describes genetic and morphologic
diversity of heather populations in mount Rainier
National Park, assesses the potential influence of
human use, and utilizes this information to
propose strategies for long-term protection of
heather populations. Allozyme analysis was used
to estimate genetic diversity of P. empetriformis
and P. glanduliflora. Study sites were
established at high and low elevation in three
locations within Mount Rainier National Park.
Analysis indicated high levels of genetic
diversity within populations and significant
differences in allele frequencies among
populations and study areas. Heather populations
are composed of multiple clones with high ratios
of local to widespread genotypes. Morphologic
variation was surveyed through measurements of
field and greenhouse populations. Significant
differences existed among field populations for
the four characters measured: annual stem
extension, plant height, leaf width and leaf
length. Greenhouse populations exhibited
significant differences among populations for leaf
width (P. empetriformis) and leaf length (P.
glanduliflora). The role of tree establishment in
long-term heather survival was examined.
Significantly more trees established in heather
populations than in plant communities dominated by
other species. Survival of Abies lasiocarpa
seedlings was significantly higher in heather
populations than populations of other species.
Low-elevation heather populations growing on
topographic convexities often provide safe-sites
for the establishment of tree seedlings. A simple
simulation model was developed to estimate the
potential impact of human sue on heather
populations. Number of genotypes, allele
frequencies, observed heterozygosity, number of
live plants, and heather crown cover were
calculated before and after simulated Human use.
No significant differences were found in allele
frequency or heterozygosity levels even after
severe impacts were modeled. Genotypic diversity
is not sensitive indicator of human use. Numbers
of genotypes only showed significant changes when
impacts were very large or when restoration
programs were initiated.
diversity of heather populations in Mount Rainier
National Park, assesses the potential influence
of human use, and utilizes this information to
propose strategies for long-term protection of
heather populations.
Populations: STUDY SITE: Study sites were
established in three areas within Mount Rainier
National Park: Paradise, Spray Park, and Panhandle
Gap (Figure 2.2, Table 2.1). Two to four
populations were sampled within each area. Paired
populations at high and low elevations were
identified within each study area in order to
sample the range of genetic diversity. Four
populations of each species (Phyllodoce
empetriformis and P. glanduliflora) were sampled
at Spray Park and Paradise, two high elevation and
two low elevation populations. At Panhandle Gap,
four populations of P. empetriformis (two high and
two low) and two populations of P. glanduliflora
(two high populations, no low populations) were
sampled. Populations were separated by greater
than 100 meters. Study plots were established
within each population; plots are 400 meters
squared (20 m x 20 m) and subdivided into 4
quarters. Fifty plants (12-13 per quarter) were
randomly selected within the large plot for
genetic analysis.
Populations: ELECTROPHORESIS: Stems with first and
second year leaves were collected for
electrophoretic analysis. Leaves were placed in
plastic bags and kept moist and cold in a
refrigerator (4"C) until they were prepared for
electrophoresis. Leaf tissues were ground with a
mortar and pestle under liquid nitrogen (Mitton et
al., 1979), and the frozen leaf powder was mixed
with a tris-HCL grinding buffer PVP solution
(Soltis et al., 1983). The homogenate was
immediately transferred into microtiter trays and
stored in an ultra-low freezer (-70C). Homogenates
were thawed, applied to paper wicks, and inserted
into 12.5% starch gels. Plant material was
initially tested for activity on 26 enzyme systems
(Appendix A). Only five loci for each species
exhibited consistent, scorable resolution. Five
putative loci were resolved on three systems for
Phyllodoce glanduliflora. Phosphoglucose isomerase
(PGI) (two loci) and uridine diphosphoglucose
pyrophosphorylase (UGP) were resolved on a
continuous morpholine citrate system, Ph 8.1
(system E in Conkle et al. 1982). Shikimate
dehydrogenase (SKD) was run on morpholine citrate,
Ph 6.1 (system D in Conkle et at. 1982).
Phosphoglucomutase (PGM) was resolved on a
discontinuous histidine citrate system, Ph 7.0
(Werth, 1985). Five putative loci were resolved
on two systems for P. empetrifonnis.
Phosphoglucose isomerase (PGI) (two·loci) was
resolved on a continuous morpholine citrate
system, Ph 8.1 (system E in Conkle et al. 1982).
Uridine diphosphoglucose pyrophosphorylase (UGP-1,
UGP-2) and phosphoglucomutase (PGM) were scored on
a discontinuous histidine-Hcl, Ph 6.5 (system 11
in Soitis et al. 1983). Only loci polymorphic at
the 95% level were utilized for genetic analysis;
PGI-1 was monomorphic for both species and
therefore was not utilized.
Populations: MORPHOLOGIC DIVERSITY: Morphologic
diversity was surveyed on field populations and
greenhouse plants grown from cuttings of field
plants. Four morphologic characters were monitored
on 25 plants in each of the field populations:
leaf width, leaf length, plant height, and annual
stem growth (i.e. extensional growth as measured
between bud scars). Plant height was measured at
the tallest portion of the plant. Annual stem
growth, leaf width, and leaf length were measured
with a micrometer; values for each character are
the average of three measurements for each plant.
Leaf width was measured at the widest portion of
each leaf. Greenhouse plants were propagated from
cuttings of plants from the Paradise field
populations. Cuttings were collected in the fall
of 1992 and rooted cuttings were transplanted into
10 cm diameter pots in the spring of 1993 and
arranged in a completely randomized design. Plants
were moved from the greenhouse to a shadehouse in
June 1994. Leaf width and length were measured in
1995 on leaves formed in 1994.
Populations: DATA ANALYSIS: Genetic data were
analyzed using Biosys-l, version 1.7 (Swofford and
Selander, 1q89) and Genestat (Lewis and Whitkus,
1989). Genetic variability was characterized by
calculating allele frequencies, observed
heterozygosity (Ho direct count), expected
heterozygosity (He Nei's 1978 unbiased estimate),
and Hamrick and Godt's Ht (1990; Nei and Chesser,
1983). Diversity within and among populations was
examined by calculating Wright's F-statistics
(1943, 1951) and Nei's genetic identity (1978).
Phenograms were produced to visualize similarities
between populations by conducting a cluster
analysis using UPGMA and Nei's genetic identity.
Correlation analysis was conducted between pairs
of genetic identity values and geographic
distances and elevation differences. Diversity
among populations and study areas was examined by
using a X2 contingency analysis to analyze
differences in allele frequencies (Snedecor and
Irwin, 1933; Workman and Niswander, 1970). In
addition, because P. glanduliflora and P.
empetriformis propagate vegetatively, genetic
diversity was examined by calculating the number
of genotypes, proportion of clones
distinguishable, and the number of local and
widespread genotypes per population (Ellstrand and
Roose, 1987). The number of genotypes was
calculated by using only those individuals for
which all four loci had been scored. The
proportion distinguishable was calculated by
dividing the number of genotypes by the sample
size. Local and widespread genotypes were
calculated within study areas and across all
populations. Local genotypes were defined as
types found in only one population within the
study area. Widespread genotypes were defined as
types found in 75% of the populations within the
study area or across all populations. Morphologic
characters were analyzed with a one-way analysis
of variance. If heteroscedasticity was
encountered, Kruskal-Wallis procedures were
followed. When the null hypothesis was rejected,
the Tukey HSD test was applied (p= 0.05). Relative
percents of among and within population variances
were calculated for each character (Sokal and
Rohlf, 1981).
Study sites were located in five 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).
METHODS: Study sites were established in 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 the 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 in order to
expand the 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.
ANALYSIS: Tree establishment and climatic 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.
MODELING AND ANALYSIS: The effect of human use on
heather populations was examined by simulating
damage within mapped, undamaged populations. I had
initially hoped to sample paired sites' of damaged
and undamaged populations, but damaged sites
usually had different environmental
characteristics than undamaged sites. Damaged
populations were frequently found on areas that
melted out earlier than undamaged sites, thus
attracting visitor use. For this reason,
simulation models provide a better appraisal of
short- term changes caused by human use. Two
populations in the Paradise study area were
selected: PG8 and PE1. Within each study plot (20
x 20 m), one quarter of the site was designated
for intense monitoring. All plants within this
block were mapped (crown cover), measured for
height, and sampled for genetic analysis. Damage
was simulated by randomly drawing social trails of
several widths (15, 56, and 102 cm) in the mapped
plot and removing all plants within the boundaries
of the plot. Simulations were conducted with a
dBase program that generated random numbers for
social trail location. Rules for the program
required straight social trails and no overlapping
of trails. For each run of the model, 1 to 20
socials trail were placed in the plot. Widths of
impacts were selected based on typical widths of
human impacts in Mount Rainier National Park
(Rochefort, 1989). Simulations allowed the
placement of 20 social trails 15 cm wide within
the study plot, but only 10 social trails 56 cm
wide and 6 or 7 trails 102 cm wide. The impact of
human use was then evaluated by calculating allele
frequencies, observed heterozygosity (H,), the
number of genotypes, the number of live plants
remaining, and heather crown cover.
Pre-disturbance and post-disturbance estimates for
each index were compared after one run of the
model for each combination of social trail width
and number of social trails (i.e., 1 social trail
of 15 cm, 2 social trails of 15 cm, up to the
maximum number of social trails for each trail
width). Significant decreases were observed in
the number of genotypes, the number of remaining
plants and heather crown cover with the addition
of increased number of social trails, but no
significant changes were observed in allele
frequencies or observed heterozygosity (H,) at the
p = 0.05. The model was then run 20 times for each
social trail width. The impact of human use was
evaluated by calculation of the number of
genotypes, number of plants, and crown cover for
each combination of social trail width and number.
However, because no significant changes in allele
frequencies and H, were observed after the initial
run of the model, these indices were calculated
only for the most severe level of impact for each
social trail width (i.e., 20 social trails of 15
cm, 10 social trails of 56 cm, and 7 social trails
of 102 cm) .
Rainier National Park
David L. Peterson, Temporal and Spatial
Distribution of Trees in Subalpine Meadows of
Mount Rainier National Park, Washington, U.S.A.,
Arctic and Alpine Research, Vol. 28, No. 1, 1996,
pp. 52-59
populations
10 populations of Phyllodoce glanduliflora and 12
populations of P. empetriformis in Mount Rainier
National Park
number for each population
number for each population
whether it was the high or low elevation site for
that local
glanduliflora
populations of P. glanduliflora.
empetriformis
populations of P. empetriformis
at four polymorphic loci in ten populations of
both P. glanduliflora and P. empetriformis in
Mount Rainier N.P.
Hardy-Weinberg expectations averaged over all
subpopulations
population
relative to that over the entire population
Phyllodoce
distance (m) for 10 populations each of P.
glanduliflora and P. empetriformis
empetriformis and 10 populations of P.
glanduliflora. Analysis and phenogram are base on
Nei's genetic identity (1978)
Phyllodoce
among populations of each Phyllodoce species
within each study area
areas for Phyllodoce
sites
of the total number
percent of the total number
loci used in calculations
the total number of genotypes
of the total number of genotypes
populations of heather
greenhouse heather
characters for greenhouse populations of
Phyllodoce
specific local
establishment
including Phyllodoce empetriformis, P.
glanduliflora, Cassiope mertensiana, and Vaccinium
deliciosum.
sitchensis, Lupinus latifolius, and Veratrum
viride
Antennaria lanata, often with lesser amounts of
Carex nigricans
spectablis, Aster alpigenus, and Antennaria lanata
park, dominated by Festuca viridula and Lupinus
latifolius
dates and climate
transformed number of trees established and
climatic variables.
specific local
analysis
coefficient
each site
themselves on each of the five study sites
intervals
each site for a specific 5 year interval
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
populations
10%, 25%, and 40% reduction in number of
genotypes, number of plants, and crown cover
within a 100 square meter study plot.
categories of number of genotypes, number of
plants, heather crown cover
modeled widths that are necessary to cause a 5%,
10%, 25% or 40% reduction in the number of
genotypes, number of plants or heather crown
cover.
>20
Phyllodoce with human impact
numbers of genotypes of Phyllodoce with increased
number of social trails
Rainier National Park
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).
dissertation may be referred to University
Microfilms, 1490 Eisenhower Place, P.O. Box 975,
Ann Arbor, MI 48106, to whom the author has
granted "the right to reproduce and sell (a)
copies of the manuscript"
Cascadia Field Station Box 352100
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