**PROJECT SUMMARY**

**The Proposed Study**

This doctoral dissertation research integrates ecological and
dendrochronological methods to investigate the extent of natural and
anthropogenic impacts on declining whitebark pine communities in Montana
and Idaho. Whitebark pine is limited in distribution to high elevations
in the mountains of western North America, where it has been present
throughout most of the Holocene. Whitebark pine is considered a keystone
species in mutualistic relationships that involve the caching of seeds
by red squirrels and the dispersal of seeds by Clark's nutcrackers.
These caches are a major food source for both grizzly bears and black
bears, which rely on the seeds for up to two-thirds of their annual
diet. The effects of 20^th^ century fire suppression, periodic mountain
pine beetle outbreaks, and white pine blister rust infestations, have
led to a dramatic decline in whitebark pine communities throughout its
native range. These factors, and a lack of research, have hindered the
response of land managers to the dramatic decline in whitebark pine
communities. Consequently, an urgency exists to implement management and
restoration practices that will slow the succession of fire-intolerant
species, and decrease the severity of decline in whitebark pine
communities in western North America.

**Intellectual Merit**

Our project will be the first dendroecological research on whitebark
pine that combines stand dynamics, disturbance regimes, and
climate-forest health interactions in the northern Rocky Mountains. To
understand the stand dynamics of whitebark pine populations, we will
inventory the size, canopy class, and health of whitebark pine and
associated species in overstory and understory fixed-radius plots. To
obtain precise information on the disturbance regime and successional
status of whitebark pine over the past few centuries, we will collect
tree cores from all live and dead standing trees. This component will
provide a temporal scale to the complex interactions in whitebark pine
communities. The results of the dissertation research will answer
theoretical forest succession and applied ecological questions: Is
subalpine fir replacing whitebark pine due to the compounding effects of
white pine blister rust, mountain pine beetle, and fire suppression
during the 20^th^ century? Is succession more rapid on mesic sites
compared to xeric subalpine sites? Can we detect annual mortality dates
of whitebark pine attributed to periodic mountain pine beetle epidemics
and white pine blister rust infestations in the 20^th^ century? And
finally, using tree rings as a proxy, do linkages exist between climate
and mountain pine beetle outbreaks or white pine blister rust intensity?

**Broader Imp acts of the Proposed Research**

*(1) Benefits to society- *Currently, a combination of techniques that
integrate the use of blister rust-resistant whitebark pine, prescribed
fire, and forestry silvicultural techniques is being used to restore
whitebark pine ecosystems. Whitebark pine dendroecological research is
critical because land managers are currently making restoration
decisions on limited ecological data from the 20^th^ century. This
keystone species will likely become extinct within the next two decades
if restoration efforts are not successful, leading to a loss in
biodiversity and population declines in grizzly bears, black bears,
Clark's nutcrackers, and red squirrels.

*(2) Results disseminated broadly and collaboration with federal
agencies- *This research addresses needs for restoration plans by the
USDA Forest Service and the Whitebark Pine Ecosystem Foundation. The
data from this dissertation research will be shared with the USDA Forest
Service, the Whitebark Pine Ecosystem Foundation, and the International
Tree-Ring Data Bank.

*(3) Promoting teaching, training, and learning- *Our project will be
incorporated into educational programs for Tennessee kindergarten-
12^th^ grade students, senior citizen groups, and YMCA summer camps.

*(4) Participation of underrepresented groups- *The principal
investigator of this research is female and we will continue to recruit
underrepresented students to participate in the fieldwork and data
analysis of this dissertation research.

**PROJECT DESCRIPTION 1. Introduction**

*1.1 Doctoral Dissertation Research*

This dissertation research investigates the ecology of whitebark pine
communities using methods of stand dynamics and dendroecology, to
provide a much-needed long-term record of disturbance and stand history
in diminishing whitebark pine communities in western Montana and central
Idaho. The impetus for this research is the growing concern by land
management agencies over the ecological status of this keystone
subalpine species.

*1.2 The Ecological Status of Pinus albicaulis*

Whitebark pine *(Pinus albicaulis *Engelm.) is a long-lived tree species
that is found in many high elevation and subalpine forest communities of
western North America (Arno and Hoff 1989). The species is restricted at
its upper elevations by severe climate conditions and at lower
elevations by competition from other tree species. It is a pioneer
species that fills a crucial niche in watershed protection, catching and
retaining snow, and stabilizing rock and soil in harsh and recently
disturbed areas (Tomback *et al. *2001).

Whitebark pine is also a food source of critical importance to Clark's
nutcrackers *(Nucifraga columbiana), *red squirrels *(Tamiasciurus
hudsonicus), *grizzly bears *(Ursus arctos), *and black bears *(Ursus
americana) *(Mattson *et al. *2001, Tomback 2001). Whitebark pine seeds
are picked and cached by nutcrackers and squirrels, and bears rely upon
these seed caches in the northern Rockies (Mattson *et al. *2001).
Cached seeds that escape predation are in turn a major source of
regeneration for whitebark pine (Tomback *et al. *2001). The
availability of whitebark pine seeds directly influences the number of
human conflicts with grizzly bears that result in management actions
(Mattson and Reinhart 1986). During low seed crop years, more conflicts
occur between grizzly bears and humans as well as high mortality rates
of grizzly bears (Mattson and Reinhart 1986).

The combination of advancing encroachment by fire-intolerant species
such as subalpine fir *(Abies lasiocarpa *(Hook.) Nutt.) and Engelmann
spruce *(Picea engelmannii *Parry ex Engelm.), infestation by mountain
pine beetle *(Dendroctonus ponderosae *Hopk.), and the epidemic of white
pine blister rust *(Cronartium ribicola *JC Fischer) have devastated
whitebark pine populations north of 45°N latitude in western North
America (Hoff *et al. *2001, Kendall and Keane 2001, Tomback *et al.
*2001). Due to natural (mountain pine beetle) and anthropogenic (fire
suppression and the introduction of white pine blister rust) impacts,
the whitebark pine ecosystem of the northern Rockies is diminishing. If
whitebark pine restoration efforts are not successful in the next 15--20
yrs, this species will become extinct (Tomback *et al. *2001).

Ecologists recognize that natural disturbance cycles involving fire,
wind, or pathogens are a normal part of most landscapes and that few
ecosystems ever achieve a steady-state climax (White 1979, Bormann and
Likens 1979, Christensen 1989). Tree reproduction in many forest
ecosystems occurs in episodes associated with major disturbances. Thus,
the distribution of ages in a population is often a sensitive indicator
of the history of disturbance in a stand (Christensen 1989).
Conventional models of succession show that whitebark pine dominates
during early stages of succession and regenerates after stand-level
fires that occur at long return intervals of 200 yrs or more (Fischer
and Bradley 1987). The long lifespan (up to 1000 yrs) of whitebark pine
makes it an important component of mid- and late-successional forests.
Research shows that this ecosystem experiences both stand-replacing
fires, which occur at long intervals and initiate forest succession, and
low-severity surface fires, which occur throughout stand development and
create and maintain an open canopy (Arno and Peterson 1983, Fischer and
Bradley 1987). Stand

replacing fires kill most trees and result in juvenile recruitment,
which alters the age structure of the stand. Low severity fires produce
fire scars on trees but do not dramatically change stand composition.

*1.3 Biogeography of White Pine Blister Rust*

White pine blister rust is an exotic pathogen first discovered in North
America on a currant plant in Geneva, New York, in 1906, and it soon
spread to the Great Lakes region and British Columbia. Eastern white
pine seedlings that had been exposed to white pine blister rust were
sold from tree nurseries in Germany and France to North America from
1890 to 1914 (Tomback *et al. *2001). Blister rust first appeared on
whitebark pine in the coastal range of British Columbia in 1926 and
spread to northern Idaho by 1938 (Childs *et al. *1938). Blanchard and
Tattar (1997) report that white pine blister rust is now found
throughout the entire range of five-needled pines in North America.
Worsening the situation, whitebark pine is also the most vulnerable
white pine species, with fewer than 1 in 10,000 trees showing resistance
to blister rust (Kendall 1994). The range of whitebark pine affected by
white pine blister rust is expanding and infection is intensifying.

White pine blister rust is a heteroecious rust fungi that produces
several spore types and requires two host types to complete its life
cycle (Blanchard and Tatter 1997, Bega 1978). White pine blister rust
alternates between five-needle pines and *Ribes *species. The disease
initially infests a tree through the needles and girdles branches, but
can travel to the main stem, where it usually leads to girdling of the
main stem and death of the tree. Currently, the degree of infestation of
whitebark pine decreases southward throughout all parts of its range,
including the Cascade-Sierra Nevada chain, the Bitterroot Mountains, and
along the Continental Divide of the Rocky Mountains (Hoff 1992).

In Washington State, northern Idaho, northwest Montana, southern
Alberta, and British Columbia, 40-- 100% of the whitebark pine are dead
in most forest stands, and 50--100% of the live trees are infested with
white pine blister rust (Tomback *et al. *2001). Keane (1995) found that
98% of the whitebark pine communities in the Columbia River Basin have
disappeared since the turn of the century. During a project to
reconstruct landscape patterns of whitebark pine in western Montana,
Arno *et al. *(1993) found that 14% of the stands were dominated by
whitebark pine around 1900, but none of these stands were dominated by
whitebark pine in the 1990's. Of the remaining live trees in these
whitebark pine stands, 80% were infected with white pine blister rust,
and more than one-third of their cone-bearing crowns were dead (Arno *et
al. *1993). Furthermore, the extent of forest stands with cone-bearing
trees had declined by half. Because whitebark pine cones form in the top
third of the tree (Kendall 1994), and blister rust kills from the top
down, a tree's ability to produce seed is eliminated long before the
tree dies.

*1.4 Mountain Pine Beetle Ecology*

Mountain pine beetle outbreaks have killed millions of *Pinus *trees
over thousands of square kilometers in the northern Rockies during the
20^th^ century (Romme *et al. *1986). Extensive mountain pine beetle
outbreaks in the northern Rockies occurred from 1920--1940 and
1970--1980 (Arno and Hoff 1989). The major hosts for mountain pine
beetle include, in addition to whitebark pine, ponderosa pine *(Pinus
ponderosa *Douglas ex C. Lawson), lodgepole pine *(Pinus contorta
*Douglas ex Loudon), and western white pine *(Pinus monticola *Douglas
ex D. Don). This native bark beetle constructs "J" shaped egg galleries
in the infected tree's inner phloem. It is the phloem tissue that the
larvae then feed on (Amman *et al. *1989). The beetle galleries impede
water and nutrient transport within the tree and can introduce a
secondary pathogen, a blue staining fungus *(Ophiostoma *spp.) that
clogs the sapwood of living trees (Kipfmueller and Swetnam 2002). The
combination of beetle galleries and blue staining fungus girdles trees
and cuts off nutrient flow, leading to the death of the tree. The
beetles tend to selectively attack larger, older trees, which have a
thicker phloem. Younger trees are usually not killed because they lack
an adequate food supply for the beetles (Cole and Amman 1980).

*1.5 Fire Suppression*

Fire deters succession of fire intolerant tree species and maintains
healthy, productive populations of whitebark pine. Changes in fire
regimes due to fire exclusion have removed fire from this ecosystem,
while increasing competition from fire-intolerant species such as
subalpine fir and Engelmann spruce (Arno 1986, Tomback *et al. *2001).
Reintroducing fire into whitebark pine ecosystems is complicated because
of abruptly changing weather patterns, scattered fuels, and short
growing seasons. In the summer of 2003, wildfires burned many portions
of Montana, including Glacier National Park, Flathead National Forest,
Lolo National Forest, and Gallatin National Forest, and threatened many
cities including Missoula, Butte, and Helena. These fires emphasize the
urgency that exists among fire managers and ecologists to retrieve,
analyze, and archive valuable dendroecological data now before wildfires
permanently erase this information.

*1.6 Research Objectives*

Knowledge of the complex stand dynamics and stand history of whitebark
pine ecosystems are essential to the long-term management and
restoration of this declining keystone species. The primary objectives
of our research are to:

1\. Quantify the changing composition of whitebark pine communities in
mesic and xeric sites and assess the successional status of subalpine
fir and Engelmann spruce in subalpine areas historically dominated by
whitebark pine.

2\. Assess the long-term impacts of human-related (fire-suppression and
the introduction of white pine blister rust) and natural (mountain pine
beetle) disturbances in high-elevation whitebark pine ecosystems.

3\. Reconstruct climate on a multi-century scale to determine the
inter-relationships between climate

change and anthropogenic and natural disturbance regimes present in
whitebark pine ecosystems.

4\. Evaluate the effects of blister rust, mountain pine beetle, and fire
suppression in whitebark pine

ecosystems on a landscape scale (study area ranging from central Idaho
to western Montana).

The Rocky Mountains provide a barrier to the flow of air across western
North America and profoundly influence local climates. Our study areas
are influenced by North Pacific weather patterns and are located in a
transitional zone between continental and maritime climates (Arno and
Hammerly 1984). Continental atmospheric patterns likely affect the
relationship between the ecology of mountain pine beetles, white pine
blister rust, and whitebark pine growth. The interacting feedbacks
between these variables and climate are important for understanding
changing ecosystems.

Tree species present in the high elevation (2000--3000 m above sea
level) study areas include whitebark pine, lodgepole pine, subalpine
fir, and Engelmann spruce. Whitebark pine occurs in open stands and
small groups with other conifers in subalpine and treeline communities
throughout the study area.

**3. Methods**

*Objective 1. Quantify the changing composition of whitebark pine
communities in mesic and xeric sites and assess the successional status
of subalpine fir and Engelmann spruce in subalpine areas historically
dominated by whitebark pine.*

At each study site, we will quantify forest composition and structure
using four 0.05 ha fixed-radius (r = 12.66 m) plots. We will sample
three strata (vertical forest structure) of woody vegetation within each
fixed-radius overstory plot. All trees greater than 5.0 cm in diameter
at breast height (measured at 1.47 m) will be recorded by species and
their dbh measured to the nearest 0.1 cm. Living tree crowns will be
classified into four categories (dominant, co-dominant, intermediate,
and suppressed), based on the amount and direction of intercepted light
(Smith 1986). We will tally all saplings less than 5.0 cm in diameter
but greater than 1.3 cm in diameter at ground level (dgl) in a nested
0.01 ha fixed radius (r = 5.66) plot. Seedlings (less than 1.3 cm dgl)
will be tallied by species in the nested fixed-radius plots. A small
number of subalpine fir saplings will be cut to determine age at ground
level within each stand. We will record site characteristics such as
percent slope and aspect at each overstory plot center. We will also
record UTM coordinates using a GPS unit to map each study site. The
length and width of dead woody debris within the overstory plots will be
measured to calculate the basal area and accumulation of fire fuels on
the forest floor.

We will determine whether succession is more rapid on mesic sites
compared to xeric whitebark pine subalpine forests, using Euclidean
distance to quantify changes in species composition through time between
subalpine forest communities (Veblen *et al. *1994). In addition, we
will calculate relative dominance and relative density (Cottam and
Curtis 1956), for each tree species to compare the importance of each
species within a forest stand. Canopy class and diameter class
distributions will also be calculated to compare both horizontal and
vertical forest structure for all species at each site.

*Objective 2. Assess the long-term impacts of anthropogenic (fire
suppression and the introduction of white pine blister rust) and natural
(mountain pine beetle) disturbances in high-elevation whitebark pine
ecosystems.*

Dendroecological techniques will help evaluate the age structure and
tree-growth patterns to document changing stand conditions related to
disturbance, stand development, or climatic variation (Payette *et al.
*1990, Foster *et al. *1996). Dendroecological techniques can also help
assess relationships between climate, site conditions, and tree growth
to evaluate factors that influence the growth of a plant community (Cook
and Kairiukstis 1989). Crossdating makes it possible to extend stand
histories beyond one generation of trees or one major disturbance event
by using narrow and wide ring-width patterns of living and dead trees
(Foster *et al. *1996). Crossdating consists of combined traditional
techniques of skeleton plotting, a graphical technique of ring-width
comparison (Stokes and Smiley 1968), and the use of a crossdating
quality control program, COFECHA (Holmes 1983, Grissino-Mayer 2001).

Dendroecological techniques and analyses will be used to provide
high-quality, temporally precise information on the ecological status of
whitebark pine ecosystems. We will sample all live and standing dead
trees in each plot for age using increment borers. Cores will be
extracted 30 cm above ground level and multiple cores will be sampled to
ensure they intercept or are near the pith. We will obtain two cores
from each standing tree (healthy, declining, and dead) in each plot.
Samples will be processed and prepared for dating using methods outlined
in Stokes and Smiley (1968). We will measure ring widths with a Velmex
measuring stage at 0.001 mm precision in conjunction with J2X software.
These measurement series will then be statistically compared to ensure
accurate crossdating using the computer program COFECHA (Holmes 1983,
Grissino-Mayer 2001). We will standardize and combine tree-ring
measurements into a master chronology using the software package ARSTAN
(Cook 1985). We will date standing snags by crossdating each unknown
sample with the master chronology derived from live trees, specific for
each species and stand. The outermost ring of each increment core from
these standing snags will provide an estimate of the year of tree death
(Daniels *et al. *2004).

Forest stand dynamics and disturbance patterns can be inferred from
suppression-release patterns in tree rings (Brubaker 1987). To quantify
the extent of the disturbance (fire, mountain pine beetle, or white pine
blister rust) on stand and landscape scales, we will calculate releases
of tree growth over time in our whitebark pine chronologies. The
occurrence of releases in trees within and between study sites can
identify the disturbance as either a regionally extensive or localized
disturbance event (Veblen *et al. *1994). Individual disturbance events
will later be disentangled by inspecting tree cores for blue stain
fungus from mountain pine beetle outbreaks and by finding declines in
tree growth over a number of decades, indicating a possible white pine
blister rust infestation. We will calculate minor and moderate release
events, which are defined as changes in growth with respect to
subsequent 10-yr growth periods (McCarthy and Bailey 1996, Nowacki and
Abrams 1997, Schuler and Fajvan 1999) using a percent growth change
equation:

\% GC = ((M2 -- M1) / M1) \* 100

where % GC is percent growth change from preceding to superceding 10-yr
radial means, M1 is the preceding 10-yr mean radial growth (exclusive of
the current year), and M2 is the superceding 10-yr mean radial growth
(inclusive of the current year) (Schuler 1999). Major releases will be
calculated using the same formula, except radial growth will be measured
over a 15-yr period. Major releases will have a percent growth change
over 100% for 15 yrs, moderate releases will have a percent growth
change over 50% for 10 yrs, and minor releases will have a growth change
greater than 25% for 10 yrs.

*Objective 3. Reconstruct climate on a multi-century scale to determine
the relationship of climate to the anthropogenic and natural disturbance
regimes present in whitebark pine ecosystems*

Meteorological data (temperature, precipitation, and the derived Palmer
Drought Severity Index) for the past 110 yrs in the central mountain
regions of Montana and Idaho are available from the National Climatic
Data Center in Asheville, North Carolina. We will use these data for
climate analysis of the whitebark pine and subalpine fir master
chronologies. We will conduct response function and correlation analysis
with state divisional climate records to assess whitebark pine and
subalpine fir annual ring-width response to monthly average temperature,
total monthly precipitation, and monthly Palmer Drought Severity Indices
(PDSI) from 1895--2004 (Grissino-Mayer 1996). Lagged variables will also
be created to calculate the relationship between ring width and
temperature, precipitation, and PDSI in the previous year. PDSI is
calculated using temperature and rainfall information in a formula
designed to determine drought conditions.

To measure the preconditioning effects of monthly climate data on tree
growth, we will use the program PRECON (Fritts and Shashkin 1995).
PRECON is software that statistically analyzes the relationship between
climate and tree-ring variation. Tree-ring chronology data are
calibrated with site-specific regional or local monthly climate data
using stepwise multiple regression, bootstrapped response function
analysis, correlation analysis, and a Palmer Drought Severity Index
growth model (Fritts and Shashkin 1995). Quantifying the preconditioning
effects of climate on tree growth will help clarify the variance
explained by climate versus anthropogenic and natural disturbance
regimes.

Moisture stress is widely regarded as an important contributor to
conifer mortality from bark beetles (Craighead 1925, Thomson and
Shrimpton 1984, Safranyik 1989). Rapid beetle population expansion can
be caused by a drought over a large area that predisposes forest stands
to beetle attacks (Safranyik 1989). Extensive mountain pine beetle
outbreaks have occurred historically in our study sites from 1920-- 1940
and 1970--1980 (Arno and Hoff 1989). We will use correlation and
regression techniques to determine the strength of temperature,
precipitation, and PDSI variables on mountain pine beetle-induced
mortality in the 20^th^ century.

*Objective 4: Evaluate the efects of blister rust, mountain pine beetle,
and fire suppression in whitebark pine ecosystems on a landscape scale
in a study area ranging from sites in central Idaho (Boise National
Forest) to sites in western Montana (Beaverhead-Deerlodge and Lolo
National Forests).*

By evaluating both live and dead trees, we will address both theoretical
and applied ecological concepts that will help land management agencies
achieve restoration goals. For all live, declining, and dead whitebark
pine trees, we will note the presence or absence of mountain pine beetle
galleries in the phloem on the stems of the trees in each plot. We will
later examine tree cores from live, declining, and dead trees for the
presence of blue stain fungus, which is inoculated by the mountain pine
beetle (Daniels *et al. *2004). If blue stain fungus is found in the
tree cores, the death of the tree was caused during a mountain pine
beetle outbreak.

The presence of white pine blister rust will be evaluated on the health
of tree crowns (brown needles, flag branches), and on the presence of
cankers on the branches and stems of whitebark pine, following the
methods of the Whitebark Pine Ecosystem Foundation (WPEF 2004). We will
compare sites that are heavily infested with white pine blister rust
(Lolo National Forest) to stands where white pine blister rust is absent
(Beaverhead-Deerlodge and Boise National Forests) to determine if there
is a unique tree-ring pattern indicating when white pine blister rust
first became present within the forest stands.

**4. Preliminary Results**

During a two-week research trip in June 2004, I led an expedition with
two fellow graduate students to collect forest composition and tree-ring
data in four whitebark pine-dominated stands in the Lolo and
Beaverhead-Deerlodge National Forests. We cored over 500 trees and
inventoried seven overstory plots, but were unable to collect data on a
more comprehensive spatial scale due to lack of funding and time
constraints.

In a 2003 pilot study, we found that whitebark pine in Montana are
sensitive to disturbance and environmental change as shown by the
variability in growth in many of our whitebark pine cross-sections. We
conducted a preliminary dendroecological assessment of 20 cross-sections
taken from whitebark pine stumps and snags at Morrell Mountain in the
Lolo National Forest, Montana. This initial chronology provides
dendroecological information for the past 475 years and has the
potential to greatly improve our understanding of disturbance regimes in
whitebark pine communities. The whitebark pine master tree-ring
chronology (N=20) ring-width index (RWI) values ranged from 0.392 to
1.484 from 1507--1982 *(Figure 2). *The interseries correlation between
the whitebark pine series (samples) was 0.522 with an average mean
sensitivity of 0.242, both strong values when compared to other research
in high mountain ecosystems *(e.g. *LaMarche 1974, Perkins and Swetnam
1996), suggesting whitebark pine, as a species, is likely to be
sensitive to disturbance events and environmental change.

We found a strong decrease in radial growth in the past 50 years in many
of the whitebark pine samples *(Figure 2) *which may be caused by
several exogenous disturbances, including mountain pine beetle
outbreaks, blue stain fungus, and white pine blister rust (McDonald and
Hoff 2001, Kendall and Keane

2001), but more data are required to determine the landscape level
impacts of these disturbance agents. We hope to investigate this decline
in whitebark pine growth with other sites in the Lolo,
Beaverhead-Deerlodge, and Boise National Forests to investigate the
spatial scale of this pattern. We can determine whether the growth
decline is caused by white pine blister rust by comparing the last 50
years of growth with our subalpine fir chronologies, which would not
exhibit this pattern. The unique 50-year tree-ring pattern in our
samples is the first whitebark pine study to record this growth decline
in western North America.

Our successful dating of these high-elevation samples indicates that
whitebark pine is a viable species to use in dendroecological
reconstructions of past disturbances and environments. Using this
species as a proxy, we will be able to build a more spatially
comprehensive disturbance history of whitebark pine at both the stand
and landscape levels in the northern Rockies.

Our whitebark pine samples were also significantly correlated (p \<
0.01) to the Palmer Drought Severity Index (PDSI) from the
previous/current year's winter. Over the past century, whitebark pine in
Montana appear to be responding primarily to snowpack conditions during
the previous December. The significant relationship found between
whitebark pine growth and PDSI may help clarify how climate relates to
the ecology of whitebark pine ecosystems (including mountain pine beetle
outbreaks and the white pine blister rust disease cycle). My
dissertation research will expand on the climate reconstruction research
of whitebark pine by comparing the climate-forest health interactions of
this threatened subalpine species on a multi-century scale. Further
research and analysis will provide a regional dendroecological
comparison between our study sites in western Montana and central Idaho
to sites in Idaho investigated by Perkins and Swetnam (1996).

*Figure 2. Master whitebark pine chronology from 150 7-1982. Sample
depth (N) throughout the 475-year chronology. indicates the 1601 frost
ring, which is a marker ring found in whitebark pine chronologies in the
northern Rockies (Perkins and Swetnam 1996, Kipfmueller 2003). The boxed
area highlights the recent 50-year decline in growth found in our
samples.*

**5. Research Structure**

Leading USDA Forest Service researchers from Missoula, Montana,
approached Henri Grissino-Mayer at the 2003 North American
Dendroecological Fieldweek to encourage a dendroecological investigation
of this keystone species. While I attended "Monitoring Whitebark Pine
for Blister Rust: A Methods Workshop," held by the Whitebark Pine
Ecosystem Foundation and in West Yellowstone in 2004, I interacted with
ecologists, foresters, and researchers from Montana and Idaho who
offered advice, resources, and research sites for further whitebark pine
ecological research. Communication channels and research logistics are
established with USDA Forest Service researchers and ecologists in the
Lolo,

Beaverhead-Deerlodge, and Boise National Forests. Over the next two
summers, it is feasible to inventory and core trees in 32 plots (eight
whitebark pine-dominated stands with four plots in each) in Montana and
Idaho.

Working closely with Federal Cooperators from the USDA Forest Service
and the Whitebark Pine Ecosystem Foundation, I will select and
investigate eight whitebark pine-dominated stands growing at 2000--3000
m elevation in the Lolo, Beaverhead-Deerlodge, and Boise National
Forests. I have been in contact with USDA Forest Service researchers and
ecologists since January 2004, developing plans for accessible research
locations, expedition accommodations, and research logistics. In the
Lolo National Forest, Bob Keane (Research Ecologist, Rocky Mountain
Research Station) suggested three research sites that are located on
remote mountain peaks accessible only by using 4-wheel drive vehicles on
unimproved Forest Service roads. Cathy Stewart (Fire Ecologist, Lolo
National Forest) signed our research permits and acquired necessary keys
for access to these remote locations. Elaine Kennedy Sutherland
(Research Biologist and Project Leader, Rocky Mountain Research Station)
provided registration fees for me to attend "Monitoring Whitebark Pine
for Blister Rust: A Methods Workshop," sponsored by the Whitebark Pine
Ecosystem Foundation, in West Yellowstone, from June 28--29, 2004. Diane
Hutton (Fire Management Officer, Beaverhead-Deerlodge National Forest)
allowed us to use Forest Service housing and took me to research sites,
some of which have been devastated by mountain pine beetle outbreaks in
the Beaverhead-Deerlodge National Forest.

This Doctoral Dissertation Research Improvement Grant would fund two
research expeditions totaling six weeks of fieldwork in July 2005 and
July 2006 *(Table 1). *I would use the requested funds to cover travel
for two graduate field assistants and myself during both expeditions.
Our research sites are isolated and located in an area of rugged terrain
with unpredictable mountain weather. We often hike 6--12 km a day, in
mountain weather, with an elevation change of at least 500 m. Although
the fieldwork is challenging, we are able to maximize our time in the
field by camping on site and by being ready for intense fieldwork. With
a three-person team, we can core 80 trees and inventory at least one
overstory plot per field day. In June 2004, we conducted fieldwork for
three days at each research site in the Lolo National Forest. In the
Beaverhead-Deerlodge National Forest, Diane Hutton supplied us with free
housing in Forest Service cabins and has promised to continue supporting
us over the next two summers. We intend to expand our study into other
subalpine sites in the Lolo, Beaverhead-Deerlodge, and Boise National
Forests to develop a better understanding of the spatio-temporal
dynamics of changes in the disturbance regimes of whitebark pine
communities.

*[Table 1. Research schedule from 2005 ]{.underline}[--]{.underline}[
2007.]{.underline}*

*Stand and Tree-Ring *X \* \* \* *Data Collection*

*Process Wood *X X *Samples*

*Measure Samples *X *Chronology*

*Development *X

*Statistical Analysis of *X

*Stand Data*

*Conference *X *Presentations*

*Write Dissertation*

*[Submit Dissertation]{.underline} * \*\*\*Beginning and end of DDR NSF
funds (24 months)

**6. Research Significance and Intellectual Merit**

The longevity of whitebark pine trees provides an opportunity for
temporally extensive tree-ring reconstructions of past disturbance
regimes and ecological events. In addition, the sensitivity of many
subalpine species to changes in their environment suggests whitebark
pine may also be an excellent indicator of global climate change
(LaMarche and Stockton 1974, Fritts 1976). The annual nature of
tree-ring formation and the ability to statistically link tree growth
with climate are important tools for understanding climate variability
on a temporal scale (Kipfmueller 2003). Perkins and Swetnam (1996)
successfully built a whitebark pine tree-ring chronology back to the
10th century, and while they found their samples difficult to date, they
suggest that whitebark pine could be used to relate disturbance regimes
and climate on a multi-millennial scale. This project will be the first
dendroecological research on whitebark pine that combines stand
dynamics, disturbance regimes, and climate-forest health interactions in
the northern Rocky Mountains. The combined results of stand and
tree-ring data will provide a much-needed long-term record of
disturbance history in whitebark pine ecosystems in Montana and Idaho.

The University of Tennessee's Laboratory of Tree-Ring Science (LTRS),
directed by Drs. Henri Grissino-Mayer, Sally Horn, and Ken Orvis, is
internationally recognized as one of the leading and best-equipped
dendrochronological research facilities in the United States. The
laboratory is funded by the Department of Geography and the College of
Arts and Sciences, and external research grants. Dendrochronological
research conducted in the LTRS involves both human and natural processes
that affect the environment, and has produced successful projects that
include archaeology, fire history, and climate reconstruction.

**7. Broader Imp acts of the Research**

***7.1 Benefits to society- ***This investigation was undertaken because
of a need for a better ecological understanding of declining whitebark
pine ecosystems. Whitebark pine restoration efforts must succeed in the
next 15--20 years in order for this species to avoid extinction, which
would dramatically impact wildlife species such as the grizzly bear,
black bear, Clark's nutcracker, and the red squirrel. Quantitative
information on the complex stand dynamics and dendroecology of whitebark
pine ecosystems are essential to the long-term management and
restoration of this declining keystone species. Whitebark pine
dendroecological research is important because land managers are
currently making restoration decisions based on limited ecological data
from the 20^th^ century.

***7.2 Results disseminated broadly and collaboration with federal
agencies- ***Whitebark pine stand data will be shared with the Whitebark
Pine Ecosystem Foundation to add to their newly-developed white pine
blister rust survey data across western North America. We will provide
copies of all data to the USDA Forest Service and the Whitebark Pine
Ecosystem Foundation. Tree-ring data in the form of raw tree-ring
measurements and chronologies will be submitted to the International
Tree-Ring Data Bank (ITRDB), which is one of the premier paleoclimatic
databases in the world. The ITRDB contains measurements and tree-ring
chronologies for over 100 tree species from over 1500 sites around the
world. This website allows researchers to easily access and download one
or more of these data sets to use in dendrochronological studies. Our
tree-ring data will add needed data points to the paleoclimate record in
North America.

***7.3 Promoting teaching, training, and learning- ***We will present
our research findings at annual meetings of the Whitebark Pine Ecosystem
Foundation and at the 7th International Conference of Dendrochronology
which will be held in June 2006, in Beijing, China. Data collected
during this research project will have a broad impact on not only the
scientific community, but on public outreach programs in Tennessee as
well. Locally, information gathered from this research will be
integrated into programs run by the Tennessee Geographic Alliance and
will be presented to a variety of audiences including
kindergarten--12^th^ grade students, senior citizen groups, and local
summer camps (YMCA). This year, I am the Coordinator of Geographic
Awareness Week (GAW) at the University of Tennessee

through the Tennessee Geographic Alliance. GAW is a national program
sponsored by The National Geographic Society to increase
kindergarten-12^th^ grade student awareness of geography-related issues.
I will also be a guide for the Tennessee Geographic Alliance at the
University of Tennessee Earth and Science Fair (7th-1 2th grade
students) in October. I have participated in outreach tree-ring programs
organized through the Laboratory of Tree-Ring Science (LTRS). This
summer I helped teach students (ages 8--16) at a YMCA summer camp how to
use tree rings as a proxy to study fire, climate, and land-use change. I
will continue to participate in outreach programs and will incorporate
our research on endangered whitebark pine ecosystems into these
interpretive programs.

***7.4 **Participation of underrepresented groups- ***This dissertation
research is being conducted by a female Ph.D. student who has extensive
field experience. Saskia van de Gevel has served as mentor and field
supervisor to ten field assistants during her graduate career at
Southern Illinois University and the University of Tennessee. We will
continue to recruit underrepresented students to participate in the
fieldwork and data analysis of this dissertation research. We will also
try to involve underrepresented students during this research from the
department's Research Experiences for Undergraduates course.

**Results from Prior NSF Support: Henri D. Grissino-Mayer**

**BCS-9809245: ***Historic expansion of western juniper on near-relict
sites: a dendroecological approach, *1998--2001, \$190,376. Co-PIs: Dr.
Paul A. Knapp, Department of Anthropology and Geography, Georgia State
University, and Dr. Peter T. Soulé, Department of Geography and
Planning, Appalachian State University. Our primary objective was to
determine mechanisms responsible for changes in western juniper
establishment, expansion, and growth rates in relict and near-relict
sites of central and southern Oregon. Our study investigated the
competing roles of land-use history, climate, biological inertia, human
and natural disturbances, and elevated CO2 on western juniper forests.
This study was designed to determine if western juniper trees growing
under natural conditions were responding to CO2 fertilization. Our
results have been disseminated at regional and national meetings and in
several peer-reviewed journals, including *Geophysical Research Letters,
Global Change Biology, Quaternary Research, Professional Geographer,
Journal of Climate, *and *Ecological Applications.*

**BCS-0327280: ***Oxygen isotope compositions of tree-ring cellulose as
a high-resolution proxy record of hurricane activity, *2003--2005,
\$149,999. Co-PI: Dr. Claudia I. Mora, Department of Earth and Planetary
Sciences, University of Tennessee. This research currently investigates
the potential use of oxygen isotope ratios of alpha cellulose in tree
rings of yellow pines from southern Georgia and South Carolina as a
proxy record of hurricane activity. We've developed a 227-year record of
oxygen isotope compositions of alpha cellulose in slash and longleaf
pine tree rings that preserves evidence of past tropical cyclone
activity, seasonal droughts, and multidecadal climate oscillations and
suggests the potential for a detailed record of tropical cyclone
occurrence extending back many centuries. The trends in hurricane
frequency closely track changes in larger-scale ocean-atmosphere
circulation features, especially the North Atlantic Oscillation, which
we believe may significantly moderate hurriance activity. We've
presented our initial findings at national meetings, including the
Association of American Geographers, the American Geophysical Union, and
the Geological Society of America, and our first manuscript was recently
accepted by the journal *Geology.*

**BCS-0242155: ***Establishing a multi-century, annual-resolution
climate record from the northeastern Caribbean, *2003--2005, \$153,072.
Co-PIs: Dr. Kennth H. Orvis and Dr. Sally P. Horn, Department of
Geography, University of Tennessee. This research project builds on the
finding that *Pinus occidentalis *Swartz, the native pine of the
Caribbean island of Hispaniola, produces excellent tree rings that are
moisture and temperature sensitive. Initial analyses, primarily of
younger samples, has yielded a well-replicated master chronology dating
to the late 1 800s and continuous, non-crossdated records to the late
1500s. Many older samples remain to be analyzed. The chronology
correlates positively (r = 0.44, p \~ 0.001, n = 51) with January--March
(late dormant season) rainfall days on the windward mountain flank, and
negatively (r = --0.46, p \< 0.001, n = 50) with February--June (early
growing season) temperature on the leeward flank. The latter represents,
to our knowledge, the first tropical-region dendroclimatological
temperature record. The primary signal apparently reflects the timing of
onset of the rainy season, which is very likely teleconnected to
large-scale global climate oscillations.

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